Taxonomia De Spionidae (Annelida) Da Bacia Do Espírito Santo E Porção Norte Da Bacia De Campos

Universidade Federal do Rio de Janeiro Instituto de Biologia Centro de Ciências da Saúde Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva

Antônio João Malafaia Peixoto

Taxonomia de Spionidae (Annelida) da Bacia do Espírito Santo e Porção Norte da Bacia de Campos

Rio de Janeiro 2020

Ficha catalográfica

Peixoto, Antônio João Malafaia. Taxionomia de Spionidae (Annelida) da Bacia do Espírito Santo e porção norte da Bacia de Campos / Antônio João Malafaia Peixoto. -- Rio de Janeiro: UFRJ / Instituto de Biologia, 2020. 524 f.: il.; 30 cm. Orientador: Paulo Cesar de Paiva. Tese (doutorado) – UFRJ, Instituto de Biologia, Programa de Pós- Graduação em Biodiversidade e Biologia Evolutiva, 2020. Referências: 515-524. 1. Classificação. 2. Anelídeos. 3. Poliquetos. 4. Biodiversidade. 5. Biologia do Desenvolvimento. Biodiversidade - Tese. I. Paiva, Paulo Cesar de. II. Universidade Federal do Rio de Janeiro, Instituto de Biologia, Programa de Pós-Graduação em Biodiversidade e Biologia Evolutiva. III. Título.

Ficha catalográfica elaborada pela equipe do Setor de Referência da Biblioteca Central do Centro de Ciências da Saúde (CCS) da Universidade Federal do Rio de Janeiro (UFRJ).

Antônio João Malafaia Peixoto

Taxonomia de Spionidae (Annelida) da Bacia do Espírito Santo e Porção Norte da Bacia de Campos

Tese de doutorado apresentada ao Programa de Pós-Graduação em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, como requisito parcial à

obtenção do título de Doutor em Ciências Biológicas (Biodiversidade e Biologia Evolutiva).

Orientador: Prof. Dr. Paulo Cesar de Paiva

Rio de Janeiro 2020

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“The journey of a thousand miles must begin with a single step.”

– Lao Tzu

“All progress takes place outside the comfort zone.”

– Michael John Bobak

“Success is not the key to happiness. Happiness is the key to success. If you love what you are doing, you will be successful.”

– Albert Schweitzer

Agradecimentos

Primeiramente, gostaria de agradecer imensamente ao meu orientador, Paulo Paiva, pela amizade, conselhos, apoio, ideias, dedicação, disponibilidade e por acreditar no meu potencial. Com a sua orientação, esse trabalho melhorou em qualidade e já gerou frutos – com artigos publicados e espécies descritas.

Aos meus amigos do Laboratório de Polychaeta: Victor Seixas, Natália Amaral, Monique dos Santos, Alana Leitão, Gustavo Mattos, Letícia Carvalho, Letícia Marinho, Stephanie Vaz, Mirian Sathler, Rodolfo Otávio, Rafael Reis, Pedro Tamanini, Renata Pedrone, Ricardo Guimarães e Ricardo Dias. Um abraço especial para a Stephanie e Natália, pela ajuda com o Photoshop.

Agradeço imensamente aos amigos Natália Ranauro, Isabela Guerra, Jaqueline Carvalho e Vinícius Miranda pela amizade, momentos de divertimento, conversas de WhatsApp, fofocas, besteirol e pelo interesse em comum nos poliquetas!

Às professoras Michelle Klautau, Alexandra Rizzo e Cinthya Santos pela participação na minha banca de qualificação – e pelos comentários, sugestões, perguntas e críticas, que resultaram num manuscrito de qualidade superior.

Um abraço para a Alexandra, Cinthya e Ana Brasil, poliquetólogas incríveis que vem acompanhando e inclusive já fizeram parte da minha evolução acadêmica.

Agradeço ao programa de Pós-Graduação em Biodiversidade e Biologia Evolutiva, ao secretário Heber Araújo e em especial aos professores do curso, pela diversidade e relevância das disciplinas oferecidas, além de constante oferta de novas disciplinas.

Ao Smithsonian National Museum of Natural History (Washington DC – USA), pela concessão da bolsa “Kenneth Jay Boss Fellowship in Invertebrate Zoology”, vital para esse trabalho, pois permitiu que o material tipo de diversas espécies desse projeto pudesse ser reexaminado. Agradeço ainda à Karen Lee, pela acomodação durante minha estadia em Washington, à Karen Osborn, por atuar como minha orientadora no Smithsonian e à Karen Reed, pela recepção e pelas orientações no Smithsonian Musem Support Center.

À Tatiana Steiner e ao Museu de Zoologia da Universidade Estadual de Campinas "Adão José Cardoso" pelo que deve ser um dos empréstimos mais longevos da coleção – 2015 e contando. Agradeço também à professora Joana Zanol e à técnica Monique dos Santos pelo empréstimo de material e por me receberem para examinar material da coleção do Museu Nacional e ao professor

Marcelo Fukuda, pela oportunidade de examinar material da coleção do Museu de Zoologia da Universidade de São Paulo.

Agradeço à toda equipe de técnicos pelo auxílio em seções de microscopia eletrônica de varredura e no preparo das amostras: Roger Silva e Taíssa de Oliveira (Plataforma de Microscopia Eletrônica Rudolf Barth – Fundação Oswaldo Cruz), Marcelo Sales (Laboratório de Protistologia – UFRJ), Otávio Pacheco (Laboratório de Ultraestrutura Celular Hertha Meyer - UFRJ) e à Camila Messias (Centro de Microscopia Eletrônica de Varredura – Museu Nacional). Agradeço especialmente ao Scott Whittaker (Scanning Electron Microscopy Lab – Smithsonian National Museum of Natural History), pelo treinamento para utilizar o microscópio eletrônico por conta própria.

À Nataliya Budaeva, pela oportunidade de participar do International Course on Annelid Systematics, Morphology and Evolution (Bergen – Noruega) e à Katharina Jörger, pela oportunidade de participar do VW-Summer School Exploring the marine meiofauna of the Azores – from discovery to scientific publication (Ponta Delgada – Portugal), cursos que me renderam parcerias, novos amigos, contato com táxons raros e experiência de pesquisa, além de conhecer lugares inesquecíveis.

Agradeço à Benthos Ambiental pela triagem e taxonomia a nível de família do material utilizado nesse trabalho. Agradeço aos amigos Thiago Rohr, Ilana Sallorenzo, Bárbara Calçado, Vanessa Berenguer, Beana Simões, Renata Pacheco, Natalia Suita, Maíra Mendes, Paula Raposo, Gustavo Albuquerque, Erika Klein, Rafael Reis, Dona Francisca e Cinthia Bianco. Agradeço especialmente à Cinthia por me passar sua experiência em taxonomia geral e pela parceria na taxonomia inicial dos Spionidae e ao Thiago e à Ilana pelas oportunidades recebidas e ao convite para trabalhar com os Spionidae.

À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) e à Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) pelo auxílio financeiro recebido.

Agradeço ainda aqueles que acompanharam boa parte da minha carreira acadêmica, mas não puderam estar fisicamente presentes nessa etapa: minha avó Eleone de Lyra Peixoto, minha tia Sheila Gabardo e meu melhor amigo Marcelo Elysio Lugarinho.

Ao Maurício Shimabukuro e ao Orlemir Carrerette (Laboratório de Ecologia e Evolução de Mar Profundo – USP), pela doação de material adequado para análises moleculares de algumas das espécies reportadas nesse projeto.

Um agradecimento especial e votos de boa sorte e muita força de vontade a todos os mestrandos e doutorandos devido aos tempos que estamos vivendo, com desvalorização da ciência,

VII obscurantismo e cortes de bolsas – incluindo um ministro da educassaum (sic) e um ministro do meio ambiente claramente ineptos e que estão atuando contra suas pastas.

Agradeço à minha família pelo total apoio desde o momento que escolhi ser biólogo, por celebrar minhas conquistas e pelo interesse pelo meu trabalho.

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Resumo

Anelídeos da família Spionidae pertencem ao táxon Sedentaria – um dos maiores grupos dentro de Annelida, atualmente incluindo os táxons Echiura – tradicionalmente tratado como um filo separado e Clitellata – habitualmente considerado uma classe de Annelida. A família é facilmente coletada em ambientes marinhos, se destacando por sua frequência de ocorrência, diversidade e abundância –potencialmente sendo o grupo dominante em ambientes eutrofizados.

Apesar da sua abundância, o conhecimento taxonômico da família na costa Brasileira ainda é muito limitado, com muitos registros de espécies descritas para outras regiões geográficas, como a Europa, América do Norte e Antártica. Os poucos estudos taxonômicos de Spionidae para a costa Brasileira correspondem a estudos em águas rasas (profundidade inferior a 50 metros) nas regiões Sul e Sudeste, enquanto extensas regiões da costa, como as regiões Norte e Nordeste permanecem praticamente desconhecidas, com escassos estudos taxonômicos e nenhuma nova espécie descrita.

Espionídeos de ambientes de plataforma continental abaixo de 50 metros de profundidade e ambientes de mar profundo são igualmente desconhecidos, com estudos geralmente associados à projetos de caracterização ambiental anterior à atividade petrolífera. O presente trabalho, realizado com material coletado na Bacia do Espírito Santo e porção Norte da Bacia de Campos, engloba diferentes substratos inconsolidados do sublitoral raso até o talude continental (25 a 3000 metros de profundidade).

Foram examinados 16.938 espécimes, pertencentes a 17 gêneros e 64 espécies. Os gêneros Spiogalea, Laubieriellus e Aonidella são novas ocorrências para a costa brasileira, enquanto 44 dessas espécies são novas para a ciência. Adicionalmente, material coletado nesse projeto de algumas espécies tradicionalmente tratadas como amplamente distribuídas – como Boccardia polybranchia, Prionospio fauchaldi e Paraprionospio tamaii foi examinado e revelou- se tratar de espécies novas para a ciência.

Palavras-chave: Taxonomia, Annelida, Spionidae, morfologia, diversidade.

Abstract

Annelids belonging to the family Spionidae are part of the taxa Sedentaria – one of the largest groups within Annelida, currently encompassing the taxa Echiura – traditionally regarded as a separate phylum and Clitellata – habitually regarded as a class of Annelida. The family is easily sampled in marine environments, standing out for its occurrence frequency, diversity and abundance – potentially being the dominant group in eutrophic environments.

Despite its abundance, the family’s taxonomic knowledge on the Brazilian coast is limited, with many records of species described to distinct geographic regions, such as Europe, North America and Antarctica. The few Spionidae taxonomic studies to the Brazilian Coast correspond to shallow-water studies (depth below 50 meters) on the South and Southeast regions, while large extension of the coast, such as the North and Northeast regions remain largely unknown, with scarce taxonomic studies and no new species described.

Spionidae from continental shelf environments below 50 meters depth and deep-sea environments are equally unknown, with studies generally associated to environmental characterization projects prior to oil drilling operations. This work, performed using samples from the Espírito Santo Basin and a portion of Campos Basin, comprises different soft-bottom substrates from the shallow sublittoral to the continental slope (25 to 3000 meters depth).

A total of 16,938 specimens, belonging to 17 genera and 64 species. The genera Spiogalea, Laubieriellus and Aonidella are new records to the Brazilian coast, while 43 species are new to science. Furthermore, material collected in this work belonging to species traditionally regarded as having a wide distribution – such as Boccardia polybranchia, Prionospio fauchaldi and Paraprionospio tamaii was examined and turned out to be new species.

Key-words: Taxonomy, Annelida, Spionidae, morphology, diversity.

Lista de Figuras

Figura 1. Morfologia geral 1 ...... 10

Figura 2. Morfologia geral 2 ...... 12

Figura 3. Morfologia geral 3...... 14

Figura 4. Área de coleta do Projeto...... 17

Figura 5. Desenho amostral e faciologia da Foz do Rio Doce...... 18

Figura 6. Desenho amostral e faciologia da Plataforma continental e talude continental...... 19

Figura 7. Desenho amostral e faciologia dos cânions Doce e Watu Norte...... 20

Sumário

1. Introdução...... 1

1.1 Histórico taxonômico e classificação de Spionidae...... 3

1.2 Diversidade de Spionidae na costa Brasileira...... 6

1.3 Morfologia geral...... 9

2. Objetivos...... 15

2.1 Objetivo geral...... 15

2.2 Objetivos específicos...... 15

3. Materiais e Métodos...... 16

3.1 Área de Estudo, coleta e processamento das amostras...... 16

3.2 Análise do Material...... 20

4. Resultados...... 23

4.1 Capítulo I………………………………………………………………………..24 A new species of Spiogalea (Polychaeta: Spionidae) from Brazil, with an amended diagnosis of the genus

4.2 Capítulo II………………………………………………………...…………….38 New Prionospio and Laubieriellus (Annelida: Spionidae) species from Southeastern Brazil

4.3 Capítulo III……………………………………………………………………...63 New apinnate Prionospio (Annelida: Spionidae) species from southeastern Brazil

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4.4 Capítulo IV…………………………………………………………………….128 Species from the Prionospio (Annelida: Spionidae) complex from southern Brazil, including new species from Prionospio and Paraprionospio

4.5 Capítulo V……………………………………………………………..……….228 Contributions to the taxonomy of Aonides, Aonidella and Laonice from southeastern Brazil, including the description of five new species

4.6 Capítulo VI…………………………………………………………..………..305 Spionidade (Annelida) from Espírito Santo Basin, southeastern Brazil: the genera Scolelepis, Dispio, Spio, Microspio and Malacoceros, including a new deep-sea Scolelepis species

4.7 Capítulo VII……………………………………………………..……………..402 A contribution to the taxonomy of Spiophanes (Annelida: Spionidae), including new species 4.8 Capítulo VIII……………….……………………………………………..…...446 Polydorids (Annelida: Spionidae) species from southern Brazil, with new records and a new Boccardia species

5. Considerações finais...... 501

6. Referências bibliográficas...... 504

Anexo 1 – Morfotipagem de espécies no projeto HABITATS...... 514

Anexo 2 – Chave para os gêneros de Spionidae reportados nesse trabalho...... 517

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De acordo com o disposto no Artigo 9 do Código Internacional de Nomenclatura Zoológica (ICZN), esta tese não constitui publicação efetiva para os nomes aqui propostos. Os táxons aqui descritos serão considerados válidos somente a partir da publicação dos manuscritos aqui apresentados.

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1. Introdução

O filo Annelida Lamarck, 1809 constitui um dos grupos mais diversos dentro de Metazoa, ocupando o ambiente terrestre, marinho e dulcícola. São conhecidas aproximadamente 21.000 espécies de anelídeos (Weigert et al., 2014).

Anelídeos possuem representantes em todos as guildas alimentares, podendo ser filtradores, depositívoros, suspensívoros, herbívoros, carnívoros, onívoros, parasitas e até mesmo osmotróficos (Jumars et al., 2015). São capazes de ocupar diferentes ambientes, desde ambientes marinhos, que concentram a maior diversidade do grupo, ambientes de água salobra, água doce e o ambiente terrestre (Rouse & Pleijel, 2001).

Os anelídeos são organismos metamerizados, formados por um primeiro segmento denominado prostômio, onde se encontra o cérebro e diferentes órgãos sensoriais sucedido por um peristômio onde se encontra a boca, seguido por um número variável de segmentos e, uma região posterior denominada pigídio (Rouse & Pleijel, 2001).

O filo foi tradicionalmente separado em três classes, Polychaeta Grube, 1850, englobando os poliquetas, Clitellata Michaelsen, 1919, englobando tanto as oligoquetas (minhocas) quanto os hirudíneos (sanguessugas) e Archiannelida Hatschek, 1878. Posteriormente os táxons

Myzostomida von Graff, 1877 (= Myzostomatidae Benham, 1896), Echiura Newby, 1940,

Sipuncula Sedgwick, 1898 e Siboglinidae (= Pogonophora Johansson, 1937 e Vestimentifera

Webb, 1969) foram adicionados ao filo, com alto grau de suporte morfológico e molecular ou apenas molecular (Halanych et al. 2002; Struck, 2017). Paralelamente, as classes Archiannelida e Clitellata foram consideradas como grupos pertencentes à classe Polychaeta (Struck, 2017).

O táxon Polychaeta corresponde à cerca de 13.000 espécies (Jumars, 2015), contanto com grande variação morfológica, guildas alimentares e formas de reprodução. Cerca de 98% das espécies se encontram no ambiente marinho (Glasby & Timm, 2008), ocupando desde o supra litoral até regiões de mar profundo. Apesar da maioria dos grupos apresentar parapódios e cerdas, utilizados na locomoção, muitos grupos intersticiais perderam essas estruturas ao longo da

evolução (Struck et al., 2015) e a única sinapomorfia do grupo são os órgãos nucais, estruturas quimiorreceptoras. O termo “Polychaeta” vem lentamente caindo em desuso, uma vez que a inclusão dos demais táxons do filo – Archiannelida, Clitellata, Echiura, Sipuncula, Myzostomida e Siboglinidae dentro de Polychaeta torna o grupo sinônimo de Annelida. Adicionalmente, se considerarmos o termo “Polychaeta” da forma como é utilizada tradicionalmente – como apenas uma classe dentro de Annelida, separada de Clitellata, ‘Polychaeta” se torna um táxon parafilético

(Struck, 2017).

“Polychaeta” foi dividido em 2 grandes grupos – Annelidae erraticae (= Errantia) e

Annelidae sedentariae (=Sedentaria) por Quatrefages (1866), classificação que foi seguida por muitos autores ao longo do século XIX e boa parte do século XX (Fauvel, 1923; Day 1967;

Hartman 1968. Essa classificação foi duramente criticada por estar mais relacionada ao modo de vida dos organismos do que ao histórico evolutivo dos mesmos. Autores subsequentes agruparam poliquetas em ordens, embora as classificações variassem entre os autores (Dales, 1963; Fauchald,

1977; Pettibone, 1982; Hartmann-Schröder, 1996; Westheide, 1996). Rouse & Fauchald (1997) dividiram os poliquetas entre as ordens Scolecida Rouse & Fauchald, 1997 e Palpata Rouse &

Fauchald,1997, enquanto uma análise molecular de Struck et al. (2011) recuperou agrupamentos muito similares à Errantia e Sedentaria, de forma que os autores reestabeleceram esses termos.

A ordem Spionida foi criada por Fauchald (1977) e incluía as subordens Cirratuliformia

Fauchald, 1977, Chaetopteriformia Fauchald, 1977 e Spioniformia Fauchald, 1977, sendo essa

última subordem constituída pelas famílias Apistobranchidae Mesnil & Caullery, 1898, Spionidae

Grube, 1850, Magelonidae Cunningham & Ramage, 1888, Trochochaetidae, Pettibone, 1963,

Poecilochaetidae Hannerz, 1956 e Longosomatidae Hartman, 1944, sendo definida por “Palpos no prostômio ou na junção do prostômio com o peristômio; faringe eversível como uma almofada ventral ou um saco axial” (Fauchald, 1977).

Em análises moleculares recentes, Spionida é recuperado num clado juntamente com

Sabellida (Struck et al., 2008; Struck et al., 2011; Weigert et al., 2014). É importante ressaltar que a definição de Spionida desses trabalhos inclui apenas as famílias Spionidae, Uncispionidae

Green, 1982, Trochochaetidae e Poecilochaetidae, sendo que Struck et al. (2008) recuperou um clado formado por Spionidae, Trochochaetidae e Poecilochaetidae com um alto grau de suporte.

Uncispionidae ainda não dispõe de sequências moleculares e o suporte para o grupo dentro de

Spionida é somente morfológico (Struck, 2017). As famílias Apistobranchidae e Magelonidae foram recuperadas por Struck et al. (2011) em posições mais basais na árvore evolutiva de

Annelida, enquanto Blake & Maciolek (2020), em uma análise molecular preliminar, posicionaram Longosomatidae próximo à Cirratuliformia.

Spionidae representa uma das famílias mais abundantes e frequentes em ambientes marinhos, podendo ser dominantes em comunidades de substrato inconsolidado, particularmente em ambientes eutrofizados. Mundialmente, são conhecidas cerca de 580 espécies distribuídas em 39 gêneros (Blake et al., 2017), embora esse número possa representar uma subestimativa, uma vez que a família possui um histórico bem documentado de complexos de espécies (Sato-Okoshi et al., 2016), espécies com distribuição aparentemente cosmopolita e gêneros que nunca foram revisados.

1.1 Histórico taxonômico e Classificação de Spionidae

A família Spionidae foi formalmente descrita por Grube (1850), como o táxon “Spiodea”, embora nenhuma espécie tenha sido descrita nesse trabalho. Na realidade, a publicação de várias espécies foi feita antes da descrição da família, como Spio seticornis (Linnaeus, 1767), Spio filicornis (Müller, 1776), Polydora cornuta Bosc, 1802, Scolelepis squamata (O.F. Muller,

1806), Malacoceros vulgaris (Johnston, 1827), Polydora calcarea (Templeton, 1836),

Malacoceros laevicornis (Rathke, 1837), Polydora ciliata (Johnston, 1838) e Malacoceros girardi Quatrefages, 1843, todas descritas para a Europa.

Ainda no século XIX, diversos gêneros novos foram descritos para a Europa, como os gêneros Spiophanes (Grube, 1860), Pygospio (Claparède, 1863), Aonides Claparède, 1864,

Laonice Malmgren, 1867, Prionospio Malmgren, 1867, Pseudopolydora Czerniavsky, 1881,

Marenzelleria Mesnil, 1896 e Microspio Mesnil, 1896. Os únicos gêneros descritos no século

XIX para outras regiões geográficas foram os gêneros Streblospio Webster, 1879, descrito para

Nova Jersey, costa Atlântica dos Estados Unidos da América, Dipolydora Verrill, 1881, descrito para Nova Inglaterra, costa Atlântica dos Estados Unidos da América e Boccardia Carazzi, 1893, descrito para costa Pacífica da Austrália.

Na primeira metade do século XX, apenas quatro novos gêneros foram descritos:

Scolecolepides Ehlers, 1907 para a Nova Zelândia, Polydorella Augener, 1914 para a costa Índica da Austrália, Paraprionospio Caullery, 1914, para o Chile e Rhynchospio Hartman, 1936, para a costa Pacífica dos Estados Unidos da América. A partir da segunda metade do século XX, um maior esforço taxonômico e campanhas de coleta focadas em ambientes de mar profundo levaram a um aumento significativo na quantidade e na frequência na qual novos gêneros eram descritos, uma vez que 15 novos gêneros foram descritos nesse período. O primeiro gênero descrito foi

Dispio Hartman, 1951, para a costa Atlântica dos Estados Unidos da América, seguido de

Tripolydora Woodwick, 1964 para as Ilhas Marshall (Oceano Pacífico) e Apoprionospio Foster,

1969 para a costa Atlântica dos Estados Unidos da América.

Na década seguinte, Blake & Kudenov (1978), num trabalho para a costa Pacífica da

Austrália, descreveram quatro gêneros: Australospio Blake & Kudenov, 1978, Boccardiella

Blake & Kudenov, 1978, Carazziella Blake & Kudenov, 1978 e Orthoprionospio Blake &

Kudenov, 1978. Na década de 1980, Nancy J. Maciolek descreveu três gêneros de mar profundo, oriundos de sua tese de doutorado (Maciolek, 1983): Aurospio Maciolek, 1981a, Laubieriellus

Maciolek, 1981b e Xandaros Maciolek, 1981b. Ainda na década de 1980, são descritos os gêneros

Amphipolydora Blake, 1983 para a Argentina e Pygospiopsis Blake, 1983 para a Ilha Geórgia,

Território Britânico Ultramarino, além de Atherospio Mackie & Duff, 1986 para a Escócia e

Aonidella López-Jamar, 1989 para a Península Ibérica.

Na década de 1990, foi descrito apenas o gênero Lindaspio Blake & Maciolek, 1992, baseado em material coletado em fontes hidrotermais próximo à costa Pacífica dos Estados Unidos da

América. No século XXI, apenas três novos gêneros foram descritos: Glyphochaeta Bick, 2005

4 para a Noruega, Spiogalea Aguirrezabalaga & Ceberio, 2005 para a costa Atlântica da França e

Glandulospio Meißner et al., 2014 para o Atlântico Nordeste, a cerca de 1200km do continente

Africano.

Em relação à classificação da família, Söderström (1920) foi o primeiro a englobar a maioria dos gêneros da época e também o primeiro a utilizar caracteres internos e caracteres reprodutivos.

Söderström (1920) estabeleceu três subfamílias: Spioninae Söderström, 1920, englobando os gêneros Spio, Microspio, Pygospio e Polydora; Nerininae Söderström, 1920, para os gêneros

Malacoceros, Scolelepis e Aonides e Laonicinae Söderström, 1920, formado apenas por Laonice.

As duas primeiras subfamílias ainda são utilizadas atualmente, enquanto Laonice foi transferido para Nerininae por Blake & Arnofsky (1999).

A primeira análise filogenética do grupo foi realizada por Sigvaldadóttir et al. (1997), que obteve resultados divergentes de Söderström (1920), embora os clados encontrados apresentassem um baixo suporte de ramo (Blake et al., 2017), devido a problemas e limitações admitidos pela própria autora, como reduzido número de gêneros analisados e escolha equivocada dos grupos-externos – devido à forte homologia de caracteres entre diversos gêneros analisados e os grupos-externos (Sigvaldadóttir et al., 1997; Blake & Arnofsky, 1999).

Posteriormente, uma análise filogenética preliminar foi feita por Blake & Arnofsky (1999) utilizando um maior número de caracteres e gêneros, incluindo outros táxons de Spionidae sensu

Fauchald, 1977. Essa análise recuperou um clado formado pela subfamília Spioninae, outro clado formado por todos os outros gêneros de Spionidae, além de Heterospio Ehlers, 1874

(Longosomatidae), Poecilochaetus Claparède in Ehlers, 1875 (Poecilochaetidae), Trochochaeta

Levinsen, 1884 (Trochochaetidae) e Uncispio Green, 1982 (Uncispionidae) e um clado menor composto apenas por Pygospiopsis e Atherospio.

Segundo Blake & Arnofsky (1999), há evidência morfológica para que as famílias Spionidae,

Uncispionidae, Poecilochaetidae e Trochochaetidae sejam reorganizadas em três clados com nível de subfamília dentro de uma família com uma definição morfológica mais ampla. O primeiro

5 clado seria formado por Pygospiopsis e Atherospio, o segundo clado seria formado pela subfamília Spioninae, incluindo os gêneros Spio, Microspio, Pygospio e o complexo-Polydora

(Polydora, Dipolydora, Tripolydora, Amphipolydora, Pseudopolydora, Polydorella, Boccardia,

Boccardiella e Carazziella) e um terceiro e maior clado, referido como subfamília Nerininae, incluindo Poecilochaetus, Trochochaeta, Uncispio, Laonice, Dispio, Scolelepis, Aonides,

Aonidella, Malacoceros, Rhynchospio, Spiophanes, Australospio, Lindaspio, Scolecolepides,

Marenzelleria e o complexo-Prionospio (Prionospio, Apoprionospio, Paraprionospio,

Orthoprionospio, Aurospio, Streblospio e Laubieriellus) (Blake & Arnofsky, 1999).

De acordo com Blake & Arnofsky (1999) e Blake (2006), mais estudos são necessários para um posicionamento de Poecilochaetus, Trochochaeta, Uncispio dentro de Spionidae, enquanto uma análise molecular preliminar de Blake e Maciolek (2020) posicionaram Heterospio próximo à Cirratuliformia. Até o momento, apenas o trabalho de Radashevsky et al. (2018) considerou um desses táxons, Trochochaeta, como parte de Spionidae.

Apesar da existência e aceitação das subfamílias, essas são amplamente negligenciadas por autores subsequentes, sendo referenciadas posteriormente apenas por Blake et al. (2017). Ainda de acordo com esse trabalho, os gêneros Spiogalea, Glandulospio, Glyphochaeta e Xandaros possuem poucas afinidades com os demais Spionidae, não tendo sido posicionados em nenhuma subfamília.

1.2 Diversidade de Spionidae na costa Brasileira

Para a costa brasileira são reconhecidas 90 espécies (Amaral et al., 2013), distribuídas em 17 gêneros considerados válidos pela plataforma on-line WoRMS (World Register of Marine

Species) (Read & Fauchald, 2019): Aonides, Aurospio, Boccardia, Boccardiella, Dipolydora,

Dispio, Laonice, Malacoceros, Microspio, Paraprionospio, Polydora, Prionospio,

Pseudopolydora, Scolelepis, Spio, Spiophanes e Streblospio. Os registros de poliqueta para a

6 costa brasileira foram compilados por Amaral et al. (2013), embora muitos registros de Spionidae sejam duvidosos, provenientes de estudos de dinâmica populacional (MacCord et al., 2011) ou de estudos de cunho ecológico, que regularmente reportam espécies ditas cosmopolitas ou conhecidos complexos de espécies.

A primeira espécie de Spionidae escrita para a costa brasileira Mandane brevicornis Kinberg,

1866, foi descrita por Kinberg (1866), a partir de material coletado no Rio de Janeiro. Essa espécie permaneceu ignorada por mais de um século até ser revalidada como Laonice brevicornis por

Sikorski (2011).

Quase um século depois, foi descrita a próxima espécie para o Brasil, Laonice antarcticae

Hartman, 1953 para a plataforma continental (80 m de profundidade) do estado do Rio Grande do Sul. A espécie foi descrita originalmente como uma subespécie de Laonice cirrata, Laonice cirrata antarcticae, enquanto a localidade tipo foi referido para o Uruguai, por um equívoco da autora. Na década de 1980, foi descrita a próxima espécie para o Brasil, Laonice branchiata

Nonato et al., 1986, para águas rasas no litoral do Estado do Paraná (Nonato et al., 1986). Ainda para o Estado do Paraná, Polydora carinhosa Radashevsky et al. (2006) foi descrita a partir de espécimes perfurando conchas de ostras.

Em 2009, Radashevsky & Migotto (2009) descreveram Pseudopolydora rosebelae

Radashevsky & Migotto, 2009 a partir de material coletado nos Estados do Rio de Janeiro e São

Paulo, enquanto Radashesvky & Lana (2009), em uma revisão de Laonice da América do Sul e

América Central, descreveram Laonice parvabranchiata Radashevsky & Lana, 2009 a partir de material dos Estados de São Paulo e Santa Catarina, além de Laonice petersenae Radashevsky &

Lana, 2009 e L. aperata Radashevsky & Lana, 2009, as quais foram posteriormente sinonimizadas com L. brevicornis por Sikorski (2011).

Trabalhos com uma maior abrangência taxonômica são pouco comuns, como Bolívar &

Lana, 1987 para o Estado do Paraná, Pardo et al. 2006 e Pardo & Peixoto in press para as regiões

Sul e Sudeste do Brasil e Paiva & Barroso, 2010 para o talude continental da Bacia de Campos,

Estado do Rio de Janeiro. Além disso, esses trabalhos não fizeram a descrição de novas espécies, apenas novos registros (ver abaixo).

Todas as demais espécies registradas no Brasil e compiladas por Amaral et al. (2013) foram descritas em outras localidades geográficas, particularmente a Europa, costa Atlântica da América do Norte e América Central, países circunvizinhos na América do Sul e Antárctica. Devido às diferenças em relação à latitude, profundidade e distância geográfica, é possível que alguns desses registros estejam equivocados. Contudo, decidiu-se não se fazer comentários a respeito da validade desses registros para a costa Brasileira até que material da localidade-tipo e material registrado no Brasil possa ser examinado e comparado. Trabalhos taxonômicos para a costa brasileira que não realizaram a descrição de espécies novas incluem Radashevsky & Nogueira

(2003), Rocha et al. (2009), Rocha & Paiva (2012) e Radashevsky & Migotto (2017).

É importante ressaltar que praticamente todas as espécies reportadas na costa brasileira foram coletadas em ambientes rasos (profundidade inferior a 50 metros), com a exceção de Prionospio fauchaldi Maciolek, 1985 e Aurospio dibranchiata Maciolek, 1981a, que são reportadas apenas no talude continental, além de Spiophanes kroyeri Grube,1860, Prionospio steenstrupi Malmgren,

1867 e Dipolydora blakei Maciolek, 1984, que ocorrem tanto na plataforma quanto no talude continental. Ambientes de mar profundo permanecem pouco conhecidos, com apenas um estudo realizado na Bacia de Campos (Paiva & Barroso, 2010), o qual reportava espécies ainda não descritas.

Quase todos os registros de espécies foram feitos para as regiões Sul e Sudeste do Brasil, com poucos trabalhos realizando o registro de espécies para a região Nordeste (Santos et al., 1994;

Netto et al., 1999; Viana et al., 2005; Costa & Nalesso, 2006; Paiva et al., 2007; Radashevsky &

Lana, 2009; Santos et al., 2009; Silva & Barros, 2011; Assis et al., 2012; Berlandi et al., 2012;

Rocha & Paiva, 2012) e nenhum trabalho reportando Spionidae para a região Norte do Brasil.

Para a Bacia do Espírito Santo são reportadas apenas cinco espécies de Spionidae: Dipolydora socialis (Schmarda, 1861) (Costa & Nalesso, 2006), Polydora cornuta Bosc, 1802 (Costa &

Nalesso, 2006; Silva & Barros, 2011), Prionospio heterobranchia Moore, 1907 (Costa & Nalesso,

2006), Apoprionospio pygmaeus Hartman, 1961 (Costa & Nalesso, 2006) e Spiophanes duplex

(Chamberlin, 1919) (Costa & Nalesso, 2006).

1.3 Morfologia geral

Os espionídeos possuem um corpo longo, cilíndrico ou achatado dorsoventralmente. O corpo é dividido numa região pré-septal formada por prostômio e peristômio, uma longa região septal e uma região pós-septal, formada pelo pigídio e a zona de crescimento.

O prostômio possui grande variação morfológica, podendo ser arredondado, retangular, cônico, subtriangular ou bilobado, sendo arredondado ou truncado na margem anterior (Figura

1A). O prostômio pode ainda ser dotado de estruturas sensoriais, como olhos, antena e picos prostomiais (Figura 1B). A parte posterior do prostômio é flanqueada por órgãos nucais ciliados e pode ainda se alongar posteriormente como uma carúncula por um número variável de setígeros

(Figura 1C).

O peristômio é um anel que circunda a boca ventralmente e o prostômio dorsalmente

(Figura 1A). O peristômio geralmente é um segmento átoco (não possui cerdas), exceto quando se encontra fundido com o primeiro setígero, como ocorre em espécies de Prionospio (Blake et al. 2017) (Figura 1A). Os palpos se encontram na margem posterior do peristômio, sendo usados na alimentação, seja através da filtração de partículas da coluna d’água (suspensívoro) ou captura de partículas de alimento no substrato (depositívoro). Os palpos são geralmente longos e sulcados

(sulco ausente em adultos do gênero Scolelepis), sendo estruturas frágeis e perdidas com extrema facilidade (Figura 1D).

Figura 1. Morfologia geral 1. A. Prionospio sp., vista dorsal. B. Prionospio delta, vista dorsal. C.

Laonice cf. antarcticae, vista dorsal. D. Boccardia sp., vista lateral. Abreviações: an – antena, br – brânquia, car – carúncula, esp – espinhos, on – órgão nucal, pa – palpo, pe – peristômio, pp – pico prostomial, pro – prostômio.

A região septal é dotada de parapódios birremes, ou seja, dotados de lobos parapodiais dorsais (notopódio) e ventrais (neuropódio) (Figura 2A). Os lobos parapodiais costumam ser pouco desenvolvidos, mas dotados de lamelas notopodiais e neuropodiais. Enquanto lamelas pós- septais (i.e., imediatamente após o feixe de cerdas) estão presentes em todos os espionídeos, lamelas pré-septais (i.e., imediatamente antes do feixe de cerdas) são observadas em poucos organismos (Figura 2A). Lamelas são mais desenvolvidas na região anterior e diminuem de tamanho e frequentemente mudam de formato em direção à região posterior. Existem algumas

10 especializações das lamelas pós-septais, como as cristas, um prolongamento fino e membranoso que conecta as lamelas pós-septais de um mesmo setígero (Figura 2B). Cristas dorsais são bem conhecidas em espécies dos gêneros Prionospio, Spiophanes e Laonice, enquanto as cristas ventrais são mais raras e presentes apenas no gênero Laubieriellus e alguma espécies de

Prionospio. Algumas espécies possuem membranas finas interligando os neuropódios de setígeros adjacentes, chamadas de bolsas interneuropodiais, sendo frequentes no gênero Laonice e ocorrendo raramente em espécies de Prionospio, Spiophanes e Aonidella (Figura 2C).

A maioria das espécies possui brânquias, podendo estas estar presentes apenas em alguns setígeros anteriores e ter um limite espécie-específico ou podem estar presentes em (quase) todo o corpo, com mais pares branquiais sendo adicionados conforme o indivíduo cresce (Figuras 1

A–C, 2B). As brânquias geralmente são cilíndricas e cirriformes, embora brânquias achatadas e triangulares sejam conhecidas no gênero Prionospio. Em Prionospio e gêneros relacionados

(Apoprionospio, Paraprionospio), alguns pares de brânquias possuem extensões digitiformes ao longo do seu comprimento, denominadas pínulas, com a função de aumentar a superfície de trocas gasosas. Em espécies de Scolelepis e Dispio, as brânquias podem estar fundidas às lamelas notopodiais pós-septais pela maior parte do seu comprimento. Em muitas espécies de Dispio, brânquias acessórias digitiformes, localizadas logo após a lamela notopodial pós-septal, podem estar presentes (Figura 2A).

Os parapódios são dotados de cerdas, geralmente em quatro categorias: cerdas capilares, ganchos, cerdas em sabre ou espinhos. Cerdas capilares são cerdas simples (não articuladas), podendo apresentar um alargamento em um dos lados (unilimbada), em ambos os lados

(bilimbadas) ou sem alargamento (não limbada). Estas são as cerdas mais comuns em Spionidae e estão presentes em todas as espécies (Figura 2C). Ganchos estão presentes no neuropódio de todas as espécies de Spionidae e frequentemente presentes no notopódio, geralmente se iniciando em um setígero específico ou apresentando uma pequena variação no setígero de início dos ganhos.

Ganchos podem possuir um único dente (unidentados), ou possuir um ou mais dentes secundários imediatamente acima do dente principal (bi- a multidentados) (Figura 2D). A maioria dos ganchos possui um capuz hialino transparente protegendo o ápice do gancho (onde estão os dentes) e algumas espécies possuem um capuz secundário logo abaixo do dente principal. No gênero Spiophanes, o capuz se encontra reduzido ou mesmo completamente ausente.

Figura 2. Morfologia geral 2. A. Dispio sp., vista lateral. B. Prionospio sp., vista dorsal. C.

Laonice cf. antarcticae, vista lateral. D. Prionospio solisi, vista lateral. Abreviações: ba – brânquias acessórias, bi – bolsa interneuropodial, br – brânquia, ca – capilar, cd – crista dorsal, ga – gancho, lapos – lamela pós-septal, lapre – lamela pré-septal, pe – peristômio, pro – prostômio, sa – sabre.

Cerdas em sabre são cerdas recurvadas e robustas, sendo sempre a cerda em posição mais ventral no feixe de cerdas do neuropódio (Figura 2D). Assim como os ganchos, as cerdas em sabre geralmente se iniciam em um setígero específico ou apresentam uma pequena variação no setígero de início. Usualmente existe apenas uma cerda em sabre por feixe, embora um número maior pode estar presente em espécies de Malacoceros, Dispio e Laonice.

Por último, os espinhos são uma classificação ampla de cerdas com morfologia e funções diferentes, como as cerdas em anzol (crook-like chaetae) presentes no neuropódio do setígero 1 do gênero Spiophanes (Figura 3A), utilizadas para ancorar o indivíduo no tubo; os espinhos do setígero 5 presentes em espécies do complexo-Polydora (Figura 3B), utilizados para desgastar estruturas calcáreas (uma vez que muitas espécies perfuram conchas de moluscos bivalves) e os espinhos notopodiais presentes na região posterior de algumas espécies do gênero Microspio e do complexo-Polydora (Figura 3C), que ficam embebidos no corpo e são protraíveis, tendo uma função ainda incerta na defesa do indivíduo.

No final do corpo, se encontra o pigídio, geralmente dotado de cirros, lobos ou uma combinação de ambos, que circundam o ânus em posição terminal. A quantidade e arranjo dos cirros pode ser fixa, como em Prionospio, que geralmente possui dois cirros curtos e um cirro mediano longo (Figura 3D) ou pode ser dependente do tamanho, com mais cirros se desenvolvendo conforme o organismo cresce.

Figura 3. Morfologia geral 3. A. Spiophanes duplex, vista dorsolateral. B. Dipolydora cf. caulleryi, vista lateral. C. Dipolydora barbilla, vista lateral. D. Prionospio solisi, vista lateral.

Abreviações: can – cerda em anzol, es noto – espinho notopodial, es set 5 – espinho do setígero 5, pig – pigídio.

2. Objetivos

2.1 Objetivo geral

. Realizar a taxonomia da família Spionidae ao menor nível taxonômico possível

2.2 Objetivos específicos

. Identificar os gêneros e espécies de Spionidae da Bacia do Espírito Santo e Porção Norte da

Bacia de Campos.

. Redescrever as espécies da família identificadas no material analisado.

. Atualizar dados de distribuição batimétrica e geográfica para as espécies e registrar novas ocorrências.

. Realizar a descrição dos morfotipos, descrevendo novos táxons, quando for o caso.

. Elaborar chaves de identificação ao nível de família para todos os Spionidae da Bacia do Espírito

Santo e Porção Norte da Bacia de Campos.

3. Materiais e Métodos

3.1 Área de Estudo, coleta e processamento das amostras

A Bacia do Espírito Santo (20.19ºS a 19ºS e 37.88ºW a 40.22ºW) abrange uma área aproximada de 123.000 km2, se estendendo desde o sul do estado do Espírito Santo até o norte deste estado, na divisa com a Bahia (Vieira et al., 1994). O limite sul dessa bacia se dá no Alto de Vitória, onde se inicia a Bacia de Campos, enquanto o limite norte, na divisa com a Bahia e onde se inicia a Bacia de Mucuri, corresponde a um limite apenas geográfico, uma vez que, segundo Rangel et al. (1994) e Vieira et al. (1994), as bacias do Espírito Santo e Mucuri compartilham comportamento estrutural e geológico similares. A bacia do Espírito Santo possui uma extensão da plataforma continental de aproximadamente 40 km no limite sul e de aproximadamente 240 km no limite norte, devido à formação Abrolhos, localizada na Bacia de

Mucuri (Vieira et al., 1994; França et al. 2007) (Figura 4). A bacia possui substratos consolidados, como rodolitos, bancos de algas calcáreas, estruturas recifais e afloramentos rochosos e uma grande variedade de substratos inconsolidado – de cascalho até lama (Figuras 5–7).

O material utilizado nesse estudo faz parte do Projeto AMBES – Caracterização Ambiental

Marinha da Bacia do Espírito Santo e Porção Norte da Bacia de Campos, coordenado pelo Centro de Pesquisas e Desenvolvimento Leopoldo Américo Miguez de Mello (CENPES) /PETROBRAS.

Houveram 2 campanhas de coleta, sendo a primeira no verão e a segunda no inverno, conforme explicado a seguir.

Figura 4 – Área de coleta do projeto AMBES.

As coletas foram realizadas da seguinte forma: na área de influência da Foz do Rio Doce, as coletas de sedimento ocorreram nos períodos de 11 a 19/12/2010 (verão) e 12 a 18/07/2011

(inverno) a bordo dos navios oceanográficos Luke Thomas e Seward Johnson, respectivamente.

Foi empregada uma malha amostral regular paralela à linha da costa, composta por 20 estações

(Figura 5). Cada estação foi amostrada em triplicata através da utilização de vanVeen (92x80x40 cm) ou Box-Corer (50x50x50 cm) com janela superior para a retirada por réplica de 4 litros correspondente à camada sedimentar de 0-10 cm.

Figura 5 – Faciologia e pontos amostrais da Foz do Rio Doce. Fonte: CENPES/ Petrobras.

Para as coletas na plataforma continental, as coletas de sedimento ocorreram a bordo do navio oceanográfico Seward Johnson nos períodos de 02/12/2011 a 02/02/2012 (verão) e 06/06 a

17/07/2013 (inverno). A malha amostral empregada na plataforma continental foi composta por

28 estações distribuídas em 7 transectos, denominados de A até G de sul para norte da Bacia do

Espírito Santo e porção norte da Bacia de Campos (Figura 6). Foram amostradas 4 estações por transecto, denominadas 01 a 04 conforme o aumento da profundidade (isóbatas de 25, 40, 50, 150 m). Cada estação foi amostrada em triplicata através do uso de vanVeen (92x80x40 cm) ou Box-

Corer (50x50x50 cm) com janela superior, onde foram coletados por réplica 4 litros correspondente a camada sedimentar de 0-10 cm.

As coletas de sedimento do talude ocorreram a bordo do navio oceanográfico Seward Johnson nos períodos de 02/12/2011 a 02/02/2012 (verão) e 06/06 a 17/07/2013 (inverno). A malha amostral empregada no talude continental foi composta por 42 estações distribuídas em 7 transectos, denominados de A até G de sul para norte da Bacia do Espírito santo e porção norte da Bacia de Campos (Figura 6). Foram amostradas 6 estações por transecto, denominadas 05 a 10

18 conforme o aumento da profundidade (isóbatas de 400, 1000, 1300, 1900, 2300 e 3000 m). Cada estação foi amostrada em triplicata através do uso de vanVeen (92x80x40 cm) ou Box-Corer

(50x50x50 cm) com janela superior, onde foram coletados por réplica 9 litros correspondente a camada sedimentar de 0-10 cm.

Figura 6 – Faciologia e pontos amostrais da plataforma continental e talude. Fonte: CENPES/ Petrobras.

Por último, as coletas de sedimento do Cânion Watu Norte (CANWN) e do Cânion do Rio Doce

(CAND) ocorreram conjuntamente às amostragens de sedimento da plataforma continental e do talude continental e a bordo do navio oceanográfico Seward Johnson nos períodos de 02/12/2011 a 02/02/2012 (verão) e 06/06 a 17/07/2013 (inverno). Em cada cânion foram amostradas 4 estações, denominadas de 04 a 07 conforme o aumento da profundidade (isóbatas de 150, 400,

1000, 1300 m) (Figura 7). Cada estação foi amostrada em triplicata através do uso de vanVeen

(92x80x40cm) ou Box-Corer (50x50x50cm) com janela superior, onde foram coletados por réplica 9 litros correspondente a camada sedimentar de 0-10 cm.

Figura 7 – Faciologia e pontos amostrais dos cânions. Fonte: CENPES/ Petrobras.

Todo o sedimento coletado nas etapas anteriores foi fixado in situ em formalina a 10%.

Posteriormente, as amostras foram transferidas para empresa de consultoria ambiental, onde foram lavadas em água doce e transferidas para etanol a 70%. Em seguida, as amostras foram elutriadas e o material foi triado e inicialmente separado em grandes grupos zoológicos (e.g. classe Polychaeta). O material de Polychaeta foi identificado a nível de família e enviado para especialistas das respectivas famílias, a fim de ser identificado ao menor nível taxonômico possível.

3.2 Análise do Material

O material foi analisado sob estereomicroscópio e microscópio óptico. Primeiramente, foi realizada a taxonomia em nível de gênero e posteriormente foi realizada a identificação em nível de espécie a partir da bibliografia disponível para a costa brasileira (e.g. Bolívar & Lana,

1987; Pardo et al., 2006, Pardo & Peixoto in press). Considerando que diversas espécies reportadas por Amaral et al. (2013) para a costa brasileira foram originalmente descritas para

20 outras regiões geográficas, foram consultados trabalhos para a América do Norte e Central (e.g.

Blake, 1971; Maciolek, 1984; 1987; 1990; 1985; 2000) e Antártica (Blake, 1983). Quando os organismos encontrados diferiam das espécies reportadas na bibliografia, eram criados morfotipos a partir de caracteres taxonômicos. Foram examinados 16.938 indivíduos. Táxons cuja morfotipagem ocorreu durante o projeto HABITATS, realizado na Bacia de Campos e que também foram reportados no atual projeto se encontram sumarizados no Anexo 1.

Os seguintes caracteres de valor taxonômico foram analisados: formato do prostômio; presença, posição, tipo e distribuição de brânquias; presença e distribuição de ganchos notopodiais e neuropodiais; morfologia dos ganchos; presença e distribuição das cerdas em sabre; morfologia das lamelas pré e pós septais; presença e distribuição de bolsas laterais, cristas dorsais e carúncula, morfologia do pigídio, morfologia das cerdas e morfologia dos espinhos modificados no 5º setígero. O corante temporário verde de metila foi utilizado para melhor visualização dos caracteres citados e observação de eventuais padrões de coloração espécie-específicos. Uma vez que os organismos foram fixados e permaneceram em formalina 10% por um período estendido de tempo, optou-se por não realizar estudos moleculares.

Os organismos foram fotografados utilizando um estereomicroscópio modelo Leica M205C equipado com sistema de captura de imagens (Laboratório de Poliquetas – UFRRJ e Laboratório de Ecologia de Insetos – UFRJ). As imagens foram tratadas e as pranchas montadas no Adobe

Photoshop CC 2019. Adicionalmente, foram feitas pranchas de desenhos para cada espécie utilizando o Adobe Illustrator CC 2019.

Sempre que possível, foram realizados estudos de microscopia eletrônica. Os espécimes foram inicialmente desidratados em série alcoólica crescente (etanol 70% até etanol 100%, 15 minutos em cada etapa, além de 2 trocas de etanol 100%), seguido de secagem em ponto crítico de CO2. Alternativamente, o processo de secagem em ponto crítico de CO2 foi substituído pelo uso de série crescente de hexametildizilazano (HMDS) (Braet et al., 1997) (2 partes de etanol

100% para uma parte de HMDS, 1:1, 1:2 e HMDS, 15 minutos em cada etapa, seguido de 2 trocas de HMDS), sendo a secagem realizada em capela de gases. Após a secagem, os organismos foram

21 montados em stubs e metalizados com ouro. Para a observação da morfologia dos ganchos, foi utilizado um limpador ultrassônico, anteriormente à série alcoólica.

Organismos foram em seguida examinados em um microscópio eletrônico de varredura modelo JEOL JSM-6390 LV (Museu Nacional e FIOCRUZ), ou JEOL JSM-6510 LV

(Departamento de Zoologia – UFRJ). As fotomicrografias foram tratadas no Adobe Photoshop

CC 2019 e as pranchas montadas no Adobe Illustrator CC 2019.

O material tipo de espécies está sendo tombado no Museu Nacional do Rio de Janeiro

(MNRJP), Universidade Federal do Rio de Janeiro; Museu de Zoologia Adão José Cardoso

(ZUEC-POL), Universidade Estadual de Campinas e Museu de Zoologia (MZUSP), Universidade de São Paulo. Material adicional (i.e., não tipo) será tombado no Museu de Zoologia Adão José

Cardoso (ZUEC-POL), Universidade Estadual de Campinas.

4. Resultados

A tese foi dividida em oito capítulos, editados para a revista alvo. Enquanto alguns gêneros com maior afinidade foram agrupados (e.g. complexo Polydora e complexo Prionospio), outros gêneros (e.g. Spiophanes) foram abordados separadamente, como é costumeiro em trabalhos recentes para esses táxons.

O capítulo I se encontra publicado no periódico Journal of the Marine Biological Association of the United Kingdom como parte de uma edição especial relativa à 12th Internacional Polychaete Conference, enquanto o capítulo II se encontra publicado no periódico Zootaxa. O capítulo III foi submetido para o periódico Zootaxa e se encontra em fase de revisão, enquanto os demais capítulos ainda serão submetidos.

A seguir o título e a organização dos capítulos:

4.1 Capítulo I – A new species of Spiogalea (Polychaeta: Spionidae) from Brazil, with an amended diagnosis of the genus.

4.2 Capítulo II – New Prionospio and Laubieriellus (Annelida: Spionidae) species from Southeastern Brazil.

4.3 Capítulo III – New apinnate Prionospio (Annelida: Spionidae) species from southeastern Brazil.

4.4 Capítulo IV – Species from the Prionospio (Annelida: Spionidae) complex from southern Brazil, including new species from Apoprionospio and Paraprionospio.

4.5 Capítulo V – Contributions to the taxonomy of Aonides, Aonidella and Laonice from southeastern Brazil, including the description of five new species.

4.6 Capítulo VI – Spionidae (Annelida) from Espírito Santo Basin, southeastern Brazil: the genera Scolelepis, Dispio, Spio, Microspio and Malacoceros, including a new deep-sea Scolelepis species.

4.7 Capítulo VII – A contribution to the taxonomy of Spiophanes (Annelida: Spionidae), including new species.

4.8 Capítulo VIII – Polydorids (Annelida: Spionidae) species from southern Brazil, with new records and a new Boccardia species.

Uma chave para os gêneros encontrados nesse projeto se encontra no Anexo 2.

Capítulo 1

Journal of the Marine Biological Association of the United Kingdom, 2017, 97(5), 955 – 960. # Marine Biological Association of the United Kingdom, 2017 doi:10.1017/S0025315417000509

A new species of Spiogalea (Polychaeta: Spionidae) from Brazil, with an amended diagnosis of the genus

ANTONIO J.M. PEIXOTO1,2 AND PAULO c. PAIVA1,2 1Laborato´rio de Polychaeta, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, CCS, Ilha do Fundaõ, Rio de Janeiro, RJ CEP 21941- 902, Brazil, 2Programa de Po´s-graduaçaõ em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, CCS, Ilha do Fundaõ, Rio de Janeiro, RJ CEP 21941-902, Brazil

A new Spiogalea species is described from South-east Brazil. The specimens were collected in muddy bottoms during a large survey (15 – 3000 m) in southern Brazil. This genus comprised, up to now, only one species described from Capbreton Canyon (Bay of Biscay). The new species differs from the type-species, mostly in the morphology of the prostomial chitinous plate, a character restricted to the genus Spiogalea. With this new species, the generic diagnosis of the genus was amended.

Keywords: Spiogalea, Spionidae, Esp´ırito Santo Basin, Doce Canyon

Zoobank identiﬁer: LSID urn:lsid:zoobank.org:pub:760FF015-7252-4F47-9726-C4991A1A27CD

Submitted 8 November 2016; accepted 17 March 2017

INTRODUCTION

Polychaetes belonging to the Family Spionidae Grube, 1850 are very common and ecologically important in many marine ecosystems, and are one of the most species-rich polychaete families. Currently there are 606 nominal species distributed in 39 genera (Read & Fauchald, 2016).

Around 90 nominal species of the family were already recorded for the Brazilian coast, distributed in 17 genera (Amaral et al., 2013), although some authors suggest that those numbers are underestimates, as many species remain undescribed, mainly in the continental slope and deep- sea (Paiva & Barroso, 2010).

Taxonomic studies on this family in the Brazilian coast are still scarce, focusing mainly on coastal and intertidal environments and only a few genera, such as Laonice Malmgren, 1867 (Nonato et

24 al., 1986; Radashevsky & Lana, 2009), Scolelepis Blainville, 1828 (Rocha et al., 2009; Rocha & Paiva, 2012), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto, 2009), Dipolydora Verrill, 1881 (Radashevsky & Nogueira, 2003) and Polydora Bosc, 1802 (Radashevsky et al., 2006), have received any detailed attention.

Large-scale studies are restricted to those of Bolívar & Lana (1987) in the continental shelf of Paraná State and Paiva & Barroso (2010) in the continental slope of Campos Basin, Rio de Janeiro State.

The genus Spiogalea Aguirrezabalaga & Ceberio, 2005 is known only by the type-species, S. vieitezi Aguirrezabalaga & Ceberio, 2005. It is characterized by two chevron-shaped chitinous plates surrounding anterior part of prostomium, lack of branchiae, notopodial chaetae all capillary. Other signiﬁcant generic characters of Spiogalea are parapodia of ﬁrst chaetiger reduced and devoid of notopodial postchaetal lobe while presenting a small neuropodial postchaetal lobe.

Along with the genera Spiophanes Grube, 1860, Spiophanella Fauchald & Hancock, 1981 and Glyphochaeta Bick, 2005, Spiogalea lack branchiae, which is unusual for Spionidae.

During the Project ‘Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin’ (AMBES), coordinated by the research Centre CENPES/PETROBRAS, focusing on the southern coast of Brazil, three specimens belonging to an undescribed species of Spiogalea were found on the continental slope of southern Brazil, at 950 m depth.

These specimens ﬁt reasonably well to the diagnosis of the genus, but noteworthy differences were observed, such as the presence of notopodial hooks on posteriormost chaetigers and a large single chitinous plate covering the dorsal and ventral sides of the prostomium. A description of this new species and an amended diagnosis are provided on this paper.

MATERIALS AND METHODS

Specimens were collected on the continental slope of Espírito Santo Basin, Doce Canyon at 950 m depth using a Box-corer sampler, during the cruises of AMBES (Figure 1). Specimens were ﬁxed in 10% formalin and preserved in 70% ethanol.

Morphological traits of specimens were observed under a Zeiss Stemi SV 11 stereomicroscope and Zeiss Axio Lab A1 microscope. Specimens were stained in a strong solution of methyl green (1 g/l) for 5s and then transferred to70% ethanol to improve the visualization of characters. The staining is temporary and fades completely in ethanol after a few minutes.

Due to the small number of specimens, it was decided not to perform SEM studies. As the specimens were ﬁxed in 10% formalin, molecular studies were also not possible.

Since these specimens display characters that differ from or were not observed on the original description of the genus, its diagnosis was therefore amended.

The holotype (MNRJP 1220) and two paratypes (MNRJP 1221) were deposited in the Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro State, Brazil.

Fig. 1. Map showing sampling location of Spiogalea capixaba sp. nov.

RESULTS

Systematics

Order SPIONIDA sensu Rouse & Fauchald, 1997

Suborder SPIONIFORMIA sensu Fauchald, 1977

Family SPIONIDAE Grube, 1850

Genus Spiogalea Aguirrezabalaga & Ceberio, 2005

Diagnosis (after Aguirrezabalaga & Ceberio, 2005, amended)

Prostomium anteriorly rounded or with short anterolateral projections, narrowing posteriorly, with caruncle. Nuchal organs absent. Two chevron-shaped chitinous plates or single chitinous plate surrounding anterior part of prostomium. Eyes absent. Peristomium well developed forming

26 posteriorly open collar that surrounds prostomium. Branchiae absent. Parapodia of ﬁrst chaetiger reduced, lacking notopodial postchaetal lobe and with small neuropodial postchaetal lobe. Subsequent parapodia larger, well-developed with rounded noto- and neuropodial postchaetal lobes. Posterior notopodial lobes connected by low dorsal ridge or ridges absent. Notopodial chaetae all capillary or capillaries and multidentate hooded hooks on posterior chaetigers. Posterior neuropodia with long-shafted, multidentate hooded hooks, with complete hood. Sabre chaetae present. Pygidium morphology uncertain, possibly rounded with a pair of small anal cirri.

Spiogalea capixaba sp. nov. (Figures 2 – 5)

Type material

Holotype: Incomplete specimen, 0.8 mm wide, palps missing. AMBES Project, Station Amb 3 CAND6R2 (19837′49.14′′S, 3983′59.7′′W), 950 m depth (MNRJP 1220); coll. V. Veloso/ PETROBRAS, 11/12/2011.

Paratypes: two incomplete specimens, 0.3 and 0.7 mm wide, palps missing (MNRJP 1221). Collection details as for holotype.

Comparative material examined

Spiogalea vieitezi Aguirrezabalaga & Ceberio, 2005. Paratype: Incomplete specimen, palps missing. (Aguirrezabalaga’s personal collection; not deposited in MNHN), Capbreton Canyon, Bay of Biscay, Station KF 50 (43º35′35′′N 18º55′15′′W); collected with Flusha Box corer, 1000 m depth, 14/09/1989.

Fig. 2. Spiogalea capixaba sp. nov., holotype: anterior region, dorsal view. Scale bar: 400 mm.

Description

Three specimens examined, all incomplete. Holotype with 44 chaetigers; 4 mm long, 0.8 mm wide at chaetiger 4 (excluding parapodia).

Body elongate, dorsoventrally ﬂattened, without regionalization. Anterior portion wide, tapering from middle body region onwards (Figure 2).

Prostomium bearing short anterolateral projections, posteriorly narrowing to an inconspicuous caruncle reaching posterior margin of ﬁrst chaetiger. Nuchal organs absent. Single large chitinous plate present on the dorsal and ventral sides of anterior part of the prostomium. Eyes absent (Figure 3).

Peristomium large, well developed, separated from chaetiger 1, encircling the prostomium. Peristomium biannulated, overlapping the ﬁrst chaetiger on posterior margin. Palps lost. Branchiae absent throughout the body.

First chaetiger reduced. Notopodial postchaetal lobe absent, bearing few long non-limbate capillaries. Neuropodial postchaetal lobe thick and rounded, reduced, bearing few long non- limbate capillaries.

Fig. 3. Spiogalea capixaba sp. nov., holotype: close-up of the prostomium, dorsal view: Scale bar: 200 mm.

Fig. 4. Spiogalea capixaba sp. nov., holotype: (A) Left notopodium of chaetiger 8, dorsal view; (B) Left neuropodium of chaetiger 9, ventral view; (C) Left neuropodium of chaetiger 16, ventral view. Scale bars: 70 mm.

Parapodial lamellae well developed starting on chaetiger 2. Notopodial postchaetal lamellae rounded from chaetigers 2 – 5, being progressively smaller between chaetiger 6 – 9 and reduced from chaetiger 10 onwards. Dorsal ridges absent. Neuropodial postchaetal lamellae from chaetigers 2 – 7 short, slender and cirriform, progressively smaller from chaetiger 4 and strongly reduced from chaetiger 8 onwards (Figure 4).

Notopodial chaetae of three types: (1) anterior row of 3 – 6 wide, short, limbate chaetae, densely granulated throughout length, decreasing in number after chaetiger 8, being absent after chaetiger 13; (2) posterior row of 5–10 long (up to 1.5 times longer than limbate chaetae) non-limbate chaetae and (3) from chaetiger 25–27, 2–4 long-shafted multidentate (3– 4 pairs of small teeth above main tooth) hooded hooks per rami, accompanied by 1–4 long non-limbate chaetae (Figure 5). Neuropodial chaetae of four types: (1) anterior row of 2 – 5 wide, short, limbate chaetae, densely granulated throughout length, decreasing in number after chaetiger 8, being absent after chaetiger 13; (2) posterior row of 4 – 8 long (up to 2 times longer than limbate chaetae) non- limbate chaetae; (3) from chaetiger 12 – 16, 2 – 5 long-shafted multidentate (3 – 4 pairs of small teeth above main tooth) hooded hooks per rami, accompanied by 1 – 4 long non-limbate chaetae and (4) lightly granulated sabre chaetae from chaetiger 9 – 10, only one per rami, almost straight at chaetigers 9 – 10 and more curved on following chaetigers (Figure 5).

Gametes not observed. Pygidium unknown.

Methyl green staining pattern: Margins of postchaetal notopodial lamellae of chaetigers 2 – 10 and whole dorsal and ventral surfaces of chaetigers 15 – 20 intensely stained.

Fig. 5. Spiogalea capixaba sp. nov., holotype: (A) sabre chaeta from neuropodium of chaetiger 14; (B) short limbate capillary chaetae; (C) long non-limbate capillary chaetae; (D) hooded hook. Scale bars: 20 mm.

Etymology

‘Capixaba’ is the common denomination given to natives of the Espı´rito Santo State, in southeastern Brazil, where the specimens were collected.

DISTRIBUTION Espírito Santo Basin, Doce Canyon, on muddy bottoms

(mainly silt), 950 m depth. The present record expands the geographic range of the genus to the Western Atlantic.

Remarks Spiogalea capixaba sp. nov. exhibited significant morphological differences from S. vieitezi, mainly the presence of a single prostomial chitinous plate instead of two, a T-shaped rather than bluntly rounded prostomium, biannulated peristomium, absence of low dorsal ridges, presence of notopodial hooded hooks and the position of neuropodial hooded hooks.

According to Aguirrezabalaga & Ceberio (2005), long non-limbate notopodial capillaries are absent, although such chaetae could be observed on the first 9 chaetigers of the paratype. The fact that notopodial hooded hooks are present in S. capixaba sp. nov. and lacking in S. vieitezi, is quite unusual, since this character is usually constant among species of the same genus. While notopodial hooded hooks are not present on all Scolelepis species (Rocha & Paiva, 2012) and absent on most Laonice species (Sikorski, 2003), in Aurospio, notopodial hooded hooks may start after the 30th chaetiger (Maciolek, 1981; Mincks et al., 2009; Paterson et al., 2016). Based on the original illustrations of S. vieitezi, it is likely that the type-specimen was regenerating the posteriormost chaetigers and the pygidium, since the authors described a pygidium of uncertain morphology that could not be seen clearly. Furthermore, the holotype had only 19 chaetigers, against 44 in S. capixaba. Thus, the absence of notopodial hooded hooks in S. vieitezi is doubtful considering that type-specimens are likely to be not complete.

According to Aguirrezabalaga & Ceberio (2005), the chitinous plates surrounding the anterior part of the prostomium are a feature that distinguishes Spiogalea from the remaining genera of Spionidae. Other features include the absence of branchiae, peristomium forming a posteriorly open collar encircling the prostomium, parapodia of the first chaetiger reduced and long-shafted multidentate neuropodial hooded hooks with complete hoods.

The presence, distribution and morphology of the branchiae are features of great taxonomic importance for spionids (Foster, 1969, 1971; Blake & Kudenov, 1978; Johnson, 1984; Maciolek, 1985; Blake, 1996; Bick, 2005; Delgado-Blas, 2009; Radashevsky, 2012). Branchiae are also useful to access phylogenetic relationships of the family, as seen in Sigvaldadóttir et al. (1997), Sigvaldadóttir (1998) and Blake & Arnofsky (1999). Absence of branchiae is an unusual character that probably evolved independently among spionid genera. It is recorded only in the genera

Spiophanes, Spiophanella, Glyphochaeta, some species of Polydorella Augener, 1914 (Williams, 2004), and a single species of Aurospio Maciolek, 1981 (Neal et al. in Paterson et al., 2016).

Spiogalea can be distinguished from Spiophanes based on the presence of a chitinous prostomial plate, absence of nuchal organs, absence of crook-like chaetae on chaetiger 1, and presence of multidentate long-shafted neuropodial hooks. Spiogalea can be distinguished from Spiophanella, a monotypic and doubtful genus (Sigvaldadóttir et al., 1997), based on the presence of a chitinous plate and by the shape of the peristomium, which is large, well-developed, separated from chaetiger 1, encircling the prostomium, and a reduced first chaetiger lacking notopodial postchaetal lobe and reduced postchaetal lobe. Spiophanella lacks notopodial multidentate hooded hooks, which are present at least in S. capixaba sp. nov. Glyphochaeta is a recently described genus, which comprises only Glyphochaeta laudieni Bick, 2005, described from Kongsfjorden, Spitsbergen. Spiogalea can be separated from G. laudieni based on the morphology of the prostomial projections, and the presence, in the former, of sabre chaetae, a chitinous plate and absence of grooved spines.

The genus Polydorella belongs to a complex of related genera informally called ‘Polydora- complex’ (Sigvaldadóttir et al., 1997), ‘polydorids’ (Blake, 1996) or ‘polydorins’ (Radashevsky, 2012), which comprises spionids with a modified 5th chaetiger bearing spines. According to Williams (2004), branchiae may be absent on some species, although Spiogalea species can be easily distinguished from Polydorella by the presence, in the former, of a chitinous plate and the absence of a modified 5th chaetiger bearing spines.

As for Aurospio, one of the main diagnostic characters are branchiae starting on chaetiger 3 (Maciolek, 1981; Mincks et al., 2009), although a new species without branchiae, Aurospio abranchiata Neal, Paterson & Soto in Paterson et al., 2016, was recently described in the Northeast Atlantic, near Portugal. Spiogalea species can be easily separated from Aurospio abranchiata based on the presence of a chitinous plate, position of notopodial hooded hooks and morphology of the postchaetal notopodial lamella of chaetiger 3, that is rounded in Spiogalea and greatly enlarged and nearly square-shaped in A. abranchiata. In conclusion, as both the lack of branchiae and presence of a prostomial chitinous plate are unusual features for the family Spionidae, the collection of more specimens of Spiogalea, including entire material suitable for morphological as well as molecular studies, would be of great relevance. The use of molecular techniques could provide valuable information on both the position of the genus and its relationships within Spionidae.

ACKNOWLEDGEMENTS

Thanks are due to Isabela Guerra for the illustrations, Natália Amaral for the map and Florencio Aguirrezabalaga for lending the paratype of Spiogalea vieitezi. We are grateful to CENPES/PETROBRAS for the opportunity and access to this material and to Benthos Ambiental and its staff for initially sorting out the specimens. The authors would also like to thank the anonymous referees for their comments and suggestions.

FINANCIAL SUPPORT

This work was supported by a fellowship to PCP (CNPq) and grants from FAPERJ and CNPq.

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Correspondence should be addressed to:

A.J.M. Peixoto Laboratório de Polychaeta, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, CCS, Ilha do Funda˜o, Rio de Janeiro, RJ CEP 21941-902, Brazil email: [email protected]

Capítulo 2

Zootaxa 4577 (3): 529 547 – ISSN 1175-5326 (print edition)

Article ZOOTAXA

https://www.mapress.com/j/zt/

New Prionospio and Laubieriellus (Annelida: Spionidae) species from Southeastern Brazil

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

1Laboratório de Polychaeta, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, CCS, Ilha do Fundão, Rio de Janeiro, RJ CEP 21941-902, Brazil.

2Programa de Biodiversidade e Biologia Evolutiva, Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, CCS, Ilha do Fundão, Rio de Janeiro, RJ CEP 21941- 902, Brazil.

3Corresponding author. E-mail: [email protected]

Abstract

Two new Prionospio species—Prionospio solisi sp. nov. and Prionospio nonatoi sp. nov. and a new Laubieriellus species—Laubieriellus decapitata sp. nov. are described from Espírito Santo Basin, Southeast Brazil. For both Prionospio species, branchial pattern differed from the genus diagnosis, which is therefore emended. For Laubieriellus, a new species is described, and attention is drawn to the notch in the ventral crests, a structure that holds taxonomic value.

Key words: Taxonomy, Polychaeta, morphology, emended diagnosis.

Introduction

The annelid family Spionidae Grube, 1850 is one of the most diverse and abundant families, frequently dominating soft-bottom communities, particularly in disturbed environments. Worldwide, there are approximately 580 species distributed in 39 genera (Blake et al. 2017), although this number may be an underestimate since the family has a long history of taxonomic problems, with many well-documented species-complexes (Sato-Okoshi et al. 2016), as well as genera that have never been revised.

Around 90 nominal species distributed in 18 genera of the family have been recorded for the Brazilian coast (Amaral et al. 2013; Peixoto & Paiva 2017), although recent surveys and taxonomic efforts suggest these numbers are underestimates, as many species remain undescribed, particularly in the deep-sea (Paiva & Barroso 2010).

Spionid taxonomic studies in Brazil are still limited, as most of them are geographically restricted to the southern and southeastern coasts of Brazil and frequently limited to shallow coastal environments (except Paiva & Barroso 2010 and Peixoto & Paiva 2017). Only a few genera have been studied in detail, such as Laonice Malmgren, 1867 (Nonato et al. 1986; Radashevsky & Lana 2009), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Polydora Bosc, 1802 (Radashevsky et al. 2006), Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018) and Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017). Large-scale studies are scarce, being restricted to Bolívar & Lana (1987) for the continental shelf of Paraná State, Southern Brazil, and Paiva & Barroso (2010) for the continental slope of Campos Basin, Rio de Janeiro State, Southeastern Brazil.

In Brazil, 14 Prionospio species have been recorded so far, mostly from shallow-water environments. Prionospio malmgreni Claparède, 1869, is also supposedly reported from Brazil, despite that it is considered as a taxon inquirendum and possibly represents more than one species (Maciolek 1985). It should be noted that most records should be used with caution, as these records include species described from distant locations and have never been confirmed, such as Prionospio steenstrupi Malmgren, 1867 (Iceland), P. cirrifera Wirén, 1883 (Russia), P. dubia Day, 1961 (South Africa), P. ehlersi Fauvel, 1928 (Morocco), P. pygmaeus Hartman, 1961 (USA), P. multibranchiata Berkeley, 1927 (Canada) and P. lighti Maciolek, 1985 (USA) (see Amaral et al. 2013 for detailed Brazilian records). The veracity of these records will be addressed in a future work.

The Prionospio generic complex represents a diverse and rich assemblage of species among the spionids, being comprised of seven currently valid genera: Apoprionospio Foster, 1969, Aurospio Maciolek, 1981a, Laubieriellus Maciolek 1981b, Orthoprionospio Blake & Kudenov, 1978, Paraprionospio Caullery, 1914, Prionospio Malmgren, 1867, and Streblospio Webster, 1879 (Blake et al. 2017). Two additional taxa, Minuspio and Aquilaspio, were erected by Foster (1971) as subgenera of Prionospio, but were latter synonymized by Sigvaldadóttir (1998), due to the lack of support for its monophyletic condition, thus making Prionospio the largest spionid genus, with over 100 species (Blake et al. 2017). The current diagnosis of Prionospio sensu lato comprises considerable morphological diversity, especially in regards to the distribution and arrangement of smooth and pinnate branchiae.

The genus Laubieriellus Maciolek 1981b was erected to include species of the Prionospio- complex that lack notopodial hooks, possess four pairs of smooth branchiae on chaetigers 2–5, and ventral crests on several anterior chaetigers, a set of characters exclusive to the genus. Only three species have been described so far: Laubieriellus salzi (Laubier 1970), from the Coast of Israel, Mediterranean Sea, L. grasslei Maciolek 1981b from the Galápagos Rift, and L. cacatua Erickson & Wilson, 2018 from Western Australia.

In the present work, two unusual Prionospio species, Prionospio nonatoi sp. nov. and P. solisi sp. nov., and a new Laubieriellus species, L. decapitata sp. nov., are presented and described.

The species were collected during the cruises of the “Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company), focusing on the southeastern coast of Brazil.

Materials and methods

Sediment samples were collected on the continental shelf using a Van Veen Grab sampler and on the continental slope using a box corer during the summer (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7), and winter (sample codes Amb2, Amb11, Amb12, Amb13 and Amb14) cruises of the AMBES project (Figure 1). Sediment was fixed in situ in 10% formaldehyde in seawater solution. In the laboratory, sediment was washed on a 300-µm-mesh sieve and transferred to 70% ethanol. Specimens were later sorted and examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope. Type material of Laubieriellus grasslei (USNM 65913 and USNM 65918) and L. salzi (USNM 42621) deposited in the Smithsonian National Museum of Natural History were also examined in this study and discrepancies found in described morphology were documented and illustrated.

Specimens were stained in an alcoholic solution of methyl green (1g/L in 70% EtOH) for 10 seconds and then transferred to distilled water, to examine the staining pattern. The stain Shirlastain A was used to enhance visualization of features such as branchial scars.

For SEM studies, specimens were dehydrated in a graded ethanol series, transferred to a graded HMDS (hexamethyldisilane) series, and subsequently dried in a fume hood. Afterwards, specimens were mounted on stubs, sputter coated with gold, and examined with a JEOL JSM-

6390LV scanning electron microscope. The HMDS approach was used because regular CO2 critical-point drying proved to be too destructive for Laubieriellus decapitata sp. nov. specimens. Two paratypes of Laubieriellus grasslei (USNM 65918) were dehydrated in a graded ethanol series, critical-point dried in CO2, mounted on stubs, sputter coated with carbon and gold- palladium and examined with a Philips XL-30 scanning electron microscope.

To examine hook morphology, prior to the treatment described above, posterior fragments were rinsed in distilled water and then sonicated in distilled water for a short time (between 15 to 40 sec) at 30 kHz to remove hoods from the hooks.

Type material was deposited in the Rio de Janeiro National Museum (MNRJP), Universidade Federal do Rio de Janeiro and the Museum of Zoology of the University of São Paulo (MZUSP), Universidade de São Paulo, both located in Brazil.

FIGURE 1. Map showing sampling area of the AMBES project (blue polygon).

Results

Taxonomy

Prionospio Malmgren, 1867

Type species: Prionospio steenstrupi Malmgren, 1867

Diagnosis (emended from Blake et al. 2017): Prostomium anteriorly rounded or truncate, sometimes weakly incised, often with peaks, without frontal horns; subtriangular, rectangular or oval in shape, caruncle extending at least to chaetiger 1; eyespots present or absent; occipital antenna absent. Peristomium at least partially fused with chaetiger 1, often surrounding prostomium with free, flattened lateral wings. Parapodia of chaetiger 1 reduced; noto- and

41 neuropodial lamellae largest in branchial region, reduced thereafter; notopodial lamellae often connected by low to high dorsal ridges or crests. Branchiae from chaetiger 2, chaetiger 3 or rarely absent. Branchiae limited to anterior chaetigers, 0–15 pairs, rarely more; branchiae all apinnate, all pinnate, or various combinations of both; pinnate branchiae with pinnules digitiform, not platelike; each branchia entirely free from dorsal lamella. Interparapodial pouches present or absent. Anterior chaetae limbate capillaries; posterior noto- and neuropodial hooded hooks present, bi-, tri-, or multidentate, with secondary hood. Neuropodial sabre chaetae present or absent. Pygidium with one long dorsomedial cirrus and two shorter ventrolateral lobes, all three sometimes fused.

Remarks: Blake et al.’s (2017) diagnosis was amended to include Prionospio solisi sp. nov. and Prionospio nonatoi sp. nov., which either lack branchiae or have only two pairs (see description and remarks for each species below). Changes in Blake et al.’s (2017) diagnosis have been highlighted in bold.

The Prionospio-complex represents one of the most morphologically diverse groups within the Spionidae and even within Prionospio sensu stricto, exhibiting remarkable branchial diversity, with many different arrangements. These different branchial arrangements were used in the past to erect other genera or subgenera within Prionospio (Foster 1971), such as Aquilaspio and Minuspio which are no longer considered as valid (e.g., Sigvaldadóttir 1998) by some authors and thus were synonymized with Prionospio, although such synonymy is not fully accepted and some researchers still consider Aquilaspio (Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011) and Minuspio (Aguirrezabalaga & Ceberio 2005; Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011; Dagli et al. 2011; Dagli 2013; Delgado-Blas & Díaz-Díaz 2013) as valid.

As for the species described in this work, despite bearing a branchial pattern (or lack of branchiae) that is similar to Aurospio, neither species could be placed in Aurospio, based on the lack of fusion of the branchiae to the notopodial lamellae (in Prionospio solisi sp. nov.) and the presence of a secondary hood on the hooded hooks.. Blake et al. (2017) noted that, over the past few years, several Prionospio species have been erroneously attributed to Aurospio. As further discussed in the Remarks section of each species, both species are morphologically similar to Prionospio. The placement of these species in Prionospio, however, should be treated as conservative, as we consider that the different branchial arrangements observed in both species do not justify the erection of a new taxon, neither within the Prionospio-complex nor within Prionospio sensu stricto.

Prionospio solisi sp. nov.

(Figures 2–5)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb6 D4R1, 19º 45' 55.39" S 39º 30' 25.74" W, 12/2011 to 02/2012, 149m, MZUSP 3390. Paratypes: Amb6 D4R1, 19º 45' 55.39" S 39º 30' 25.74" W, 12/2011 to 02/2012, 149m, MNRJP 1826 (6 ind), MZUSP 3391 (3 ind); Amb6 D4R2, 19º 45' 54.88" S 39º 30' 26.22" W, 12/2011 to 02/ 2012, 142m, MNRJP 1825 (2 ind);

Amb6 D4R3, 19º 45' 54.96" S 39º 30' 26.46" W, 12/2011 to 02/2012, 144m, MNRJP 1827 (1 ind).

Additional material examined. Brazil. Espírito Santo Basin. Amb 3 CAND4, 19º 31' 51.66" S 39º 3' 4.04" W, 171m (114 ind); Amb 3 CAND7, 19º 42' 18.42" S 39º 5' 52.41" W, 1307m (1 ind); Amb4 B8, 20º 41' 33.45" S 39º 35' 14.76" W, 1914m (1 ind); Amb5 C7, 20º 17' 41.07" S 39º 42' 38.02" W, 1358m (1 ind); Amb6 D4, 19º 45' 55.39" S 39º 30' 25.74" W, 149m (56 ind); Amb6 E4, 19º 36' 5.17" S 39º 10' 32.93" W, 153m (1 ind); Amb11 A5, 21º 4' 4.67" S 40º 13' 6.06" W, 415m (1 ind); Amb11 C7, 20º 17' 37.38" S 39º 42' 36.72" W, 1355m (1 ind); Amb11 E8, 20º 15' 55.44" S 38º 40' 48.23" W, 1892m, (1 ind); Amb11 F8, 20º 16' 38.17" S 38º 27' 26.52" W, 1904m (3 ind); Amb12 CAND4, 19º 31' 51.68" S 39º 3' 4.79" W, 171m (40 ind); Amb12 CAND6, 19º 37' 45.14" S 39º 3' 58.75" W, 1050m (12 ind).

Diagnostic features: Branchiae only on chaetigers 3–4, apinnate; dark yellow pigment present on anterior region.

Description. A small-sized spionid, largest individual about 7.5 mm long, 0.25 mm wide for 81 chaetigers, holotype 7.3 mm long, 0.23 mm wide for 65 chaetigers. Body cylindrical, slightly dorsoventrally compressed after branchial region and tapered towards the pygidium. Body color whitish in alcohol with conspicuous dark yellow pigment inclusions present on dorsal and ventral sides from chaetiger 4 or 5 to chaetigers8–12 (strongest on chaetigers 8–10), including parapodia (Fig 2A–C).

Prostomium anteriorly rounded, extending posteriorly as a narrow keel to the posterior margin of chaetiger 1 (Figs 2A–C; 3A; 4A–B). Two pairs of black eyes in trapezoidal arrangement (posterior pair faded in alcohol) present or eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lacking lateral wings. Grooved palps reaching up to chaetiger 8, lost in most individuals.

Chaetiger 1 with only a few short chaetae in both rami. Postchaetal lamellae rounded, reduced. Prechaetal lamellae absent.

Notopodial postchaetal lamellae triangular on chaetigers 2–4, largest on chaetigers 3 and 4, rounded with a pointed tip on chaetiger 5, rounded on chaetiger 6, and reduced to a low flap from chaetiger 7 onwards (Figs 2A–C; 3A; 4A–B). Dorsal crests absent. Prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae triangular with rounded corners on chaetiger 2, rounded on chaetigers 3 and 4 and flaplike from chaetiger 5, drastically reduced in size from chaetiger 7

43 onwards (Figs 2A–B; 3B; 4A); lamellae slightly more developed in both rami on last 4–7 chaetigers. Prechaetal lamellae absent throughout.

FIGURE 2. Prionospio solisi sp. nov., dorsal view (MZUSP 3391, paratypes). A. Anterior chaetigers. B. Anterior chaetigers and mid-body, showing characteristic pigmentation in ethanol. C. Anterior chaetigers stained with Shirlastain A. D. Posterior chaetigers and pygidium. Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Chaetae organized in two rows of unilimbate and sparsely granulated capillaries (Fig. 5A). In anterior notopodia, capillaries of posterior rows up to 1.3 times longer than capillaries in anterior row (Fig. 5B). Towards the posterior region, capillaries progressively become elongate, non- limbate, thin and less numerous (Fig. 5C).

Hooks in notopodia from chaetigers 48–61, up to two per fascicle, accompanied by 1–4 short non- limbate capillaries. Hooks in neuropodia from chaetigers 9–14, up to eight per fascicle, accompanied by 3–6 non-limbate capillaries. Neuropodial hooks slightly curved near the tip. Hooks multidentate, with 8–10 secondary teeth arranged in two rows above main tooth (Figs 3C; 5E). Small secondary hood present (Fig. 5E). Hooks accompanied by 3–10 short non-limbate capillaries.

Sabre chaetae with light granulations along the shaft, from chaetiger 9–11 (usually chaetiger 10) (Figs 3B; 5D). Two pairs of branchiae on chaetigers 3 and 4, of the same length or slightly longer than notopodial postchaetal lamellae (longer on chaetiger 3), completely free from notopodial lamellae. Branchiae triangular, flattened and densely ciliated (Figs 2A, C; 3A; 4A–B). Branchiae absent on chaetigers 2 and 5 (Figs 2A, C; 3A; 4A–B).

Pygidium with one long mid-dorsal cirrus and a pair of shorter ventral cirri (Figs 2D; 3D).

Oocytes from chaetigers 11–14, measuring up to 100 µm.

Methyl green pattern: Intense staining on prostomium, dorsal side of peristomium, dorsum of chaetigers 8– 16 and lateral sides of chaetigers 8–20.

Remarks: The placement of this species in any current genus is problematic since, based on branchiae starting on chaetiger 3 and pigmentation of the anterior chaetigers, it resembles Aurospio species. However, the branchiae are morphologically similar to the apinnate branchiae observed in chaetigers 3 and 4 of Prionospio sensu stricto (i.e., robust, flattened, densely ciliated, and free from the notopodial postchaetal lamellae), as well as the presence of a secondary hood on the hooded hooks, places the species closer to the genus Prionospio.

FIGURE 3. SEM of Prionospio solisi sp. nov. A. Anterior chaetigers, dorso-lateral view. B. Mid-body neuropodium, lateral view. C. Posterior neuropodium, dorso-lateral view. D. Pygidium, dorso-lateral view. Abbreviations: br, branchia; hh, hooded hook; py, pygidium, sa, sabre chaeta.

FIGURE 4. SEM of Prionospio solisi sp. nov., dorsal view. A. Anterior chaetigers and mid-body. B. Anterior chaetigers. Abbreviations: br, branchia; bs, branchial scars; nol, notopodial lamella; pe, peristomium; pr, prostomium.

Boundaries between both genera are still discussed, although, according to Blake et al. (2017), in the last years, several Prionospio species have been erroneously attributed to Aurospio (Sigvaldadóttir 2002; Mincks et al. 2009; Patterson et al. 2016). According to the current valid diagnosis, only species bearing branchiae starting from chaetiger 3 and partially fused to the notopodial postchaetal lamellae and lacking a secondary hood on the hooded hooks should be included in Aurospio, a view also followed in this work. Aurospio pilkena (Wilson, 1990), a species with a similar branchial arrangement and morphology, was described as Prionospio pilkena Wilson 1990 and placed in Aurospio by Sigvaldadóttir (1998), although Dagli & Çinar (2011) treat it as Prionospio (Minuspio) pilkena. An amended diagnosis of the genus Prionospio is proposed in this work to include species with branchiae starting from chaetiger 3 and not fused to the notopodial lamellae, rather than erecting a new taxon within the Prionospio-complex, which is beyond the scope of this work as it would require a review of all related taxa. Prionospio solisi sp. nov. differs from the remaining Prionospio species by the presence of conspicuous dark yellow pigment inclusions on the anterior region and the presence of only two pairs of branchiae, on chaetigers 3 and 4. According to Radashevsky (2012), late development of branchiae on anterior chaetigers is common in Prionospio, which could suggest that the species represents a juvenile stage. However, the same branchial pattern was observed in all individuals, regardless of size or sexual maturity.

FIGURE 5. Prionospio solisi sp. nov. A. Anterior capillary chaeta from posterior row. B. Anterior capillary chaeta from anterior row. C. Posterior capillary chaeta. D. Sabre chaeta. E. Hooded hook (as seen in light microscopy).

Regarding species recorded in Brazil, Prionospio solisi sp. nov. shares the absence of dorsal crests with P. dubia, P. lighti, P. perkinsi Maciolek, 1985 and P. pygmaeus. Prionospio solisi sp. nov.

46 differs from P. lighti and P. perkinsi by the lack of prostomial peaks, lack of lateral wings, starting chaetiger of notopodial hooded hooks and number of branchial pairs, up to 12 pairs in P. lighti and 10 pairs in P. perkinsi, starting chaetiger of sabre chaetae (in P. lighti) and absence of sabre chaetae (in P. perkinsi) (Maciolek, 1985). Prionospio solisi sp. nov. differs from P. pygmaeus by the prostomial shape, branchial pattern and starting chaetiger of notopodial and neuropodial hooded hooks. The species is most similar to P. dubia, sharing a similar prostomium shape, lack of prostomial peaks and lateral wings, but the species can be distinguished by branchial pattern, the starting chaetiger of neuropodial hooded hooks, and sabre chaetae.

Prionospio solisi sp. nov. is similar to Aurospio dibranchiata Maciolek 1981a and some individuals of A. banyulensis Laubier, 1966 due to the presence of only two pairs of branchiae, on chaetigers 3 and 4. However, P. solisi sp. nov. can be distinguished from A. dibranchiata by the morphology of branchiae—robust and completely free from the notopodial postchaetal lamellae, by the absence of dorsal crests, starting chaetiger of notopodial and neuropodial hooded hooks and presence of a secondary hood on the hooded hooks. Prionospio solisi sp. nov. can be distinguished from A. banyulensis by the starting chaetiger of notopodial and neuropodial hooded hooks, the shape of postchaetal neuropodial lamellae on anterior chaetigers and absence of dorsal crests.

Etymology: The species name, solisi, refers to the dark yellow to orange pigment observed on anterior chaetigers (solis, Latin for sun).

Habitat: muddy sand to mud, 142–1914 m depth.

Distribution: Northeastern Brazil (Alagoas and Sergipe states) and Southeastern Brazil (Espírito Santo and Campos basins).

Prionospio nonatoi sp. nov.

(Figures 6–9)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 D4, 19º 45' 54.56" S 39º 30' 25.23" W, 12/2011 to 02/2012, 144m, MZUSP 3387. Paratypes: Amb7 B4R1, 20º 35' 25.16" S 39º 54' 58.31" W, 12/2011 to 02/2012, 157m, MNRJP 1828 (3 ind), MZUSP 3388 (2 ind), MZUSP 3389 (2 ind).

Additional material: Amb7 B4, 20º 35' 25.16" S 39º 54' 58.31" W, 157m (47 ind); Amb7 D4, 19º 45' 54.56" S 39º 30' 25.23" W, 144m (15 ind); Amb7 E4, 19º 36' 4.32" S 39º 10' 34.07" W, 147m (6 ind).

Diagnostic feature: Branchiae absent.

Description: A small-sized spionid, largest individual about 4.8 mm long, 0.18 mm wide for 65 chaetigers, holotype 4.6 mm long, 0.18 mm wide for 62 chaetigers. Body cylindrical, slightly dorsoventrally compressed throughout body, tapered towards the pygidium (Fig. 6A–D; 8A). Color in alcohol white. Pigmentation absent.

Prostomium narrow, slightly widened towards the anterior margin, rounded anteriorly, extending posteriorly as a narrow keel to the posterior margin of chaetiger 1 (Figs 6A–D; 7A; 8A–B). Eyes absent. Prostomial peaks not observed. Prostomium and peristomium well-delimitated by a deep incision (Figs 6A–D; 8A–B). Peristomium short, surrounding prostomium and partially fused to chaetiger 1, lacking lateral wings. Palps lost in all individuals.

Chaetiger 1 with few and short chaetae on both rami. Postchaetal lamellae auricular, reduced. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–5, largest on chaetiger 3 (Figs 6A–D; 7A; 8A–B) and smaller on chaetigers 4 and 5. Lamellae rounded from chaetiger 6 to chaetiger 11–14 and reduced to a low flap afterwards. Notopodial prechaetal lamella absent throughout. Dorsal crests low, from chaetiger 8 to chaetiger 10– 15 (Fig. 8A).

Neuropodial postchaetal lamellae of chaetiger 2 well-developed, triangular and elongated ventrally, triangular and not elongated ventrally on chaetiger 3, rounded on chaetigers 4–11 and reduced to a low flap afterwards (Fig. 6D; 8A). Neuropodial prechaetal lamellae absent throughout.

Chaetae organized in two rows of sparsely granulated non-limbate capillaries (Fig. 9A). Towards the posterior region, capillaries progressively become elongate, thinner and less numerous (Fig. 9B).

Hooks in notopodia from chaetigers 32–44, up to three per fascicle, accompanied by 1–4 short, non-limbate capillaries. Hooks in neuropodia from chaetigers 11–12, up to seven per fascicle, accompanied by 1–4 non-limbate capillaries. Hooks multidentate, with 8 secondary teeth arranged in two rows above the main tooth (appearing as a single row of 4 secondary teeth in light microscopy) (Figs 7A–B; 9D). Small secondary hood present (Fig. 9D). Hooks accompanied by 3–10 short non-limbate capillaries.

Non-limbate and sparsely granulated sabre chaetae consistently from chaetiger 10 (Fig. 9C).

FIGURE 6. Prionospio nonatoi sp. nov. (MNRJP 1828, paratypes). A. Anterior chaetigers, dorsal view. B. Anterior chaetigers and mid-body, dorsal view. C. Anterior chaetigers and mid-body stained with Shirlastain A, dorsal view. D. Anterior chaetigers and mid-body stained with Shirlastain A, dorso-lateral view. E. Pygidium, dorsal view. Abbreviations: nol, notopodial lamella; pe, peristomium; pr, prostomium.

FIGURE 7. SEM of Prionospio nonatoi sp. nov. A. Anterior chaetigers, dorsal view. B. Hooded hooks, dorso-lateral view. C. Hooks (hoods removed), lateral view. D. Pygidium, lateral view. Abbreviations: hh, hooded hooks; py, pygidium.

Branchiae absent in all individuals (Figs 6A–D; 7A; 8A–B). Pygidium with a pair of short rounded ventral cirri and a slightly longer mid-dorsal cirrus (Figs 6E; 7D).

Oocytes from chaetigers 10–11, measuring up to 80 µm.

Methyl green pattern: Intense staining on prostomium and peristomium.

Remarks: According to Radashevsky (2012), the late development of branchiae on anterior chaetigers is common in Prionospio, which could lead to the assumption that the species represents a juvenile stage. However, the complete absence of branchiae was observed in all 76 individuals, regardless of size (from 2.5 to 4.8 mm long; from 39 to 65 chaetigers) or sexual maturity.

Owing to the lack of branchiae, the placement of this species in any current genus is problematic, since branchial morphology and distribution are characters of great significance for spionid taxonomy (Foster 1969, 1971; Blake & Kudenov 1978; Johnson 1984; Maciolek 1985; Blake 1996; Bick 2005; Delgado-Blas 2009; Radashevsky 2012; Blake et al. 2017). For the Prionospio- complex, lack of branchiae is unusual, shared only with Aurospio abranchiata Neal, Paterson & Soto in Paterson et al., 2016, although its placement in Aurospio is questioned by Blake et al. (2017), stating that, recently, several Prionospio species have been erroneously attributed to Aurospio.

FIGURE 8. Prionospio nonatoi sp. nov., dorsal view. A. Anterior segments and mid-body. B. Anterior segments. Abbreviations: dc, dorsal crest; nol, notopodial lamella; pe, peristomium; pr, prostomium.

FIGURE 9. Prionospio nonatoi sp. nov. A. Anterior capillary chaeta. B. Posterior capillary chaeta. C. Sabre chaetae. D. Hooded hooks (as seen in light microscopy).

Among Brazilian species, Prionospio nonatoi sp. nov. is similar to P. cirrifera, P. delta Hartman, 1965, P. fauchaldi Maciolek, 1985 and P. multibranchiata in having only low dorsal crests and significant overlapping in starting chaetiger of notopodial and neuropodial hooded hooks. However, not only do these species possess branchiae, but P. cirrifera, P. delta, and P. multibranchiata present multiple pairs of smooth apinnate branchiae, while P. fauchaldi presents distinctly wrinkled branchiae on chaetigers 2–5. Even if branchiae are completely lost in these species, they can still be separated from P. nonatoi sp. nov. by prostomial shape.

As for Aurospio abranchiata, both species are similar in having an enlarged notopodial postchaetal lamellae on chaetiger 3, distribution of dorsal crests, lack of branchiae, starting chaetiger of sabre chaetae and hooded hooks and presence of a secondary hood on the hooks. However, they can be separated based on the shape of notopodial postchaetal lamellae from chaetigers 2–5, the shape of neuropodial postchaetal lamellae from chaetigers 2–4 and by bathymetrical distribution.

Etymology: The species name, nonatoi, is a tribute to Edmundo Ferraz Nonato (1920–2014), who dedicated his life to the study of marine worms and is considered the “father” of Brazilian polychaetology. Habitat: Fine sand to muddy sand, at 144–153 m depth.

Distribution: Southeast Brazil (Espírito Santos and Campos Basins), and only found during the summer.

Genus Laubieriellus Maciolek, 1981b

Type-species: Laubieriellus grasslei Maciolek, 1981b

Diagnosis (emended from Erickson & Wilson, 2018): Prostomium anteriorly rounded, or with slight medial incision, extended posteriorly as a caruncle, occipital tentacle absent. Peristomium distinct from chaetiger 1, partly fused to prostomium. Four pairs of branchiae from chaetiger 2; branchiae elongate, cylindrical, smooth and distinct from notopodial lamellae. Neuropodial lamellae connected by ventral crests from chaetiger 2, rarely 1. Post-branchial notopodial lamellae connected in dorsal crests. Anterior chaetae all capillaries, multidentate hooded hooks present in posterior neuropodia. Notopodial hooks absent. Pygidium with two short ventrolateral lobes or cirri and one dorsomedial cirrus, or three subequal lobes or an undifferentiated ring.

Remarks: Maciolek (1981b) described Laubieriellus based on deep-sea specimens found on the Galápagos Rift and also placed Prionospio salzi Laubier, 1970 in Laubieriellus. Laubier (1970) considered his specimens as late-stage larvae, despite an adult morphology, as noted by Maciolek (1981b). The reexamination of the holotype (USNM 42621) also revealed the presence of oocytes from chaetiger 9–10.

Blake et al. (2017) considered Laubieriellus as part of the Prionospio-complex, as although morphologically similar to Prionospio, Laubieriellus species lack notopodial hooks and present ventral crests on several anterior chaetigers. The ventral crest, however, is not exclusive to Laubieriellus and has been observed in Prionospio rugosa Sigvaldadóttir, 1997 and Prionospio cristaventralis, Delgado-Blas et al. 2018. The genus diagnosis was emended to include a ventral crest on chaetiger 1, as observed in Laubieriellus grasslei (Fig. 10).

FIGURE 10. SEM of Laubieriellus grasslei (USNM 65918, paratypes). A. Anterior and median chaetigers, dorsal view. B. Anterior and median chaetigers, ventral view. C. Close-up of the ventral crests, ventral view. Abbreviations: br, branchia; cha1, chaetiger 1; cn, complete notch; dc, dorsal crests; pe, peristomium; pr, prostomium; vc, ventral crests.

The notch on the ventral crests, a character generally neglected, is present in Laubieriellus grasslei (Maciolek 1981b: Fig. 4A), L. cacatua Erickson & Wilson, 2018, L. decapitata sp. nov. (Fig. 11B; 12C) and L. salzi (Laubier 1970; Dagli 2013: Fig. 3A), as indicated in Laubier’s (1970) description: “Enfin, en avant du neuropode une crête fine se prolonge ventralement presque jusqu'h la ligne médioventrale.” (Finally, in front of the neuropodium, a fine crest extends ventrally almost to the medioventral line). This notch can be shallow and incomplete, as seen in L. cacatua and L. decapitata sp. nov., or a complete notch, as in L. salzi and L. grasslei (Fig 10B– C).

Laubieriellus decapitata sp. nov.

(Figures 11–13)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb1 Foz 14R3, 19º 42' 34.9" S 39º 39' 3.42" W, 12/11– 19/2010, 39m, MZUSP 3383. Paratypes: Amb6 D4R2, 19º 45' 54.88" S 39º 30' 26.22" W, 12/2011 to 02/2012, 142m, MNRJP 1830 (2 ind), MZUSP 3386 (2 ind); Amb6 E4R3, 19º 36' 5.08" S 39º 10' 32.85" W, 12/2011 to 02/ 2012, 149m, MNRJP 1829 (1 ind), MZUSP 3384 (1 ind), MZUSP 3385 (1 ind); Amb7 E3R3, 19º 26' 4.07" S 39º 17' 38.2" W, 12/2011 to 02/2012, 50m, MNRJP 1831 (5 ind).

Additional material: Amb1 Foz6, 19º 54' 56.16" S 39º 56' 40.83" W, 34m (1 ind); Amb1 Foz11, 19º 57' 32.89" S 39º 53' 30.69" W, 47m (1 ind); Amb1 Foz14, 19º 42' 32.21" S 39º 38' 57.36" W, 42m (4 ind); Amb1 Foz15, 19º 37' 48.27" S 39º 35' 25.83" W, 42m (15 ind); Amb1 Foz16, 20º 1' 3.73" S 39º 50' 13.76" W, 52m (5 ind); Amb1 Foz17, 19º 55' 44.66" S 39º 45' 38.7" W, 52m (3 ind); Amb2 Foz11, 19º 57' 32.36" S 39º 53' 33.01" W, 47m (2 ind); Amb2 Foz16, 20º 1' 2.6" S39º 50' 18.72" W, 51m (1 ind); Amb2 Foz17, 19º 55' 45.59" S39º 45' 41.35" W, 51m (3 ind); Amb2 Foz19, 19º 46' 10.69" S 39º 34' 55.84" W, 50m (1 ind); Amb2 Foz20b, 19º 41' 29.14" S 39º 31' 18.18" W, 53m (1 ind); Amb6 D4, 19º 45' 55.39" S 39º 30' 25.74" W, 132m (79 ind); Amb6 E4, 19º 36' 5.17" S 39º 10' 32.93" W, 153m (10 ind); Amb6 CAND4, 19º 31' 51.68" S 39º 3' 4.79" W, 171m (2 ind); Amb6 CANWN4, 19º 49' 7.27" S 39º 36' 8.52" W, 158m (2 ind); Amb7 A2, 21º 2' 47.31" S 40º 32' 28.94" W, 40m (1 ind); Amb7 A3, 21º 4' 1.29" S 40º 18' 50.11" W, 50m (2 ind); Amb7 B3, 20º 34' 53.42" S 40º 6' 27.43" W, 50 m (13 ind); Amb7 B4, 20º 35' 25.16" S 39º 54' 58.31" W, 157m (2 ind); Amb7 C2, 20º 11' 25.35" S 40º 2' 16.02" W, 39m (5 ind);Amb7

C3, 20º 12' 20.26" S 39º 57' 59.7" W, 50m (4 ind); Amb7 D1, 19º 35' 37.21" S 39º 41' 19.68" W, 26m (1 ind); Amb7 D2, 19º 40' 26.04" S 39º 36' 19.65" W, 40m,(4 ind); Amb7 D3, 19º 43' 14.34" S 39º 33' 34.86" W, 50m (6 ind); Amb7 D4, 19º 45' 54.56" S 39º 30' 25.23" W, 145m (6 ind); Amb7 E2, 19º 18' 5.9" S 39º 23' 23.3" W, 39m (3 ind); Amb7 E3, 19º 26' 5" S 39º 17' 38.92" W, 50m (33 ind); Amb7 E4, 19º 36' 4.32" S 39º 10' 34.07" W, 147m (2 ind); Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 39m (10 ind); Amb7 F3, 18º 53' 29.72" S 39º 6' 23.3" W, 52m (2 ind); Amb7 F4, 19º 33' 2.92" S38º 42' 52.26" W, 152m (1 ind); Amb7 G2, 18º 36' 31.68" S 39º 9' 33" W, 39m (5 ind); Amb7 G3, 18º 40' 55.3" S 38º 55' 41.48" W, 55m (11 ind); Amb12 D4 P300, 19º 45' 53.43" S 39º 30' 25.97" W, 146m (1 ind); Amb12 D4, 19º 45' 53.43" S 39º 30' 25.97" W, 146m (3 ind); Amb12 D4 T300,19º 45' 53.43" S 39º 30' 25.97" W, 144m (1 ind); Amb12 E4, 19º 36' 3.57" S 39º 10' 33.64" W, 152m (3 ind); Amb14 A2, 21º 3' 31.13" S 40º 22' 59.88" W, 40m (1 ind); Amb14 E2, 19º 18' 6.12" S 39º 23' 23.35" W, 38m (1 ind); Amb 14 F3, 18º 53' 31.97" S 39º 6' 21.78" W, 51m (13 ind); Amb13 G2, 18º 36' 32.45" S 39º 9' 32.83" W, 40m (9 ind); Amb13 G3, 18º 40' 57.41" S 38º 55' 39.92" W, 53m (30 ind).

Diagnostic features: Large auricular notopodial prechaetal and postchaetal lamellae on chaetiger 1; prostomium often obscured by peristomium. Description: A small-sized spionid, largest individual about 5 mm long, 0.3 mm wide for 45 chaetigers, holotype 3.5 mm long, 0,28 mm wide for 37 chaetigers. Body translucent and fragile, dorsoventrally flattened throughout. Pigmentation absent. Prostomium truncate, weakly incised anteriorly, eyes absent. Caruncle extending posteriorly to end of chaetiger 3. Prostomial peaks absent. Peristomium encircling the prostomium and partially fused with chaetiger 1, lateral wings absent (Figs 11; 12A–B). Grooved palps extending up to chaetiger 8, lost in most specimens. Chaetiger 1 with short chaetae on both rami. Notopodial and neuropodial postchaetal lamellae large, auricular-shaped, well-developed in adults, although smaller than those on following chaetigers (Figs 11; 12A–C). Notopodial prechaetal lamellae large and auricular-shaped. Notopodial postchaetal lamellae foliaceous and largest on chaetigers 2–5, extending to mid- dorsum (Figs 11A; 12A–B; 13A–D). Postchaetal lamellae round on chaetigers 6–7, progressively smaller on chaetigers 8–9 and reduced from chaetiger 10 onwards. Prechaetal lamellae absent. Dorsal crests high from chaetiger 7 to 9 and low crest from chaetiger 10 to chaetigers 11–12 (Figs 11A; 12A–B).

Neuropodial postchaetal lamellae rounded from chaetigers 2–9, broad in the branchial region, largest on chaetiger 3 and reduced from chaetiger 10 onwards. Prechaetal lamellae absent. Ventral crests with median notch from chaetiger 2 to chaetigers 9–11, shallow on chaetigers 2 and 3 (Figs 11B; 12C). Chaetae organized in two rows of non-limbate and lightly granulated capillaries. In anterior notopodia, capillaries of the posterior rows up to two times longer than capillaries from the anterior row (Fig. 13F–G). Towards the posterior region, capillaries progressively become elongate, thin and less numerous.

FIGURE 11. Laubieriellus decapitata sp. nov. (MNRJP 1831, paratypes). A. Anterior chaetigers, dorsal view. B. Anterior chaetigers, ventral view. Abbreviations: dc, dorsal crest; pe, peristomium; pr, prostomium; vc, ventral crest.

Neuropodial hooded hooks from chaetiger 9–10, up to eight per fascicle. Notopodial hooks absent. Hooks multidentate, with 6–8 secondary teeth arranged in two rows above main tooth (Figs 12D; 13H). Small secondary hood present. Hooks accompanied by 3–10 short non-limbate capillaries. Granulated sabre chaetae from chaetiger 10 (Fig. 13I). Branchiae on chaetigers 2–5, longer than postchaetal notopodial lamellae. Branchiae apinnate, cirriform on chaetigers 2 and 5 and robust, flattened and densely ciliated on chaetigers 3 and 4, all pairs narrowing abruptly at tips (Figs 11A; 12A–B; 13A–D). Pygidium with one long mid-dorsal cirrus and a pair of shorter ventral cirri (Fig. 13E). Oocytes from chaetigers 9–12, measuring up to 120 µm. Variation: Around 20% of all specimens presented two sabre chaetae per rami on chaetigers 10 and 11.

Methyl green pattern: Diffused throughout the body. Margins of lamellae slightly more stained from chaetiger 1 to chaetiger 10.

Remarks: The genus Laubieriellus was erected by Maciolek (1981b) to include species of the Prionospio-complex that bear ventral crests on anterior chaetigers, have cylindrical apinnate branchiae, and lack notopodial hooded hooks. Only three Laubieriellus species are known: L. grasslei, from hydrothermal vents on the Galápagos Islands (Ecuador), L. salzi, from the coast of Israel, Mediterranean Sea, and L. cacatua, from Western Australia. Laubieriellus species present remarkable character overlapping. Thus, characters such as caruncle length, first chaetiger with hooded hooks and sabre chaetae may not be helpful to distinguish species (Table 1). Laubieriellus decapitata sp. nov. is similar to L. cacatua in the distribution of dorsal and ventral crests, but these species can be distinguished by prostomial shape, shape of postchaetal neuropodial lamellae (especially on chaetiger 1), arrangement and number of secondary teeth on the hooded hooks, and pygidial shape. Laubieriellus decapitata sp. nov. can be readily

55 distinguished from L. grasslei by prostomial shape, distribution of dorsal and ventral crests, notch morphology, pygidial shape, and bathymetry.

FIGURE 12. SEM of Laubieriellus decapitata sp. nov. A. Anterior chaetigers, dorsal view. B. Anterior chaetigers, dorso-lateral view. C. Anterior chaetigers, ventral view. D. Hooks (hoods removed), lateral view. Abbreviations: br, branchia; dc, dorsal crest; pr, prostomium; vc, ventral crest.

Laubieriellus decapitata sp. nov. is most similar to L. salzi in prostomial shape, but the two species can be distinguished by the distribution of dorsal and ventral crests, shape of parapodial lamellae on chaetiger 1, shape of notopodial lamellae after chaetiger 10 (reduced in L. decapitata and digitiform in L. salzi) and number of secondary teeth on the hooded hooks.

Etymology: The epithet decapitata (Latin for beheaded) refers to the prostomium, which is difficult to distinguish as it is translucent and does not stand out from the surrounding structures, and is also encircled by the peristomium for the most part. This led many students to consider specimens as fragments with the prostomium absent. Habitat: Coarse to fine sand. Distribution: Espírito Santo and Campos basins, 25–150 m deep. The present record expands the geographic range of the genus to the Southern Atlantic Ocean.

FIGURE 13. Laubieriellus decapitata sp. nov. A. Parapodium from chaetiger 2. B. Parapodium from chaetiger 3. C. Parapodium from chaetiger 4. D. Parapodium from chaetiger 5. E. Pygidium. F. Anterior capillary chaetae from posterior row. G. Anterior capillary chaeta from anterior row. H. Hooded hooks (as seen in light microscopy). I. Sabre chaeta.

Acknowledgments

We are grateful to CENPES/PETROBRAS for the opportunity and access to this material and to Benthos Ambiental and its staff for initially sorting out the specimens. Thanks are due to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) for providing laboratory financial support. This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, granted to AJMP. AJMP was awarded a fellowship provided by the National Museum of Natural History, Smithsonian Institution, U.S.A. (Kenneth Jay Boss Fellowship in Invertebrate Zoology, 2018). PCP also has a research fellowship from CNPq. Finally, we would also like to thank Nancy J. Maciolek and an anonymous referee, whose insightful comments greatly improved the quality of this paper.

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Table

Capítulo 3

New apinnate Prionospio (Annelida: Spionidae) species from southeastern Brazil

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista alvo: Zootaxa

Status: Submetido, em processo de revisão

New apinnate Prionospio (Annelida: Spionidae) species from southeastern Brazil

Running title: NEW APINNATE PRIONOSPIO FROM BRAZIL.

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract

Eight apinnate Prionospio Malmgren, 1867 species were recorded from southeastern Brazil, both from coastal environments and the continental slope. Seven of these species — Prionospio acutus sp. nov., P. fosterae sp. nov., P. kinbergi sp. nov., P. mutatus sp. nov., P. cinthyae sp. nov., P. corrugatus sp. nov., and P. hartmanae sp. nov. are new to science, whereas P. cf. delta Hartman, 1965 have been previously recorded in Brazil. All novel species, except for P. cinthyae sp. nov. bear a similar number of branchiae, but can be distinguished based on the prostomial shape, presence of dorsal crests, branchiae morphology, presence of sabre chaetae, morphology of parapodial lamellae, and hook morphology. An identification key and a comparison table to all eight species recorded in the area are provided.

Keywords: Taxonomy, Polychaeta, morphology, biodiversity, Minuspio.

Introduction

The Spionidae Grube, 1850 are one of the most abundant and speciose families of annelid polychaetes, often dominating soft-bottom communities, especially in organically enriched environments. Worldwide, there are roughly 580 described species distributed in 39 genera (Blake et al. 2017), although this number most likely is underestimated, as the family has a history of taxonomic problems with many well-documented species-complexes (Sato-Okoshi et al. 2016), species with dubious distributions, and genera that have never been revised.

The first apinnate Prionospio species, Prionospio cirrifera, was described by Wirén (1883) from the Kara Sea, Arctic Ocean, northern Russia, but was subsequentially (and doubtingly) reported from France (Fauvel, 1927), Chile (as a synonym of P. patagonica Augener, 1923), the Pacific coast of Canada (as a synonym of P. multibranchiata Berkeley, 1927), Suriname (as a synonym of P. delta Hartman, 1965) (synonyms by Foster 1971), Australia, the Pacific coast of the USA, the Gulf of Mexico, the Mediterranean sea, and Brazil (Pardo et al. 2006; Amaral et al. 2013).

In a review of the genus Prionospio (Foster, 1971), the author synonymized three apinnate species with Prionospio cirrifera and transferred all seven remaining apinnate Prionospio species to the newly erected genus Minuspio. This genus included mostly multibranchiate species, although Foster’s (1971) diagnosis included apinnate species bearing four to 40 pairs of branchiae.

Minuspio was considered a subgenus of Prionospio by Day (1973), a placement followed by Blake & Kudenov (1978) and by Maciolek (1981; 1985), who described additional species belonging to the subgenus. Minuspio, alongside the subgenus Aquilaspio Foster, 1971 and Apoprionospio Foster, 1969 were later synonymized with Prionospio by Sigvaldadóttir (1998), who argued that, despite recognized shortcomings such as the small number of characters and taxa examined, apinnate branchiae is a plesiomorphy and may represent a non-homologous character. Because of these synonyms, the current diagnosis of Prionospio sensu lato includes a considerable morphological diversity regarding the number and arrangement of branchiae and over 100 known species (Blake et al. 2017). These synonyms, however, are not fully accepted, as both Aquilaspio (Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011) and Minuspio (Aguirrezabalaga & Ceberio 2005; Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011; Dagli et al. 2011; Dagli 2013; Delgado-Blas & Díaz-Díaz 2013) are frequently treated as valid, whereas Blake et al. (2017) reinstated Apoprionospio and raised it back to genus status.

To date, approximately 90 spionid species arranged in 19 genera have been recorded from Brazil (Amaral et al. 2013; Peixoto & Paiva 2019), but these numbers may be underestimated as a greater taxonomic effort coupled with molecular studies and recent surveys in previously unexplored areas revealed many undescribed species, especially from the deep sea (Paiva & Barroso 2010).

The current state of knowledge of spionid diversity in Brazil is uneven, as most studies are limited to shallow-water and coastline environments (except for Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019) and geographically restricted to the southern and southeastern coast of Brazil. A small number of genera have been studied in greater detail (but still restricted to South and Southeast Brazil) such as Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Polydora Bosc, 1802 (Radashevsky et al. 2006), Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017), Laonice Malmgren, 1867 (Kinberg 1866, Nonato et al. 1986; Radashevsky & Lana 2009), Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018), Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), and Laubieriellus Maciolek, 1981 (Peixoto & Paiva 2019). Wide-scale surveys are rare, restricted to the continental shelf of the state of Paraná, southern Brazil (Bolívar & Lana 1987) and the continental slope of Campos Basin, state of Rio de Janeiro, southeastern Brazil (Paiva & Barroso 2010).

To date, 16 Prionospio species have been recorded in Brazil, mostly from shallow-water environments (Peixoto & Paiva 2019). An additional species that is treated as a taxon inquirendum, Prionospio malmgreni Claparède, 1869 is also supposedly reported from the states of Rio de Janeiro and São Paulo, southeastern Brazil (Amaral et al. 2013). Six apinnate Prionospio species have been recorded in Brazil: P. cirrifera Wirén, 1883, P. delta Hartman, 1965, described from Suriname, P. fauchaldi Maciolek, 1985, described from Massachusetts (USA), P. lighti Maciolek, 1985, described from Washington (USA), P. multibranchiata Berkeley, 1927, described from British Columbia, Canada, and P. perkinsi Maciolek, 1985, described from the Gulf of Mexico, Florida (USA). Brazilian records, however, should be viewed with caution, as many species recorded (compiled by Amaral et al. 2013) are known species- complexes or species described from rather distant locations, whose records have never been confirmed. The veracity of these records will be addressed in a future work.

In the present work, eight Prionospio species are recorded based on material collected off the state of Espírito Santo during the cruises of the “Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company) and focusing on the southeastern coast of Brazil. Seven of these species are new to science, whereas the remaining species was previously recorded from nearby locations.

Materials and Methods

During the cruises of the AMBES Project, sediment samples were collected during the summer (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and winter (sample codes Amb2,

Amb11, Amb12, Amb13, and Amb14) from 2010 to 2013 on Espírito Santo Basin and the northern portion of Campos Basin (Fig 1), southeastern Brazil. Samples were collected on the continental shelf using a Van Veen Grab sampler and on the continental slope using a box corer and fixed in 10% formalin-seawater solution in situ. Samples were taken to the laboratory, washed, elutriated on a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm-mesh sieve (for samples collected on the continental slope and on the Watu Norte and Doce canyons), and transferred to 70% ethanol. Next, specimens were sorted and examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope. Measurements were taken using a calibrated eyepiece attached to the microscope. Main taxonomic characters are summarized in Table 1. An identification key is also provided for all species recorded in this study. Because the specimens were fixed in 10% formalin, molecular studies were not possible.

Specimens lots deposited in the Smithsonian National Museum of National History (USNM), Smithsonian Institution, Washington, D.C., USA, corresponding to type material of apinnate Prionospio species recorded in Brazil — Prionospio perkinsi (USNM 67679 – holotype and USNM 67681 – paratypes) and P. lighti (USNM 74740 – holotype and USNM 74741 – paratypes) — were examined and compared with material collected in this study. Non-type material of Prionospio delta (USNM 80299 and USNM 80300) and P. fauchaldi (USNM 67700 and USNM 67701) were also examined.

Specimens were stained in a methyl green solution (1 g/L in 70% EtOH) for 5‒15 s and transferred to distilled water to enhance visualization of features such as branchial scars and to observe any species-specific staining patterns.

Prior to scanning electron microscope (SEM) studies, specimens were dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each) followed by two changes of 100% ethanol. Specimens were then transferred to an ascending hexamethyldisilane (HMDS) series (2 parts 100% EtOH:1 part HMDS, 1:1, 1:2, pure HMDS, followed by a second change of pure HMDS, 15 min each), and dried in a fume hood. Next, specimens were mounted on stubs, sputter coated with gold, and viewed in a JEOL JSM-6390LV SEM. The HMDS method was used because regular CO2 critical-point drying repeatedly proved to be too damaging for our specimens, causing the loss of some branchiae. Regarding the material deposited in the USNM, two specimens of Prionospio fauchaldi (USNM 67701) were dehydrated in a graded ethanol series, critical-point dried in CO2, mounted on stubs, sputter coated with carbon and gold-palladium (10 µm thick each), and examined in a Philips XL-30 SEM.

To examine hook morphology using the SEM, before the technique described above, posterior fragments were rinsed in contact lens cleaning solution and sonicated in distilled water for 5‒15 s at 30 kHz to remove hoods from the hooks.

Type material was deposited in the Museu Nacional do Rio de Janeiro (MNRJP), Universidade Federal do Rio de Janeiro; Museu de Zoologia Adão José Cardoso (ZUEC-POL), Universidade Estadual de Campinas; and in the Museu de Zoologia (MZUSP), Universidade de São Paulo, all located in Brazil.

Results

Taxonomy

Phylum Annelida Lamarck, 1818

Order Spionida Fauchald, 1977

Family Spionidae Grube, 1850

Genus Prionospio Malmgren, 1867

Type species: Prionospio steenstrupi Malmgren, 1867.

Diagnosis (from Peixoto & Paiva 2019): Prostomium anteriorly rounded or truncate, sometimes weakly incised, often with peaks, without frontal horns; subtriangular, rectangular or oval in shape, caruncle extending at least to chaetiger 1; eyespots present or absent; occipital antenna absent. Peristomium at least partially fused with chaetiger 1, often surrounding prostomium with free, flattened lateral wings. Parapodia of chaetiger 1 reduced; noto- and neuropodial lamellae largest in branchial region, reduced thereafter; notopodial lamellae often connected by low to high dorsal ridges or crests. Branchiae from chaetiger 2, chaetiger 3, or rarely absent. Branchiae limited to anterior chaetigers, 0‒15 pairs, rarely more; branchiae all apinnate, all pinnate, or various combinations of both; pinnate branchiae with pinnules digitiform, not platelike; each branchia entirely free from dorsal lamella. Interparapodial pouches present or absent. Anterior chaetae limbate capillaries; posterior noto- and neuropodial hooded hooks present, bi-, tri-, or multidentate, with secondary hood. Neuropodial sabre chaetae present or absent. Pygidium with one long dorsomedial cirrus and two shorter ventrolateral lobes, all three sometimes fused.

Prionospio acutus sp. nov.

(Figures 2‒5)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 B5, 20° 35’ 15.16” S 39° 53’ 46.36” W, 02 Dec 2011 to 02 Feb 2012, 991 m, MZUSP XXXX. Paratypes: Amb3 E5, 19° 36’ 25.08” S 39° 10’ 20.15” W, 02 Dec 2011 to 02 Feb 2012, 352 m, MZUSP XXXX (4 ind); Amb11 B5, 20° 35’ 13.87” S 39° 53’ 45.78” W, 06 Jun 2013 to 17 Jul 2013, 382 m, MNRJP XXXX (4 ind).

Diagnostic features: Prostomium bearing two short antero-lateral projections, dorsal crests low from chaetigers 10‒11 to chaetigers 15‒17, and lack of sabre chaetae.

Description. A medium-sized Prionospio, largest specimen about 8 mm long, 0.35 mm wide at the widest point for 63 chaetigers (incomplete), holotype 7 mm long, 0.3 mm wide at the widest point for 49 chaetigers (incomplete). Body dorsoventrally flattened on the branchial region and cylindrical afterwards, tapering towards the pygidium. Body color yellow to whitish in alcohol (Fig. 2).

Prostomium triangular, rounded anteriorly, bearing two short antero-lateral projections, extending as a narrow keel almost to the posterior margin of chaetiger 2, flanked by conspicuous nuchal organs extending up to the margin of chaetiger 2 (Figs. 2; 3A). Eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lacking lateral wings. Grooved palps reaching up to chaetiger 10, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on both rami, smaller than lamellae on succeeding chaetigers, especially on the neuropodium (Figs. 3A; 4A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2‒15 (Fig. 4A–D), rounded with a pointed tip directed upwards from chaetiger 16 to chaetigers 20‒21 (depending on specimen size) and rounded afterwards, gradually reducing in size towards the posterior region, present as a low flap on the last chaetigers. Notopodial prechaetal lamellae absent throughout (Fig. 3B). Dorsal crests low, from chaetigers 10‒11 to chaetigers 15‒17. Ciliary bands present on branchiate chaetigers (Fig. 3A–B).

Neuropodial postchaetal lamellae rounded on chaetigers 2‒15 and rounded with a pointed tip directed downwards (Fig. 4A–D), from chaetiger 16 to chaetigers 18–20 (depending on specimen size), gradually becoming rounder and reducing in size towards the posterior region, present as a low flap on the last chaetigers. Neuropodial prechaetal lamellae absent throughout (Fig. 3B).

Chaetae from notopodia and neuropodia organized in two rows of unilimbate (limbation almost imperceptible) and intensely granulated capillaries (Fig. 5A). Chaetae from both rows of equal length, although neuropodial chaetae are slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner, and less numerous (Fig. 5B).

Hooks in notopodia starting from chaetigers 24‒42, up to two per fascicle, accompanied by row of 1‒4 short non-limbate capillaries (Fig. 5C–D). Hooks in neuropodia starting from chaetigers 13‒24, up to five per fascicle, accompanied by 3‒6 non-limbate capillaries. Hooks multidentate, with six small secondary teeth arranged in two rows above the main tooth (Fig. 5C). Secondary hood absent (Fig. 5C). Sabre chaetae absent throughout.

Up to 11 pairs of smooth, robust and flattened branchiae, starting from chaetiger 2, up to 2.5 times longer than notopodial postchaetal lamellae, gradually reducing in length towards last branchial pair. Branchiae moderately ciliated towards the tip, completely free from notopodial postchaetal lamellae (Fig. 5E).

Pygidium of unknown morphology.

Oocytes from chaetiger 13, measuring up to 120 µm.

Methyl green staining pattern: No specific pattern observed; light green pigment diffused throughout the body, except for anterior prostomial margin and branchial tips.

Remarks. The prostomial shape of Prionospio acutus sp. nov. is uncommon among Prionospio species, but similar to the prostomium observed in specimens of Spiogalea Aguirrezabalaga & Ceberio, 2005 (although lacking chitinous plates), forming antero-lateral projections rather than typical antero-lateral horns, as found in the genera Malacoceros Quatrefages, 1843, Rhynchospio Hartman, 1936, and Scolecolepides Ehlers, 1907.

Among Prionospio species, only Prionospio cornuta Hylleberg & Nateewathana, 1991, described from Thailand and P. paradisea Imajima, 1990a, described from Japan also bear antero-lateral projections, although both species bear sabre chaetae (lacking in Prionospio acutus sp. nov.) and only four pairs of branchiae, pinnate on chaetigers 2 and 5. In addition, the antero-lateral projections (which the authors referred to as horns) of P. cornuta are actually short and rounded (Hylleberg & Nateewathana, 1991, Fig. 5 K–N).

The lack of sabre chaetae is uncommon among Prionospio species, being observed only in Prionospio perkinsi, P. fosterae sp. nov., P. cerastae Radashevsky, 2015, described from Lizard Island, Australia, and P. hermesia Neal & Paterson in Paterson et al., 2016, described from Portugal. Prionospio acutus sp. nov. is most similar to P. perkinsi and P. fosterae sp. nov. in

71 having a similar number of branchiae (10 branchial pairs in P. perkinsi and 11 pairs in P. fosterae sp. nov. and P. acutus sp. nov.), but different morphology, as P. acutus sp. nov. bears robust flattened branchiae whereas P. perkinsi and P. fosterae sp. nov. bear cirriform branchiae. Additionally, P. acutus sp. nov. bears dorsal crests, present from chaetigers 10‒11 to chaetigers 15‒17, which are absent in P. fosterae sp. nov. and P. perkinsi.

Prionospio acutus sp. nov. is similar to P. hermesia in having low dorsal crests, although those occur on different chaetigers — from chaetigers 10‒11 to chaetigers 15‒17 in P. acutus sp. nov. and from chaetiger 8 to about chaetiger 25 in P. hermesia. However, both species can be readily distinguished based on the number of branchiae, only two pairs in P. hermesia and up to 11 pairs in P. acutus sp. nov. As for Prionospio cerastae, which is the only known Prionospio species to bear antero-lateral horns, specimens can be distinguished by the lack of dorsal crests in P. cerastae and the branchial morphology and distribution, as this species possesses only four pairs of branchia, the last pair bearing digitiform pinnules, whereas P. acutus sp. nov. bears up to 11 pairs of apinnate branchiae.

Etymology. The specific epithet, acutus, refers to the antero-lateral projections on the prostomium (acutus, Latin for sharp or pointed).

Habitat: muddy sand to mud, 349‒1327 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), South Atlantic Ocean.

Prionospio fosterae sp. nov.

(Figures 6‒10)

Prionospio perkinsi: Pardo & Peixoto (in press). Not Maciolek, 1985.

Prionospio lighti: Pardo & Peixoto (in press). Not Maciolek, 1985.

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 D1, 19° 35’ 37.21” S 39° 41’ 19.68” W, 02 Dec 2011 to 02 Feb 2012, 21 m, MZUSP XXXX. Paratypes: Amb3 F5, 19° 34’ 21.01” S 38° 41’ 16.99” W, 02 Dec 2011 to 02 Feb 2012, 438 m, MZUSP XXXX (2 ind), ZUEC- POL XXXX (2 ind).

Diagnostic features: Tridentate hooded hooks, lack of dorsal crests, and lack of sabre chaetae.

Description. A medium-sized Prionospio, largest complete specimen about 8 mm long, 0.3 mm wide at the widest point for 60 chaetigers, holotype complete, 7.5 mm long, 0.3 mm wide at the widest point for 56 chaetigers. Body dorsoventrally flattened throughout, tapering towards the pygidium. Body color whitish in alcohol (Fig. 6).

Prostomium narrow and rounded anteriorly, slightly widened towards the anterior margin, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2. Nuchal organs reaching posterior margin of chaetiger 1 (Fig. 7A–B). One or two pairs of minute reddish eyes generally present. Prostomial peaks absent. Peristomium surrounding prostomium, partially fused to the first chaetiger. Lateral wings absent. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, especially on the notopodium, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on both rami, much smaller than lamellae on succeeding chaetigers (Fig. 8A). Prechaetal lamellae absent.

Well-developed foliaceous notopodial postchaetal lamellae on chaetigers 2‒9 (Fig. 8A–D), square-shaped and drastically reducing in size towards the posterior region (Fig. 7A–B). Notopodial prechaetal lamella absent throughout. Dorsal crests absent. Ciliary bands present on branchiate chaetigers (Fig. 7B).

Neuropodial postchaetal lamellae rounded on the branchial region — from chaetiger 2 to chaetigers 8‒12 (depending on specimen size) (Fig. 8A–D), drastically reduced afterwards, present as a low flap on last chaetigers. Neuropodial prechaetal lamellae absent throughout (Fig. 7A).

Chaetae from notopodia and neuropodia organized in two rows, anterior row of short, robust and unilimbate capillaries and posterior row of long (up to two times longer than anterior row), thin and narrowly unilimbate capillaries (Fig. 9A–B). Towards the posterior region, chaetae of both rows progressively become elongate, non-limbate, thinner, and less numerous (Fig. 9C).

Hooks in notopodia starting from chaetigers 20‒23, up to five per fascicle, accompanied by 1‒5 short non-limbate capillaries (Figs. 7C; 9D–E). Hooks in neuropodia starting from chaetigers 13‒ 16, up to nine per fascicle, accompanied by 3‒7 non-limbate capillaries. Hooks tridentate, with a single pair of teeth arranged side by side above the main tooth (Fig. 9D). Small secondary hood present (Fig. 9D). Sabre chaetae absent throughout.

Up to 11 pairs of smooth and cirriform branchiae, tapered at the tips. Branchiae starting from chaetiger 2, up to four times longer than notopodial lamellae, gradually reducing in length towards last branchial pair. Branchiae densely ciliated throughout the length, completely free from notopodial postchaetal lamellae (Fig. 10A).

Pygidium bearing one dorsal cirrus and two short ventral cirri (Fig. 10B).

Gametes not observed.

Methyl green staining pattern: anterior margin of the prostomium and ventral side of the peristomium intensely stained; postchaetal lamellae slightly stained throughout the body.

Remarks. Prionospio fosterae sp. nov. is unusual among Prionospio species for lacking sabre chaetae — a character shared only with P. acutus sp. nov. (described above) and Prionospio perkinsi, in addition to P. cerastae and P. hermesia, which have never been recorded in Brazil.

Prionospio fosterae sp. nov. can be promptly distinguished from P. acutus sp. nov. by the lack of antero-lateral prostomial projections, the morphology of the hooded hooks (one pair of secondary teeth in P. fosterae sp. nov. and three pairs in P. acutus sp. nov.), branchiae morphology (branchiae up to 2.5 times longer than notopodial lamellae in P. acutus sp. nov. and up to four times longer in P. fosterae sp. nov.), and lack of dorsal crests in P. fosterae sp. nov., present from chaetigers 10‒11 to chaetigers 15‒17 in P. acutus sp. nov.

The lack of sabre chaetae and tridentate hooded hooks of P. fosterae sp. nov. makes it similar to P. hermesia (which, according to the authors, bears bidentate, tridentate, and possibly multidentate hooded hooks), although both species can be distinguished by the prostomial shape (narrowed in P. fosterae sp. nov. and oval in P. hermesia), branchial morphology and arrangement (up to 11 pairs of flattened branchiae in P. fosterae sp. nov. and two pairs of cirriform branchiae in P. hermesia), and lack of dorsal crests in P. fosterae sp. nov.

The delicate sabre chaetae of P. fauchaldi and P. corrugatus sp. nov. could lead researchers to conclude that these species lack sabre chaetae, but both species can be distinguished from P. fosterae sp. nov. by the presence of long neuropodial chaetae on the posterior row of chaetiger 3, by the prostomial shape (narrowed in P. fosterae sp. nov. and rectangular in P. fauchaldi and P. corrugatus sp. nov.), and branchiae morphology (up to 11 pairs of smooth flattened branchiae in P. fosterae sp. nov.; four pairs of branchiae, sculptured on chaetigers 2 and 5 in P. fauchaldi; and six pairs of branchiae, sculptured on chaetiger 2 and chaetigers 5‒7 in P. corrugatus sp. nov.). Sabre chaetae are supposedly lacking in P. kaplani Altamira, Glover, & Paterson in Paterson et al. 2016, described from Cape Verde Abyssal Plain and P. branchilucida Altamira, Glover, & Paterson in Paterson et al., 2016, although the authors only examined anterior fragments reaching 20 chaetigers at most.

Prionospio fosterae sp. nov. is remarkably similar to P. perkinsi in the prostomial shape (narrow and rounded anteriorly), lack of sabre chaetae, lack of dorsal crests, and branchial morphology and distribution (up to 11 pairs in P. fosterae sp. nov. and up to 10 pairs in P. perkinsi, apinnate and cirriform in both species). The starting chaetiger of notopodial and neuropodial hooded hooks also exhibit considerable overlapping (Prionospio fosterae sp. nov.: chaetigers 20‒23 [notopodial hooks] and 13‒16 [neuropodial hooks]; Prionospio perkinsi: 16‒23 [notopodial hooks] and 13‒ 18 [neuropodial hooks]). Due to those similarities, P. perkinsi has been mistakenly recorded from the Brazilian coast (Amaral et al. 2013; Pardo & Peixoto in press). However, P. fosterae sp. nov. can be distinguished from P. perkinsi by the absence of prostomial peaks, the shape of notopodial

74 postchaetal lamellae on post-branchial chaetigers (square-shaped in P. fosterae sp. nov. and triangular in P. perkinsi), the shape of neuropodial postchaetal lamellae (rounded in P. fosterae sp. nov. and square-shaped in P. perkinsi), and the number of secondary teeth above the main tooth (one pair in P. fosterae sp. nov. and two pairs in P. perkinsi).

Prionospio fosterae sp. nov. is also similar to P. lighti, as both species possess an anteriorly rounded prostomium, branchiae of similar distribution (up to 11 pairs in P. fosterae sp. nov. and up to 12 pairs in P. lighti), an overlap on the starting chaetiger of neuropodial hooded hooks (chaetigers 13‒16 in P. fosterae sp. nov. and chaetigers 13‒18 in P. lighti), and a lack of dorsal crests. Those similarities probably led to Brazilian records of P. lighti (Amaral et al. 2013), a species originally described from the Pacific coast of the USA. Despite the similarities, the two species can be separated based on the lack of prostomial peaks and sabre chaetae in P. fosterae sp. nov., starting chaetiger of notopodial hooded hooks (chaetigers 20‒23 in P. fosterae sp. nov. and chaetigers 25‒40 in P. lighti), and number of secondary teeth on the hooded hooks (one pair in P. fosterae sp. nov. and three pairs in P. lighti).

The occurrence of tridentate hooded hooks is unusual among Prionospio species, documented only in Prionospio tridentata Blake & Kudenov, 1978, described from Australia and P. fosterae sp. nov., but the two species differ on the prostomial morphology (narrowed in P. fosterae sp. nov. and trapezoidal in P. tridentata), branchial morphology and arrangement (up to 11 pairs of apinnate cirriform branchiae in P. fosterae sp. nov. and four pairs of branchiae, the last pair being pinnate in P. tridentata), and in the presence of sabre chaetae in P. tridentata. In some species, the hooded hooks may look tridentate, as in P. hermesia (which may bear tridentate hooks among bidentate and multidentate hooded hooks) and P. cerastae, but a closer inspection or SEM images may reveal the actual number of secondary teeth and its arrangement on the hooded hooks.

Etymology. The specific epithet, fosterae, is a tribute to Nancy M. Foster (1941‒2000), a polychaetologist who erected the (now synonymized) genus Minuspio (Foster, 1971), which included Prionospio species bearing only smooth branchiae.

Habitat: muddy sand to mud, 11‒438 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), South Atlantic Ocean.

Prionospio kinbergi sp. nov.

(Figures 11–15)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb2 Foz7, 19º 49' 50.65" S 39º 52' 22.92" W, 12 Jul 2011 to 18 Jul 2011, 28m, MZUSP XXXX. Paratypes: Amb7 A2, 21º 3' 26.62"

S 40º 23' 0.67" W, 02 Dec 2011 to 02 Feb 2012, 36 m, MZUSP XXXX (4 ind); Amb12 CAND4, 19º 31' 51.68" S 39º 3' 4.79" W, 06 Jun 2013 to 17 Jul 2013, 163 m, ZUEC-POL XXXX (5 ind); Amb14 C2, 20º 11' 25.75" S 40º 2' 15.87" W, 06 Jun 2013 to 17 Jul 2013, 33m, MNRJP XXXX (5 ind).

Additional material examined. Amb1 Foz7, 19º 49' 54.12" S 39º 52' 11.79" W, 29m (3 ind); Amb1 Foz10, 19º 35' 11.01" S 39º 38' 37.65" W, 29m (1 ind);; Amb1 Foz14, 19º 42' 32.21" S 39º 38' 57.36" W, 35m (1 ind); Amb1 Foz16, 20º 1' 3.73" S 39º 50' 13.76" W, 48m (1 ind); Amb1 Foz17, 19º 55' 44.66" S 39º 45' 38.7" W, 46m (1 ind); Amb2 Foz1, 19º 52' 15.25" S 39º 59' 42.87" W, 19m (1 ind); Amb2 Foz7, 19º 49' 50.65" S 39º 52' 22.92" W, 28m (11 ind); Amb2 Foz11, 19º 57' 30.39" S 39º 53' 35.28" W, 46m (10 ind); Amb2 Foz14, 19º 42' 28.53" S 39º 39' 4.25" W, 36m (6 ind); Amb2 Foz16, 20º 1' 1.92" S 39º 50' 18.19" W, 49m (1 ind); Amb2 Foz17, 19º 55' 43.7" S 39º 45' 39.68" W, 43m (2 ind); Amb2 Foz18, 19º 50' 16.34" S 39º 40' 10.8" W, 46m (3 ind); Amb3 CAND4, 19º 31' 51.66" S 39º 3' 4.04" W, 140m (1 ind); Amb6 D4, 19º 45' 54.96" S 39º 30' 26.46" W, 121m (2 ind); Amb7 A2, 21º 3' 26.62" S 40º 23' 0.67" W, 36 m (11 ind); Amb7 A3, 21º 4' 1.29" S 40º 18' 50.11" W, 46m (2 ind); Amb7 A4, 21º 4' 4" S 40º 14' 15.31" W, 142m (1 ind); Amb7 B2, 20º 34' 46.54" S 40º 11' 30.67" W, 35m (1 ind); Amb7 B3, 20º 34' 53.69" S 40º 6' 27.35" W, 45m (8 ind); Amb7 C2, 20º 11' 25.35" S 40º 2' 16.02" W, 35m (5 ind); Amb7 C3, 20º 12' 20.26" S 39º 57' 59.7" W, 44m (1 ind); Amb7 F1, 18º 42' 56.07" S 39º 31' 28.8" W, 20m (1 ind); Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 34m (47 ind); Amb7 F3, 18º 53' 29.72" S 39º 6' 23.3" W, 43m (6 ind); Amb7 G2, 18º 36' 31.68" S 39º 9' 33" W, 30m (10 ind); Amb12 E4, 19º 36' 3.57" S 39º 10' 33.64" W, 142m (1 ind); Amb12 CAND4, 19º 31' 51.68" S 39º 3' 4.79" W, 163 m (7 ind); Amb13 G2, 18º 36' 31.38" S 39º 9' 33.56" W, 28m (1 ind); Amb13 G3, 18º 40' 57.41" S 38º 55' 39.92" W, 44m (7 ind); Amb14 A4, 21º 4' 4.81" S 40º 14' 13.86" W, 147m (1 ind); Amb14 B1, 20º 34' 29.6" S 40º 20' 54.56" W, 21m (5 ind); Amb14 B3, 20º 34' 53.05" S 40º 6' 27.68" W, 43m (1 ind); Amb14 B4, 20º 35' 26.19" S 39º 54' 59.91" W, 156m (2 ind); Amb14 C2, 20º 11' 25.75" S 40º 2' 15.87" W, 33m (10 ind); Amb14 E2, 19º 18' 8.36" S 39º 23' 24.69" W, 33m (3 ind); Amb14 F2, 18º 52' 32.42" S 39º 8' 41.41" W, 33m (22 ind); Amb14 F3, 18º 53' 33.65" S 39º 6' 20.73" W, 43m (17 ind).

Diagnostic features: First and last pair of branchiae cirriform, remaining branchiae flattened; dorsal crests low from chaetiger 12 to chaetigers 15–18.

Description. A large-sized Prionospio, largest complete specimen 12 mm long, 0.45 mm wide at the widest point for 94 chaetigers, holotype 10 mm long, 0.4 mm wide at the widest point for 74 chaetigers (incomplete). Body dorsoventrally flattened on the branchial region and cylindrical afterwards, tapering towards the pygidium. Body color whitish in alcohol (Fig. 11).

Prostomium rounded, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2. U-shaped nuchal organs reaching posterior margin of chaetiger 1 (Fig. 12A). One pair of small eyes or eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, especially on the notopodium, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae auricular and shifted dorsally on the notopodium, and rounded on the neuropodium, both much smaller than lamellae on succeeding chaetigers (Figs. 12A; 13A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–15 (Fig. 13A–D), rounded afterwards and drastically reducing in size, present as a low flap on the posterior region. Notopodial prechaetal lamellae small and rounded from chaetiger 2 to chaetigers 15–21 (depending on specimen size). Dorsal crests low from chaetiger 12 to chaetigers 15–18, incomplete crests for about 10 more chaetigers (Fig. 12A).

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetiger 20 (Fig. 13A–D), abruptly reduced to a low flap afterwards. Neuropodial prechaetal small and rounded from chaetiger 2 to chaetiger 15.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate and granulated capillaries (Fig. 14A–B). Chaetae from the posterior row up to 1.5 times longer than chaetae from the anterior row; neuropodial chaetae slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non- granulated, thinner and less numerous (Fig. 14C).

Hooks in notopodia starting from chaetigers 21–41, up to four per fascicle, accompanied by 1–5 short non-limbate capillaries (Fig. 14D–E). Hooks in neuropodia starting from chaetigers 12–19, up to eight hooks per fascicle, accompanied, by 1–7 short non-limbate capillaries. Hooks multidentate, with 14 secondary teeth organized in two rows above the main tooth (Figs. 12C; 14D). Rudimentary secondary hood present (Fig. 14D). Sabre chaetae starting from chaetigers 10–14. Sabre chaetae unilimbate, with sparsely light granulations along the shaft (Fig. 14F).

Up to 10 pairs of smooth branchiae, cirriform on the first and last chaetigers (rarely cirriform also on the penultimate chaetiger) and robust and flattened on the remaining chaetigers (Figs. 12A–B; 15A–B). Branchiae starting from chaetiger 2, up to two times longer than notopodial lamellae, not significantly reduced in length towards the last branchial pair. First and last pair of branchiae sparsely ciliated, remaining pairs of branchiae densely ciliated throughout the length (except the tip). All branchiae completely free from notopodial postchaetal lamellae (Fig. 12A–B).

Pygidium bearing a single long dorsal cirrus and two short ventro-lateral lobes (Fig. 15C).

Oocytes from chaetiger 10, measuring up to 150 µm.

Variation: In specimens with less than 10 pairs of branchiae, the last pair is always cirriform, regardless of the occurring chaetiger.

In juveniles, the ventral pygidial lobes were not observed and the pygidium possessed a single dorsal cirrus.

Methyl green staining pattern: anterior margin of the prostomium, dorsal and lateral sides of the peristomium and ventral midline intensely stained, postchaetal lamellae from the branchial region slightly stained.

Remarks. Prionospio kinbergi sp. nov. is unusual in having a combination of cirriform and flattened branchiae, a character observed only in P. cirrifera, P. elongata Imajima, 1990b, described from Japan and P. aluta Maciolek, 1985, described from Massachusetts (USA).

All four species share a similar hook morphology (seven pairs of secondary teeth in P. kinbergi sp. nov., six to seven pairs of secondary teeth in P. elongata, five to six pairs in P. cirrifera and six pairs in P. aluta) and an overlap in the starting chaetiger of neuropodial hooded hooks (chaetigers 10–20 in P. kinbergi sp. nov., chaetiger 19 in P. elongata, chaetigers 10–17 in P. cirrifera and chaetigers 14–17 in P. aluta).

However, species can be separated based on the starting chaetiger of sabre chaetae (chaetigers 10–14 in P. kinbergi sp. nov. and P. aluta, chaetiger 16 in P. elongata and chaetiger 10 in P. cirrifera), distribution of dorsal crests (from chaetiger 12 to chaetigers 15–18 in P. kinbergi sp. nov., from chaetiger 14 to several chaetigers [distribution not mentioned] in P. elongata and from chaetiger 10 almost the end of the body in P. cirrifera; distribution not clearly mentioned in P. aluta).

Species can also be distinguished based on the shape of postchaetal lamellae from chaetiger 1 (auricular on the notopodium and digitiform on the neuropodium in P. kinbergi sp. nov., foliaceous on both rami in P. elongata, triangular on the notopodium and rounded on the neuropodium in P. cirrifera and rounded on both rami in P. aluta),branchial distribution (10 pairs in P. kinbergi sp. nov., 13 pairs in P. elongata, six to eight pairs in P. cirrifera [Mackie, 1984; Maciolek, 1985] and six pairs in P. aluta) and branchial morphology (first and last pairs cirriform in P. kinbergi sp. nov. and P. cirrifera, first and last four pairs cirriform in P. elongata and only first pair cirriform in P. aluta).

Apart from the characters discussed above, P. kinbergi sp. nov. can be readily distinguished from P. aluta due to the lack of interparapodial pouches, a character rarely observed among Prionospio species.

Prionospio kinbergi sp. nov. is similar to P. acutus sp. nov. and P. mutatus sp. nov. in having robust flattened branchiae, although these species bear only robust branchiae and not a combination of cirriform and flattened branchiae, as in P. kinbergi sp. nov. All three species share a similar distribution of dorsal crests (from chaetigers 10–11 to chaetigers 15–17 in P. acutus sp. nov., from chaetiger 10 to chaetigers 18–20 in P. mutatus sp. nov. and from chaetiger 12 to chaetigers 15–18 in P. kinbergi sp. nov.), although P. kinbergi sp. nov. can be distinguished by the prostomial shape (rounded in P. mutatus sp. nov. and P. kinbergi sp. nov. and triangular in P. acutus sp. nov.), the presence of sabre chaetae appearing abruptly (absent in P. acutus sp. nov. and appearing gradually in P. mutatus sp. nov.) and presence of incomplete crests after the complete dorsal crests, from chaetiger 25 to chaetiger 28.

In specimens of Prionospio kinbergi sp. nov. with less than 10 pairs of branchiae, the last pair is always cirriform, regardless of the occurring chaetiger. This hints to a gradual branchial transformation from cirriform to tapered, as the penultimate chaetigers sometimes is also cirriform, but more robust than the last branchial pair.

According to Radashevsky (2012), in the majority of species from the Prionospio-complex, the number of branchiae is constant and species-specific, increasing only to a certain point, when new branchial pairs will not develop regardless of specimen growth. The sequence of branchial development, however, is not clear, since branchiae from chaetiger 2 remain cirriform in adults, while the remaining branchiae gradually transform into flattened branchiae – except for the last branchial pair, which remains cirriform in adults.

Etymology. The specific epithet, kinbergi, is a tribute to Johan G. H. Kinberg (1820–1908), who described the first Brazilian spionid, Laonice brevicornis Kinberg, 1866.

Habitat: medium sand to fine sand, 19–163 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Prionospio hartmanae sp. nov.

(Figures 16–20)

Prionospio cirrifera: Pardo et al. (2006). Not Wirén, 1883.

Prionospio multibranchiata: Pardo & Peixoto (in press). Not Berkeley, 1927.

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 02 Dec 2011 to 02 Feb 2012, 34m, MZUSP XXXX. Paratypes: Amb1 Foz7, 19º 49' 54.12" S 39º 52' 11.79" W, 11 Dec 2010 to 19 Dec 2010, 29m, MZUSP XXXX (4 ind); Amb2 Foz11,

19º 57' 30.39" S 39º 53' 35.28" W, 12 Jul 2011 to 18 Jul 2011, 46m, ZUEC-POL XXXX (3 ind); Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 02 Dec 2011 to 02 Feb 2012, 34m, MNRJP XXXX (3 ind).

Additional material examined. Amb1 Foz7, 19º 49' 54.12" S 39º 52' 11.79" W, 29m (5 ind); Amb1 Foz11, 19º 57' 34.65" S 39º 53' 26.1" W, 43m (2ind); Amb1 Foz14, 19º 42' 32.21" S 39º 38' 57.36" W, 35m (2 ind); Amb1 Foz16, 20º 1' 3.73" S 39º 50' 13.76" W, 48m (2 ind); Amb2 Foz7, 19º 49' 50.65" S 39º 52' 22.92" W, 28m (4 ind); Amb2 Foz8, 19º 44' 40.28" S 39º 46' 32.55" W, 29m (2 ind); Amb2 Foz11, 19º 57' 30.39" S 39º 53' 35.28" W, 46m (3 ind); Amb2 Foz14, 19º 42' 28.53" S 39º 39' 4.25" W, 36m (2 ind); Amb2 Foz15, 19º 37' 41.83" S 39º 35' 31.52" W, 35m (1 ind); Amb2 Foz17, 19º 55' 43.7" S 39º 45' 39.68" W, 43m (1 ind); Amb2 Foz18, 19º 50' 16.34" S 39º 40' 10.8" W, 46m (1 ind); Amb7 A2, 21º 3' 26.62" S 40º 23' 0.67" W, 36 m (2 ind); Amb7 B3, 20º 34' 53.69" S 40º 6' 27.35" W, 45m (3 ind); Amb7 C2, 20º 11' 25.35" S 40º 2' 16.02" W, 35m (4 ind); Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 34m (7 ind); Amb7 F3, 18º 53' 29.72" S 39º 6' 23.3" W, 43m (4 ind); Amb14 B3, 20º 34' 53.05" S 40º 6' 27.68" W, 43m (2 ind); Amb14 F2, 18º 52' 32.42" S 39º 8' 41.41" W, 33m (1 ind).

Diagnostic features: Up to 11 pairs of cirriform branchiae, lack of prostomial peaks and dorsal crests, sabre chaetae from chaetigers 13–14 and ventrally pointed postchaetal neuropodial lamella on chaetiger 2.

Description. A small-sized Prionospio, largest complete specimen 6.5 mm long, 0.3 mm wide at the widest point for 57 chaetigers, holotype complete, 6 mm long, 0.3 mm wide at the widest point for 50 chaetigers. Body widest and cylindrical on the branchial region, slightly dorsoventrally flattened afterwards, tapering towards the pygidium. Body color whitish in alcohol (Fig. 16).

Prostomium narrow, rounded anteriorly, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2 (Fig. 17A). Prostomial peaks absent. One or two small eye pairs usually present or eyes absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lacking lateral wings. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded with a pointed tip on the notopodium and digitiform on the neuropodium (Figs. 17B; 18A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–13 (Fig. 18A–D), rounded from chaetiger 14 and gradually reducing in size towards the posterior region, present as a low flap on the posterior region, slightly more developed and triangular on the last 3–8 chaetigers. Notopodial prechaetal lamellae absent throughout. Dorsal crests absent (Fig. 17A).

Neuropodial postchaetal lamellae triangular with pointed edges and ventrally pointed tip on chaetiger 2 (Figs. 17B; 18B). Lamellae elliptical from chaetigers 3–8 (Fig. 18A–D), rounded from chaetigers 9–20, present as a low flap on the posterior chaetigers. Neuropodial prechaetal lamellae absent throughout (Fig. 17B).

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbated capillaries, bearing light granulations along the shaft (Fig. 19A). Chaetae from both rows of equal length, although neuropodial chaetae are slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig. 19B).

Hooks in notopodia from chaetigers 23–41, up to four per fascicle, accompanied by 1–4 short non-limbate capillaries (Figs. 17C; 19C–D). Hooks in neuropodia starting from chaetigers 17–22, up to five per fascicle, accompanied by 1–4 short non-limbate capillaries. Hooks multidentate, with 10 secondary teeth organized in two rows above the main tooth (Figs. 17C; 19C). Small secondary hood present. Sabre chaetae starting from chaetigers 13–14 (usually chaetiger 13). Sabre chaetae non-limbate, with granulations along the upper part of the shaft (Fig. 19E).

Up to 11 pairs of smooth and cirriform branchiae, tapered at the tips. Branchiae starting from chaetiger 2, up to three times longer than notopodial lamellae, slightly reducing in length towards last branchial pair. Branchiae densely ciliated throughout the length (except the tip), completely free from notopodial postchaetal lamellae (Figs. 17A–B; 20A).

Pygidium bearing one dorsal cirrus and two short ventro-lateral lobes (Fig. 20B).

Oocytes from chaetiger 15, measuring up to 90 µm.

Methyl green pattern: Prostomium, dorsal side of the peristomium and margins of lamellae up to chaetiger 20 intensely stained. Stain diffused on the ventral side up to chaetiger 15.

Remarks. Prionospio hartmanae sp. nov. shares similarities with P. fosterae sp. nov., P. perkinsi and P. lighti in having a rounded prostomium, similar branchial number and morphology (up to 11 pairs in P. hartmanae sp. nov., up to 10 pairs in P. perkinsi and up to 12 pairs in P. lighti, apinnate and cirriform in all species) and lack of dorsal crests, although species can be separated based on the lack of prostomial peaks in P. hartmanae sp. nov. (present in P. lighti and P. perkinsi), presence of sabre chaetae (absent in P. fosterae and P. perkinsi), number of secondary teeth on the hooded hooks (five pairs in P. hartmanae sp. nov., one pair in P. fosterae sp. nov., three pairs in P. lighti and two pairs in P. perkinsi) and morphology of the postchaetal lamellae from chaetiger 1 (rounded with a pointed tip on the notopodium and digitiform on the neuropodium in P. hartmanae sp. nov., digitiform on the notopodium and rounded on the neuropodium in P. fosterae sp. nov., rounded on the notopodium and digitiform on the

81 neuropodium in P. perkinsi and absent on the notopodium and digitiform on the neuropodium in P. lighti).

Prionospio hartmanae sp. nov. is most similar to P. multibranchiata, a species originally described from the British Columbia, Canada, Pacific Ocean, although currently regarded as a cosmopolitan species (as discussed by Mackie, 1994 and Maciolek, 1985), with documented Brazilian records (São Paulo State) (Amaral et al. 2013). Prionospio multibranchiata was redescribed by Mackie (1984), based on material from Scotland, England and Sweden and by Maciolek (1985), based on material from Mexico, Florida (USA) and Washington (USA), the latter being the closest to the type-locality.

Based on Mackie’s (1984) redescription, both species share a rounded prostomium lacking prostomial peaks, long cirriform branchiae, overlap in the starting chaetiger of sabre chaetae (chaetigers 13–14 in P. hartmanae sp. nov. and chaetigers 12–17 in P. multibranchiata), considerable overlap in the starting chaetiger of notopodial hooded hooks (chaetigers 23–41 in P. hartmanae sp. nov. and chaetigers 20–52 in P. multibranchiata) and hooks of similar morphology (five pairs of secondary teeth in P. hartmanae sp. nov. and four to five pairs in P. multibranchiata), but species can be separated based on the branchial distribution (up to 11 pairs in P. hartmanae sp. nov. and up to 13 pairs in P. multibranchiata), morphology of postchaetal lamellae from chaetiger 1 (rounded with a pointed tip on the notopodium and digitiform on the neuropodium in P. hartmanae sp. nov. and rounded on both rami in P. multibranchiata) and lack of dorsal crests in P. hartmanae sp. nov., which are present from chaetigers 13–15 on a variable number of chaetigers in P. multibranchiata. Some specimens examined by Mackie (1984) also exhibited interparapodial pouches, absent in P. hartmanae sp. nov.

When compared with Maciolek’s redescription (1985), species share a similar prostomium morphology (narrow and rounded anteriorly), lack of prostomial peaks, same branchial number and morphology (up to 11 pairs of apinnate cirriform branchiae) and a considerable overlap in the starting chaetiger of notopodial hooded hooks (chaetigers 23–41 in P. hartmanae sp. nov. and chaetigers 26–34 in P. multibranchiata) and sabre chaetae (chaetigers 13–14 in P. hartmanae sp. nov. and chaetigers 10–16 in P. multibranchiata), but species can be separated based on the number of secondary teeth on the hooded hooks (five pairs of secondary teeth in P. hartmanae sp. nov. and three pairs in P. multibranchiata), shape of the postchaetal neuropodial lamellae from chaetiger 1 (digitiform in P. hartmanae sp. nov. and rounded in P. multibranchiata), lack of dorsal crests in P. hartmanae sp. nov. (present on several postbranchial chaetigers in P. multibranchiata) and lack of notopodial postchaetal lamellae from chaetiger 1 in P. multibranchiata (rounded with a pointed tip in P. hartmanae sp. nov.). Prionospio hartmanae sp.

82 nov. also possess a ventrally pointed neuropodial lamellae on chaetiger 2, which is absent in P. multibranchiata.

The ventrally pointed postchaetal neuropodial lamella on chaetiger 2 is similar to the lamellae reported in P. cirrifera (Mackie, 1984) and P. aluta (Maciolek, 1985). However, P. hartmanae sp. nov. can be readily distinguished from these species due to the lack of interparapodial pouches present from chaetigers 4–5 to chaetigers 37–38 in P. aluta), lack of dorsal crests (present from chaetigers 10–11 to almost the end of the body in P. cirrifera and present of several postbranchial chaetigers in P. aluta) and by the branchial morphology (up to 11 pairs of apinnate and cirriform branchiae in P. hartmanae sp. nov., up to eight pairs of apinnate and cirriform branchiae in P. cirrifera and six pairs of robust subtriangular branchiae in P. aluta). According to Delgado-Blas & Salazar-Silva (2011), both P. cirrifera and P. aluta belong to a species-complex that includes species bearing a ventrally pointed neuropodial postchaetal lamellae on chaetiger 2, which would include P. hartmanae sp. nov.

Etymology. The specific epithet, hartmanae, is a tribute to Olga Hartman (1900–1974), an invertebrate zoologist who made invaluable contributions to the study of polychaetes.

Habitat: Gravel sand to fine sand, 28–48 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Prionospio mutatus sp. nov.

(Figures 21–25)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 B6, 20º 36' 2.03" S 39º 51' 35.37" W, 02 Dec 2011 to 02 Feb 2012, 991m, MZUSP XXXX. Paratypes: Amb5 B6, 20º 36' 2.03" S 39º 51' 35.37" W, 02 Dec 2011 to 02 Feb 2012, 991m, MZUSP XXXX (2 ind), ZUEC-POL (1 ind); Amb12 E7, 19º 47' 2.44" S 39º 3' 14.62" W, 06 Jun 2013 to 17 Jul 2013, 1242m, MNRJP (2 ind).

Additional material examined. Amb3 E5, 19º 36' 25.08" S 39º 10' 20.15" W, 352m (2 ind); Amb3 E6, 19º 40' 5.6" S 39º 7' 15.87" W, 1010m (12 ind); Amb4 F7, 20º 4' 8.18" S 38º 31' 27.32" W, 1288m (2 ind); Amb5 B5, 20º 35' 15.16" S 39º 53' 46.36" W, 382m (1 ind); Amb5 B6, 20º 36' 2.03" S 39º 51' 35.37" W, 991m (24 ind); Amb5 B7, 20º 36' 41.55" S 39º 49' 24.15" W, 1315m (2 ind); Amb5 C5, 20º 14' 22.9" S 39º 48' 33.06" W, 416m (5 ind); Amb5 C6, 20º 15' 34.21" S 39º 46' 15.69" W, 1031m (2 ind); Amb5 C7, 20º 17' 35.61" S 39º 42' 35.67" W, 1309m (1 ind); Amb5 D6, 19º 50' 1.87" S 39º 26' 30.04" W, 1053m (1 ind); Amb5 D8, 20º 8' 43.27" S 39º 7' 22.23" W, 1905m (1 ind); Amb11 B6, 20º 35' 57.27" S 39º 51' 38.12" W, 994m (4 ind); Amb11

B7, 20º 36' 49.74" S 39º 49' 29.01" W, 1327m (6 ind); Amb11 C5, 20º 14' 19.79" S 39º 48' 36.57" W, 418m (6 ind); Amb11 C7, 20º 17' 37.38" S 39º 42' 36.72" W, 1347m (1 ind); Amb12 A5, 21º 4' 8.5" S 40º 13' 7.35" W, 383m (1 ind); Amb12 D7, 19º 54' 4.77" S 39º 22' 29.46" W, 1330m (1 ind); Amb12 E6, 19º 40' 1.68" S 39º 7' 23.79" W, 1018m (1 ind); Amb12 E7, 19º 47' 2.44" S 39º 3' 14.62" W, 1242m (11 ind); Amb11 E8, 20º 15' 53.31" S 38º 40' 53.51" W, 1886m (1 ind); Amb12 F5, 19º 34' 20.47" S 38º 41' 19.8" W, 445m (1 ind); Amb12 G7, 19º 3' 31.11" S 37º 48' 42.65" W, 1361m (1 ind); Amb12 CANWN5, 19º 49' 36.4" S 39º 35' 41.77" W, 363m (1 ind).

Diagnostic features: Dorsal crests low from chaetiger 10 to chaetigers 18–20; sabre chaetae appearing as a gradual transformation of capillary chaeta.

Description. A small-sized Prionospio, largest complete specimen 2 mm long, 0.15 mm wide at the widest point for 34 chaetigers, holotype 5 mm long, 0.2 mm wide at the widest point for 45 chaetigers (incomplete). Body dorsoventrally flattened on the branchial region and cylindrical afterwards, tapering towards the pygidium. Body color whitish in alcohol (Fig. 21).

Prostomium elongated, rounded anteriorly, not widened towards the anterior margin and extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2. U-shaped nuchal organs reaching the anterior margin of chaetiger 2 (Fig. 22A). Eyes absent. Prostomial peaks generally present in adults (Fig. 21). Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent. Grooved palps reaching up to chaetiger 8, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, especially on the notopodium, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on the notopodium and digitiform on the neuropodium, both smaller than lamellae on succeeding chaetigers (Fig. 23A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–10 (Fig. 23A–D), square-shaped with a pointed tip on chaetigers 11–12 and broadly round starting on chaetiger 13, reduced to a low flap after chaetigers 22–25 (depending on specimen size). Notopodial prechaetal lamellae absent throughout. Dorsal crests low from chaetiger 10 to chaetigers 18–20 (Fig. 22A, C).

Neuropodial postchaetal lamellae broadly rounded on chaetigers 2–11(Fig. 23A–D), elliptical from chaetiger 12 to chaetigers 20–23 (depending on specimen size) and drastically reduced afterwards, present as a low flap. Neuropodial prechaetal lamellae absent throughout (Fig. 22A– B).

Chaetae from notopodia and neuropodia organized in two rows of unilimbate (narrow limbation) and sparsely granulated capillaries (Fig. 24A). Chaetae from both rows of equal length, although neuropodial chaetae are slightly shorter than notopodial chaetae. Towards the posterior region,

84 capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig. 24B).

Hooks in notopodia starting from chaetigers 26–34, up to three per fascicle, accompanied by 1–4 short non-limbate capillaries (Fig. 24C–D). Hooks in neuropodia starting from chaetigers 13–19, up to seven per fascicle, accompanied by 2–6 short non-limbate capillaries. Hooks multidentate, with 10 secondary teeth organized in two rows above the main tooth (Fig. 24C). Secondary hood present (Fig. 24C). Sabre chaetae starting from chaetigers 12–17, appearing as a gradual transformation of the inferior-most capillary chaeta from 2–3 previous chaetigers. Sabre chaetae non-limbate, with dense light granulations along the shaft (Fig. 24E–G).

Up to 11 pairs of smooth, robust and cirriform branchiae, tapered at the tips. Branchiae starting from chaetiger 2, up to two times longer than notopodial lamellae, gradually reducing in length towards last branchial pair. Branchiae densely ciliated throughout the length (except the tip), completely free from notopodial postchaetal lamellae (Figs. 22A–C; 25A).

Pygidium bearing one dorsal cirrus and two short digitiform ventro-lateral lobes (Fig. 25B).

Oocytes from chaetiger 12, measuring up to 100 µm.

Methyl green staining pattern: Prostomium, dorsal and lateral sides of the peristomium intensely stained, postchaetal lamellae from chaetigers 2–5 slightly stained.

Remarks. The gradual transformation of capillary chaetae into sabre chaetae had never been reported in Prionospio, as, according to Radashevsky (2012), it is known only in Microspio Mesnil, 1896 and Spio Fabricius, 1785, genera that belong to Spioninae Söderström, 1920 a different subfamily. Lack of sabre chaetae have been reported only in a few species – Prionospio perkinsi, P. hermesia, P. cerastae, P. fosterae sp. nov. and P. acutus sp. nov., while it is present and appearing abruptly in the remaining Prionospio species.

Prionospio mutatus sp. nov. can be readily distinguished from P. cerastae and P. hermesia by branchial number and morphology – 11 pairs of cirriform branchiae in P. mutatus sp. nov., two pairs of cirriform branchiae in P. hermesia and four pair of branchiae, the last pair bearing pinnules in P. cerastae, and presence and distribution of dorsal crests - from chaetiger 10 to chaetigers 18–20 in P. mutatus sp. nov., from chaetiger 8 to chaetiger 25 in P. hermesia and lacking in P. cerastae.

The presence of up to 11 pairs of cirriform branchiae is reported in Prionospio mutatus sp. nov., P. fosterae sp. nov. and P. hartmanae sp. nov., although species can be distinguished based on the presence of dorsal crests from chaetiger 10 to chaetigers 18–20 in P. mutatus sp. nov., shape of lamellae on chaetiger 1 (rounded on the notopodium and digitiform on the neuropodium of P.

85 mutatus sp. nov., rounded on both rami in P. fosterae sp. nov. and rounded with a pointed tip on the notopodium and digitiform on the neuropodium of P. hartmanae sp. nov.) and sabre chaetae, lacking in P. fosterae sp. nov., appearing abruptly in P. hartmanae sp. nov. and appearing gradually in P. mutatus sp. nov.

Prionospio mutatus sp. nov.is similar to P. fosterae sp. nov., P. hartmanae sp. nov. and P. kinbergi sp. nov. in having a rounded prostomium, but P. mutatus sp. nov. can be separated from these species based on the presence of prostomial peaks, presence and distribution of dorsal crests (absent in P. fosterae sp. nov. and P. hartmanae sp. nov., from chaetiger 10 to chaetigers 18–20 in P. mutatus sp. nov. and from chaetiger 12 to chaetigers 15–18 in P. kinbergi sp. nov.) and branchial number and morphology (up to 11 pairs of cirriform branchiae in P. fosterae sp. nov. and P. hartmanae sp. nov. and up to 10 pairs of branchiae, cirriform on the first and last pairs and flattened on the remaining pairs in P. kinbergi sp. nov.).

Etymology. The specific epithet, mutatus, refers to the gradual transformation of the inferior- most capillary chaeta into sabre chaetae (mutatus, Latin for change; alteration).

Habitat: mud to silt, 352–1905 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Prionospio cinthyae sp. nov.

(Figures 26–30)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 E3, 19º 26' 5" S 39º 17' 38.92" W, 02 Dec 2011 to 02 Feb 2012, 26m, MZUSP XXXX. Paratypes: Amb1 Foz15, 19º 37' 48.27" S 39º 35' 25.83" W, 11 Dec 2010 to 19 Dec 2010, 41m, MZUSP (2 ind), ZUEC-POL (3 ind); Amb7 E3, 19º 26' 5" S 39º 17' 38.92" W, 02 Dec 2011 to 02 Feb 2012, 26m, MZUSP (3 ind), ZUEC- POL (2 ind), MNRJP (4 ind).

Additional material examined. Amb1 Foz1, 19º 52' 22.74" S 39º 59' 38.05" W, 28m (1 ind); Amb1 Foz2, 19º 47' 22.34" S 39º 55' 6.96" W, 24m (1 ind); Amb1 Foz5, 19º 32' 32.82" S 39º 41' 34.89" W, 20m (1 ind); Amb1 Foz7, 19º 49' 57.38" S 39º 52' 14.02" W, 33m (15 ind); Amb1 Foz8, 19º 44' 45.48" S 39º 46' 25.78" W, 32m (2 ind); Amb1 Foz10, 19º 35' 10.73" S 39º 38' 35.79" W, 32m (2 ind); Amb1 Foz11, 19º 57' 32.89" S 39º 53' 30.69" W, 47m (4 ind); Amb1 Foz12, 19º 52' 35.48" S 39º 49' 5.63" W, 43m (7 ind); Amb1 Foz15, 19º 37' 48.27" S 39º 35' 25.83" W, 41m (23 ind); Amb1 Foz20, 19º 41' 33.92" S 39º 31' 17.74" W, 54m (5 ind); Amb2 Foz1, 19º 52' 14.57" S 39º 59' 41.06" W, 27m (8 ind); Amb2 Foz3, 19º 42' 8.87" S 39º 49' 29.72" W, 16m (1 ind); Amb2 Foz7, 19º 49' 52.15" S 39º 52' 24.51" W, 30m (5 ind); Amb2 Foz10, 19º

35' 2.68" S 39º 38' 40.25" W, 31m (4 ind); Amb2 Foz12, 19º 52' 29.19" S 39º 49' 12.78" W, 45m (3 ind); Amb2 Foz20, 19º 41' 26.47" S 39º 31' 19.38" W, 53m (3 ind); Amb2 Foz20b, 19º 41' 26.14" S 39º 31' 20.62" W, 53m (1 ind); Amb7 A2, 21º 3' 27.14" S 40º 22' 59.61" W, 40m (1 ind); Amb7 B1, 20º 34' 32.47" S 40º 20' 52.37" W, 25m (20 ind); Amb7 D1, 19º 35' 37.21" S 39º 41' 19.68" W, 25m (13 ind); Amb7 D3, 19º 43' 14.34" S 39º 33' 34.86" W, 50m (27 ind); Amb7 E2, 19º 18' 5.9" S 39º 23' 23.3" W, 39m (6 ind); Amb7 E3, 19º 26' 5" S 39º 17' 38.92" W, 26m (138 ind); Amb7 F2, 18º 52' 32.61" S 39º 8' 42.82" W, 40m (1 ind); Amb14 B1, 20º 34' 29.6" S 40º 20' 54.56" W, 26m (1 ind); Amb14 E2, 19º 18' 6.12" S 39º 23' 23.35" W, 38m (4 ind); Amb14 E3, 19º 26' 4.81" S 39º 17' 38.64" W, 50m (8 ind); Amb12 E4, 19º 36' 4,74" S 39º 10' 34,59" W, 150m (3 ind).

Diagnostic features: Four pairs of apinnate branchiae, first and last pairs wrinkled, first pair up to three times longer than last branchial pair, dorsal crests from chaetiger 6 to chaetigers 7–9.

Description. A medium-sized Prionospio, largest complete specimen about 7.5 mm long, 0.25 mm wide at the widest point for 55 chaetigers, holotype complete, 7.5 mm long, 0.25 mm wide at the widest point for 54 chaetigers. Body dorsoventrally flattened throughout, tapering towards the pygidium. Body color light yellow to whitish in alcohol (Fig. 26A–B).

Prostomium triangular, rounded anteriorly, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2. Nuchal organs reaching anterior margin of chaetiger 2. (Fig. 27A– C). A single pair of minute blackish eyes generally present or eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium, partially fused to the first chaetiger. Lateral wings absent. Grooved palps reaching up to chaetiger 10, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae square-shaped on the notopodium and rounded on the neuropodium, slightly reduced (Fig. 28A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 7–8 (largest on chaetigers 3–5) (Fig. 28A–D), rounded afterwards and gradually reducing in size towards the posterior region, present as a low flap on the last chaetigers, Notopodial prechaetal lamellae absent throughout. Dorsal crests from chaetiger 6 to chaetigers 7–9 (usually chaetiger 7), high on chaetiger 7 and low on the remaining chaetigers.

Neuropodial postchaetal lamellae square-shaped on chaetigers 2 and 3 and rounded from chaetiger 4 afterwards (Fig. 28A–D), gradually reducing in size towards the posterior region, present as a low flap on the last chaetigers. Neuropodial prechaetal lamellae absent throughout (Figs. 26B; 27A).

Chaetae from notopodia and neuropodia organized in two rows of unilimbate (narrow limbation) and sparsely granulated capillaries (Fig. 29A–C). Chaetae from both rows of equal length in notopodia, while in neuropodia the posterior row of chaetae is up to 1.5 times longer than anterior row (Fig. 29B–C). Neuropodial chaetae slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated and less numerous (Fig. 29D).

Hooks in notopodia starting from chaetigers 21–33, up to three per fascicle, accompanied by 1–4 short non-limbate capillaries (Fig 29. E–F). Hooks in neuropodia starting from chaetigers 10–12 (generally chaetiger 10), up to five per fascicle, accompanied by 1–5 non-limbate capillaries. Hooks multidentate, with 3 pairs of teeth arranged side by side above the main tooth (Fig. 29E). Secondary hood absent. Sabre chaetae consistently from chaetiger 10. Sabre chaetae non-limbate, with sparsely light granulations along the shaft (Fig. 29G).

Four pairs of apinnate branchiae, subtriangular, robust and flattened on chaetigers three and four and long, cirriform and sculptured (“wrinkled”) on chaetigers two and five, all longer than notopodial lamellae and tapered at the tips. Branchiae from chaetiger two up to eight times longer than notopodial lamellae and up to three times longer than branchiae from chaetiger five. All branchiae completely free from notopodial postchaetal lamellae (Fig. 30A–B).

Pygidium bearing a single long dorsal cirrus and two short and robust ventro-lateral lobes (Fig. 30C).

Oocytes from chaetiger 10, measuring up to 100 μm.

Methyl green staining pattern: Light green pigment diffused throughout the body, margins of the prostomium, peristomium, ventral side pf the peristomium and chaetiger 1 intensely stained.

Remarks. Prionospio cinthyae sp. nov. differs from most apinnate species – and, particularly, from all apinnate species recorded in Brazil in having only four pairs of branchiae. While this pattern is common among pinnate Prionospio species, it is somewhat rare among apinnate species, being observed only in P. laciniosa Maciolek, 1985, described from Angola, P. fauchaldi Maciolek, 1985, P. elegantula Imajima, 1990a and P. japonicus Okuda, 1935.

Prionospio cinthyae sp. nov. can be easily distinguished from P. elegantula based on branchial morphology, as the latter species bear extremely long, thin and smooth branchiae on chaetigers 2–5, while P. cinthyae sp. nov. bears wrinkled branchiae on chaetigers 2 and 5 and flattened branchiae on chaetigers 3 and 4.

Prionospio cinthyae and P. japonicus are similar in having a triangular prostomium and long branchiae on chaetiger 2, but species can be distinguished by the lack of dorsal crests and presence

88 of prostomial peaks in P. japonicus and by branchial morphology, as P. japonicus possesses a long, cirriform branchiae on chaetiger 2 and shorter branchiae on the remaining chaetigers, while P. cinthyae sp. nov. possess wrinkled branchiae on chaetigers 2 and 5 and flattened branchiae on chaetigers 3 and 4.

Only five additional Prionospio species bear wrinkled branchiae: P. fauchaldi, P. laciniosa, P. sandersi Maciolek, 1981, P. vallensis Neal & Paterson in Paterson et al., 2016, described from Portugal and P. corrugatus sp. nov. Although Prionospio sandersi and P. corrugatus sp. nov. also bear wrinkled branchiae, both can be separated from P. cinthyae sp. nov. based on the branchial number, always four pairs in P. cinthyae sp. nov., up to six pairs in P. corrugatus sp. nov. and up to nine pairs in P. sandersi and by the prostomium morphology, triangular in P. cinthyae sp. nov., rectangular in P. corrugatus sp. nov. and rounded in P. sandersi.

Prionospio cinthyae sp. nov. is most similar to P. fauchaldi, P. vallensis and P. laciniosa in having branchiae on chaetigers 2–5, a combination of smooth and wrinkled apinnate branchiae and long branchiae in chaetiger 2 (present in P. cinthyae sp. nov., P. vallensis and P. laciniosa). However, Prionospio cinthyae sp. nov. can be distinguished from the remaining species based on the presence of robust sabre chaetae (delicate in P. fauchaldi), unmodified dorsal crests on chaetigers 6–9 (present from chaetigers 8–11 in P. fauchaldi, from chaetiger 4 to beyond chaetiger 20 in P. vallensis and modified into semicircular flaps on chaetigers 5–13 in P. laciniosa), hook morphology (three pairs of secondary teeth in Prionospio cinthyae sp. nov., four pairs in P. fauchaldi and six pairs in P. laciniosa and P. vallensis) and by the prostomium morphology (triangular in P. cinthyae sp. nov. and P. laciniosa, rectangular in P. fauchaldi and P. vallensis).

Etymology. The specific epithet, cinthyae, is a homage to Dr. Cinthya S. G. Santos, a polychaete researcher and former advisor of the first author.

Habitat: Gravel sand to mud, 16–150 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Prionospio corrugatus sp. nov.

(Figures 31–35)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 B5, 20º 35' 15.16" S 39º 53' 46.36" W, 02 Dec 2011 to 02 Feb 2012, 991m, MZUSP XXXX. Paratypes: Amb11 C5, 20º 14' 19.79" S 39º 48' 36.57" W, 06 Jul 2013 to 17 Jul 2013, 418m, MNRJP XXXX (5 ind); Amb5 B6, 20º 36' 2.03" S 39º 51' 35.37" W, 02 Dec 2011 to 02 Feb 2012, 991m, MZUSP XXXX (4 ind); Amb12

D6, 19º 50' 6.01" S 39º 26' 34.62" W, 02 Dec 2011 to 02 Feb 2012, 1048m, MZUSP XXXX (2 ind), ZUEC-POL XXXX (5 ind), MNRJP (3 ind).

Additional material examined. Amb3 E5, 19º 36' 25.08" S 39º 10' 20.15" W, 352m (19 ind); Amb3 E6, 19º 40' 13.13" S 39º 7' 20.92" W, 1010m (27 ind); Amb3 F5, 19º 34' 21.01" S 38º 41' 16.99" W, 438m (1 ind); Amb3 CAND4, 19º 31' 51.13" S 39º 3' 4.99" W, 140m (1 ind); Amb3 CAND5, 19º 33' 20.99" S 39º 2' 36.2" W, 384m (34 ind); Amb3 CAND7, 19º 42' 20.31" S 39º 5' 55.46" W, 1289m (1 ind); Amb4 B8, 20º 41' 33.45" S 39º 35' 14.76" W, 1902m (1 ind); Amb4 F7, 20º 4' 8.18" S 38º 31' 27.32" W, 1288m (7 ind); Amb5 A5, 21º 4' 5.57" S 40º 13' 2.63" W, 396m (49 ind); Amb5 A6, 21º 4' 43.84" S 40º 8' 31.76" W, 990m (26 ind); Amb5 A7, 21º 4' 51.67" S 40º 4' 14.88" W, 1316m (16 ind); Amb5 B5, 20º 35' 15.16" S 39º 53' 46.36" W, 382m (5 ind); Amb5 B6, 20º 36' 2.03" S 39º 51' 35.37" W, 991m (43 ind); Amb5 B7, 20º 36' 47.44" S 39º 49' 22.5" W, 1315m (18 ind); Amb5 C5, 20º 14' 19.45" S 39º 48' 36.67" W, 416m (5 ind); Amb5 C6, 20º 15' 34.21" S 39º 46' 15.69" W, 1031m (24 ind); Amb5 C7, 20º 15' 39.85" S 39º 46' 10.03" W, 1309m (15 ind); Amb5 D6, 19º 50' 4.42" S 39º 26' 33.42" W, 1053m (23 ind); Amb5 D7, 19º 54' 5.01" S 39º 22' 20.04" W, 1333m (12 ind); Amb5 D8, 20º 8' 42.11" S 39º 7' 29.06" W, 1905m (1 ind); Amb6 D5, 19º 46' 32.67" S 39º 30' 3.54" W, 402m (36 ind); Amb6 E4, 19º 36' 5.17" S 39º 10' 32.93" W, 145m (1 ind); Amb6 CANWN5, 19º 49' 38.59" S 39º 35' 44.43" W, 352m (68 ind); Amb6 CANWN6, 19º 53' 29.3" S 39º 32' 59.04" W, 955m (7 ind); Amb6 CANWN7, 19º 58' 13.68" S 39º 31' 43.28" W, 1284m (4 ind); Amb8 E7, 19º 47' 5.96" S 39º 3' 11.96" W, 1223m (4 ind); Amb8 G6, 19º 3' 32.9" S 37º 49' 4.82" W, 1011m (1 ind); Amb8 G7, 19º 3' 30.21" S 37º 48' 42.41" W, 1245m (2 ind); Amb11 A5, 21º 4' 8.5" S 40º 13' 7.35" W, 383m (43 ind); Amb11 A6, 21º 4' 43.95" S 40º 8' 34.11" W, 997m (18 ind); Amb11 A7, 21º 4' 40.63" S 40º 4' 10.65" W, 1331m (8 ind); Amb11 B5, 20º 35' 13.87" S 39º 53' 45.78" W, 382m (7 ind); Amb11 B6,20º 35' 57.27" S 39º 51' 38.12" W, 994m (29 ind); Amb11 B7, 20º 36' 49.74" S 39º 49' 29.01" W, 1327m (13 ind); Amb11 C5, 20º 14' 17.95" S 39º 48' 34.35" W, 418m (3 ind); Amb11 C6, 20º 15' 32.22" S 39º 46' 9.28" W, 1029m (17 ind); Amb11 C7, 20º 17' 39.63" S 39º 42' 38.76" W, 1347m (16 ind); Amb12 D5, 19º 46' 32.84" S 39º 30' 3.65" W, 431m (18 ind); Amb12 D6, 19º 50' 6.01" S 39º 26' 34.62" W, 1048m (34 ind); Amb12 D7, 19º 54' 5.24" S 39º 22' 28.64" W, 1330m (1 ind); Amb12 E5, 19º 36' 30.6" S 39º 10' 19.39" W, 349m (10 ind); Amb12 E6, 19º 40' 1.46" S 39º 7' 21.99" W, 1018m (20 ind); Amb12 E7, 19º 47' 2.24" S 39º 3' 13.72" W, 1242m (8 ind); Amb11 E8, 20º 15' 53.31" S 38º 40' 53.51" W, 1886m (9 ind); Amb12 G7, 19º 3' 31.11" S 37º 48' 42.65" W, 1361m (1 ind); Amb12 CAND5, 19º 33' 23.09" S 39º 2' 35.67" W, 446m (12 ind); Amb12 CAND6, 19º 37' 46.26" S 39º 3' 59.65" W, 1036m (4 ind); Amb12 CANWN5, 19º 49' 36.67" S 39º 35' 43.43" W, 363m (38 ind); Amb12 CANWN6, 19º 53' 27.27" S 39º 32' 59.82" W, 960m (7 ind); Amb12 CANWN7, 19º 58' 12.82" S 39º 31' 42.22" W, 1305m (15 ind).

Diagnostic features: Three rows of capillary chaetae on notopodia from chaetiger 2 to chaetigers 10–12; neuropodium of chaetiger 3 bearing very long chaetae on the posterior row; sculptured branchiae on chaetigers 2 and 5–7, pygidium bearing a single dorsal cirrus.

Description. A medium-sized Prionospio, largest complete specimen 6 mm long, 0.5 mm wide for 91 chaetigers, holotype complete, 5.5 mm long, 0.40 mm wide at the widest point for 74 chaetigers. Body widest and dorsoventrally flattened on the branchial region, cylindrical afterwards, tapering towards the pygidium. Body color yellow to whitish in alcohol (Fig. 31).

Prostomium rectangular, narrow, truncated on the anterior margin, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2 (Figs. 31; 32A). Small prostomial peaks might be present. Eyes absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, forming low lateral wings. Grooved palps reaching up to chaetiger 10, lost in most specimens.

Chaetiger 1with only a few chaetae on both rami, especially on the notopodium, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae elliptical on the notopodium and rounded on the neuropodium. (Figs. 32A–B; 33A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–7 (Fig. 33A–D), square-shaped on chaetigers 8 and 9 (largest on chaetiger 8) and rounded afterwards, gradually reducing in size towards the posterior region, present as a low flap on last chaetigers. Notopodial prechaetal lamellae absent throughout. Dorsal crests low from chaetigers 8–9 to chaetigers 14–15 (Fig. 32A).

Neuropodial postchaetal lamellae elliptical on chaetigers 2–4 (Fig. 33A–D), including a pointed tip on chaetiger 3. Lamellae rounded from chaetiger 5, gradually reduced in size towards the posterior region, present as a low flap on the last chaetigers. Neuropodial prechaetal lamellae absent throughout (Fig. 32B).

Chaetae from notopodia and neuropodia unilimbate (narrow limbation), non-granulated capillaries, organized in three rows on notopodia from chaetiger 2 to chaetiger 10–12 and two rows in neuropodia and remaining notopodial chaetigers (Fig. 34A–B). Chaetae from each row of different lengths, anterior row shortest. Neuropodial chaetae slightly shorter than notopodial chaetae. Neuropodium of chaetiger 3 bearing very long chaetae on the posterior row (up to four times longer than anterior row). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig. 34C).

Hooks in notopodia starting from chaetigers 22–44, up to five per fascicle, accompanied by 1–4 short non-limbate capillaries (Fig. 34D–E). Hooks in neuropodia starting from chaetigers 10–12, up to 12 per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks multidentate, with eight secondary teeth organized in two rows above the main tooth (Fig. 34D). Small secondary

91 hood present (Fig. 34D). Sabre chaetae starting from chaetigers 10–17. Sabre chaetae morphologically identical to remaining capillaries – albeit shorter, only recognizable by a slight curvature and by its ventral-most position (Fig. 34F).

Up to six pairs of branchiae, subtriangular, robust and flattened on chaetigers 3 and 4 and cirriform, sculptured (“wrinkled”) on the remaining chaetigers, all tapered at the tips (Fig. 35A– B). Branchiae present from chaetiger 2, up to three times longer than notopodial lamellae, slightly reduced in length towards the last branchial pair. Branchiae on chaetigers 3 and 4 heavily ciliated throughout, branchiae on the remaining chaetigers ciliated at the base. All branchiae completely free from notopodial postchaetal lamellae (Fig. 32A).

Pygidium rounded, bearing a single long dorsal cirrus (Fig. 35C).

Gametes not observed.

Variation: In juveniles (up to 2.5 mm long), the prostomium is square-shaped, rather than rectangular.

Methyl green staining pattern: prostomium, lateral and ventral sides of the peristomium, postchaetal neuropodial lamellae from chaetiger 2 to chaetiger 6, postchaetal notopodial lamellae from chaetiger 2 to chaetigers 12–15 and dorsal crests intensely stained; tips of the last branchial pair slightly stained.

Remarks. Prionospio corrugatus sp. nov. is easily recognizable from the remaining apinnate Brazilian species by its set of characters: a rectangular prostomium, sculptured branchiae (except on chaetigers 2 and 3), three rows of capillary chaetae on notopodia from chaetiger 2 to chaetigers 10– 2, row of long neurochaetae on chaetiger three and pygidium bearing a single dorsal cirrus.

The narrow rectangular prostomium is most similar to P. ehlersi Fauvel, 1928, described from Morocco, P. kaplani, P. amarsupiata Neal & Altamira in Paterson et al., 2016, described from Portugal and P. branchilucida Altamira, Glover, & Paterson in Paterson et al. 2016, described from the Kaplan Clarion-Clipperton Fracture Zone, East Pacific Ocean, but these species lack a row of long neurochaetae on chaetiger 3 and possess different branchial distribution (four pairs in P. ehlersi and P. amarsupiata; two pairs in P. branchilucida and P. kaplani).

While the row of long neurochaetae on chaetiger 3 and a third row of capillaries on some anterior notopodia are known only from Prionospio corrugatus sp. nov. and P. fauchaldi (discussed below), sculptured branchiae have only been observed in P. cinthyae sp. nov. (described earlier in this work), P. sandersi, described from the Galapagos Islands, Ecuador, P. laciniosa and P. vallensis, although these species can be distinguished from P. corrugatus sp. nov. by the prostomium morphology (rectangular and truncated anteriorly in P. corrugatus sp. nov., P.

92 fauchaldi, triangular in P. cinthyae sp. nov. and P. vallensis, broadly rounded in P. sandersi and triangular in P. laciniosa), presence of typical sabre chaetae (except for P. fauchaldi, see discussion below), branchial distribution (up to six pairs in P. corrugatus sp. nov., four pairs in P. fauchaldi, P. laciniosa, P. cinthyae sp. nov. and P. vallensis, and nine pairs in P. sandersi) and dorsal crests presence, morphology and distribution, as dorsal crests are lacking in P. sandersi, present as semicircular flaps on chaetigers 5–13 in P. laciniosa, from chaetigers 8–9 to chaetigers 14–15 in P. corrugatus sp. nov., from chaetiger 6 to beyond chaetiger 20 in P. vallensis, from chaetigers 8–11 in P. fauchaldi (distribution not mentioned) and from chaetiger 6 to chaetigers 7–9 in P. cinthyae sp. nov.

Prionospio corrugatus is very similar to P. fauchaldi, as both species bear a rectangular prostomium, a long row of neurochaetae on chaetiger 3, three rows of chaetae on several anterior segments, hook with four pairs of secondary teeth and delicate sabre chaetae. However, species differed on important characters, such as the number of branchiae, up to six pairs in P. corrugatus sp. nov. and four pairs in P. fauchaldi, shape of the postchaetal lamellae on chaetiger 1 (elliptical on the notopodium and rounded on the neuropodium of P. corrugatus sp. nov. and rounded on the notopodium and elliptical on the neuropodium of P. fauchaldi), chaetigers bearing three rows of capillaries in the notopodium (from chaetiger 2 to chaetigers 10–12 P. corrugatus sp. nov. and from chaetiger 2 to chaetigers 15–17 in P. fauchaldi), distribution of dorsal crests (from chaetigers 8–9 to chaetigers 14–15 in P. corrugatus sp. nov. and starting from chaetigers 8–11 in P. fauchaldi [distribution not mentioned]), shape of notopodial postchaetal lamellae on various anterior chaetigers (larger on chaetigers 2–8 and broad, square-shaped on chaetiger 8 in P. corrugatus sp. nov.), starting chaetiger of neuropodial hooded hooks (chaetigers 10–12 in P. corrugatus sp. nov. and chaetigers 12–13 in P. fauchaldi) and presence of sabre chaetae on all chaetigers from chaetigers 10–17 in P. corrugatus sp. nov. which are frequently lacking in P. fauchaldi. Unfortunately, all type material of P. fauchaldi was incomplete, thus making it impossible to compare the starting chaetiger of notopodial hooded hooks and pygidium morphology.

Etymology. The specific epithet, corrugatus, derives from the wrinkled appearance of branchiae from chaetigers 2 and 5 (corruga, Latin for wrinkled and the suffix -atus, meaning possessing, bearing).

Habitat: muddy sand to mud, 140–1905 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Prionospio cf. delta Hartman, 1965

(Figures 36–40)

Minuspio minor Fauchald & Hancock, 1981

Prionospio (Minuspio) delta Hartman, 1965

Prionospio longibranchiata Reish, 1968

Prionospio delta in Pardo & Peixoto (in press).

Material examined. Amb3 E5, 19º 36' 26.24" S 39º 10' 17.35" W, 352m (7 ind); Amb3 F5, 19º 34' 20.51" S 38º 41' 18.75" W, 438m (4 ind); Amb3 CAND5, 19º 33' 20.99" S 39º 2' 36.2" W, 374m (10 ind); Amb5 B5, 20º 35' 16.23" S 39º 53' 47.1" W, 382m (2 ind); Amb5 C5, 20º 14' 19.45" S 39º 48' 36.67" W, 416m (2 ind); Amb5 D6, 19º 50' 4.42" S 39º 26' 33.42" W, 1053m (2 ind); Amb6 D5, 19º 46' 31.83" S 39º 30' 3.38" W, 402m (5 ind); Amb6 CANWN4, 19º 49' 10.21" S 39º 36' 10.05" W, 124m (1 ind); Amb6 CANWN5, 19º 49' 37.21" S 39º 35' 41.25" W, 352m (47 ind); Amb11 A5, 21º 4' 4.67" S 40º 13' 6.06" W, 383m (2 ind); Amb11 B5, 20º 35' 15.33" S 39º 53' 45.22" W, 382m (2 ind); Amb11 C5, 20º 14' 17.95" S 39º 48' 34.35" W, 418m (8 ind); Amb12 D5, 19º 46' 32.84" S 39º 30' 3.65" W, 431m (7 ind); Amb12 E5, 19º 36' 30.6" S 39º 10' 19.39" W, 349m (5 ind); Amb12 CAND5, 19º 33' 23.09" S 39º 2' 35.67" W, 446m (1 ind); Amb12 CANWN4, 19º 49' 6.26" S 39º 36' 9.34" W, 181m (1 ind); Amb12 CANWN5, 19º 49' 36.9" S 39º 35' 42.69" W, 363m (16 ind); Amb14 A4, 21º 4' 4.81" S 40º 14' 13.86" W, 147m (1 ind).

Comparative material examined. USNM 80299 (1 ind); USNM 80300 (1 ind).

Diagnostic features: Up to eight pairs of thin strap-like branchiae, single median prostomial peak, sabre chaetae consistently from chaetiger 12 and lack of dorsal crests.

Description. A large-sized Prionospio, largest complete specimen 16 mm long, 0.45 mm wide at the widest point for 82 chaetigers. Body dorsoventrally flattened throughout, especially on the branchial region, tapering towards the pygidium. Body color yellow to whitish in alcohol (Fig. 36).

Prostomium triangular, inflated on the mid-length, widened towards the anterior margin, extending posteriorly as a narrow keel reaching the posterior margin of chaetiger 1. Median prostomial peak present. One or two pairs of eyes present in most specimens. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent. Grooved palps reaching up to chaetiger 14 (Figs. 36; 37A).

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae absent on the notopodium and digitiform on the neuropodium, greatly reduced and barely visible (Fig. 37A–B). Prechaetal lamellae absent.

Notopodial lamellae foliaceous from chaetiger 2 to chaetigers 11–13 (depending on specimen size) (Fig. 38A–D), rounded afterwards and gradually reducing in size towards the posterior region, present as a low flap on the last chaetigers. Notopodial prechaetal lamellae absent throughout. Dorsal crests very low from chaetigers 12–14 to chaetigers 19–20, not easily visible. (Fig. 37A).

Neuropodial postchaetal lamellae rounded on chaetiger 2 and elliptical from chaetiger 3 (Fig. 38A–D), gradually reduced in size towards the posterior region, present as a low flap on last chaetigers. Neuropodial prechaetal lamellae absent throughout (Fig. 37B).

Chaetae from notopodia and neuropodia organized in two rows of unilimbate and granulated capillaries (Fig. 39A). Chaetae from both rows of equal length, although neuropodial chaetae are slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig. 39B).

Hooks in notopodia starting from chaetigers 23–39, up to five per fascicle, accompanied by 1–4 short non-limbate capillaries (Fig. 39C–D). Hooks in neuropodia starting from chaetigers 14–22, up to seven per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks multidentate, with eight secondary teeth organized in two rows above the main tooth (Figs. 37C; 39C). Secondary hood absent (Fig. 39C). Sabre chaetae consistently from chaetiger 12. Sabre chaetae unilimbate (with broad limbation) and densely dark granulations along the shaft (Fig. 39E).

Up to eight pairs of smooth branchiae, cirriform, strap-like. Branchiae from chaetiger 2, up to 10 times longer than notopodial lamellae, slightly reduced in length towards the last branchial pair. Branchiae apparently non-ciliated, but with small annulations throughout the length (except at the base) (Figs. 37A–B; 40A). All branchiae completely free from notopodial postchaetal lamellae (Fig. 37A).

Pygidium bearing one thin dorsal cirrus and two short ventral lobes (Fig. 40B).

Gametes not observed.

Methyl green staining pattern: prostomium, peristomium, postchaetal lamellae from chaetiger 2 up to chaetigers 15–20 intensely stained; ventral side of branchial region slightly stained.

Remarks. Prionospio cf. delta is easily recognized among Brazilian apinnate Prionospio species by its cirriform, strap-like branchiae and triangular prostomium, inflated on the mid-length. Our

95 specimens are similar to the original description (Hartman, 1965) and its redescription (Maciolek, 1985), differing in the number of branchiae and number of secondary teeth on the hooded hooks.

While in the descriptions of P. delta specimens had up to six pairs of branchiae, our specimens bore eight pairs of branchiae, although the last pairs of branchiae could have been lost or had not yet developed in specimens examined on the previous descriptions. As for the number of secondary teeth, three pairs in the descriptions and four pairs in our specimens, the last pair of teeth could have been overlooked using light microscopy, due to its small size. However, such discrepancies to the original descriptions cannot be ignored, justifying the ‘cf.’ adoption.

Only a few species possess branchiae of similar morphology, such as Prionospio pulchra Imajima, 1990a, described from Japan, P. elegantula Imajima, P. perkinsi, P. anatolica Dagli & Çinar, 2011, described from the Levantine Sea, Turkey and P. yuriel Wilson, 1990, described from Australia, all which possess a similar branchial distribution (eight pairs in P. cf. delta, up to 10 pairs in P. pulchra and P. perkinsi, four pairs in P. elegantula, seven pairs in P. anatolica and up to nine pairs in P. yuriel), except for P. elegantula, which possess only four pairs of branchiae. Interestingly, all mentioned species (except for P. perkinsi) lack notopodial postchaetal lamella on chaetiger 1.

Prionospio cf. delta can be quickly distinguished from P. perkinsi by the prostomial shape – triangular with a single median prostomial peak in P. cf. delta and rounded with several small peaks in P. perkinsi, starting chaetigers of hooded hooks (from chaetigers 23–39 in notopodia and from chaetigers 14–22 in neuropodia in P. cf. delta and from chaetigers 16–23 in notopodia and from chaetigers 13–18 in neuropodia in P. perkinsi and by the presence of sabre chaetae and dorsal crests in P. cf. delta.

Although P. cf. delta shares some similarities with P. yuriel and P. anatolica – an overlap on the starting chaetiger of sabre chaetae (chaetiger 12 in P. cf. delta, chaetigers 11–13 in P. yuriel and chaetiger 11 in P. anatolica), starting chaetiger of hooded hooks (from chaetigers 23–39 in notopodia and from chaetigers 14–22 in neuropodia in P. cf. delta, from chaetigers 34–42 in notopodia and from chaetigers 14–19 in neuropodia in P. yuriel and from chaetigers 27–39 in notopodia and from chaetigers 14–15 in neuropodia in P. anatolica), morphology of the hooded hooks (four pairs of secondary teeth in P. cf. delta and P. anatolica and four to five pairs in P. yuriel) and similar number of branchiae, these species can be distinguished from P. cf. delta by the prostomial shape – prostomium triangular, inflated on the mid-length in P. cf. delta, narrow and elongated anteriorly in P. yuriel and inflated, rounded anteriorly in P. anatolica) and lack of dorsal crests, present in P. cf. delta.

Prionospio cf. delta is also similar to P. pulchra, but these species can be readily distinguished by the prostomial shape – prostomium triangular, inflated on the mid-length in P. cf. delta and

96 subtriangular in P. pulchra and number of prostomial peaks – five peaks in P. pulchra and a single median peak in P. cf. delta.

Although not described as wrinkled, branchiae of Prionospio cf. delta on the SEM micrographs seem sculptured, just as branchiae found in Prionospio fauchaldi and P. corrugatus sp. nov. (Fig 27A; Fig X), but these species differ on the branchial number (eight pairs of strap-like branchiae in P. cf. delta, four pairs in P. fauchaldi and up to six pairs in P. corrugatus sp. nov., shape of the prostomium (triangular in P. cf. delta and rectangular in P. fauchaldi and P. corrugatus sp. nov., presence of typical sabre chaetae in P. cf. delta and lack of long row of neurochaetae on chaetiger 3 in P. cf. delta.

Habitat: muddy sand to mud, 124–1053 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean. According to Maciolek (1985), this species has been recorded from Northwestern Atlantic Ocean (USA), Central America (Mexico), Southwestern Atlantic Ocean (Suriname, type locality), Southeastern Atlantic Ocean (Namibia and Angola) and Northeastern Pacific Ocean (USA and Mexico).

Identification key to apinnate species of Prionospio recorded in this work.

1 Sabre chaetae absent…………..………………………..……………………………………2 - Sabre chaetae present……………………….………………………………………………..3 2(1) Prostomium rounded, dorsal crests lacking………………..………….P. fosterae sp. nov. - Prostomium bearing two short antero-lateral projections, dorsal crests present in some postbranchial chaetigers……………………………………………………..P. acutus sp. nov. 3(1) Sabre chaetae appearing abruptly…………..…………………………………………….4 - Sabre chaetae appearing as a gradual transformation of inferior capillary chaetae…………………………………………………………………….P. mutatus sp. nov. 4(3) Sabre chaetae robust……………………….……………………………………………..5 - Sabre chaetae thin, delicate, similar to capillary chaetae…...………....P. corrugatus sp. nov. 5(4) Dorsal crests present…………………..…...……………………………………………..6 - Dorsal crests absent………………..…………………………………..P. hartmanae sp. nov. 6(5) Branchia up to twice as long as notopodial lamellae, cirriform on first and last chaetigers…………………………………………………………………..P. kinbergi sp. nov. - Branchia more than twice as long as notopodial lamellae…………………………………...7 7(6) Up to eight pairs of extremely long branchiae, up to 10 times longer than notopodial lamellae……………………………………………………………………………...P. cf. delta

- Only four pairs of branchiae, first and last pairs sculpted………...……...P. cinthyae sp. nov.

Acknowledgments

The authors would like to thank Dr. Michelle Klautau (Universidade Federal do Rio de Janeiro), Dr. Cinthya Santos (Universidade Federal Fluminense), and Dr. Alexandra Rizzo (Universidade do Estado do Rio de Janeiro) for interesting discussions and insightful comments on the manuscript.

This paper is part of the D. Sc. requirements of Antônio João Malafaia Peixoto at the Biodiversity and Evolutionary Biology Graduate Program of the Federal University of Rio de Janeiro. Financial support was provided by the National Council for Scientific and Technological Development (CNPq) (Grant 43900/2014-0).

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001 (Process: 1654206). AJMP was supported by Carlos Chagas Filho Research Support Foundation (FAPERJ) (Registration: 2019.00407.5).

Thanks are due to the Smithsonian Institution for the Kenneth Jay Boss Fellowship in Invertebrate Zoology (National Museum of Natural History) in 2018. PCP received a fellowship from CNPq (Process: 304321/2017-6). References

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Radashevsky, V.I., Lana, P.C. & Nalesso, R.C. (2006) Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa, 1353, 1–37.

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Reish, D.J. (1968). A biological survey of Bahia de Los Angeles, California, Mexico. II. Benthic polychaetous annelids. Transactions of the San Diego Society of Natural History, 15, 67–106.

Rocha, M.B. & Paiva, P.C. (2012) Scolelepis (Polychaeta: Spionidae) from the Brazilian coast with a diagnosis of the genus. Zoologia, 29, 385–393.

Rocha, M.B., Radashevsky, V. & Paiva, P.C. (2009) Espécies de Scolelepis (Polychaeta, Spionidae) de praias do Estado do Rio de Janeiro, Brasil. Biota Neotropica, 9, 101–108.

Sigvaldadóttir, E. (1998) Cladistic analysis and classification of Prionospio and related genera (Polychaeta, Spionidae). Zoologica Scripta, 27, 175–187.

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Wilson, R.S. (1990) Prionospio and Paraprionospio (Polychaeta: Spionidae) from Southern Australia. Memoirs of the Museum of Victoria, 50, 243–274.

Wirén, A. (1883) Chaetopoder från Sibiriska Ishafvet och Berings Haf Insamlade under Vega- Expeditionen 1878-1879. Vega-Expeditionens Vetenskapliga Iakttagelser bearbetade af deltagare i resan och andra forskare. F&G Beijers Förlag, Stockholm, 2, 383–428.

Zhou, J. & Li, X. (2009) Report of Prionospio complex (Annelida: Polychaeta: Spionidae) from China's waters, with description of new species. Acta Oceanologica Sinica, 1, 116–127.

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Legends of illustrations

Figure 1. Map showing sampling area of the AMBES project (blue polygon).

Figure 2. Prionospio acutus sp. nov., anterior chaetigers and mid-body, dorsal view (MZUSP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 3. SEM of Prionospio acutus sp. nov., dorsal view. A. Anterior chaetigers. B. Mid-body chaetigers. Abbreviations: br, branchia; dc, dorsal crest; pe, peristomium; pr, prostomium.

Figure 4. Prionospio acutus sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 5. Prionospio acutus sp. nov. A. Anterior capillary chaeta. B. Posterior capillary chaetae. C. Hooded hook. D. Companion capillary chaetae. E. Branchiae.

Figure 6. Prionospio fosterae sp. nov., anterior chaetigers and mid-body, dorsal view (MZUSP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Figure 7. SEM of Prionospio fosterae sp. nov., dorsal view. A. Anterior chaetigers and mid- body. B. Close-up of the anterior chaetigers. C. Hooks (hoods removed). Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Figure 8. Prionospio fosterae sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 9. Prionospio fosterae sp. nov. A. Anterior capillary chaeta from anterior row. B. Anterior capillary chaeta from posterior row. C. Posterior capillary chaetae. D. Hooded hook. E. Companion capillary chaetae.

Figure 10. Prionospio fosterae sp. nov. A. Branchia. B. Posterior-most chaetigers and pygidium.

Figure 11. Prionospio kinbergi sp. nov., dorso-lateral view (MZUSP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: Cbr, cirriform branchia; dc, dorsal crest; Fbr, flattened branchia; pe, peristomium; pr, prostomium.

Figure 12. SEM of Prionospio kinbergi sp. nov., dorsal view. Anterior chaetigers and mid- body. B. Close-up of flattened branchiae. C. Hooks (hoods removed). Abbreviations: br, branchia; dc, dorsal crest, nol, notopodial lamella; pe, peristomium; pr, prostomium.

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Figure 13. Prionospio kinbergi sp. nov., anterior chaetigers, frontal view (prechaetal lamellae omitted). A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 14. Prionospio kinbergi sp. nov. A. Anterior capillary chaeta from anterior row. B. Anterior capillary chaeta from posterior row. C. Posterior capillary chaetae. D. Hooded hook. E. Companion capillary chaetae. F. Sabre chaetae.

Figure 15. Prionospio kinbergi sp. nov. A. Flattened branchiae. B. Cirriform branchiae. C. Posterior-most chaetigers and pygidium.

Figure 16. Prionospio hartmanae sp. nov., dorso-lateral view (MZUSP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; ey, eye; pe, peristomium; pr, prostomium.

Figure 17. SEM of Prionospio hartmanae sp. nov. A. Anterior chaetigers and mid-body, dorsal view. B. Close-up of anterior chaetigers, lateral view. Hook (hood removed). Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Figure 18. Prionospio hartmanae sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 19. Prionospio hartmanae sp. nov. A. Anterior capillary chaeta. B. Posterior capillary chaetae. C. Hooded hook. D. Companion capillary chaetae. E. Sabre chaetae.

Figure 20. Prionospio hartmanae sp. nov. A. Branchia. B. Posterior-most chaetigers and pygidium.

Figure 21. Prionospio mutatus sp. nov., dorsal view (MZUSP XXXX, paratype), showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Figure 22. SEM of Prionospio mutatus sp. nov., dorsal view. Anterior chaetigers and mid- body. B. Close-up of the anterior region. C. Mid-body. Abbreviations: br, branchia; dc, dorsal crest; nol, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 23. Prionospio mutatus sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 24. Prionospio mutatus sp. nov. A. Anterior capillary chaeta. B. Posterior capillary chaetae. C. Hooded hook. D. Companion capillary chaetae. E–F. Gradual transformation of capillary chaetae in sabre chaetae. G. Sabre chaetae.

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Figure 25. Prionospio mutatus sp. nov. A. Branchia. B. Posterior-most chaetigers and pygidium.

Figure 26. Prionospio cinthyae sp. nov., showing characteristic pigmentation in ethanol (MNRJP XXXX, paratype). Abbreviations: br, branchia; nol, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 27. SEM of Prionospio cinthyae sp. nov. A. whole animal, dorsal view. B. Anterior chaetigers and mid-body, dorsal view. C. Hooded hooks (hoods removed). Abbreviations: br, branchia; dc, dorsal crest; nol, notopodial lamella; pe, peristomium; pr, prostomium.

Figure 28. Prionospio cinthyae sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 29. Prionospio cinthyae sp. nov. A. Anterior capillary notochaetae. B. Anterior capillary neurochaeta from anterior row. C. Anterior capillary neurochaeta from posterior row. D. Posterior capillary chaetae. E. Hooded hook. F. Companion capillary chaetae. G. Sabre chaetae.

Figure 30. Prionospio cinthyae sp. nov. A. Sculptured branchia. B. Flattened branchia. C. Posterior-most chaetigers and pygidium.

Figure 31. Prionospio corrugatus sp. nov., showing characteristic pigmentation in ethanol (MNRJP XXXX, paratype). Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

Figure 32. SEM of Prionospio corrugatus sp. nov. A. Anterior chaetigers and mid-body, dorsal view. B. Anterior chaetigers, ventral view. C. Hooded hooks (hoods partially removed). Abbreviations: br, branchia; cha3, chaetiger 3; nol, notopodial lamella; pe, peristomium; pr, prostomium.

Figure 33. Prionospio corrugatus sp. nov., anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 34. Prionospio corrugatus sp. nov. A. Anterior capillary chaeta from anterior row. B. Anterior capillary chaeta from posterior row. C. Posterior capillary chaetae. D. Hooded hook. E. Companion capillary chaetae. F. Sabre chaetae.

Figure 35. Prionospio corrugatus sp. nov. A. Sculptured branchia. B. Flattened branchia. C. Posterior-most chaetigers and pygidium.

Figure 36. Prionospio cf. delta, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; pa, palp; pe, peristomium; pr, prostomium.

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Figure 37. SEM of Prionospio cf. delta. A. Anterior chaetigers, dorsal view. B. Anterior chaetigers and mid-body, ventro-lateral view. Hooks (hoods removed). Abbreviations: br, branchia; pa, palp; pe, peristomium; pp, prostomial peak; pr, prostomium.

Figure 38. Prionospio cf. delta, anterior chaetigers, frontal view. A. Chaetiger 1. B. Chaetiger 2. C. Chaetiger 3. D. Chaetiger 4.

Figure 39. Prionospio cf. delta. A. Anterior capillary chaeta. B. Posterior capillary chaetae. C. Hooded hook. D. Companion capillary chaetae. E. Sabre chaetae.

Figure 40. Prionospio cf. delta. A. Branchia. B. Posterior-most chaetigers and pygidium.

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Tables

Table 1. Comparison among Prionospio species examined in this work.

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Table 1 (continued). Comparison among Prionospio species examined in this work.

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Table 1 (continued). Comparison among Prionospio species examined in this work.

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Capítulo 4

Species from the Prionospio complex (Annelida: Spionidae) from southern Brazil, including new species from Prionospio and Paraprionospio

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista alvo: Zootaxa Status: A submeter

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Species from the Prionospio complex (Annelida: Spionidae) from southern Brazil, including new species from Prionospio and Paraprionospio

Running title: NEW PRIONOSPIO FROM BRAZIL

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract 13 species belonging to four genera of the Prionospio-complex were recorded from Espírito Santo Basin, southeastern Brazil. Nine of these species – Prionospio brevipinnulata sp. nov., Prionospio capixaba sp. nov., Prionospio quadrilamellata sp. nov., Prionospio alexandrae sp. nov., Prionospio triangularis sp. nov., Prionospio posterocristata sp. nov., Prionospio absensi sp. nov., Prionospio biancoi sp. nov. and Paraprionospio anterocristata sp. nov. are new to science. Species can be distinguished by the prostomium morphology, parapodial lamellae morphology, branchiae morphology, presence and distribution of dorsal crests as well as the starting chaetiger and morphology of hooded hooks. An identification key and a comparison table to Prionospio species recorded in this work are provided.

Keywords: morphology, biodiversity, Prionospio-complex, Annelida.

Introduction

The Spionidae Grube, 1850 are one of the largest families of annelids, often dominating soft- bottom communities, especially in organically enriched environments. There are approximately 580 known species organized in 39 genera (Blake et al. 2017), although this most likely represents an underestimate, as the family is insufficiently known in deep-sea environments (as seen in Paiva & Barroso 2010), possess well-documented species-complexes (Sato-Okoshi et al. 2016), as well as species with suspiciously large distributions and genera that were never revised.

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In Brazil, most spionid taxonomic studies focus on shallow-waters environments (except for Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019) and are restricted to the southern and southeastern coast of Brazil. In fact, while only a few studies are known from the Northeast coast (checklists and ecological works), no taxonomic study is known from the North coast of Brazil (based on data compiled by Amaral et al. 2013). Only a handful of taxa are better known, such as Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Laonice Malmgren, 1867 (Kinberg 1866; Nonato et al. 1986; Radashevsky & Lana 2009), Laubieriellus Maciolek, 1981b (Peixoto & Paiva 2019), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), Polydora Bosc, 1802 (Radashevsky et al. 2006), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017) and Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018). Studies encompassing several genera are scarce, such as Bolívar & Lana (1987) to the continental shelf of the state of Paraná, southern Brazil, Paiva & Barroso (2010) to the continental slope of Campos Basin, state of Rio de Janeiro, southeastern Brazil, as well as and Pardo et al. (2006) and Pardo & Peixoto (in press) to southern and southeastern Brazil, although none of this aforementioned works described new species.

According to Blake et al. (2017), the Prionospio complex is an informal taxonomic unity within the subfamily Nerininae Söderström, 1920 that includes several morphologically similar taxa, possessing slender bodies; branchiae of varied morphology, present on anterior part of the body (rarely absent, as in Prionospio nonatoi Peixoto & Paiva 2019); prostomium of simple morphology, lacking horns or occipital antennae, bi- to multidentate hooded hooks preset on both rami (except for Laubieriellus, which bears hooks only on neuropodia); well-developed anterior parapodial lamellae and hard-membraned eggs (Blake et al. 2017). The Prionospio complex includes the following genera: Prionospio, Apoprionospio Foster, 1969, Orthoprionospio Blake & Kudenov, 1978, Aurospio Maciolek, 1981a, Laubieriellus, Paraprionospio Caullery, 1914 and Streblospio Webster, 1879, all of which were previously recorded in Brazil, except for Orthoprionospio. Two additional taxa, Minuspio Foster, 1971 and Aquilaspio Foster, 1971, which were described as subgenera of Prionospio (Foster, 1971), were latter synonymized by Sigvaldadóttir (1998), due to the lack of support for monophyly. These genera can be separated mostly on prostomium morphology and branchiae morphology and distribution.

The genus Prionospio represents one of the most morphologically diverse groups within Spionidae, with over than 100 known species, being the family’s most specious genus. The genus possesses great diversity on the branchial morphology and arrangement, characters that bear taxonomic value and might be species-specific. This branchial diversity was used in the past to erect the following genus, closely related to Prionospio: Apoprionospio, Minuspio and Aquilaspio. These taxa were sequentially treated as subgenus of Prionospio (Light, 1977; Blake

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& Kudenov, 1978) and posteriorly synonymized with Prionospio (Sigvaldadóttir, 1998), although such synonym is not widely accepted, since some researchers still consider Aquilaspio (Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011) and Minuspio (Aguirrezabalaga & Ceberio 2005; Zhou & Li 2009; Dagli & Çinar 2009; Dagli & Çinar 2011; Dagli et al. al. 2011; Dagli 2013; Delgado-Blas & Díaz-Díaz 2013) as valid taxa, while Maciolek (1985) and Blake et al. (2017) reestablished Apoprionospio.

The genus Aurospio was described based on material collected from the deep sea and the type species, A. dibranchiata is considered an amphi-Atlantic species (Maciolek 1981a). Blake et al. (2017) considers Aurospio as part of the Prionospio-complex due to morphological similarities, such as cylindrical body, branchiae limited to the anterior part of the body, multidentate hooded hooks, well-developed parapodial lamellae on the anterior part of the body and pygidium bearing long median cirrus and a pair of short ventrolateral cirri or lobes. Aurospio can be separated from Prionospio by branchiae morphology – usually from chaetiger 3, flattened and shorter or about the same length than notopodial lamellae in Aurospio and usually from chaetiger 2, cylindrical or triangular, usually longer than notopodial lamellae in Prionospio; branchiae basally fused to the notopodial lamellae in Aurospio and branchiae free from the notopodial lamellae in Prionospio and lack on secondary hood on the hooded hooks in Aurospio and secondary hood usually present in Prionospio. Curiously, during a preliminary phylogenetic analysis, Mincks et al. (2009) reported a polytomy of different Aurospio and Prionospio clades using the 16S gene and a mixed clade of Aurospio and Prionospio species using the 18S gene.

The genus Apoprionospio was erected by Foster, 1969 to include Prionospio species bearing three pairs of apinnate and cirriform branchiae starting on chaetigers 2–4 and one pair of pinnate branchiae from chaetiger 5, bearing only platelike pinnules. Apoprionospio was synonymized with Prionospio by Blake & Kudenov (1978), but was resurrected by Maciolek (1985), only to be synonymized a second time by Sigvaldadóttir (1998) when the author made a cladistic analysis on Prionospio and related taxa. Currently the genus was reinstated by Blake et al. (2017), as the authors considered that the platelike pinnules found in Apoprionospio are more similar to those found in Paraprionospio than digitiform pinnules found in Prionospio, thus synonymizing Apoprionospio would also require a revaluation of Paraprionospio.

The genus Paraprionospio is a part of the Prionospio-complex, being characterized by a fusiform prostomium, peristomium fused with achaetous segment, prominent lateral wings, branchiae starting from chaetiger 1 and branchiae bearing plate-like pinnules. This genus was described as a subgenus of Prionospio by Caullery (1914), who also reallocated Prionospio pinnata to the subgenus Paraprionospio. The subgenus remained largely neglected until Foster (1971) raised it to genus status. The genus Laubieriellus was erected by Maciolek (1981b), based on specimens

132 from the deep-sea. Blake et al. (2017) treat Laubieriellus as part of the Prionospio-complex, differing mainly on the lack of notopodial hooks and presence of ventral crests on a variable number of anterior chaetigers. Laubieriellus was addressed by Peixoto & Paiva (2019), while no Streblospio species was recorded in this work.

Currently, 14 Prionospio species have been recorded in the Brazilian coast: P. cristata Foster, 1971, described to the Atlantic coast of the USA, P. cirrifera Wirén (1883) from the Kara Sea, Arctic Ocean, northern Russia, P. heterobranchia Moore, 1907 to the Atlantic coast of the USA, P. delta Hartman, 1965, described from Suriname, P. fauchaldi Maciolek, 1985, described from Massachusetts (USA), P. dubia Day, 1961, described to South Africa, P. lighti Maciolek, 1985, described from Washington (USA), P. orensanzi Blake, 1983, described to Argentina, P. multibranchiata Berkeley, 1927, described from British Columbia, Canada, P. perkinsi Maciolek, 1985, described from the Gulf of Mexico, Florida (USA), P. ehlersi Fauvel, 1928 to Morocco, P. steenstrupi Malmgren, 1867, described to Iceland, P. nonatoi Peixoto & Paiva, 2019, described from southeastern Brazil and P. solisi Peixoto & Paiva, 2019, also described from southeastern Brazil (Amaral et al. 2013; Pardo et al. 2006; Peixoto & Paiva 2019; Pardo & Peixoto in press). Seven additional Prionospio species are being described by Peixoto & Paiva (ms, Chapter 3).

Remaining species from the Prionospio-complex that were recorded in Brazil include Aurospio dibranchiata Maciolek, 1981a, described from Argentina (although recorded on most Atlantic Ocean), Apoprionospio dayi, Foster, 1969, described from the Atlantic coast of the USA, A. pygmaeus (Hartman, 1961), described from the Pacific coast of the USA, Laubieriellus decapitata Peixoto & Paiva, 2019, described to southeastern Brazil, Paraprionospio pinnata Ehlers, 1901, described to Chile and Streblospio benedicti Webster, 1879, described from the Atlantic coast of the USA (Amaral et al. 2013; Pardo et al. 2006; Peixoto & Paiva 2019; Pardo & Peixoto in press). However, it should be noted that most Brazilian records belong to species described to distant locations, are known-species complexes and were never confirmed. The status of these records will be reviewed in future works.

13 species have been recorded in this work, based on material collected on Espírito Santo Basin, southeastern Brazil, during the oceanographic cruises of the “Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company). Nine of these species are new to science, while two species were previously recorded from nearby locations and the last two species may represent new species, although additional material will be necessary to make a taxonomical decision.

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Material and Methods

Sediment samples were collected during the oceanographic cruises of the AMBES Project on the Espírito Santo Basin and northern portion of Campos Basin between 2010 and 2013 (Fig 1). Sampling efforts were divided in summer months (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and winter months (sample codes Amb2, Amb11, Amb12, Amb13, and Amb14).

Sediment samples were collected using a Van Veen Grab sampler on the continental shelf and a box corer in the continental slope. Samples were immediately fixed in situ in a 10% formalin- seawater solution, then taken to the laboratory, washed in freshwater and then transferred to 70% ethanol. Afterwards, samples were elutriated using a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm-mesh sieve (for samples collected on the continental slope and on the Watu Norte and Doce canyons) and specimens were sorted out. Specimens were examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope and measurements were taken through a calibrated eyepiece attached to the microscope.

Specimens were stained in a methyl green solution (1 g/L in 70% EtOH) for 10‒30 s and transferred to distilled water to avoid stain fading, in order to enhance visualization of features such as branchial scars and to observe any species-specific staining patterns. As specimens were fixed and kept in formalin for several months, molecular studies were not possible. For Prionospio species, main taxonomic characters are summarized in Table 1 and an identification key is provided for species recorded in this study, while for Paraprionospio and Apoprionospio, comparison tables are provided including species previously recorded in Brazil by Amaral et al. (2013) (Tables 2 and 3, respectively).

Specimen lots deposited in the Smithsonian National Museum of National History (USNM), Smithsonian Institution, Washington, D.C., USA were examined and compared with material collected in this study. Type material of Apoprionospio dayi USNM 39487 (holotype) and USNM 39488 (5 paratypes), Aurospio dibranchiata USNM 60326 (holotype) and USNM 60327 (35 paratypes), Prionospio cristata USNM 43001 (holotype) and USNM 43002 (2 paratypes), as well as non-type lots of Aurospio dibranchiata USNM 162820 (20 specimens) and USNM 1106015 (9 specimens), Paraprionospio alata USNM 1260223 (1 specimen), Prionospio cristata USNM 1171111 (74 specimens) and Prionospio dubia USNM 65932 (two specimens) were examined.

Before scanning electron microscope (SEM) studies, specimens were dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each, followed by three changes of 100% EtOH), then critical-point dried using CO2 or transferred to an ascending hexamethyldisilane (HMDS) series (2 parts 100% EtOH:1 part HMDS, 1:1, 1:2, pure HMDS, followed by a second change of pure

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HMDS, 15 min each), and dried in a fume hood. Specimens were then mounted on stubs, sputter coated with gold and viewed in a JEOL JSM-6510 LV SEM or a JEOL JSM-6390 LV SEM. In order to visualize fine hook morphology during SEM studies, prior to the graded ethanol series, posterior fragments were sonicated in distilled water for 15‒25 s at 30 kHz to remove hoods from the hooks. Specimens of Prionospio cristata (USNM 1171111) and Apoprionospio dayi (USNM 1273385) deposited in the USNM were dehydrated in a graded ethanol series, critical-point dried in CO2, mounted on stubs, sputter coated with carbon and gold-palladium (10 µm thick each), and examined in a Philips XL-30 SEM.

Results

Taxonomy

Phylum Annelida Lamarck, 1809

Order Spionida Fauchald, 1977

Family Spionidae Grube, 1850

Genus Prionospio Malmgren, 1867

Diagnosis (Peixoto & Paiva 2019): Prostomium anteriorly rounded or truncate, sometimes weakly incised, often with peaks, without frontal horns; subtriangular, rectangular or oval in shape, caruncle extending at least to chaetiger 1; eyespots present or absent; occipital antenna absent. Peristomium at least partially fused with chaetiger 1, often surrounding prostomium with free, flattened lateral wings. Parapodia of chaetiger 1 reduced; noto- and neuropodial lamellae largest in branchial region, reduced thereafter; notopodial lamellae often connected by low to high dorsal ridges or crests. Branchiae from chaetiger 2, chaetiger 3 or rarely absent. Branchiae limited to anterior chaetigers, 0–15 pairs, rarely more; branchiae all apinnate, all pinnate, or various combinations of both; pinnate branchiae with pinnules digitiform, not platelike; each branchia entirely free from dorsal lamella. Interparapodial pouches present or absent. Anterior chaetae limbate capillaries; posterior noto- and neuropodial hooded hooks present, bi-, tri-, or

135 multidentate, with secondary hood. Neuropodial sabre chaetae present or absent. Pygidium with one long dorsomedial cirrus and two shorter ventrolateral lobes, all three sometimes fused.

Prionospio cf. ehlersi Fauvel, 1928

(Figs 2–4E)

Material examined. Amb3 CAND5, 19°33'20,99" S 39°2'36,2" W, 420m (3 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (7 ind); Amb5 D6, 19°50'1,87" S 39°26'30,04" W, 1055m (1 ind); Amb6 CANWN5, 19°49'37,21" S 39°35'41,25" W, 410m (9 ind); Amb11 A5, 21°4'4,67" S 40°13'6,06" W, 415m (20 ind); Amb11 B5, 20°35'15,33" S 39°53'45,22" W, 415m (3 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (3 ind); Amb12 D5, 19°46'32,84" S 39°30'3,65" W, 416m (1 ind); Amb12 E5, 19°36'30,6" S 39°10'19,39" W, 360m (2 ind); Amb12 CAND5, 19°33'22,17" S 39°2'36,03" W, 414m (8 ind); Amb12 CANWN5, 19°49'36,9" S 39°35'42,69" W, 378m (2 ind).

Description: A large-sized Prionospio species, largest complete specimen 28 mm long, 1.1 mm wide at the widest point for 73 chaetigers (Fig 2). Body slightly dorsoventrally flattened on anterior region and cylindrical on middle and posterior regions, tapering towards the pygidium. Body color whitish to yellow in alcohol (Fig 2).

Prostomium narrowed, widened and rounded anteriorly, extending posteriorly as a narrow keel reaching the posterior margin of chaetiger 2. U-shaped nuchal organs reaching posterior margin of chaetiger 2. One pair of eyes or eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 2, 3A–D). Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae oval on notopodium and rounded on neuropodium, only slightly smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetiger 6, subtriangular from chaetiger 7 to chaetiger 10 and rounded from chaetiger 11 onwards, gradually reducing in size towards posterior region, becoming subtriangular and shifted laterally on last 9–15 chaetigers. Notopodial prechaetal lamellae as a low flap from chaetiger 2 to chaetigers 10–13, absent afterwards. Dorsal crests low from chaetiger 6 to chaetiger 29 (Figs 2, 3A–D).

Neuropodial postchaetal lamellae rounded throughout, well developed on anterior and median regions and present as a low flap on posterior region. Neuropodial prechaetal lamellae as a low

136 flap from chaetiger 2 to chaetigers 8–10, absent afterwards. Interneuropodial pouches from chaetiger 4, present on anterior half of the body (Figs 2, 3B–C).

Chaetae from notopodia organized in three rows of widely unilimbate and granulated capillaries; anterior row shortest and posterior row longest (Fig 4A). From middle region onwards, only two rows of chaetae, posterior row up to 1.4 times longer than anterior row. Chaetae from neuropodia organized in two rows of unilimbate and granulated capillaries (Fig 4B). Notopodial chaetae slightly longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 4C).

Hooks in notopodia starting from chaetigers 36–45, up to nine per fascicle, accompanied by 2–9 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 17–21, up to 15 per fascicle, accompanied, by 3–12 non-limbate capillaries. Hooks multidentate, with secondary teeth arranged in two rows above main tooth (Fig 4D). Secondary hood present. Sabre chaetae starting from chaetigers 18–21, up to three per fascicle. Sabre chaetae thick, non-limbate and intensely granulated throughout length (Fig 4E).

Branchiae present on chaetigers 2–5, cirriform and up to 2.5 times longer than notopodial postchaetal lamellae on chaetigers 2 and 5; robust, flattened and up to 1.5 times longer than notopodial postchaetal lamellae on chaetigers 3 and 4. Branchiae bearing pinnules on entire length on chaetiger 2 and apinnate on chaetigers 3–5 (Fig 2, 3A–D).

Pygidium bearing single long dorsal cirrus and two short ventral lobes.

Oocytes from chaetiger 24, measuring up to 150 μm.

Juvenile morphology: In juveniles, the interneuropodial pouches are absent.

Methyl green staining pattern: Prostomium and peristomium strongly stained; dorsal crests and margins of lamellae weakly stained.

Remarks. Prionospio ehlersi was described by Fauvel (1928) based on material collected on Morocco, eastern Atlantic Ocean. Since then, the species has been doubtfully and repeatedly recorded on tropical and subtropical areas on the deeper parts of the continental shelf, continuing to the shallow continental slope. The species was reported from the western Atlantic, Pacific Ocean, Indian Ocean, Antarctic Ocean and the Mediterranean Sea, with records ranging from full descriptions (Maciolek 1985; Mackie & Hartley 1990; Imajima 1990a; Dagli & Çinar 2009; Delgado-Blas & Salazar-Silva 2011; Delgado-Blas et al. 2018) to simple reports (Aguirrezabalaga & Ceberio 2005; Simon 2012; Çinar 2015).

Blake & Kudenov (1978), while describing material from Australia, described P. ehlersi specimens with interneuropodial pouches starting on more anterior chaetigers when compared to

137 the original description – starting on chaetiger 5 in Fauvel (1928) and starting on chaetiger 2 in Blake & Kudenov (1978), but made no comments on this discrepancy. Mackie & Hartley (1990) redescribed the species based on syntypes from Morocco and the Ivory Coast, described another Prionospio species bearing interneuropodial pouches – P. saccifera Mackie & Hartley, 1990 and also viewed the cosmopolitism of P. ehlersi as erroneous.

Delgado-Blas (2015) described an additional Prionospio species bearing interneuropodial pouches – P. crassumbranchiata Delgado-Blas, 2015. Mackie & Hartley (1990) and Delgado- Blas (2015) used characters such as prostomium shape, presence of eyes, lamellae morphology, branchiae morphology, starting chaetiger of interneuropodial pouches, capillary chaetae morphology, hooded hooks morphology and starting chaetiger of sabre chaetae to differentiate species.

Interneuropodial pouches have been reported in two additional species – Prionospio cirrifera Wirén, 1883 (in which pouches may be present or absent) and P. aluta Maciolek, 1985, although these species can be promptly distinguished from P. ehlersi based on prostomium shape - narrowed, widened and rounded anteriorly in P. ehlersi and subtriangular, truncate anteriorly on the two remaining species, as well as branchiae morphology and distribution – four pairs, pinnate only on the first pair in P. ehlersi, up to eight pairs of apinnate branchiae in P. cirrifera and up to six pairs of apinnate branchiae in P. aluta.

Specimens examined in this work are similar to the original description (Fauvel 1928) and to Mackie & Hartley (1990) redescription in having a prostomium of similar shape, same branchiae morphology and the presence of interneuropodial pouches, but differ in several characters, such as the starting chaetiger of interneuropodial pouches – from chaetiger 5 in Fauvel (1928) and from chaetiger 4 in Mackie & Hartley (1990) and this work; shape of notopodial postchaetal lamellae on posterior chaetigers – subtriangular and shifted laterally in this work and rounded, ventrally directed in Mackie & Hartley (1990) and shape of neuropodial postchaetal lamellae – rounded throughout in this work and rounded, gradually becoming elliptical in Mackie & Hartley (1990).

Differences were also noted on chaetae morphology and distribution, as the starting chaetiger of notopodial hooded hooks, from chaetigers 36–45 in this work and from chaetigers 25–32 in Mackie & Hartley (1990); maximum number of notopodial hooded hooks – up to nine per fascicle in this work and not mentioned (fewer than neuropodial hooks) in Mackie & Hartley (1990); starting chaetiger of neuropodial hooded hooks, from chaetigers 17–21 in this work and from chaetigers 18–20 in Mackie & Hartley (1990); maximum number of neuropodial hooded hooks – up to 15 per fascicle in this work and up to 10 per fascicle in Mackie & Hartley (1990); and starting chaetiger of sabre chaetae – from chaetigers 18–21 in this work and from chaetigers 17– 19 in Mackie & Hartley (1990), justifying the “cf.” adoption.

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Given the extent of recognized differences from the syntypes, and taking into account that some character discrepancies observed here were used to distinguish Prionospio ehlersi from P. saccifera and P. crassumbranchiata, it is highly likely that specimens from this work could represent a new species. However, it was decided not to erect a new taxon, since no comparative material was examined by the authors and the ontogenetic and intra-specific variation of P. ehlersi remains unknown. This issue might be resolved in the future if more material from the Brazilian coast becomes available.

Due to its combination of characters – prostomium narrowed, widened and rounded anteriorly; presence of interneuropodial pouches, sabre chaetae starting in more posterior chaetigers and branchiae bearing pinnules only on chaetiger 2, Prionospio cf. ehlersi can be readily distinguished from the remaining Brazilian Prionospio species.

Habitat: coarse silt to fine silt, 360–1055 m depth.

Distribution: Atlantic Ocean: Morocco (type-locality), Ivory Coast, Spain, Angola, Namibia, Mexico, USA, Suriname, Brazil (Paraná, São Paulo, Rio de Janeiro and Espírito Santo states); Mediterranean Sea: Cyprus, Turkey; Pacific Ocean: Japan, Vietnam, Mexico, Australia, Solomon Islands; Indian Ocean: Mozambique and Antarctic Ocean.

Prionospio cristata Foster, 1971

(Figs 5–7E)

Synonym list:

Prionospio (Prionospio) cristata Foster, 1971 (superseded subgenus)

Material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (6 ind); Amb1 Foz6, 19°54'56,16" S 39°56'40,83" W, 34m (5 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (2 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (1 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (1 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (5 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (12 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (14 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (4 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (2 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (9 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (7 ind); Amb2 Foz8, 19°44'44,06" S 39°46'32,01" W, 31m (1 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (6 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (2 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (2 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (12 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (5 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (12 ind);

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Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (13 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (16 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (2 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (8 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (2 ind); Amb7 B2, 20°34'45,78" S 40°11'30,74" W, 41m (4 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (16 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (3 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (2 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (12 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (57 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (33 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (21 ind); Amb7 F1, 18°42'56,07" S 39°31'28,8" W, 25m (2 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (17 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (20 ind); Amb7 G1, 18°32'3,6" S 39°23'16,91" W, 25m (5 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (22 ind); Amb13 G2, 18°36'32,45" S 39°9'32,83" W, 40m (18 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (8 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (8 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (2 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (11 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (7 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (1 ind); Amb14 F1, 18°42'54,91" S 39°31'28,99" W, 25m (2 ind); Amb14 F2, 18°52'31,35" S 39°8'41,34" W, 40m (3 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (1 ind).

Comparative material examined. Prionospio cristata USNM 43001 (holotype); USNM 43002 (2 paratypes); USNM 1171111 (74 specimens, topotypes).

Description: A medium-sized Prionospio species, largest complete specimen 8 mm long, 0.6 mm wide at the widest point for 73 chaetigers. Body slightly dorsoventrally flattened on anterior region and cylindrical on middle and posterior regions, tapering towards the pygidium. Body color whitish in alcohol (Fig 5).

Prostomium subtriangular and rounded anteriorly, extending posteriorly as a narrow and elevated keel reaching the posterior margin of chaetiger 2. Large U-shaped nuchal organs reaching the anterior margin of chaetiger 2. Two pairs of eyes in trapezoidal arrangement, posterior pair large and kidney-shaped. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 5, 6A–B). Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and oval on neuropodium, not significantly smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–7, square-shaped on chaetigers 8 and 9 and rounded from chaetiger 10 onwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent. Dorsal crests high on chaetigers 7 and 9 and low on chaetiger 8 (Figs 5, 6A, 6C).

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Neuropodial postchaetal lamellae square-shaped from chaetiger 2 to chaetigers 3–5 (depending on specimen size) and rounded afterwards, gradually reducing in size, present as a low flap from middle region onwards. Neuropodial prechaetal lamellae absent (Fig 6A–B).

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate and granulated capillaries (Fig 7A). Notopodial chaetae up to 1.5 times longer than neuropodial chaetae (Fig 7B). Towards the posterior region, capillaries progressively become elongate, non- limbate, non-granulated, thinner and less numerous (Fig 7C).

Hooks in notopodia starting from chaetigers 17–33, up to four per fascicle, accompanied by 1–4 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 10–13, up to eight per fascicle, accompanied by 1–8 non-limbate capillaries. Hooks multidentate, with six or eight secondary teeth arranged in two rows above main tooth (Figs 6D, 7D). Secondary hood absent. Sabre chaetae consistently from chaetiger 10. Sabre chaetae limbate and intensely granulated throughout length (Fig 7E).

Branchiae present on chaetigers 2–5, up to 3 times longer than notopodial lamellae on chaetigers 2 and 5 and up to 1.5 times longer than notopodial lamellae on chaetigers 3 and 4. Branchiae from chaetigers 3 and 4 triangular, robust, apinnate, ciliated and flattened; branchiae from chaetigers 2 and 5 cirriform, bearing pinnules throughout, except at the tips (Fig 6A–C).

Pygidium bearing a single long dorsal cirrus and two short ventral lobes.

Sperm plates from chaetiger 10.

Juvenile morphology: In juveniles, the prostomium is shorter, with a rounded appearance.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio cristata was described by Foster (1971), based on material collected on North Carolina (USA), northwestern Atlantic Ocean and also recorded on the Gulf of Mexico. The species was record on northeastern Brazil by Berlandi et al. (2012) and southeastern Brazil – adjacent to the study area of this work by Zalmon et al. (2013).

Sigvaldadóttir & Mackie (1993) considers all Prionospio species bearing branchiae on chaetigers 2–5, cirriform and pinnate on chaetigers 2 and 5 and flattened and apinnate on chaetigers 3 and 4 as part of the “Prionospio steenstrupi – group”, including P. cristata.

Regardless of species belonging to this group displaying highly variable morphological characters and despite the fact that P. steenstrupi was recorded on deeper parts of the continental shelf of Iceland, it is still recorded worldwide, including in Brazil. During visits to the Museu Nacional (Rio de Janeiro state), Museu de Zoologia da Universidade de São Paulo (São Paulo state) and Museu de Zoologia "Adão José Cardoso" (São Paulo state), all located in Brazil, no P. steenstrupi

141 specimens were found among Brazilian material, while the examined material corresponded to Prionospio cristata, P. cf. heterobranchia Moore, 1907, P. orensanzi, P. triangularis sp. nov. and P. posterocristata sp. nov. (pers. obs.).

Specimens recorded in this work are very similar to Foster (1971) description, while presenting small differences from Johnson (1984), Maciolek (1985) and Delgado-Blas (2015) descriptions. Our specimens agree with all descriptions in having a subtriangular prostomium; two pairs of eyes in trapezoidal arrangement; shape of lamellae from chaetiger 1; foliaceous notopodial postchaetal lamellae on anterior chaetigers; branchiae on chaetigers 2–5, cirriform and pinnate on chaetigers 2 and 5 and flattened and apinnate on chaetigers 3 and 4 and sabre chaetae consistently starting from chaetiger 10.

The only differences found from Foster (1971) description and our specimens are the presence of large nuchal organs accompanying the caruncle in our specimens, a character not mentioned by Foster (1971), but readily observed during reexamination of the holotype (pers. obs.) and also noted by Maciolek (1985) and Delgado-Blas (2015); and the starting chaetiger of notopodial hooded hooks – from chaetigers 17–33 in specimens from this work and from chaetigers 21-37 in Foster (1971) and the starting chaetiger of neuropodial hooded hooks – from chaetigers 10–13 in specimens from this work and from chaetigers 11–12 in Foster (1971). Subtle variations on the starting chaetiger of hooded hooks have been reported by Johnson (1984), Maciolek (1985) and Delgado-Blas (2015) and could be caused by ontogenetic variability or due to the small number of specimens examined to describe Prionospio cristata.

Among Brazilian Prionospio species, P. cristata is most similar to P. triangularis sp. nov. and P. posterocristata sp. nov. in having a subtriangular and anteriorly rounded prostomium; branchiae on chaetigers 2–5, cirriform and pinnate on chaetigers 2 and 5 and flattened and apinnate on chaetigers 3 and 4; presence of dorsal crests and sabre chaetae consistently starting from chaetiger 10.

All three may co-occur on shallow depths, but can be distinguished based on dorsal crests distribution and morphology, high on chaetigers 7 and 9 and low on chaetiger 8 in P. cristata, high on chaetiger 7 and low from chaetiger 8 to chaetigers 13–18 in P. triangularis sp. nov. and low from chaetigers 11–12 to chaetigers 16–18 in P. posterocristata sp. nov.; lamellae morphology from chaetiger 1, rounded on notopodium and oval on neuropodium in P. cristata, square-shaped on notopodium and subtriangular on neuropodium in P. triangularis sp. nov. and rounded on notopodium and digitiform on neuropodium in P. posterocristata sp. nov. and shape of neuropodial postchaetal lamellae from chaetiger 2, rounded in P. cristata, triangular with greatly elongated and ventrally pointed tip in P. triangularis sp. nov. and triangular, slightly elongated ventrally and bearing a rounded tip in P. posterocristata sp. nov.

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All three species can also be distinguished by the starting chaetiger of notopodial hooded hooks, from chaetigers 17–33 in P. cristata, from chaetigers 24–54 in P. triangularis sp. nov. and from chaetigers 25–32 in P. posterocristata sp. nov. and starting chaetiger of neuropodial hooded hooks, from chaetigers 10–13 in P. cristata, from chaetigers 13–20 in P. triangularis sp. nov. and from chaetigers 12–17 in P. posterocristata sp. nov.

Habitat: coarse sand to fine sand, 25–54 m depth.

Distribution: Atlantic Ocean: USA (type-locality), Mexico, Puerto Rico, Trinidad and Tobago and Brazil (Rio de Janeiro, Espírito Santo, Bahia, Sergipe and Alagoas states).

Prionospio brevipinnulata sp. nov.

(Figs 8–10F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 02 Dec 2011 to 02 Feb 2012, 1300m, MNRJP XXXX. Paratypes: Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 02 Dec 2011 to 02 Feb 2012, 1335m, MNRJP XXXX (2 ind), MZUSP XXXX (2 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m, 06 Jun 2013 to 17 Jul 2013 ZUEC- POL XXXX (4 ind).

Additional material examined. Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 1035m (8 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (1 ind); Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 2426m (1 ind); Amb3 CAND5, 19°33'20,99" S 39°2'36,2" W, 420m (2 ind); Amb3 CAND6, 19°37'50,08" S 39°3'56,16" W, 970m (13 ind); Amb3 CAND7, 19°42'18,42" S 39°5'52,41" W, 1307m (7 ind); Amb4 A10, 21°11'4,9" S 38°28'2,78" W, 3006m (1 ind); Amb4 B8, 20°41'33,45" S 39°35'14,76" W, 1914m (7 ind); Amb4 E9, 20°35'50,48" S 38°27'7,64" W, 2494m (1 ind); Amb4 F6, 19°52'56,9" S 38°35'8,88" W, 1022m (4 ind); Amb4 F7, 20°4'8,18" S 38°31'27,32" W, 1302m (3 ind); Amb4 F8, 20°16'35,72" S 38°27'18,98" W, 1902m (6 ind); Amb4 F9, 20°29'3,31" S 38°23'15,5" W, 2504m (5 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (2 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (6 ind); Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 1300m (5 ind); Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 1889m (3 ind); Amb5 A9, 21°9'40,3" S 38°52'25,04" W, 2498m (1 ind); Amb5 B5, 20°35'16,23" S 39°53'47,1" W, 410m (1 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (9 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (10 ind); Amb5 B9, 20°54'44,14" S 38°56'10,72" W, 2519m (6 ind); Amb5 C5, 20°14'19,45" S 39°48'36,67" W, 381m (4 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (5 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (7 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (13 ind); Amb5 C9, 20°48'39,87" S 38°45'23,86" W, 2496m (1 ind); Amb5 D6, 19°50'1,87" S 39°26'30,04" W, 1055m (6 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W,

143

1335m (4 ind); Amb5 D8, 20°8'42,82" S 39°7'29,5" W, 1922m (2 ind); Amb5 D9, 20°34'36,32" S 38°40'53,6" W, 2460m (3 ind); Amb5 E8, 20°15'59,97" S 38°40'53,86" W, 1887m (2 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (15 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (20 ind); Amb8 E7, 19°47'5,96" S 39°3'11,96" W, 1258m (1 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (2 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (11 ind); Amb8 G8, 19°3'45,82" S 37°47'28,26" W, 1928m (3 ind); Amb11 A6, 21°4'37,64" S 40°8'32,68" W, 1015m (3 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (1 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (6 ind); Amb11 B6, 20°36'1,61" S 39°51'39,15" W, 1003m (5 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (7 ind); Amb11 B8, 20°41'33,93" S 39°35'22,06" W, 1908m (4 ind); Amb11 B9, 20°54'43,79" S 38°56'10,85" W, 2520m (3 ind); Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 1040m (4 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (5 ind); Amb11 C8, 20°25'13,22" S 39°27'19,49" W, 1920m (5 ind); Amb11 C9, 20°48'37,26" S 38°45'28,85" W, 2465m (5 ind); Amb11 C10, 20°57'57,99" S 38°27'52,41" W, 2996m (1 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (6 ind); Amb11 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (1 ind); Amb11 E8, 20°15'55,63" S 38°40'45,57" W, 1892m (8 ind); Amb11 E9, 20°35'51,99" S 38°27'13,04" W, 2501m (1 ind); Amb11 F6, 19°52'52,59" S 38°35'10,48" W, 1021m (1 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (7 ind); Amb11 F8, 20°16'38,17" S 38°27'26,52" W, 1904m (6 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (3 ind); Amb11 F10, 20°46'17,79" S 38°17'16,01" W, 3002m (2 ind); Amb12 D6, 19°50'6,01" S 39°26'34,62" W, 1050m (5 ind); Amb12 D7, 19°54'4,77" S 39°22'29,46" W, 1335m (3 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (6 ind); Amb12 E7, 19°47'2,44" S 39°3'14,62" W, 1250m (5 ind); Amb12 G7, 19°3'30,62" S 37°48'46,66" W, 1347m (2 ind); Amb12 G8, 19°3'39,78" S 37°47'39,35" W, 1867m (3 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (2 ind); Amb12 CAND6, 19°37'45,14" S 39°3'58,75" W, 1050m (1 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (1 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (20 ind).

Description: A small-sized Prionospio species, largest complete specimen 5 mm long, 0.3 mm wide at the widest point for 40 chaetigers, holotype 4.5 mm long, 0.25 mm wide at the widest point for 34 chaetigers. Body cylindrical throughout, tapering towards the pygidium. Body color whitish in alcohol (Fig 8).

Prostomium broadly rounded, extending posteriorly as a narrow and slightly elevated keel reaching the posterior margin of chaetiger 2. U-shaped nuchal organs reaching anterior margin of chaetiger 2 (Fig 9A–C). Eyes absent. Prostomial peaks absent. Peristomium surrounding

144 prostomium and partially fused to the first chaetiger, lateral wings absent (Fig 8, 9B–C). Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae low and rounded on both rami, both much smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–6, quadrangular from chaetiger 7 to chaetigers 10–14 (depending on specimen size) (Fig 9A), gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent. Dorsal crests absent (Fig 8, 9A).

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetigers 8–11, abruptly reduced to a low flap afterwards (Fig 9D). Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of wide, narrowly unilimbate and intensely granulated capillaries (Fig 10A). On chaetigers 2–4, neuropodial capillaries extremely long, up to 2.5 times longer than capillaries of chaetiger 5 (Fig 10B). On remaining chaetigers, neuropodial chaetae slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 10C).

Hooks in notopodia starting from chaetigers 18–32, up to six per fascicle, accompanied by 1–4 long non-limbate capillaries. Hooks in neuropodia starting from chaetigers 10–16, up to 10 per fascicle, accompanied, by 1–8 long non-limbate capillaries. Hooks tridentate, with two secondary teeth arranged side by side above main tooth (Fig 10D). Secondary hood absent. Sabre chaetae consistently from chaetiger 10. Sabre chaetae non-limbate and intensely granulated throughout length, strongly curved on first 2–4 chaetigers (Fig 10E).

Branchiae present on chaetigers 2–5, twice as long as notopodial lamellae on chaetigers 2 and 5 and about the same length of notopodial lamellae on chaetigers 3 and 4. Branchiae from chaetigers 3–4 robust and flattened (Fig 9B–C); branchiae from chaetigers 2 and 5 cirriform, bearing few and very short pinnules on lower half of branchial stem (Fig 10F).

Pygidium bearing a single long dorsal cirrus and two short and thin ventral cirri.

Oocytes from chaetiger 9, measuring up to 80 μm.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio brevipinnulata sp. nov. is similar to P. fauchaldi Maciolek, 1985 and P. alexandrae sp. nov.in having elongated capillaries on anterior neuropodia, although both species can be readily distinguished from P. brevipinnulata sp. nov. based on the prostomium shape –

145 broadly rounded in P. brevipinnulata sp. nov. and rectangular in P. fauchaldi and alexandrae sp. nov.; branchiae morphology – pairs 2 and 5 pinnate in P. brevipinnulata sp. nov. and all pairs apinnate in P. fauchaldi and alexandrae sp. nov. and distribution of elongated capillaries on anterior neuropodia – on chaetigers 2–4 in P. brevipinnulata sp. nov. and P. alexandrae sp. nov. and only on chaetiger 3 in P. fauchaldi (Maciolek, 1985).

The morphology of hooded hooks – tridentate, is uncommon in Prionospio, as most species bear multidentate hooded hooks. Tridentate hooded hooks are known only from Prionospio tridentata Blake & Kudenov, 1978, described from Australia, P. hermesia Neal & Paterson in Paterson et al., 2016, described from Portugal and Prionospio fosterae Peixoto & Paiva (ms), from the same locality of this work. Prionospio brevipinnulata sp. nov. can be distinguished from the aforementioned species by the prostomium shape – broadly rounded in P. brevipinnulata sp. nov., trapezoidal in P. tridentata, oval in P. hermesia and narrow, rounded anteriorly in P. fosterae; lack of dorsal crests in P. brevipinnulata sp. nov. (also absent in P. fosterae); starting chaetiger of neuropodial hooded hooks – from chaetiger 10–16 in P. brevipinnulata sp. nov., chaetiger 19 in P. tridentata, chaetigers 13–14 in P. hermesia and chaetigers 13‒16 in P. fosterae; starting chaetiger of sabre chaetae – chaetiger 10 in P. brevipinnulata sp. nov., chaetiger 11 in P. tridentata and lacking in both P. hermesia and P. fosterae and branchiae morphology – four pairs, pinnate on chaetigers 2 and 5 in P. brevipinnulata sp. nov., four pairs, pinnate only on chaetiger 5 in P. tridentata, two apinnate pairs in P. hermesia and up to 11 pairs of apinnate branchiae in P. fosterae (Blake & Kudenov, 1978, Paterson et al., 2016; Peixoto & Paiva, ms).

The lack of dorsal crests is also unusual in Prionospio, observed only in P. dubia Day, 1961, P. jonatani Delgado-Blas (2015), P. hartmanae Peixoto & Paiva (ms), P. absensi sp. nov., P. alexandrae sp. nov. and P. biancoi sp. nov., Prionospio brevipinnulata sp. nov. can be distinguished from the aforementioned species in the prostomium shape, broadly rounded in P. brevipinnulata sp. nov., rounded and posteriorly narrowed in P. dubia, P. jonatani and P. absensi sp. nov., narrow and anteriorly rounded in P. hartmanae, rectangular in P. alexandrae sp. nov. and rectangular in P. biancoi sp. nov.; branchiae morphology – four pairs, with very short pinnules on chaetigers 2 and 5 in P. brevipinnulata sp. nov., four pairs, with long pinnules on chaetigers 2 and 5 (branchiae from chaetiger 2 extremely long) in P. dubia, P. jonatani and P. absensi sp. nov., up to 11 pairs of apinnate branchiae in P. hartmanae, two pairs of apinnate branchiae in P. alexandrae sp. nov. and four pairs of apinnate branchiae in P. biancoi sp. nov.; hook morphology – tridentate in P. brevipinnulata sp. nov. and multidentate on all remaining species and the starting chaetiger of sabre chaetae – consistently from chaetiger 10 in P. brevipinnulata sp. nov., from chaetiger 18 in P. dubia, from chaetigers 14–18 in P. jonatani, from chaetigers 10–16 in P. absensi sp. nov., from chaetigers 13–14 in P. hartmanae, lacking in P.

146 alexandrae sp. nov. and from chaetigers 14–20 in P. biancoi sp. nov. (Day 1961; Sigvaldadóttir & Mackie 1993; Delgado-Blas 2015; Peixoto & Paiva ms).

This species was very common on the continental slope, even at greater depths and was also collected on the states of Sergipe and Alagoas (pers. Obs.).

Etymology. The specific epithet, brevipinnulata (“brevi”, Latin for short), refers to the very short pinnules on branchiae.

Habitat: very fine sand to clay, 381–3006 m depth.

Distribution: Brazil (Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states), Atlantic Ocean.

Prionospio capixaba sp. nov.

(Figs 11–13E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb3 E5, 19°36'26,24" S 39°10'17,35" W, 02 Dec 2011 to 02 Feb 2012, 392m, MZUSP XXXX. Paratypes: Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 02 Dec 2011 to 02 Feb 2012, 153m, MZUSP XXXX (4 ind), MNRJP XXXX (4 ind), ZUEC-POL XXXX (4 ind).

Additional material examined. Amb3 E5, 19°36'26,24" S 39°10'17,35" W, 392m (33 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (19 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (2 ind); Amb5 B5, 20°35'16,23" S 39°53'47,1" W, 410m (1 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (5 ind); Amb5 C5, 20°14'19,45" S 39°48'36,67" W, 381m (83 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (1 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (49 ind); Amb6 D5, 19°46'34,99" S 39°30'4,65" W, 428m (1 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (202 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (1 ind); Amb6 CANWN5, 19°49'37,21" S 39°35'41,25" W, 410m (19 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (6 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (2 ind); Amb11 A5, 21°4'4,67" S 40°13'6,06" W, 415m (5 ind); Amb11 B5, 20°35'15,33" S 39°53'45,22" W, 415m (11 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (1 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (43 ind); Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 1040m (3 ind); Amb12 D5, 19°46'32,84" S 39°30'3,65" W, 416m (3 ind); Amb12 E5, 19°36'30,6" S 39°10'19,39" W, 360m (23 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (2 ind); Amb12 F5, 19°34'20,47" S 38°41'19,8" W, 449m (15 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (1 ind); Amb12 CAND5, 19°33'22,17" S 39°2'36,03" W, 414m (7 ind); Amb12 CANWN5, 19°49'36,9" S 39°35'42,69" W, 378m (3 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (2 ind).

147

Description: A small-sized Prionospio species, largest complete specimen 7 mm long, 0.3 mm wide at the widest point for 77 chaetigers, holotype 6 mm long, 0.25 mm wide at the widest point for 58 chaetigers. Body slightly dorsoventrally flattened on anterior and middle regions and cylindrical on posterior region, tapering towards the pygidium. Body color whitish in alcohol (Fig 11).

Prostomium narrow, rounded anteriorly, extending posteriorly as a narrow and slightly elevated keel reaching the posterior margin of chaetiger 2 (Fig 12A–B). U-shaped nuchal organs reaching anterior margin of chaetiger 2. One pair of minute eyes or eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 11, 12A–C). Grooved palps reaching up to chaetiger 8, lost in most specimens.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 3–5 (depending on specimen size), rounded from chaetigers 4–6 onwards and gradually reducing in size towards posterior region, present as a low flap from middle region onwards (Fig 12A). Notopodial prechaetal lamellae absent. Dorsal crests low from chaetiger 8 to chaetiger 13 (Figs 11, 12A–B).

Neuropodial postchaetal lamellae rounded and elongated ventrally on chaetiger 2, rounded from chaetiger 3 to chaetigers 9–12 and abruptly reduced to a low flap afterwards (Figs 11, 12D). Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of unilimbate and intensely granulated capillaries. Notopodial chaetae (Fig 13A) up to twice as long as neuropodial chaetae (Fig 13B). Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 13C).

Hooks in notopodia starting from chaetigers 19–50, up to five per fascicle, accompanied by 1–4 long non-limbate capillaries. Hooks in neuropodia starting from chaetigers 10–14, up to six per fascicle, accompanied, by 1–6 long non-limbate capillaries. Hooks multidentate, with secondary teeth arranged in two rows above main tooth (Fig 13D). Secondary hood absent. Sabre chaetae starting from chaetigers 10–12. Sabre chaetae narrowly unilimbate and granulated throughout length (Fig 13E).

Branchiae present on chaetigers 2–5, twice as long as notopodial lamellae on chaetigers 2 and 5 and slightly longer than notopodial lamellae on chaetigers 3 and 4. Branchiae from chaetigers 3–

148

4 robust and flattened; branchiae from chaetigers 2 and 5 cirriform, bearing pinnules on entire length.

Pygidium bearing single long and thin dorsal cirrus and two short ventral cirri.

Oocytes from chaetigers 10–11, measuring up to 100 μm.

Juvenile morphology: In juveniles, notopodial postchaetal lamella is absent on chaetiger 1.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio capixaba sp. nov. is similar to P. brevipinnulata sp. nov. in having an overlap on the starting chaetiger of sabre chaetae, as well as notopodial and neuropodial hooded hooks and same lamellae morphology on chaetiger 1, although species can be distinguished based on the presence of dorsal crests in P. capixaba sp. nov., hook morphology – multidentate in P. capixaba sp. nov. and tridentate in P. brevipinnulata sp. nov., lack of elongated neuropodial capillaries on anterior chaetigers in P. capixaba sp. nov. and morphology of pinnate branchiae, with pinnules on entire branchial stem in P. capixaba sp. nov. and short pinnules present only on the lower half of branchial stem in P. brevipinnulata sp. nov.

The species is also similar to P. posterocristata sp. nov., P. quadrilamellata sp. nov., Prionospio cf. delta Hartman, 1965, Prionospio corrugatus Peixoto & Paiva, ms, Prionospio mutatus Peixoto & Paiva, ms, Prionospio acutus Peixoto & Paiva, ms and Prionospio kinbergi Peixoto & Paiva, ms in having dorsal crests beginning after chaetiger 7. However, P. capixaba sp. nov. can be readily distinguished from Prionospio corrugatus sp. nov., Prionospio mutatus sp. nov., Prionospio acutus sp. nov. and Prionospio kinbergi sp. nov. in having only four pairs of branchiae, first and fourth pair bearing pinnules, while the aforementioned species bears 8–11 pairs of apinnate branchiae. As for P. posterocristata sp. nov. and P. quadrilamellata sp. nov., species can be separated based on the prostomium shape – broadly rounded in P. capixaba sp. nov., subtriangular in P. posterocristata sp. nov. and rectangular in P. quadrilamellata sp. nov.; branchiae morphology – four pairs of branchiae, first and fourth pair bearing pinnules in both P. capixaba sp. nov. and P. posterocristata sp. nov. and all four pairs apinnate in P. quadrilamellata sp. nov.; dorsal crests distribution – from chaetigers 8–13 in P. capixaba sp. nov., from chaetigers 9–13 in P. quadrilamellata sp. nov. and from chaetigers 11–12 to chaetigers 16–18 in P. posterocristata sp. nov. and starting chaetiger of sabre chaetae – from chaetigers 10–12 in P. capixaba sp. nov., from chaetigers 9–11 in P. quadrilamellata sp. nov. and consistently from chaetiger 10 in P. posterocristata sp. nov. (Peixoto & Paiva, ms).

Etymology. The species epithet, “capixaba” is the common denomination of natives from the state of Espírito Santo, southeastern Brazil, where the specimens were collected.

149

Habitat: very fine sand to fine silt, 153–1324 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Prionospio quadrilamellata sp. nov.

(Figs 14–16F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 02 Dec 2011 to 02 Feb 2012, 1040m, MNRJP XXXX. Paratypes: Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 06 Jun 2013 to 17 Jul 2013, 1355m, MNRJP XXXX (2 ind), MZUSP XXXX (2 ind), ZUEC-POL XXXX (1 ind).

Additional material examined. Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 1035m (1 ind); Amb3 CAND6, 19°37'50,08" S 39°3'56,16" W, 970m (1 ind); Amb4 F6, 19°52'56,9" S 38°35'8,88" W, 1022m (1 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (2 ind); Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 1300m (3 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (4 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (1 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (2 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (1 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 1335m (1 ind); Amb8 E7, 19°47'5,96" S 39°3'11,96" W, 1258m (1 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (2 ind); Amb11 B6, 20°36'1,61" S 39°51'39,15" W, 1003m (1 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (2 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (3 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (1 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (2 ind); Amb12 D6, 19°50'6,01" S 39°26'34,62" W, 1050m (2 ind); Amb12 D7, 19°54'4,77" S 39°22'29,46" W, 1335m (1 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (1 ind); Amb12 F5, 19°34'20,47" S 38°41'19,8" W, 449m (1 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (2 ind).

Description: A small-sized Prionospio species, largest complete specimen 5 mm long, 0.2 mm wide at the widest point for 43 chaetigers, holotype 3.5 mm long, 0.15 mm wide at the widest point for 33 chaetigers. Body slightly dorsoventrally flattened on anterior region and cylindrical on middle and posterior regions (Fig 15B), tapering towards the pygidium. Body color whitish in alcohol (Fig 14).

Prostomium rectangular, narrow, truncated on the anterior margin, extending posteriorly as a narrow keel reaching the posterior margin of chaetiger 1 (Fig15A–C). U-shaped nuchal organs reaching anterior margin of chaetiger 1. One or two pairs of small eyes present. Prostomial peaks

150 absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 14, 15A–C). Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and digitiform in neuropodium, both much smaller than lamellae on succeeding chaetigers, especially in neuropodium. Prechaetal lamellae absent (Fig 15C).

Notopodial postchaetal lamellae quadrangular from chaetiger 2 to chaetigers 4–6 (depending on specimen size), triangular from chaetigers 5–7 to chaetigers 6–9 and rounded afterwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent. Dorsal crests low from chaetiger 9 to chaetiger 13 (Figs 14, 15A).

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetigers 5–7, abruptly reduced to a low flap afterwards. Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate and granulated capillaries (Fig 16A). Notopodial chaetae slightly longer than neuropodial chaetae (Fig 16B). Towards the posterior region, capillaries progressively become elongate, non-limbate, non- granulated, thinner and less numerous (Fig 16C).

Hooks in notopodia starting from chaetigers 20–32, up to five per fascicle, accompanied by 1–5 non-limbate capillaries (Fig 15D). Hooks in neuropodia starting from chaetigers 9–13, up to seven per fascicle, accompanied, by 1–7 non-limbate capillaries. Hooks multidentate, with secondary teeth arranged in two rows above main tooth (Fig 16D). Secondary hood absent. Sabre chaetae starting from chaetigers 9–11 (Fig 15D). Sabre chaetae non-limbate and granulated throughout length, robust on first 2–3 chaetigers and thinner on remaining chaetigers (Fig 16E).

Branchiae present on chaetigers 2–5, about the same length of notopodial lamellae on chaetigers 2–4 and slightly shorter than notopodial lamellae on chaetiger 5. All branchiae apinnate, robust and flattened (Fig 16F).

Pygidium bearing a single thin and long dorsal cirrus and two short ventral cirri.

Juvenile morphology: In juveniles, neuropodial postchaetal lamellae is absent on chaetiger 1.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio quadrilamellata sp. nov. is similar to P. fauchaldi, P. corrugatus, P. alexandrae sp. nov., P. vallensis, P. kaplani and P. amarsupiata in bearing a rectangular prostomium, although species can be distinguished based on branchiae morphology and distribution – four pairs of flattened and apinnate branchiae in P. quadrilamellata sp. nov., up to six pairs of apinnate branchiae in both P. fauchaldi and P. corrugatus, two pairs of apinnate

151 branchiae in both P. alexandrae sp. nov. and P. kaplani, four pairs of wrinkled branchiae in P. vallensis and four pairs of branchiae, first (pinnate) and fourth (apinnate) pairs cirriform in P. amarsupiata (Maciolek 1985; Paterson et al. 2016; Peixoto & Paiva ms).

The species also differs from the aforementioned species in the distribution of dorsal crests – on chaetigers 9–13 in P. quadrilamellata sp. nov., on chaetigers 8 – 11 in P. fauchaldi, from chaetigers 8–9 to chaetigers 14–15 in P. corrugatus, absent in P. alexandrae sp. nov., from chaetiger 6 to beyond chaetiger 20 in P. vallensis, on chaetigers 7–17 in P. kaplani and on chaetigers 5–20 in P. amarsupiata. Species can be further distinguished based on the starting chaetiger of sabre chaetae – from chaetigers 9–11 in P. quadrilamellata sp. nov., not mentioned in P. fauchaldi, from chaetigers 10–17 in P. corrugatus, absent in P. alexandrae sp. nov., chaetiger 10 in P. vallensis, not observed in P. kaplani and from chaetigers 18–19 in P. amarsupiata. Unfortunately, characters such as the starting chaetiger of notopodial and neuropodial hooded hooks presented a considerable overlap among species, not being particularly useful in separating species (Maciolek 1985; Paterson et al. 2016; Peixoto & Paiva ms).

At last, Prionospio quadrilamellata sp. nov. is also similar to P. cinthyae (Peixoto & Paiva ms) and P. biancoi sp. nov. in bearing four pairs of apinnate branchiae, although species can be easily distinguished due to prostomium shape – rectangular in P. quadrilamellata sp. nov. and P. biancoi sp. nov. and triangular in P. cinthyae, branchiae morphology – all pairs flattened in P. quadrilamellata sp. nov., first and fourth pairs wrinkled in P. cinthyae and first and last pair cirriform in P. biancoi sp. nov.; starting chaetiger of notopodial hooded hooks – chaetigers 20– 32 in P. quadrilamellata sp. nov., chaetigers 21–33 in P. cinthyae and chaetiger 67 in P. biancoi sp. nov.; starting chaetiger of neuropodial hooded hooks – chaetigers 9–13 in P. quadrilamellata sp. nov., chaetigers 10–12 in P. cinthyae and chaetigers 16–22 in P. biancoi sp. nov. and starting chaetiger of sabre chaetae – chaetigers 9–11 in P. quadrilamellata sp. nov., consistently from chaetiger 10 in P. cinthyae and chaetigers 14–20 in P. biancoi sp. nov. (Peixoto & Paiva ms).

Etymology. The specific epithet, quadrilamellata, refers to the shape of notopodial postchaetal lamellae from the anterior region (quadri is term for quadrangular, while lamellata refers to the lamellae).

Habitat: coarse silt to fine silt, 395–1926 m depth.

Distribution: Brazil (Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states), Atlantic Ocean.

152

Prionospio alexandrae sp. nov.

(Figs 17–19F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 02 Dec 2011 to 02 Feb 2012, 2426m, ZUEC-POL XXXX. Paratypes: Amb4 F10, 20°46'23,65" S 38°17'17,65" W, 02 Dec 2011 to 02 Feb 2012, 3004m, MNRJP XXXX (3 ind), MZUSP XXXX (3 ind), ZUEC-POL XXXX (3 ind).

Additional material examined. Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 2426m (27 ind); Amb3 G10, 19°3'55,8" S 37°45'8,27" W, 2993m (12 ind); Amb4 A9, 21°9'35,19" S 38°52'4,93" W, 2502m (2 ind); Amb4 A10, 21°11'4,9" S 38°28'2,78" W, 3006m (4 ind); Amb4 B8, 20°41'33,45" S 39°35'14,76" W, 1914m (8 ind); Amb4 B10, 21°4'40,47" S 38°26'13,68" W, 3007m (3 ind); Amb4 C10, 20°59'0,28" S 38°28'36,56" W, 2997m (3 ind); Amb4 D10, 20°53'33,64" S 38°21'17,35" W, 3004m (19 ind); Amb4 E9, 20°35'50,48" S 38°27'7,64" W, 2494m (29 ind); Amb4 E10, 20°49'23,58" S 38°17'11,07" W, 2997m (13 ind); Amb4 F7, 20°4'8,18" S 38°31'27,32" W, 1302m (2 ind); Amb4 F8, 20°16'35,72" S 38°27'18,98" W, 1902m (23 ind); Amb4 F9, 20°29'3,31" S 38°23'15,5" W, 2504m (30 ind); Amb4 F10, 20°46'23,65" S 38°17'17,65" W, 3004m (17 ind); Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 1889m (9 ind); Amb5 A9, 21°9'40,3" S 38°52'25,04" W, 2498m (7 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (2 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (5 ind); Amb5 B9, 20°54'44,14" S 38°56'10,72" W, 2519m (21 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (2 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (14 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (11 ind); Amb5 C9, 20°48'39,87" S 38°45'23,86" W, 2496m (9 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 1335m (3 ind); Amb5 D8, 20°8'42,82" S 39°7'29,5" W, 1922m (29 ind); Amb5 D9, 20°34'36,32" S 38°40'53,6" W, 2460m (7 ind); Amb5 E8, 20°15'59,97" S 38°40'53,86" W, 1887m (17 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (2 ind); Amb8 E7, 19°47'5,96" S 39°3'11,96" W, 1258m (5 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (2 ind); Amb8 G8, 19°3'45,82" S 37°47'28,26" W, 1928m (24 ind); Amb11 A5, 21°4'4,67" S 40°13'6,06" W, 415m (1 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (3 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (14 ind); Amb11 A9, 21°9'39,38" S 38°52'7,25" W, 2502m (6 ind); Amb11 A10, 21°10'59,11" S 38°28'4,99" W, 2987m (3 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (2 ind); Amb11 B8, 20°41'33,93" S 39°35'22,06" W, 1908m (9 ind); Amb11 B9, 20°54'43,79" S 38°56'10,85" W, 2520m (14 ind); Amb11 B10, 21°4'33,4" S 38°26'19,34" W, 3001m (8 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (3 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (19 ind); Amb11 C8, 20°25'13,22" S 39°27'19,49" W, 1920m (6 ind); Amb11 C9, 20°48'37,26" S 38°45'28,85" W, 2465m (18 ind); Amb11 C10, 20°57'57,99" S

153

38°27'52,41" W, 2996m (7 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (30 ind); Amb11 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (9 ind); Amb11 D10, 20°53'28,94" S 38°21'21,75" W, 3000m (14 ind); Amb11 E8, 20°15'55,63" S 38°40'45,57" W, 1892m (17 ind); Amb11 E9, 20°35'51,99" S 38°27'13,04" W, 2501m (9 ind); Amb11 E10, 20°49'19,95" S 38°17'8,76" W, 2994m (12 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (5 ind); Amb11 F8, 20°16'38,17" S 38°27'26,52" W, 1904m (25 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (27 ind); Amb11 F10, 20°46'17,79" S 38°17'16,01" W, 3002m (13 ind); Amb12 D6, 19°50'6,01" S 39°26'34,62" W, 1050m (2 ind); Amb12 D7, 19°54'4,77" S 39°22'29,46" W, 1335m (9 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (1 ind); Amb12 E7, 19°47'2,44" S 39°3'14,62" W, 1250m (3 ind); Amb12 G7, 19°3'30,62" S 37°48'46,66" W, 1347m (1 ind); Amb12 G8, 19°3'39,78" S 37°47'39,35" W, 1867m (23 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (33 ind); Amb12 G10, 19°3'10,22" S 37°45'28,45" W, 2950m (31 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (5 ind).

Description: A small-sized Prionospio species, largest complete specimen 7 mm long, 0.25 mm wide at the widest point for 52 chaetigers, holotype 5 mm long, 0.15 mm wide at the widest point for 40 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium. Body color whitish in alcohol (Fig 17).

Prostomium rectangular, narrow, truncated on the anterior margin, extending posteriorly as a narrow keel reaching the anterior margin of chaetiger 2 (Fig 18A–B). U-shaped nuchal organs reaching posterior margin of chaetiger 1. Eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 17, 18A–D). Grooved palps reaching up to chaetiger 6, lost in most specimens. Palps bearing minute papillae along groove margin (Fig 18E–F).

Chaetiger 1 with only a few chaetae in both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae absent on both rami. Prechaetal lamellae absent (Fig 18D).

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 6–10 (depending on specimen size), longest on chaetiger 3 (twice as long as surrounding lamellae) and rounded afterwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent. Dorsal crests absent (Figs 17, 18A, C).

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetigers 8–10, abruptly reduced to a low flap afterwards. Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate and lightly granulated capillaries (Figs 19A, B). On chaetigers 2–4, neuropodial capillaries extremely long,

154 up to twice as long as capillaries from chaetiger 5 (Fig 19C). On remaining chaetigers, neuropodial chaetae slightly shorter than notopodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 19D).

Hooks in notopodia starting from chaetigers 26–46, up to six per fascicle, accompanied by 1–6 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 10–20, up to nine per fascicle, accompanied, by 1–8 non-limbate capillaries. Hooks multidentate, with secondary teeth arranged in two rows above main tooth (Fig 19E). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae present on chaetigers 2 and 3, up to twice as long as notopodial lamellae on chaetiger 2 and shorter than notopodial lamellae on chaetiger 3. Both branchiae cirriform, apinnate and sparsely ciliated (Figs 18B, D, 19F).

Pygidium rounded, lacking cirri.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio alexandrae sp. nov. is similar to P. fauchaldi, P. corrugatus, P. quadrilamellata sp. nov., P. vallensis, P. kaplani and P. amarsupiata in bearing a rectangular prostomium, although P. alexandrae sp. nov. differs from all aforementioned species (except P. kaplani, which is discussed below) in having only two pairs of branchiae, on chaetigers 2 and 3, while P. quadrilamellata sp. nov. bears four pairs of flattened and apinnate branchiae, both P. fauchaldi and P. corrugatus bears up to six pairs of apinnate branchiae, P. vallensis bears four pairs of wrinkled branchiae and P. amarsupiata bears four pairs of branchiae, first (pinnate) and fourth (apinnate) pairs cirriform (Maciolek 1985; Paterson et al. 2016; Peixoto & Paiva ms).

Species can also be distinguished based on the lack of dorsal crests in P. alexandrae sp. nov., present on chaetigers 8 – 11 in P. fauchaldi, from chaetigers 8–9 to chaetigers 14–15 in P. corrugatus, on chaetigers 9–13 in P. quadrilamellata sp. nov., from chaetiger 6 to beyond chaetiger 20 in P. vallensis and on chaetigers 5–20 in P. amarsupiata. Species can further be distinguished by the lack of sabre chaetae in Prionospio alexandrae sp. nov., present on all species – except for P. kaplani, in which sabre chaetae were not observed in any fragment. (Maciolek 1985; Paterson et al. 2016).

The occurrence of only two pairs of branchia is unusual among Prionospio species, observed only in P. hermesia, P. kaplani and P. solisi Peixoto & Paiva, 2019, although species can be distinguished by branchiae morphology and distribution – on chaetigers 2–3, smooth and cirriform in P. alexandrae sp. nov., on chaetigers 2–3, smooth to slightly wrinkled and cylindrical in P. hermesia, digitiform on chaetiger 2 and conical on chaetiger 3 in P. kaplani and on chaetigers

155

3–4, both pairs robust and flattened in P. solisi. Species also differ from P. alexandrae sp. nov. in having, rather than lacking sabre chaetae (apparently absent in P. kaplani) and in lacking dorsal crests (also lacking in P. solisi) (Paterson et al. 2016).

The presence of long neuropodial chaetae on anterior chaetigers is known from P. brevipinnulata sp. nov., as well as P. fauchaldi, P. corrugatus, both which were already discussed earlier. Prionospio alexandrae sp. nov. can be distinguished from P. brevipinnulata sp. nov. in the prostomium shape – rectangular in P. alexandrae sp. nov. and broadly rounded in P. brevipinnulata sp. nov.; branchiae morphology and distribution – two pairs of apinnate branchiae in P. alexandrae sp. nov. and four pairs of branchiae, first and fourth pairs bearing pinnules in P. brevipinnulata sp. nov. and lack of postchaetal lamellae on chaetiger 1, sabre chaetae and dorsal crests in P. alexandrae sp. nov. (Maciolek 1985; Peixoto & Paiva ms).

The pygidium morphology is unusual, as no cirri nor lobes were observed. Although Blake et al. (2017) diagnosis of the genus Prionospio includes a pygidium in which dorsal cirrus and ventral cirri are fused, this does not seem to be the case, as the pygidium ends abruptly, lacking elongated structures. Since over 50 complete specimens were examined, all with the same pygidium morphology, we believe our observations to be correct.

This species was also observed on samples collected in the continental slope of Sergipe and Alagoas states, northeastern Brazil (pers. obs.).

Etymology. The specific epithet, alexandrae, is a homage to Dr. Alexandra E. Rizzo, a fellow polychaete researcher and a friend.

Habitat: medium silt to clay, 395–3007m depth.

Distribution: Brazil (Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states), Atlantic Ocean.

Prionospio triangularis sp. nov.

(Figs 20–22F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 02 Dec 2011 to 02 Feb 2012, 40m, MZUSP XXXX. Paratypes: Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 11 Dec 2010 to 19 Dec 2010, 41m, MZUSP XXXX (2 ind), MNRJP XXXX (3 ind), ZUEC-POL XXXX (3 ind).

Additional material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (10 ind); Amb1 Foz6, 19°54'56,16" S 39°56'40,83" W, 34m (7 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (3 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (8 ind); Amb1

156

Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (17 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (2 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (2 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (18 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (5 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (6 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (2 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (1 ind); Amb1 Foz19, 19°46'14,99" S 39°34'50,94" W, 52m (7 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (2 ind); Amb2 Foz2, 19°47'12,95" S 39°55'15,42" W, 23m (1 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 34m (3 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (2 ind); Amb2 Foz8, 19°44'44,06" S 39°46'32,01" W, 31m (2 ind); Amb2 Foz9, 19°39'47,16" S 39°42'13,45" W, 28m (12 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (3 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (9 ind); Amb2 Foz13, 19°47'22,52" S 39°43'20,72" W, 41m (5 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (2 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (1 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (4 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (1 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (3 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (12 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (8 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (2 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (7 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (2 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (2 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (15 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (3 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (14 ind); Amb7 C1, 20°10'2,35" S 40°8'31,07" W, 25m (2 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (10 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (10 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (19 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (2 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (5 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (7 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (3 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (4 ind); Amb7 G3, 18°40'55,3" S 38°55'41,48" W, 55m (11 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (1 ind); Amb12 CANWN4, 19°49'6,26" S 39°36'9,34" W, 181m (3 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (24 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (1 ind); Amb14 C1, 20°10'2,67" S 40°8'31,96" W, 26m (2 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (1 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (15 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (1 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (1 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (3 ind).

157

Description: A large-sized Prionospio species, largest complete specimen 18 mm long, 1.1 mm wide at the widest point for 113 chaetigers, holotype 14 mm long, 0.8 mm wide at the widest point for 97 chaetigers. Body slightly dorsoventrally flattened on anterior region and cylindrical on middle and posterior regions, tapering towards the pygidium. Body color whitish in alcohol (Fig 20).

Prostomium subtriangular and rounded anteriorly, extending posteriorly as a narrow and elevated keel reaching the posterior margin of chaetiger 2. U-shaped nuchal organs reaching posterior margin of chaetiger 2. Two pairs of eyes in trapezoidal arrangement, posterior pair large and kidney-shaped. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 20, 21A–C). Grooved palps reaching up to chaetiger 25, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae square-shaped on notopodium and subtriangular on neuropodium, not significantly smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–10 and rounded from chaetiger 11 onwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae present as a low flap from chaetiger 2 to chaetigers 8–11 (depending on specimen size), absent afterwards. Dorsal crests high on chaetiger 7 and low from chaetiger 8 to chaetigers 13–18 (depending on specimen size) (Figs 20, 21A).

Neuropodial postchaetal lamellae triangular with greatly elongated and ventrally pointed tip on chaetiger 2 (Figs 21B–C, 22A), triangular with pointed edges on chaetiger 3 and rounded on remaining chaetigers, gradually reducing in size, present as a low flap from middle region onwards. Neuropodial prechaetal lamellae present as a low flap from chaetiger 2 to chaetigers 7– 9 (depending on specimen size), absent afterwards (Fig 21C).

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, rows of unilimbate and intensely granulated capillaries (Fig 22B), anterior row short and posterior row long (up to 1.5 times longer than anterior row). On neuropodia, rows of narrowly unilimbate and lightly granulated capillaries (Fig 22C). Notopodial chaetae slightly longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non- granulated, thinner and less numerous (Fig 22D).

Hooks in notopodia starting from chaetigers 24–54, up to six per fascicle, accompanied by 1–5 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 13–20, up to nine per fascicle, accompanied by 1–8 non-limbate capillaries. Hooks multidentate, with eight secondary teeth arranged in two rows above main tooth (Fig 22E). Secondary hood present. Sabre chaetae

158 consistently from chaetiger 10, up to two per fascicle. Sabre chaetae narrowly limbate and intensely granulated, except at the base (Fig 22F).

Branchiae present on chaetigers 2–5, 1.5 times longer than notopodial postchaetal lamellae on chaetigers 2 and 5, and slightly longer than notopodial postchaetal lamellae on chaetigers 3 and 4. Branchiae robust, smooth and flattened on chaetigers 3 and 4 and branchiae cirriform, bearing pinnules throughout, except at the tips on chaetigers 2 and 5.

Pygidium bearing a single long dorsal cirrus and two short ventral cirri.

Oocytes from chaetiger 13, measuring up to 120 µm.

Juvenile morphology: In juveniles, the prostomium is shorter, with a rounded appearance. Additionally, the dorsal crests possess a shorter distribution, from chaetiger 7 to chaetiger 10–11.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio triangularis sp. nov. is similar to P. cristata, P. posterocristata sp. nov. and P. cinthyae Peixoto & Paiva, in press, in having a subtriangular and anteriorly rounded prostomium, narrow and elevated keel reaching chaetiger 2, prominent U-shaped nuchal organs, four pairs of branchiae, second and third pairs being flattened, multidentate hooded hooks and sabre chaetae consistently from chaetiger 10. Prionospio triangularis sp. nov. can be easily distinguished from P. cinthyae due to branchiae morphology on first and fourth branchial pairs – pinnate in P. triangularis sp. nov. and apinnate, wrinkled in P. cinthyae (Peixoto & Paiva ms).

However, species can be distinguished based on the morphology of lamellae from chaetiger 1, square-shaped on notopodium and subtriangular in neuropodium in P. triangularis sp. nov., rounded on notopodium and oval in neuropodium in P. cristata, rounded on notopodium and digitiform in neuropodium in P. posterocristata sp. nov. and square-shaped on notopodium and rounded in neuropodium in P. cinthyae and distribution of dorsal crests – high on chaetiger 7 and low from chaetiger 8 to chaetigers 13–18 in P. triangularis sp. nov., high on chaetigers 7 and 9 and low on chaetiger 8 in P. cristata, low from chaetigers 11–12 to chaetigers 16–18 in P. posterocristata sp. nov. and high on chaetiger 7 and low on chaetigers 6, 8 and 9 in P. cinthyae (Foster 1971; Maciolek 1985; Peixoto & Paiva ms).

Species can be further distinguished based on the starting chaetiger of notopodial hooded hooks – from chaetigers 24–54 in P. triangularis sp. nov., chaetigers 17–33 in P. cristata, chaetigers 25–32 in P. posterocristata sp. nov. and chaetigers 21–33 in P. cinthyae, as well as the starting chaetiger of neuropodial hooded hooks – from chaetigers 13–20 in P. triangularis sp. nov., chaetigers 10–13 in P. cristata, chaetigers 12–17 in P. posterocristata sp. nov. and chaetigers10– 12 in P. cinthyae. Despite an overlap on the distribution of notopodial and neuropodial hooded

159 hooks, adult specimens of P. triangularis sp. nov. exhibit an upper range of characters that sets it apart from the remaining aforementioned species (Foster 1971; Maciolek 1985; Peixoto & Paiva ms).

In Maciolek (1985) description of P. cristata, the author reports a low dorsal crest extending after chaetiger 10, in contrast with Foster (1971) original description and Delgado-Blas (2015) description, which reported crests high on chaetigers 7 and 9 and a low crest on chaetiger 8. The dorsal crests distribution of Prionospio cristata reported by Maciolek (1985) is similar to P. triangularis sp. nov., and most likely represents more than one species.

This species, alongside P. cristata, P. cf. heterobranchia Moore, 1907, P. orensanzi Blake, 1983 and P. posterocristata sp. nov., was responsible for all P. steenstrupi records in the collections of the Museu Nacional (Rio de Janeiro state), Museu de Zoologia da Universidade de São Paulo (São Paulo state) and Museu de Zoologia "Adão José Cardoso" (São Paulo state), all located in Brazil (pers. obs.).

This species was also observed on samples collected in the continental slope of Sergipe and Alagoas states, northeastern Brazil (pers. obs.).

Etymology. The specific epithet, triangularis, refers to the shape of neuropodial postchaetal lamellae from chaetiger 2, triangular with pointed edges.

Habitat: medium sand to coarse silt, 23–181 m depth.

Distribution: Brazil (Espírito Santo, Sergipe and Alagoas states), Atlantic Ocean.

Prionospio posterocristata sp. nov.

(Figs 23–25F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 E3, 19°26'5" S 39°17'38,92" W, 02 Dec 2011 to 02 Feb 2012, 50m, MZUSP XXXX. Paratypes: Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 02 Dec 2011 to 02 Feb 2012,149m, MZUSP XXXX (3 ind), MNRJP XXXX (4 ind), ZUEC-POL XXXX (4 ind).

Additional material examined. Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (7 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (9 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (11 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (1 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (1 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (6 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (3 ind); Amb2 Foz13, 19°47'22,52" S 39°43'20,72" W, 41m (1 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m

160

(1 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (9 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (8 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (7 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (2 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (4 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (1 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (3 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (2 ind); Amb5 C5, 20°14'19,45" S 39°48'36,67" W, 381m (1 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (30 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (8 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (2 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (4 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (4 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (6 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (18 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (2 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (3 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (2 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (10 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (17 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (2 ind); Amb7 G3, 18°40'55,3" S 38°55'41,48" W, 55m (3 ind); Amb8 F4, 19°32'57,67" S 38°42'57,83" W, 152m (1 ind); Amb11 B5, 20°35'15,33" S 39°53'45,22" W, 415m (1 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (2 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (7 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (12 ind); Amb12 E5, 19°36'30,6" S 39°10'19,39" W, 360m (1 ind); Amb12 F5, 19°34'20,47" S 38°41'19,8" W, 449m (6 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (1 ind); Amb12 CANWN4, 19°49'6,26" S 39°36'9,34" W, 181m (1 ind); Amb13 G2, 18°36'32,45" S 39°9'32,83" W, 40m (1 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (2 ind); Amb14 A4, 21°4'4,56" S 40°14'14,08" W, 141m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (9 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (6 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (2 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (3 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (2 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (1 ind); Amb14 F4, 19°33'2,92" S 38°42'52,26" W, 160m (2 ind).

Description: A medium-sized Prionospio species, largest complete specimen 9 mm long, 0.7 mm wide at the widest point for 64 chaetigers, holotype 7 mm long, 0.5 mm wide at the widest point for 52 chaetigers. Body slightly dorsoventrally flattened on anterior region and cylindrical on middle and posterior regions, tapering towards the pygidium. Body color whitish in alcohol (Fig 23).

Prostomium subtriangular and rounded anteriorly, extending posteriorly as a narrow and elevated keel reaching the posterior margin of chaetiger 2. U-shaped nuchal organs reaching anterior

161 margin of chaetiger 2. Two pairs of eyes in trapezoidal arrangement, posterior pair slightly enlarged and kidney-shaped. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 23, 24A–B). Grooved and sheathed palps reaching up to chaetiger 15, lost in most specimens. Sheaths covering about half of palp length.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and digitiform on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–11 (largest on chaetigers 3 and 4), subtriangular on chaetigers 12 and 14 and rounded afterwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent throughout. Dorsal crests low from chaetigers 11–12 to chaetigers 16–18 (Figs 23, 24A).

Neuropodial postchaetal lamellae triangular, slightly elongated ventrally and bearing a rounded tip on chaetiger 2 (Fig 25A), square-shaped on chaetiger 3, elliptical on chaetigers 4–9 and rounded afterwards, gradually reducing in size, present as a low flap from middle region onwards (Figs 23, 24C). Neuropodial prechaetal lamellae throughout.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, rows of thick, unilimbate and intensely granulated capillaries (Fig 25B). On neuropodia, rows of unilimbate and lightly granulated capillaries (Fig 25C). Notopodial chaetae slightly longer than neuropodial chaetae, both with a yellow hue. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 25D).

Hooks in notopodia starting from chaetigers 25–32, up to five per fascicle, accompanied by 1–5 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 12–17, up to 10 per fascicle, accompanied by 2–8 non-limbate capillaries. Hooks multidentate, with eight secondary teeth arranged in two rows above main tooth (Figs 24D, 25E). Small secondary hood present. Sabre chaetae consistently from chaetiger 10, up to two per fascicle. Sabre chaetae narrowly limbate and intensely granulated, except at the base (Fig 25F).

Branchiae present on chaetigers 2–5, twice as long as notopodial lamellae on chaetiger 2, up to 1.5 times longer than notopodial lamellae on chaetigers 3 and 4 and up to three times longer than notopodial lamellae on chaetiger 5. Branchiae robust, smooth and flattened on chaetigers 3 and 4 and branchiae cirriform, bearing long pinnules throughout, except at the tips on chaetigers 2 and 5.

Pygidium bearing a single long dorsal cirrus and two short ventral lobes.

162

Juvenile morphology: In juveniles, the prostomium is shorter, with a rounded appearance.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio posterocristata sp. nov. share similarities with P. cristata, P. cinthyae and P. triangularis sp. nov. in having a subtriangular and anteriorly rounded prostomium, narrow and elevated keel reaching chaetiger 2, prominent U-shaped nuchal organs, four pairs of branchiae, second and third pairs being flattened, multidentate hooded hooks and sabre chaetae consistently from chaetiger 10. Prionospio posterocristata sp. nov. can be readily distinguished from P. cinthyae in having pinnate branchiae on chaetigers 2 and 5, instead of apinnate and wrinkled branchiae as in P. cinthyae (Peixoto & Paiva ms).

Prionospio posterocristata sp. nov. can be distinguished from the aforementioned species based on the morphology of lamellae from chaetiger 1 – rounded on notopodium and digitiform in neuropodium in P. posterocristata sp. nov., rounded on notopodium and oval in neuropodium in P. cristata, square-shaped on notopodium and rounded in neuropodium in P. cinthyae and square- shaped on notopodium and subtriangular in neuropodium in P. triangularis sp. nov., as well as distribution of dorsal crests – low from chaetigers 11–12 to chaetigers 16–18 in P. posterocristata sp. nov., high on chaetigers 7 and 9 and low on chaetiger 8 in P. cristata, high on chaetiger 7 and low on chaetigers 6, 8 and 9 in P. cinthyae and high on chaetiger 7 and low from chaetiger 8 to chaetigers 13–18 in P. triangularis sp. nov. (Foster 1971; Maciolek 1985; Peixoto & Paiva ms).

Except for P. triangularis sp. nov., a considerable overlap was observed on the starting chaetiger of hooded hooks, which was not particularly informative in separating species – chaetigers 25– 32 in P. posterocristata sp. nov., chaetigers 17–33 in P. cristata, chaetigers 21–33 in P. cinthyae and chaetigers 24–54 in P. triangularis sp. nov., while the starting chaetiger of neuropodial hooded hooks was more useful in separating species (except for P. triangularis sp. nov.) – from chaetigers 12–17 in P. posterocristata sp. nov., chaetigers 10–13 in P. cristata, chaetigers10–12 in P. cinthyae and chaetigers 13–20 in P. triangularis sp. nov. (Foster 1971; Maciolek 1985; Peixoto & Paiva ms).

Prionospio posterocristata sp. nov. is very similar to P. triangularis sp. nov., although species can be further distinguished by the shape of neuropodial postchaetal lamellae on anterior chaetigers – triangular, slightly elongated ventrally and with a rounded tip on chaetiger 2, square- shaped on chaetiger 3 and elliptical on chaetigers 4–9 in P. posterocristata and triangular with greatly elongated and ventrally pointed tip on chaetiger 2, triangular with pointed edges on chaetiger 3 and rounded on remaining anterior chaetigers in P. triangularis sp. nov., besides specimen size, as adults of P. posterocristata are smaller than adults of P. triangularis sp. nov. (pers. obs.).

163

This species, alongside P. cristata, P. cf. heterobranchia, P. orensanzi and P. triangularis sp. nov., was responsible for all P. steenstrupi records in the collections of the Museu Nacional (Rio de Janeiro state), Museu de Zoologia da Universidade de São Paulo (São Paulo state) and Museu de Zoologia "Adão José Cardoso" (São Paulo state), all located in Brazil. Despite being a species described from the deeper regions of the continental shelf of Iceland, this species is the most recorded Prionospio species in Brazil, unfortunately still being recorded nowadays (Amaral et al., 2013). In a review of the Prionospio steenstrupi species group, Sigvaldadóttir & Mackie (1993) listed additional species bearing dorsal crests starting on posterior chaetigers, although a species comparison is not possible, as most of the species cited are poorly known.

This species was also present on samples collected in the continental slope of Sergipe and Alagoas states, northeastern Brazil (pers. obs.).

Etymology. The species epithet, posterocristata, refers to the late start of dorsal crests, on chaetigers 11–12.

Habitat: medium sand to fine silt, 39–450 m depth.

Distribution: Brazil (Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states).

Prionospio absensi sp. nov.

(Figs. 26–28E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 02 Dec 2011 to 02 Feb 2012,144m, MNRJP XXXX. Paratypes: Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 02 Dec 2011 to 02 Feb 2012,144m, MNRJP XXXX (7 ind), MZUSP (6 ind), ZUEC-POL XXXX (7 ind).

Additional material examined. Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (2 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (16 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (12 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (15 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (9 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (20 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (11 ind); Amb2 Foz10, 19°35'3,5" S 39°38'39,06" W, 31m (2 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (14 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (3 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (1 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (17 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (12 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (28 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (2 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m

164

(2 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (8 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (7 ind); Amb3 CAND5, 19°33'20,99" S 39°2'36,2" W, 420m (1 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (1 ind); Amb5 B5, 20°35'16,23" S 39°53'47,1" W, 410m (12 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (1028 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (214 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (898 ind); Amb6 CANWN5, 19°49'37,21" S 39°35'41,25" W, 410m (1 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (11 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (3 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (14 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (23 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (15 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (2 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (181 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (8 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (90 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (10 ind); Amb7 F4, 19°32'57,67" S 38°42'57,83" W, 152m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (3 ind); Amb11 B5, 20°35'15,33" S 39°53'45,22" W, 415m (2 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (56 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (25 ind); Amb12 F5, 19°34'20,47" S 38°41'19,8" W, 449m (1 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (2 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (4 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (6 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (2 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (7 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (5 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (4 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind); Amb14 F4, 19°33'2,92" S 8°42'52,26" W, 160m (4 ind).

Comparative material examined. Prionospio dubia USNM 65932 (2 specimens; non-type).

Description: A medium-sized Prionospio species, largest complete specimen 23 mm long, 1.1 mm wide at the widest point for 90 chaetigers, holotype 15 mm long, 0.6 mm wide at the widest point for 65 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium. Body color whitish in alcohol (Fig 26).

Prostomium narrow, rounded anteriorly and extending posteriorly as a keel reaching the anterior margin of chaetiger 2. U-shaped nuchal organs reaching anterior margin of chaetiger 2. Two pairs of eyes in trapezoidal arrangement, anterior pair large and kidney-shaped or a single pair of eyes present. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 26, 27A–B). Grooved palps reaching up to chaetiger 40, lost in most specimens.

165

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and digitiform on neuropodium, both much smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–6, gradually becoming rounded on chaetiger 7–9 and rounded from chaetiger 10 onwards, gradually reducing in size, present as a low flap from middle region onwards. Notopodial prechaetal lamellae absent. Dorsal crests absent (Figs 26, 27A–C).

Neuropodial postchaetal lamellae triangular with small dorsally pointed tip on chaetiger 2, triangular on chaetigers 3–5 and rounded from chaetiger 6 onwards, gradually reducing in size, present as a low flap from middle region onwards. Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of narrowly bilimbate capillaries, intensely granulated on notopodia (Fig 28A) and granulated on neuropodia (Fig 28B). Notopodial chaetae slightly longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, non-granulated, thinner and less numerous (Fig 28C).

Hooks in notopodia starting from chaetigers 25–64, up to four per fascicle, accompanied by 0–4 non-limbate capillaries. Hooks in neuropodia starting from chaetigers 10–19, up to eight per fascicle, accompanied by 1–8 non-limbate capillaries. Hooks multidentate, with 10 secondary teeth arranged in two rows above main tooth (Figs 27D, 28D). Secondary hood absent. Sabre chaetae starting from chaetigers 10–19. Sabre chaetae thick, narrowly bilimbate and intensely granulated throughout length (Fig 28E).

Branchiae present on chaetigers 2–5, up to twelve times longer than notopodial lamellae on chaetiger 2, up to 1.5 times longer than notopodial lamellae on chaetigers 3 and 4 and twice as long as notopodial lamellae on chaetiger 5. Branchiae from chaetigers 3–4 robust and flattened; branchiae from chaetigers 2 and 5 cirriform, bearing elongated pinnules throughout, except at the tips. Branchiae from chaetiger 2 around three times longer than branchiae from chaetiger 5.

Pygidium bearing a single long dorsal cirrus and two short ventral cirri.

Sperm plates from chaetiger 15.

Juvenile morphology: In juveniles, branchia from chaetiger 2 is up to three times longer than notopodial postchaetal lamellae.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio absensi sp. nov. is a member of the hereby defined Prionospio dubia Day, 1961 – species group. This group is characterized by an anteriorly narrowed prostomium,

166 widening and becoming rounded in midregion, posterior part of the prostomium extending to chaetiger 2, branchiae on chaetigers 2 to 5, flattened and apinnate on chaetigers 3 and 4 and cirriform and pinnate on chaetigers 2 and 5, branchial pair from chaetiger 2 from 2.5 times (in P. dubia) to five times (in P. jonatani Delgado-Blas, 2015) longer than branchiae from chaetiger 5, long pinnules and lack of dorsal crests. Currently, P. dubia, from Agulhas Bank, South Africa, P. coorilla Wilson, 1990 from Southern Australia, P. jonatani, from the Gulf of Mexico, Mexico and P. absensi sp. nov. are part of this group, besides an undescribed species collected in Northeastern Brazil (Day 1961; Delgado-Blas 2015; pers. obs.). Prionospio oshimensis Imajima 1990b, from Japan, possess a similarly shaped prostomium, long pinnules and lack of dorsal crests, but differs from the remaining species in this group in having a shorter prostomium – extending up to chaetiger 1 and in having branchiae from chaetiger 2 only slightly longer than branchiae from chaetiger 5.

This striking morphological resemblance among species probably led to Brazilian records of P. dubia, compiled by Amaral et al. (2013). According to Sigvaldadóttir & Mackie (1993), this species is reported from Northwestern Atlantic (USA to Surinam), Northeastern Atlantic (North Sea and Mediterranean Sea), Australia and Japan. Delgado-Blas & Salazar-Silva (2011) stated that the identity of material from the Grand Caribbean region should be confirmed, while Delgado-Blas (2015) described P. jonatani, from the Gulf of Mexico.

Additionally, the morphological diversity of P. dubia is still insufficiently known, as all specimens examined by Day (1961) were anterior fragments, which was also the case of topotypes examined by Sigvaldadóttir & Mackie (1993). All complete specimens examined by the authors were collected on the North Sea, far from the type locality. Nevertheless, Sigvaldadóttir & Mackie (1993) description of P. dubia will be used for comparison among species, since Maciolek (1985) description differs from the description of P. dubia from South Africa, as noted by Wilson (1990), and may actually represent more than one species (Wilson 1990; Sigvaldadóttir & Mackie 1993).

Prionospio absensi sp. nov. can be distinguished from P. dubia, P. coorilla and from P. jonatani in the shape of notopodial lamellae from chaetiger 1, rounded on notopodium and digitiform on neuropodium in P. absensi sp. nov., large and rounded on notopodium and narrow, rounded and rectangular in P. dubia, rounded on notopodium and neuropodium rami in both P. coorilla and P. jonatani; starting chaetiger of notopodial hooded hooks – from chaetigers 25–64 in P. absensi sp. nov., chaetiger 48 in a single South African P. dubia specimen (chaetigers 25–70 in European specimens), chaetigers 42–50 in P. coorilla and chaetigers 35–57 in P. jonatani and starting chaetiger of neuropodial hooded hooks – from chaetigers 10–19 in P. absensi sp. nov., chaetigers 18–20 in South African P. dubia specimens (chaetigers 13–22 in European specimens), chaetigers

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15–18 in P. coorilla and chaetigers 16–28 in P. jonatani (Wilson, 1990; Sigvaldadóttir & Mackie,1993; Delgado-Blas, 2015).

Species can also be distinguished based on the starting chaetiger of sabre chaetae – from chaetigers 10–19 in P. absensi sp. nov., chaetigers 18–20 in South African P. dubia specimens (chaetigers 11–22 in European specimens), chaetigers 12–14 in P. coorilla and chaetigers 14–18 in P. jonatani. Prionospio absensi sp. nov. can be further distinguished from P. coorilla and P. jonatani in having notopodial capillaries from the anterior region organized in two rows, instead of three (Wilson 1990; Sigvaldadóttir & Mackie 1993; Delgado-Blas 2015).

Among Brazilian species, P. solisi, P. fosterae, P. hartmanae, P. brevipinnulata sp. nov., P. alexandrae sp. nov. and P. biancoi sp. nov. also lack dorsal crests, although species can be readily distinguished based on the prostomium shape – narrow and rounded anteriorly in P. absensi sp. nov., rounded in P. solisi, narrow, rounded anteriorly in both P. fosterae and P. hartmanae, broadly rounded in P. brevipinnulata sp. nov., narrow and rectangular in P. alexandrae sp. nov. and rectangular P. biancoi sp. nov., and branchiae morphology – cirriform and pinnate on chaetigers 2 and 5 (bearing long pinnules); flattened and apinnate on chaetigers 3 and 4 in P. absensi sp. nov., two pairs of flattened branchiae in P. solisi, up to 11 pairs of apinnate branchiae in both P. fosterae and P. hartmanae, cirriform and pinnate on chaetigers 2 and 5 (bearing very short pinnules); flattened and apinnate on chaetigers 3 and 4 in P. brevipinnulata sp. nov., two pairs of cirriform and apinnate branchiae in P. alexandrae sp. nov. and four pairs of apinnate branchiae in P. biancoi sp. nov. (Peixoto & Paiva 2019; Peixoto & Paiva ms).

This species was the most abundant Prionospio species of this work, with nearly 3,000 specimens collected and over 1,000 specimens in a single station.

Etymology. The species epithet, absensi (“absens”, Latin for lacking) refers to the lack of dorsal crests.

Habitat: medium sand to coarse silt, 30–450 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Prionospio biancoi sp. nov.

(Figs. 29–31E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 C7,20°17'41,07" S 39°42'38,02" W, 02 Dec 2011 to 02 Feb 2012,1358m, MNRJP XXXX. Paratypes: Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 02 Dec 2011 to 02 Feb 2012, 1300m, MZUSP XXXX (1 ind), ZUEC-POL XXXX (1 ind).

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Additional material examined. Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (1 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (1 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (1 ind).

Description: A large-sized Prionospio species, largest specimen 15 mm long, 1 mm wide at the widest point for 72 chaetigers (incomplete), holotype 9 mm long, 0.7 mm wide at the widest point for 40 chaetigers (incomplete). Body slightly dorsoventrally flattened on entire fragment, tapering towards end of fragment. Body color light yellow to yellow in alcohol, with golden inclusions from chaetigers 5–6 to chaetigers 11–13 (Fig 29).

Prostomium rectangular, extending posteriorly as a narrow keel reaching the posterior margin of chaetiger 2. Nuchal organs not observed. Eyes absent. Prostomial peaks absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lateral wings absent (Figs 29, 30B–C). Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae absent on notopodium and auricular on neuropodium, both much smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 5–8 (depending on specimen size), quadrangular and gradually reducing in size from chaetigers 6–9 to chaetigers 13–17, present as a low flap afterwards (Figs 29, 30A). Notopodial prechaetal lamellae absent. Dorsal crests absent. Lamellae from posterior region unknown.

Neuropodial postchaetal lamellae rounded and gradually reducing in size from chaetiger 2 to chaetigers 24–31, reduced to a low flap afterwards. Neuropodial prechaetal lamellae absent. Lamellae from posterior region unknown.

Chaetae from notopodia and neuropodia organized in two rows of wide, unilimbate and granulated capillaries, intensely granulated near the base and sparsely granulated near the tip (Fig 31A). Notopodial capillaries up to 1.4 times longer than neuropodial capillaries (Fig 31B). Towards end of fragment, capillaries progressively become elongate, non-limbate, non- granulated, thinner and less numerous (Fig 31C).

Hooks in notopodia starting from chaetiger 67, up to three per fascicle, accompanied by 1–4 non- limbate capillaries. Hooks in neuropodia starting from chaetigers 16–22, up to six per fascicle, accompanied, by 1–6 non-limbate capillaries. Hooks multidentate, with 10 secondary teeth arranged in two rows above main tooth (Figs 30D, 31D). Secondary hood present. Sabre chaetae starting from chaetigers 14–20. Sabre chaetae non-limbate and intensely granulated throughout length (Fig 31E).

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Branchiae present on chaetigers 2–5, about the same length of notopodial lamellae on chaetiger 2, slightly shorter than notopodial lamellae on chaetigers 3 and 4 and about half the length of notopodial lamellae on chaetiger 5. Branchiae from chaetigers 3–4 robust, flattened and densely ciliated; branchiae from chaetigers 2 and 5 smooth and cirriform.

Pygidium of unknown morphology.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Prionospio biancoi sp. nov. is unusual for its large size in deep sea samples, being much larger than its congenerics. The yellowish body color and lack of dorsal crests are similar to P. solisi, although species can be readily distinguished by the prostomium morphology – rectangular in P. biancoi sp. nov. and rounded in P. solisi, shape of lamellae from chaetiger 1 – absent on notopodium and auricular on neuropodium in P. biancoi sp. nov. and rounded on both rami in P. solisi; branchiae distribution – on chaetigers 2–5 in P. biancoi sp. nov. and chaetigers 3 and 4 in P. solisi; starting chaetiger of notopodial hooded hooks – from chaetiger 67 in P. biancoi sp. nov. and chaetigers 48–61 in P. solisi; starting chaetiger of neuropodial hooded hooks – from chaetigers 16–22 in P. biancoi sp. nov. and chaetigers 9–14 in P. solisi and starting chaetiger of sabre chaetae – from chaetigers 14–20 in P. biancoi sp. nov. and chaetigers 9–11 in P. solisi (Peixoto & Paiva 2019).

The species is also similar to P. fosterae, P. hartmanae, P. brevipinnulata sp. nov., P. alexandrae sp. nov. and P. absensi sp. nov. in lacking dorsal crests, although species differ on the prostomium shape – rectangular in P. biancoi sp. nov., anteriorly rounded in both P. fosterae and P. hartmanae, broadly rounded in P. brevipinnulata sp. nov., narrow and rectangular in P. alexandrae sp. nov. and narrow and rounded anteriorly in P. absensi sp. nov. Species can also be distinguished by the branchial morphology – four pairs of apinnate branchiae in P. biancoi sp. nov., up to 11 pairs of apinnate branchiae in both P. fosterae and P. hartmanae, cirriform and pinnate on chaetigers 2 and 5 (bearing very short pinnules); flattened and apinnate on chaetigers 3 and 4 in P. brevipinnulata sp. nov. two pairs of cirriform and apinnate branchiae in P. alexandrae sp. nov. and cirriform and pinnate on chaetigers 2 and 5 (bearing long pinnules); flattened and apinnate on chaetigers 3 and 4 in P. absensi sp. nov. (Peixoto & Paiva ms).

Prionospio biancoi sp. nov. is similar to P. quadrilamellata sp. nov. in having four pairs of apinnate branchiae, although species can be distinguished by the prostomial morphology – rectangular, rounded on the anterior margin in P. biancoi sp. nov. and rectangular, narrow, truncated on the anterior margin in P. quadrilamellata sp. nov., branchiae morphology – first and fourth branchial pairs cirriform and second and third pairs flattened in P. biancoi sp. nov. and all pairs flattened in P. quadrilamellata sp. nov., lack of dorsal crests in P. biancoi sp. nov.; starting chaetiger of notopodial hooded hooks – from chaetiger 67 in P. biancoi sp. nov. and chaetigers

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20–32 in P. quadrilamellata sp. nov., starting chaetiger of neuropodial hooded hooks – from chaetigers 16–22 in P. biancoi sp. nov. and chaetigers 9–13 in P. quadrilamellata sp. nov. and starting chaetiger of sabre chaetae – from chaetigers 14–20 in P. biancoi sp. nov. and chaetigers 9–11 in P. quadrilamellata sp. nov.

Unfortunately, only a few anterior fragments were collected, thus characters such as starting chaetiger of notopodial hooded hooks remain poorly known while the morphology of the pygidium remains unknown.

Etymology. The specific epithet, biancoi, is a homage to Cinthia X. Bianco, who helped the first author immensely during the initial sorting and identification of Spionidae specimens.

Habitat: very fine sand to fine silt, 988 –1358 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Identification key to Prionospio species recorded in this work.

1 Sabre chaetae absent...... P. alexandrae sp. nov. - Sabre chaetae present………………………………………………………………………...2 2(1) Branchiae all apinnate…………………………………………………………………….3 - Branchiae pinnate and apinnate……………………………………………………………...4 3(2) Cirriform and flattened branchiae, dorsal crests absent…...………...P. biancoi sp. nov. - All branchiae flattened, dorsal crests present...... P. quadrilamellata sp. nov. 4(2) Dorsal crests absent...... 5 - Dorsal crests present...... 6 5(4) Prostomium broadly rounded, row of long neurochaetae on chaetigers 2–4………………………………………………………..P. brevipinnulata sp. nov - Prostomium narrow, neurochaetae from chaetigers 2–4 not elongated…...P. absensi sp. nov. 6(4) Dorsal crests starting after chaetiger 10……………………..P. posterocristata sp. nov. - Dorsal crests starting before chaetiger 10……………………………………………………7 7(6) Branchiae pinnate only on chaetiger 2……………………………...Prionospio cf. ehlersi - Branchiae pinnate on chaetigers 2 and 5……………………………………………………..8 8(7) Dorsal crests from chaetiger 8, sabre chaetae from chaetigers 10–12…………………………………………………………..P. capixaba sp. nov. - Dorsal crests from chaetiger 7, sabre chaetae consistently from chaetiger 10……………….9 9(8) Dorsal crests on chaetiger 7–9, neuropodial postchaetal lamellae from chaetiger 2 square-shaped……………………………………………………………………..…P. cristata

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- Dorsal crests from chaetiger 7 to chaetigers 13–18, neuropodial postchaetal lamellae from chaetiger 2 triangular and ventrally pointed…………………………...P. triangularis sp. nov.

Genus Aurospio Maciolek, 1981a

Type species: Aurospio dibranchiata Maciolek, 1981a

Diagnosis (modified from Blake et al. 2017): Prostomium broadly rounded anteriorly, prolonged posteriorly as a keel; eyespots 0–2 pairs, occipital tentacle absent. Peristomium partly fused to chaetiger 1, not developed into lateral wings or hood. Notopodial postchaetal lamellae enlarged on chaetigers 3 and 4. Branchiae two pairs on chaetigers 3–4, short, fingerlike partly fused to notopodial lamellae. Interramal or interparapodial pouches absent. Anterior chaetae all capillaries; neuro- and notopodial multidentate hooded hooks from chaetigers 9–11 and 24–38 respectively in type-species; hooks multidentate without secondary hood; neuropodial sabre chaetae present from chaetigers 9–11. Pygidium with one long medial cirrus and two short lateral cirri.

Aurospio dibranchiata Maciolek, 1981a

(Figs 32–34H)

Material examined. Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 1035m (50 ind); Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 2426m (7 ind); Amb3 G10 19°3'55,8" S 37°45'8,27" W, 2993m (2 ind); Amb3 CAND6 19°37'50,08" S 39°3'56,16" W, 970m, (3 ind); Amb3 CAND7, 19°42'18,42" S 39°5'52,41" W, 1307m (28 ind); Amb4 A9 21°9'35,19" S 38°52'4,93" W, 2502m (1 ind); Amb4 A10, 21°11'4,9" S 38°28'2,78" W, 3006m (1 ind); Amb4 B8, 20°41'33,45" S 39°35'14,76" W, 1914m (4 ind); Amb4 B10, 21°4'40,47" S 38°26'13,68" W, 3007m (1 ind); Amb4 D10, 20°53'33,64" S 38°21'17,35" W, 3004m (1 ind); Amb4 E9, 20°35'50,48" S 38°27'7,64" W, 2494m (3 ind); Amb4 E10, 20°49'23,58" S 38°17'11,07" W, 2997m (1 ind); Amb4 F6, 19°52'56,9" S 38°35'8,88" W, 1022m (17 ind); Amb4 F7, 20°4'8,18" S 38°31'27,32" W, 1302m (56 ind); Amb4 F8, 20°16'35,72" S 38°27'18,98" W, 1902m (53 ind); Amb4 F9, 20°29'3,31" S 38°23'15,5" W, 2504m (3 ind); Amb4 F10, 20°46'23,65" S 38°17'17,65" W, 3004m (4 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (28 ind); Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 1300m (38 ind); Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 1889m (4 ind); Amb5 A9, 21°9'40,3" S 38°52'25,04" W, 2498m (1 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (34 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (48 ind); Amb5 B9,

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20°54'44,14" S 38°56'10,72" W, 2519m (1 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (41 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (42 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (4 ind); Amb5 C9, 20°48'39,87" S 38°45'23,86" W, 2496m (2 ind); Amb5 D6, 19°50'1,87" S 39°26'30,04" W, 1055m (41 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 1335m (60 ind); Amb5 D8, 20°8'42,82" S 39°7'29,5" W, 1922m (9 ind); Amb5 D9, 20°34'36,32" S 38°40'53,6" W, 2460m (2 ind); Amb5 E8, 20°15'59,97" S 38°40'53,86" W, 1887m (16 ind); Amb6 D5, 19°46'34,99" S 39°30'4,65" W, 428m (7 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (50 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (35 ind); Amb8 E7, 19°47'5,96" S 39°3'11,96" W, 1258m (19 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (24 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (60 ind); Amb8 G8, 19°3'45,82" S 37°47'28,26" W, 1928m (23 ind); Amb11 A6, 21°4'37,64" S 40°8'32,68" W, 1015m (12 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (16 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (4 ind); Amb11 A9, 21°9'39,38" S 38°52'7,25" W, 2502m (2 ind); Amb11 A10, 21°10'59,11" S 38°28'4,99" W, 2987m (1 ind); Amb11 B6, 20°36'1,61" S 39°51'39,15" W, 1003m (13 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (37 ind); Amb11 B8, 20°41'33,93" S 39°35'22,06" W, 1908m (2 ind); Amb11 B9, 20°54'43,79" S 38°56'10,85" W, 2520m (1 ind); Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 1040m (35 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (48 ind); Amb11 C8, 20°25'13,22" S 39°27'19,49" W, 1920m (5 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (9 ind); Amb11 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (1 ind); Amb11 E8, 20°15'55,63" S 38°40'45,57" W, 1892m (46 ind); Amb11 E9, 20°35'51,99" S 38°27'13,04" W, 2501m (1 ind); Amb11 E10, 20°49'19,95" S 38°17'8,76" W, 2994m (3 ind); Amb11 F6, 19°52'52,59" S 38°35'10,48" W, 1021m (7 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (20 ind); Amb11 F8, 20°16'38,17" S 38°27'26,52" W, 1904m (51 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (5 ind); Amb11 F10, 20°46'17,79" S 38°17'16,01" W, 3002m (2 ind); Amb12 D6, 19°50'6,01" S 39°26'34,62" W, 1050m (56 ind); Amb12 D7, 19°54'4,77" S 39°22'29,46" W, 1335m (38 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (50 ind); Amb12 E7, 19°47'2,44" S 39°3'14,62" W, 1250m (22 ind); Amb12 G7, 19°3'30,62" S 37°48'46,66" W, 1347m (54 ind); Amb12 G8, 19°3'39,78" S 37°47'39,35" W, 1867m (21 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (2 ind); Amb12 G10, 19°3'10,22" S 37°45'28,45" W, 2950m (1 ind); Amb12 CAND6, 19°37'45,14" S 39°3'58,75" W, 1050m (10 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (31 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (53 ind).

Comparative material examined. Aurospio dibranchiata USNM 60326 (holotype); USNM 60327 (35 paratypes); USNM 162820 (20 specimens; non-type); USNM 1106015 (9 specimens; non-type).

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Description: A medium-sized Aurospio species, largest complete specimen 5.5 mm long, 0.3 mm wide at the widest point for 43 chaetigers. Body rounded throughout, tapering towards the pygidium. Body color yellow to whitish in alcohol, with dark yellow or golden inclusions on the prostomium, dorsal side of the peristomium and first chaetigers (Fig 32).

Prostomium broadly rounded, extending as a narrow keel almost to the posterior margin of chaetiger 1. One pair of minute reddish eyes or eyes absent. Peristomium surrounding prostomium and partially fused to the first chaetiger, lacking lateral wings (Figs 32, 33A–B, 34A). Grooved palps reaching up to chaetiger 12, lost in most specimens. Saccular and highly ciliated proboscis everted on some specimens (Fig 33A–B).

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on both rami, much smaller than lamellae on succeeding chaetigers; neuropodial lamellae displaced to a slightly dorsal position (Figs 33A–B, 34A). Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–6 (Fig 34B), rounded from chaetiger 7 and gradually reducing in size towards the posterior region, present as a low flap on the last chaetigers. Notopodial prechaetal lamellae absent throughout. Low dorsal crests usually present, from chaetigers 10–11 to chaetigers 15–17 (Figs 32, 33A, 34A).

Neuropodial postchaetal lamellae rounded and well developed on chaetigers 2‒5, gradually reducing in size towards the posterior region (Fig 34C), present as a low flap on the last chaetigers. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of unilimbate and lightly granulated capillaries (Fig 34D). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner, and less numerous (Fig 34E).

Hooks in notopodia starting from chaetigers 18‒38, up to four per fascicle, accompanied by row of 1‒3 short non-limbate capillaries. Hooks in neuropodia starting from chaetigers 9–11 (almost always chaetiger 10), up to nine per fascicle, accompanied by 1‒5 non-limbate capillaries. Hooks multidentate, with six small secondary teeth arranged in two rows above the main tooth (Figs 33D–E, 34F). Secondary hood absent. Sabre chaetae from chaetiger 9–11 (almost always chaetiger 10) (Fig 34G).

Branchiae present on chaetigers 3–4, robust, short, digitiform and basally fused to the notopodial lamellae. Branchiae longer on chaetiger 3, both shorter than notopodial lamellae (about one third of the notopodial lamellae length at most) (Fig 33C).

Pygidium bearing one dorsal cirrus and two short ventral cirri (Figs 33F, 34H).

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Juvenile morphology: In juveniles less than 20 chaetigers long, notopodial hooded hooks are absent.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Aurospio dibranchiata is an easily recognized species due to its yellow to golden inclusions, broadly rounded prostomium and very short and flattened branchiae on chaetigers 3 and 4.

Specimens recorded in this work fit the description and illustrations of the original description (Maciolek 1981a), differing only in the absence of notopodial hooded hooks in small specimens, but this can be due to ontogeny, as the notopodial hooded hooks may still develop as the specimen grows.

Aurospio dibranchiata is similar to Prionospio solisi in having two pairs of apinnate branchiae on chaetigers 3 and 4, starting chaetiger of sabre chaetae (chaetigers 9–11; usually chaetiger 10), and dark yellow pigment on the prostomium, although both species can be distinguished based on the branchial morphology – flattened and basally fused to and shorter than notopodial lamellae in A. dibranchiata and triangular, free from and about the same size or slightly longer than notopodial lamellae. The two species also differ on the starting chaetiger of notopodial hooded hooks (chaetigers 24–38 in A. dibranchiata and chaetigers 48–61 in P. solisi), starting chaetiger of neuropodial hooded hooks (chaetigers 9–11 in A. dibranchiata and chaetigers 9–14 in P. solisi) and distribution of dark yellow inclusions, on chaetiger 1 and posterior margin of prostomium in A. dibranchiata and on dorsal and ventral sides of chaetigers 4–5 to chaetigers 8–12 in P. solisi.

Only three Aurospio species co-occur with A. dibranchiata in Atlantic Ocean: A. tribranchiata Neal, Paterson & Soto in Paterson et al., 2016, from Porcupine Abyssal Plain, Northeastern Atlantic; A. abranchiata Neal, Paterson & Soto in Paterson et al., 2016, from Canyons of the Iberian Margin Northeastern Atlantic; and A. banyulensis (Laubier, 1966), from the Mediterranean coast of France.

Aurospio dibranchiata can be readily distinguished from A. abranchiata as the latter species lacks branchiae and can be separated from remaining species based on branchial morphology, as both A. tribranchiata and A. banyulensis bear three pairs of branchiae on chaetigers 3–5, all free from notopodial lamellae, while A. dibranchiata possess only two pairs of branchiae on chaetigers 3 and 4, which are robust, flattened and basally fused to notopodial lamellae.

The species was one of the most common taxa on the continental slope, found on most samples below the 400 m isobath. The species is also considered widespread in the Atlantic Ocean – with records from North and South America, Europe and Africa (Maciolek, 1981a), Antarctica (Mincks et al. 2009) and the Pacific Ocean (Maciolek 1981a; Mincks et al. 2009). Despite the

175 wide geographical record, the author considers notable the consistency of morphological characters, treating all records as the same species, although a recent study off Nova Scotia, Northwestern Atlantic Ocean (Neal et al. 2018), revealed that Aurospio dibranchiata is a species complex, with bathyal and abyssal clades.

Habitat: muddy sand to mud,428 – 3006 m depth.

Distribution: Atlantic Ocean –North (Bermuda, USA, Canada) and South America (Suriname, Brazil, Argentina), Europe (Ireland, off France and Spain) and Africa (Canary Islands, Angola) (Maciolek 1981a), Antarctica (Mincks et al., 2009) and the Pacific Ocean (Ecuador, Clarion- Clipperton Fracture Zone) (Maciolek 1981a; Mincks et al. 2009).

Genus Paraprionospio Caullery, 1914

Type species: Prionospio pinnata Ehlers, 1901, designated by Caullery, 1914 (superseded original combination).

Diagnosis (from Delgado-Blas & Carrera-Parra 2018): Prostomium fusiform with rounded, truncate to slightly rounded or bluntly pointed anterior end, occasionally with slight medial peaks; with a raised, narrow ridge on the posterior region of the prostomium; caruncle absent. Eyes present or absent. Peristomium fused with first achaetous segment, forming large lateral membranous wings enclosing prostomium. A pair of grooved palpi with membranous basal sheath. Eversible bilobate proboscis. Muscular gizzard present. Chaetiger 1 well developed, distinct from preceding segments. Two or three pairs of branchiae. When three pairs of branchiae are present, these are on chaetigers 1-3, all with densely packed lamellar plates attached serially on inner to posterior face of shaft, excepting in the basal region and distal tips. When two pairs of branchiae are present, these are located on chaetigers 1-2; first pair of branchiae pinnate with few plate-like pinnules or lamellae, the second one cirriform. Each branchia entirely free from notopodial lamella. Two thread- like filaments at the base of third notopodium present in some species. Distinct transverse ridge between branchial bases of chaetiger 1 or chaetigers 1-3 as in P. africana. Dorsal ridges in middle segment present. Interparapodial pouches present or absent. Notopodial and neuropodial capillaries limbate and granulated in anterior chaetigers, replaced by non-limbate, slender capillaries without granulation in middle and posterior chaetigers. Sabre chaetae starting from chaetiger 9. Neuropodial multidentate hooded hooks from chaetiger 9. Notopodial hooded hooks appearing from middle body region. Notopodial and neuropodial hooks with conspicuous striated secondary internal hooded, absent in some species. Shallow ventral furrow presents longitudinally from middle to the end of the body. Dorsal crest present or absent.

176

Pygidium with one long and thin mid-dorsal cirrus, and two shorter ventrolateral cirri present in some species.

Paraprionospio anterocristata sp. nov.

(Figs 35–37F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 11 Dec 2010 to 19 Dec 2010, 33m, MNRJP XXXX. Paratypes: Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 06 Jun 2013 to 17 Jul 2013, 53m, MNRJP XXXX (3 ind), MZUSP XXXX (3 ind), ZUEC-POL XXXX (3 ind).

Additional material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (1 ind); Amb1 Foz3, 19°42'12,86" S 39°49'19,48" W, 16m (1 ind); Amb1 Foz5, 19°32'32,82" S 39°41'34,89" W, 20m (1 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (1 ind); Amb1 Foz10, 19°35'12,39" S 39°38'33,16" W, 32m (2 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (3 ind); Amb2 Foz1, 19°52'14,57" S 39°59'41,06" W, 27m (4 ind); Amb2 Foz3, 19°42'8,87" S 39°49'29,72" W, 16m (3 ind); Amb2 Foz5, 19°32'28,16" S 39°41'35,15" W, 20m (1 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (1 ind); Amb2 Foz10, 19°35'3,5" S 39°38'39,06" W, 31m (6 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (5 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (3 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (2 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (1 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (2 ind); Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (5 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (5 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (1 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (15 ind); Amb14 B1, 20°34'34,37" S 40°20'50,77" W, 26m (2 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (2 ind); Amb14 D1, 19°35'35,96" S 39°41'20,61" W, 26m (3 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (4 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (4 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (9 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (3 ind).

Comparative material examined. Paraprionospio alata USNM 1260223 (one specimen, non- type material).

Description: A medium-sized Paraprionospio species, largest complete specimen 15 mm long, 0.8 mm wide at the widest point for 61 chaetigers, holotype 13 mm long, 0.7 mm wide at the widest point for 53 chaetigers. Body cylindrical on the anterior region and slightly dorsoventrally flattened afterwards, tapering towards the pygidium. Body color light yellow to whitish in alcohol (Fig 35).

177

Prostomium fusiform, rounded anteriorly, extending posteriorly as a narrow keel almost to the posterior margin of chaetiger 1. Two pairs of eyes in trapezoidal arrangement on the posterior half of the prostomium or eyes absent. Peristomium well-developed, surrounding prostomium and forming large lateral wings, not fused to first chaetiger (Fig 35, 36A–B). Grooved palps reaching up to chaetiger 9, lost in most specimens. Palp sheath smooth, covering palps for two thirds of its length.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae foliaceous on notopodium and lanceolate on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 3–6 (depending on specimen size), triangular on following chaetigers; becoming lanceolate on middle region and gradually reducing in size towards the posterior region, although remaining lanceolate until last chaetiger. Notopodial prechaetal lamellae as a low flap from chaetiger 2 to chaetigers 12–15 (depending on specimen size) (Figs 35, 36A–B, D). Transverse ridge between branchial bases on chaetiger 1 (Fig 36B). Dorsal crests low from chaetiger 7 to chaetiger 11. Semi-transparent dorsal plates bearing several small rounded concavities from chaetiger 23 to chaetiger 27 (Figs 35, 36E).

Neuropodial postchaetal lamellae lanceolate on chaetigers 2 and 3, rounded with small median tip on chaetiger 4 and rounded from chaetiger 5 onwards, rapidly decreasing in size and present as a low flap from chaetiger 9 onwards. Neuropodial prechaetal lamellae absent throughout. Ventral crests present from chaetiger 4 to chaetigers 13–17 (depending on specimen size) (Fig 36C).

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate capillaries, intensely granulated on notopodia (Fig 37A) and lightly granulated on neuropodia (Fig 37B). Chaetae from both rows of equal length, although notopodial chaetae are up to 1.4 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 37C).

Hooks in notopodia starting from chaetigers 34–45, up to two per fascicle, accompanied by row of 2–5 short non-limbate capillaries. Hooks in neuropodia consistently from chaetiger 9, up to 13 per fascicle, accompanied by 1–9 short non-limbate capillaries. Hooks slightly bent near distal margin; with six or eight small secondary teeth arranged in two rows above the main fang (Figs 36F, 37D). Secondary hood present. Intensely granulated sabre chaetae consistently from chaetiger 9 (Fig 37E).

Branchiae present on chaetigers 1–3, longest on chaetiger 1 and shorter on chaetiger 3. Branchiae up to six times longer than notopodial postchaetal lamellae on chaetiger 1, up to three times longer

178 than notopodial postchaetal lamellae on chaetiger 2 and up to 1.5 times longer than notopodial postchaetal lamellae on chaetiger 3. Branchiae from chaetiger 1 basally fused to dorsal ridge (transverse ridge) (Fig 36B). Branchiae bearing densely packed plate-like pinnules, absent on branchial base (Figs 36A–B, 37F). Pinnules fused through branchial stem from midregion to branchial tip.

Pygidium bearing a single long dorsal cirrus and two short and thin ventro-lateral cirri.

Internal anatomy: Gizzard-like structure on digestive tract on anterior chaetigers, extending for four or five chaetigers (Fig 36D).

A single specimen had a single large parasite on the coelomic cavity. The parasite resembled a nematode, but was unfortunately broken during sample preparation for SEM studies (Fig 36E).

Juvenile morphology: In juveniles, notopodial hooded hooks are absent on specimen with less than 34 chaetigers.

Methyl green staining pattern: Prostomium, peristomium and notopodial postchaetal lamellae from middle and posterior regions strongly stained.

Remarks. Paraprionospio anterocristata sp. nov. shares many similarities with P. africana Augener, 1918, P. pinnata (Ehlers, 1901) and P. tamaii Delgado-Blas, 2004, species that were previously recorded to the Brazilian coast (Amaral et al. 2013). Species possess a fusiform prostomium, well-developed peristomium forming large lateral wings that do not touch dorsally, lack of neuropodial prechaetal lamellae, neuropodial hooded hooks and sabre chaetae starting from chaetiger 9, presence of a secondary hood and pygidium bearing a single long dorsal cirrus and two short and thin ventro-lateral cirri.

This species can be most easily distinguished from P. africana, as species differ on the anterior shape of the prostomium – rounded in P. anterocristata sp. nov. and blunt in P. africana; two pairs of eyes in P. anterocristata sp. nov. and eyes not observed in P. africana; shape of notopodial postchaetal lamellae from chaetiger 1 – foliaceous in P. anterocristata sp. nov. and elongate subtriangular in P. africana; shape of notopodial postchaetal lamellae on middle and posterior regions – lanceolate in P. anterocristata sp. nov. and low and rounded in P. africana; presence of notopodial prechaetal lamellae, lacking in P. africana; distribution of dorsal ridge – only on chaetiger 1 in P. anterocristata sp. nov. and on chaetigers 1–3 in P. africana; distribution of dorsal crests – on chaetigers 7–11 in P. anterocristata sp. nov. and on chaetigers 21–36 in P. africana; shape of neuropodial postchaetal lamellae from chaetiger 4 onwards (Table 2), hook morphology – with six secondary teeth above main fang in P. anterocristata sp. nov. and 6 – 10 secondary teeth in P. africana and branchiae morphology – lack of additional branchial filament on chaetiger 3 in P. anterocristata sp. nov., which is present in P. africana (Yokoyama 2007).

179

Paraprionospio anterocristata sp. nov. is most similar to P. pinnata and P. tamaii in having an anteriorly rounded prostomium, presence of eyes, palp sheaths and presence of dorsal ridge only on chaetiger 1, although species can be distinguished based on length of palp sheaths – covering up to two thirds of palp length in P. anterocristata sp. nov., one third of palp length in P. pinnata and half of palp length in P. tamaii; shape of lamellae from chaetiger 1 – foliaceous on notopodium and lanceolate on neuropodium in P. anterocristata sp. nov., elongate subtriangular on notopodium and rounded with a pointed tip on neuropodium in P. pinnata and oval on notopodium and lanceolate on neuropodium in P. tamaii; shape of notopodial postchaetal lamellae (Table 2) and presence of notopodial prechaetal lamellae, absent in P. pinnata and P. tamaii. (Yokoyama 2007; Delgado-Blas & Carrera-Parra 2018).

Species also differ on the distribution of dorsal crests – on chaetigers 7–11 in P. anterocristata sp. nov., absent in P. pinnata (although there is a ridge on chaetigers 21–26) and from chaetigers 20–29, present in five to nine chaetigers in P. tamaii; distribution of dorsal plates – on chaetigers 23–27 in P. anterocristata sp. nov., on chaetigers 21–26 in P. pinnata and chaetigers 6–17 (Delgado-Blas, 2004) or chaetigers 21–28 (Yokoyama 2007) in P. tamaii; shape of neuropodial postchaetal lamellae (Table 2); distribution of notopodial hooded hooks – from chaetigers 34–45 in P. anterocristata sp. nov., chaetigers 32–34 in P. pinnata and chaetigers 40–56 in P. tamaii and morphology of hooded hooks – bearing 6–8 secondary teeth above main fang in both P. pinnata and P. anterocristata sp. nov., and 7 secondary teeth above main fang in P. tamaii.

Paraprionospio anterocristata sp. nov. is unique in having ventral crests from chaetiger 4 to chaetigers 13–17, as this character is present only in chaetiger 9 in P. cordifolia Yokoyama, 2007 and P. cristata Zhou et al., 2008, being completely absent on the remaining Paraprionospio species.

Etymology. The species epithet, anterocristata, refers to the anterior position of dorsal crests, present on chaetigers 7–11.

Habitat: coarse sand to coarse silt,16–53 m depth.

Distribution: Atlantic Ocean – Brazil (Rio de Janeiro and Espírito Santo states).

Genus Apoprionospio Foster, 1969

Type species: Apoprionospio dayi Foster, 1969, by original designation.

Diagnosis (from Blake et al. 2017): Prostomium subtriangular, broad anteriorly, often with medial peak, caruncle extends posteriorly to end of chaetiger 1; occipital antenna absent. Peristomium reduced, fused with chaetiger 1, surrounding prostomium posteriorly as a collar; lateral wings

180 absent. Notopodial postchaetal lamellae of anterior chaetigers lateral rather than dorsomedial. Neuropodial postchaetal lamellae enlarged on chaetiger 2. Branchiae from chaetiger 2, with four pairs, first three pairs apinnate, fourth pair larger, with flattened platelike pinnules. Anterior chaetae capillaries; multidentate hooded hooks present in posterior noto- and neuropodia neuropodial sabre chaetae present. Pygidium with one long dorsomedial cirrus and two shorter ventrolateral lobes.

Apoprionospio cf. dayi Foster, 1969

(Figs. 38–40G)

Material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (3 ind); Amb1 Foz2, 19°47'21,11" S 39°55'7,58" W, 24m (1 ind); Amb1 Foz3, 19°42'12,86" S 39°49'19,48" W, 16m (1 ind); Amb1 Foz4, 19°37'21,13" S 39°45'6,41" W, 15m (99 ind); Amb1 Foz5, 19°32'32,82" S 39°41'34,89" W, 20m (11 ind); Amb1 Foz6, 19°54'56,16" S 39°56'40,83" W, 34m (4 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (1 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (6 ind); Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (1 ind); Amb1 Foz10, 19°35'12,39" S 39°38'33,16" W, 32m (2 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (6 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (2 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (5 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (9 ind); Amb2 Foz1, 19°52'14,57" S 39°59'41,06" W, 27m (1 ind); Amb2 Foz3, 19°42'8,87" S 39°49'29,72" W, 16m (1 ind); Amb2 Foz4, 19°37'12,91" S 39°45'11,36" W, 13m (24 ind); Amb2 Foz5, 19°32'28,16" S 39°41'35,15" W, 20m (8 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 34m (1 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (6 ind); Amb2 Foz13, 19°47'22,52" S 39°43'20,72" W, 41m (1 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (9 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (1 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (5 ind); Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (13 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (4 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (1 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (11 ind); Amb7 F1, 18°42'56,07" S 39°31'28,8" W, 25m (6 ind); Amb7 G1, 18°32'3,6" S 39°23'16,91" W, 25m (2 ind); Amb13 G1, 18°32'3,21" S 39°23'17,35" W, 25m (1 ind); Amb13 G2, 18°36'32,45" S 39°9'32,83" W, 40m (1 ind); Amb14 B1, 20°34'34,37" S 40°20'50,77" W, 26m (3 ind); Amb14 D1, 19°35'35,96" S 39°41'20,61" W, 26m (2 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (1 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (1 ind); Amb14 E1, 19°9'51,36" S 39°29'20,53" W, 27m (1 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (1 ind); Amb14 F1, 18°42'54,91" S 39°31'28,99" W, 25m (1 ind).

181

Comparative material examined. Apoprionospio dayi USNM 39487 (holotype), USNM 39488 (5 paratypes).

Description: A large-sized Apoprionospio species, largest complete specimen 11mm long, 0.5 mm wide at the widest point for 56 chaetigers. Body dorsoventrally flattened for about 10 anterior chaetigers and rounded afterwards, tapering towards the pygidium. Body color light yellow to whitish in alcohol (Fig 38).

Prostomium triangular, blunt anteriorly, extending posteriorly as a narrow keel to the posterior margin of chaetiger 1. Two pairs of eyes in trapezoidal arrangement present on the midregion of the prostomium. Median prostomial peak usually present. Peristomium low, surrounding prostomium and partially fused to the first chaetiger, lacking lateral wings (Figs 38, 39A–B). Grooved palps reaching up to chaetiger 14, lost in most specimens.

Notopodial postchaetal lamellae well-developed and foliaceous from chaetiger 2 to chaetigers 9– 11 (depending on specimen size) (Fig 40A), rounded on following 2–4 chaetigers and subtriangular afterwards, gradually reducing in size towards the posterior region, although remaining subtriangular until last chaetiger (Figs 38, 39A–C). Notopodial prechaetal lamellae well developed from chaetiger 2 to chaetigers 11–15 (depending on specimen size), absent on more posterior chaetigers. Dorsal crest high on chaetiger 7, dorsal crests low on last 10–15 chaetigers (Fig 39C).

Neuropodial postchaetal lamellae triangular with an elongated ventral part on chaetiger 2, rounded on chaetigers 3–5 and elliptical from chaetiger 6 onwards, progressively elongate until chaetigers 12–16, gradually reducing from chaetigers 13–17 onwards, although remaining elliptical until last chaetiger. Neuropodial prechaetal lamellae present as a low flap from chaetiger 2 to chaetigers 11–13.

Chaetae from notopodia organized in three rows from chaetiger 2 to chaetigers 10–14, anterior row of short, narrowly unilimbate and intensely granulated capillaries; middle and posterior rows of long (up to 1.4 times longer than anterior row), narrowly unilimbate and intensely granulated capillaries (Fig 40C). From chaetigers 11–15 onwards, middle row is lacking. Chaetae from neuropodia organized in two rows of narrowly unilimbate and lightly granulated capillaries, posterior row up to twice as long as anterior row (Fig 40D). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner, and less numerous (Fig 40E).

182

Hooks in notopodia starting from chaetigers 15‒39, up to six per fascicle, accompanied by 1–6 non-limbate capillaries, up to two times longer than neuropodial hooded hooks. Hooks in neuropodia starting from chaetigers 11‒18, up to 12 per fascicle, accompanied by 1‒8 non-limbate capillaries. Hooks long-shafted and heptadentate (tridentate in light microscopy), with six very small secondary teeth arranged side by side above main tooth (Figs 39D, 40F). Secondary hood absent. Sabre chaetae consistently from chaetiger 11. Sabre chaetae wide, non-limbate and intensely granulated (except at the base) (Fig 40G).

Branchia present on chaetigers 2–5, apinnate, and cirriform on chaetigers 2–4 and pinnate on chaetiger 5. Branchiae smooth, about the same length of notopodial postchaetal lamellae on chaetiger 2, heavily ciliated and up to twice as long as notopodial postchaetal lamellae on chaetiger 3, heavily ciliated and up to 2.5 times longer than notopodial postchaetal lamellae on chaetiger 4 and up to five times longer than notopodial postchaetal lamellae on chaetiger 5. Pinnate branchiae bearing stacked plate-like pinnules on entire length, except at the tips (Fig 40B). All branchiae completely free from notopodial postchaetal lamellae.

Pygidium bearing one dorsal cirrus and two short and thin ventral cirri.

Sperm plates or oocytes (measuring up to 130 µm), consistently from chaetiger 13, present on all chaetigers afterwards.

Juvenile morphology: In juveniles, the elongated ventral part of the neuropodial postchaetal lamellae from chaetiger 2 is lacking (as in Fig 39A). Juveniles frequently lack eyes.

Methyl green staining pattern: Margins of prostomium, peristomium and neuropodial postchaetal lamellae from posterior region strongly stained; margins of noto- and neuropodial postchaetal lamellae from anterior region weakly stained.

Remarks. Apoprionospio dayi was described by Foster (1969), based on material collected on North Carolina, Northwestern Atlantic Ocean and also recorded on the Gulf of Mexico and the Caribbean Sea. The species was recorded in Brazil by Amaral (1980) to São Paulo state.

A second Apoprionospio species, A. pygmaeus was recorded by Petti & Nonato (2000) to southeast coast of Brazil. Apoprionospio dayi can be distinguished from A. pygmaeus by the lack of dorsal crests, digitiform neuropodial lamellae on anterior chaetigers, which are oval to elliptical in A. dayi and sabre chaetae starting from chaetigers 11–21 (Hartman 1961) or chaetiger 11 (Maciolek 1985), consistently starting from chaetiger 11 in A. dayi (Foster 1969; Maciolek 1985). (Table 3).

Specimens recorded in this work are very similar to Foster (1969) description and Maciolek (1985) redescription of Apoprionospio dayi. Specimens examined in this work agree with all

183 descriptions in having a triangular and anteriorly blunt prostomium, a peristomium partially fused to chaetiger 1 and not forming lateral wings, notopodial lamellae rounded and neuropodial lamellae digitiform on chaetiger 1, notopodial lamellae from anterior chaetigers well-developed and foliaceous, neuropodial lamellae from anterior chaetigers oval to elliptical, high dorsal crest on chaetiger 7, overlap on the starting chaetiger of notopodial and neuropodial hooded hooks, tridentate hooded hooks (see below), sabre chaetae consistently starting from chaetiger 11 and pygidium bearing one dorsal cirrus and two short and thin ventral cirri.

However, noteworthy differences were observed from Foster (1969) and Maciolek (1985) descriptions, such as presence of median prostomial peak – always present in Foster (1969) and usually present in Maciolek (1985) and this work; posterior extension of the prostomium – to the end of chaetiger 1 in Maciolek (1985) and this work and to the end of chaetiger 2 in Foster (1969); distribution of dorsal crests – high on chaetiger 7 and low on last 10–15 chaetigers in this work, present only on chaetiger 7 in Foster (1969) and Maciolek (1985); morphology of neuropodial postchaetal lamellae from chaetiger 2, elongated ventrally in this study and Foster (1969), not elongated in Maciolek (1985); hook morphology – heptadentate in our specimens, tri- to quadridentate in Foster (1969) and tri- to hexadentate in Maciolek (1985), besides a secondary hood on the hooks, absent in specimens from this study and present, albeit reduced in Foster (1969) and Maciolek (1985) (Table 3). Due to the discrepancies observed on Brazilian material, it was decided to use adopt the “cf.” until a taxonomic decision can be reached.

Habitat: coarse sand to fine silt, 15–50 m depth.

Distribution: Northwestern Atlantic Coast, Gulf of Mexico, Caribbean Sea, Southwestern Atlantic Ocean (São Paulo to Espírito Santo states).

Acknowledgments

The authors would like to thank the National Council for Scientific and Technological Development (CNPq) (Grant 43900/2014-0) for the provided financial support. PCP received a fellowship from CNPq (Process: 304321/2017-6). Thanks are due to the Smithsonian Institution for the Kenneth Jay Boss Fellowship in Invertebrate Zoology (National Museum of Natural History) and to Scott Whittaker (National Museum of Natural History), who helped with sample preparation prior to SEM examination.

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Legends of illustrations Figure 1. Map showing sampled area (blue polygon).

Figure 2. Prionospio cf. ehlersi, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 3. SEM of Prionospio cf. ehlersi. A. Whole animal, dorsal view. B. Anterior and middle region, dorsal view. C. Anterior region, dorsal view. D. Anterior fragment, dorsal view. Abbreviations: br, branchiae; dc, dorsal crest; ip, interneuropodial pouch; nola, notopodial lamella; pe, peristomium; pi, pinnule; pr, prostomium.

Figure 4. Prionospio cf. ehlersi. A. Notochaetae from anterior region, third row. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 5. Prionospio cristata, anterior chaetigers lateral view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 6. SEM of Prionospio cristata. A. Anterior and middle regions, dorsolateral view. B. Anterior region, dorsolateral view. C. Anterior region, dorsal view. D. Hooks (hoods removed). Abbreviations: br, branchiae; dc, dorsal crest; ho, hook; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium.

Figure 7. Prionospio cristata. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 8. Prionospio brevipinnulata sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 9. SEM of Prionospio brevipinnulata sp. nov. A. Anterior and middle region, dorsal view. B. Anterior fragment, dorsal view. C. Anterior region, dorsal view. D. Anterior fragment, dorsolateral view. Abbreviations: br, branchia; nola, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 10. Prionospio brevipinnulata sp. nov. A. Chaetae from anterior region. B. Row of long neurochaetae from chaetigers 2–4. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae. F. Pinnate branchiae.

Figure 11. Prionospio capixaba sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; pe, peristomium; pr, prostomium.

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Figure 12. SEM of Prionospio capixaba sp. nov. A. Anterior and middle region, dorsal view. B. Anterior region, dorsal view. C. Anterior region, dorsal view. D. Anterior region, lateral view. Abbreviations: dc, dorsal crest; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium.

Figure 13. Prionospio capixaba sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 14. Prionospio quadrilamellata sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 15. SEM of Prionospio quadrilamellata sp. nov. Anterior region, dorsal view. B. Anterior fragment, dorsal view. C. Anterior region, dorsal view. D. Posterior region, dorsolateral view. Abbreviations: dc, dorsal crest; hh, hooded hook; nola, notopodial lamella; pe, peristomium; pr, prostomium; as, sabre chaetae.

Figure 16. Prionospio quadrilamellata sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae. F. Apinnate branchiae.

Figure 17. Prionospio alexandrae sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 18. SEM of Prionospio alexandrae sp. nov. A. Anterior and middle region, dorsal view. B. Anterior region, dorsal view. C. Anterior region, dorsolateral view. D. Close-up of anterior region, dorsal view. E. Anterior and middle region, lateral view. F. Palp, lateral view. Abbreviations: br, branchiae; nola, notopodial lamella; pa, palp; pe, peristomium; pp, palp papillae; pr, prostomium.

Figure 19. Prionospio alexandrae sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Row of long neurochaetae from chaetigers 2–4. D. Capillary from posterior region. E. Hooded hook. F. Apinnate branchiae.

Figure 20. Prionospio triangularis sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; dc, dorsal crest; pe, peristomium; pr, prostomium.

Figure 21. SEM of Prionospio triangularis sp. nov. A. Anterior region, dorsal view. B. Anterior and middle regions, lateral view. C. Anterior region, lateral view. D. Hooks (hoods removed).

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Abbreviations: br, branchiae; dc, dorsal crest; ho, hook; nela2, neuropodial lamellae from chaetiger 2; nola, notopodial lamellae; pr, prostomium.

Figure 22. Prionospio triangularis sp. nov. A. Neuropodial postchaetal lamellae from chaetiger 2. B. Notochaetae from anterior region. C. Neurochaetae from anterior region. D. Capillary from posterior region. E. Hooded hook. F. Sabre chaetae.

Figure 23. Prionospio posterocristata sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; dc, dorsal crests; pe, peristomium; pr, prostomium.

Figure 24. SEM of Prionospio posterocristata sp. nov. A. Anterior and middle region, dorsal view. B. Anterior region, dorsal view. C. Anterior region, lateral view. D. Hook (hood removed). Abbreviations: br, branchiae; dc, dorsal crest; ho, hook; neuropodial lamellae from chaetiger 2; nola, notopodial lamellae; pr, prostomium.

Figure 25. Prionospio posterocristata sp. nov. A. Neuropodial postchaetal lamellae from chaetiger 2. B. Notochaetae from anterior region. C. Neurochaetae from anterior region. D. Capillary from posterior region. E. Hooded hook. F. Sabre chaetae.

Figure 26. Prionospio absensi sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 27. SEM of Prionospio absensi sp. nov. A. Anterior and middle regions, dorsal view. B. Anterior region, dorsal view. C. Middle region, dorsal view. D. Hooks (hoods removed). Abbreviations: br, branchiae; ho, hook; nola, notopodial lamella; pe, peristomium; pin, pinnula; pr, prostomium.

Figure 28. Prionospio absensi sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 29. Prionospio biancoi sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 30. SEM of Prionospio biancoi sp. nov. A. Whole fragmente, dorsal view. B. Anterior region, dorsal view. C. Close-up of anterior region, dorsal view. D. Hook (hood removed). br, branchiae; ho, hook; pe, peristomium; pr, prostomium.

Figure 31. Prionospio biancoi sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

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Figure 32. Aurospio dibranchiata, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 33. SEM of Aurospio dibranchiata. A. Whole fragment, dorsal view. B. Anterior region, dorsal view. C. Branchiae. D. Hooks and sabre chaetae. E. Hooks (hoods removed). F. Pygidium. Abbreviations: br, branchiae; hh, hooded hooks; ho, hooks; pe, peristomium; pr, prostomium; prob, proboscis; pyg, pygidium; sa, sabre chaetae.

Figure 34. Aurospio dibranchiata. A. Anterior region. B. Notopodial postchaetal lamellae from chaetigers 3 and 4. C. Parapodia from posterior region. D. Chaetae from anterior region. E. Capillary from posterior region. F. Hooded hook. G. Sabre chaetae. H. Pygidium. Figures A, B, F, G, and H redrawn from Maciolek (1981a); figure C modified from Maciolek (1981a).

Figure 35. Paraprionospio anterocristata sp. nov., anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 36. SEM of Paraprionospio anterocristata sp. nov. A. Anterior region, dorsal view. B. Close-up of anterior region, dorsal view. C. Anterior and middle regions, ventral view. D. Anterior region, lateral view. E. Middle region, dorsal view. F. Hooks (hoods removed). Abbreviations: br, branchiae; dc, dorsal crest; dp, dorsal plate; gi, gizzard-like structure; ho, hook; par, parasite; pe, peristomium; pr, prostomium; tr, transverse ridge; vc, ventral crest.

Figure 37. Paraprionospio anterocristata sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae. F. Pinnate branchiae.

Figure 38. Apoprionospio cf. dayi, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 39. SEM of Apoprionospio cf. dayi. A. Anterior region, lateral view. B. Anterior region, dorsal view. C. Posterior region, dorsal view. D. Hook (hood removed). Abbreviations: br, branchiae; dc, dorsal crest; hh, hooded hook; ho, hook; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium; pyg, pygidium; sa, sabre chaetae.

Figure 40. Apoprionospio cf. dayi. A. Chaetiger 2. B. Pinnate branchiae. C. Notochaetae from anterior region, third row. D. Neurochaetae from anterior region. E. Capillary from posterior region. F. Hooded hook. G. Sabre chaetae.

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Tables

Table 1 – Comparative diagnostic characters of Prionospio species from this work.

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Table 1 (continued) – Comparative diagnostic characters of Prionospio species from this work.

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Table 1 (continued) – Comparative diagnostic characters of Prionospio species from this work.

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Table 1 (continued) – Comparative diagnostic characters of Prionospio species from this work.

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Table 2 – Comparative diagnostic characters of Paraprionospio species reported to Brazil.

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Table 2 (continued) – Comparative diagnostic characters of Paraprionospio species reported to Brazil.

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Table 2 (continued) – Comparative diagnostic characters of Paraprionospio species reported to Brazil.

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Table 3 – Comparative diagnostic characters of Apoprionospio species reported to Brazil.

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Table 3 (continued) – Comparative diagnostic characters of Apoprionospio species reported to Brazil.

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Capítulo 5

Contributions to the taxonomy of Aonides, Aonidella and Laonice from southeastern Brazil, including the description of five new species

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista-alvo: Zootaxa Status: A submeter

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Contributions to the taxonomy of Aonides, Aonidella and Laonice from southeastern Brazil, including the description of five new species

Running title: AONIDES, AONIDELLA AND LAONICE FROM BRAZIL

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract 11 species belonging to the spionid genera Aonides, Aonidella and Laonice were recorded from the Espírito Santo Basin, southeastern Brazil. Five of these species – Aonides sarsi sp. nov., A. curvus sp. nov., Laonice kinbergi sp. nov., L. profunda sp. nov. and L. brevicarunculata sp. nov. are new to science. Species can be separated based on prostomium shape, parapodial lamellae shape, branchiae distribution, presence and distribution of dorsal crests, starting chaetiger of interneuropodial pouches and starting chaetiger and morphology of hooded hooks. An identification key and a comparison table to Laonice species recorded in this work are provided. This work represents the first record of the genus Aonidella to the Brazilian coast.

Keywords: Annelida, morphology, biodiversity, Espírito Santo Basin.

Introduction

Spionidae Grube, 1850, is one of the most diverse and speciose annelid families, potentially being the doming group in deep-sea sediments (Blake et al. 2017) or disturbed environments. There are approximately 580 described species organized in 39 genera (Blake et al. 2017), although the family is most likely much more diverse, as spionids possess a history of species-complexes (Sato-Okoshi et al. 2016), species with doubtful cosmopolitan status, genera that never have been revised and the family itself might not be monophyletic, as it has been suggest that the families Uncispionidae Green, 1982, Trochochaetidae Pettibone, 1963 and Poecilochaetidae Hannerz, 1956 are also part of Spionidae (Struck 2017).

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To the Brazilian coast, 90 species distributed in 18 valid genera have been recorded (Amaral et al. 2013), with most taxonomic works focused on shallow environments (except for Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019, ms) and are restricted to southeastern and southern Brazil (Lana et al. 2017). A few genera have been better studied: Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Polydora Bosc, 1802 (Radashevsky et al. 2006), Laonice Malmgren, 1867 (Kinberg 1866; Nonato et al. 1986; Radashevsky & Lana 2009), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017), Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), and Laubieriellus Maciolek, 1981 (Peixoto & Paiva 2019).

Only a few taxonomic works focus on multiple genera, although lacking the description of new species : Bolívar & Lana (1987) to the continental shelf of the state of Paraná, southern Brazil, Paiva & Barroso (2010) to the continental slope of Campos Basin, state of Rio de Janeiro, southeastern Brazil, as well as and Pardo et al. (2006) and Pardo & Peixoto (in press) to southern and southeastern Brazil.

Aonides Claparède, 1864 is a small genus, comprising only nine species found in all Oceans, from the subtidal to abyssal depths. Besides the reproductive biology, little is known about the biology of most species, which never were redescribed since their original description. The genus is easily identified due to the prostomial shape, rarely being mistaken for other spionid genus, although Aonides species can be confused with Levinsenia Mesnil, 1897, a Paraonidae Cerruti, 1909 (pers. obs.). According to Blake et al. (2017), the type species, A. oxycephala (Sars, 1862), described to Norway, was reported to the Pacific Ocean, Eastern Coast of Africa, Mediterranean Sea, Black Sea and Brazil (Amaral et al. 2013) and most likely comprehends many morphologically similar and undescribed species. Four Aonides species are reported from the Brazilian Coast: Aonides californiensis Rioja, 1947, A. mayaguezensis Foster, 1969, A. oxycephala and A. paucibranchiata Southern, 1914 (Amaral et al., 2013).

The genus Aonidella López-Jamar, 1989 is superficially similar to Prionospio, although Aonidella can be distinguished from Prionospio due to parapodial lamellae of similar size and morphology throughout body length; triangular postchaetal notopodial lamellae and slender postchaetal neuropodial lamellae; prostomium flat, lacking caruncle; very wide angle between the main tooth and smaller teeth on the hooded hooks; sabre chaetae usually absent; rows of holes on the dorsum of branchiate chaetigers and pygidium bearing 4–6 cirri of unequal length. Aonidella shares characters with Aonides, such as branchiae morphology, pygidium morphology and the wide angle between the main tooth and smaller teeth on the hooded hooks. However, Aonides species bear a conical prostomium, tapered on both ends.

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The genus authorship is disputed, since Maciolek (1983) described the genus in her Ph.D. dissertation, transferring Prionospio cirrobranchiata Day, 1961 to Aonidella and describing Aonidella dayi López-Jamar, 1989, although she only published her findings and designated type material years later (Maciolek 2000). The species A. dayi was recorded to Cádiz Gulf, Iberian Peninsula by López-Jamar (1989), who attributed the genus and species’ authority to Maciolek (1983), the first report of both genus and species in published scientific records.

Since López-Jamar (1989) referenced her thesis, Maciolek (2000) defends that the genus authorship should be “Maciolek in López-Jamar, 1989” (Maciolek 2000; Blake et al. 2017). However, as noted by the World Register of Marine Species webpage (Read & Fauchald 2020), López-Jamar (1989) only referenced unpublished data from Maciolek (1983), but did not use the diagnostic characters present in this work, thus advocating that the correct authorship should be attributed to López-Jamar (1989). Only three species are known, found on all Oceans, from the continental shelf to the continental slope.

The genus Laonice is superficially similar to Prionospio, albeit Laonice species typically are larger and more robust, with branchiae present until more posterior chaetigers, possess occipital tentacle and presence of lateral pouches in almost all species.

Worldwide there are 36 described species, ranging from sublitoral to abyssal depths. According to records compiled by Amaral et al. (2013), eight species are recorded for the Brazilian Coast: Laonice antarcticae Hartman, 1953, L. brevicornis (Kinberg, 1866) (including L. aperata Radashevsky & Lana, 2009 and L. petersenae Radashevsky & Lana, 2009, as subjective synonyms), L. branchiata Nonato et al., 1986, L. cirrata (Sars, 1851), L. japonica (Moore, 1907), L. parvabranchiata Radashevsky & Lana, 2009, L. pinnulata Radashevsky & Lana, 2009 and L. weddellia Hartman, 1978.

Material and Methods Sediment samples were collected during the oceanographic cruises of the AMBES Project on the Espírito Santo Basin and northern portion of Campos Basin, southeastern Brazil, from 2010 to 2013 (Fig 1). Samples were collected during the summer (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and winter (sample codes Amb2, Amb11, Amb12, Amb13, and Amb14) using a Van Veen Grab sampler on the continental shelf and a box corer on the continental slope and fixed in situ in 10% formalin-seawater solution. Then, samples were washed, elutriated on a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm-mesh sieve (for samples collected on the continental slope

231 and on the Watu Norte and Doce canyons), and kept in 70% ethanol. Since specimens were fixed in 10% formalin, molecular studies were not possible.

Afterwards, specimens were sorted out and examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope. Measurements were taken using a calibrated eyepiece attached to the microscope. For Aonides species recorded on the Brazilian coast in this study and on previous works (Amaral et al. 2013), main diagnostic characters were summarized in Table 1. A comparison table for Laonice species recorded in this work is also provided (Table 2). For Laonice species, an identification key is also provided.

Specimens lots deposited in the Smithsonian National Museum of National History (USNM), Smithsonian Institution, Washington, D.C., USA, corresponding to type material of Aonidella dayi (USNM 80476, holotype), Aonides mayaguezensis (USNM 39486, holotype), Laonice brevicornis (USNM 100447, 1 paratype of L. aperata) and Laonice weddellia (USNM 46700, holotype; USNM 46701, 20 paratypes) were examined and compared with material collected in this study.

Specimens from this work were stained in a methyl green solution (1 g/L in 70% EtOH) for 10‒ 30 s and examined in distilled water (to slow down stain fading). The stain enhances the visualization of morphological features, such as dorsal crests, lateral pouches and branchial scars, as well as reveal species-specific patterns.

Some specimens were prepared for scanning electron microscope (SEM) studies, being dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each) followed by two changes of 100% ethanol. Specimens were critical point dried in CO2 or dried in an ascending hexamethyldisilane (HMDS) series (2 parts 100% EtOH:1 part HMDS, 1:1, 1:2, pure HMDS, followed by a second change of pure HMDS, 15 min each), and kept in a fume hood until completely dry. The HMDS ascending series was used to avoid damage to small or delicate specimens, which sometimes occurred when using CO2 drying. Next, specimens were mounted on stubs, sputter coated with gold, and viewed in a JEOL JSM-6390LV SEM or a JEOL JSM- 6510 LV SEM.

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Results

Taxonomy

Phylum Annelida Lamarck, 1809

Order Spionida Fauchald, 1977

Family Spionidae Grube, 1850

Genus Aonides Claparède, 1864

Type species: Aonides auricularus Claparède, 1864 [= A. oxycephala (Sars, 1862)], by monotypy.

Synonym list: Paranerine Czerniavsky, 1881 (subjective synonym).

Diagnosis (modified from Blake et al. 2017) Prostomium conical, tapered both anteriorly and posteriorly; lacking pointed tip; eyespots present or absent; occipital antenna present or absent.

Peristomium poorly developed, not fused to chaetiger 1. Postchaetal lamellae developed on posterior chaetiger, usually cirriform. Branchiae from chaetiger 2, limited to a variable number of anterior chaetigers, absent posteriorly; all apinnate, smooth, cirriform, separated from notopodial postchaetal lamellae. Anterior chaetae all capillaries; posterior chaetae include capillaries and bi- or tridentate hooded hooks. Hooks lacking secondary hooded, sometimes with strongly curved shaft. Sabre chaetae present or absent. Pygidium bearing 4–12 anal cirri.

Aonides sarsi sp. nov.

(Figs 2–4G)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 11 Dec 2010 to 19 Dec 2010, 33m, MZUSP XXXX. Paratypes: Amb7 A1, 21°2'45,81" S 40°32'29,21" W, 02 Dec 2011 to 02 Feb 2012, 25m, MZUSP XXXX (2 ind), MNRJP XXXX (2 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 02 Dec 2011 to 02 Feb 2012, 26m, ZUEC-POL XXXX (3 ind).

233

Additional material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (1 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (5 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (5 ind); Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (1 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (1 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (2 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (3 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (9 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (5 ind); Amb1 Foz19, 19°46'14,99" S 39°34'50,94" W, 52m (1 ind); Amb2 Foz2, 19°47'12,95" S 39°55'15,42" W, 23m (2 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 34m (2 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (2 ind); Amb2 Foz8, 19°44'44,06" S 39°46'32,01" W, 31m (2 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (2 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (1 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (3 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (5 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (5 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (1 ind); Amb7 A1, 21°2'45,81" S 40°32'29,21" W, 25m (6 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (4 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (1 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (11 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (4 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (1 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (2 ind); Amb14 C1, 20°10'2,67" S 40°8'31,96" W, 26m (1 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (3 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (6 ind); Amb14 E1, 19°9'51,36" S 39°29'20,53" W, 27m (2 ind).

Comparative material examined. USNM 39486 (paratype of Aonides mayaguezensis, 1 specimen).

Description: A large-sized Aonides species, largest complete specimen 9.5 mm long, 0.5 mm wide at the widest point for 167 chaetigers, holotype 9 mm long, 0.45 mm wide at the widest point for 134 chaetigers. Body rounded on the branchial region and slightly dorsoventrally flattened on the remaining chaetigers, tapering towards the pygidium. Body color light yellow to whitish in alcohol (Fig 2).

Prostomium conical, tapered in both ends, narrow anteriorly, but lacking an anteriorly pointed tip; widest at midregion; posterior end reaching end of chaetiger 1. Caruncle absent. Two pairs of eyes in trapezoidal arrangement on the posterior half of the prostomium, anterior pair barely visible. Digitiform occipital antenna present on the posterior-most part of the prostomium (Fig 2, 3B–C, 4A). Peristomium weakly developed, surrounding prostomium laterally and forming a ring around the mouth, not fused with chaetiger 1. Palps lost in all specimens.

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Chaetiger 1 bearing only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae foliaceous on notopodium and digitiform on neuropodium, smaller than lamellae on succeeding chaetigers, especially on the neuropodium. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous, broad and greatly elongated on the branchial region, neuropodial postchaetal lamellae cirriform on chaetiger 2 (Fig 4B) and foliaceous on the remaining branchiate chaetigers, although narrower and less developed than notopodial lamellae (Figs 2, 3A–B, 4A).

On the postbranchial region, notopodial postchaetal lamellae abruptly reduced, foliaceous, although short and narrow; neuropodial postchaetal lamellae foliaceus, less developed than notopodial lamellae. Both lamellae progressively smaller towards the posterior region. Prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated unilimbate capillaries, notopodial chaetae (Fig 4C) up to 1.2 times longer than neuropodial chaetae (Fig 4D). Towards the posterior region, chaetae of both rami progressively become non-limbate and less numerous (Fig 4E).

Hooks in notopodia from chaetigers 18–44, up to four per fascicle, accompanied by 1–5 non- limbate capillaries. Hooks in neuropodia from chaetigers 12–35, up to six per fascicle, accompanied by 2–5 non-limbate capillaries and 1–3 inferior non-limbate and lightly granulated capillaries, longer and more robust than companion capillaries. Hooks bidentate, slightly curved near the tip. Secondary hood absent (Fig 4F). Sabre chaetae absent throughout.

Branchiae starting from chaetiger 2, from six to 17 pairs (depending on specimen size). Branchiae smooth, cirriform, tapered at the tips and completely free from the notopodial lamellae. Branchiae up to 2 times longer than notopodial postchaetal lamellae, except on the last branchiate chaetiger, where branchia is slightly shorter or as long as notopodial postchaetal lamellae.

Pygidium bearing two pairs of cirri, ventral pair slightly shorter than dorsal pair (Figs 3D, 4G).

Gametes not observed.

Methyl green staining pattern: Margins of lamellae and branchiae weakly stained.

Remarks. Among species recorded in Brazil, Aonides sarsi sp. nov. is similar to A. mayaguezensis, described from the Atlantic Coast of Puerto Rico, A. oxycephala, described from Norway and A. paucibranchiata, described from Ireland and the North Sea in having a conical prostomium, tapered in both ends, narrow anteriorly, lacking an anteriorly pointed tip; foliaceous notopodial lamellae on the branchial region and foliaceous to subtriangular notopodial lamellae

235 on the postbranchial region (Southern, 1914; Foster, 1969; Ramos, 1976; Imajima, 1989) (Table 1).

However, A. sarsi sp. nov. can be distinguished from the aforementioned species by the branchial distribution, up to chaetigers 7–18 in A. sarsi sp. nov., up to chaetigers 15–16 in A. mayaguezensis, up to chaetigers 18–23 in A. oxycephala and up to chaetigers 11–12 in A. paucibranchiata and shape of neuropodial lamellae on both branchial and postbranchial chaetigers, cirriform on chaetigers 2 and foliaceous on the remain chaetigers (reduced in posterior chaetigers) in A. sarsi sp. nov. and from elliptical to subtriangular on the remaining species.

Aonides sarsi sp. nov. further differs from A. oxycephala in the starting chaetigers of neuropodial hooded hooks, from chaetigers 19–23 in A. sarsi sp. nov. and from chaetigers 32–42 in A. oxycephala and in the pygidium morphology bearing four cirri in A. sarsi sp. nov. and 6–10 cirri in A. oxycephala (Sars, 1862). The species also differs from A. paucibranchiata in having bidentate hooded hooks (shared with A. mayaguezensis and A. oxycephala) instead of tridentate hooded hooks (Southern, 1914; Brito et al. 2006).

Among species recorded in this work, A. sarsi sp. nov. differs from A. curvus sp. nov. in being much larger; having an anteriorly narrow prostomium; presence of occipital tentacle; branchiae distributed on more chaetigers; shape of neuropodial lamellae on the branchial region; starting chaetiger of notopodial and neuropodial hooded hooks and presence of slightly curved bidentate hooded hooks.

The species is unusual in having inferior chaetae accompanying the neuropodial hooded hooks. According to Radashevsky (2015), these chaetae can be referred as sabre chaetae in posterior neuropodia of large specimens, present only in A. orensanzi Radashevsky, 2015, described from Australia. However, species can be distinguished by the distribution of notopodial hooded hooks – from chaetigers 18–44 in A. sarsi sp. nov. and from chaetigers 20–25 in A. orensanzi; distribution of neuropodial hooded hooks – from chaetigers 12–35 A. sarsi sp. nov. and from chaetigers 16–22 in A. orensanzi; branchial distribution – up to 17 pairs in A. sarsi sp. nov. and up to 14 pairs in A. orensanzi and pygidium morphology – four cirri in A. sarsi sp. nov. and five cirri in A. orensanzi.

Etymology. The specific epithet, sarsi, is a tribute to Michael Sars (1809–1869), a Norwegian naturalist who described the first Aonides species (as Nerine oxycephala).

Habitat: coarse sand to fine sand, 23–55 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and São Paulo states), Atlantic Ocean.

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Aonides curvus sp. nov.

(Figs 5–7F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 02 Dec 2011 to 02 Feb 2012,153m, MNRJP XXXX. Paratypes: Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 02 Dec 2011 to 02 Feb 2012,153m, MNRJP XXXX (3 ind), MZUSP XXXX (3 ind), ZUEC-POL XXXX (4 ind).

Additional material examined. Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (1 ind); Amb2 Foz2, 19°47'12,95" S 39°55'15,42" W, 23m (1 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 34m (3 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (2 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (15 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (1 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (25 ind); Amb7 A1, 21°2'45,81" S 40°32'29,21" W, 25m (1 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (6 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (7 ind); Amb8 F4, 19°32'57,67" S 38°42'57,83" W, 152m (5 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (11 ind); Amb12 E5, 19°36'30,6" S 39°10'19,39" W, 360m (1 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (2 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (1 ind); Amb14 F4, 19°33'2,92" S 38°42'52,26" W, 160m (3 ind).

Description: A small-sized Aonides species, largest complete specimen 2 mm long, 0,07 mm wide at the widest point for 59 chaetigers, holotype complete, 1.8 mm long, 0.65 mm wide at the widest point for 47 chaetigers. Body slightly dorsoventrally flattened on the branchial region and rounded on the remaining chaetigers, tapering towards the pygidium. Body color whitish or translucent white in alcohol (Fig 5).

Prostomium slightly conical, tapered in both ends, rounded at the tip; widest at midregion; posterior end reaching end of chaetiger 1 (Fig 5, 6A–B, 7A). Caruncle, eyes and occipital antennae absent. Peristomium weakly developed, surrounding prostomium laterally and forming a ring around the mouth, not fused with chaetiger 1. Grooved palps reaching up to chaetiger 8, lost in most specimens.

Chaetiger 1 bearing only a few chaetae in both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and digitiform on neuropodium, much smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

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On the branchial region, notopodial postchaetal lamellae foliaceous, narrowing from chaetiger 6 onwards, neuropodial postchaetal lamellae rounded on chaetigers 2–3 (Fig 7B), cirriform on chaetigers 4–6, foliaceous and narrow starting on chaetiger 7, less developed than notopodial lamellae.

On the postbranchial region, both postchaetal lamellae short, narrow and foliaceous, abruptly reduced and then gradually reducing towards the posterior region (Fig 6A–B); neuropodial postchaetal lamellae less developed than notopodial lamellae. Prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated unilimbate capillaries (narrow limbation), notopodial and neuropodial chaetae of the same length (Fig 7C). Towards the posterior region, chaetae of both rami progressively become non-limbate and less numerous (Fig 7D).

Hooks in notopodia from chaetigers 13–21, up to three per fascicle, accompanied by 1–3 non- limbate capillaries (Fig 6C). Hooks in neuropodia from chaetigers 9–17, up to four per fascicle, accompanied by 1–3 non-limbate capillaries. Hooks tridentate, with very small secondary teeth; strongly bent near the base (Fig 7E). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae starting from chaetiger 2, from three to six pairs (depending on specimen size). Branchiae smooth, cirriform, tapered at the tips and completely free from the notopodial lamellae. Branchiae short, about the same length of the notopodial lamellae, much shorter on the last branchiate chaetiger.

Pygidium bearing two pairs of cirri, ventral pair slightly shorter than dorsal pair (Fig 7F).

Gametes not observed.

Variation: In juveniles, neuropodial postchaetal lamellae may be absent on last 3–5 chaetigers.

Methyl green staining pattern: Prostomium, peristomium and notopodial lamella from branchial region strongly stained, body weakly stained.

Remarks. Aonides curvus sp. nov. differs from most Aonides species recorded in Brazil in having an anteriorly rounded prostomium; lack of occipital tentacle; very short branchial distribution – up to only six pairs of branchiae; morphology of neuropodial postchaetal lamellae on the branchial region – rounded on chaetigers 2–3 and cirriform on chaetigers 4–6; notopodial and neuropodial hooded hooks starting from more anterior chaetigers, being tridentate and strongly bent near the base.

Regarding species recorded in Brazil, A. curvus sp. nov. is most similar to A. paucibranchiata, described from the Belgian part of the North Sea, in lacking an occipital tentacle and bearing

238 tridentate hooded hooks, although species can be distinguished based on prostomium morphology – rounded in A. curvus sp. nov. and narrowed, lacking a pointed tip in A. paucibranchiata; branchial distribution – up to six pairs of branchiae in A. curvus sp. nov. and up to 11 pairs of branchiae in A. paucibranchiata, morphology of neuropodial postchaetal lamellae on the branchial region; starting chaetiger of notopodial hooded hooks – from chaetigers 13–21 in A. curvus sp. nov. and chaetiger 36 in A. paucibranchiata; starting chaetiger of neuropodial hooded hooks – from chaetigers 9–17 in A. curvus sp. nov. and chaetiger 31 in A. paucibranchiata and hook morphology – strongly bent in A. curvus sp. nov. and straight A. paucibranchiata (Southern 1914) (Table 1).

Interestingly, two additional Aonides species also possess an anteriorly rounded prostomium, lack of eyes, lack of occipital tentacle, very short branchial distribution, notopodial and neuropodial hooded hooks starting from more anterior chaetigers and strongly bent tridentate hooded hooks – Aonides selvagensis Brito et al. 2006, from the Macaronesian region of Portugal and Spain and A. nodosetosa Störch, 1966 from Red Sea Coast of Egypt. Despite sharing many characters not observed in other Aonides species, it is unclear it these species represent a clade within Aonides.

Species can be distinguished by the morphology of notopodial postchaetal lamellae – foliaceous on branchial region and foliaceous, short, narrow and reduced on the postbranchial region in A. curvus sp. nov., cirriform on branchial region and conical on the postbranchial region in A. selvagensis and elongated throughout in A. nodosetosa and morphology of neuropodial postchaetal lamellae – rounded on chaetigers 2–3 and cirriform on chaetigers 4–6 on branchial region and foliaceous, short and reduced on the postbranchial region in A. curvus sp. nov., rounded on branchial region and conical on the postbranchial region in A. selvagensis and elongated in both rami in A. nodosetosa (Brito et al. 2006).

These species can also be distinguished based on the starting chaetiger of notopodial hooded hooks – from chaetigers 13–21 in A. curvus sp. nov., from chaetiger 13 in A. selvagensis and from chaetigers 9–13 in A. nodosetosa; starting chaetiger of neuropodial hooded hooks – from chaetigers 9–17 A. curvus sp. nov., from chaetiger 11 in A. selvagensis and from chaetigers 8–13 in A. nodosetosa and pygidium morphology – bearing four cirri in A. curvus sp. nov. and five cirri in A. selvagensis and A. nodosetosa.

Etymology. The specific epithet, curvus (Latin adjective for curved or bent), refers to the morphology of hooded hooks, strongly bent near the base.

Habitat: coarse sand to fine silt, 23–410 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

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Genus Aonidella López-Jamar, 1989

Type species: Aonidella cirrobranchiata (Day, 1961) (superseded original combination)

Diagnosis (modified from Blake et al. 2017): Prostomium broad, flattened anteriorly, sometimes weakly notched on anterior margin, lacking posterior caruncle but with posterior margin completely fused with dorsum; paired nuchal organs present extending posteriorly from prostomium on to chaetiger 21; occipital tentacle absent; eyespots present or absent. Peristomium partly fused to chaetiger 1. Dorsum flat, without dorsal crests. Branchiae from chaetiger 2, limited to anterior chaetigers, 10–16 pairs; all simple, apinnate, elongate, separate from notopodial lamellae. Notopodial postchaetal lamellae triangular, similar along body; neuropodial postchaetal lamellae narrower. Prechaetal lamellae absent. Interparapodial lateral pouches present or absent.

Chaetae of two types: limbate capillaries anteriorly; bi-, tri- or quadridentate hooded hooks with somewhat perpendicular apical teeth and small secondary hood in posterior noto- and neuropodia.

Ventral sabre chaetae usually absent. Dorsal crests absent. Rows of small holes or pits on dorsum of chaetigers of the branchiate region or adjacent to it. Pygidium with 4–6 subequal anal cirri.

Aonidella cf. cirrobranchiata (Day, 1961)

(Figs 8–9F)

Material examined. Amb4 F10, 20°46'23,65" S 38°17'17,65" W, 3004M (4 ind); Amb5 C9, 20°48'39,87" S 38°45'23,86" W, 2496m (1 ind); Amb6 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (1 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (1 ind) Amb11 A9, 21°9'39,38" S 38°52'7,25" W, 2502m (2 ind).

Description: A small-sized Aonidella species, largest incomplete specimen 3.2 mm long, 0.3 mm wide at the widest point for 26 chaetigers. Body slightly dorsoventrally flattened throughout. Body color light white in alcohol (Fig 8).

Prostomium broad, flattened anteriorly, laterally separated from the peristomium, with a by a small median furrow on the anterior margin and bifid projection extending posteriorly to the end of chaetiger 1. Eyes lacking. Nuchal organs not observed. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 8, 9A). Palps lost in all specimens.

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Chaetiger 1 bearing only a few chaetae on both rami. Postchaetal lamellae triangular on both rami, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent. Interparapodial pouches absent.

Postchaetal lamellae foliaceous on both rami, more developed on the branchial region (Fig 9B). Lamellae gradually reducing in size, although preserving its shape in postbranchial chaetigers. Prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of slightly granulated and narrowly limbate (limbation almost imperceptible) capillaries, anterior row short (Fig 9C) and posterior row long (up to 1.5 times longer than anterior row) (Fig 9D). Towards the posterior region, chaetae of both rows progressively become non-granulated, elongate, non-limbate and less numerous (Fig 9E).

Hooks not observed in notopodia, absent for at least 26 chaetigers (longest fragment). Hooks in neuropodia from chaetiger 16–17, up to three per fascicle, accompanied by 1–4 non-limbate capillaries. Hooks bidentate, with an obtuse angle between the main tooth and secondary tooth (Fig 9F). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae starting from chaetiger 2, from eight to 10 pairs (depending on specimen size). Branchiae smooth, cirriform, delicate, tapered at the tips and completely free from the notopodial lamellae. First and last branchial pairs reduced, about the same size of the notopodial postchaetal lamellae; remaining branchiae up to two times longer than notopodial postchaetal lamellae.

Pygidium of unknown morphology.

Gametes not observed.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Aonidella cirrobranchiata (Day, 1961) was described based on material collected on the continental shelf of South Africa. The species was described as Prionospio cirrobranchiata by Day (1961), who noted that the species differed from all multibranchiate Prionospio species in having bidentate hooded hooks and postchaetal lamellae of the same morphology throughout the body. The species was placed in the subgenus Minuspio by Foster (1971) and posteriorly referred to Aonidella by Maciolek (1983, 2000).

Aonidella cirrobranchiata was recorded in South Africa (Day 1961), Antarctica (López 2010), Grand Caribbean (Delgado-Blas & Salazar-Silva 2011) and Bay of Biscay (Maciolek 2000). Specimens collected in this study represent the deepest record for this species, 2503m, if our specimens in fact corresponds to A. cirrobranchiata, which might not be the case. Not only there

241 is a low number of collected specimens – all short anterior fragments in poor condition, but there are also discrepancies from the original description and posterior records.

While our specimens agree with all descriptions (Day 1961; Maciolek 2000; López 2010) regarding the presence of foliaceus postchaetal lamellae on both rami throughout the body and in having bidentate hooded hooks, our specimens differed on the starting chaetiger of neuropodial hooded hooks and branchiae distribution.

In all aforementioned descriptions, neuropodial hooded hooks started on chaetiger 18–19, while in our specimens, neuropodial hooks started on chaetiger 16–17. As for the branchial distribution, Day (1961) reported 11–12 pairs of branchiae, Maciolek (2000) reported 10–12 pairs, López (2010) reported 10 pairs, whereas 8–10 pairs were reported in this study. López (2010) also reported a red-pigmented notopodial postchaetal lamellae on chaetiger 2, which was lacking in all the remaining descriptions.

These differences could be explained by the specimen’s small size – our specimens are thinner than other records and could represent A. cirrobranchiata juveniles or our specimens could represent a new species, due to the morphological differences coupled with a deeper depth of occurrence and geographic distance from the remaining A. cirrobranchiata records (Day 1961; Maciolek 2000; López 2010; Delgado-Blas & Salazar-Silva 2011). Unfortunately, additional material – preferably complete specimens in good condition would have to be obtained before the status of this taxon could be resolved.

There is a second Aonidella species from the Atlantic Ocean, A. dayi, which also exhibits a wide geographic and bathymetric distribution (see Remarks of Aonidella cf. dayi for details), but species can be separated mainly on hook morphology – bidentate hooded hooks in A. cirrobranchiata and tri-to quinquedentate hooded hooks in A. dayi. Reports of Aonidella cirrobranchiata and A. dayi from this work corresponds to the first record of the genus Aonidella to the Brazilian coast.

Habitat: muddy sand to mud, 1300‒3004 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), South Atlantic Ocean, South Africa (Day, 1961), Antarctica (López, 2010), Grand Caribbean (Delgado-Blas & Salazar- Silva, 2011) and Bay of Biscay (Maciolek, 2000).

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Aonidella cf. dayi López-Jamar, 1989

(Figs 10–12G)

Material examined. Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (5 ind); Amb2 Foz 20, 19°41'24,99" 39°31'20,42" W, 53m (3 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (1 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (1 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (1 ind); Amb7 F4, 19°32'57,67" S 38°42'57,83" W, 152m (11 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (1 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (4 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (6 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (1 ind); Amb14 F4, 19°33'2,92" S 38°42'52,26" W, 160m (10 ind).

Comparative material examined. USNM 80476 (holotype).

Description: A large-sized Aonidella species, largest specimen 19mm long, 0.65 mm wide at the widest point for 68 chaetigers. Body dorsoventrally flattened throughout, especially on the branchial region, tapering towards the pygidium. Body color light yellow to whitish in alcohol (Fig 10).

Prostomium broad, rounded and flattened anteriorly, laterally separated from the peristomium by a small median furrow on the anterior margin and bifid projection extending posteriorly to the end of chaetiger 1 (Figs 10, 11A–B, 12A). Two pairs of eyes in trapezoidal arrangement on the posterior half of the prostomium, anterior pair slightly larger than posterior one. Nuchal organs present as “U” shaped ciliary bands reaching anterior margin of chaetiger 2. Peristomium partly fused to chaetiger 1, lacking lateral wings. Grooved palps reaching up to chaetiger 14, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae triangular on both rami, smaller than lamellae on succeeding chaetigers, especially on the neuropodium. Prechaetal lamellae absent.

Postchaetal lamellae triangular on the branchial region (Figs 11C, 12B), neuropodial lamellae less developed than notopodial lamellae. On the postbranchial region, notopodial postchaetal lamellae subtriangular and neuropodial postchaetal lamellae cirriform, less developed than notopodial lamellae. Lamellae gradually reducing in size, although preserving its shape, towards the posterior region (Fig 11D). Prechaetal lamellae absent throughout. Rows of holes on dorsal side absent.

Chaetae from notopodia and neuropodia organized in two rows of slightly granulated capillaries, anterior row of short unilimbate capillaries (Fig 12C) and posterior row of long (up to 1.5 times

243 longer than anterior row) and narrowly unilimbate capillaries (Fig 12D). Towards the posterior region, chaetae of both rows progressively become non-granulated, elongate, non-limbate and less numerous (Fig 12E).

Hooks in notopodia starting from chaetigers 18–28, up to five per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks in neuropodia starting from chaetiger 13–19, up to six per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks tridentate or quadridentate, with an obtuse angle between the main tooth and secondary teeth (Fig 12F). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae starting from chaetiger 2, from six to 12 pairs (depending on specimen size). Branchiae smooth, cirriform, tapered at the tips and completely free from the notopodial lamellae. First and last branchial pairs reduced, about half the length of the notopodial postchaetal lamellae; remaining branchiae of the same size or slightly shorter than notopodial postchaetal lamellae.

Pygidium bearing two pairs of cirri, ventral pair slightly shorter than dorsal pair (Fig 12G).

Gametes not observed.

Variation: a single incomplete specimen, 0.65 mm wide at the widest point for 49 chaetigers possessed sabre chaetae on the two last chaetigers.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Aonidella dayi was described by Maciolek (1983) on her Ph.D. thesis based on incomplete specimens collected on the east coast of the USA, Canary Islands and the Mediterranean Sea. However, according to The International Code of Zoological Nomenclature (ICZN) (Chapter 3, Articles 8 and 9), this description is not considered valid since it is treated as unpublished work. Posteriorly, López-Jamar (1989) found specimens similar to the description of A. dayi on the Gulf of Cadíz, Iberic Peninsula and described these specimens, including complete individuals, thus being considered as the specie’s author, despite never designating a holotype. Posterior studies of Imajima (1992) and Sigvaldadóttir et al. (1997) considered the genus authorship as Maciolek in López-Jamar, 1989, but never justified its use.

Maciolek (2000) published a part of her Ph.D. thesis, which included an extended description of A. dayi, including complete specimens and attributing the species’ authorship to Maciolek in López-Jamar, 1989, based on the ICZN, an authorship followed by posterior works (Delgado- Blas 2009; López 2010; Delgado-Blas & Salazar-Silva 2011; Simon 2012; Meiβner et al. 2014; Çinar 2005), although, according to the World Register of Marine Species’ website (Read & Fauchald 2020), the authorship is attributed to López-Jamar, 1989, since, according to comments

244 made by the website’s editor, López-Jamar (1989) described his own material from the Gulf of Cadíz, which presented morphological differences from Maciolek’s description (1983), thus treating the authorship attributed to Maciolek in López-Jamar, 1989 as incorrect.

Aonidella dayi was described to the Gulf of Cadíz (López-Jamar 1989) and supposedly is widely distributed, with records from the east coast of the USA, Canary Islands and the Mediterranean Sea (Maciolek 1983); Gulf of Mexico - as Prionospio (Minuspio) cirrobranchiata (Johnson 1984), South Africa and Japan (Imajima 1992); Grand Caribbean (as A. cirrobranchiata; Delgado-Blas & Salazar-Silva 2011); seamounts on northeast Atlantic and Brazil (Meiβner et al. 2014).

The species most likely represents a species-complex, due to discrepancies on the branchiae distribution (Meiβner et al. 2014), postchaetal lamellae morphology, hook morphology, as well as the presence of rows of holes on the dorsum and presence of sabre chaetae on some specimens, besides a wide geographic and bathymetric distribution.

Postchaetal lamellae are described as triangular and of similar size throughout the body (López- Jamar 1989; Maciolek 1983, 2000), while Imajima (1992) described notopodial postchaetal lamellae as triangular on the first two chaetigers and foliaceus on the remaining chaetigers and neuropodial postchaetal lamellae cirriform on the branchial region and triangular on postbranchial chaetigers. In this work, the postchaetal lamellae is triangular on both rami in the branchial region, whereas, in the postbranchial region, lamellae are subtriangular on notopodia and cirriform on neuropodia.

As for the hooded hooks, hook morphology varied from tridentate only (López-Jamar 1989; Imajima 1992; Meiβner et al. 2014), tri- and tetradentate (this work) and tri-, tetra- or quinquedentate (Maciolek 1983, 2000). Notopodial hooded hooks were reported starting from chaetiger 26–37 (López-Jamar 1989), chaetiger 20 (Maciolek 1983, 2000), chaetiger 22–24 (Imajima 1992), chaetiger 19–37 (Meiβner et al., 2014) and chaetigers 18–28 (this study), whereas neuropodial hooded hooks started from chaetiger 16–19 (López-Jamar 1989), chaetiger 14–20 (Maciolek 1983, 2000), chaetiger 18–20 (Imajima 1992), chaetiger 15-22 (Meiβner et al. 2014) and chaetiger 13–19 (this study).

Sabre chaetae were reported by López-Jamar (1989) on the last 10–12 chaetigers of all specimens from the Iberic Peninsula, while Meiβner et al. (2014) reported sabre chaetae on a single specimen, present on the last three chaetigers of a 52-chaetiger fragment.

Branchiae distribution ranged from 12–16 pairs (López-Jamar 1989), 10–12 pairs (Maciolek 1983, 2000), 14–16 pairs (Imajima 1992), 6–12 pairs (this study) and up to 16 pairs, reported by

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Meiβner et al. (2014), which states that branchiae distribution is size-dependent, correlating body width (measured at 10th chaetiger) to the number of branchia. Although the authors only examined two large Brazilian species, both had 12 pairs of branchiae, the upper range found in this study.

Due to differences in the starting chaetiger of hooded hooks, branchiae distribution, postchaetal lamellae morphology and presence of sabre chaetae, it was decided to adopt the “cf.” as it represents a more conservative view. It should be noted, however, that this adoption is a temporary classification, until specimens from the type locality could be examined. Our material also differs from Aonidella cf. dayi described by Meiβner et al. (2014) in having an anteriorly narrow prostomium, laterally widened in Meiβner et al. (2014) and in lacking rows of holes on the dorsum, present on several anterior and middle chaetigers in specimens examined by Meiβner et al. (2014, Fig 4).

A second Aonidella species, A. cirrobranchiata (Day, 1961) is known from the Atlantic Ocean, also with a wide bathymetric and geographic distribution (South Africa [Day, 1961], Bay of Biscay [Maciolek, 1983, 2000] and Antarctica [López, 2010]), but this species can be separated from A. dayi based on hook morphology – bidentate in A. cirrobranchiata and tri- to quinquedentate in A. dayi.

Habitat: medium sand to muddy sand, 50‒171 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), South Atlantic Ocean, Gulf of Cadíz (López-Jamar 1989), east coast of the USA, Canary Islands and the Mediterranean Sea (Maciolek 1983); Gulf of Mexico - as Prionospio (Minuspio) cirrobranchiata (Johnson 1984), South Africa and Japan (Imajima 1992); Grand Caribbean (as A. cirrobranchiata; Delgado-Blas & Salazar-Silva 2011); seamounts on northeast Atlantic and Brazil (Meiβner et al. 2014).

Genus Laonice Malmgren, 1867

Type species: Nerine cirrata Sars, 1851, designated by Malmgren, 1867 [=Laonice cirrata (Sars,

1851)], by monotypy.

Synonym list: Mandane Kinberg, 1866 (subjective synonym)

Spionides Webster & Benedict, 1887 (subjective synonym)

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Aricideopsis Johnson, 1901 (subjective synonym)

Diagnosis (modified from Blake et al. 2017): Prostomium anteriorly rounded to slightly incised, or bell-shaped; nuchal organ(s) extending posteriorly for variable number of chaetigers; occipital tentacle usually present. 1–2 pairs of eyes por eyes absent. Peristomium with reduced lateral wings absent or poorly developed and separated from the prostomium and moderately developed or, in other species enlarged and fused to anterior margin of prostomium. Cirriform branchiae from chaetiger 2, apinnate or with digitiform pinnules, separated from or partially fused to notopodial postchaetal lamellae, continuing posteriorly for at least one-half of body length.

Auricular lobe sometimes present. Notopodial and neuropodial postchaetal lamellae large, expanded in anterior chaetigers, reduced posteriorly. Dorsal crests present or absent.

Interparapodial lateral pouches usually present. Anterior chaetae all capillary, posterior chaetae include capillaries, bi- to multidentate hooded hooks – present only on the neuropodia or in both rami and sabre chaetae. Hooded hooks lacking secondary hood. Pygidium bearing a pair of ventral cirri and a variable number of dorsal cirri.

Laonice cf. antarcticae Hartman, 1953

(Figs 13–15D)

Synonym list: Laonice cirrata antarcticae Hartman, 1953 (superseded original combination)

Material examined. Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (3 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (4 ind); Amb5 B5, 20°35'16,23" S 39°53'47,1" W, 410m (1 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (3 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (1 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (1 ind).

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Description: A medium-sized Laonice species, largest complete specimen 16 mm long, 2.1 mm wide at the widest point for 98 chaetigers. Body slightly dorsoventrally flattened on the branchial region and cylindrical afterwards, tapering towards the pygidium. Dark pigmented stripes on branchiae of the anterior region (Fig 13). Body color whitish to light yellow in alcohol.

Prostomium bell-shaped, broadly rounded anteriorly, with pointed lateral margins, lacking frontal horns, extending posteriorly as a low caruncle to the end of chaetiger 12 (Fig 14C), flanked by U- shaped nuchal organs extending through entire caruncle length. Two pairs of eyes in trapezoidal arrangement, anterior pair small and posterior pair large and recurved or presence of a single large pair of eyes or eyes absent. Digitiform occipital antenna with a pointed tip present on posterior half of the prostomium, near the posterior margin of chaetiger 1 (Figs 13, 14B). Peristomium well-separated from prostomium for most of its length, fused only on the anterior margin; not fused to chaetiger 1 and bearing low lateral wings. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamella foliaceous on notopodium and auricular on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae wide and foliaceous on anterior and middle regions (Figs 13, 14A), rounded on the posterior region, greatly reduced on last chaetigers. Notopodial prechaetal lamellae absent throughout. Dorsal crests absent.

Neuropodial postchaetal lamellae foliaceous and inferiorly fused to the body on anterior and middle regions, gradually reducing afterwards, low and rounded on the posterior region, greatly reduced on last chaetigers. Neuropodial prechaetal lamellae absent throughout. Interneuropodial pouches starting from chaetigers 4–5, present on most of the body, absent on last 6–9 chaetigers (Figs 13, 14D).

Chaetae from notopodia and neuropodia organized in two rows of lightly granulated and widely bilimbate capillaries (Fig 15A). Chaetae from both rows of equal length, although notopodial chaetae are up to 1.2 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 15B).

Hooks in neuropodia only, starting from chaetigers 21–38, up to 12 per fascicle, accompanied by 2–8 non-limbate capillaries. Hooks short and tridentate, with two secondary teeth arranged side by side above main tooth (Fig 15C). Secondary hood absent. Sabre chaetae starting from chaetigers 20–25, up to two per fascicle. Sabre chaetae narrowly limbate and lightly granulated (granulation absent at the base) (Fig 15D).

Branchiae from chaetiger 2, present on most of the anterior region, 14–35 pairs, short on first chaetigers and reaching full size around chaetiger 10, decreasing in size on last 5 chaetigers.

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Branchiae short (about the same size or slightly shorter than notopodial postchaetal lamellae), cirriform, moderately ciliated and completely free from notopodial postchaetal lamellae. Accessory branchiae absent.

Pygidium rounded, bearing one pair of short ventral cirri and 2 pairs of long (up to twice as long as ventral cirri) dorsal cirri.

Juvenile morphology: In juveniles, the caruncle is shorter, reaching up to chaetigers 8–11.

Methyl green staining pattern: Anterior half of the prostomium and postchaetal lamellae from anterior and middle regions strongly stained; ventral side of anterior and middle regions weakly stained.

Remarks. This species was originally described as Laonice cirrata antarcticae Hartman, 1953 off the coast of Rio Grande do Sul state, Southern Brazil (type locality mistakenly identified as Uruguay on the original description), being raised to species rank by Hartman (1965).

Blake (1983) expanded the species distribution to Argentina, while also recording the species in the entire Western Atlantic, from North America to Argentina, and Antarctic and subantarctic seas, from 27 to 3697 m depth. Radashevsky & Lana (2009) briefly redescribed the syntypes and also doubted this wide distribution, as the species’ individual and ontogenetic variability is poorly known and no material collected in Brazil during previous research programs entirely matched the species original description (see below), hypothesizing that morphological characters may be more variable than previously thought or more than one species is present in deeper waters.

Specimens in this work are similar to Hartman (1953) original description in having interneuropodial pouches starting from chaetigers 4–5; hooded hooks from chaetigers 21–38 in this work and chaetiger 35 in the original description; first branchial pairs shorter than notopodial postchaetal lamellae; similar branchial length – about the same length of notopodial postchaetal lamellae) and similar number of branchiae 14–35 pairs in this work and 31 pairs in the original description. Our specimens differ from the original description on the starting chaetiger of sabre chaetae – chaetigers 20–25 in this work and from chaetiger 34 in the original description, justifying the “cf” adoption.

Material examined in this work is also similar to Blake (1983) and Radashevsky & Lana (2009) descriptions, based on syntypes; differing in the prostomium morphology – fused on the anterior margin with peristomium on the anterior margin in specimens from this work and well-separated from the peristomium in the syntypes; starting chaetiger of neuropodial hooded hooks – from chaetigers 21–38 in this work and from chaetiger 35–40 in the syntypes; maximum number of branchiae–up to 35 branchial pairs in this work and up to 29 pairs in the syntypes and number of sabre chaetae per rami – up to two in this work and only one in the syntypes.

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The anterior fusion of prostomium and peristomium was noted by Sikorski (2011), who latter (Sikorski et al. 2017) erected four Laonice subgenus and placed Laonice antarcticae in the subgenus Sarsiana Sikorski et al., 2017. This subgenus is characterized by the anterior fusion of prostomium and peristomium, nuchal organ short and presence of notopodial hooded hooks in some specimens (absent in Laonice antarcticae). This subgenus was preliminarily recovered in a COI analysis by Bogantes et al. (2018), but it was decided not to use this subgenus yet, as this analysis included only a few taxa and two other subgenera erected by Sikorski et al., 2017 were not recovered.

Among Brazilian species, L. antarcticae is similar to L. branchiata and L. brevicornis in having a bell-shaped prostomium; prostomium fused to the peristomium; chaetae from the anterior region organized in two rows and a partial overlap on the starting chaetiger of interneuropodial pouches, neuropodial hooded hooks, sabre chaetae and branchial distribution. Laonice antarcticae can be separated from L. brevicornis based on the degree of fusion of prostomium and peristomium – only on the anterior margin in L. antarcticae and completely fused in L. brevicornis; caruncle length – to chaetiger 12 in L. antarcticae and to chaetiger 26 in L. brevicornis and branchiae length – about the same length of notopodial lamellae in L. antarcticae and twice as long as notopodial lamellae in L. brevicornis.

Laonice antarcticae is most similar to L. branchiata as both species possess a prostomium fused with peristomium only on the anterior margin and same caruncle length, although species can be distinguished based on the lack of dorsal crests and auricular lobes in L. antarcticae and branchiae length – about the same length of notopodial lamellae in L. antarcticae and twice as long as notopodial lamellae in L. branchiata.

Habitat: medium sand to fine silt, 49–410 m depth.

Distribution: Southeastern Brazil and Southern Brazil (Rio Grande do Sul to Espírito Santo states), Atlantic Ocean; Antarctic Ocean: Bellingshausen Sea.

Laonice branchiata Nonato et al., 1986

(Figs 16–18E)

Material examined. Amb1 Foz2, 19°47'21,11" S 39°55'7,58" W, 24m (1 ind); Amb2 Foz1, 19°52'14,57" S 39°59'41,06" W, 27m (20 ind); Amb2 Foz5, 19°32'28,16" S 39°41'35,15" W, 20m (3 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (1 ind); Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (3 ind); Amb14 B1, 20°34'34,37" S 40°20'50,77" W, 26m (5 ind); Amb14 D1, 19°35'35,96" S 39°41'20,61" W, 26m (2 ind).

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Comparative material examined. USNM 100451 (1 specimen, paratype).

Description: A large-sized Laonice species, largest complete specimen 15 mm long, 1.9 mm wide at the widest point for 115 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium. Body color whitish to light yellow in alcohol (Fig 16).

Prostomium bell-shaped, broadly rounded anteriorly, lacking frontal horns, extending posteriorly as a low caruncle to the end of chaetiger 12, flanked by U-shaped nuchal organs extending through entire caruncle length (Fig 17B). Two pairs of eyes in trapezoidal arrangement, anterior pair small and posterior pair large and recurved. Occipital antenna present on posterior half of the prostomium, around chaetiger 1 (Figs 16, 17A). Peristomium well-separated from prostomium for most of its length, fused only on the anterior margin; not fused to chaetiger 1 and bearing moderate lateral wings. Palps lost in all specimens.

Notopodial postchaetal lamellae foliaceous on branchial region and triangular on postbranchial region, greatly reduced on posterior region (Fig 17A–D). Notopodial prechaetal lamellae as a low flap on anterior region. Low dorsal crests present on postbranchial region of large specimens, from chaetigers 37–44 to chaetigers 46–59.

Neuropodial postchaetal lamellae auricular on branchial region and rounded on postbranchial region, reduced to a low flap on posterior region. Neuropodial prechaetal lamellae as a low flap on anterior chaetigers lacking interneuropodial pouches. Interneuropodial pouches starting from chaetigers 4–10 (depending on specimen size), present on whole body, except last 5–8 chaetigers (Fig 17D).

Chaetae from notopodia and neuropodia organized in two rows of granulated and narrowly unilimbate capillaries (Fig 18A). Chaetae from both rows of equal length, although notopodial chaetae are up to 1.4 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 18B).

Hooks in neuropodia only, starting from chaetigers 17–45, up to 12 per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks tridentate, with two secondary teeth arranged side by side above main tooth (Fig 18C). Secondary hood absent. Sabre chaetae starting from chaetigers 10–39, up to two per fascicle. Sabre chaetae non-limbate and intensely granulated (except at the base) (Fig 18D).

251

Branchiae from chaetiger 2, present on most of the anterior region, 10–45 pairs, short on first chaetigers and reaching full size around chaetigers 15–25 (Fig 17C), decreasing in size on last 10 chaetigers. Branchiae cirriform, moderately ciliated, completely free from and up to two times longer than notopodial postchaetal lamellae. Accessory branchiae present as auricular lobes connected basally to anterior part of branchiae, small on first chaetigers, reaching full size around chaetigers 5–10 (Figs 17A–B, 18E), much reduced on last branchiate chaetigers.

Pygidium rounded, bearing one pair of long ventral cirri (up to three times longer than dorsal cirri) and 3–4 pairs of short dorsal cirri.

Juvenile morphology: In juveniles, the caruncle is shorter, reaching up to chaetigers 8–11.

Methyl green staining pattern: Prostomium, peristomium and postchaetal lamellae from anterior region strongly stained; ventral side of anterior and middle regions weakly stained.

Remarks. Laonice branchiata was described by Nonato et al. (1986) based on material collected on the states of Paraná, São Paulo and Rio de Janeiro, southern and southeastern Brazil, from 15– 37 m depth.

Specimens collected in this work agree almost completely with the original description and with Radashevsky & Lana (2009) redescription based on type material and topotypes, except for a small variation on the starting chaetiger of neuropodial hooded hooks – chaetigers 17–45 in specimens examined in this work and chaetigers 14–46 in Radashevsky & Lana (2009); distribution of sabre chaetae – chaetigers 10–39 in specimens examined in this work and chaetigers 9–38 in Radashevsky & Lana (2009) and distribution of dorsal crests – from chaetigers 37–44 to chaetigers 46–59 in specimens examined in this work and from chaetigers 45–48 to chaetigers 56–60 in Radashevsky & Lana (2009), although such small discrepancies could be explained by specimen size and individual variability.

The main difference among descriptions is the fusion of prostomium and peristomium – prostomium fused with the peristomium on the anterior margin in this work, also noted by Sikorski (2011), and prostomium completely free from peristomium, according to Nonato et al. (1986) and Radashevsky & Lana (2009). The fusion is very subtle, visible only on the ventral side and was probably overlooked on previous descriptions.

Sikorski et al., 2017 erected four Laonice subgenera, placing L. branchiata in the subgenus Laonice Sikorski et al., 2017. This subgenus is characterized by the clear fusion of prostomium and peristomium on the anterior margin and capillaries in anterior region of the body arranged in two vertical rows. The validity of this subgenus could not be verified by Bogantes et al. (2018), since the authors only had access to only had COI sequences of a single species of this subgenus, thus being unable to comment on the genus validity. It was decided not to use this subgenus yet,

252 as the subgenus validity could not be tested and two other subgenera erected by Sikorski et al., 2017 were not recovered in a COI analysis.

The auricular lobes seems to act as accessory branchiae, as the structure is highly vascularized (Nonato et al. 1986). This character seems exclusive of L. branchiata, although Radashevsky & Lana (2009), while examining material from Espírito Santo state, southeastern Brazil, found a single specimen of an apparently unknown species close to L. branchiata that also bear auricular lobes but were unable to make further comments due to the poor condition of the material. Laonice pinnulata Radashevsky & Lana, 2009, described from the Pacific coast of Costa Rica, also bears accessory branchiae, but these branchiae are pectinate and composed of pinnules, instead of forming an auricular lobe as in L. branchiata.

Among Brazilian species, L. branchiata is most similar to L. antarcticae and L. brevicornis in having a bell-shaped prostomium; prostomium fused to the peristomium; chaetae from the anterior region organized in two rows and a partial overlap on the starting chaetiger of interneuropodial pouches, neuropodial hooded hooks, sabre chaetae and branchial distribution. Laonice branchiata can be separated from L. brevicornis based on the degree of fusion of prostomium and peristomium – only on the anterior margin in L. branchiata and completely fused in L. brevicornis; caruncle length – to chaetiger 12 in L. branchiata and to chaetiger 26 in L. brevicornis and lack of dorsal crests in L. brevicornis.

Laonice branchiata is most similar to L. antarcticae as both species possess a prostomium fused with peristomium only on the anterior margin and same caruncle length, although species can be distinguished based on the presence of dorsal crests and auricular lobes in L. branchiata and branchiae length – twice as long as notopodial lamellae in L. branchiata and about the same length of notopodial lamellae in L. antarcticae.

The specimens examined in this works corresponds to the northern-most record of this species, extending its distribution to the Espírito Santo state.

Habitat: fine sand to clay, 20–27 m depth.

Distribution: Southeastern Brazil and Southern Brazil (Santa Catarina to Espírito Santo states), Atlantic Ocean.

253

Laonice brevicornis (Kinberg, 1866)

(Figs 19–21E)

Synonym list:

Aricidea alata Treadwell, 1902 (subjective synonym)

Laonice aperata Radashevsky & Lana, 2009 (subjective synonym)

Laonice petersenae Radashevsky & Lana, 2009 (subjective synonym)

Mandane brevicornis Kinberg, 1866 (superseded original combination)

Material examined. Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (2 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (1 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (4 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (1 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (1 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (2 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (8 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (4 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (2 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (3 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (1 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (2 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (2 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (2 ind); Amb14 C1, 20°10'2,67" S 40°8'31,96" W, 26m (1 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (3 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (2 ind).

Comparative material examined. USNM 100447 (1 specimen, paratype of Laonice aperata).

Description: A large-sized Laonice species, largest complete specimen 43 mm long, 1.6 mm wide at the widest point for 103 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium. Body color whitish to light yellow in alcohol (Fig 19).

Prostomium bell-shaped, broadly rounded anteriorly, lacking frontal horns, extending posteriorly as a low caruncle up to the end of chaetiger 26, flanked by U-shaped nuchal organs extending through the entire caruncle length (Fig 20A). Two pairs of eyes in trapezoidal arrangement, anterior pair small and posterior pair large. Occipital antenna present on posterior half of the prostomium, near the posterior margin of chaetiger 1 (Figs 19, 20B). Peristomium fused with prostomium on the anterior margin (Fig 20B–C), not fused to chaetiger 1 and bearing moderate lateral wings. Palps lost in all specimens.

254

Notopodial postchaetal lamellae wide and foliaceous on branchial region and triangular on postbranchial region, greatly reduced on posterior region (Figs 19, 20A). Notopodial prechaetal lamellae as a low flap on anterior region. Dorsal crests absent.

Neuropodial postchaetal lamellae foliaceous on anterior region, gradually becoming elliptical on middle region and low and rounded on posterior region, greatly reduced on last chaetigers. Neuropodial prechaetal lamellae as a low flap on anterior chaetigers lacking interneuropodial pouches. Interneuropodial pouches starting from chaetigers 3–12, present for most of the body (absent on last 6–13 chaetigers) (Fig 20D) or interneuropodial pouches completely absent.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, non-limbate and non-granulated capillaries (Fig 21A), posterior row up to 2.5 times longer than anterior row; on neuropodia, widely unilimbate and lightly granulated capillaries (Fig 21B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 21C).

Hooks in neuropodia only, starting from chaetigers 12–36, up to 10 per fascicle, accompanied by 2–8 short non-limbate capillaries. Hooks tridentate, with two secondary teeth arranged side by side above main tooth (Fig 21D). Secondary hood absent. Sabre chaetae starting from chaetigers 13–30 up to two per fascicle. Sabre chaetae lightly granulated throughout (Fig 21E).

Branchiae from chaetiger 2, present on most of the anterior region, 13–34 pairs, short on first chaetigers and reaching full size around chaetigers 5–10, decreasing in size on last 10 chaetigers. Branchiae cirriform, moderately ciliated, completely free from and up to 2.5 times longer than notopodial postchaetal lamellae. Accessory branchiae absent.

Pygidium rounded, bearing one pair of short ventral cirri and 3–4 pairs of long (up to two times longer than ventral cirri) and thin dorsal cirri.

Juvenile morphology: In juveniles, the caruncle is shorter, reaching up to chaetigers 4–25 and branchiae may start on chaetiger 3.

Methyl green staining pattern: Anterior margin of prostomium strongly stained; margins of notopodial postchaetal lamellae weakly stained.

Remarks. Laonice brevicornis was originally described as Mandane brevicornis Kinberg, 1866 from Rio de Janeiro state, southeastern Brazil. The type material was reexamined by Söderström (1920) who noted that this species probably belonged to Laonice. Sikorski (2011) transferred the

255 species to Laonice and also synonymized Aricidea alata Treadwell, 1902, Laonice petersenae Radashevsky & Lana, 2009 and Laonice aperata Radashevsky & Lana, 2009 with Laonice brevicornis. According to Sikorski (2011), L. petersenae and L. aperata were both described from the type-locality of Laonice brevicornis and closely resembles the holotype of this species.

Sikorski (2011) examined material from the type locality of L. petersenae and L. aperata, noting that these species were mainly separated based on the lack of interneuropodial pouches in L. aperata, a non-reliable character, since it is variable and there are records of Prionospio cf. cirrifera populations in which some specimens presented interneuropodial pouches, while this structure was completely absent in other specimens. Furthermore, while in Radashevsky & Lana (2009) L. petersenae and L. aperata were separated by bathymetry, Sikorski (2011) examined samples where both species co-occurred, what was also observed in this work. The caruncle length seems to differ between species, to chaetiger 16 in L. aperata and to chaetiger 26 in L. petersenae; although in this study, some specimens identified as L. aperata presented a longer caruncle, up to chaetiger 19, while many L. petersenae had shorter caruncles, up to chaetigers 20– 21.

The methyl green staining pattern was also the same in forms with interneuropodial pouches (L. petersenae) and forms lacking interneuropodial pouches (L. aperata), further supporting this synonym. Ontogeny and genetic studies would be beneficial to support the synonym of both species with L. brevicornis.

Sikorski et al., 2017 erected four Laonice subgenera, placing L. brevicornis in the subgenus Laonice. This subgenus is characterized by the clear fusion of prostomium and peristomium on the anterior margin and capillaries in anterior region of the body arranged in two vertical rows. The validity of this subgenus could not be verified by Bogantes et al. (2018), since the authors only had access to only had COI sequences of a single species of this subgenus, thus being unable to comment on the genus validity. It was decided not to use this subgenus yet, as the subgenus validity could not be tested and two other subgenera erected by Sikorski et al., 2017 were not recovered in a COI analysis.

Among Brazilian species, Laonice brevicornis is most similar to L. antarcticae and L. branchiata in having a bell-shaped prostomium, prostomium fused with the peristomium, two rows of capillaries in anterior chaetigers and a partial overlap on the starting chaetiger of interneuropodial pouches, neuropodial hooded hooks, sabre chaetae and branchial distribution. However, species can be separated based on the degree of fusion between prostomium and peristomium – completely fused in L. brevicornis and fused only on the anterior margin in L. antarcticae and L. branchiata; caruncle length, to chaetiger 26 in L. brevicornis and to chaetiger 12 in L. antarcticae and L. branchiata and lack of interneuropodial pouches in some L. brevicornis specimens.

256

Habitat: coarse sand to medium silt, 25–157 m depth.

Distribution: Southeastern Brazil and Southern Brazil (Santa Catarina to Bahia states), Atlantic Ocean.

Laonice cf. weddellia Hartman, 1978

(Figs 22–24G)

Material examined. Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (38 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (2 ind); Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (2 ind); Amb1 Foz10, 19°35'12,39" S 39°38'33,16" W, 32m (1 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (10 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (1 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (4 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (9 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (14 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (11 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (4 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (1 ind); Amb1 Foz19, 19°46'14,99" S 39°34'50,94" W, 52m (4 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (2 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W, 30m (31 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (9 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (4 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (4 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (6 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (9 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (1 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (6 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (1 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (1 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (2 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (2 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (3 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (21 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (4 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (5 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (1 ind); Amb7 B2, 20°34'45,78" S 40°11'30,74" W, 41m (3 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (8 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (11 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (18 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (2 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (3 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (1 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (1 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (1 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (2 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (11 ind); Amb14 B1, 20°34'34,37" S 40°20'50,77" W, 26m (2 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (4 ind); Amb14

257

B3, 20°34'53,05" S 40°6'27,68" W, 49m (6 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (4 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (6 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (1 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (1 ind).

Comparative material examined. USNM 46700 (holotype), USNM 46701 (paratypes, 20 specimens).

Description: A medium-sized Laonice species, largest complete specimen 20 mm long, 1.5 mm wide at the widest point for 98 chaetigers. Body slightly dorsoventrally flattened on anterior and middle regions and cylindrical on posterior region, tapering towards the pygidium. Red pigmented spots on ventral side of chaetiger 8. Body color light yellow in alcohol (Fig 22A).

Prostomium wide, broadly rounded anteriorly, lacking frontal horns, extending posteriorly as a low caruncle to the end of chaetigers 12–20, flanked by U-shaped nuchal organs extending through entire caruncle length (Fig 23A–B) . Two pairs of eyes in trapezoidal arrangement, anterior pair small and posterior pair large. Occipital antenna present on posterior half of the prostomium, near the posterior margin of chaetiger 1 (Fig 22). Peristomium free from prostomium, not fused to chaetiger 1 and bearing moderate lateral wings. Grooved palps reaching up to chaetiger 10, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamella cirriform to lanceolate on notopodium and square-shaped on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae wide and foliaceous throughout, greatly reduced on posterior region (Figs 22A, 23A, C). Notopodial prechaetal absent throughout. Dorsal crests low from chaetiger 30 to chaetiger 35, not observed in small specimens.

Neuropodial postchaetal lamellae foliaceous throughout, greatly reduced on posterior region. Neuropodial prechaetal absent throughout. Interneuropodial pouches starting from chaetigers 3– 6, present throughout entire specimen (except last 10–15 chaetigers) (Fig 23C–D).

Chaetae from notopodia and neuropodia organized in five rows from chaetiger 2 to chaetiger 15, two anterior rows of narrowly unilimbate (Fig 24A) and lightly granulated capillaries, followed by three rows of widely unilimbate and lightly granulated capillaries (Fig 24B). From chaetiger 16 onwards, only two rows of capillaries, anterior row narrowly unilimbate and lightly granulated (Fig 24C) and posterior row of widely unilimbate and lightly granulated capillaries (Fig 24D). Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region,

258 chaetae of both rami progressively become elongate, non-limbate, thinner, and less numerous (Fig 24E).

Hooks in neuropodia only, starting from chaetigers 15–22, up to eight per fascicle, accompanied by 1–6 short non-limbate capillaries. Hooks tridentate, with two small secondary teeth arranged side by side above main tooth (Fig 24F). Secondary hood absent. Sabre chaetae starting from chaetigers 9–13. Sabre chaetae granulated throughout (Fig 24G).

Branchiae from chaetiger 2, present on most of the anterior region, 15–52 pairs short on first chaetigers and reaching full size around chaetigers 10–15, decreasing in size on last 10 chaetigers (Fig 23A, C). Branchiae cirriform, moderately ciliated, completely free from and up to 2.5 times longer than notopodial postchaetal lamellae. Accessory branchiae absent.

Pygidium rounded, bearing up to eight cirri of equal length.

Sperm plates from chaetiger 17.

Juvenile morphology: In juveniles, the caruncle is shorter, reaching up to chaetigers 7–11 and in early juveniles, red pigmented spots may be present on the ventral side of chaetigers 8 and 9.

Methyl green staining pattern: No pattern observed, whole specimen weakly stained.

Remarks. Laonice weddellia was described by Hartman (1978) from material collected in the Weddell Sea, Antarctica, from 400–659 m depth. Blake (1983) redescribed the species and expanded its distribution to Chile, Strait of Magellan, Falkland Islands, South Orkney Islands and South Shetland Islands, from 44 to 3111 m depth. Radashevsky & Lana (2009) recorded the species in southern and southeastern Brazil, from Rio Grande do Sul to Rio de Janeiro states, while at the same time being cautious about Brazilian records, which did not entirely fit the species’ original description, while Pardo & Peixoto (in press) extended the species distribution to Espírito Santo state. The species was also recorded in southwestern Australia, Indian Ocean, by Greaves et al. (2011), who also noted discrepancies from the original description.

Specimens in this work are similar to Hartman (1978) original description and Blake (1983) redescription in having a bell-shaped and anteriorly rounded prostomium; prostomium not fused with peristomium; chaetae from anterior chaetigers organized in up to five rows; pygidium bearing up to eight cirri; up to eight neuropodial hooded hooks per fascicle; similar maximum number of branchiae – up to 49 pairs (Hartman, 1978), up to 50 pairs (Blake, 1983) and up to 52 pairs (this work); overlap in the starting chaetiger of hooded hooks – chaetigers 20–21 in the descriptions and chaetigers 15–22 in this work; overlap in the starting chaetiger of sabre chaetae – chaetiger 12 in the descriptions and chaetigers 9–13 in this work and similar caruncle length – to chaetiger 12 (Hartman, 1978) and to chaetigers 12–20 (Blake, 1983) and this work.

259

However, significant differences were found, such as the presence of red pigmented spots on ventral side of chaetiger 8 (sometimes also present in chaetiger 9), lacking in Hartman (1978) and Blake (1983) descriptions; starting chaetiger of interneuropodial pouches, from chaetigers 8–9 in Hartman (1978), chaetigers 8–10 in Blake (1983) and chaetigers 3–6 in this work and presence of dorsal crests on chaetigers 30–35 in specimens examined in this work, absent in Hartman (1978), who only mentioned the occurrence of ciliated ridges and intersegmental grooves in posterior chaetigers, while Blake (1983) does not report the presence of these structures. Due to these differences, it was decided to be cautious, justifying the “cf” adoption.

Sikorski et al., 2017 erected four Laonice subgenera, placing L. weddellia in the subgenus Appelloefia Sikorski et al., 2017. This subgenus is characterized by the prostomium totally free from peristomium, capillaries arranged in more than two vertical rows in several anterior chaetigers, genital pouches generally present on very limited number of segments (or absent), hooded hooks with more than two teeth in side view, and frequent widening of the body on several anterior segments. Our specimens do not fit quite well in this group, as interneuropodial pouches are present on most of the body, the body is not widened and hooded hooks look bidentate in lateral view. Greaves et al. (2011) noted that some specimens of L. weddellia had quadridentate hooded hooks, with a superior and very small unpaired teeth above the secondary teeth. It was decided not to use this subgenus, since it was not recovered in a COI analysis by Bogantes et al. (2018), as some taxa that were placed in the subgenus Appelloefia appeared in a different region of the tree, separated by multiple nodes.

Among Brazilian species, L. weddellia is most similar to Laonice kinbergi sp. nov., L. profunda sp. nov. and L. brevicarunculata sp. nov., as all species possess a prostomium completely separated from peristomium. Laonice weddellia can be readily distinguished from L. profunda sp. nov. and L. brevicarunculata sp. nov. by the presence of an occipital tentacle, presence of dorsal crests and in having up to five rows of chaetae in anterior chaetigers (Table 2). The species is most similar to L. kinbergi sp. nov. in having an occipital tentacle; partial overlap of caruncle length, number of branchial pairs, starting chaetiger of interneuropodial pouches, neuropodial hooded hooks and sabre chaetae, although species can be distinguished by distribution of dorsal crests – from chaetiger 20 to chaetiger 27 in L. kinbergi sp. nov. and from chaetiger 30 to chaetiger 35 in L. weddellia, as well as presence of up to five rows of chaetae in anterior chaetigers in L. weddellia.

Habitat: medium sand to fine silt, 25–450 m depth.

Distribution: Atlantic Ocean: Brazil (Rio Grande do Sul to Espírito Santo states); Antarctic Ocean: Weddell Sea (type-locality), Strait of Magellan, Falkland Islands, South Orkney Islands and South Shetland Islands; Indian Ocean: Australia.

260

Laonice kinbergi sp. nov.

(Figs 25–27G)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 06 Jun 2013 to 17 Jul 2013, 970m, MNRJP XXXX. Paratypes: Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 06 Jun 2013 to 17 Jul 2013, 970m, ZUEC-POL XXXX (2 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 02 Dec 2011 to 02 Feb 2012, 1333m, MNRJP XXXX (1 ind), MZUSP XXXX (1 ind).

Additional material examined. Amb4 F7, 20°4'8,18" S 38°31'27,32" W, 1302m (1 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (1 ind); Amb5 C5, 20°14'19,45" S 39°48'36,67" W, 381m (1 ind); Amb5 D6, 19°50'1,87" S 39°26'30,04" W, 1055m (2 ind); Amb11 C9, 20°48'37,26" S 38°45'28,85" W, 2465m (1 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (1 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (1 ind); Amb12 E7, 19°47'2,44" S 39°3'14,62" W, 1250m (1 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (1 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (1 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (1 ind).

Description: A medium-sized Laonice species, largest specimen 13 mm long, 1.3 mm wide at the widest point for 57 chaetigers (incomplete), holotype 10 mm long, 1 mm wide at the widest point for 35 chaetigers (incomplete). Body slightly dorsoventrally through whole fragment. Body color whitish to light yellow in alcohol (Fig 25).

Prostomium bell-shaped, broadly rounded anteriorly, with rounded lateral margins, lacking frontal horns, extending posteriorly as a low caruncle to the end of chaetiger 14, flanked by U- shaped nuchal organs extending through entire caruncle length. Small pair of subdermal eyes present or eyes absent. Digitiform occipital antenna with a rounded tip present on posterior half of the prostomium, near the posterior margin of chaetiger 1. Peristomium well-separated from prostomium; not fused to chaetiger 1 and bearing very low lateral wings (Fig 26A–B). Palps lost in all specimens.

Chaetiger 1 bearing numerous short chaetae. Postchaetal lamella foliaceous on notopodium and auricular on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae short and foliaceous, bearing a rounded tip on chaetigers 2–7, auricular, with greatly expanded dorsal part on the remaining anterior and middle chaetigers, gradually becoming foliaceous, with a widened based on last chaetigers of the fragment (Figs 25,

261

26A). Notopodial prechaetal lamellae triangular, well-developed through entire fragment. Notopodial lamellae unknown on posterior region. Dorsal crests low from chaetiger 20 to chaetiger 27 (Fig 26C).

Neuropodial postchaetal lamellae auricular on chaetigers 2–8, bearing an elongated inferior tip from chaetiger 9 onwards; tip fused to the body, well-developed on anterior region and shorter on last chaetigers of the fragment. Near the end of the fragment, lamellae develops a laterally pointed tip. Neuropodial prechaetal lamellae auricular on anterior region and reduced to a low flap from middle region onwards. Neuropodial lamellae unknown on posterior region. Interneuropodial pouches starting from chaetigers 5–14 (usually on chaetiger 5), present until end of fragment.

Chaetae from notopodia and neuropodia organized in two rows of unilimbate capillaries, posterior row up to 1.5 times longer than anterior row. In notopodia, anterior row narrowly limbate and intensely granulated (Fig 27A) and posterior row narrowly limbate, bearing numerous transversal bars (Fig 27B). In neuropodia, anterior row widely unilimbate and intensely granulated (Fig 27C), posterior row narrowly limbate and lightly granulated (Fig 27D). Towards the posterior region, chaetae of both rami progressively become elongate, non-limbate, thinner, and less numerous (Fig 27E).

Hooks in neuropodia only, starting from chaetigers 18–35, up to six per fascicle, accompanied by 6–9 non-limbate capillaries. Hooks very long and tridentate, with two secondary teeth arranged side by side above main tooth (Fig 27F). Secondary hood absent. Sabre chaetae starting from chaetigers 10–29, up to two per fascicle. Sabre chaetae non-limbate and intensely granulate throughout (Fig 27G).

Branchiae from chaetiger 2, present on most of the anterior region, 17–36 pairs, shorter than notopodial postchaetal lamellae on chaetigers 2 and 3 and slightly longer than notopodial postchaetal lamellae on remaining chaetigers, reaching full size around chaetiger 8, decreasing in size on last 5 chaetigers. Branchiae cirriform and delicate, sparsely ciliated and completely free from notopodial postchaetal lamellae. Accessory branchiae absent.

Pygidium of unknown morphology.

Methyl green staining pattern: anterior half of the prostomium strongly stained; entire fragment weakly stained, except for caruncle, interneuropodial pouches and branchiae.

Remarks. Laonice kinbergi sp. nov. is distinguished from all Brazilian species in having a bell- shaped prostomium that is not fused with the peristomium. In L. antarcticae, L. branchiata and L. brevicornis the prostomium is also bell-shaped, but the prostomium and peristomium are fused (at least in the anterior margin) (Sikorski 2011); while in Laonice weddellia L. profunda sp. nov.

262 and L. brevicarunculata sp. nov., the prostomium is widened, but it is completely free from the peristomium.

This seems to be the most obvious character to distinguish this species, as most of Laonice kinbergi sp. nov. morphological characters overlap among species (Table 2), such as caruncle length (except for L. brevicornis and L. brevicarunculata sp. nov.); starting chaetiger of interneuropodial pouches (except for L. profunda sp. nov.); two rows of capillaries in anterior chaetigers (except for L. weddellia); staring chaetiger of neuropodial hooded hooks (except for L. brevicarunculata sp. nov.); starting chaetiger of sabre chaetae (except for L. profunda sp. nov.) and branchiae distribution (except for except for L. profunda sp. nov.). Also, digitiform antennae are present on most species (except for L. profunda sp. nov. and L. brevicarunculata sp. nov.) (Radashevsky & Lana 2009; Sikorski 2011).

Branchiae morphology is another unique character in L. kinbergi sp. nov., as branchiae are delicate, cirriform and only slightly longer than notopodial postchaetal lamellae. However, in some deep-sea specimens, branchiae may be entirely missing, further complicating identification.

Laonice kinbergi sp. nov. is most similar to L. antarcticae in having a bell-shaped and anteriorly rounded prostomium, digitiform antennae, two rows of anterior capillaries, similar starting chaetiger of neuropodial hooded hooks – from chaetigers 18–35 in L. kinbergi sp. nov. and from chaetigers 21–38 in L. antarcticae; similar branchial distribution – from 17 to 36 pairs in L. kinbergi sp. nov. and from 14 to 35 pairs in L. antarcticae; lack of auricular lobes and slight overlap on the caruncle length, starting chaetiger of interneuropodial pouches, starting chaetiger of sabre chaetae. However, species can be distinguished based on the lateral margins in the prostomium – rounded in L. kinbergi sp. nov. and pointed in L. antarcticae; degree of fusion between prostomium and peristomium – unfused in L. kinbergi sp. nov. and anteriorly fused in L. antarcticae and branchiae morphology – thin and slightly longer than notopodial postchaetal lamellae in L. kinbergi sp. nov. and slightly shorter than notopodial postchaetal lamellae in L. antarcticae, and presence of dorsal crests in L. kinbergi sp. nov. (Hartman 1953; Radashevsky & Lana 2009).

Etymology: The specific epithet, kinbergi, is a tribute to Johan G. H. Kinberg (1820–1908), who described the first Laonice species, Laonice brevicornis Kinberg, 1866 (described as Mandane brevicornis).

Habitat: medium silt to clay, 381–2770 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

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Laonice profunda sp. nov.

(Figs 28–30E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 02 Dec 2011 to 02 Feb 2012,1889m, MNRJP XXXX. Paratypes: Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 06 Jun 2013 to 17 Jul 2013, 2770m, MNRJP XXXX (2 ind), MZUSP XXXX (2 ind), ZUEC-POL XXXX (2 ind).

Additional material examined. Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 2426m (2 ind); Amb3 G10, 19°3'55,8" S 37°45'8,27" W, 2993m (1 ind); Amb4 B10, 21°4'40,47" S 38°26'13,68" W, 3007m (2 ind); Amb4 C10, 20°59'0,28" S 38°28'36,56" W, 2997m (1 ind); Amb4 E9, 20°35'50,48" S 38°27'7,64" W, 2494m (1 ind); Amb4 F9, 20°29'3,31" S 38°23'15,5" W, 2504m (4 ind); Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 1889m (4 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (2 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (1 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (8 ind); Amb5 E8, 20°15'59,97" S 38°40'53,86" W, 1887m (1 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (4 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (5 ind); Amb8 G8, 19°3'45,82" S 37°47'28,26" W, 1928m (13 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (1 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (6 ind); Amb11 A9, 21°9'39,38" S 38°52'7,25" W, 2502m (3 ind); Amb11 B8, 20°41'33,93" S 39°35'22,06" W, 1908m (6 ind); Amb11 B9, 20°54'43,79" S 38°56'10,85" W, 2520m (1 ind); Amb11 B10, 21°4'33,4" S 38°26'19,34" W, 3001m (1 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (1 ind); Amb11 C8, 20°25'13,22" S 39°27'19,49" W, 1920m (6 ind); Amb11 C9, 20°48'37,26" S 38°45'28,85" W, 2465m (3 ind); Amb11 C10, 20°57'57,99" S 38°27'52,41" W, 2996m (2 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (6 ind); Amb11 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (1 ind); Amb11 D10, 20°53'28,94" S 38°21'21,75" W, 3000m (3 ind); Amb11 E8, 20°15'55,63" S 38°40'45,57" W, 1892m (3 ind); Amb11 E9, 20°35'51,99" S 38°27'13,04" W, 2501m (6 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (4 ind); Amb11 F8, 20°16'38,17" S 38°27'26,52" W, 1904m (7 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (10 ind); Amb11 F10, 20°46'17,79" S 38°17'16,01" W, 3002m (4 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (1 ind); Amb12 G7, 19°3'30,62" S 37°48'46,66" W, 1347m (1 ind); Amb12 G8, 19°3'39,78" S 37°47'39,35" W, 1867m (6 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (2 ind); Amb12 G10, 19°3'10,22" S 37°45'28,45" W, 2950m (5 ind).

Description: A small-sized Laonice species, largest specimen 12 mm long, 0.3 mm wide at the widest point for 29 chaetigers (incomplete), holotype being the largest specimen. Body slightly

264 dorsoventrally flattened throughout, tapering towards end of fragment. Body color light yellow in alcohol (Fig 28).

Prostomium wide, anteriorly rounded, lacking frontal horns, widest in middle region, narrowed posteriorly and extending as a low caruncle up to chaetiger 13, flanked by U-shaped nuchal organs extending through entire caruncle length (Figs 28, 29A, B). Eyes absent. Occipital antenna absent. Presence of a prominent elevation on posterior half of the prostomium, near the posterior margin of chaetiger 1. Peristomium free from prostomium not fused to chaetiger 1 and bearing low lateral wings. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae triangular on both rami, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous from chaetiger 2 to chaetigers 4–7 (depending on specimen size), triangular on following chaetigers and greatly reduced on postbranchial region (Figs 28, 29A, C). Notopodial prechaetal lamellae present as a low flap on branchiate chaetigers. Notopodial lamellae unknown on posterior region. Dorsal crests absent.

Neuropodial postchaetal lamellae triangular on branchial region and greatly reduced afterwards, although remaining triangular until end of fragment (Fig 29D). Neuropodial prechaetal lamellae absent. Neuropodial lamellae unknown on posterior region. Interneuropodial pouches starting from chaetiger 4, present until end of fragment.

Chaetae from notopodia (Fig 30A) and neuropodia (Fig 30B) organized in two rows of non- limbate and intensely granulated capillaries. Chaetae from both rows of same length in notopodia and posterior row twice as long as anterior row in neuropodia. Towards end of fragment, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 30C).

Hooks in neuropodia only, starting from chaetigers 15–19, up to five per fascicle, accompanied by 1–4 short non-limbate capillaries. Hooks tridentate, with two very small secondary teeth arranged side by side above main tooth (Fig 30D). Secondary hood absent. Sabre chaetae starting from chaetigers 6–10, up to two per fascicle. Sabre chaetae narrowly unilimbate and intensely granulated throughout (Fig 30E).

Branchiae from chaetiger 2, present on most of the anterior region, 6–14 pairs, shorter than notopodial postchaetal lamellae on first 3–6 chaetigers and about the same length of notopodial postchaetal lamellae on following chaetigers (Fig 29C), abruptly reducing in size on last branchial pair. Branchiae cirriform, moderately ciliated and completely free from notopodial postchaetal lamellae. Accessory branchiae absent.

265

Pygidium of unknown morphology.

Methyl green staining pattern: No specific pattern observed, whole specimen weakly stained.

Remarks. Laonice profunda sp. nov. is unique among Brazilian Laonice species in having an anteriorly rounded prostomium, widest at midregion and completely free from peristomium, lack of occipital antenna and interneuropodial pouches starting from chaetiger 4.

As in many Laonice species, most numerical characters overlap and are not helpful in distinguishing species, such as the starting chaetiger of interneuropodial pouches (except for L. kinbergi sp. nov. and L. brevicarunculata sp. nov.), starting chaetiger of neuropodial hooded hooks (except for L. antarcticae and L. brevicarunculata sp. nov.) and the starting chaetiger of sabre chaetae (except for L. antarcticae, L. brevicornis and L. brevicarunculata sp. nov.) (Table 2). Unfortunately, all specimens were anterior fragments, thus pygidium morphology remains unknown.

Laonice profunda sp. nov. differs from Laonice antarcticae, L. branchiata, L. brevicornis and L. kinbergi sp. nov. in having a wide and anteriorly rounded prostomium, instead of a bell-shaped prostomium (Radashevsky & Lana 2009). The species further differs from Laonice antarcticae, L. branchiata and L. brevicornis in having a prostomium completely free from the peristomium (Sikorski 2011).

Laonice profunda sp. nov. is most similar to L. brevicarunculata sp. nov. in having a wide and anteriorly rounded prostomium, prostomium completely free from peristomium and lack of occipital antennae, although species can be distinguished based on caruncle length – to the end of chaetiger 13 in L. profunda sp. nov. and to the end of chaetiger 4 in L. brevicarunculata sp. nov.; starting chaetiger of interneuropodial pouches – starting from chaetiger 4 in L. profunda sp. nov. and starting from chaetiger 6 in L. brevicarunculata sp. nov.; starting chaetiger of neuropodial hooded hooks – from chaetigers 15–19 in L. profunda sp. nov. and from chaetiger 36 in L. brevicarunculata sp. nov.; starting chaetiger of sabre chaetae – from chaetigers 6–10 in L. profunda sp. nov. and starting from chaetiger 20 in L. brevicarunculata sp. nov.; branchiae distribution – from 6 to 14 pairs in L. profunda sp. nov. and 34 pairs in L. brevicarunculata sp. nov. and branchiae length – about the same size as notopodial postchaetal lamellae in L. profunda sp. nov. and up to 1.5 times longer than notopodial postchaetal lamellae in L. brevicarunculata sp. nov.

Etymology: The specific epithet, profunda, refers to the great depths in which this species was collected.

Habitat: medium silt to clay, 1000–3007 m depth.

266

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Laonice brevicarunculata sp. nov.

(Figs 31–32E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 02 Dec 2011 to 02 Feb 2012, MNRJP XXXX. Paratype: Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 06 Jun 2013 to 17 Jul 2013, 1040m, MNRJP XXXX (1 ind).

Additional material examined. Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (1 ind);

Description: A large-sized Laonice species, largest specimen 8 mm long, 1 mm wide at the widest point for 78 chaetigers (incomplete), holotype being the largest specimen. Body slightly dorsoventrally flattened throughout, tapering towards end of fragment. Body color whitish in alcohol (Fig 31).

Prostomium wide, broadly rounded anteriorly, lacking frontal horns, extending posteriorly as a low caruncle up to the end of chaetiger 4, flanked by U-shaped nuchal organs extending through entire caruncle length. Eyes absent. Occipital antenna absent (Fig 31). Peristomium free from prostomium and partially fused to chaetiger 1. Lateral wings absent. Palps lost.

Notopodial postchaetal lamellae wide and foliaceous on branchial region and triangular on postbranchial region (Fig 31). Notopodial prechaetal lamellae present as a low flap from chaetiger 2 to chaetiger 15. Notopodial lamellae unknown on posterior region. Dorsal crests absent.

Neuropodial postchaetal lamellae triangular with ventrally pointed tip on branchial region and auricular on postbranchial region. Neuropodial prechaetal lamellae absent. Neuropodial lamellae unknown on posterior region. Interneuropodial pouches starting from chaetiger 6, present until end of fragment.

Chaetae from notopodia (Fig 32A) and neuropodia (Fig 32B) organized in two rows of narrowly unilimbate and lightly granulated capillaries. Posterior row up to 1.5 times longer than anterior row. Towards end of fragment, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 32C).

267

Hooks in neuropodia only, starting from chaetiger 36, up to nine per fascicle, accompanied by 3– 7 short non-limbate capillaries. Hooks tridentate, with two secondary teeth arranged side by side above main tooth (Fig 32D). Secondary hood absent. Sabre chaetae starting from chaetiger 20, up to two per fascicle. Sabre chaetae non-limbate and intensely granulated throughout (Fig 32E).

Branchiae from chaetiger 2, present on most of the anterior region, 34 pairs, short on first chaetigers and reaching full size on chaetiger 5, gradually decreasing in size on last 15 chaetigers. Branchiae cirriform, moderately ciliated, completely free from and up to 1.5 times longer than notopodial postchaetal lamellae. Accessory branchiae absent.

Pygidium of unknown morphology.

Methyl green staining pattern: Prostomium, dorsal side of peristomium and margins of postchaetal lamellae from branchiate chaetigers strongly stained, ventral side of branchiate chaetigers weakly stained.

Remarks. Laonice brevicarunculata sp. nov. can be separated from the remaining Brazilian Laonice species in having a wide an anteriorly rounded prostomium, prostomium completely free from peristomium, lack of occipital antennae and a very short caruncle, reaching to the end of chaetiger 4.

The caruncle length not only readily distinguishes L. brevicarunculata sp. nov. from other Brazilian Laonice species, being also unusual among Laonice species (Sikorski 2011), present only in L. asaccata Sigvaldadóttir & Desbruyères, 2003, from the Mid-Atlantic vent in Azores (Portugal) triple junction and in L. papillibranchiae Ward, 1981, from Hawaii. Laonice brevicarunculata sp. nov. can be separated from both species in lacking an occipital antenna (short in L. asaccata and long in L. papillibranchiae); lacking dorsal crests (present from chaetiger 17 in L. papillibranchiae and lacking in L. asaccata); starting chaetiger of interneuropodial pouches – from chaetiger 6 in L. brevicarunculata sp. nov., from chaetiger 12 in L. papillibranchiae and completely absent in L. asaccata; starting chaetiger of neuropodial hooded hooks – chaetiger 36 in L. brevicarunculata sp. nov., from chaetigers 16–19 in L. papillibranchiae and from chaetigers 30–32 in L. asaccata; starting chaetiger of sabre chaetae – from chaetiger 20 in L. brevicarunculata sp. nov., from chaetigers 20–23 in L. asaccata and from chaetigers 9–12 in L. papillibranchiae and number of branchial pairs – 34 pairs in L. brevicarunculata sp. nov., 30 pairs in L. asaccata and present on two thirds of body length in L. papillibranchiae (Sigvaldadóttir & Desbruyères 2003; Ward 1981). Laonice brevicarunculata sp. nov. further differs from L. papillibranchiae in lacking papillae on inner side of branchiae (Ward 1981).

268

As pointed out in previous remarks, numerical characters overlap in Brazilian species, not being informative to distinguish species. A character overlap is observed in the starting chaetiger of interneuropodial pouches (except for L. antarcticae and L. profunda sp. nov.), starting chaetiger of neuropodial hooded hooks (except for L. weddellia, L. kinbergi sp. nov. and L. profunda sp. nov.), starting chaetiger of sabre chaetae (except for L. weddellia and L. profunda sp. nov.) and branchiae distribution (except for L. profunda sp. nov.) (Radashevsky & Lana 2009; Sikorski 2011). (Table 2).

Among Brazilian species, L. brevicarunculata sp. nov. is most similar to L. profunda sp. nov. in having a wide prostomium, prostomium and peristomium completely separated and lack of occipital antennae, but species can be distinguished based on caruncle length – to the end of chaetiger 4 in L. brevicarunculata sp. nov. and to the end of chaetiger 13 in L. profunda sp. nov., starting chaetiger of interneuropodial pouches – from chaetiger 6 in L. brevicarunculata sp. nov. and from chaetiger 4 in L. profunda sp. nov.; starting chaetiger of neuropodial hooded hooks – from chaetiger 36 in L. brevicarunculata sp. nov. and from chaetiger 15–19 in L. profunda sp. nov.; starting chaetiger of sabre chaetae – from chaetiger 20 in L. brevicarunculata sp. nov. and from chaetigers 6–10 in L. profunda sp. nov.; branchial distribution – 34 pairs in L. brevicarunculata sp. nov. and 6–14 pairs in L. profunda sp. nov. and branchiae length – up to 1.5 times longer than notopodial postchaetal lamellae in L. brevicarunculata sp. nov. and about the same length of notopodial postchaetal lamellae in L. profunda sp. nov. Unfortunately, all specimens were anterior fragments, thus the pygidium morphology remains unknown.

Etymology: The specific epithet, brevicarunculata (from “brevis”, Latin adjective for short, plus “caruncula” and the suffix “-atus”, meaning bearing), refers to the short caruncle present in this species.

Habitat: medium silt to fine silt, 1035–1295 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Identification key to Laonice species recorded in this work.

1. Up to five rows of capillaries on anterior chaetigers……………………………L. weddellia - Two rows of capillaries on anterior chaetigers………………………………………………2 2(1). Caruncle extending after chaetiger 20……………………………………...L. brevicornis - Caruncle extending to chaetiger 20 at most (usually shorter)………………………………3 3(2). Dorsal crests present……………………………………………………………………..4 - Dorsal crests absent…………………………………………………………………………..5

269

4(3). Prostomium fused with peristomium on the anterior margin, auricular lobes present…………………………………………………………………………….L. branchiata - Prostomium completely free from peristomium, auricular lobes absent…L. kinbergi sp. nov. 5(3). Caruncle extending to chaetiger 4…………………………..L. brevicarunculata sp. nov. - Caruncle extending after chaetiger 4………………………………………………………...6 6(5). Prostomium bell-shaped, anteriorly fused with peristomium……………...L. antarcticae - Prostomium wide, anteriorly rounded, completely free from peristomium……………………………………………………………….L. profunda sp. nov.

Acknowledgments

References

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Legends of illustrations Figure 1. Map showing sampled area (blue polygon).

Figure 2. Aonides sarsi sp. nov., anterior chaetigers and mid-body, lateral view (MZUSP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 3. SEM of Aonides sarsi sp. nov. A. Anterior and middle regions, lateral view. B. Anterior region, dorsal view. C. Close-up of anterior region, dorsal view. D. Pygidium, ventrolateral view. Abbreviations: an, antenna; br, branchia; nela, neuropodial lamellae; nolo, notopodial lamellae; pe, peristomium; pr, prostomium; pyg, pygidium.

Figure 4. Aonides sarsi sp. nov. A. Anterior region. B. Anterior chaetiger. C. Notopodial chaetae from anterior region. D. Neuropodial chaetae from anterior region. E. Capillary from posterior region. F. Hooded hook. G. Pygidium.

Figure 5. Aonides curvus sp. nov., whole specimen, lateral view (MNRJP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: pr, prostomium.

Figure 6. SEM of Aonides curvus sp. nov. A. Anterior region, dorsal view. B. Anterior and middle regions, dorsal view. C. Hooded hook, lateral view. Abbreviations: br, branchia; hh, hooded hook; nolo, notopodial lamellae; pe, peristomium; pr, prostomium.

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Figure 7. Aonides curvus sp. nov. A. Anterior region. B. Anterior chaetiger. C. Notopodial and neuropodial chaetae from anterior region. D. Capillary from posterior region. E. Hooded hook. F. Pygidium.

Figure 8. Aonidella cf. cirrobranchiata, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 9. Aonidella cf. cirrobranchiata. A. Anterior region. B. Anterior chaetiger. C. Noto- and neurochaetae from anterior region, anterior row. D. Noto- and neurochaetae from anterior region, posterior row. E. Capillary from posterior region. F. Hooded hook.

Figure 10. Aonidella cf. dayi, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 11. SEM of Aonidella cf. dayi. A. Anterior region, dorsal view. B. Close-up of anterior region, dorsal view. C. Middle and posterior regions, dorsolateral view. D. Posterior region, ventral view. Abbreviations: br, branchia; hh, hooded hook; nela, neuropodial lamellae; nolo, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 12. Aonidella cf. dayi. A. Anterior region. B. Anterior chaetiger. C. Noto- and neurochaetae from anterior region, anterior row. D. Noto- and neurochaetae from anterior region, posterior row. E. Capillary from posterior region. F. Hooded hook. G. Pygidium.

Figure 13. Laonice cf. antarcticae, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; ip, interneuropodial pouch; pe, peristomium; pr, prostomium.

Figure 14. SEM of Laonice cf. antarcticae. A. Anterior and middle regions, dorsal view. B. Anterior region, dorsal view. C. Close-up of anterior region showing end of caruncle, dorsal view. D. Middle region, lateral view. Abbreviaions: an, antenna; br, branchia; car, caruncle; ip, interneuropodial pouch; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 15. Laonice cf. antarcticae. A. Noto- and neurochaetae from anterior region. B. Capillary from posterior region. C. Hooded hook. D. Sabre chaetae.

Figure 16. Laonice branchiata, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; ip, interneuropodial pouch; pe, peristomium; pr, prostomium.

Figure 17. SEM of Laonice branchiata. A. Anterior region, dorsal view. B. Close-up of anterior region showing end of caruncle, dorsal view. C. Anterior region, lateral view. D. Middle region, lateral view. Abbreviations: al, auricular lobe; an, antenna; br, branchia; car, caruncle; ip,

276 interneuropodial pouch; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 18. Laonice branchiata. A. Noto- and neurochaetae from anterior region. B. Capillary from posterior region. C. Hooded hook. D. Sabre chaetae. E. Anterior chaetiger.

Figure 19. Laonice brevicornis, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; pe, peristomium; pr, prostomium.

Figure 20. SEM of Laonice brevicornis. A. Anterior and middle regions, dorsal view. B. Anterior region, dorsal view. C. Anterior region, ventral view. D. Anterior region, lateral view. Abbreviations: an, antenna; br, branchia; car, caruncle; ip, interneuropodial pouch; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium; prob, proboscis.

Figure 21. Laonice brevicornis. A. Notopodial chaetae from anterior region. B. Neuropodial chaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 22. Laonice cf. weddellia, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; pe, peristomium; pr, prostomium.

Figure 23. SEM of Laonice cf. weddellia. A. Anterior and middle regions, dorsal view. B. Close- up of anterior region, dorsal view. C. Anterior and middle regions, lateral view. Middle region, lateral view. Abbreviations: br, branchia; car, caruncle; ip, interneuropodial pouch; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 24. Laonice cf. weddellia. A. Noto- and neuro chaetae from rows 1 and 2, anterior region. B. Noto- and neuro chaetae from rows 3 to 5, anterior region. C. Noto- and neuro chaetae fro9m chaetiger 16 onwards, anterior row. D. Noto- and neuro chaetae fro9m chaetiger 16 onwards, posterior row. E. Capillary from posterior region. F. Hooded hook. G. Sabre chaetae.

Figure 25. Laonice kinbergi sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; dc, dorsal crest; ip, interneuropodial pouch; pe, peristomium; pr, prostomium.

Figure 26. SEM of Laonice kinbergi sp. nov. A. Anterior and middle regions, dorsal view. B. Close-up of anterior region, dorsal view. C. Middle region, dorsal view. Abbreviations: br, branchia; car, caruncle; dc, dorsal crest; nola, notopodial lamellae; pr, prostomium.

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Figure 27. Laonice kinbergi sp. nov. A. Notopodial chaetae from anterior region, anterior row. B. Notopodial chaetae from anterior region, posterior row. C. Neuropodial chaetae from anterior region, anterior row. D. Neuropodial chaetae from anterior region, posterior row. E. Capillary from posterior region. F. Hooded hook. G. Sabre chaetae.

Figure 28. Laonice profunda sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 29. SEM of Laonice profunda sp. nov. A. Whole fragment, dorsal view. B. Anterior region, dorsal view. C. Whole fragment, dorsal view. D. Anterior and middle regions, lateral view. Abbreviations: br, branchia; car, caruncle; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium.

Figure 30. Laonice profunda sp. nov. A. Notopodial chaetae from anterior region. B. Neuropodial chaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 31. Laonice brevicarunculata sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; car, caruncle; pe, peristomium; pr, prostomium.

Figure 32. Laonice brevicarunculata sp. nov. A. Notopodial chaetae from anterior region. B. Neuropodial chaetae from anterior region. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

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Tables

Table 1 – Comparative diagnostic characters of Aonides species reported to Brazil.

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Table 1 (continued) – Comparative diagnostic characters of Aonides species reported to Brazil.

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Table 2 – Comparative diagnostic characters of Laonice species from this work.

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Table 2 (continued) – Comparative diagnostic characters of Laonice species from this work.

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Capítulo 6

Spionidae (Annelida) from Espírito Santo Basin, southeastern Brazil: the genera Scolelepis, Dispio, Spio, Microspio and Malacoceros, including a new deep-sea Scolelepis species

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista-alvo: Zootaxa Status: A submeter

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Spionidae (Annelida) from Espírito Santo Basin, southeastern Brazil: the genera Scolelepis, Dispio, Spio, Microspio and Malacoceros, including a new deep-sea Scolelepis species

Running title: NEW SPIONIDAE FROM BRAZIL

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract In this work, 17 species belonging to the genera Scolelepis, Dispio, Microspio, Spio and Malacoceros are reported to the Espírito Santo Basin, southeastern Brazil. Eight species are new to Science, including a deep-sea Scolelepis species – Scolelepis profunda sp. nov., Dispio levisi sp. nov., Microspio alexandrae sp. nov., Microspio tridentata sp. nov., Microspio quadridentata sp. nov., Spio anaclaudiae sp. nov., Spio capixaba sp. nov. and Malacoceros aureus sp. nov. Species can be distinguished mostly by the prostomium shape, branchiae morphology, presence and starting chaetiger of sabre chaetae and distribution and morphology of hooded hooks. A dichotomic key to Spio and Microspio species recorded in this work is provided.

Keywords: Morphology, Annelida, branchiae, deep-sea.

Introduction The annelid family Spionidae Grube, 1850 are one of the most diverse and abundant families of annelids, found mostly on soft-bottom sediments, particularly in eutrophic environments. There approximately 580 valid species distributed in 39 genera (Blake et al. 2017), although the group’s diversity is possibly much higher, as the family has a history of taxonomic problems with many well-documented species-complexes (Sato-Okoshi et al. 2016), specimens with suspiciously

306 global distributions, genera that never were revised and many insufficiently studies areas, both from coastal and deep-sea environments.

Currently, about 90 spionid species arranged in 19 genera are known to Brazil (Amaral et al. 2013; Peixoto & Paiva 2019), although spionid knowledge in Brazil is incipient, as most studies are restricted to the southern and southeastern coasts of Brazil, areas where most Brazilian polychaetologists are centered (Lana et al., 2017) and also are restricted to the upper part of the continental shelf (above 50 m depth), with only a few works focused on the continental slope (Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019, ms). Only a few taxa have been studied in greater details, such as Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Laonice Malmgren, 1867 (Kinberg 1866; Nonato et al. 1986; Radashevsky & Lana 2009), Laubieriellus Maciolek, 1981 (Peixoto & Paiva 2019), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), Polydora Bosc, 1802 (Radashevsky et al. 2006; Radashevsky & Migotto, 2017), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012) and Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017), Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018). Spionid Brazilian surveys including multiple genera are scarce, focused on the continental shelf of the state of Paraná, southern coast (Bolívar & Lana 1987), on the continental slope of Campos Basin, state of Rio de Janeiro, southeastern coast (Paiva & Barroso 2010) and on both southern and southeastern coasts (Pardo et al. 2006; Pardo & Peixoto in press). However, none of the aforementioned works described new taxa.

According to Blake et al. (2017), the genera Dispio Hartman, 1951/ Scolelepis; Spio Fabricius, 1785/ Microspio Mesnil, 1896 and Malacoceros Quatrefages, 1843/ Rhynchospio Hartman, 1936 represents generic pairs in which one of the genera possess branchiae starting from chaetiger 1 while the other possess branchiae starting from chaetiger 2.

The genus Scolelepis was erected by Blainville (1828), who transferred Lumbricus squamata Müller, 1806 to Scolelepis, based on material from Denmark. Scolelepis is one of the most specious spionid genus, with over 80 species, all collected in shallow depths – from the intertidal to continental shelf depths. The genus was reviewed by Maciolek (1987), who erected two subgenera, based mainly on hook morphology – species bearing uni- to tridentate hooded hooks and a straight shaft were assigned to Scolelepis (Scolelepis), while species with multidentate hooded hooks and a curved shaft were assigned to Scolelepis (Parascolelepis). So far, nine Scolelepis species are reported from Brazil: S. (S.) andradei, S. (S.) chilensis, S. (S.) gaucha, S. (S.) goodbodyi, S. (S.) lighti, S. (Parascolelepis) quinquidentata, S (S.) squamata, S. (P.) texana and Scolelepis (Scolelepis) acuta (Rocha et al. 2009; Rocha & Paiva 2012; Amaral et al. 2013).

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This last species has a long story of taxonomic problems, such as a very brief original description – only three sentences in Latin, according to Radashevsky (2012) and incorrect synonymizing of valid species (see Pettibone 1963 and Foster 1971a), which led to a supposedly immense morphological variability. Currently, S. squamata is considered a cosmopolitan species, with records from China (Zhou et al. 2009), the USA (Pettibone 1963), Jamaica (Pettibone 1963), France (Pettibone 1963), South Africa (Pettibone 1963) and Brazil (Amaral et al. 2013), being the Scolelepis species with most records in the Brazilian Coast – almost 100 records. The veracity of the Brazilian records will be addressed in a future work.

Its generic pair, the genus Dispio, is found in shallow sublittoral environments, such as the surf zone of sandy beaches, frequently co-occurring with Scolelepis. The genus Dispio and Scolelepis are superficially similar due to a fusiform and anteriorly narrowed prostomium and branchiae partially fused to the postchaetal notopodial lamellae, but the genus differs on the starting chaetiger of branchiae – chaetiger 1 in Dispio and chaetiger 2 in Scolelepis, occurrence of hooded hooks – only in the neuropodia in Dispio and on both rami in Scolelepis and hook morphology – uni- or bidentate in Dispio and uni- to multidentate in Scolelepis. Besides these characters, Dispio presents lobed anterior postchaetal lamellae, long row of notochaetae on chaetiger 1 and pygidial cirri, not found in Scolelepis. Three species are known to the Brazilian Coast: Dispio magna, D. remanei and D. uncinata the latter being considered as a species-complex, as, after being described to the Gulf of Mexico (Hartman 1951), it was supposedly recorded in Panama, California (Pacific Coast of the USA), Chile, Brazil and even Japan (Delgado-Blas & Díaz-Díaz 2016).

As for the second generic pair, Spio and Microspio are often found in small numbers in marine soft-bottom substrates, although some species (such as Spio limicola Verrill, 1879) can reach densities of thousands of specimens/m2. The main difference between these genera is the starting chaetiger of branchiae – from chaetiger 1 in Spio and from chaetiger 2 in Microspio. The genera were separated by Söderström (1920) based on the number of ciliary bands on the metameric dorsal ciliated organs – two bands in Microspio and four bands in Spio, although at least one Spio species, Spio goniocephala possess only two ciliated bands on the metameric ciliated organs. According to Bick and Meiβner (2011), most Microspio species bear bacillary glands on the postchaetal lamellae and anal cirri, while Spio do not possess such glands.

Foster (1971) treated Microspio as a subgenus of Spio, as she considered that these genera had a nebulous distinction and even admitted that it might be more practical to synonymize them in the future. According to Radashevsky (2012), in both Spio and Microspio larvae, branchiae first develop on chaetiger 3, appearing on chaetiger 2 in juveniles and later on chaetiger 1 in case of Spio, difficulting the identification of juvenile forms.

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Maciolek (1990) reviewed both genera and transferred some Microspio species to Spio and vice versa. According to the author, Spio species are mainly found in continental shelf depths, while Microspio can be found on the continental shelf and continental slope depths. So far, four Spio species have been recorded in the Brazilian coast (Amaral et al. 2013): Spio filicornis (Müller, 1776), S. pettiboneae Foster, 1971, S. quadrisetosa Blake, 1983 and S. setosa Verrill, 1873, while three Microspio species have been previously recorded in Brazil (Amaral et al. 2013; Pardo & Peixoto, in press): M. microcera (Dorsey, 1977), M. minuta (Hartmann-Schröder, 1962) and M. pigmentata (Reish, 1959). Nevertheless, these records should be used with caution, as some species referred to the Brazilian coast have been described to distant regions, with very contrasting environmental parameters (e.g. bathymetry, temperature, daylight length, etc.).

The last generic pair, Malacoceros and Rhynchospio, represents two small genera that are mostly found in shallow soft-bottom substrates. The genus Malacoceros is remarkably similar to Rhynchospio due to its large size, smooth and cirriform branchiae and presence of frontal horns. The genus Rhynchospio have been considered as a subgenus of Malacoceros (Pettibone 1963; Foster 1971), although currently the genera are separated based on reproductive characters (Radashevsky et al. 2014, 2016) and starting chaetiger of branchiae – chaetiger 1 in Malacoceros and chaetiger 2 in Rhynchospio. Only two Malacoceros species have been reported to the Brazilian coast: M. vanderhorsti (Augener, 1927) and Malacoceros indicus (Fauvel, 1928). This latter species, described to the Gulf of Mannar (India), Indian Ocean, is doubtfully considered a cosmopolitan species, being recorded from Caribe, (Foster 1971), Chile (Williams 2007), Australia (Williams 2007), Japan (Imajima 1991) and Brazil (Amaral et al. 2013). No Rhynchospio species are known to the Brazilian coast, although a new species from Guanabara Bay, Rio de Janeiro state, southeastern Brazil is being described by Mendes et al. (in preparation).

17 species have been recorded in this work, based on material from the Espírito Santo Basin, Espírito Santo state, southeastern Brazil, during the “Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company). Eight species are new to science, while two species are new records to the Brazilian coast, and six additional taxa may represent new species, although additional material is needed to make a final taxonomic decision.

Material and Methods Samples were collected during the AMBES Project on the Espírito Santo Basin and northern portion of Campos Basin between 2010 and 2013 (Figure 1). Sediment samples were taken using a Van Veen Grab sampler on the continental shelf and Doce River mouth, and using a box corer

309 in the continental slope and canyons. Sampling was divided in two campaigns: summer months (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and winter months (sample codes Amb2, Amb11, Amb12, Amb13, and Amb14).

Sediment samples were then fixed in situ in an unbuffered solution of 10% formalin, washed in the laboratory using a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm-mesh sieve (for samples collected on the continental slope and on the Watu Norte and Doce canyons) and stored in 70% EtOH. Samples were then elutriated (using the same mesh size as described above), specimens were sorted out, examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope, and were measured using a calibrated eyepiece attached to the microscope.

Specimens were stained in a methyl green solution (1 g/L in 70% EtOH) for 10‒15 s, then examined in distilled water to avoid stain fading, in order to enhance visualization of features such as branchial scars and to observe any species-specific staining patterns. As specimens were fixed and kept in formalin for several months, it was decided not to undertake molecular studies.

For Scolelepis species recorded from the Brazilian coast in this study and on previous works (Rocha et al. 2009; Rocha & Paiva 2012, Amaral et al. 2013), main diagnostic characters were summarized in Table 1, while similar comparison tables were provided for Dispio (Table 2), Spio (Table 4) and Malacoceros (Table 5). A comparison table for Microspio species recorded in this work is also provided (Table 3). For Spio and Microspio species, an identification key is provided.

Specimen lots deposited in the Smithsonian National Museum of National History (USNM), Smithsonian Institution, Washington, D.C., USA were examined and compared with material collected in this study. Type material of Dispio latilamella Williams, 2007 USNM 1096794 (holotype), Dispio brachychaeta Blake, 1983 USNM 67512 (holotype), Microspio profunda Maciolek, 1990 USNM 80480 (holotype), Microspio tetrabranchia Maciolek, 1990 USNM 129435 (holotype) and Spio quadrisetosa Blake, 1983 USNM 67520 (holotype), as well as non- type material of Microspio profunda USNM 80481 (25 specimens) were examined.

Prior to scanning electron microscope (SEM) studies, studies, specimens were dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each, followed by three changes of 100%

EtOH), then critical-point dried using CO2. Then, dried specimens were mounted on stubs sputter coated with gold and viewed in a JEOL JSM-6510 LV SEM. To study the hook morphology, middle or posterior fragments were sonicated in distilled water for 20 s at 40 kHz, in order to rupture the hoods. A single specimen of Microspio profunda deposited in the USNM (USNM

80481), was dehydrated in a graded ethanol series, critical-point dried in CO2, mounted on stubs,

310 sputter coated with carbon and gold-palladium (10 µm thick each) and examined using a Philips XL-30 SEM.

Results Taxonomy

Phylum Annelida Lamarck, 1809

Order Spionida Fauchald, 1977

Family Spionidae Grube, 1850

Genus Scolelepis Blainville, 1828

Type species: Lumbricus squamata Müller, 1806 [= Scolelepis squamata (Müller, 1806)], by monotypy.

List of synonyms: Aonis sensu Audouin & Milne Edwards, 1833

Asetocalamyzas Tzetlin, 1985 (type by original designation)

Nerine Johnston, 1838 (subjective synonym)

Nerinides Mesnil, 1896 (subjective synonym)

Pseudomalacoceros Czerniavsky, 1881 (subjective synonym)

Pseudonerine Augener, 1926 (junior homonym, subjective synonym)

Scolecolepis Malmgren, 1867 (unjustified spelling emendation of Scolelepis Blainville, 1828)

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Diagnosis (modified from Blake et al. 2017): Prostomium fusiform, pointed anteriorly, extending posteriorly as narrow caruncle; caruncle attached or detached posteriorly; occipital tentacle present or absent. Peristomium well developed, lateral wings present or absent; palps without ciliated groove, often with thickened basal sheath. Branchiae from chaetiger 2 to near posterior end; anterior branchiae either completely fused to notopodial lamellae, partially fused, or entirely free; posterior branchiae usually free from notopodial lamellae; accessory branchiae present or absent. Anterior chaetae all limbate capillaries, posterior chaetae include capillaries and uni- to tridentate hooks with slightly curved shaft (subgenus Scolelepis) or multidentate hooks with strongly curved shaft (subgenus Parascolelepis). Hooks lacking secondary hood. Pygidium as an oval disk (sometimes referred as cushion) or multilobed.

Scolelepis profunda sp. nov.

(Figs 2–3D)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 06 Jun to 17 Jul 2013, 970m, MNRJP XXXX. Paratypes: Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 02 Dec 2011 to 02 Feb 2012, 1023m, MNRJP XXXX (1 ind).

Additional material examined. Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (2 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (1 ind); Amb7 D5, 19°46'34,99" S 39°30'4,65" W, 428m (1 ind).

Description: A medium-sized Scolelepis, largest specimen 5.5 mm long, 2.2 mm wide at the widest point for 23 chaetigers (incomplete), holotype being the largest specimen. Fragment cylindrical on anterior and middle regions, laterally widened on first 12–14 chaetigers and tapering towards the posterior region. Segmental ciliary bands on the anterior region. Body color light yellow in alcohol (Fig 2).

Prostomium fusiform with anteriorly pointed tip, extending posteriorly as an inflated caruncle to the posterior margin of chaetiger 1, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 3. Eyes absent. Occipital antenna absent. Peristomium well- developed, surrounding prostomium for about four fifths of its total length, lacking lateral wings

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(Fig 2). Palps grooved in juveniles, reaching up to chaetiger 10, lost in adult specimens. Palps with inflated and rounded bases, palp sheaths not observed.

Chaetiger 1 lacking notopodial chaetae. Neuropodium with only a few chaetae, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae conical on notopodium and rounded with a pointed tip on neuropodium, smaller than lamellae on succeeding chaetigers, especially on notopodium. Prechaetal lamellae absent.

Notopodial postchaetal lamellae narrow with pointed tip on anterior and middle regions, rounded with a dorsal cirriform projection after lamellae separates from branchiae (see below); morphology unknown on posterior region. Postchaetal lamellae well-developed on anterior region, gradually reducing in size towards middle region. Postchaetal lamellae smooth, not ruffled nor serrated (Fig 2). Notopodial prechaetal lamellae present as a low flap on chaetigers 3–19. Lateral ciliated organs conical, present from chaetiger 1 to end of fragment (chaetiger 23).

Neuropodial postchaetal lamellae rounded with a pointed tip on chaetigers 2–18, largest on chaetigers 4–6 (Fig 2). Shallow notch present from chaetiger 13, completely separating lamellae on chaetiger 19. On chaetiger 19, postchaetal lamellae divided in superior and inferior lobes; superior lobe subtriangular, inferior lobe rounded, both gradually reducing in size on middle region. Neuropodial prechaetal lamellae absent.

Chaetae from notopodia and neuropodia organized in two rows of granulated and widely bilimbate capillaries (Fig 3A). Chaetae from both rows of equal length, although notopodia chaetae are up to 1.4 times longer than neuropodial chaetae (Fig 3B). Towards the end of the fragment, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 3C). Chaetae of posterior region unknown.

Hooks in neuropodia from chaetiger 18, up to 10 per fascicle, accompanied by 2–5 short non- limbate capillaries, absent on middle region. Inferior limbate capillaries absent. Hooks tridentate, with secondary teeth arranged side by side above main tooth (Fig 3D). Secondary hood absent. Notopodial hooded hooks not observed (specimen incomplete).

Branchiae from chaetiger 2, present throughout entire fragment, completely fused with notopodial postchaetal lamellae until chaetiger 17. Towards end of fragment (chaetiger 23), branchiae digitiform, fused only basally with notopodial postchaetal lamellae. Branchiae intensely ciliated on the internal margin, connected by ciliary bands to the branchiae of the same chaetiger, bands prominent on anterior region. Accessory branchiae absent.

Pygidium of unknown morphology.

Sperm plates from chaetiger 15.

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Methyl green staining pattern: Postchaetal lamellae from chaetigers 2–15 weakly stained.

Remarks. Scolelepis profunda sp. nov. can be readily distinguished from all Scolelepis species by its bathymetric range, being found on the continental slope only, while all other species occur on depths less than 100m, with most species inhabiting the intertidal zone.

Among Brazilian species, S. profunda sp. nov. is only superficially similar to S. goodbodyi and S. acuta in having a prostomium extending posteriorly to the end of chaetiger 1 and tridentate hooded hooks, although species can be readily distinguished by the lack of notochaetae on chaetiger 1, present in S. goodbodyi and S. acuta; starting chaetiger of neuropodial hooded hooks – chaetiger 18 in S. profunda sp. nov., chaetiger 28 in S. acuta and chaetigers 24–28 in S. goodbodyi; as well as branchiae completely fused to notopodial lamellae on anterior chaetigers and presence of occipital antenna in S. profunda sp. nov. (Jones 1962; Delgado-Blas 2006; Rocha & Paiva 2012).

Scolelepis profunda sp. nov. is also superficially similar to S. squamata, a species with a complicated taxonomic history, being recorded worldwide since several species were synonymized with S. squamata (Pettibone 1963; Blake et al. 2017). Species are similar in having a prostomium extending posteriorly to the end of chaetiger 1, partial overlap on the starting chaetiger of neuropodial hooded hooks – chaetiger 18 in S. profunda sp. nov. and chaetigers 14– 43 in S. squamata and tridentate hooded hooks (see below) and neuropodial lamellae divided on middle chaetigers, although species can be easily differentiated by the lack of notochaetae on chaetiger 1, present in S. squamata; morphology of neuropodial hooded hooks – as S. squamata can also bear uni- and bidentate hooded hooks; degree of fusion between branchiae and lamellae on anterior chaetigers – completely fused in S. profunda sp. nov. and almost completely fused in S. squamata and presence of occipital antennae in S. profunda sp. nov., lacking in S. squamata (Delgado-Blas 2006; Surugiu 2016).

Material from this species preserved in 96% EtOH was kindly donated by the Laboratory of Deep- Sea Ecology and Evolution (Oceanographic Institute, Universidade de São Paulo, São Paulo, Brazil) and will be addressed in future works (Peixoto et al., in preparation).

Etymology. The specific epithet, profunda, refers to the depth where this species was collected, being the deepest record for the genus.

Habitat: very fine sand to fine silt, 410–1024 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

314

Scolelepis sp. 1

(Figs 4–5E)

Material examined. Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (2 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind).

Description: A medium-sized Scolelepis species, largest complete specimen 4.5 mm long, 0.5 mm wide at the widest point for 21 chaetigers. Body wide and cylindrical on anterior and middle regions, slightly flattened dorsoventrally on posterior region, tapering towards the pygidium. Segmental ciliary bands present on anterior and middle segments. Body color light yellow to whitish in alcohol (Fig 4).

Prostomium fusiform with anteriorly pointed tip, extending posteriorly as an inflated caruncle to the anterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 2. Eyes absent. Occipital antenna absent. Peristomium well- developed, surrounding prostomium for about two thirds of its total length, lacking lateral wings (Fig 4). Palps lost in all specimens.

Chaetiger 1 lacking notopodial chaetae. Neuropodium with only a few chaetae, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae conical on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae narrow with pointed tip throughout, well-developed on anterior and middle regions, gradually reducing in size towards the posterior region. Postchaetal lamellae smooth, not ruffled nor serrated. Notopodial prechaetal lamellae rounded from chaetiger 2 to chaetigers 13–20 (depending on specimen size), reduced to a low flap afterwards (Fig 4). Lateral ciliated organs absent between notopodia and neuropodia.

Neuropodial postchaetal lamellae rounded on anterior region, triangular with a digitiform dorsal projection on middle region and gradually reducing in size towards the posterior region (Fig 4). Neuropodial prechaetal lamellae rounded from chaetiger 2 to chaetigers 12–18 (depending on specimen size), reduced to a low flap afterwards.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated unilimbate capillaries. On notopodia, limbation narrow and rows of the same length (Fig 5A), while in neuropodia, anterior row short and narrowly unilimbate (Fig 5B) and posterior row long (up to 1.5 times longer than anterior row) and widely limbate capillaries (Fig 5C). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 5D).

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Hooks in neuropodia only, starting from chaetigers 15–17, up to 12 per fascicle, accompanied by 1–3 short non-limbate capillaries, absent on posterior chaetigers. Inferior limbate capillaries absent. Hooks quinquedentate, with secondary teeth arranged in two rows above main tooth (Fig 5E). Superior row of teeth very small, usually not observed in light microscopy. Secondary hood absent.

Branchiae from chaetiger 2, present almost on the whole body, absent on last 3 chaetigers, completely fused with notopodial postchaetal lamellae until chaetigers 11–16 (depending on specimen size). On last branchiate chaetiger, branchiae fused with notopodial postchaetal lamellae for half of its length. Branchiae intensely ciliated on the internal margin, connected by ciliary bands to the branchiae of the same chaetiger, bands prominent on anterior and middle regions, interspersed with segmental ciliary bands. Accessory branchiae absent.

Pygidium rounded, with entire margin, cushion-like.

Methyl green staining pattern: Margins of pre- and postchaetal notopodial lamellae intensely stained.

Remarks. Scolelepis sp. 1 belongs to the subgenus Parascolelepis Maciolek, 1987, as it bears multidentate hooded hooks, occipital antenna, lack of division on neuropodial postchaetal lamellae from posterior chaetigers and lack of notopodial hooded hooks. Another character commonly associated with this subgenus, the presence of palp sheaths, could not be observed, since all specimens had lost its palps.

Scolelepis sp. 1 is similar to S. texana Foster, 1971, from Texas (USA) in having a prostomium posteriorly extended to the start of chaetiger 2, presence of occipital antenna, lack of notochaetae on chaetiger 1, slight overlap on the starting chaetiger of neuropodial hooded hooks – chaetigers 15–17 in Scolelepis sp. 1 and chaetiger 13–27 in S. texana and branchiae completely fused with notopodial postchaetal lamellae on anterior chaetigers, although species can be separated based on the lack of division of neuropodial lamellae in S. sp. 1, present from chaetiger 28 in S. texana and hook morphology – quinquedentate in S. sp. 1 and heptadentate in S. texana.

Scolelepis sp. 1 is most similar to S. quinquedentata Hartmann-Schröder, 1965, from Chile, as both species possess a prostomium posteriorly extended to the start of chaetiger 2, presence of occipital antenna, lack of notochaetae on chaetiger 1, remarkable overlap on the starting chaetiger of neuropodial hooded hooks – chaetigers 15–17 in Scolelepis sp. 1 and chaetiger 16 in S. quinquedentate and same hook morphology – quinquedentate. However, species can be distinguished by fusion of branchial and notopodial lamellae on anterior chaetigers – completely fused in Scolelepis sp. 1 and only partially fused in S. quinquedentata. Furthermore, no complete specimen was described by Hartmann-Schröder (1965), hence important characters such as the

316 presence of notopodial hooded hooks and division of neuropodial lamellae (undivided, according to Maciolek [1987]) are unknown.

Despite similarities with S. quinquedentata, which was previously recorded in Brazil (Lana et al. 2006), it was decided not to formally describe this species until additional material becomes available, as a very small number of specimens was collected and the single complete specimen is in poor condition.

Habitat: fine sand to medium silt, 25–41 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Scolelepis sp. 2

(Figs 6–7F)

Material examined. Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (1 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (1 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W, 46m (1 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (1 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (1 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (1 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (1 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (1 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (1 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (1 ind).

Description: A medium-sized Scolelepis, largest specimen 6 mm long, 0.65 mm wide at the widest point for 29 chaetigers (incomplete). Fragment cylindrical on anterior region slightly flattened dorsoventrally on middle region, tapering towards posterior region. Segmental ciliary bands absent. Body color light yellow to whitish in alcohol (Fig 6).

Prostomium fusiform with anteriorly pointed tip and two short lateral protuberances, extending posteriorly as an inflated caruncle to the anterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 3. Two pairs of eyes in trapezoidal arrangement. Occipital antenna absent. Peristomium well-developed, surrounding prostomium for about four fifths of its total length, lacking lateral wings (Fig 6). Palps lost in all specimens.

Chaetiger 1 lacking notopodial chaetae. Neuropodium with only a few chaetae, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and rounded on

317 neuropodium, smaller than lamellae on succeeding chaetigers, especially on notopodium. Prechaetal lamellae absent.

Notopodial postchaetal lamellae narrow with pointed tip on anterior and middle regions and square-shaped on the posterior region. Postchaetal lamellae well-developed on anterior and middle regions, gradually reducing in size towards the posterior region (Fig 6). Postchaetal lamellae smooth, not ruffled nor serrated. Notopodial prechaetal lamellae rounded from chaetiger 2 to chaetigers 14–19 (depending on specimen size), reduced to a low flap afterwards. Lateral ciliated organs absent between notopodia and neuropodia.

Neuropodial postchaetal lamellae rounded on chaetigers 2–5 and elliptical on chaetigers 6–15 (Fig 6). On chaetiger 16, postchaetal lamellae divided in superior and inferior elliptical lobes; superior lobe reduced to a low flap after chaetiger 20, inferior lobe absent after chaetiger 18. Neuropodial prechaetal lamellae as a low flap from chaetiger 13 to chaetiger 19.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, anterior row of long (up to 1.2 times longer than posterior row), unilimbate and striated capillaries (Fig 7A); posterior row of short and widely unilimbate capillaries (Fig 7B). On neuropodia, anterior row of short, unilimbate and granulated capillaries (Fic 7C); posterior row of long (up to two times longer than anterior row), narrowly limbate and non-granulated capillaries (Fig 7D). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 7E).

Hooks in neuropodia from chaetigers 13–15, up to 10 per fascicle, accompanied by 1–3 short non- limbate capillaries, absent on posterior chaetigers. Inferior limbate capillaries absent. Hooks quadridentate, bearing a main tooth, secondary teeth arranged side by side and an additional unpaired tooth at the top (Fig 7F). Secondary hood absent. Notopodial hooded hooks not observed (no complete specimen examined).

Branchiae from chaetiger 2, present throughout entire fragment, completely fused with notopodial postchaetal lamellae until chaetigers 15–19 (depending on specimen size). Towards end of longest fragment (chaetiger 29), branchiae digitiform, and fused with notopodial postchaetal lamellae for half of its length. Branchiae intensely ciliated on the internal margin, connected by ciliary bands to the branchiae of the same chaetiger, bands prominent on anterior region. Accessory branchiae absent.

Pygidium of unknown morphology.

Oocytes (measuring up to 300 µm), consistently from chaetiger 13. Sperm plates consistently from chaetiger 13.

318

Methyl green staining pattern: Prostomium (except inflated caruncle), peristomium and margins of postchaetal lamellae from chaetigers 2–16 strongly stained; ventral side from chaetigers 2–13 weakly stained.

Remarks. Scolelepis sp. 2 is similar to S. quinquedentata in having a prostomium extending posteriorly to the start of chaetiger 2, presence of occipital antenna, lack of notochaetae on chaetiger 1 and branchiae completely fused to notopodial postchaetal lamellae, although species can be distinguished based on the starting chaetiger of neuropodial hooded hooks – chaetigers 13–15 in S. sp. 2 and chaetiger 16 in S. quinquedentata, as well as hook morphology – quadridentate in S. sp. 2 and quinquedentate in S. quinquedentata. However, the description of S. quinquedentata was based on incomplete specimens, thus characters such as the presence of notopodial hooded hooks and division of neuropodial lamellae are unknown (Hartmann-Schröder 1965).

Scolelepis sp. 2 presents division on neuropodial lamellae on chaetiger 16, which is unusual, as this character usually occurs on more posterior chaetigers (usually with a slight intraspecific variation). Among Brazilian species, only S. goodbodyi (Jones 1962) presents a similar starting chaetiger – from chaetigers 16–19. However, species can be readily distinguished as Scolelepis sp. 2 lacks notochaetae on chaetiger 1, present in S. goodbodyi; starting chaetiger of neuropodial hooded hooks – from chaetigers 13–15 in S. sp. 2 and chaetigers 24–28 in S. goodbodyi; morphology of hooded hooks – quadridentate in S. sp. 2 and uni- to tridentate in S. goodbodyi, fusion of branchiae and notopodial lamellae on anterior chaetigers – completely fused in S. sp. 2 and partially fused in S. goodbodyi and lack of occipital antennae in S. goodbodyi.

All specimens examined in this work were incomplete specimens in less than ideal conditions, so it was decided not to describe a new species until additional material becomes available.

Habitat: medium sand to coarse silt ,38–158 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Scolelepis sp. 3

(Figs 8–9E)

Material examined. Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (1 ind).

Description: A medium-sized Scolelepis species, specimen 4.3 mm long, 1 mm wide at the widest point for 25 chaetigers (incomplete). Fragment cylindrical on anterior and middle regions,

319 tapering towards the posterior region. Segmental ciliary bands present on anterior and middle segments. Body color light yellow in alcohol (Fig 8).

Prostomium fusiform with anteriorly pointed tip, extending posteriorly as an inflated caruncle to the posterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 3. Two pairs of small eyes in trapezoidal arrangement. Occipital antenna absent. Peristomium well-developed, surrounding prostomium for about four fifths of its total length, lacking lateral wings (Fig 8). Palps lost.

Chaetiger 1 lacking notopodial chaetae. Neuropodium with only a few chaetae, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae narrow with pointed tip on anterior and middle regions, reduced and square-shaped after lamellae separates from branchiae (see below); morphology unknown on posterior region. Postchaetal lamellae well-developed on anterior region, gradually reducing in size on middle region. Postchaetal lamellae smooth, not ruffled nor serrated (Fig 8). Notopodial prechaetal lamellae as a low flap on chaetigers 2–13, absent on remaining chaetigers. Lateral ciliated organs absent between notopodia and neuropodia.

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetiger 10 and broadly rounded afterwards. Shallow notch present from chaetiger 16, completely separating lamellae on chaetiger 17 (Fig 8). On chaetiger 17, postchaetal lamellae divided in superior and inferior lobes; superior lobe square-shaped on chaetigers 17–19 and elongate, subtriangular from chaetiger 20 onwards, gradually reducing in size on middle region; inferior lobe rounded on chaetigers 17–22, absent from chaetiger 23 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, rows of narrowly limbate, non-granulated and striated capillaries (Fig 9A). On neuropodia, anterior row of short, narrowly unilimbate and lightly granulated capillaries (Fig 9B) and posterior row of log (up to 1.5 times longer than anterior row), widely unilimbate and lightly granulated capillaries (Fig 9C). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 9D). Chaetae of posterior region unknown.

Hooks in neuropodia from chaetiger 17, up to 13 per fascicle, accompanied by 1–3 short non- limbate capillaries, absent on middle region. Inferior limbate capillaries absent. Hooks tridentate, with secondary teeth arranged in a vertical row above main tooth (Fig 9E). Secondary hood absent. Notopodial hooded hooks not observed (specimen incomplete).

Branchiae from chaetiger 2, present throughout entire fragment, completely fused with notopodial postchaetal lamellae until chaetiger 18. Towards end of fragment (chaetiger 26), branchiae

320 digitiform, completely separated from notopodial postchaetal lamellae. Branchiae intensely ciliated on the internal margin, connected by ciliary bands to the branchiae of the same chaetiger, bands prominent on anterior region. Accessory branchiae absent.

Pygidium of unknown morphology.

Sperm plates from chaetiger 16.

Methyl green staining pattern: Inflated caruncle and margins of lamellae from chaetigers 1–15 intensely stained.

Remarks. Scolelepis sp. 3 differs from all Brazilian Scolelepis species in having a prostomium extending posteriorly to the end of chaetiger 2, nuchal organs extending to the end of chaetiger 3, lack of notochaetae on chaetiger 1, neuropodial lamellae divided from chaetiger 19, tridentate hooded hooks from chaetiger 17, branchiae completely fused to notopodial lamellae on anterior chaetigers and presence of occipital tentacle.

Since a single incomplete specimen was collected, important taxonomic characters such as the presence of palp sheaths and presence of notopodial hooded hooks remain unknown, thus it was decided not to describe this species until additional material becomes available.

Habitat: medium sand, 32 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Genus Dispio Hartman, 1951

Type species: Dispio uncinata Hartman, 1951 (by monotypy)

Diagnosis (modified from Blake et al. 2017): Prostomium fusiform pointed anteriorly, with short caruncle extending posteriorly; eyes present or absent. Occipital tentacle absent. Peristomium moderately developed, forming low lateral wings. Anterior postchaetal lamellae lobed or entire.

Prechaetal lamellae present or absent. Prechaetal notopodial and neuropodial lobes present or absent. Single row of long notopodial capillary chaetae usually present on chaetiger 1. Branchiae from chaetiger 1, fused to notopodial lamellae for half or more of length. Accessory branchiae present along posterior margin of notopodia in middle and posterior chaetigers. Notochaetae

321 capillaries; neurochaetae include capillaries, hooded hooks and sabre chaetae. Hooded hooks uni or bi-dentate. Pygidium with mid-ventral flap and prominent anal cirri.

Dispio levisi sp. nov.

(Figs 10–12G)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 11 Dec to 19 Dec 2010, 41m, MZUSP XXXX. Paratypes: Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 12 Jul to 18 Jul 2011, 34m, MZUSP XXXX (2 ind), MNRJP XXXX (2 ind).

Additional material examined. Amb1 Foz6, 19°54'56,16" S 39°56'40,83" W, 34m (1 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (1 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (3 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (1 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W, 34m (4 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (3 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (15 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (1 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (1 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (1 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (1 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (2 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (1 ind).

Description: A medium-sized Dispio species, largest complete specimen 3.5 mm long, 0.9 mm wide at the widest point for 65 chaetigers, holotype, 25 mm long, 1.7 mm wide at the widest point for 70 chaetigers (incomplete). Body dorsoventrally flattened on the anterior region and cylindrical afterwards, tapering towards the pygidium. J-shaped metameric dorsal ciliated organs from chaetiger 2, easily visible on anterior chaetigers. Body color whitish in alcohol (Fig 10).

Prostomium fusiform, with elongated and anteriorly pointed tip, extending posteriorly to the posterior margin of chaetiger 1. Two pairs of eyes in trapezoidal arrangement. Peristomium surrounding prostomium for most of its length, not fused to the first chaetiger, and lacking lateral wings (Figs 10, 11A–C). Peristomium well separated from prostomium by a shallow groove. Palps lost in all specimens.

Chaetiger 1 with fewer chaetae than succeeding chaetigers. Row of long capillaries on the notopodium (Figs 10, 11A–C, 12A). Postchaetal lamellae foliaceous and fused with branchia for

322 most of its length on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous throughout, partially fused with branchiae, gradually reducing in size towards the posterior region (Figs 10, 11A–B). On middle segments, notopodial postchaetal lamellae develops a triangular ventral edge. Postchaetal lamellae fused with branchiae for most of its length on the anterior region and fused for half of its length on the posterior region. Postchaetal lamellae smooth, not ruffled nor serrated. Notopodial prechaetal present as a low flap on anterior chaetigers, becoming foliaceous with a pointed tip towards the posterior region. Dorsal crests very low, starting from chaetigers 19–29 (Fig 10). Between notopodia and neuropodia, presence of lateral organs (interramal organs) from chaetiger 1, throughout anterior and middle regions, organs not observed in the posterior region (Fig 11C).

Neuropodial postchaetal lamellae rounded throughout, gradually reducing in size towards the posterior region (Figs 10, 11C). On middle segments, neuropodial postchaetal lamellae develops a triangular dorsal edge. Postchaetal lamellae smooth, not ruffled nor serrated. Neuropodial prechaetal lamellae present as a low flap on anterior chaetigers, becoming rounded (fan-shaped) towards the posterior region.

Chaetae from notopodia and neuropodia organized in two rows, anterior row of intensely granulated and widely unilimbate capillaries (Fig 12B) and posterior row of long (up to 1.2 times length of anterior row), lightly granulated and narrowly unilimbate capillaries (Fig 12C). Above the notopodial rows, additional tuft of 2–4 long, wide, intensely granulated and distally unilimbate capillaries (Fig 12D). Notopodial chaetae up to two times longer than neuropodial chaetae. Towards the posterior region, chaetae of both rami progressively become elongate, non-limbate, thinner, and less numerous (Fig 12E).

Hooks in neuropodia only, starting from chaetigers 16–25, up to eight per fascicle, accompanied by 1–8 short non-limbate capillaries. Hooks unidentate (Fig 12F). Secondary hood absent. Sabre chaetae from chaetigers 10–14, up to three per fascicle (Fig 12G).

Branchiae from chaetiger 1 onwards. Branchiae long, cirriform, with a pointed tip, fused to the notopodial postchaetal lamellae for most of its length on anterior chaetigers and for about half of its length on posterior chaetigers. Accessory branchiae present from chaetigers 15–24, on the dorsolateral side of body, behind notopodia base. Secondary branchiae appearing as a single digitiform projection, increasing in number on middle chaetigers, up to 5 digitiform projections (Fig 11D); absent on last 10–20 chaetigers (depending on specimen size).

Pygidium rounded, bearing two short lateral cirri.

Oocytes from chaetiger 33, measuring up to 170 µm.

323

Methyl green staining pattern: Anterior and middle regions of the prostomium (except pointed tip), peristomium and margins of all lamellae intensely stained; margins of J-shaped metameric dorsal ciliated organs weakly stained.

Remarks. Dispio levisi sp. nov. differs from all Dispio species due to its set of characters – a fusiform and anteriorly pointed prostomium; row of long notopodial chaetae on chaetiger 1; postchaetal lamellae smooth throughout; sabre chaetae from chaetigers 10–14; branchiae fused to notopodial postchaetal lamellae for most of its length on anterior chaetigers and for about half of its length on posterior chaetigers, accessory branchiae from chaetigers 15–24 and pygidium bearing two pairs of short cirri.

Among species recorded in Brazil, Dispio levisi sp. nov. is similar to D. magna and Dispio cf. brachychaeta in lacking serrations or ruffles on anterior postchaetal lamellae, although Dispio levisi sp. nov. can be distinguished from the aforementioned species by the presence of a long row of notochaetae on chaetiger 1; presence of accessory branchiae; pygidium bearing two short cirri; maximum number of neuropodial hooded hooks – up to eight per fascicle in D. levisi sp. nov., up to 10 per fascicle in D. magna and up to five per fascicle in Dispio cf. brachychaeta and degree of fusion between branchiae and notopodial postchaetal lamellae on anterior chaetigers – fused for most of its length in D. levisi sp. nov. and completely fused in D. magna and Dispio cf. brachychaeta (Blake 1983; Delgado-Blas & Díaz-Diáz 2016).

The species shares similarities with D. uncinata and D. sp. 2 in having a long row of notochaetae on chaetiger 1 and up to eight neuropodial hooded hooks per fascicle. Dispio levisi sp. nov. can be distinguished from both species in having smooth postchaetal lamellae on anterior chaetigers (Hartman 1951; Delgado-Blas & Díaz-Diáz 2016). Dispio levisi sp. nov. can also be separated from D. sp. 2 by the starting chaetiger of accessory branchiae, from chaetigers 15–24 in D. levisi sp. nov. and chaetiger 33 in D. sp. 2.

Dispio levisi sp. nov. can be separated from D. uncinata based on prostomium morphology – fusiform and anteriorly pointed in D. levisi sp. nov. and bearing a pair of lateral projections in D. uncinata; starting chaetiger of sabre chaetae – from chaetigers 10–14 in D. levisi sp. nov. and from chaetiger 2 in D. uncinata; degree of fusion between branchiae and notopodial postchaetal lamellae – fused most of its length on anterior chaetigers and for about half of its length on posterior chaetigers in D. levisi sp. nov. and completely fused to the notopodial postchaetal lamellae on anterior chaetigers and with free tips on the posterior region in D. uncinata and pygidium morphology – bearing a pair of cirri in D. levisi sp. nov. (Delgado-Blas & Díaz-Díaz 2016).

Etymology: The specific epithet, levisi (from “levis”, Latin adjective for smooth), refers to the morphology of postchaetal lamellae on anterior chaetigers, lacking ruffles and serrations.

324

Habitat: coarse sand to coarse silt, 26–54 m depth.

Distribution: Southeastern Brazil (Espírito Santo state), South Atlantic Ocean.

Dispio cf. brachychaeta Blake, 1983

(Figs 13–14F)

Material examined. Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (1 ind).

Comparative material examined. USNM 67512 (holotype).

Description: A medium-sized Dispio species, specimen 5 mm long, 0.8 mm wide at the widest point for 47 chaetigers (incomplete). Body dorsoventrally flattened on the anterior region and cylindrical afterwards, tapering towards the pygidium. Metameric dorsal ciliated organs not observed. Body color light yellow in alcohol (Fig 13).

Prostomium fusiform, with elongated and anteriorly pointed tip, extending posteriorly to the posterior margin of chaetiger 1, flanked by conspicuous nuchal organs extending up to the end of chaetiger 2. Two pairs of eyes in trapezoidal arrangement. Peristomium surrounding prostomium for about two thirds of its length, not fused to the first chaetiger, and lacking lateral wings (Fig 13). Peristomium well separated from prostomium by a shallow groove. Palps lost.

Chaetiger 1 bearing fewer chaetae than succeeding chaetigers. Row of long notopodial capillaries absent. Postchaetal lamellae foliaceous and completely fused with branchia on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous throughout, completely fused with branchiae on anterior chaetigers, gradually reducing in size towards the posterior region (Fig 13). On middle segments, ventral part of notopodial postchaetal lamellae develops a triangular edge with a ventrally pointed tip. Postchaetal lamellae completely fused with branchiae until chaetiger 25, separating gradually – although still fused for half of its length on the middle region. Postchaetal lamellae smooth, not ruffled nor serrated. Notopodial prechaetal present as a low flap on anterior chaetigers, low and rounded on middle chaetigers and gradually reducing towards the posterior region. Dorsal crests absent. Lateral organs present between postchaetal lamellae of notopodia and neuropodia.

Neuropodial postchaetal lamellae rounded throughout, gradually reducing in size towards the posterior region (Fig 13). On middle segments, neuropodial postchaetal lamellae develops a

325 dorsally pointed tip. Postchaetal lamellae smooth, not ruffled nor serrated. Neuropodial prechaetal absent throughout.

Chaetae from notopodia and neuropodia organized in two rows, anterior row of short, intensely granulated and widely unilimbate capillaries (Fig 14A) and posterior row of long (up to 1.4 times length of anterior row), lightly granulated and narrowly unilimbate (limbation almost imperceptible) capillaries (Fig 14B). Above the notopodial rows, additional tuft of 2–4 long, wide, intensely granulated and distally unilimbate capillaries (Fig 14C). Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region, chaetae of both rami progressively become elongate, non-limbate, thinner, and less numerous (Fig 14D).

Hooks in neuropodia only, starting from chaetiger 24, up to five per fascicle, accompanied by 1– 5 short non-limbate capillaries. Hooks unidentate (Fig 14E). Secondary hood absent. Sabre chaetae from chaetiger 5, unilimbate and granulated, up to three per fascicle (Fig 14F).

Branchiae from chaetiger 1 onwards. Branchiae long, cirriform, with a blunt tip, completely fused to the notopodial postchaetal lamellae on anterior chaetigers and for about half of its length on the middle region. Branchia intensely ciliated on the internal margin, connected by ciliary bands to the branchiae of the same chaetiger, bands prominent on middle region. Accessory branchiae absent.

Pygidium of unknown morphology.

Methyl green staining pattern: Anterior and middle regions of the prostomium (except pointed tip) intensely stained.

Remarks. Dispio brachychaeta is a peculiar species, as it lacks elongated notopodial chaetae on chaetiger 1, lacks accessory branchiae and lacks ruffled or serrated margins on anterior chaetigers, characters present in most Dispio species (Blake 1983). The prostomial morphology – fusiform, with elongated and anteriorly pointed tip and branchiae starting on chaetiger 1 are characters that place this species in the genus Dispio. The specimen recorded in this work also exhibits these characters, as well as other characters observed in D. brachychaeta, such as notopodial capillaries organized in two rows, with anterior row short and widely limbate and posterior row long and narrowly unilimbate, and presence of an additional tuft of capillaries above the notopodial rows. However, some differences were found from the original description, such as the presence of eyes, present in our and Pardo & Peixoto (in press) specimens, but lacking in the original description; starting chaetiger of hooded hooks – chaetiger 32 in the original description and chaetiger 25 in specimens from this study, number of hooded hooks – up to three in the original description and up to five in specimens from this study; starting chaetiger of sabre chaetae – from chaetiger 1 in the original description and from chaetiger 5 in specimens from this study, and

326 number of sabre chaetae – up to 6 six in the original description and up to four in specimens from this study. It is unclear if these differences are size-related or if our specimens could be a new species. This issue very much likely won’t be solved in the near future, as there are only three D. brachychaeta specimens ever recorded, the holotype, the specimen examined in this study and a specimen deposited in the “ Museu de Zoologia da Universidade Estadual de Campinas "Adão José Cardoso"” (ZUEC-POL 9873), all anterior fragments. If the specimen described in this work indeed corresponds to Dispio brachychaeta, this is the northern-most record of the species, and its first record to Brazil, as the species was described from off Rio de la Plata, Argentina.

Dispio brachychaeta shares the lack of accessory branchiae and lack of elongated notopodial chaetae on chaetiger 1 with D. magna (Day, 1955), described to South Africa and D. oculata Imajima, 1990, described to Japan, as well as branchiae completely fused with notopodial lamellae on anterior chaetiger and for about half of total length from middle region onwards. However, species can be separated based on morphology of notopodial lamellae – smooth in D. brachychaeta and D. oculata and ruffled in D. magna, starting chaetiger of hooded hooks – chaetigers 24–32 in D. brachychaeta, chaetigers 27–33 in D. oculata and chaetiger 44 in D. magna, starting chaetiger of sabre chaetae – chaetiger 1 (Blake, 1983) or chaetiger 55 in D. brachychaeta (this study), chaetiger 2 D. oculata (Imajima, 1990) and chaetiger 14 in D. magna (Day, 1955) and number of sabre chaetae – from two to six sabre chaetae in D. brachychaeta, 1– 2 sabre in D. oculata and a group of sabre-like chaetae (number not mentioned) in D. magna.

Habitat: Coarse sand, 26 m depth.

Distribution: Atlantic Ocean: Argentina (type locality), Brazil (São Paulo and Espírito Santo states).

Dispio sp. 1

(Figs 15–16H)

Material examined. Amb2 Foz2, 19°47'12,95" S 39°55'15,42" W, 23m (1 ind).

Description: A large -sized Dispio species, specimen 14 mm long, 1.1 mm wide at the widest point for 54 chaetigers (incomplete). Body dorsoventrally flattened on the anterior region and cylindrical afterwards, tapering towards the pygidium. Inverted J-shaped metameric dorsal ciliated organs from chaetiger 2, easily visible on anterior chaetigers. Body color white in alcohol (Fig 15).

327

Prostomium fusiform, with elongated and anteriorly pointed tip, extending posteriorly to the posterior margin of chaetiger 1. Eyes absent. Peristomium surrounding prostomium for most of its length, not fused to the first chaetiger, and lacking lateral wings. Grooved palps reaching up to chaetiger 20, lost in most specimens (Fig 15). Palp sheaths present, large and fused to palp base.

Chaetiger 1 with fewer chaetae than succeeding chaetigers. Row of long capillaries on the notopodium (Figs 15, 16A). Postchaetal lamellae foliaceous and fused with branchia for most of its length on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Both lamellae bearing short serrations. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous throughout, partially fused with branchiae; external margin greatly enlarged, gradually reducing in size towards the posterior region (Fig 15). On middle segments, notopodial postchaetal lamellae develops a triangular ventral edge. Postchaetal lamellae fused with branchiae for most of its length on the anterior region and fused for half of its length on the posterior region. Postchaetal lamellae bearing a single serration on chaetiger 2.

Notopodial prechaetal lamellae reduced and rectangular on anterior chaetigers, becoming foliaceous with a pointed tip towards the posterior region (Fig 15). On the anterior region, notopodial and neuropodial prechaetal lamellae fused. Dorsal crests absent. Lateral organs present between postchaetal lamellae of notopodia and neuropodia, barely visible on anterior chaetigers due to overlapping of notopodial and neuropodial postchaetal lamellae and fused prechaetal lamellae.

Neuropodial postchaetal lamellae rounded on chaetigers 2–6, rectangular from chaetiger 7 onwards, gradually reducing in size towards the posterior region (Fig 15). On middle segments, neuropodial postchaetal lamellae develops a tapered dorsal projection. Postchaetal lamellae serrated on chaetiger 2. Neuropodial prechaetal lamellae well-developed and fan-shaped on anterior and middle chaetigers, gradually reducing in size towards the posterior region, present as a low flap on posterior chaetigers.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, anterior row of short, robust, curved, intensely granulated and widely bilimbate capillaries (Fig 16B) and posterior row of long (up to two times longer than anterior row), lightly granulated and narrowly unilimbate capillaries (Fig 16C). Above the notopodial rows, additional tuft of 2–3 long, wide, intensely granulated and distally unilimbate capillaries (Fig 16D). Notopodial chaetae up to two times longer than neuropodial chaetae. On neuropodia, chaetae from both rows of same morphology – lightly granulated and narrowly unilimbated capillaries, posterior row up to two times longer than anterior row (Fig 16E). Towards the posterior region, chaetae of both rami progressively become elongate, non-limbate, thinner, and less numerous (Fig 16F).

328

Hooks in neuropodia only, starting from chaetiger 23, up to eight per fascicle, accompanied by 1–8 short non-limbate capillaries. Hooks unidentate (Fig 16G). Secondary hood absent. Sabre chaetae from chaetiger 13, up to three per fascicle, intensely granulated throughout its length (Fig 16H).

Branchiae from chaetiger 1 onwards. Branchiae long, cirriform, with a pointed tip, fused to the notopodial postchaetal lamellae for most of its length on anterior chaetigers and for about half of its length on posterior chaetigers. Accessory branchiae present from chaetiger 33, on the dorsolateral side of body, behind notopodia base. Accessory branchiae appearing as a single digitiform projection, increasing in number on middle chaetigers, up to 6 digitiform projections; present until the last chaetiger of the fragment.

Pygidium of unknown morphology.

Methyl green staining pattern: Anterior and middle regions of the prostomium (except pointed tip), dorsal and lateral sides of the peristomium, palp sheaths, and margins of postchaetal lamellae intensely stained; margins of J-shaped metameric dorsal ciliated organs weakly stained.

Remarks. Dispio sp. 1 is readily distinguished from Dispio levisi sp. nov., D. magna and D. cf. brachychaeta in having serrations on anterior postchaetal lamellae. The species is similar to D. uncinata in having a long row of notochaetae on chaetiger 1 and up to eight neuropodial hooded hooks per fascicle, but species can be distinguished based on the prostomium shape – fusiform and anteriorly pointed in D. sp. 1 and bearing lateral projections in D. uncinata; starting chaetiger of neuropodial hooded hooks – from chaetiger 23 in D. sp. 1 and from chaetigers 25–27 in D. uncinata, starting chaetiger of sabre chaetae – from chaetiger 13 in D. sp. 1 and from chaetiger 2 in D. uncinata; degree of fusion between branchiae and notopodial postchaetal lamellae, fused most of its length on anterior chaetigers and for about half of its length on posterior chaetigers in D. sp. 1 and completely fused to the notopodial postchaetal lamellae on anterior chaetigers and with free tips on the posterior region in D. uncinata and starting chaetiger of accessory branchiae, from chaetiger 33 in in D. sp. 1 and from chaetigers 24–28 in D. uncinata.

The species is also similar to D. remanei in having serrations on anterior postchaetal lamellae and branchia fused with notopodial postchaetal lamellae for most of its length on anterior chaetigers, although D. remanei description (Friedrich, 1956) was very brief and contained few characters, leading Foster (1971) to synonymize the species with D. uncinata, which is not listed as a synonym by Read & Fauchald (2020), while Bolívar & Lana (1987) noted that the type material was lost. Currently the only known character to distinguish D. sp. 1 from D. remanei is the starting chaetiger of accessory branchiae, from chaetiger 33 in D. sp. 1 and from chaetiger 30 in D. remanei.

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Since a single incomplete specimen was collected in this work, it is uncertain if this record represents a new species or could represent a juvenile of D. remanei, as most characters of this species remain obscure, its type material was lost and the species ontogenetic and intraspecific variability remain completely unknown. Before additional material attributed to Dispio sp. 1 can be obtained, it is not possible to make a final taxonomic decision.

Habitat: Medium sand, 23 m depth.

Distribution: Southeastern Brazil (Espírito Santo state), South Atlantic Ocean.

Genus Microspio Mesnil, 1896

Type species: Spio mecznikowianus Claparède, 1869, designated by Söderström, 1920.

Synonym: Mesospio Gravier, 1911 (subjective synonym)

Diagnosis (modified from Blake et al. 2017): Prostomium rounded, bifid or bilobed anteriorly, frontal or lateral horns absent; eyes present or absent; occipital antenna present or absent. Nuchal organs typically as ciliated bands lateral to caruncle; metameric dorsal ciliated organs often present. Transverse ciliary bands present between bases of branchiae sometimes present.

Branchiae always from chaetiger 2, limited to anterior region of body or continuing to posterior end, basally fused to postchaetal notopodial lamellae. Ventral epidermal glands present or absent.

Notochaetae capillaries of various; notopodial spines present or absent; neurochaetae include capillaries and bi, -tri or multidentate hooded hooks. Sabre chaetae usually present. Pygidium bearing 2–4 anal cirri, usually with bacillary glands.

Microspio profunda Maciolek, 1990

(Figs 17–19F)

Material examined. Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 1035m (2 ind); Amb3 CAND7, 19°42'18,42" S 39°5'52,41" W, 1307m (1 ind); Amb4 A10, 21°11'4,9" S 38°28'2,78" W, 3006m (1 ind); Amb4 F6, 19°52'56,9" S 38°35'8,88" W, 1022m (1 ind); Amb4 F7, 20°4'8,18" S

330

38°31'27,32" W, 1302m (1 ind); Amb4 F8, 20°16'35,72" S 38°27'18,98" W, 1902m (1 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (1 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (2 ind); Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 1300m (6 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (3 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (2 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (20 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (7 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (1 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (1 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (1 ind); Amb11 B6, 20°36'1,61" S 39°51'39,15" W, 1003m (1 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (2 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (3 ind).

Comparative material examined. USNM 80480 (holotype); USNM 80481 (25 specimens; non- type).

Description: A small Microspio species, largest complete specimen 3.8 mm long, 0.9 mm wide at the widest point for 29 chaetigers. Body wide on the anterior region, tapered abruptly on the middle region and gradually tapering towards the pygidium. Body dorsoventrally flattened throughout. Pigmentation patterns absent. Body color dark yellow to whitish in alcohol (Fig 17).

Prostomium narrow and anteriorly bifid, with two knob-like tips usually directed ventrally, extending posteriorly to the posterior margin of chaetiger 1, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 2 (Fig 18A–B). Metameric dorsal ciliated organs (ciliated bands) present on anterior and middle chaetigers. Eyes absent. Occipital antenna absent. Peristomium well-developed, not clearly separated from and surrounding prostomium; lacking lateral wings (Figs 17, 18B). Grooved palps reaching up to chaetiger 9, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae triangular on notopodium and oval on neuropodium, both slightly shifted dorsally and smaller than lamellae on succeeding chaetigers (Fig 18C). Prechaetal lamellae absent.

Notopodial postchaetal lamellae rounded on the anterior region, digitiform on the middle region and cirriform on the posterior region (Figs 17, 18A–C). Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded on the anterior region, digitiform on the middle region and cirriform on the posterior region (Fig 18C). Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows, anterior row of short, bilimbate and granulated capillaries (Fig 19A) and posterior row of long (up to 1.2 times longer than anterior

331 row), non-limbate and non-granulated capillaries (Fig 19B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 19C). On last 5–9 chaetigers, notopodial capillaries exceedingly long and thin, needle-like (Figs 18D, 19D).

Hooks in neuropodia only, starting from chaetiger 9–13, up to five per fascicle (Fig 19E), accompanied by row of 2–10 short non-limbate capillaries and 1–4 inferior narrowly unilimbate and lightly granulated capillaries (Fig 19F). Hooks tridentate, with two secondary teeth arranged side by side above the main tooth. Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 2, present almost on the whole body, absent on last 3–7 chaetigers, completely free from notopodial postchaetal lamellae. Branchiae slightly tapered, long on anterior region – up to four times longer than notopodial lamellae, gradually reducing in length towards last branchial pair.

Pygidium bearing a pair of ventral cirri.

Oocytes from chaetiger 11, measuring up to 90 µm.

Methyl green staining pattern: Prostomium and branchiae weakly stained, ventral side of the anterior region, notopodial lamellae from chaetigers 1–6 and noto- and neuropodial lamellae from posteriormost chaetigers intensely stained.

Remarks. Microspio profunda was described by Maciolek (1990) based on material from northwestern Atlantic Ocean (Bahamas and the USA). The species is characterized by a wide body on the anterior region, anteriorly bifid prostomium, lack of eyes, presence of notochaetae on chaetiger 1, lamellae from chaetiger 1 slightly shifted dorsally and needle-like chaetae on posterior notopodia.

The species is similar to Microspio alexandrae sp. nov. and M. pigmentata (Reish, 1959) in having branchiae on most of the body (except a variable number of posterior chaetigers); branchiae long and slightly tapered; lack of sabre chaetae; partial overlap on the starting chaetiger of neuropodial hooded hooks – from chaetigers 9–13 in M. profunda, from chaetiger 8–12 in M. alexandrae sp. nov. and from chaetigers 11–12 in M. pigmentata and presence of posterior notopodial needle-like spines. These spines seem to be exclusive of Microspio profunda, M. alexandrae sp. nov. and M. pigmentata.

Specimens collected in this work agree completely with the original description by Maciolek (1990) in morphological and reproductive characters – as both the holotype and some of the specimens examined in this work had oocytes starting from chaetiger 11. The only character that could not be properly evaluated is the pygidium morphology, which was damaged in all specimens from the type series, presenting 1–3 cirri, while specimens examined in this work

332 presented a pair of ventral cirri. Although specimens from this work seem to belong to Microspio profunda, the species distribution is worth investigating, as no Microspio profunda specimens were found on deep sea samples from northeastern Brazil (Sergipe and Alagoas states) (pers. obs.), thus making its distribution disjunct.

These species can be separated based on prostomial shape – anteriorly bifid in Microspio profunda and anteriorly rounded in M. alexandrae sp. nov. and M. pigmentata; hook morphology – tridentate in Microspio profunda and multidentate in M. alexandrae sp. nov. (with six secondary teeth above main fang) and M. pigmentata (with seven secondary teeth above main fang); pygidium morphology – bearing two ventral cirri in M. profunda (1 – 3 cirri in Maciolek 1990), a pair of dorsal cirri and a pair of ventral lobes in M. alexandrae sp. nov. and two pairs of lobes (one pair dorsal and one pair ventral) in M. pigmentata and the presence of inferior capillaries accompanying the hooded hooks in M. profunda.

Specimens recorded in this work corresponds to the first report of Microspio profunda in Brazil.

Habitat: coarse silt to clay, 410–3006 m depth.

Distribution: Atlantic Ocean: Bahamas, USA, Brazil (Rio de Janeiro and Espírito Santo states).

Microspio alexandrae sp. nov.

(Figs 20–22E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 12 Jul to 18 Jul 2011 53m, MZUSP XXXX. Paratypes: Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 06 Jun to 17 Jul 2013, 53m, MZUSP XXXX (2 ind), ZUEC- POL XXXX (2 ind).

Additional material examined. Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (1 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (1 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (1 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (1 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (3 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (1 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (5 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (1 ind); Amb7 A1, 21°2'45,81" S 40°32'29,21" W, 25m (2 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (1 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (3 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (3 ind); Amb14 B3, 20°34'53,05" S

333

40°6'27,68" W, 49m (2 ind); Amb14 F1, 18°42'54,91" S 39°31'28,99" W, 25m (1 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (2 ind).

Description: A medium-sized Microspio species, largest complete specimen 4.5 mm long, 0.9 mm wide at the widest point for 44 chaetigers, holotype 4 mm long, 0.8 mm wide at the widest point for 40 chaetigers. Body wide and dorsoventrally flattened, tapering towards the pygidium. Light to dark brown pigmented spots present as an oval spot on anterior part of prostomium, in front of eyes; and between notopodia and neuropodia on chaetiger 1–3. Body color light yellow to whitish in alcohol (Fig 20).

Prostomium narrow, anteriorly rounded and slightly expanded laterally, extending posteriorly to the anterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the anterior margin of chaetiger 2. Prostomium separated from peristomium by a furrow. Parentheses-shaped (round bracket) metameric dorsal ciliated organs present on anterior chaetigers. One pair of faded eyes or eyes absent. Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings (Fig 21A–B) . Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and rounded with a pointed tip on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae rounded with a pointed tip throughout, gradually reducing in size towards the posterior region (Fig 20). Notopodial prechaetal lamellae rounded on chaetigers 4– 25, absent on remaining chaetigers.

Neuropodial postchaetal lamellae broadly rounded on chaetigers 2–13, gradually reducing in size towards the posterior region and absent on last 4–8 chaetigers (Fig 21C). Neuropodial prechaetal lamellae as a low flap on chaetigers 3–14, absent on remaining chaetigers.

Chaetae from notopodia and neuropodia organized in two rows of unilimbate capillaries. Capillaries non-granulated and striated on notopodia (Fig 22A) and lightly granulated and non- striated on neuropodia (Fig 22B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 22C). On last 8–12 chaetigers, notopodial capillaries exceedingly long and thin, needle-like (Fig 22D).

Hooks in neuropodia only, starting from chaetigers 8–12, up to 10 per fascicle, accompanied by 2–12 short non-limbate capillaries (Fig 21D). Inferior limbate capillaries absent. Hooks heptadentate, with one large median upper tooth immediately above main tooth, two pairs of teeth

334 arranged in two vertical rows and one very small uppermost unpaired tooth (Fig 22E). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 2, present almost on the whole body, absent on last 4–7 chaetigers, completely free from notopodial postchaetal lamellae. Branchiae slightly tapered, long on anterior region – up to four times longer than notopodial lamellae, not much reduced in length towards last branchial pair (Fig 21 A–D).

Pygidium bearing a pair of dorsal cirri and pair of ventral lobes.

Methyl green staining pattern: Prostomium and dorsal part of peristomium strongly stained; dorsal and ventral body surface, margins of both lamellae and branchiae weakly stained.

Remarks. Microspio alexandrae sp. nov. can be distinguished from all remaining Microspio species in having a slightly expanded prostomium, branchiae up to four times longer than and completely free from notopodial lamellae, neuropodial hooded hooks with six secondary hooks above main fang, lack of inferior capillaries in neuropodia and sabre chaetae and presence of needle-like chaetae on posterior chaetigers. The presence of needle-like chaetae is uncommon in Microspio, found only in M. profunda and M. pigmentata.

These species can be separated based on prostomial shape – anteriorly rounded and slightly expanded laterally in M. alexandrae sp. nov., anteriorly bifid in Microspio profunda and anteriorly rounded in M. pigmentata; hook morphology – with six secondary teeth above main fang in M. alexandrae sp. nov., tridentate in Microspio profunda and with seven secondary teeth above main fang in M. pigmentata; pygidium morphology – bearing a pair of dorsal cirri and a pair of ventral lobes in M. alexandrae sp. nov., two ventral cirri in M. profunda and two pairs of lobes (one pair dorsal and one pair ventral) in M. pigmentata, besides lack of inferior capillaries accompanying the hooded hooks, present in M. profunda (Maciolek 1990).

Among Brazilian species, M. alexandrae sp. nov. can be readily distinguished from the remaining species (except Microspio profunda) in having branchiae completely free from the notopodial postchaetal lamellae and multidentate hooded hooks with one large median upper tooth immediately above main tooth, two pairs of teeth arranged in two vertical rows and one very small uppermost unpaired tooth (Table 3). The species is most similar to M. profunda, as most species bear branchiae almost on the whole body (absent on last chaetigers), branchiae free from notopodial postchaetal lamellae, overlap on the starting chaetiger of neuropodial hooded hooks, lack of sabre chaetae and presence of needle-like chaetae on posterior chaetigers, as discussed in M. profunda (Maciolek, 1990).

The lack of both sabre chaetae and inferior capillaries in neuropodia is uncommon, present only in M. alexandrae sp. nov. and M. quadridentata sp. nov., although species can be distinguished

335 by branchiae morphology – free from and up to four times longer than notopodial lamellae in M. alexandrae sp. nov. and basally fused to and up to three times longer than notopodial lamellae in M. quadridentata sp. nov., as well as hook morphology – heptadentate in M. alexandrae sp. nov. and quadridentate in M. quadridentata sp. nov.

Etymology. The specific epithet, alexandrae, is a homage to Dr. Alexandra E. Rizzo, a fellow polychaete researcher and a friend.

Habitat: coarse sand to coarse silt, 25–153m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Microspio tridentata sp. nov.

(Figs 23–25E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 02 Dec 2011 to 02 Fev 2012, 26m, ZUEC-POL XXXX. Paratypes: Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 06 Jun to 17 Jul 2013,50m, MNRJP XXXX (1 ind), MZUSP XXXX (2 ind).

Additional material examined. Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (2 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (1 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (1 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (4 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (2 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (1 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (1 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (1 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (1 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (3 ind); Amb7 F2, 18°52'32,61" S 39°8'42,82" W, 40m (1 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (1 ind); Amb7 G1, 18°32'3,6" S 39°23'16,91" W, 25m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (1 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (2 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (1 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (3 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind).

Description: A small-sized Microspio species, largest complete specimen 2 mm long, 0.3 mm wide at the widest point for 31 chaetigers, holotype 1.5 mm long, 0.2 mm wide at the widest point for 28 chaetigers. Body dorsoventrally flattened, tapering towards the pygidium. Brownish

336 pigmented spot present as an oval spot on anterior margin of chaetiger 2, immediately behind nuchal organs. Body color whitish in alcohol (Fig 23).

Prostomium narrow and anteriorly bifid, with tips pointed forward, extending posteriorly to the anterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the anterior margin of chaetiger 2. Prostomium separated from peristomium by a furrow. Metameric dorsal ciliated organs not observed. Two pairs of eyes in trapezoidal arrangement (Fig 23). Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings (Fig 24A). Grooved palps reaching up to chaetiger 8, lost in most specimens.

Chaetiger 1 lacking notopodial chaetae. Neuropodium with only a few chaetae, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae digitiform on chaetiger 2, subtriangular on chaetiger 3 to chaetigers 9–12 (depending on specimen size) and abruptly reduced to a low flap afterwards (Figs 23, 24B). Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded on chaetiger 2 to chaetigers 8–9 (depending on specimen size) and abruptly reduced to a low flap afterwards. Neuropodial prechaetal lamellae absent throughout (Fig 24B).

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, long (up to 1.5 times longer than neuropodial capillaries), lightly granulated and narrowly unilimbate (limbation almost imperceptible) capillaries (Fig 25A); on neuropodia, short, intensely granulated and widely bilimbate capillaries (Fig 25B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 25C). Modified notopodial capillaries absent on posterior region.

Hooks in neuropodia only, starting from chaetigers 8–11, up to five per fascicle and accompanied by 2–5 short non-limbate capillaries (Fig 24C). Inferior limbate capillaries absent. Hooks tridentate, with secondary teeth arranged side by side above main tooth (Fig 25D). Secondary hood absent. Sabre chaetae from chaetigers 11–14, appearing as a gradual transformation of the inferior-most capillary chaeta from 2–3 previous chaetigers. Sabre chaetae non-limbate, granulated along the shaft (Fig 25E).

Branchiae from chaetiger 2, continuing to last chaetiger. Branchiae slightly tapered, short (up to 2.5 times longer than notopodial lamellae), basally fused to notopodial postchaetal lamellae and reducing in size towards last chaetiger.

Pygidium bearing a pair of dorsal cirri and pair of ventral lobes.

337

Oocytes from chaetiger 12, measuring up to 250 µm. Sperm plates from chaetiger 11.

Methyl green staining pattern: Peristomium, margins of palp grooves, margins of notopodial and neuropodial lamellae, and pygidium, all weakly stained.

Remarks. Among Brazilian species, Microspio tridentata sp. nov. is similar to M. profunda in having a narrowed and anteriorly bifid prostomium, overlap on the starting chaetiger of neuropodial hooded hooks – from chaetigers 8–11 in M. tridentata sp. nov. and from chaetigers 9–13 in M. profunda and same hook morphology – tridentate with pair of secondary teeth arranged side by side above main fang. However, species can be distinguished based on branchial morphology and distribution – basally fused to and up to 2.5 times longer than notopodial lamellae, continuing to last chaetiger in M. tridentata sp. nov. and free from and up to four times longer than notopodial lamellae, absent on some posterior chaetigers in M. profunda; lack of inferior chaetae in neuropodia and presence of sabre chaetae in M. tridentata sp. nov.; lack of needle-like chaetae in posterior notopodia in M. tridentata sp. nov. and pygidium morphology – bearing a pair of dorsal cirri and pair of ventral lobes in M. tridentata sp. nov. and a pair of ventral cirri in M. profunda.

A few Microspio species also possess tridentate hooded hooks, such as Microspio microcera (Dorsey, 1977), described to California (Pacific Coast of the USA), M. tetrabranchia Maciolek, 1990, from the Atlantic Coast of the USA, M. granulata Blake & Kudenov, 1978, from the Pacific Coast of Australia (Maciolek, 1990). However, these species can be distinguished from M. tridentata sp. nov. by the arrangement of secondary teeth above the main fang – arranged as a vertical row of unpaired teeth in all aforementioned species and arranged side by side in M. tridentata sp. nov. Microspio minuta (Hartmann-Schröder, 1962) also possess tridentate hooks with secondary teeth arranged side by side, but will be addressed latter on, due to other similarities with M. tridentata sp. nov.

Some Spio species, such as Spio limnicola Verrill, 1879, S. picta Zachs, 1933, S. pettiboneae Foster, 1971, S. blakei Maciolek, 1990 and S. thulini Maciolek, 1990 also possess tridentate hooded hooks, but Spio species can be readily separated from Microspio species based on the starting chaetiger of branchiae – chaetiger 1 in Spio and chaetiger 2 in Microspio.

The presence of sabre chaetae is somewhat uncommon among Microspio, present only in M. tetrabranchia Maciolek, 1990 from the Atlantic Coast of the USA, M. kussakini Chlebovitsch, 1959, from Kuril Islands, Pacific Coast of Russia. Microspio tridentata sp. nov. can be readily distinguished from M. tetrabranchia by branchial distribution, present on the whole body in M. tridentata sp. nov. and only four pairs in M. tetrabranchia (Maciolek, 1990). As for M. kussakini, species can be distinguished by the prostomium morphology – anteriorly bifid in M. tridentata sp. nov. and anteriorly rounded, with short lateral projections in M. kussakini; lack of notopodial

338 chaetae in chaetiger 1 in M. tridentata sp. nov.; starting chaetiger of neuropodial hooded hooks – from chaetigers 8–11 in M. tridentata sp. nov. and from chaetigers 18–23 in M. kussakini; hook morphology – tridentate in M. tridentata sp. nov. and bidentate in M. kussakini and starting chaetigers of sabre chaetae, from chaetigers 11–14 in M. tridentata sp. nov. and from chaetigers 18–23 in M. kussakini. Microspio minuta also bears sabre chaetae (see below).

Microspio tridentata sp. nov. is most similar to Microspio minuta, described to Northern Chile and latter recorded in Galápagos Islands, Ecuador (Blake, 1983), as both species possess an anteriorly bifid prostomium, lack of notopodial chaetae in chaetiger 1, notopodial postchaetal lamellae subtriangular and neuropodial lamellae rounded in anterior chaetigers; overlap on the starting chaetiger of neuropodial hooded hooks – from chaetigers 8–11 in M. tridentata sp. nov. and from chaetiger 9 in M. minuta, same hooks morphology, lack of inferior capillaries in neuropodia and presence of sabre chaetae (Hartmann-Schröder 1962; Blake 1983). Despite great morphological similarity, species can be distinguished by the presence of notopodial postchaetal lamellae on chaetiger 1 – absent in M. minuta; branchiae morphology – basally fused to notopodial postchaetal lamellae in M. tridentata sp. nov. and free from notopodial postchaetal lamellae in M. minuta; branchiae length – up to 2.5 times longer than notopodial postchaetal lamellae in M. tridentata sp. nov. and up to twice as long as notopodial postchaetal lamellae in M. minuta and starting chaetiger of sabre chaetae – from chaetigers 11–14, appearing as a gradual transformation of the inferior-most capillary chaeta and abruptly appearing on chaetiger 9 in M. minuta.

Etymology. The specific epithet, tridentata, refers to the morphology of neuropodial hooded hooks, which are tridentate.

Habitat: coarse sand to very fine sand, 25–143 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Microspio quadridentata sp. nov.

(Figs 26–28D)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb12 D4,19°45'53,43" S 39°30'25,97" W, 06 Jun to 17 Jul 2013, 143m, MZUSP XXXX. Paratype: Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 06 Jun to 17 Jul 2013, 143m, MNRJP XXXX (1 ind).

Additional material examined. Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (1 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (1 ind).

339

Description: A medium-sized Microspio species, largest specimen 1.2 mm long, 0.2 mm wide at the widest point for 24 chaetigers (incomplete), holotype 1 mm long, 0.18 mm wide at the widest point for 17 chaetigers (incomplete). Body dorsoventrally flattened, tapering towards the posterior region. Pigmentation patterns absent. Body color whitish in alcohol (Fig 26).

Prostomium narrow and anteriorly rounded, extending posteriorly to the posterior margin of chaetiger 1, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 1. Prostomium separated from peristomium by a furrow. Metameric dorsal ciliated organs not observed. Two pairs of eyes in trapezoidal arrangement. Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings (Fig 26, 27A–B). Grooved palps reaching up to chaetiger 15, lost in most specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded with a pointed tip on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae rounded with a pointed tip on chaetigers 2–10, largest on chaetigers 7–9 and reduced to a low flap from chaetiger 11 onwards (Fig 26, 27A). Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded on chaetigers 2–9, largest on chaetigers 6 and 7 and reduced to a low flap from chaetiger 10 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate capillaries, lightly granulated on notopodia (Fig 28A) and non-granulated on neuropodia (Fig 28B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 28C). Modified notopodial capillaries absent on posterior region.

Hooks in neuropodia only, starting from chaetiger 10, up to seven per fascicle, accompanied by 7–12 short non-limbate capillaries. Inferior limbate capillaries absent. Hooks quadridentate, with three single secondary teeth arranged vertically above main fang (Fig 28D). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 2, extending through entire fragment, basally fused to notopodial lamellae. Branchiae slightly tapered, longest pair up to three times longer than notopodial postchaetal lamellae, not reducing in size towards end of fragment (chaetiger 24) (Fig 27B).

Pygidium of unknown morphology.

Oocytes from chaetiger 12, measuring up to 130 µm.

340

Methyl green staining pattern: Prostomium and peristomium strongly stained; branchiae and margins of lamellae weakly stained.

Remarks. Microspio quadridentata sp. nov. is similar to M. sp. 2 in having an anteriorly rounded prostomium, branchiae basally fused to notopodial postchaetal lamellae and lack of sabre chaetae, although species can be distinguished by branchiae length – up to three times longer than notopodial postchaetal lamellae in M. quadridentata sp. nov. and up to twice as long as notopodial postchaetal lamellae in M. sp. 2; starting chaetiger of neuropodial hooded hooks – chaetiger 10 in M. quadridentata sp. nov. and chaetiger 11 in M. sp. 2; hook morphology – quadridentate in M. quadridentata sp. nov. and tridentate in M. sp. 2 and lack of inferior capillaries in neuropodia in M. quadridentata sp. nov.

Unfortunately, all collected specimens were anterior fragments, hence characters such as branchial distribution, presence of needle-like notopodial chaetae on posterior chaetigers and pygidium morphology remain unknown. However, the presence of quadridentate hooded hooks, with three single secondary teeth arranged vertically above main fang, is not known in any other Microspio species, being an exclusive character of M. quadridentata sp. nov. A redescription of the species will likely be published if complete specimens are found.

Etymology. The specific epithet, quadridentata, refers to the morphology of neuropodial hooded hooks, which are quadridentate.

Habitat: medium sand to very fine sand, 50–171 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Microspio sp. 1

(Figs 29–30E)

Material examined. Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (1 ind).

Description: A medium-sized Microspio species, specimen 2.8 mm long, 0.9 mm wide at the widest point for 15 chaetigers (incomplete). Fragment dorsoventrally flattened, tapering towards the posterior region. Yellow pigmented spots present as an oval spot on anterior part of prostomium, and on ventral side of peristomium, close to chaetiger 1. Body color whitish in alcohol (Fig 29).

341

Prostomium narrow, widening towards rounded anterior margin, with slightly expanded antero- lateral margins. Prostomium extending posteriorly to the anterior margin of chaetiger 2, flanked by C-shaped nuchal organs extending up to the anterior margin of chaetiger 2. Metameric dorsal ciliated organs not observed. Eyes and occipital antenna absent. Peristomium moderately developed, lacking lateral wings (Fig 29). Peristomium separated from prostomium by a furrow, surrounding prostomium for most of its length. Palps lost.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and fan-shaped on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae foliaceous on chaetigers 2–12, largest on chaetigers 4–10 and remarkably reduced in size from chaetiger 13 onwards (Fig 29). Notopodial prechaetal lamellae well-developed, digitiform on chaetigers 4–15 (end of fragment), largest on chaetigers 10–11.

Neuropodial postchaetal lamellae fan-shaped on chaetiger 2–11, remarkably reduced in size from chaetiger 12 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, narrowly unilimbate and striated capillaries (Fig 30A); on neuropodia anterior row of short, bilimbate and granulated capillaries (Fig 30B) and posterior row of long (up to 1.2 times longer than anterior row), narrowly unilimbate, thin and non-granulated capillaries (Fig 30C). Chaetae of middle and posterior regions unknown.

Hooks in neuropodia only (Fig 30D), starting from chaetiger 11, up to 10 per fascicle and accompanied by 8–12 short non-limbate capillaries and 1–2 inferior narrowly unilimbate and lightly granulated capillaries (Fig 30E). Hooks multidentate, with four secondary teeth arranged in two rows above main tooth. Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 2, extending through entire fragment, basally fused to notopodial postchaetal lamellae. Branchiae slightly tapered, longest pair up to four times longer than notopodial postchaetal lamellae, not reducing in size towards end of fragment (chaetiger 15).

Pygidium of unknown morphology.

Methyl green staining pattern: Prostomium intensely stained, margins of lamellae and branchiae weakly stained.

Remarks. Microspio sp. 1 is most similar to M. alexandrae sp. nov. in having a slightly expanded prostomium, branchiae um to four times longer than notopodial prechaetal lamellae, partial overlap on the starting chaetiger of neuropodial hooded hooks – from chaetiger 11 in Microspio sp. 1 and from chaetigers 8–12 in M. alexandrae sp. nov. and lack of sabre chaetae. However,

342 species can be distinguished by branchial morphology – basally fused to notopodial postchaetal lamellae in M. sp. 1 and free from notopodial lamellae in M. alexandrae sp. nov., morphology of neuropodial hooded hooks – bearing eight secondary teeth above main fang in M. sp. 1 and six secondary teeth above main fang in M. alexandrae sp. nov. and presence of inferior capillary chaetae in neuropodia in M. sp. 1.

Microspio sp. 1 is similar to M. tridentata sp. nov., M. sp. 2 in having branchiae basally fused to notopodial postchaetal lamellae, although species can be separated based on branchial length – up to four times longer than notopodial postchaetal lamellae in M. sp. 1., up to 2.5 times longer than notopodial postchaetal lamellae in M. tridentata sp. nov., up to twice as long as notopodial postchaetal lamellae in M. sp. 2 and up to three times longer than notopodial postchaetal lamellae in M. quadridentata sp. nov.

Microspio sp. 1 corresponds to a new species, although, since only one incomplete specimen was collected, the morphology of the species is poorly known, since characters such as branchiae distribution, presence of needle-like chaetae in posterior notopodia and pygidium morphology are unknown. Due to these shortcomings, it was decided not to formally describe the species until additional material becomes available.

Habitat: fine sand, 40m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Microspio sp. 2

(Figs 31–32E)

Material examined. Amb11 F10, 20°46'17,79" S 38°17'16,01" W, 3002m (1 ind).

Description: A small-sized Microspio species, complete specimen 3.8 mm long, 0.4 mm wide at the widest point for 24 chaetigers. Body dorsoventrally flattened, wide on the anterior region, tapered abruptly on chaetiger 11 and gradually tapering towards the pygidium. Body dorsoventrally flattened throughout. Pigmentation patterns absent. Body color light yellow in alcohol (Fig 31).

343 not observed. Eyes absent. Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings. Palps lost.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and rounded on neuropodium, not significantly smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae digitiform on chaetigers 2–6 and reduced to a low flap from chaetiger 7 onwards (Fig 31). Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded on chaetigers 2–7 reduced to a low flap from chaetiger 8 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate capillaries, lightly granulated on notopodia (Fig 32A) and intensely granulated on neuropodia (Fig 32B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 32D). Modified notopodial capillaries absent on posterior region.

Hooks in neuropodia only, starting from chaetiger 11, up to 13 per fascicle and accompanied by 7–11 short non-limbate capillaries and 1–4 long, non-limbate capillaries and 1–4 lightly granulated capillaries on inferior-most position (Fig 32C); short companion chaetae absent from chaetiger 16 onwards. Hooks multidentate, with secondary teeth arranged in two rows above main tooth (Fig 32E). Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 2, present almost on the whole body, absent on last two chaetigers, basally fused to notopodial lamellae. Branchiae slightly tapered, short (up to twice as long as notopodial lamellae), gradually reducing in length towards last branchial pair.

Pygidium bearing a pair of dorsal cirri and a pair of ventral lobes.

Oocytes from chaetiger 9, measuring up to 150 µm.

Methyl green staining pattern: Prostomium and peristomium strongly stained; branchiae, lamellae and ventral part of first 6 chaetigers weakly stained.

Remarks. Microspio sp. 2 is similar to Microspio quadridentata sp. nov. in having a narrow and anteriorly rounded prostomium, branchiae basally fused to notopodial postchaetal lamellae and lack of sabre. However, species can be distinguished based on branchiae length – up to twice as long as notopodial postchaetal lamellae in Microspio sp. 2 and up to three times longer than notopodial postchaetal lamellae in Microspio quadridentata sp. nov.; starting chaetiger of neuropodial hooded hooks – from chaetiger 11 in Microspio sp. 2 and – from chaetiger 10 in M. quadridentata sp. nov.; hook morphology – multidentate, in M. sp. 2 and quadridentate in M. quadridentata sp. nov.

344

The species is also similar to M. profunda and M. sp. 1 in having inferior capillaries in neuropodia, although species can be distinguished by prostomium shape – anteriorly rounded in M. sp. 2, anteriorly bifid in M. profunda and with slightly expanded antero-lateral margins in M. sp. 1 and branchiae length – up to twice as long as notopodial postchaetal lamellae in M. sp. 2 and up to four times longer than notopodial postchaetal lamellae in M. profunda and M. sp. 1 (Maciolek 1990).

Microspio sp. 2 is also similar to M. alexandrae sp. nov. and M. tridentata sp. nov. as all three species possess a pygidium with a pair of dorsal cirri and pair of ventral lobes, although species can be distinguished by prostomium shape – anteriorly round in M. sp. 2, slightly expanded laterally in M. alexandrae sp. nov. and anteriorly bifid in M. tridentata sp. nov.; branchiae length – up to twice as long as notopodial postchaetal lamellae in M. sp. 2, up to four times longer than notopodial postchaetal lamellae in M. alexandrae sp. nov and up to 2.5 times longer than notopodial postchaetal lamellae in M. tridentata sp. nov., besides the presence of inferior capillaries in neuropodia in M. sp. 2.

Given the characters discussed here, Microspio sp. 2 most likely represents a new species, however, since a single complete species was collected during the entire project, it was decided not to formally describe a new species until additional material becomes available, as the ontogenetic and intraspecific variation of this species are currently unknown.

Habitat: fine silt, 3002 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and Espírito Santo states), Atlantic Ocean.

Genus Spio Fabricius, 1785

Type species: Nereis filicornis Müller, 1776, accepted as Spio filicornis (Müller, 1776) (type by superseded original combination)

Synonym list: Paraspio Czerniavsky, 1881 (subjective synonym)

Euspio McIntosh, 1915 (subjective synonym)

Diagnosis (from Blake et al. 2017): Prostomium anteriorly rounded, truncate, or slightly incised, lacking frontal or lateral horns; eyespots present or absent; occipital antenna absent, but posterior

345 portion of prostomium may be raised or inflated. Nuchal organ with short median and long lateral ciliary bands, extending to chaetiger 2 or 3. Metameric dorsal ciliated organs usually present; transverse dorsal ciliated bands present. Branchiae present from chaetiger 1, continuing almost throughout the body, completely separate from or basally fused with notopodial lamella, often reduced in size on chaetiger 1. Ventral epidermal glands usually present in anterior and middle chaetigers. Notochaetae and anterior neurochaetae all capillaries; capillaries, hooded hooks and inferior sabre chaetae on middle and posterior neuropodia. Pygidium with four anal cirri.

Spio anaclaudiae sp. nov.

(Figs 33–35E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 02 Dec 2011 to 02 Feb 2011, 50m, MZUSP XXXX. Paratypes: Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 02 Dec 2011 to 02 Feb 2011, 52m (4 ind); MNRJP XXXX (2 ind), ZUEC-POL XXXX (2 ind).

Additional material examined. Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (1 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (2 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (1 ind); Amb7 B2, 20°34'45,78" S 40°11'30,74" W, 41m (2 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (1 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (1 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (4 ind); Amb7 G1, 18°32'3,6" S 39°23'16,91" W, 25m (2 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (1 ind); Amb13 G2, 18°36'32,45" S 39°9'32,83" W, 40m (2 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (1 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (4 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (1 ind).

Description: A medium-sized Spio species, largest complete specimen 2.1 mm long, 0.2 mm wide at the widest point for 37 chaetigers, holotype 2 mm long, 0.2 mm wide at the widest point for 34 chaetigers. Body cylindrical on anterior and middle regions and slightly dorsoventrally flattened on the posterior region, tapering towards the pygidium. Pigmentation patterns present as a transversal band of dark pigment in front of eyes and as brownish middorsal melanophores form

346 chaetiger 3 to about chaetiger 18 (most intense on chaetigers 4– 14). Body color light yellow in alcohol (Fig 33).

Prostomium narrow, slightly bifid anteriorly with tips pointed forward, extending posteriorly to the posterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 2. Prostomium separated from peristomium by a furrow (Fig 33, 34A–B). Metameric dorsal ciliated organs not observed. Two pairs of eyes in trapezoidal arrangement. Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings. Palps lost in all specimens.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and rounded on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Notopodial postchaetal lamellae rounded from chaetiger 2 to chaetigers 9–14 (Fig 33, 34B–C) (depending on specimen size), reduced to a low flap afterwards. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae fan-shaped from chaetiger 2 to chaetigers 9–14 (depending on specimen size), reduced to a low flap afterwards (Fig 34C). Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows. On notopodia, long (up to 1.5 times longer than neuropodial capillaries), lightly granulated and narrowly unilimbate capillaries (Fig 35A); on neuropodia, short, intensely granulated and widely bilimbate capillaries (Fig 35B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 35C). Modified notopodial capillaries absent on posterior region.

Hooks in neuropodia only, starting from chaetigers 10–11, up to seven per fascicle and accompanied by 3–5 short non-limbate capillaries. Inferior limbate capillaries absent. Hooks tridentate, with secondary teeth in a vertical line above main tooth (Fig 35D). Secondary hood absent. Sabre chaetae from chaetigers 19–21, appearing as a gradual transformation of the inferior-most capillary chaeta from two previous chaetigers. Sabre chaetae thin, non-limbate and lightly granulated (Fig 35E).

Branchiae from chaetiger 1, present on almost the whole body, absent on last 8–13 chaetigers. Branchiae slightly tapered, long (up to three times longer than notopodial lamellae), free from notopodial lamellae and reducing in size towards last chaetiger (Fig 34B, D).

Pygidium bearing two pairs of cirri, dorsal and ventral pairs of the same size and morphology (Fig 34D).

347

Oocytes from chaetiger 13, measuring up to 180 µm.

Juvenile morphology: In juveniles, the prostomium frequently is rounded anteriorly, instead of slightly bifid. In most juvenile specimens, branchiae start in chaetiger 2 instead of chaetiger 1 (see Remarks below).

Methyl green staining pattern: Prostomium and peristomium strongly stained, lamellae and branchiae weakly stained.

Remarks. Spio anaclaudiae sp. nov. is similar to Spio filicornis in having a slight median incision (prostomium bifid), similar starting chaetiger of neuropodial hooded hooks – from chaetigers 10– 11 in S. anaclaudiae sp. nov. and chaetiger 11 in S. filicornis, branchiae morphology – slightly tapered and pygidial morphology – bearing two pairs of cirri of the same size. However, species can be distinguished based on branchiae morphology – slightly longer in S. anaclaudiae sp. nov. and free from notopodial postchaetal lamellae on all chaetigers and branchiae basally fused to notopodial lamellae on anterior chaetigers in S. filicornis, hook morphology – tridentate in S. anaclaudiae sp. nov. and bidentate in S. filicornis, lack of inferior capillaries in neuropodia in S. anaclaudiae sp. nov., which are present in S. filicornis and starting chaetiger of sabre chaetae – from chaetigers 19–21 in S. anaclaudiae sp. nov. and from chaetiger 18 in S. filicornis (Meiβner et al., 2011).

Among Brazilian Spio species, S. capixaba sp. nov., S. sp. 1, S. quadrisetosa and S. pettiboneae also present tridentate hooded hooks, although in S. capixaba sp. nov. and S. quadrisetosa possess secondary teeth organized in a vertical row, instead of secondary teeth arranged side by side, as in S. sp. 1 and S. pettiboneae. Spio anaclaudiae sp. nov. can be distinguished from S. sp. 1 and S. pettiboneae in the prostomial shaped – anteriorly bifid in S. anaclaudiae sp. nov. and anteriorly rounded with constriction in S. sp. 1 and S. pettiboneae; branchiae morphology – up to three times longer than and free from notopodial lamellae in S. anaclaudiae sp. nov., up to three times longer than and free from notopodial lamellae in S. sp. 1 and up to three times longer than and fused to notopodial lamellae in anterior chaetigers in S. pettiboneae; starting chaetiger of neuropodial hooded hooks – from chaetigers 10–11 in S. anaclaudiae A sp. nov., chaetiger 18 in S. sp. 1 and chaetigers 12–16 in S. pettiboneae and starting chaetiger of sabre chaetae – from chaetigers 19– 21 in S. anaclaudiae A sp. nov., chaetiger 18 in S. sp. 1 and chaetigers 10–15 in S. pettiboneae (Foster, 1971; Maciolek, 1990).

Etymology. The specific epithet, anaclaudiae, is a homage to Dr. Ana Cláudia dos Santos Brasil, a fellow polychaete researcher who have been accompanying the first author since his undergraduate years.

Habitat: coarse sand to very fine sand, 25–53 m depth.

348

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Spio capixaba sp. nov.

(Figs 36–38E)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 02 Dec 2011 to 02 Feb 2011, 26m, MZUSP XXXX. Paratypes: Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 11 Dec to 19 Dec, 32m, MNRJP XXXX (2 ind), ZUEC-POL XXXX (1 ind).

Additional material examined. Amb1 Foz4, 19°37'21,13" S 39°45'6,41" W, 15m (2 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (1 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (1 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (1 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (2 ind).

Description: A medium-sized Spio species, largest complete specimen 2.5 mm long, 0.3 mm wide at the widest point for 33 chaetigers, holotype 2 mm long, 0.25 mm wide at the widest point for 29 chaetigers. Body dorsoventrally flattened, tapering towards the pygidium. Pigmentation patterns present as a transversal band of dark pigment in front of eyes and as brownish paired ventral melanophores on anterior edge of chaetiger 2 to chaetigers 8–10 (most intense on chaetigers 3– 6). Body color light yellow to whitish in alcohol (Fig 36).

Prostomium narrow, rounded anteriorly and bearing short lateral projections, extending posteriorly to the posterior margin of chaetiger 2, flanked by J-shaped nuchal organs extending up to the posterior margin of chaetiger 2. Prostomium separated from peristomium by a furrow (Figs 36, 37A–B, D). Metameric dorsal ciliated organs extending at least to chaetiger 12 (Fig 37C). Two pairs of eyes in trapezoidal arrangement or eyes absent. Occipital antenna absent; posterior part of prostomium inflated. Peristomium moderately developed, surrounding prostomium; lacking lateral wings. Palps lost in all specimens.

Notopodial postchaetal lamellae rounded with a pointed tip from chaetiger 2 to chaetigers 7–9 (depending on specimen size), rounded from chaetigers 8–10 to chaetigers 16–21 (depending on

349 specimen size), and reduced to a low flap afterwards (Figs 36, 37A). Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded from chaetiger 2 to chaetigers 17–20 (depending on specimen size), reduced to a low flap afterwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows, anterior row of short, intensely granulated and widely bilimbate capillaries (Fig 38A) and posterior row of long (up to 2.5 times longer than anterior row on notopodia and up to 1.5 times longer than anterior row on neuropodia), non-granulated and unilimbate capillaries (Fig 38B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 38C). Modified notopodial capillaries absent on posterior region.

Hooks in neuropodia only (Fig 38D), starting from chaetiger 11, up to eight per fascicle and accompanied by 5–8 short non-limbate capillaries and 1–2 inferior narrowly unilimbate and lightly granulated capillaries (Fig 38E). Hooks tridentate, with secondary teeth in a vertical line above main tooth. Secondary hood absent. Sabre chaetae absent throughout.

Branchiae from chaetiger 1, present almost on the whole body, absent on last 10–12 chaetigers. Branchiae slightly tapered, long (up to three times longer than notopodial lamellae), free from notopodial lamellae (Figs 37A–B, D) and reducing in size towards last chaetiger.

Pygidium bearing two pairs of cirri, dorsal pair longer and thinner than ventral pair.

Oocytes from chaetiger 13, measuring up to 130 µm.

Juvenile morphology: In juveniles, the prostomium is rounded anteriorly, lacking lateral projections. In most juvenile specimens, branchiae start in chaetigers 2–3 instead of chaetiger 1 (see Remarks below).

Methyl green staining pattern: Prostomium (except for inflated part of prostomium) and peristomium strongly stained, margins of lamellae and branchiae weakly stained.

Remarks. Spio capixaba sp. nov. is unique among Brazilian Spio species in having a prostomium bearing two short lateral projections.

The species is similar to S. quadrisetosa in having neuropodial hooded hooks from chaetiger 11 with secondary teeth arranged on a vertical line and lack of inferior capillaries in neuropodia, although species can be distinguished by prostomial shape, branchial morphology – slightly longer and free from notopodial lamellae in Spio capixaba sp. nov. and shorter and fused to notopodial lamellae in S. quadrisetosa and lack of sabre chaetae in Spio capixaba sp. nov., present from chaetiger 12 in S. quadrisetosa (Blake 1983).

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Spio capixaba sp. nov. is most similar to S. anaclaudiae sp. nov. in having branchiae up to three times longer than and completely free from notopodial lamellae, similar starting chaetiger of neuropodial hooded hooks – chaetiger 11 in S. capixaba sp. nov. and chaetigers 10–11 in S. anaclaudiae sp. nov., tridentate hooded hooks and lack of inferior capillaries in neuropodia, although, besides the prostomial shape, species can be distinguished by morphology of the hooks – with secondary teeth arranged on a vertical line in S. capixaba sp. nov. and secondary teeth arranged side by side in S. anaclaudiae sp. nov.; lack of sabre chaetae in S. capixaba sp. nov., present from chaetigers 19–21 in S. anaclaudiae sp. nov. and pygidial morphology – bearing two pairs of cirri, dorsal pair longer in S. capixaba sp. nov. and dorsal and ventral pairs of the same length in S. anaclaudiae sp. nov.

Etymology. The species epithet, “capixaba” is the common denomination of natives from the state of Espírito Santo, southeastern Brazil, where the specimens were collected.

Habitat: medium sand to medium silt, 15–144 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Spio sp. 1

(Figs 39–40E)

Material examined. Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (1 ind).

Description: A medium-sized Spio species, specimen 2.3 mm long, 0.35 mm wide at the widest point for 24 chaetigers (incomplete). Body cylindrical on anterior region and dorsoventrally flattened afterwards, tapering towards the posterior region. Pigmentation patterns absent. Body color light yellow in alcohol (Fig 39).

Prostomium narrow, anteriorly rounded and constricted laterally in front of the anterior pair of eyes, extending posteriorly to the posterior margin of chaetiger 2, flanked by U-shaped nuchal organs extending up to the posterior margin of chaetiger 2. Prostomium separated from peristomium by a furrow (Fig 39). Metameric dorsal ciliated organs not observed. Two pairs of eyes in trapezoidal arrangement. Occipital antenna absent. Peristomium moderately developed, surrounding prostomium; lacking lateral wings. Palps lost.

Chaetiger 1 with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and foliaceous on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

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Notopodial postchaetal lamellae well-developed, foliaceous on chaetigers 2–12, largest on chaetigers 5–11 and absent from chaetiger 13 onwards (Fig 39). Notopodial prechaetal lamellae absent throughout whole fragment.

Neuropodial postchaetal lamellae foliaceous throughout whole fragment, reducing in size from chaetiger 9 onwards. Neuropodial prechaetal lamellae absent throughout whole fragment.

Chaetae from notopodia and neuropodia organized in two rows of narrowly unilimbate and non- granulated capillaries. Chaetae from both rows of equal length, although notopodial chaetae (Fig 40A) are up to 1.5 times longer than neuropodial chaetae (Fig 40B). Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 40C). Modified notopodial capillaries absent throughout whole fragment.

Hooks in neuropodia only, starting from chaetiger 18, up to four per fascicle and accompanied by 3–5 short non-limbate capillaries. Inferior limbate capillaries absent. Hooks tridentate, with secondary teeth arranged side by side above main tooth (Fig 40D). Secondary hood absent. Sabre chaetae from chaetiger 18. Sabre chaetae non-limbate and lightly granulated (Fig 40E).

Branchiae from chaetiger 1, extending through entire fragment, free from notopodial lamellae. Branchiae slightly tapered, longest on chaetiger 1 (up to four times longer than notopodial lamellae), not reducing in size towards end of fragment (chaetiger 20).

Pygidium of unknown morphology.

Methyl green staining pattern: Prostomium intensely stained, peristomium, margins of both lamellae and of branchiae weakly stained.

Remarks. Among species reported to Brazil, Spio sp. 1 is similar to S. pettiboneae in having a narrow and anteriorly rounded prostomium bearing a constriction, long branchiae – up to four times longer than notopodial lamellae in Spio sp. 1 and up to five times longer than notopodial lamellae in S. pettiboneae, tridentate hooded hooks with secondary teeth arranged side by side and lack of inferior capillaries in neuropodia, although species can be distinguished due to branchiae morphology – free from notopodial lamellae in Spio sp. 1 and basally fused to notopodial lamellae on anterior chaetigers in S. pettiboneae; starting chaetiger of neuropodial hooded hooks – from chaetiger 18 in Spio sp. 1 and chaetigers 12–16 in S. pettiboneae and starting chaetiger of sabre chaetae – chaetiger 18 in Spio sp. 1 and chaetigers 10–15 in S. pettiboneae (Foster, 1971; Maciolek, 1990).

Given differences from the remaining Brazilian species, it is believed that Spio sp. 1 represents a new species. However, since a single incomplete species was collected, it was decided not to formally described a new species until additional material becomes available.

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Habitat: fine sand, 41 m depth.

Distribution: Southeastern Brazil (Espírito Santo and Campos basins), Atlantic Ocean.

Identification key to Spio and Microspio species recorded in this work.

1. Branchiae from chaetiger 1……………………………………………...………………….2 - Branchiae from chaetiger 2………………………………………………..…………………4 2. Neuropodial hooded hooks from chaetiger 18…………………………………….Spio sp. 1 - Neuropodial hooded hooks from chaetigers 10……………………………………………3 3. Sabre chaetae from chaetigers 19–21, appearing as a gradual transformation of inferior- most capillaries………………………………….………………….Spio anaclaudiae sp. nov. - Sabre chaetae absent………………………………………………….Spio capixaba sp. nov. 4. Prostomium anteriorly bifid………………………………………...………………………5 - Prostomium anteriorly rounded…………………………………...…………………………6 5. Needle-like spines present on posterior notopodia…………………Microspio cf. profunda - Needle-like spines absent on posterior notopodia…….……….Microspio tridentata sp. nov. 6. Neuropodial hooded hooks quadridentate...... Microspio quadridentata sp. nov. - Neuropodial hooded hooks multidentate...... 7 7. Branchiae not basally fused to notopodial lamellae, neuropodial hooded hooks from chaetigers 8–12……………………………………..………….Microspio alexandrae sp. nov. Branchiae basally fused to notopodial lamellae, neuropodial hooded hooks consistently from chaetiger 11……………………………………………………………………………………8 8. Prostomium with slightly expanded antero-lateral margins, branchiae up to four times longer than notopodial lamellae………………………………………………..Microspio sp. 1 - Prostomium short and anteriorly rounded, branchiae up to two times longer than notopodial lamellae………………………………………………………………………...Microspio sp. 2

Genus Malacoceros Quatrefages, 1843

Type species: Spio vulgaris Johnson, 1827, accepted as Malacoceros vulgaris (Johnson, 1827)

(type by subsequent designation)

Synonym list: Colobranchus Schmarda, 1861 (subjective synonym)

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Uncinia Quatrefages, 1866 (subjective synonym)

Diagnosis (from Blake et al. 2017): Prostomium broad anteriorly, triangular, T- or bell-shaped; frontal horns typically present; occipital antenna absent. eyespots present in pairs, irregularly arranged or eyespots absent. Caruncle entire, trilobed or buttonlike. Nuchal organs as two small ciliated grooves posterolateral to the caruncle. Palps ventrally grooved. Peristomium reduced to moderately developed. Branchiae from chaetiger 1 to end or near end of body; free from or basally fused to notopodial lamellae. Transverse ciliated bands across the dorsum present. Notochaetae all capillaries, neurochaetae includes capillaries, scalpel chaetae and uni-, bi-, tri- or quadridentate hooded hooks. Sabre chaetae present. Pygidium with 2–30 anal cirri or with two anal cirri and a rounded or spatuliform dorsal lobe.

Malacoceros aureus sp. nov.

(Figs 41–43F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 02 Dec 2011 to 02 Feb 2012, 39m, MZUSP XXXX. Paratypes: Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 02 Dec 2011 to 02 Feb 2012, 39m, MNRJP XXXX (2 ind), ZUEC-POL XXXX (1 ind).

Additional material examined. Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (2 ind); Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (1 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (2 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (1 ind); Amb7 B2, 20°34'45,78" S 40°11'30,74" W, 41m (1 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (3 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (3 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (2 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (1 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (9 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (1 ind).

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Description: A medium-sized Malacoceros species, largest complete specimen 7 mm long, 0.6 mm wide for 60 chaetigers, holotype 12 mm long, 0.9 mm wide at the widest point for 72 chaetigers (incomplete). Body slightly flattened dorsoventrally throughout, tapering towards the pygidium. Body color light yellow to golden in alcohol, markedly from chaetigers 7–45 to chaetigers 14–80 (depending on specimen size), strongest on the posterior region (Fig 41).

Prostomium bell-shaped, rounded anteriorly, bearing two short antero-lateral horns; extending as a narrow keel almost to the posterior margin of chaetiger 2. Posterior half of prostomium slightly elevated. Eyes present as two groups of 4–8 small eyespots distributed irregularly, located in the posterior half of the prostomium. Peristomium poorly developed, separated from chaetiger 1 and lacking lateral wings (Figs 41, 42A–B). Grooved palps reaching up to chaetiger 40, lost in most specimens. Proboscis saccular and highly ciliated, everted on some specimens.

Chaetiger 1 reduced and positioned dorsally, with only a few chaetae on both rami, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae well-developed, lanceolate on notopodium and elongated with a tapered tip on neuropodium. Prechaetal lamellae as a low flap on notopodium and absent on neuropodium. Chaetiger 2 slightly positioned dorsally.

Notopodial postchaetal lamellae lanceolate throughout, gradually reducing in size from the middle region onwards, always shorter than branchiae (except on chaetiger 1) (Figs 41, 42C–D). Notopodial prechaetal lamellae present as a prominent flap on anterior chaetigers, absent from chaetiger 15–22 (depending on specimen size) onwards. Interramal budge present from chaetigers 2–5 to chaetigers 13–32 (depending on specimen size), absent on remaining chaetigers. Between notopodia and neuropodia, presence of lateral organs on anterior and middle regions, more developed on middle region (Fig 42C–D).

Neuropodial postchaetal lamellae rounded on the anterior region and elliptical from middle region onwards, gradually reducing in size after the middle region, present as a low flap on last chaetigers. Neuropodial postchaetal lamellae with small median nipple-like projection from chaetiger 2 to chaetigers 3–8 (depending on specimen size) (Fig 42D), indistinct on chaetigers 4– 9 (depending on specimen size) and readily recognizable from chaetigers 12–30 to chaetigers 19– 50. Neuropodial prechaetal lamellae present as a very low flap on the anterior region.

Chaetae from notopodia and neuropodia organized in two rows. In notopodia, anterior row of short, non-limbate and lightly granulated capillaries (Fig 43A) and posterior row of long (up to 1.5 longer than anterior row), narrowly unilimbate and non-granulated capillaries (Fig 43B). In neuropodia, two rows of unilimbate and non-granulated capillaries (Fig 43C). Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner and less numerous (Fig 43D).

355

Hooks in neuropodia only, starting from chaetigers 19–39, up to 11 per fascicle, accompanied by row of 1–5 non-limbate capillaries. Hooks tridentate, with 2 small secondary teeth arranged side by side above the main tooth. Secondary hood absent. (Fig 43E). Sabre chaetae from chaetigers 5–23 (depending on specimen size), up to five per fascicle (Figs 42D, 43F).

Branchiae from chaetiger 1 onwards, present on most of the body, except last 8–13 chaetigers. Branchiae cirriform, ciliated on outer and inner margins, tapered at the tip and basally fused with notopodial postchaetal lamellae. Branchiae about the same size or shorter than notopodial postchaetal lamellae on chaetiger 1 and longer than notopodial lamellae on the remaining chaetigers, longest and overlapping at midline on the anterior region (up to four times longer than notopodial lamellae) and reducing in size on the posterior region.

Pygidium rounded, bearing 4 cirri.

Juvenile morphology: In juveniles, the prostomium possess short antero-lateral projections instead of horns. Juveniles also bear less eyespots, from 1–3 in each group.

Methyl green staining pattern: No discernible pattern observed. Whole specimens faintly stained.

Remarks. Malacoceros aureus sp. nov. is unusual for its light yellow to golden color on several chaetigers. Maciolek (1983) and Imajima (1991) reported a yellow tan in M. indicus specimens from the Atlantic Coast of the USA and Japan, respectively, while Blake (1996) reported specimens with a light tan with brown pigment on middle region on specimens from the Pacific Coast of the USA and Foster (1971b) observed pigmented granules, but made no further comments on the pigment’s color.

Malacoceros aureus sp. nov. is similar to M. vanderhorsti in having a bell-shaped prostomium, numerous small eyespots, branchiae present on most of the body and tridentate hooded hooks (although, according to Foster [1971a], M. vanderhorsti possess quadridentate hooks on posterior chaetigers. The two species can be readily distinguished by the presence and distribution of the interramal bulge – from chaetigers 2–5 to chaetigers 13–32 in M. aureus sp. nov. and from chaetigers1–2, present on most of the body in M. vanderhorsti; lack of nipple-like projection on neuropodial postchaetal lamellae in M. vanderhorsti; starting chaetiger of neuropodial hooded hooks – from chaetigers 19–39 in M. aureus sp. nov. and from chaetigers 70–90 in M. vanderhorsti and pygidium morphology – bearing four cirri in M. aureus sp. nov. and bearing two cirri in M. vanderhorsti.

The distinction of M. aureus sp. nov. and M. indicus, which is the most reported Malacoceros species in Brazil, is more complex since M. indicus was originally described to the Gulf of Mannar (India), but was subsequentially recorded in multiple locations, such as the Gulf of Mexico (Foster

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1971a), Atlantic Coast of the USA (Maciolek 1983), Japan (Imajima 1991), Pacific Coast of the USA (Blake 1996) and Australia (Meiβner & Gotting 2015). Each record presents morphological differences from the type material and from the syntype (Foster 1971b) (Table 5), which in many cases is just regarded as intraspecific variation. Meiβner & Gotting (2015) considers M. indicus a widespread species, while also recognizing the great morphological variability among different descriptions of the species and acknowledging the need of integrative taxonomic studies to access the status of these records.

Due to this uncertainty regarding M. indicus reports outside its type locality, which may represent more than one new species, it was decided to discuss M. aureus sp. nov. with Foster (1971b) description of a syntype of M. indicus from the type locality. M. aureus sp. nov. is similar to M. indicus in having multiple eyespots, interramal bulge starting in anterior chaetigers (although starting chaetiger and distribution are not clearly mentioned in M. indicus), overlap on the starting chaetiger of neuropodial hooded hooks – from chaetigers 19–39 in M. aureus sp. nov. and from chaetiger 37 in M. indicus and presence of tridentate hooded hooks (although M. indicus also possess quadridentate hooded hooks in posterior chaetigers).

Species can be distinguished based on the prostomium shape – bell-shaped and bearing two short antero-lateral horns in M. aureus sp. nov. and T-shaped, with prominent frontal horns in M. indicus. However, Foster (1971b) only listed the presence of morphological characters instead of their actual distribution, thus characters as the starting chaetiger and distribution of interramal bulge and nipple-like neuropodial projection and the starting chaetiger of sabre chaetae are unknown.

These gaps in Foster (1971b) description are troublesome, as, while M. indicus is treated as widely distributed, the species is poorly known and not sufficiently well-defined in its type locality. A study by Delgado-Blas & Salazar-Siva (2011) highlighted morphological differences between the original description and material collected from Caribe, questioning M. indicus’ cosmopolitism, while Meißner & Götting (2015) discussed intra-specific variety of M. indicus and sequenced genetic markers (COI, 16S and 18S), pointing the need of integrative taxonomic studies.

Etymology. The specific epithet, aureus (Latin adjective for golden), refers to the light yellow to golden coloration observed on specimens.

Habitat: coarse sand to very fine sand, 29–51 m depth.

Distribution: Southeastern Brazil (Rio de Janeiro and São Paulo states), Atlantic Ocean.

357

Acknowledgments

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Legends of illustrations

Figure 1. Map showing sampled area (blue polygon).

Figure 2. Scolelepis profunda sp. nov., whole fragment, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 3. Scolelepis profunda sp. nov. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from middle region. D. Hooded hook.

Figure 4. Scolelepis sp. 1, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 5. Scolelepis sp. 1. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae from anterior row. C. Neuropodial capillary chaetae from posterior row. D. Capillary from posterior region. E. Hooded hook.

Figure 6. Scolelepis sp. 2, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: ga, gametes; pe, peristomium; pr, prostomium.

Figure 7. Scolelepis sp. 2. A. Notopodial capillary chaetae from anterior row. B. Notopodial capillary chaetae from posterior row. C. Neuropodial capillary chaetae from anterior row. D. Neuropodial capillary chaetae from posterior row. E. Capillary from posterior region. F. Hooded hook.

Figure 8. Scolelepis sp. 3, anterior chaetigers and mid-body, lateral view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchia; ga, gametes; pe, peristomium; pr, prostomium.

Figure 9. Scolelepis sp. 3. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae from anterior row. C. Neuropodial capillary chaetae from posterior row. D. Capillary from posterior region. E. Hooded hook.

Figure 10. Dispio levisi sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 11. SEM of Dispio levisi sp. nov. A. Anterior region, dorsal view. B. Close-up of anterior region, dorsal view. C. Anterior region, lateral view. D. Middle region, lateral view. Abbreviations: ab, accessory branchiae; br, branchia; ca1, capillary from chaetiger 1; lo, lateral organ; nela, neuropodial lamellae; nola, notopodial lamellae; pe, peristomium; pr, prostomium;

366 prenola, prechaetal notopodial lamellae; posnola, postchaetal notopodial lamellae; sa, sabre chaeta.

Figure 12. Dispio levisi sp. nov. A. Row of long notochaetae from chaetiger 1. B. Capillary chaetae from anterior row. C. Capillary chaetae from posterior row. D. Tuft of notochaetae above notopodial rows. E. Capillary from posterior region. F. Hooded hook. G. Sabre chaetae.

Figure 13. Dispio cf. brachychaeta, anterior chaetigers and mid-body, lateral view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 14. Dispio cf. brachychaeta. A. Capillary chaetae from anterior row. B. Capillary chaetae from posterior row. C. Tuft of notochaetae above notopodial rows. D. Capillary from posterior region. E. Hooded hook. F. Sabre chaetae.

Figure 15. Dispio sp. 1, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 16. Dispio sp. 1. A. Row of long notochaetae from chaetiger 1. B. Notopodial capillary chaetae from anterior row. C. Notopodial capillary chaetae from posterior row. D. Tuft of notochaetae above notopodial rows. E. Neuropodial capillary chaetae. F. Capillary from posterior region. G. Hooded hook. H. Sabre chaetae.

Figure 17. Microspio cf. profunda, anterior chaetigers and mid-body, lateral view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 18. SEM of Microspio cf. profunda. A. Anterior fragment, dorsal view. B. Close-up of anterior region, dorsal view. C. Anterior fragment, lateral view. D. Posterior region (pygidium damaged), dorsal view. Abbreviation: br, branchiae; cb, ciliary band; en, elongated notochaetae; hh, hooded hook; no, nuchal organ; pe, peristomium; pr, prostomium.

Figure 19. Microspio cf. profunda. A. Capillary chaetae from anterior row. B. Capillary chaetae from posterior row. C. Capillary from posterior region. D. Needle-like capillaries from posterior notopodia. E. Hooded hooks. F. Inferior capillary chaetae.

Figure 20. Microspio alexandrae sp. nov., anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 21. Figure 18. SEM of Microspio alexandrae sp. nov. A. Anterior fragment, dorsal view. B. Close-up of anterior region, dorsal view. C. Anterior fragment, lateral view. D. Close-up of anterior fragment, lateral view. Abbreviation: br, branchiae; hh, hooded hook; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium.

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Figure 22. Microspio alexandrae sp. nov. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from posterior region. D. Needle-like capillaries from posterior notopodia. E. Hooded hook.

Figure 23. Microspio tridentata sp. nov., anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 24. SEM of Microspio tridentata sp. nov. A. Anterior fragment, dorsal view. B. Anterior fragment, lateral view. C. Neuropodia, lateral view. Abbreviations: br, branchia; hh, hooded hook; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium.

Figure 25. Microspio tridentata sp. nov., Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from posterior region. D. Hooded hooks. E. Hooded hooks.

Figure 26. Microspio quadridentata sp. nov., anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 27. SEM of Microspio quadridentata sp. nov. A. Anterior fragment, dorsal view. B. Close- up of anterior region, dorsal view. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 28. Microspio quadridentata sp. nov. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from posterior region. D. Hooded hook.

Figure 29. Microspio sp. 1, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 30. Microspio sp. 1. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae from anterior row. C. Neuropodial capillary chaetae from posterior row. D. Hooded hook. E. Inferior capillary chaetae.

Figure 31. Microspio sp. 2, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 32. Microspio sp. 2. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Inferior capillary chaetae. D. Capillary from posterior region. E. Hooded hook.

Figure 33. Spio anaclaudiae sp. nov., anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 34. SEM of Spio anaclaudiae sp. nov. A. Complete animal, dorsal view (anterior and middle regions) and ventral view (pygidium). B. Anterior region, dorsal view. C. Anterior and

368 middle regions, lateral view. D. Posterior region, ventral view. Abbreviations: br, branchiae; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium; pyg, pygidium.

Figure 35. Spio anaclaudiae sp. nov. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 36. Spio capixaba sp. nov., whole fragment, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 37. SEM of Spio capixaba sp. nov. A. Anterior fragment, dorsal view. B. Close-up of anterior region, dorsal view. C. End of fragment, dorsal view. D. Anterior fragment, dorsal view. Abbreviations: br, branchiae; cb, ciliary band; hh, hooded hook; pe, peristomium; pr, prostomium.–

Figure 38. Spio capixaba sp. nov. A. Capillary chaetae from anterior row. B. Capillary chaetae from posterior row. C. Capillary from posterior region. D. Hooded hook. E. Inferior capillary chaetae.

Figure 39. Spio sp. 1, whole fragment, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 40. Spio sp. 1. A. Notopodial capillary chaetae. B. Neuropodial capillary chaetae. C. Capillary from posterior region. D. Hooded hook. E. Sabre chaetae.

Figure 41. Malacoceros aureus sp. nov., anterior and middle chaetigers, lateral view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 42. SEM of Malacoceros aureus sp. nov. A. Anterior fragment, dorsal view. B. Anterior fragment, dorsal view. C. Anterior and middle regions, lateral view. Anterior region, lateral view. Abbreviations: br, branchiae; lo, lateral organ; nela, neuropodial lamella; nola, notopodial lamella; pe, peristomium; pr, prostomium; as, sabre chaetae.

Figure 43. Malacoceros aureus sp. nov. A. Notopodial capillary chaetae from anterior row. B. Notopodial capillary chaetae from posterior row. C. Neuropodial capillary chaetae. D. Capillary from posterior region. E. Hooded hook. F. Sabre chaetae.

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Tables

Table 1 – Comparative diagnostic characters of Scolelepis species from the Brazilian coast.

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Table 1 (continued) – Comparative diagnostic characters of Scolelepis species from the Brazilian coast.

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Table 2 – Comparative diagnostic characters of Dispio species from the Brazilian coast.

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Table 3 - Comparative diagnostic characters of Microspio species from this work.

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Table 4 – Comparative diagnostic characters of Spio species from the Brazilian coast.

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Table 4 (continued) – Comparative diagnostic characters of Spio species from the Brazilian coast.

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Table 5 – Comparative diagnostic characters of Malacoceros species from the Brazilian coast.

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Table 5 (continued) – Comparative diagnostic characters of Malacoceros species from the Brazilian coast.

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Capítulo 7

A contribution to the taxonomy of Spiophanes (Annelida: Spionidae), including new species

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista alvo: Zootaxa Status: A submeter

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A contribution to the taxonomy of Spiophanes (Annelida: Spionidae), including new species

Running title: NEW SPIOPHANES FROM BRAZIL

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract Spiophanes is a genus commonly found in Brazil waters, although no Brazilian species are known. Four Spiophanes species Grube, 1860 were recorded from southeastern Brazil, both from the continental shelf and the continental slope, up to 3000m depth. Three of these species – Spiophanes grubei sp. nov., S. aureus sp. nov. and S. profundus sp. nov. are new to science. The last species, Spiophanes duplex, is routinely recorded from Brazilian coast. All species are very morphologically similar in having the same body division, but species can be distinguished mainly on the morphology of frontal horns, antennae presence and morphology, length of nuchal organs, morphology of lamellae from the anterior region, presence/absence of hooded on the hooks, presence of modified notopodial chaetae on posterior most chaetigers and morphology of neuropodial chaetae from chaetigers 9–14. Spiophanes profundus sp. nov. is also unique to the Brazilian fauna in having interneuropodial pouches on the posterior region. An identification key and a comparison table to all four species recorded in this work are provided.

Keywords: Taxonomy, Annelida, deep-sea, morphology.

Introduction The Spionidae Grube, 1850 are one of the most abundant and speciose families of annelids, often dominating soft-bottom communities, especially in eutrophic environments. Globally, there are approximately 580 described species distributed in 39 genera (Blake et al. 2017), although this number most likely is underestimated, as the family has a history of taxonomic problems with

402 many well-documented species-complexes (Sato-Okoshi et al. 2016), species with dubious distributions, genera that have never been revised and Spionidae might even be paraphyletic, as the family most likely includes the families Uncispionidae Green, 1982, Trochochaetidae Pettibone, 1963 and Poecilochaetidae Hannerz, 1956 (Struck 2017).

Spiophanes Grube, 1860 is easily distinguished among spionids in having crook-like chaetae on neuropodia from chaetiger 1, fronto-lateral horns, body divided in three regions, nuchal organs of variable length and lack of branchiae, a character shared only with Spiogalea Aguirrezabalaga & Ceberio, 2005, Spiophanella Fauchald & Hancock, 1981, Glyphochaeta Bick, 2005, Amphipolydora Blake, 1983, some species of Polydorella Augener, 1914, as well as a single species of Prionospio Malmgren, 1867, P. nonatoi Peixoto & Paiva, 2019 and a single Aurospio species Maciolek, 1981a, Aurospio abranchiata Neal et al. in Paterson et al., 2016 (Williams 2004; Paterson et al. 2016 Peixoto & Paiva 2017; 2019). The first described Spiophanes species, S. kroyeri Grube, 1860, described to the Greenland Sea, Northwest Atlantic Ocean, was subsequentially recorded from the Atlantic coast of North America, Atlantic and Mediterranean Coast of Europe, the Red Sea and South and Southeast Brazil (Pardo et al. 2006; Amaral et al. 2013; Read & Fauchald 2020).

The genus Spiophanes is among the most specious genus within Spionidae, with 33 described species (Read & Fauchald 2020), besides a subjective synonym, Morants Chamberlin, 1919. A monotypic genus, Spiophanella was originally described as a Spiophanes species lacking crook- like chaetae, but was considered superficially similar to Prionospio or Laonice species (Pettibone 1962), while Blake et al. (2017) considers Spiophanella as a taxon with little or no affiliation to remaining spionid taxa.

Spiophanes is divided in three body regions, an anterior region from chaetiger 1 to chaetiger 4, a middle region from chaetiger 5 to last neuropodial chaetiger bearing capillary chaetae (usually chaetiger 14, although varying from chaetiger 13 to chaetiger 15) and a posterior region, characterized by the first neuropodia bearing neuropodial hooded hooks (Meiβner 2005). A new diagnostic character, the chaetal spreader morphology, was introduced by Meiβner & Hutchings (2003), corresponding to a glandular opening that is species-specific (Meiβner & Hutchings 2003) and possibly plays a role in tube construction (Meiβner et al. 2012). A specific chaeta, called bacillary chaetae, protrudes from these openings, being similar among different species. The type of chaetal spreader of most Spiophanes species was provided by Meiβner (2005), who revised the entire genus. Radashevsky (2012) considers the chaetal spreader morphology confusing, since Meiβner & Hutchings (2003) did not refer to the morphology of the spreaders, but to the number of projections on the margins of the gland openings, while showing illustrations with the anterior end in different positions.

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Currently, around 90 spionid species arranged in 19 genera have been recorded in Brazil (Amaral et al. 2013; Peixoto & Paiva 2019), but these numbers may be underestimated as a greater taxonomic effort coupled with molecular studies and recent surveys in previously unexplored areas revealed many undescribed species, especially from the deep sea (Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019).

The current state of knowledge of spionid diversity in Brazil is scarce in most areas, in most cases limited to shallow-water and coastline environments (except for Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019) and geographically restricted to the southern and southeastern coast of Brazil, as no contributions regarding Spionidae taxonomy have been published in the north and northeast coasts, besides species records from ecological papers, which were compiled by Amaral et al. (2013).

A few taxa have been studied in greater detail such as Dipolydora Verrill, 1881 (Radashevsky & Nogueira 2003), Polydora Bosc, 1802 (Radashevsky et al. 2006), Laonice Malmgren, 1867 (Kinberg 1866; Nonato et al. 1986; Radashevsky & Lana 2009), Pseudopolydora Czerniavsky, 1881 (Radashevsky & Migotto 2009), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017), Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), and Laubieriellus Maciolek, 1981b (Peixoto & Paiva 2019). Wide-scale surveys are rare, restricted to the continental shelf of the state of Paraná, southern Brazil (Bolívar & Lana 1987) and the continental slope of Campos Basin, state of Rio de Janeiro, southeastern Brazil (Paiva & Barroso 2010), which described no new species.

Five Spiophanes species have been recorded on the Brazilian coast (Amaral et al. 2013): Spiophanes anoculata Hartman, 1960, described from California (Pacific coast of the USA), S. berkeleyorum Pettibone, 1962, described from Vancouver Island (Pacific coast of Canada), S. bombyx (Claparède, 1870), described from the Mediterranean coast of Italy, S. duplex (Chamberlin, 1919), described from California (Pacific coast of the USA) and S. kroeyeri (Grube, 1860), described from the Greenland Sea, northwestern Atlantic Ocean. Two additional records, Spiophanes missionensis Hartman, 1941 and S. soederstroemi Hartman, 1953, were synonymized with S. duplex by Blake (1996) and Meiβner (2005), respectively. Researchers should interpret these records with caution, as many species recorded in the Brazilian coast (compiled by Amaral et al. 2013) are known species-complexes or were described from rather distant locations, and its presence in Brazil still needs to be verified.

In the present work, four Spiophanes species are recorded based on material collected off the state of Espírito Santo during the cruises of the “Marine Environmental Characterization of Espírito

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Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company) and focusing on the southeastern coast of Brazil. Three of these species – Spiophanes aureus sp. nov., Spiophanes profundus sp. nov. and Spiophanes grubei sp. nov. are new to science, whereas the remaining species was previously recorded from nearby locations.

Material and Methods

Sediment samples were collected from 2010 to 2013 during the oceanographic cruises of the AMBES Project. Two sampling efforts were carried out: the first during the summer (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and the second during the winter (sample codes Amb2, Amb11, Amb12, Amb13, and Amb14) on the Espírito Santo Basin and the northern portion of Campos Basin (Fig 1), southeastern Brazil. Sediment was sampled using a Van Veen Grab sampler on the continental shelf and a box corer on the continental slope, then fixed in situ with 10% formalin in seawater. Since specimens were fixed in 10% formalin, molecular studies were not possible.

Afterwards, sediment samples were washed in the laboratory, elutriated through a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm- mesh sieve (for samples collected on the continental slope and on the Watu Norte and Doce canyons) and stored in 70&% EtOH. Specimens were sorted and examined under a Leica S8 APO stereomicroscope and Nikon Eclipse E200 microscope. Photographs and measurements were made using a Leica M205C equipped with a camera using the Leica Application Suite. Main taxonomic characters were summarized in Table 1. An identification key is also provided for all species recorded in this study.

In order to reveal species-specific staining patterns and enhance the visualization of features, specimens were stained in a methyl green solution (1 g/L in 70% EtOH) for 20 s and then examined in distilled water to avoid stain fading.

Specimens used for scanning electron microscope (SEM) studies were dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each), followed by three changes of 100% ethanol.

Afterwards, specimens were critical point dried in CO2, mounted in stubs, sputter-coated in gold and examined under a JEOL JSM-6510 LV SEM or a JEOL JSM-6390 LV SEM.

Type material is being deposited in the Museu Nacional do Rio de Janeiro (MNRJP), Universidade Federal do Rio de Janeiro; Museu de Zoologia Adão José Cardoso (ZUEC-POL),

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Universidade Estadual de Campinas and in the Museu de Zoologia (MZUSP), Universidade de São Paulo, all located in Brazil. Non type material will be deposited in the Museu de Zoologia Adão José Cardoso (ZUEC-POL), Universidade Estadual de Campinas.

Results

Taxonomy

Phylum Annelida Lamarck, 1809

Family Spionidae Grube, 1850

Genus Spiophanes Grube, 1860

Type species: Spiophanes kroyeri Grube, 1860, by monotypy.

Synonym list: Morants Chamberlin, 1919 (subjective synonym)

Diagnosis (from Meiβner 2005): Body elongate; prostomium subtriangular, bell-shaped or rarely rounded, anterior margin never incised; frontal or lateral horns present or absent; eyespots present or absent; occipital antenna present or absent. Nuchal organs as two ciliated bands along dorsum, maximally extending to chaetiger 17, or as pair of dorsal loops not extending beyond chaetiger 6. Branchiae absent. Dorsal and transverse ciliated crests or bands usually present. Body divided into three regions: anterior region extending to chaetiger 4, with well-developed parapodial lamellae; middle body region from chaetiger 5 to last chaetiger bearing capillary chaetae rather than hooks in neuropodia (chaetigers 13–15); posterior region with neuropodial hooks. Middle chaetigers usually with parapodial glandular organs on chaetigers 5–7(8) and with chaetal spreaders of different types; from chaetiger 9 glands open as simple vertical slit. Ventrolateral intersegmental (or interneuropodial) pouches present or absent between neuropodia. Chaetiger 1 with 1–2 conspicuous crook-like chaetae in neuropodium; otherwise neurochaetae in anterior and middle body region all capillaries, arranged in 1–2 rows; posterior region with quadridentate hooks, hood present or absent. Notochaetae all capillaries, in middle body region usually arranged in three rows; otherwise in two rows or indistinct rows. Bacillary chaetae may be present from chaetigers 5–8. One to two ventral sabre chaetae usually from chaetiger 4, rarely from chaetigers

406

5 or 10, or sometimes not present until neuropodial hooks appear. Stout, curved notochaetae often present in far posterior parapodia. Pygidium with two or more anal cirri.

Spiophanes grubei sp. nov.

(Figs 2–4H)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 11 Dec 2010 to 19 Dec 2010, 53m, MNRJP XXXX. Paratypes: Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 12 Jul 2011 to 18 Jul 2011, 47m, MNRJP XXXX (2 ind), MZUSP XXXX (3 ind), ZUEC-POL XXXX (3 ind).

Additional material examined. Amb1 Foz4, 19°37'21,13" S 39°45'6,41" W, 15m (10 ind); Amb1 Foz10, 19°35'12,39" S 39°38'33,16" W, 32m (1 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (18 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (7 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (2 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (1 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (1 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (15 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (24 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (4 ind); Amb1 Foz19, 19°46'14,99" S 39°34'50,94" W, 52m (8 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (18 ind); Amb2 Foz4, 19°37'12,91" S 39°45'11,36" W, 13m (1 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W, 47m (5 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W, 39m (1 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W, 41m (2 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W, 51m (13 ind); Amb2 Foz17, 19°55'45,59" S 39°45'41,35" W, 52m (14 ind); Amb2 Foz18, 19°50'16,39" S 39°40'11,23" W, 53m (7 ind); Amb2 Foz19, 19°46'10,69" S 39°34'55,84" W, 51m (15 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W, 53m (7 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (7 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (3 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (91 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (37 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W, 40m (4 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W, 49m (11 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (15 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (6 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (34 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (10 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (2 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (37 ind); Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (4 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (13 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (22 ind); Amb7 E1, 19°9'48,26" S 39°29'19,79" W, 26m (2 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (2 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (20 ind); Amb7 F1, 18°42'56,07" S 39°31'28,8" W, 25m (5 ind); Amb7 F4, 19°32'57,67" S 38°42'57,83" W, 152m (7 ind); Amb7 G1, 18°32'3,6" S

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39°23'16,91" W, 25m (2 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (20 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (40 ind); Amb12 CANWN4, 19°49'6,26" S 39°36'9,34" W, 181m (1 ind); Amb14 A4, 21°4'4,56" S 40°14'14,08" W, 141m (7 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (13 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (12 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (2 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (13 ind); Amb14 E1, 19°9'51,36" S 39°29'20,53" W, 27m (2 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (1 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (1 ind); Amb14 F4, 19°33'2,92" S 38°42'52,26" W, 160m (5 ind).

Description: A medium-sized Spiophanes species, largest complete specimen 9.5 mm long, 1 mm wide at the widest point for 63 chaetigers, holotype 8 mm long, 8 mm wide at the widest point for 55 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium and divided in three regions: anterior region from chaetiger 1 to chaetiger 4, middle region from chaetiger 5 to chaetiger 14 and posterior region from chaetiger 15 onwards. Parapodia from chaetiger 1 shifted dorsally; parapodia from chaetiger 2–4 shifted dorsolaterally. Body color whitish in alcohol, with small pigmented yellow spots on lateral side of chaetigers 8–14 (most intense on chaetigers 10–13) and yellow to orange spots on notopodial lamellae from last 3–5 chaetigers (Fig 2).

Prostomium bell-shaped, rounded anteriorly and bearing short and thin anterolateral horns (Figs 2, 3A–B) Prostomium narrowing posteriorly, extending to the posterior margin of chaetiger 1. Continuous nuchal organs reaching posterior margin of chaetiger 3, metameric nuchal organs reaching chaetiger 18 (Fig 3A). Two pairs of eyes in trapezoidal arrangement or a single pair of eyes present. Occipital antenna as a small papilla on the posterior part of the prostomium (Fig 3B). Peristomium weakly developed, surrounding prostomium for about half of its length; lacking lateral wings and not fused to chaetiger 1 (Figs 2, 3A–B). Grooved palps reaching up to chaetiger 11, lost in most specimens. Proboscis saccular and highly ciliated, everted on some specimens (Fig 3A–B).

Chaetiger 1 with lamellae cirriform on both rami, slightly more developed on neuropodia. On chaetigers 2–5, lamellae cirriform on both rami, slightly more developed on notopodia. From chaetiger 6 to chaetiger 14, notopodial lamellae elliptical or rounded and gradually smaller on notopodia and reduced in neuropodia. Chaetal spreader semicircular (type 0 +1) on chaetigers 5– 8 (Fig 3C). Opening of glandular organs as simple vertical slits on chaetigers 9–13. From chaetiger 15 onwards, notopodial lamellae gradually longer, becoming cirriform with a pointed

408 tip; neuropodial lamellae reduced. Very low dorsal crests starting from chaetiger 4, present on 15–20 following chaetigers (Figs 2, 3D).

Chaetae from anterior region organized in two rows of non-limbate and non-granulated capillaries on both rami (Fig 4A); presence of 1–2 robust crook-like chaetae on neuropodium of chaetiger 1 (Figs 3B, 4B) and a row of long notopodial capillaries on chaetiger 2 (Figs 3B, 4C). On middle region, chaetae organized in three rows of narrowly unilimbate and non-granulated capillaries on notopodia (Figs 3C, 4D) – anterior row shortest and posterior row longest, and two rows of short, thick, unilimbate and granulated capillaries on neuropodia (Fig 4E). Bacillary chaetae inside glandular opening from chaetiger 6 (Figs 3C, 4F). On posterior region, narrowly unilimbate and non-granulated capillaries on notopodia (Fig 4G) – gradually becoming elongate, non-limbate, thinner and less numerous, and solely hooded hooks in neuropodia (Fig 4H) (described below). On last 1–6 chaetigers, presence of 1–2 robust and bent notopodial chaetae (Fig 4J), accompanied by non-limbate and non-granulated capillaries.

Hooks in neuropodia only, consistently starting from chaetiger 15, up to nine per fascicle, not accompanied by capillaries (Fig 3E–F). Hooks quadridentate, with single unpaired tooth immediately above main tooth and upper-most pair of minute secondary teeth organized side by side (Fig 4H). Hood reduced, present only below main tooth. Sabre chaetae consistently starting from chaetiger 4. Sabre chaetae thick, non-limbate and intensely granulated throughout length (Figs 3E, 4I).

Pygidium bearing pair of dorsal cirri and a single ventral papilla.

Oocytes from chaetiger 16, with well-defined nucleus and measuring up to 280 µm.

Glandular organs from chaetiger 5 to chaetiger 14, largest on chaetigers 7 and 8.

Muscular pharynx observed through light microscopy, from chaetiger 18 to chaetiger 21.

Methyl green staining pattern: Glandular organs from chaetigers 5–14 strongly stained, lamellae from anterior and middle regions weakly stained.

Remarks. Spiophanes grubei sp. nov. is similar to S. bombyx and S. anoculata in having a bell- shaped prostomium, thin anterolateral horns, weakly developed peristomium, presence of papilliform antenna (in S. anoculata), hooks starting from chaetiger 15 (chaetiger 14 in some juveniles of S. bombyx), hooks possessing a partial hood, sabre chaetae starting from chaetiger 4 (in S. anoculata) and presence of robust and bent notopodial chaetae in posterior chaetigers (in S. bombyx) (Meiβner 2005).

However, species can be distinguished by horn length – short in S. grubei sp. nov. and S. anoculata and long in S. bombyx, presence of antennae in S. grubei sp. nov. and S. anoculata,

409 absent in S. bombyx; length of dorsal ciliated organs – extending to chaetiger 18 in S. grubei sp. nov. (as segmental ciliated grooves), chaetiger 22 in S. anoculata (as segmental ciliated grooves) and chaetiger 2 in S. bombyx; shape of notopodial lamellae from anterior region – cirriform in S. grubei sp. nov. and subulate in S. anoculata and S. bombyx; as well as the shape of notopodial lamellae on middle and posterior region – cirriform on chaetiger 5, elliptical on chaetigers 6–14 and gradually longer, becoming cirriform with a pointed tip in S. grubei sp. nov., subtriangular on chaetigers 5–8 and subtriangular to cirriform with broad base from chaetiger 9 onwards in S. bombyx and subulate from chaetiger 5 onwards in S. anoculata (Hartman 1960; Meiβner 2005).

Species can also be distinguished based on dorsal crests – from chaetiger 4 in S. grubei sp. nov. and chaetiger 3 in S. anoculata and S. bombyx; shape of neuropodial lamellae from anterior region – cirriform in S. grubei sp. nov., subulate on chaetiger 1 and subtriangular to rounded on chaetigers 2–4 in S. bombyx and subulate in S. anoculata; presence of robust and bent notopodial chaetae in posterior chaetigers in S. grubei sp. nov. and S. bombyx, lacking in S. anoculata; starting chaetiger of sabre chaetae – chaetiger 4 in S. grubei sp. nov. and S. anoculata and chaetiger 10 or 15 (depending on locality) in S. bombyx (Hartman 1960; Meiβner 2005) and distribution and morphology of glandular openings – semicircular and on chaetigers 5–8 in both S. grubei sp. nov. and S. anoculata and undulate, on chaetigers 5, 7 and 8 in S. bombyx. Given superficial similarities in the prostomium morphology, it is possible that Brazilian records of S. bombyx and S. anoculata correspond to S. grubei sp. nov. In fact, no material from these species was recorded in this work, and Pardo et al. (2006) record of S. bombyx and Pardo & Peixoto (in press) record of S. anoculata corresponds to S. grubei sp. nov.

Among species described in this work, Spiophanes grubei sp. nov. is unique due to its set of characters – bell-shaped prostomium, short and thin anterolateral horns, presence of papilliform antenna, dorsal ciliated organs reaching to chaetiger 18, neuropodial lamellae cirriform on chaetigers 1–5, lack of interneuropodial pouches, reduced hoods present on the hooks and presence of modified notochaetae on the posterior region.

Etymology. The species epithet, grubei, is a tribute to Adolph E. Grube (1812–1880), a German zoologist who described the genus Spiophanes.

Habitat: coarse sand to medium silt, 13–181 m depth.

Distribution: Southeastern Brazil (Espírito Santo state), Atlantic Ocean.

410

Spiophanes duplex (Chamberlin, 1919)

(Figs 5–7K)

Synonym list:

Morants duplex Chamberlin, 1919 (superseded original combination)

Spiophanes chilensis Hartmann-Schröder, 1965 (subjective synonym)

Spiophanes missionensis Hartman, 1941 (subjective synonym)

Spiophanes söderströmi Hartman, 1953 (subjective synonym)

Material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (3 ind); Amb1 Foz4, 19°37'21,13" S 39°45'6,41" W, 15m (3 ind); Amb1 Foz6, 19°54'56,16" S 39°56'40,83" W, 34m (2 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m (22 ind); Amb1 Foz8, 19°44'45,48" S 39°46'25,78" W, 32m (3 ind); Amb1 Foz9, 19°39'45,36" S 39°42'29,16" W, 29m (1 ind); Amb1 Foz10, 19°35'12,39" S 39°38'33,16" W, 32m (5 ind); Amb1 Foz11, 19°57'32,89" S 39°53'30,69" W, 47m (27 ind); Amb1 Foz12, 19°52'35,48" S 39°49'5,63" W, 43m (6 ind); Amb1 Foz13, 19°47'32,83" S 39°43'15,08" W, 41m (4 ind); Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (7 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (25 ind); Amb1 Foz16, 20°1'3,73" S 39°50'13,76" W, 53m (1 ind); Amb1 Foz17, 19°55'44,66" S 39°45'38,7" W, 51m (4 ind); Amb1 Foz18, 19°50'22,01" S 39°40'6,1" W, 54m (1 ind); Amb1 Foz19, 19°46'14,99" S 39°34'50,94" W, 52m (1 ind); Amb1 Foz20, 19°41'33,92" S 39°31'17,74" W, 54m (37 ind); Amb2 Foz1, 19°52'14,57" S 39°59'41,06" W, 27m (1 ind); Amb2 Foz6, 19°54'50,43" S 39°56'47,01" W,34m (2 ind); Amb2 Foz7, 19°49'52,15" S 39°52'24,51" W,30m (29 ind); Amb2 Foz11, 19°57'32,36" S 39°53'33,01" W,47m (8 ind); Amb2 Foz12, 19°52'29,66" S 39°49'8,1" W,46m (44 ind); Amb2 Foz13, 19°47'22,52" S 39°43'20,72" W,41m (1 ind); Amb2 Foz14, 19°42'26,81" S 39°39'5,27" W,39m (4 ind); Amb2 Foz15, 19°37'41,83" S 39°35'31,52" W,41m (18 ind); Amb2 Foz16, 20°1'2,6" S 39°50'18,72" W,51m (2 ind); Amb2 Foz20, 19°41'24,99" S 39°31'20,42" W,53m (18 ind); Amb2 Foz20b, 19°41'29,14" S 39°31'18,18" W, 53m (11 ind); Amb7 A2, 21°3'27,14" S 40°22'59,61" W,40m (2 ind); Amb7 A3, 21°4'1,29" S 40°18'50,11" W,49m (2 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W,153m (34 ind); Amb7 B1, 20°34'32,47" S 40°20'52,37" W, 25m (21 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (9 ind); Amb7 C2, 20°11'25,35" S 40°2'16,02" W, 39m (25 ind); Amb7 C3, 20°12'20,26" S 39°57'59,7" W, 50m (1 ind); Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (26 ind); Amb7 D2, 19°40'26,04" S 39°36'19,65" W, 40m (6 ind); Amb7 D3, 19°43'14,34" S 39°33'34,86" W, 50m (37 ind); Amb7 E1, 19°9'48,26" S

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39°29'19,79" W, 26m (1 ind); Amb7 E2, 19°18'5,9" S 39°23'23,3" W, 39m (5 ind); Amb7 E3, 19°26'5" S 39°17'38,92" W, 50m (33 ind); Amb7 F1, 18°42'56,07" S 39°31'28,8" W, 25m (1 ind); Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (4 ind); Amb7 F4, 19°32'57,67" S 38°42'57,83" W, 152m (1 ind); Amb7 G2, 18°36'31,68" S 39°9'33" W, 39m (2 ind); Amb7 G3, 18°40'55,3" S 38°55'41,48" W, 55m (5 ind); Amb13 G2, 18°36'32,45" S 39°9'32,83" W, 40m (14 ind); Amb13 G3, 18°40'57,41" S 38°55'39,92" W, 53m (4 ind); Amb14 A2, 21°3'31,13" S 40°22'59,88" W, 39m (1 ind); Amb14 B1, 20°34'34,37" S 40°20'50,77" W, 26m (3 ind); Amb14 B2, 20°34'47,13" S 40°11'31,1" W, 41m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (1 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (2 ind); Amb14 C3, 20°12'21,46" S 39°58'0,3" W, 50m (8 ind); Amb14 D1, 19°35'35,96" S 39°41'20,61" W, 26m (1 ind); Amb14 D2, 19°40'25,29" S 39°36'21,96" W, 41m (4 ind); Amb14 D3, 19°43'14,33" S 39°33'34,78" W, 50m (17 ind); Amb14 E1, 19°9'51,36" S 39°29'20,53" W, 27m (1 ind); Amb14 E2, 19°18'6,12" S 39°23'23,35" W, 38m (15 ind); Amb14 E3, 19°26'4,81" S 39°17'38,64" W, 50m (19 ind); Amb14 F3, 18°53'31,97" S 39°6'21,78" W, 51m (2 ind); Amb14 F4, 19°33'2,92" S 38°42'52,26" W, 160m (1 ind).

Description: A medium-sized Spiophanes species, largest complete specimen 14 mm long, 1.5 mm wide at the widest point for 68 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium and divided in three regions: anterior region from chaetiger 1 to chaetiger 4, middle region from chaetiger 5 to chaetiger 14 and posterior region from chaetiger 15 onwards. Parapodia from chaetiger 1 shifted dorsally; parapodia from chaetiger 2–4 shifted dorsolaterally. Body color whitish in alcohol (Fig 5).

Prostomium narrow, rounded anteriorly and bearing short and fleshy anterolateral horns (Fig 5, 6A–B) Prostomium extending posteriorly to the posterior margin of chaetiger 1. Continuous nuchal organs reaching posterior margin of chaetiger 2, metameric nuchal organs reaching chaetiger 15 (Fig 6A). One to two pairs of eyes or eyes absent. Occipital antenna absent. Peristomium weakly developed, surrounding prostomium for about half of its length; lacking lateral wings and not fused to chaetiger 1 (Figs 5, 6A–B). Grooved palps reaching up to chaetiger 20, lost in most specimens. Proboscis saccular and highly ciliated, everted on some specimens.

Chaetigers 1–2 with lamellae cirriform on both rami, slightly more developed on neuropodia. On chaetigers 3–5, lamellae cirriform on both rami, slightly more developed on notopodia (Fig 5). From chaetiger 6 to chaetiger 14, notopodial lamellae digitiform to rounded and gradually smaller on notopodia and reduced in neuropodia (Fig 6A). Chaetal spreader undulate, type “2 + 3”, present on chaetigers 5–7 (Fig 6D). Opening of glandular organs as simple vertical slits on chaetigers 9– 14. From chaetiger 15 onwards, notopodial lamellae gradually longer, subtriangular to rounded

412 with long and fragile cirriform projection (Fig 6E); neuropodial lamellae reduced. Very low dorsal crests starting from chaetigers 16–20, present on 8–10 following chaetigers (Fig 5).

Chaetae from anterior region organized in two rows of non-granulated capillaries, narrowly unilimbate on notopodia (Fig 7A) and widely unilimbate on neuropodia (Fig 7B), besides 1–2 robust crook-like chaetae on neuropodium of chaetiger 1 (Figs 6C, 7C). On middle region, chaetae organized in three rows of short, lightly granulated and bilimbated capillaries on notopodia (Fig 7D) – anterior row shortest and posterior row longest, and two rows of lightly granulated and widely unilimbate capillaries on neuropodia (Fig 7E). Bacillary chaetae present on chaetigers 5– 7 (Fig 7F). On posterior region, narrowly unilimbate and non-granulated capillaries on notopodia (Fig 7G) – gradually becoming elongate, non-limbate, thinner and less numerous (Fig 7H), and solely hooded hooks in neuropodia (Figs 6F, 7I) (described below). On last 6–13 chaetigers, presence of 1–3 robust and bent notopodial chaetae (Figs 6E, 7K), accompanied by non-limbate and non-granulated capillaries.

Hooks in neuropodia only, consistently starting from chaetiger 15, up to eight per fascicle, not accompanied by capillaries. Hooks quadridentate, with single unpaired tooth immediately above main tooth and upper-most pair of minute secondary teeth organized side by side (Fig 7I). Hood completely absent. Sabre chaetae consistently starting from chaetiger 4. Sabre chaetae thick, non- limbate and non-granulated (Fig 7J).

Pygidium bearing pair of dorsal cirri.

Oocytes from chaetigers 16–19, measuring up to 220 µm; at first occupying only lateral sides of body, progressively increasing occupied area, last chaetigers completely filled with oocytes.

Glandular organs from chaetiger 5 to chaetiger 7.

Methyl green staining pattern: Prostomium, margins of peristomium and parapodia from chaetigers 1–4 strongly stained; parapodia and dorsal side from chaetigers 5–14, ventral side of chaetigers 10–14 weakly stained.

Remarks. Spiophanes duplex was originally described to California (Northern Pacific Coast of the USA), being posteriorly reported to the Southern Pacific Ocean (off Chile, as Spiophanes chilensis Hartmann-Schröder,1965), the Northern Atlantic Ocean (Gulf of Mexico) and the Southern Atlantic Ocean (Brazil, Uruguay and Argentina, as Spiophanes duplex and Spiophanes söderströmi Hartman, 1953). The species was originally referred to Brazil by Flynn et al. (1999), although Spiophanes missionensis Hartman, 1941 and Spiophanes söderströmi, which are currently synonymized with Spiophanes duplex (Blake 1996; Meiβner 2005), were previously recorded from the Brazilian coast (Tommasi 1967; Orensanz & Gianuca 1974 respectively).

413

Despite being described to a distant locality, specimens collected in this study agree with Meiβner (2005) redescription of Spiophanes duplex. Specimens share a bell-shaped with short and fleshy horns, lack of antennae, similar morphology of notopodial and neuropodial lamellae, same starting chaetiger of neuropodial hooks and sabre chaetae and presence of modified notochaetae on posterior chaetigers. The only differences from the original description were the peristomium morphology – always weakly developed in specimens from this work and weakly to moderately developed in Meiβner (2005); length of dorsal ciliated organs – reaching the anterior end of chaetiger 15 in specimens from this work and chaetigers 16–17 in Meiβner (2005); morphology of dorsal ciliated organs near the end of length – as slightly diverging bands in our specimens and alternately diverging and converging on chaetigers 15–17 in Meiβner (2005, Fig 18B); starting chaetiger of dorsal crests – chaetigers 16–20 in specimens from this work and chaetigers 17–18 in Meiβner (2005) and pygidium morphology –bearing a pair of dorsal cirri in specimens from this work and bearing a bearing pair of lateral cirri and a medioventral inconspicuous papilla in Meiβner (2005). Unfortunately, the chaetal spreader was not observed in our specimens. Genetic studies are highly needed for this species, which could reveal if specimens from Brazil really belong to Spiophanes duplex or a cryptic species.

The species can be distinguished from the remaining Brazilian Spiophanes species due to its set of characters – narrow and anteriorly rounded prostomium bearing short and fleshy anterolateral horns, lack of antenna, dorsal ciliated organs reaching to chaetiger 18, weakly developed peristomium, shape of notopodial and neuropodial lamellae, distribution of dorsal crests, lack of interneuropodial pouches, morphology of neurochaetae from chaetigers 9–14, lack of hood on the hooks and presence of modified notochaetae on posterior chaetigers (Table 1).

Habitat: coarse sand to coarse silt, 15–160 m depth.

Distribution: Brazil, continuous from Santa Catarina to Bahia states and Ceará state, Atlantic Ocean.

Spiophanes aureus sp. nov.

(Figs 8A–10L)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb3 E5, 19°36'26,24" S 39°10'17,35" W, 02 Dec 2011 to 02 Feb 2012, 392m, MNRJP XXXX. Paratypes: Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 02 Dec 2011 to 02 Feb 2012, 450m, MNRJP XXXX (8 ind), MZUSP XXXX (8 ind), ZUEC-POL XXXX (8 ind).

414

Additional material examined. Amb3 E5, 19°36'26,24" S 39°10'17,35" W, 392m (138 ind); Amb3 F5, 19°34'20,42" S 38°41'18,43" W, 450m (170 ind); Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (105 ind); Amb3 CAND5, 19°33'20,99" S 39°2'36,2" W, 420m (14 ind); Amb5 A5, 21°4'9,61" S 40°13'7,38" W, 410m (2 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (3 ind); Amb5 B5, 20°35'16,23" S 39°53'47,1" W, 410m (218 ind); Amb5 C5, 20°14'19,45" S 39°48'36,67" W, 381m (172 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (404 ind); Amb6 D5, 19°46'34,99" S 39°30'4,65" W, 428m (14 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (28 ind); Amb6 CANWN4, 19°49'7,27" S 39°36'8,52" W, 158m (129 ind); Amb6 CANWN5, 19°49'37,21" S 39°35'41,25" W, 410m (6 ind); Amb7 A4, 21°4'4,76" S 40°14'14,14" W, 153m (64 ind); Amb7 B4, 20°35'25,16" S 39°54'58,31" W, 157m (87 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (105 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (17 ind); Amb11 A5, 21°4'4,67" S 40°13'6,06" W, 415m (2 ind); Amb11 B5, 20°35'15,33" S 39°53'45,22" W, 415m (225 ind); Amb11 C5, 20°14'17,95" S 39°48'34,35" W, 395m (185 ind); Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 1040m (1 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (252 ind); Amb12 D5, 19°46'32,84" S 39°30'3,65" W, 416m (52 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (8 ind); Amb12 E5, 19°36'30,6" S 39°10'19,39" W, 360m (61 ind); Amb12 F5, 19°34'20,47" S 38°41'19,8" W, 449m (221 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (72 ind); Amb12 CAND5, 19°33'22,17" S 39°2'36,03" W, 414m (6 ind); Amb12 CANWN4, 19°49'6,26" S 39°36'9,34" W, 181m (87 ind); Amb12 CANWN5, 19°49'36,9" S 39°35'42,69" W, 378m (12 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (2 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (36 ind).

Description: A medium-sized Spiophanes species, largest complete specimen 10 mm long, 0.9 mm wide at the widest point for 52 chaetigers, holotype 8 mm long, 0.8 mm wide at the widest point for 48 chaetigers. Body slightly dorsoventrally flattened throughout, tapering towards the pygidium and divided in three regions: anterior region from chaetiger 1 to chaetiger 4, middle region from chaetiger 5 to chaetiger 14 and posterior region from chaetiger 15 onwards. Parapodia from chaetiger 1 shifted dorsally. Body color whitish in alcohol, with noto- and neuropodial lamellae from anterior region and notopodial lamellae from middle region stained in yellow or golden pigment (Fig 8A–B).

Prostomium narrow and elongate, truncated anteriorly and bearing fleshy anterolateral horns. Prostomium extending posteriorly to the posterior margin of chaetiger 1 (Fig 8A, 9A–B). Continuous nuchal organs reaching posterior margin of chaetiger 6 (Fig 9A–B). One pair of small eyes or eyes absent. Occipital antenna short, near the posterior margin of chaetiger 1. Peristomium

415 well-developed, surrounding prostomium for about two thirds of its length, forming low and wide lateral wings, not fused to chaetiger 1 (Fig 8A). Grooved palps reaching up to chaetiger 16, lost in most specimens. Proboscis saccular and highly ciliated, everted on some specimens.

Chaetiger 1 bearing cirriform lamella on notopodium and foliaceous with cirriform tip on neuropodium. On chaetigers 2–4, lamellae foliaceous, widened at the base and abruptly tapered (with pointed tip) on notopodia and foliaceous, lacking pointed tip on neuropodia (Fig 8A, 9A– B). From chaetiger 5 to chaetiger 8, notopodial lamellae rounded, lacking cirriform projection; from chaetiger 9 onwards, notopodial lamellae rounded and progressively smaller towards the posterior region, bearing long and delicate dorsal projection (Fig 9A). Chaetal spreader kidney- shaped (type 0 + 1) on chaetigers 5–7 (Fig 9D–E). Opening of glandular organs as simple vertical slits on chaetigers 8–14. Neuropodial lamellae absent from chaetiger 5 onwards. Prechaetal lamellae absent on both rami. Very low dorsal crests from chaetigers 16–17 to about chaetiger 50. Interneuropodial pouches absent.

On anterior region, chaetae organized in two rows, granulated and widely bilimbate capillaries on notopodia (Fig 10A) and lightly granulated and narrowly bilimbate on neuropodia (Fig 10B), besides very long row of non-limbate notopodial capillaries on chaetiger 1 (Fig 10C) as well as 1–2 robust crook-like chaetae on neuropodium of chaetiger 1 (Figs 9C, 10D). On middle region, chaetae organized in three rows of short, granulated and bilimbated capillaries on notopodia (Fig 10E) – anterior row longest and posterior row shortest, and two rows of lightly granulated and bilimbate capillaries on neuropodia (Fig 10F). Neuropodial capillaries from chaetigers 9–14 wide and bearing pointed tip (Fig 10G). Bacillary chaetae present on chaetigers 5–7 (Fig 10H). On posterior region, non-limbate and non-granulated capillaries on notopodia (Fig 10I), gradually becoming thinner and less numerous (Fig 10J) and solely hooded hooks in neuropodia (Fig 10 K) (described below). Lack of robust notochaetae on posterior-most chaetigers.

Hooks in neuropodia only, consistently starting from chaetiger 15, up to four per fascicle, not accompanied by capillaries. Hooks long and slightly constricted near the tip. Hooks quadridentate, with single unpaired tooth immediately above main tooth and upper-most pair of minute secondary teeth arranged side by side (Figs 9F, 10K). Hood completely absent. Sabre chaetae consistently starting from chaetiger 4, up to two per rami. Sabre chaetae non-limbate, bearing numerous transversal columns near the base and intensely granulated near the tip (Figs 9F, 10L).

Pygidium bearing pair of dorsal cirri.

Glandular organs large, from chaetiger 5 to chaetiger 7.

416

Methyl green staining pattern: Prostomium, dorsal side of peristomium and notopodial lamellae from middle region strongly stained; pigmentation weakly diffused throughout whole body.

Remarks. Spiophanes aureus sp. nov. can be distinguished from remaining Brazilian Spiophanes species as it bears a narrow, elongate and anteriorly rounded prostomium, fleshy anterolateral horns, presence of a short occipital antenna, dorsal ciliated organs reaching chaetiger 6, low peristomium forming wide lateral wings, shape of notopodial and neuropodial lamellae (specially on the anterior region), distribution of dorsal crests, lack of interneuropodial pouches, wide and distally pointed capillaries on chaetigers 9–14, lack of hood on the hooks and lack of modified notochaetae on posterior chaetigers (Table 1).

Among species reported in Brazil, S. berkeleyorum, S. anoculata, S. grubei sp. nov. and S. profundus sp. nov. also possess pigmentation patterns, although species can be readily distinguished based on position of pigmented spots and pigment color – noto- and neuropodial lamellae from anterior region and notopodial lamellae from middle region stained in yellow or golden pigment in S. aureus sp. nov., orange to brownish pigment on chaetigers 9–15 at the base of the parapodia and ventrolaterally on the segments in S. berkeleyorum, orange to brownish pigment visible laterally on chaetigers 10–14 in S. anoculata, yellow spots on lateral side of chaetigers 8–14 and yellow to orange spots on notopodial lamellae from last 3–5 chaetigers in S. grubei sp. nov. and brownish pigment at the base of notopodia from middle region in S. profundus sp. nov. (Meiβner 2005).

The species is most similar to S. anoculata, S. berkeleyorum, S. kroeyeri, S. grubei sp. nov. and S. profundus. sp. nov. in having an occipital antenna, although S. aureus can be readily distinguished from S. kroeyeri and S. profundus. sp. nov. in lacking interneuropodial pouches, present on the two aforementioned species. Spiophanes aureus sp. nov. can be separated from S. anoculata and S. berkeleyorum based on the prostomium shape, narrow and elongated in S. aureus sp. nov.; shape of anterolateral horns – fleshy in S. aureus sp. nov. and S. berkeleyorum and thin in S. anoculata and S. grubei sp. nov.; antenna morphology – short in S. aureus sp. nov., papilliform in both S. anoculata and S. grubei sp. nov. and long in S. berkeleyorum; peristomium morphology – well-developed in S. aureus sp. nov. and S. berkeleyorum and weakly developed in S. anoculata and S. grubei sp. nov.; shape of notopodial and neuropodial lamellae; starting chaetiger of dorsal crests – chaetigers 16–17 in S. aureus sp. nov., chaetigers 17–18 in S. berkeleyorum, chaetiger 4 in S. grubei sp. nov. and chaetiger 3 in S. anoculata; lack of modified notochaetae on posterior chaetigers in S. aureus sp. nov. and S. anoculata, present in S. berkeleyorum and in S. grubei sp. nov.; and pygidium morphology – bearing a pair of dorsal cirri in S. aureus sp. nov., bearing a pair of dorsal cirri and a single ventral papilla in S. grubei sp.

417 nov., 8–12 cirri in S. berkeleyorum, being of unknown morphology in S. anoculata (Hartman 1960; Meiβner 2005).

This was the most abundant Spiophanes species in this work, with over 2,900 collected specimens, found mainly in sediment samples from the deep continental shelf and shallow continental slope.

Etymology. The species epithet, aureus (from aurum, Latin word for gold), refers to the yellow or golden pigment present on the anterior and middle regions.

Habitat: very fine sand to fine silt, 143–1040 m depth.

Distribution: Southeastern Brazil (Espírito Santo state).

Spiophanes profundus sp. nov.

(Figs 11A–13J)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb3 CAND7, 19°42'18,42" S 39°5'52,41" W, 02 Dec 2011 to 02 Feb 2012, 1307m, MNRJP XXXX. Paratypes: Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 02 Dec 2011 to 02 Feb 2012, 1300m, MNRJP XXXX (2 ind), MZUSP XXXX (2 ind); Amb11 F6, 19°52'52,59" S 38°35'10,48" W, 06 Jun 2013 to 17 Jul 2013, 1021m, ZUEC-POL XXXX (4 ind).

Additional material examined. Amb3 E6, 19°40'8,03" S 39°7'22,1" W, 1035m (23 ind); Amb3 G9, 19°3'13,43" S 37°45'37,49" W, 2426m (2 ind); Amb3 CAND6, 19°37'50,08" S 39°3'56,16" W, 970m (5 ind); Amb3 CAND7, 19°42'18,42" S 39°5'52,41" W, 1307m (17 ind); Amb4 B8, 20°41'33,45" S 39°35'14,76" W, 1914m (6 ind); Amb4 E9, 20°35'50,48" S 38°27'7,64" W, 2494m (2 ind); Amb4 E10, 20°49'23,58" S 38°17'11,07" W, 2997m (1 ind); Amb4 F6, 19°52'56,9" S 38°35'8,88" W, 1022m (34 ind); Amb4 F7, 20°4'8,18" S 38°31'27,32" W, 1302m (17 ind); Amb4 F8, 20°16'35,72" S 38°27'18,98" W, 1902m (7 ind); Amb4 F9, 20°29'3,31" S 38°23'15,5" W, 2504m (2 ind); Amb5 A6, 21°4'43,84" S 40°8'31,76" W, 1024m (5 ind); Amb5 A7, 21°4'51,67" S 40°4'14,88" W, 1300m (5 ind); Amb5 A8, 21°6'30,57" S 39°38'36,43" W, 1889m (2 ind); Amb5 A9, 21°9'40,3" S 38°52'25,04" W, 2498m (1 ind); Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (18 ind); Amb5 B7, 20°36'42,03" S 39°49'25,36" W, 1333m (16 ind); Amb5 B9, 20°54'44,14" S 38°56'10,72" W, 2519m (3 ind); Amb5 C6, 20°15'36,86" S 39°46'15,05" W, 1040m (8 ind); Amb5 C7, 20°17'41,07" S 39°42'38,02" W, 1358m (5 ind); Amb5 C8, 20°25'16,2" S 39°27'20" W, 1918m (3 ind); Amb5 D6, 19°50'1,87" S 39°26'30,04" W, 1055m (13 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 1335m (9 ind); Amb5 D8, 20°8'42,82" S 39°7'29,5" W, 1922m (2 ind); Amb5 D9, 20°34'36,32" S 38°40'53,6" W, 2460m (3 ind); Amb5 E8, 20°15'59,97" S 38°40'53,86" W, 1887m (3 ind); Amb6 D5, 19°46'34,99" S 39°30'4,65" W, 428m (16 ind);

418

Amb6 CANWN5, 19°49'37,21" S 39°35'41,25" W, 410m (1 ind); Amb6 CANWN6, 19°53'31,53" S 39°32'56,35" W, 1023m (47 ind); Amb6 CANWN7, 19°58'11,44" S 39°31'38,29" W, 1300m (10 ind); Amb8 E7, 19°47'5,96" S 39°3'11,96" W, 1258m (2 ind); Amb8 G6, 19°3'32,9" S 37°49'4,82" W, 988m (15 ind); Amb8 G7, 19°3'29,3" S 37°48'39,27" W, 1302m (8 ind); Amb8 G8, 19°3'45,82" S 37°47'28,26" W, 1928m (4 ind); Amb11 A6, 21°4'37,64" S 40°8'32,68" W, 1015m (3 ind); Amb11 A7, 21°4'43,08" S 40°4'12,96" W, 1295m (7 ind); Amb11 A8, 21°6'38,26" S 39°38'31,44" W, 1889m (3 ind); Amb11 A9, 21°9'39,38" S 38°52'7,25" W, 2502m (1 ind); Amb11 B6, 20°36'1,61" S 39°51'39,15" W, 1003m (10 ind); Amb11 B7, 20°36'48,64" S 39°49'32,61" W, 1324m (6 ind); Amb11 B8, 20°41'33,93" S 39°35'22,06" W, 1908m (4 ind); Amb11 C6, 20°15'32,18" S 39°46'12,38" W, 1040m (6 ind); Amb11 C7, 20°17'37,38" S 39°42'36,72" W, 1355m (1 ind); Amb11 C9, 20°48'37,26" S 38°45'28,85" W, 2465m (1 ind); Amb11 D8, 20°8'45,23" S 39°7'31,74" W, 1926m (2 ind); Amb11 D9, 20°34'41,91" S 38°40'57,73" W, 2496m (1 ind); Amb11 D10, 20°53'28,94" S 38°21'21,75" W, 3000m (1 ind); Amb11 E8, 20°15'55,63" S 38°40'45,57" W, 1892m (3 ind); Amb11 E9, 20°35'51,99" S 38°27'13,04" W, 2501m (2 ind); Amb11 F6, 19°52'52,59" S 38°35'10,48" W, 1021m (19 ind); Amb11 F7, 20°4'9,68" S 38°31'29,01" W, 1295m (12 ind); Amb11 F8, 20°16'38,17" S 38°27'26,52" W, 1904m (6 ind); Amb11 F9, 20°29'3,85" S 38°23'18,56" W, 2507m (2 ind); Amb12 D6, 19°50'6,01" S 39°26'34,62" W, 1050m (16 ind); Amb12 D7, 19°54'4,77" S 39°22'29,46" W, 1335m (5 ind); Amb12 E6, 19°40'1,46" S 39°7'21,99" W, 1050m (21 ind); Amb12 E7, 19°47'2,44" S 39°3'14,62" W, 1250m (4 ind); Amb12 G7, 19°3'30,62" S 37°48'46,66" W, 1347m (9 ind); Amb12 G8, 19°3'39,78" S 37°47'39,35" W, 1867m (2 ind); Amb12 G9, 19°3'10,76" S 37°45'34,37" W, 2770m (4 ind); Amb12 CAND6, 19°37'45,14" S 39°3'58,75" W, 1050m (6 ind); Amb12 CANWN6, 19°53'27,27" S 39°32'59,82" W, 970m (38 ind); Amb12 CANWN7, 19°58'12,82" S 39°31'42,22" W, 1316m (22 ind).

Description: A medium-sized Spiophanes species, largest complete specimen 15 mm long, 1.3 mm wide at the widest point for 44 chaetigers (incomplete), holotype 13 mm long, 1.1 mm wide at the widest point for 40 chaetigers (incomplete). Body slender and slightly dorsoventrally flattened on chaetigers 1–4 and considerably flattened on remaining chaetigers, tapering towards posterior region and divided in three regions: anterior region from chaetiger 1 to chaetiger 4, middle region from chaetiger 5 to chaetiger 14 and posterior region from chaetiger 15 onwards. Parapodia from chaetigers 1–4 shifted dorsally. Parapodia fleshy and well developed on anterior region, short and fleshy on middle region and greatly reduced from chaetiger 16 onwards. Body color whitish in alcohol, with brownish pigment at the base of notopodia from middle region (Fig 11A–B).

419

Prostomium narrow and short (quadrangular), truncate anteriorly and bearing short and angular anterolateral horns. Prostomium extending posteriorly to the posterior margin of chaetiger 1 (Fig 11A, 12A–B). Continuous nuchal organs reaching posterior margin of chaetiger 10 (Fig 12A). Eyes absent. Occipital antenna long and thin, near the posterior margin of chaetiger 1 (Fig 12B). Peristomium weakly developed, reduced to a ring around the mouth, surrounding prostomium for about half of its length, not forming lateral wings and not fused to chaetiger 1 (Fig 11A, 12B). Palps lost in all specimens.

On anterior region, notopodial lamellae cirriform and gradually tapered on notopodia and foliaceous with cirriform tip on neuropodia. From chaetiger 5 to chaetiger 8, notopodial lamellae oval and well-developed, foliaceous and bearing long and delicate dorsal projection from chaetiger 9 onwards (Fig 11, 12A, C). Chaetal spreader not observed. Opening of glandular organs not observed. Neuropodial lamellae rounded and reduced on chaetigers 5–6 and absent from chaetiger 7 onwards. Prechaetal lamellae absent on both rami. Very low dorsal crests from chaetigers 16–17 to end of fragment (chaetiger 30) (Fig 11A, 12C). Interneuropodial pouches present from chaetiger 15, progressively smaller and more delicate towards end of the fragment (Figs 11B, 12C).

Chaetae from anterior region organized in two rows, granulated and narrowly bilimbate capillaries on both rami (Fig 13A), as well as 1–2 robust crook-like chaetae on neuropodium of chaetiger 1 (Fig 13B). On middle region, chaetae organized in three rows of granulated and narrowly bilimbate capillaries on notopodia (Fig 13C) – anterior row longest and posterior row shortest, and two rows of short, narrowly bilimbate and non-granulated capillaries on neuropodia (Fig 13D). Neuropodial capillaries from chaetigers 9–14 wide and bearing a twisted tip (Figs 12D, 13E). Bacillary chaetae present on chaetigers 5–7 (Fig 13F). On posterior region, non-limbate and non-granulated capillaries on notopodia (Fig 13G), gradually becoming thinner and less numerous (Fig 13H) and solely hooded hooks in neuropodia (Fig 13I) (described below). Morphology of posterior-most chaetae unknown.

Hooks in neuropodia only, consistently starting from chaetiger 15, up to five per fascicle, not accompanied by capillaries. Hooks short and slightly constricted near the tip. Hooks quadridentate, with single unpaired tooth immediately above main tooth and upper-most pair of minute secondary teeth arranged side by side (Fig 13I). Hood completely absent (Fig 12E–F). Sabre chaetae consistently starting from chaetiger 4, up to two per rami. Sabre chaetae non- limbate, bearing numerous transversal columns (Figs 12E, 13J).

Pygidium of unknown morphology.

Oocytes from chaetiger 15, measuring up to 230 µm.

420

Glandular organs large, from chaetiger 5 to chaetiger 7 (especially on chaetigers 5–6).

Methyl green staining pattern: Prostomium, ventral side of anterior and middle regions and notopodial lamellae from middle region strongly stained; pigmentation weakly diffused throughout whole body.

Remarks. Spiophanes profundus sp. nov. differs from all remaining Spiophanes species recorded in Brazil in having interneuropodial pouches, a character present only in S. kroeyeri Grube,1860, S. kimballi Meiβner, 2005, S. afer Meiβner, 2005, S. viriosus Meißner &Hutchings, 2003, S. algidus Meiβner, 2005, S. fimbriata Moore,1923 and S. mediterraneus Meiβner, 2005 (Meiβner & Hutchings 2003; Meiβner 2005). The presence of neurochaetae on chaetigers 9–14 with a twisted tip (Fig. 13E) is also unique to Spiophanes profundus sp. nov.

The species can be promptly distinguished from S. viriosus, S. fimbriata, S. afer and S. mediterraneus in having interneuropodial pouches starting from chaetiger 15, instead of chaetiger 14. As for S. kimballi, which may present interneuropodial pouches from chaetiger 15, interneuropodial pouches are present only on some mature specimens from greater depths and may start after chaetiger 18 or develop on only one side of the body (Meiβner & Hutchings 2003; Meiβner 2005).

Besides Spiophanes profundus sp. nov., both S. kroeyeri and S. algidus also possess interneuropodial pouches starting from chaetiger 15, although species can be distinguished by prostomium shape – quadrangular in S. profundus sp. nov., bell-shaped in S. kroeyeri and S. algidus; peristomium – weakly developed in S. profundus sp. nov., moderately developed in S. kroeyeri and poorly to moderately developed in S. algidus; shape of notopodial lamellae from anterior region – cirriform in S. profundus sp. nov. and S. algidus and foliaceous in S. kroeyeri; shape of notopodial lamellae from middle and posterior regions – oval and well-developed on chaetigers 5–8, foliaceous and bearing long and delicate dorsal projection from chaetiger 9 onwards in S. profundus sp. nov., rounded on chaetigers 5–8 and with broad triangular base and slender tip from chaetiger 9 onwards in S. kroeyeri and rounded on chaetigers 5–8, cirriform to subulate on chaetigers 9–14, basally bulbous on chaetigers 15–18 and subulate with slender tip from chaetiger 19 onwards in S. algidus; shape of neuropodial lamellae from anterior region – foliaceous with cirriform tip in S. profundus sp. nov., foliaceous, tapering to subulate in S. kroeyeri and subulate to subtriangular, with slender tip on chaetigers 1–2 in S. algidus and lack of dorsal crests in S. profundus sp. nov., present in S. kroeyeri and S. algidus (Meiβner, 2005).

Etymology. The species epithet, profundus, refers to the depths where this species was collected – only on the continental slope and most records from depths greater than 1000 m.

Habitat: medium silt to clay, 410–3000 m depth.

421

Distribution: Southeastern Brazil (Espírito Santo state), Atlantic Ocean.

Identification key to Spiophanes species recorded in this work.

1. Interneuropodial pouches present……...... ……...Spiophanes profundus sp. nov.

- Interneuropodial pouches absent...... 2

2(1). Antenna absent………………………………………….………………..Spiophanes duplex

- Antennae present………………………………………………………………………………..3

3(2). Prostomium narrow, truncated anteriorly and bearing fleshy anterolateral horns; hooks lacking hoods………………………………………………………Spiophanes aureus sp. nov.

- Prostomium bell-shaped, bearing thin anterolateral projections; hooks with hoods……………………………………………………………...….Spiophanes grubei sp. nov.

Acknowledgments

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Legends of illustrations

Figure 1. Map showing sampled area (blue polygon).

Figure 2. Spiophanes grubei sp. nov., anterior chaetigers, dorsal view (MNRJP XXXX, holotype), showing characteristic pigmentation in ethanol. Abbreviations: cr, crook-like chaetae; pe, peristomium; pr, prostomium.

Figure 3. SEM of Spiophanes grubei sp. nov. A. Anterior, middle and posterior regions (in part), dorsal view. B. Anterior and middle regions (in part), dorsal view. C. Middle region, lateral view. D. Posterior region, dorsolateral view. E. Neuropodia from posterior region, dorsal view. F. Hooded hooks, dorsal view. Abbreviations: an, antenna; bc, bacillary chaetae; car, caruncle; dc, dorsal crest; go, glandular opening; hh, hooded hook; pe, peristomium; pr, prostomium; prob, proboscis; sa, sabre chaetae.

Figure 4. Spiophanes grubei sp. nov. A. Chaetae from anterior region. B. Crook-like chaetae from neuropodia of chaetiger 1. C. Elongated capillary from chaetiger 2. D. Notochaetae from middle region. E. Neurochaetae from middle region. F. Bacillary chaetae. G. Notochaetae from posterior region. H. Neuropodial hooded hook. I. Sabre chaetae. J. Modified notochaetae from posterior- most chaetigers.

Figure 5. Spiophanes duplex, anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: bc, bacillary chaetae; pe, peristomium; pr, prostomium.

Figure 6. SEM of Spiophanes duplex. A. Anterior, middle and posterior regions (in part), dorsal view. B. Anterior and middle regions (in part), dorsal view. C. Close-up of anterior region, dorsal view. D. Middle region, lateral view. E. Posterior region, dorsolateral view. F. Posterior

427 neuropodia, lateral view. Abbreviations: bc, bent chaetae; car, caruncle; cr, crook-like chaetae; go, glandular opening; ho, hook; pe, peristomium; pr, prostomium; ps, palp scar; sa, sabre chaetae.

Figure 7. Spiophanes duplex. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Crook-like chaetae from neuropodia of chaetiger 1. D. Notochaetae from middle region. E. Neurochaetae from middle region. F. Bacillary chaetae. G. Notochaetae from posterior region. H. Non-limbate notochaetae from posterior region. I. Neuropodial hook. J. Sabre chaetae. K. Modified notochaetae from posterior-most chaetigers.

Figure 8. Spiophanes aureus sp. nov., MNRJP XXXX, holotype, showing characteristic pigmentation in ethanol. A. Anterior chaetigers and mid-body, dorsal view. B. Anterior chaetigers and mid-body, lateral view. Abbreviations: pe, peristomium; pr, prostomium.

Figure 9. SEM of Spiophanes aureus sp. nov. A. Anterior, middle and posterior regions (in part), dorsal view. B. Anterior and middle regions (in part), dorsal view. C. Close-up of anterior region, dorsal view. D. Middle region, lateral view. E. Close-up of parapodium from middle region, lateral view. F. Posterior neuropodia, lateral view. Abbreviations: an, antenna; car, caruncle; cr, crook-like chaetae; go, glandular opening; ho, hook; pr, prostomium; prob, proboscis; sa, sabre chaetae.

Figure 10. Spiophanes aureus sp. nov. A. Notochaetae from anterior region. B. Neurochaetae from anterior region. C. Non-limbate notochaetae from chaetiger 1. D. Crook-like chaetae from neuropodia of chaetiger 1. E. Notochaetae from middle region. F. Neurochaetae from middle region. G. Neurochaetae from chaetigers 9–14. H. Bacillary chaetae. I. Notochaetae from posterior region. J. Non-limbate notochaetae from posterior region. K. Neuropodial hook. L. Sabre chaetae.

Figure 11. Spiophanes profundus sp. nov., MNRJP XXXX, holotype, showing characteristic pigmentation in ethanol. A. Whole fragment, dorsal view. B. Middle and posterior regions (in part), ventral view. Abbreviations: ip, interneuropodial pouch; pe, peristomium; pr, prostomium.

Figure 12. SEM of Spiophanes profundus sp. nov. A. Anterior, middle and posterior regions (in part), dorsal view. B. Anterior and middle regions (in part), dorsal view. C. Posterior region (in part), dorsolateral view. D. Parapodia from middle region, dorsolateral view. E. Neuropodium from posterior region, dorsal view. F. Hook, dorsal view. Abbreviations: an, antenna; ca1; capillary from chaetiger 1; car, caruncle; dc, dorsal crest; ho, hook; ip, interneuropodial pouch; pr, prostomium; sa, sabre chaetae; tc, twisted tip chaetae.

Figure 13. Spiophanes profundus sp. nov. A. Chaetae from anterior region. B. Crook-like chaetae from neuropodia of chaetiger 1. C. Notochaetae from middle region. D. Neurochaetae from middle region. E. Neurochaetae from chaetigers 9–14. F. Bacillary chaetae. G. Notochaetae from

428 posterior region. H. Non-limbate notochaetae from posterior region. I. Neuropodial hook. J. Sabre chaetae.

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Tables

Table 1. Comparison among Spiophanes species examined in this work.

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Table 1 (continued). Comparison among Spiophanes species examined in this work.

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Capítulo 8

Polydorids (Annelida: Spionidae) species from southern Brazil, with new records and a new Boccardia species

ANTÔNIO JOÃO MALAFAIA PEIXOTO & PAULO CESAR DE PAIVA

Revista alvo: Zootaxa Status: A submeter

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Polydorids (Annelida: Spionidae) species from southern Brazil, with new records and a new Boccardia species

Running title: NEW POLYDORIDS FROM BRAZIL

ANTÔNIO JOÃO MALAFAIA PEIXOTO1,2,3 & PAULO CESAR DE PAIVA1,2

3 Corresponding author. E-mail: [email protected].

Abstract Seven Polydorid species have been reported to Espírito Santo Basin, southeastern Brazil, from both shallow and deep-water environments. Six of these species are already known to science– Dipolydora armata, D. barbilla, D. blakei, D. socialis, D. tetrabranchia and D. cf. caulleryi, while Boccardia capixaba sp. nov. represents a new species. This study represents the first record of D. cf. caulleryi to the Brazilian coast and the first record of D. tetrabranchia to southeastern Brazil. Species can be distinguished based on prostomium morphology, caruncle length, branchiae distribution, morphology of spines from chaetiger 5, presence of posterior modified chaetae and pygidium morphology. An identification key is provided to Dipolydora species recorded in Espírito Santo basin.

Keywords: Polydorid, Annelida, morphology, Polydora-complex.

Introduction

The annelid family Spionidae Grube, 1850 is one of the most diverse and abundant marine benthic groups, capable of dominating soft-bottom sediments, particularly in organically rich sediments, whether naturally occurring or eutrophic. Worldwide, there are 39 valid genera and approximately 580 described species (Blake et al. 2017) however, spionids have a known history of species-complexes (Sato-Okoshi et al. 2016), species with suspicious cosmopolitan status and genera that never were revised. Furthermore, the deep-sea, a poorly known environment (Spalding et al. 2007) corresponds to an environment where spionids are often the dominant taxa (Blake et al. 2017).

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To the Brazilian coast, most spionid taxonomic studies are focused on the shallow continental shelf, on depths above 50 m (except for Paiva & Barroso 2010; Peixoto & Paiva 2017, 2019, ms) and are geographically restricted to southeastern and southern Brazil (Lana et al. 2017), while, based on data compiled by Amaral et al. (2013) only a few studies are known from the Northeast coast (checklists and ecological works) and no study is known from the North coast of Brazil. A few genera have been studied in greater detail: Laonice Malmgren, 1867 (Kinberg 1866; Nonato et al. 1986; Radashevsky & Lana 2009), Laubieriellus Maciolek, 1981 (Peixoto & Paiva 2019), Prionospio Malmgren, 1867 (Peixoto & Paiva 2019), Scolelepis Blainville, 1828 (Rocha et al. 2009; Rocha & Paiva 2012), Spiogalea Aguirrezabalaga & Ceberio, 2005 (Peixoto & Paiva 2017) and Trochochaeta Levinsen, 1884 (Radashevsky et al. 2018).

There only a small number of studies focusing on multiple genera, for instance Bolívar & Lana (1987) to the continental shelf of the state of Paraná, southern Brazil, Paiva & Barroso (2010) to the continental slope of Campos Basin, state of Rio de Janeiro, southeastern Brazil, as well as and Pardo et al. (2006) and Pardo & Peixoto (in press) to southern and southeastern Brazil. It is important to notice, however, that none of these studies described new species.

The term “Polydorid” (sometimes referred as Polydorin or Polydora-complex) refers to members of the subfamily Spioninae Söderström, 1920 that possess a modified fifth chaetiger. This group includes the following genera: Amphipolydora Blake, 1983, Boccardia Carazzi, 1893, Boccardiella Blake & Kudenov, 1978, Carazziella Blake & Kudenov, 1978, Dipolydora Verrill, 1881, Polydora Bosc, 1802, Polydorella Augener, 1914, Pseudopolydora Czerniavsky, 1881 and Tripolydora Woodwick, 1964.

The genus Dipolydora was originally described by Verrill (1881) to include Dipolydora concharum (Verrill, 1880), described as a Polydora species. Although the genus was erected in 1881, no other species were attributed to Dipolydora until the genus was synonymized with Polydora by Fauchald (1977) without further explanation. The genus was revalidated by Blake (1996), who reviewed the genus Polydora and transferred several Polydora species to Dipolydora.

In his review, Blake (1996) organized Dipolydora species in five groups of morphologically similar species: A) group of Dipolydora giardi (Mesnil, 1893); B) group of D. concharum; C) group of D. barbilla (Blake, 1981); D) group of de D. armata (Langerhans, 1880) and E) group of D. commensalis (Andrews, 1891). This organization, although convenient, was not adopted nor mentioned in posterior works. The main characters to separate Dipolydora form Polydora include the presence of notochaetae on chaetiger 1; bidentate hooded hooks lacking constriction or manubrium; hooks with main fang forming a wide angle with the shaft and a narrower, acute angle with apical tooth; pygidium bearing lobes of various shapes and presence, at least in some

447 species, of a muscular enlargement of the digestive tract (gizzard). Around 43 Dipolydora species are known, despite many species remain poorly studied concerning their biology and even morphology.

The genus Boccardia was described by Carazzi (1893), differing from the remaining genus of the subfamily Spioninae in having branchiae anterior to chaetiger 5, bidentate hooded hooks lacking constriction and bearing two types of modified spines on chaetiger 5. The genus Boccardiella is similar to Boccardia in having branchiae anterior to chaetiger 5 and bidentate hooded hooks lacking constriction, but these genera can be distinguished based on the morphology of the modified spines on chaetiger 5, two types of spines – smooth spines and expanded apex spines in Boccardia and only one type of spines – smooth spines in Boccardiella. Boccardia is one of the most specious spionid genus, with around 24 described species, among shell-boring and tube- dwelling species. Most species were described from the Pacific and Indian Oceans (Blake et al. 2017), except for Boccardia semibranchiata Guerin, 1990, from the Coast of France, Mediterranean Sea and B. salazari Delgado-Blas, 2008, from Mexico, Caribbean Sea, besides accidental introductions of B. proboscidea in Europe (Martínez et al. 2006), Africa (Simon et al. 2010) and South America (Jaubet et al. 2014).

The Polydorids are one of the more studied groups in Brazil, with the description of Polydora carinhosa Radashesvky et al. (2006) and Pseudopolydora rosebelae (Radashevsky & Migotto, 2009), as well as records of Dipolydora armata (Radashevsky & Nogueira 2003), D. socialis (Schmarda, 1861), D. barbilla, D. blakei (Maciolek, 1984), Polydora ecuadoriana Blake, 1983, P. cf. haswelli Blake & Kudenov, 1978, P. rickettsi Woodwick, 1961, P. hoplura Claparède, 1868 (Radashesvky et al. 2006; Radashevsky & Migotto 2017), P. cornuta (Bosc, 1802), P. neocaeca Williams & Radashesky, 1999, P. nuchalis Woodwick, 1953, P. colonia Moore, 1907, P. websteri Hartman in Loosanoff & Engle, 1943, Boccardia polybranchia (Haswell, 1885), Boccardia proboscidea Hartman, 1940, Boccardiella hamata (Webster, 1879), B. ligerica (Ferronière, 1898), Pseudopolydora achaeta Radashevsky & Hsieh, 2000, P. antennata (Claparède, 1869) and P. primigenia Blake, 1983 (Pardo et al. 2006; Amaral et al. 2013; Pardo & Peixoto in press). However, researchers should use these records with caution, since some species reported to Brazil were described to distinct biogeographical provinces or are part of known species-complexes. The genus Amphipolydora, Polydorella and Tripolydora have not been recorded in Brazil so far.

Six Dipolydora species and one Boccardia species were recorded in this work, from material collected on Espírito Santo basin (southeastern Brazil), during the oceanographic cruises of the “Marine Environmental Characterization of Espírito Santo Basin and Northern Portion of Campos Basin” (AMBES) Project, coordinated by CENPES/PETROBRAS (Research Center of the Brazilian Energy Company). Only one species – Boccardia capixaba sp. nov. is new to science,

448 while Dipolydora cf. caulleryi is reported to Brazil for the first time, while D. tetrabranchia is reported to southeastern Brazil.

Material and Methods During the oceanographic cruises of the AMBES Project on the Espírito Santo Basin and northern portion of Campos basin between 2010 and 2013, sediment samples were collected using Van Veen Grab sampler on the continental shelf and a box corer in the continental slope (Fig 1). There were two sampling campaigns: summer season (sample codes Amb1, Amb3, Amb4, Amb5, Amb6, and Amb7) and winter season (sample codes Amb2, Amb11, Amb12, Amb13, and Amb14).

Sediment samples were fixed in situ in a 10% formalin-seawater solution without prior use of relaxing agents, taken to the laboratory, washed in freshwater and kept in 70% EtOH. Samples were then elutriated using a 500-µm-mesh sieve (for samples collected on the continental shelf and on the Doce River mouth) or a 300-µm-mesh sieve (for samples collected on the continental slope and on the Watu Norte and Doce canyons) and sorted out. Afterwards, specimens were examined under a Leica S8 APO and Nikon Eclipse E200 microscope and measured using a calibrated eyepiece attached to the microscope.

In order to reveal species-specific patterns and facilitate visualization of morphological features such as caruncle length and branchial scars, specimens were stained in a methyl green solution (1 g/L in 70% EtOH) for 10‒30s and examined in distilled water to slow stain fading. Since specimens were kept in formalin for several months before being transferred to ethanol, molecular studies were not possible. For Dipolydora species, main taxonomic characters are summarized in Table 1 and an identification key is provided for species recorded in this study, while for Boccardia, a comparison table using species previously recorded in Brazil by Amaral et al. (2013) is provided (Table 2).

Specimen lots deposited in the Smithsonian National Museum of National History (USNM), Smithsonian Institution, Washington, D.C., USA were examined and compared with material collected in this study. Type material of Dipolydora barbilla USNM 58978 (5 paratypes), as well as non-type material of Dipolydora cf. armata USNM 73652 (36 specimens, non-type), D. blakei USNM 81928 (20 specimens; non-type material), D. socialis USNM 187537 (1 specimen, non- type), D. tetrabranchia USNM 53839 (12 specimens, non-type material) and Boccardia polybranchia USNM 67602 (35 specimens, non-type material) were also examined.

Prior to scanning electron microscope (SEM) studies, specimens were dehydrated in an ascending ethanol series (70% to 100% EtOH, 15 min each, followed by three changes of 100% EtOH), then

449 critical-point dried using CO2. Alternatively, specimens were dehydrated in an ascending ethanol series, transferred to an ascending hexamethyldisilane (HMDS) series (2 parts 100% EtOH:1-part HMDS, 1:1, 1:2, pure HMDS, followed by a second change of pure HMDS, 15 min each), and dried in a fume hood. Specimens were then mounted on stubs, sputter coated with gold and viewed in a JEOL JSM-6510 LV SEM or a JEOL JSM-6390 LV SEM.

Type material of Boccardia capixaba sp. nov. was deposited in the Museu Nacional do Rio de Janeiro (MNRJP), Universidade Federal do Rio de Janeiro and in the Museu de Zoologia (MZUSP), Universidade de São Paulo, located in Brazil, whereas non-type material from all species will be deposited in the Museu de Zoologia Adão José Cardoso (ZUEC-POL), Universidade Estadual de Campinas.

Results

Taxonomy

Phylum Annelida Lamarck, 1809

Order Spionida Fauchald, 1977

Family Spionidae Grube, 1850

Genus Dipolydora Verrill, 1881

Type species: Polydora concharum Verrill, 1880, designated by Verrill (1881).

Synonym list: Leipoceras Möbius, 1874 (subjective synonym).

Diagnosis (from Blake et al. 2017): Prostomium entire or incised anteriorly, extending posteriorly as caruncle; eyespots present or absent. Chaetiger 1 with notochaetae. Chaetiger 5 modified, with major spines of one type, with or without accompanying companion chaetae; spines arranged in single curved row; spines simple, falcate, with lateral flanges, teeth and/or apical bristles with superior notochaetae dorsal to modified spines and companion chaetae. Posterior notopodial spines present or absent. Neuropodial hooded hooks bidentate, usually with recurved shaft without constriction or manubrium, main fang forming wide angle with shaft and narrow, acute

450 angle with apical tooth; hooks first present from chaetiger 7–17. Pygidium disclike, cuff-shaped, with 2, 3, or 4 lobes of various forms, or with four or more small papillae. Anterior part of digestive tract sometimes with enlarged, thick, gizzard-like structure.

Dipolydora armata (Langerhans, 1880)

(Figs 2–4E)

Synonym list:

Dipolydora rogeri (Martin, 1996) (recombination of subjective synonym)

Polydora armata Langerhans, 1880 (superseded original combination)

Polydora monilaris Ehlers, 1904 (subjective synonym)

Polydora rogeri Martin, 1996 (subjective synonym)

Material examined. Amb7 F3, 18°53'29,72" S 39°6'23,3" W, 52m (8 ind).

Comparative material examined. Dipolydora cf. armata USNM 73652 (36 specimens, non- type).

Description: a medium-sized Dipolydora species, largest specimen, 3 mm long, 0.25 mm wide for 35 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region, tapering towards the pygidium. Pigmentation patterns absent. Body color light yellow to whitish in alcohol (Fig 2).

Prostomium narrow, with shallow anterior incision, extending posteriorly as a low caruncle to the end of chaetiger 2. Eyes absent. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 2, 3A–B). Grooved palps reaching up to chaetiger 12, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami– especially on the notopodium; shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on both rami, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent (Fig 3B).

Chaetiger 5 modified, up to 2.5 times longer than chaetigers 4 or 6 (Figs 2, 3C). Presence of 2–3 dorsal geniculated chaetae (Fig 4A), curved row of 2–3 spines (Fig 4B) and ventral row of 2–4 bilimbate capillaries (Fig 4C). Companion chaetae (pennoned chaetae) absent. Spines bearing

451 main fang, lateral tooth and flattened accessory flange; teeth bases covered in fine bristles (Figs 3C, 4C).

Notopodial postchaetal lamellae rounded on chaetigers 2–4 and absent from chaetiger 5 onwards. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae rounded on chaetigers 2–4 and absent from chaetiger 5 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and narrowly unilimbate capillaries. Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, narrowly unilimbated (barely perceptible), thinner, and less numerous. Notopodial posterior spines present from chaetiger 27–28 almost to the end of the body, inserted in the notopodia or protracted, 7–20 spines per chaetiger. Spines wide, smooth and with pointed tip (Figs 3D, 4D), accompanied by 3–5 long non-limbate capillaries.

Hooks in neuropodia only, consistently starting from chaetiger 7, up to four per fascicle, accompanied by 1–2 long non-limbate capillaries throughout the entire body. Hooks bidentate, slightly curved near the distal end, lacking constriction (Fig 4E).

Branchiae starting from chaetiger 7, 7–9 pairs (depending on specimen size). Branchiae smooth and slightly flattened, longest on chaetigers 9–10 and gradually reducing in length towards last branchial pair (Fig 3A–B).

Pygidium bilobed.

Methyl green staining pattern: Prostomium, ventral part of peristomium, ventral side of chaetigers 6–11 and region above notopodia from chaetigers 12–24 strongly stained.

Remarks: Dipolydora armata was originally described from Madeira Island (Portugal) as Polydora armata Langerhans, 1880, being posteriorly transferred to Dipolydora by Blake (1996) and repeatedly recorded on tropical and subtropical areas, despite that, according to Radashevsky & Nogueira (2003), many older records cannot be verified. Blake (1983) considers that this species corresponds to two distinct species – a typical form found in tropical and subtropical areas bearing a bristled hood connecting the main fang and accessory tooth on the spines from chaetiger 5 and thickened notopodial posterior spines arranged in a rosette; and a second form found in colder (and possibly deeper) waters, with spines from chaetiger 5 curved and bearing flattened tips, besides delicate notopodial posterior spines in fan-like arrangement.

Dipolydora armata is similar to D. cf. caulleryi in having a shallow incised prostomium, caruncle extending to the posterior margin of chaetiger 2, smooth posterior notopodial spines and hooded

452 hooks accompanied by capillaries throughout the body. However, species can be separated based on branchial distribution (7–9 pairs in D. armata and present on about two thirds of the specimen length in D. cf. caulleryi), starting chaetiger of notopodial posterior spines – starting on chaetigers 27–28 in D. armata and on chaetigers 16–21 in D. cf. caulleryi), morphology of spines from chaetiger 5 – bearing main fang, lateral tooth and flattened accessory flange in D. armata and curved and bearing distinctive crest of bristles in D. cf. caulleryi, and lack of companion chaetae in D. armata (Blake 1996; Radashevsky & Nogueira 2003).

The species is also similar to D. blakei in having a narrowly incised prostomium, similar branchial distribution (7–9 pairs in D. armata and 6–9 pairs in D. blakei), smooth posterior notopodial spines, lack of companion chaetae on chaetiger 5 and hooded hooks accompanied by capillaries throughout the body, but species can be distinguished based on caruncle morphology – extended laterally in D. blakei, starting chaetiger of notopodial posterior spines – starting on chaetigers 27– 28 in D. armata and on chaetigers 17–23 in D. blakei, as well as spine morphology from chaetiger 5 – bearing main fang, lateral tooth and flattened accessory flange in D. armata and possessing pointed main fang and large posterolateral tooth in D. blakei (Maciolek 1984).

Dipolydora armata differs from all Brazilian Dipolydora species – and from most Dipolydora species in lacking companion chaetae (pennoned chaetae) alternating with spines on chaetiger 5.

Habitat: Coarse silt, 52 m depth.

Distribution: Dipolydora armata is considered an amphitropical species. Atlantic Ocean: Belize, Brazil (São Paulo, Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states), Portugal – Madeira Island (type locality), Mexico; Mediterranean Sea: Italy, Spain; Indian Ocean: Australia and the Pacific Ocean: Taiwan, Vietnam, New Zealand, Thailand, Ecuador, Chile, Japan and USA (Hawaii and California).

Dipolydora barbilla (Blake, 1981)

(Figs 5–7F)

Synonym list:

Polydora barbilla Blake, 1981 (superseded original combination).

Material examined. Amb1 Foz14, 19°42'32,21" S 39°38'57,36" W, 38m (1 ind); Amb1 Foz15, 19°37'48,27" S 39°35'25,83" W, 41m (3 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (1 ind); Amb7 B3, 20°34'53,42" S 40°6'27,43" W, 50m (1 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (1 ind); Amb14 B3, 20°34'53,05" S 40°6'27,68" W, 49m (1 ind); Amb14

453

C1, 20°10'2,67" S 40°8'31,96" W, 26m (1 ind); Amb14 C2, 20°11'25,75" S 40°2'15,87" W, 40m (1 ind).

Comparative material examined. Dipolydora barbilla USNM 58978 (5 specimens, paratypes).

Description: a medium-sized Dipolydora species, largest specimen 4 mm long, 0.25 mm wide for 56 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region, tapering towards the pygidium. Pigmentation patterns absent. Body color light yellow to whitish in alcohol (Fig 5).

Prostomium narrow, incised anteriorly, extending posteriorly as a low caruncle to the end of chaetiger 3. One pair of eyes or eyes absent. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 5, 6A–B). Grooved palps reaching up to chaetiger 12, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami – especially on the notopodium; shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on both rami, smaller than lamellae on succeeding chaetigers, especially on the neuropodium. Prechaetal lamellae absent.

Chaetiger 5 modified, twice as long as chaetigers 4 or 6 (Figs 5, 6C–D). Presence of 3–6 dorsal geniculated chaetae (Fig 7A), curved row of 3–5 spines (Fig 7B) alternated with bilimbate companion chaetae (pennoned chaetae) (Fig 7C) and ventral row of 1–3 bilimbate capillaries (Fig 7D). Spines falcate, bearing bristled collar on the convex side (Fig 6D).

Notopodial postchaetal lamellae digitiform on chaetigers 2–4, absent from chaetiger 5 onwards. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae digitiform on chaetigers 2–4, absent from chaetiger 5 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and bilimbate capillaries, anterior row of short and posterior row of long (up to two times longer than anterior row) capillaries. Towards the posterior region, capillaries progressively become elongate, narrowly bilimbated, thinner, and less numerous. Presence of 2–5 notopodial spines starting from chaetigers 28–40, present to almost end of the body, inserted in the notopodia or protracted. Spines bearing minute barbs (Figs 6F, 7E) and accompanied by 3–5 long non-limbate capillaries.

Hooks in neuropodia only, consistently starting from chaetiger 7, up to five per fascicle, accompanied by 1 – 3 long non-limbate capillaries present only on the first 3–6 chaetigers after the appearance of the hooded hooks. Hooks bidentate, slightly curved near the distal end, lacking constriction (Fig 7F). Hook shaft elongated on midbody, progressively shorter towards the posterior region.

454

Branchiae starting from chaetiger 9–10, up to 35 pairs and restricted to the anterior half of the body (Fig 6B). Branchiae smooth and slightly flattened, longest on chaetigers 25–30, gradually reducing in length towards last branchial pair; afferent and efferent blood vessels easily visible using light microscopy.

Pygidium quadrilobate, dorsal pair of lobes slightly smaller than ventral pair (Fig 6C, E).

Methyl green staining pattern: Prostomium and caruncle weakly stained, dorsum of parapodia from chaetiger 8–10 onwards markedly stained.

Remarks: Specimens examined in this work agree completely with Blake’s (1981) original description based on specimens from the Gulf of California and Maciolek’s (1984) description based on Atlantic specimens, referred as Polydora barbilla. Blake (1996) reestablished Dipolydora, transferred Polydora barbilla to Dipolydora and divided species in five groups according to morphological characters, grouping D. barbilla with D. bidentata (Zachs, 1933), D. langerhansi (Mesnil, 1896), D. pilocollaris (Blake & Kudenov, 1978), while Simon (2011) attributed a fifth species, D. keulderae Simon, 2011 to this group.

Blake (1996) defined this group as lacking constriction on the hooded hooks (which is a character shared among all Dipolydora species), branchiae starting from chaetigers 7–10, lack of occipital tentacle, presence of notopodial chaetae on chaetiger 1 (absent in D. pilocollaris), spines from chaetiger 5 falcate, bearing bristled collar on the convex side and quadrilobated pygidium.

Dipolydora barbilla can be distinguished from D. pilocollaris, D. keulderae and D. bidentata by the starting chaetiger of branchiae, from chaetiger 7 in D. pilocollaris and D. keulderae, from chaetiger 8 in D. bidentata and from chaetigers 9–10 in D. barbilla, besides presence and morphology of posterior notopodial spines, absent in D. pilocollaris and D. keulderae, present as smooth spines in D. bidentata and present as barbed spines in D. barbilla (Simon 2011).

Among species recorded in this work, D. barbilla is most similar to D. socialis in having capillaries accompanying hood hooks for a limited number of chaetigers (3–6 chaetigers in D. barbilla and 5–6 chaetigers in D. socialis), branchiae starting after chaetiger 7 (chaetigers 9–10 in D. barbilla and chaetiger 8 in D. socialis) and broad branchial distribution (up to 35 pairs in D. barbilla and extending for most of the body in D. socialis. Species can be separated based on prostomium morphology – deeply incised in D. socialis, caruncle extension – to the end of chaetiger 3 in D. barbilla and end of chaetigers 4–5 in D. socialis, absence of posterior notopodial spines in D. socialis and pygidium morphology – quadrilobate in D. barbilla and unilobate in D. socialis.

Dipolydora barbilla differs from all Brazilian Dipolydora species – and from most Dipolydora species in having posterior notopodial spines bearing barbs.

455

Habitat: coarse sand to fine sand, 26–171 m depth.

Distribution: Atlantic Ocean: USA and Brazil (Paraná, São Paulo, Rio de Janeiro and Espírito Santo states); Pacific Ocean: Mexico (Gulf of California; type locality).

Dipolydora blakei (Maciolek, 1984)

(Figs 8–10E)

Synonym list:

Polydora blakei Maciolek, 1984 (superseded original combination).

Material examined. Amb3 CAND4, 19°31'51,66" S 39°3'4,04" W, 171m (3 ind); Amb6 D4, 19°45'55,39" S 39°30'25,74" W, 149m (23 ind); Amb6 E4, 19°36'5,17" S 39°10'32,93" W, 153m (41 ind); Amb6 CANWN, 19°49'7,27" S 39°36'8,52" W, 158m (1 ind); Amb7 D4, 19°45'54,56" S 39°30'25,23" W, 144m (10 ind); Amb7 E4, 19°36'4,32" S 39°10'34,07" W, 147m (3 ind); Amb12 D4, 19°45'53,43" S 39°30'25,97" W, 143m (12 ind); Amb12 E4, 19°36'3,57" S 39°10'33,64" W, 143m (5 ind); Amb12 CAND4, 19°31'51,68" S 39°3'4,79" W, 171m (6 ind); Amb14 B4, 20°35'23,09" S 39°55'1,18" W, 146m (6 ind).

Comparative material examined. Dipolydora blakei USNM 81928 (20 specimens; non-type material).

Description: a medium-sized Dipolydora species, largest complete specimen 3.2 mm long, 0.3 mm wide for 39 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region, tapering towards the pygidium (Fig 9C). Pigmentation patterns absent. Body color yellow to whitish in alcohol (Fig 8).

Prostomium narrow, weakly incised anteriorly, caruncle extending laterally to chaetiger 1 and posteriorly to the end of chaetiger 2; triangular-shaped (Fig 9A). Eyes absent. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 8, 9A, D). Grooved palps reaching up to chaetiger 12, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami – especially on the notopodium; shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on both rami, smaller than lamellae on succeeding chaetigers, especially on the neuropodium. Prechaetal lamellae absent.

Chaetiger 5 modified, twice as long as chaetigers 4 or 6 (Figs 8, 9B, D). Presence of 3–5 dorsal geniculated chaetae (Fig 10A), curved row of 2–4 spines (Fig 10B) and ventral row of 3–6 thick

456 and unilimbate capillaries (Fig 10C). Spines falcate, possessing pointed main fang and large posterolateral tooth; convex side of main fang bearing fine bristles (Fig 9E).

Notopodial postchaetal lamellae rounded on chaetigers 2–4 and absent from chaetiger 5 onwards. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae digitiform on chaetigers 2–4, absent on chaetiger 5 onwards, digitiform on chaetiger 6 and absent from chaetiger 7 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and narrowly unilimbate capillaries. Notopodial chaetae up to two times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner, and less numerous. Notopodial posterior spines present from chaetiger 17–23 almost to the end of the body, inserted in the notopodia or protracted, 6–15 spines per chaetiger (Figs 9C, 10D). Spines wide, smooth and with pointed tip, accompanied by 2–7 long non-limbate capillaries (Fig 9F).

Hooks in neuropodia only, consistently starting from chaetiger 7, up to five per fascicle, accompanied by 1–2 non-limbate capillaries throughout the entire body. Hooks bidentate, slightly curved near the distal end from the midbody onwards, lacking constriction (Fig 10E).

Branchiae starting from chaetiger 7, 6–9 pairs (depending on specimen size). Branchiae smooth, slightly flattened, basally fused to the notopodial lamellae, longest on chaetigers 9–10 and gradually reducing in length towards last branchial pair.

Pygidium bilobed, bearing median dorsal gap (Fig 9C).

Juvenile morphology: In juveniles with less than 30 chaetigers, notopodial posterior spines were observed on chaetigers 12–15.

Methyl green staining pattern: Prostomium and caruncle weakly stained, dorsal side from chaetiger 7 onwards, almost to last chaetiger.

Remarks: Specimens examined in this work agree completely with Maciolek’s (1984) description, based on specimens from the Northeastern coast of the USA, which was originally described as Polydora blakei.

Blake (1996) reestablished Dipolydora, transferred Polydora blakei to Dipolydora and divided species in five groups according to morphological characters, grouping D. blakei with D. aciculata (Blake and Kudenov, 1978), D. akaina Blake, 1996, D. armata, D. caulleryi, D. notialis (Blake & Kudenov, 1978) and D. quadrilobata (Jacobi, 1883), while Meiβner et al. (2014) attributed an eighth species, D. paracaulleryi Meiβner et al. 2014 to this group. 457

Blake (1996) defined this group as lacking constriction on the hooded hooks (which is a character shared among all Dipolydora species), branchiae starting from chaetiger 7, presence of notopodial chaetae on chaetiger 1, spines from chaetiger 5 falcate, always covered in fine bristles, pennoned chaetae reduced or absent and large notopodial posterior spines, usually arranged into a rosette.

Among this group, D. blakei is most similar to D. armata (leading Martin [1996] to consider species as a possible synonym) in having a caruncle extending to the posterior margin of chaetiger 2, branchiae of similar distribution, lack of pennoned chaetae, posterior notopodial spines of similar morphology and hooks accompanied by capillaries throughout the body, as discussed in D. armata’s Remarks section. However, species can be separated based on caruncle morphology – triangular in D. blakei and narrow in D. armata, starting chaetiger of notopodial posterior spines and morphology of spines from chaetiger 5, as discussed in D. armata’s Remarks section.

The unusual caruncle shape is also present in D. paracaulleryi, but species can be distinguished by the number and morphology of spines from chaetiger 5, 3–4 spines possessing pointed main fang and large posterolateral tooth in D. blakei and two spines with a lateral flange in D. paracaulleryi; number of geniculated chaetae, 3–6 in D. blakei and two in D. paracaulleryi and pygidium morphology, with two lateral lobes in D. blakei and two dorsal and two ventral lobes in D. paracaulleryi (Maciolek 1984; Meiβner et al. 2014).

The species is also similar to D. caulleryi in having a caruncle of similar extension; smooth and thin posterior notopodial spines and hooks accompanied by capillaries throughout the body, but differ on the caruncle shape, extended laterally in D. blakei; morphology of spines from chaetiger 5 – falcate, possessing pointed main fang and large posterolateral tooth in D. blakei and bearing only the main fang in D. caulleryi; lack of pennoned chaetae in D. blakei and pygidium morphology, with two lateral lobes in D. blakei and two dorsal and two ventral lobes in D. caulleryi.

Dipolydora blakei differs from all Brazilian Dipolydora species – and from most Dipolydora species in having a caruncle extending laterally to chaetiger 1 and posteriorly to the end of chaetiger 2.

Habitat: very fine sand to coarse silt, 143–171 m depth.

Distribution: Atlantic Ocean: USA (Gay Head-Bermuda transect, MA; type locality), Brazil (Rio de Janeiro, Espírito Santo, Sergipe and Alagoas states); Mediterranean Sea: Greece.

458

Dipolydora cf. caulleryi (Mesnil, 1897)

(Figs 11–13F)

Synonym list:

Polydora (Polydora) caulleryi Mesnil, 1897 (superseded combination)

Polydora brachycephala Hartman, 1936 (subjective synonym)

Polydora carazzi McIntosh, 1909 (subjective synonym)

Polydora caulleryi Mesnil, 1897 (superseded original combination)

Material examined. Amb5 B6, 20°36'2,03" S 39°51'35,37" W, 1000m (6 ind); Amb5 D7, 19°54'5,01" S 39°22'20,04" W, 1335m (2 ind).

Description: a large-sized Dipolydora species, largest complete specimen 6.5 mm long, 0.7 mm wide for 42 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region, tapering towards the pygidium. Glandular pouches between notopodia and neuropodia present from chaetiger 6 onwards. Pigmentation patterns absent. Body color whitish in alcohol (Fig 11).

Prostomium narrow, entire or with shallow anterior incision, extending posteriorly as a low caruncle to the end of chaetiger 2. One or two pairs of eyes in trapezoidal arrangement or eyes absent. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 11, 12A–B). Palps reaching up to chaetiger 15 (Fig 12B), lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami– especially on the notopodium; non-limbate and shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on both rami, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Chaetiger 5 modified, up to two times longer than chaetigers 4 or 6 (Figs 11, 12B–C). Presence of 3–8 dorsal geniculated chaetae (Fig 13A), curved row of 4–7 spines (Fig 13B) alternated with non-limbate companion chaetae (pennoned chaetae) (Fig 13C) and ventral row of 5–12 unilimbate capillaries (Fig 13D). Spines large, curved and bearing distinctive crest of fine bristles (Fig 12C).

Notopodial postchaetal lamellae rounded on chaetigers 2–4, absent on chaetiger 5, rounded on chaetigers 6–10, absent from chaetiger 11 onwards. Notopodial prechaetal lamellae absent throughout.

459

Neuropodial postchaetal lamellae rounded on chaetigers 2–4, absent on chaetiger 5, rounded on chaetigers 6–8 and absent from chaetiger 9 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and narrowly unilimbate capillaries. Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, non-limbate, thinner, and less numerous. Notopodial posterior spines present from chaetiger 16–21 almost to the end of the body, inserted in the notopodia or protracted, 4–8 spines per chaetiger (Fig 13E). Spines thin, smooth and with sharp tip, accompanied by 3–5 long non-limbate capillaries.

Hooks in neuropodia only, consistently starting from chaetiger 7, up to 12 per fascicle, accompanied by 1–5 long non-limbate capillaries throughout the entire body. Hooks bidentate, slightly curved near the distal end, lacking constriction (Fig 13F).

Branchiae starting from chaetiger 7, present on about two thirds of the total specimen length. Branchiae smooth and slightly flattened (Fig 12A), longest on chaetigers 10–12 and gradually reducing in length towards last branchial pair.

Pygidium quadrilobate, all lobes of the same size.

Juvenile morphology: In juveniles, the notopodial posterior spines are present from chaetiger 13.

Methyl green staining pattern: Prostomium weakly stained; dorsal side of chaetigers 7–20, ventral side of chaetiger 6–8 and glandular pouches strongly stained.

Remarks: Dipolydora caulleryi was originally described from France as Polydora caulleryi Mesnil, 1897 and was subsequentially recorded on different areas in the Northeastern Atlantic Ocean, Mediterranean Sea and the Pacific and Atlantic coasts of the USA. Blake (1996) reestablished Dipolydora, transferred Polydora caulleryi to Dipolydora and divided species in five groups according to morphological characters, grouping D. caulleryi with D. blakei, D. aciculata, D. akaina, D. armata, D. notialis and D. quadrilobata, while Meiβner et al. (2014) attributed an eighth species, D. paracaulleryi to this group.

Dipolydora caulleryi shares similarities with D. blakei and D. armata, discussed in the Remarks section of each species. The species is also similar to D. paracaulleryi in having a shallow incised prostomium, spines from chaetiger 5 falcate, bearing distinctive crest of fine bristles, branchiae starting from chaetiger 7 (as all species in the group established by Blake [1996]), hooks accompanied by capillaries throughout the body, smooth notopodial posterior spines and a quadrilobate pygidium. However, species can be distinguished by the caruncle morphology – extending laterally to chaetiger 1 in D. paracaulleryi, presence of pennoned chaetae in D.

460 caulleryi and by branchial distribution – on about two thirds of the total specimen length in D. caulleryi and 5–6 pairs in D. paracaulleryi (Meiβner et al. 2014).

Specimens described in this work do not entirely fit the original description nor Blake’s (1971) redescription, as our specimens exhibited an entire or shallow incised prostomium and a caruncle extending to the end of chaetiger 2, while the D. caulleryi typically possess a deeply incised prostomium and a caruncle extending to chaetigers 3–4 (Blake 1971; Maciolek 1984; Johnson 1984). Due to these differences, it was decided to adopt the “cf.”, which is provisory until material from the type locality can be examined and different developmental stages can be studied.

Habitat: medium silt, 1000–1335 m depth.

Distribution: Atlantic Ocean: Brazil (Rio de Janeiro and Espírito Santo states), USA, Europe (France; type locality); Pacific Ocean: USA; Mediterranean Sea.

Dipolydora socialis (Schmarda, 1861)

(Figs 14–16E)

Synonym list:

Leucodore socialis Schmarda, 1861 (superseded original combination)

Polydora (Polydora) socialis (Schmarda, 1861) (synonym)

Polydora socialis (Schmarda, 1861) (superseded original combination)

Material examined. Amb1 Foz1, 19°52'21,52" S 39°59'33,54" W, 28m (2 ind); Amb1 Foz7, 19°49'57,38" S 39°52'14,02" W, 33m 3 (ind).

Comparative material examined. Dipolydora socialis USNM 187537 (1 specimen, non-type).

Description: a medium-sized Dipolydora species, largest complete specimen 7.5 mm long, 0.4 mm wide for 84 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region, tapering towards the pygidium (Fig 15A). Pigmentation patterns absent. Body color whitish in alcohol (Fig 14).

Prostomium narrow, deeply incised anteriorly, extending posteriorly as a low caruncle to the end of chaetiger 4–5 (Fig 15B). Two or three pairs of eyes in trapezoidal arrangement. Peristomium

461 partly fused to chaetiger 1, lacking lateral wings (Figs 14, 15A–B). Grooved palps reaching up to chaetiger 15, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami – especially on the notopodium (2–4 capillaries); shorter than chaetae on succeeding chaetigers. Postchaetal lamellae digitiform on notopodium and triangular on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Chaetiger 5 modified, up to 2.5 times longer than chaetigers 4 or 6 (Figs 14, 15B, D). Presence of 2–4 dorsal geniculated chaetae (Fig 16A), curved row of 2–4 spines (Fig 16B) alternated with bilimbate companion chaetae (pennoned chaetae) (Fig 16C) and ventral row of 1–3 bilimbate capillaries (Fig 16D). Spines falcate, bearing subterminal protuberance (Fig 15D).

Notopodial postchaetal lamellae conical on chaetigers 2–4, absent on chaetiger 5, digitiform and much reduced from chaetiger 6 onwards, absent on the last chaetigers. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae triangular on chaetigers 2–4, absent on chaetiger 5, rounded and much reduced on chaetigers 6–7, absent after chaetiger 8–9. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and narrowly bilimbate capillaries, anterior row of short and posterior row of long (up to two times longer than anterior row) capillaries. Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, narrowly bilimbated (barely perceptible), thinner, and less numerous. Notopodial posterior spines absent.

Hooks in neuropodia only, consistently starting from chaetiger 7, up to five per fascicle, accompanied by 1 – 3 long non-limbate capillaries present only on the first 5–6 chaetigers after the appearance of the hooded hooks. Hooks bidentate, slightly curved near the distal end, lacking constriction (Fig 16E).

Branchiae starting from chaetiger 8, extending for most of the body, absent on last 15 chaetigers. Branchiae smooth and slightly flattened, longest on chaetigers 10–25 (Fig 15B–C) and gradually reducing in length towards last branchial pair.

Pygidium cuff-like, with slight dorsal gap.

Gizzard-like structure on digestive tract from chaetiger 17–20, extending for two or three chaetigers.

Methyl green staining pattern: Prostomium intensely stained.

462

Remarks: Dipolydora socialis was originally described as Leucodore socialis Schmarda, 1861 from Viña del Mar (Chile) and then transferred to Polydora by Hartman (1959), being subsequentially recorded on different latitudes and depths in the Atlantic and Pacific Oceans, although many older records cannot be verified and the type material is apparently lost. Blake (1996) reestablished Dipolydora transferred Polydora socialis to Dipolydora and divided species in five groups according to morphological characters, grouping Dipolydora socialis with D. concharum (Verrill, 1880), D. alborectalis (Radashevsky, 1993), D. cardalia (Berkeley, 1927), D. capensis (Day, 1955), D. coeca (Oersted, 1843), D. flava (Claparède, 1870), D. goreensis (Augener, 1918), D. normalis (Day, 1957), D. peristomialis (Hartman, 1976), D. pilikia (Ward, 1981), D. protuberata (Blake & Kudenov, 1978), D. saintjosephi (Eliason, 1920), D. tentaculata (Blake and Kudenov, 1978) and D. vulcanica (Radashevsky, 1994).

Blake (1996) defined this group as lacking constriction on the hooded hooks (which is a character shared among all Dipolydora species), occipital tentacle present or absent, branchiae usually starting from chaetigers 7–8, presence of notopodial chaetae on chaetiger 1, spines from chaetiger 5 falcate and posterior notopodial spines absent, needle-like or curved.

Among species recorded in this work, D. socialis is most similar to D. barbilla (as discussed on the species’ Remarks section). Specimens examined in this work agrees with previous descriptions (Blake 1971; Blake & Kudenov 1978; Johnson 1984; Maciolek 1984; Sato-Okoshi, 2000; Williams 2001; Sato-Okoshi & Takatsuka 2001; Bone & Viéitez 2002). The only varying characters among these descriptions are the caruncle extension, position of the gizzard-like structure and pygidium morphology.

Among descriptions, caruncle extended from chaetiger 3 up to chaetiger 9, although this variation is acknowledged as size-dependent, as, according to Sato-Okoshi & Takatsuka (2001), larger worms possess longer caruncles. As for the gizzard-like structure, which was described in detail by Blake (1971), this structure may be present from chaetigers 10–11 (Maciolek 1983), chaetiger 16 (Blake 1996), chaetigers 14–18 (Williams 2001), chaetigers 18–19 (Blake & Kudenov 1978) and chaetigers 17–20 (this study), although no explanation was found for this variation. At last, pygidium has been described as disk-like (Blake & Kudenov 1978; Sato-Okoshi 2000), cuff- shaped (Williams 2001, this study), bearing large ventral lobe and two smaller dorsal lobes separated by a notch (or notch absent) (Blake 1971; Maciolek 1983; Blake 1996; Sato-Okoshi 2000) or three continuous lobes (Blake 1971). The significance of this variation, however, have not been discussed.

According to Blake (1996), D. socialis is capable of boring calcareous substrates or inhabiting soft sediments.

Habitat: coarse sand to medium sand, 28–33 m depth.

463

Distribution: Atlantic Ocean: Brazil (Santa Catarina, Paraná. São Paulo, Rio de Janeiro and Espírito Santo states), Venezuela, USA, Trinidad and Tobago, Argentina, Luanda, France and Spain; Pacific Ocean: Chile (type locality), USA, Australia, Philippines, Mexico, Canada, New Zealand, Ecuador, Japan.

Dipolydora tetrabranchia (Hartman, 1945)

(Figs 17–19E)

Synonym list:

Polydora tetrabranchia Hartman, 1945 (superseded original combination).

Material examined. Amb1 Foz14 19°42'32,21" S 39°38'57,36" W, 38m (1 ind); Amb7 A2 21°3'27,14" S 40°22'59,61" W, 40m (3 ind); Amb7 F2 18°52'32,61" S 39°8'42,82" W, 40m (3 ind); Amb13 G2 18°36'32,45" S 39°9'32,83" W, 40m (7 ind); Amb14 F2 18°52'31,35" S 39°8'41,34" W, 40m (4 ind).

Comparative material examined. Dipolydora tetrabranchia USNM 53839 (12 specimens, non- type material).

Description: a small-sized Dipolydora species, largest complete specimen 1.4 mm long, 0.15 mm wide for 29 chaetigers. Body dorsoventrally flattened throughout, tapering towards the pygidium. Pigmentation patterns absent. Body color whitish in alcohol (Fig 17).

Prostomium narrow, bearing shallow median incision, extending posteriorly as a low caruncle to the end of chaetiger 1. Eyes absent. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 17, 18A). Grooved palps reaching up to chaetiger 15, lost in most specimens.

Chaetiger 1 bearing only a few chaetae on both rami – especially on the notopodium, shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on notopodium and digitiform on neuropodium, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent.

Chaetiger 5 modified, up to two times longer than chaetigers 4 or 6 (Figs 17, 18C–D). Presence of 2 dorsal geniculated chaetae (Fig 19A), curved row of 1–3 spines (Fig 19B) alternated with bilimbate companion chaetae (pennoned chaetae) (Fig 19C) and ventral row of 1–3 bilimbate capillaries (Fig 19D). Spines bifurcate, bearing two robust and distally tapered tips (Fig 18D).

464

Notopodial postchaetal lamellae rounded on chaetigers 2–4 and 6, absent on chaetiger 5, rounded and much reduced from chaetiger 6 onwards, absent after chaetiger 10. Notopodial prechaetal lamellae absent throughout.

Neuropodial postchaetal lamellae digitiform on chaetigers 2–4, absent on chaetiger 5, rounded and much reduced on chaetigers 6 and absent from chaetiger 7 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and narrowly bilimbate capillaries. On notopodia, anterior row of short and posterior row of long (up to 1.5 times longer than anterior row) capillaries. Notopodial chaetae up to 1.5 times longer than neuropodial chaetae. Chaetae scarce, 3–10 per rami; only 1–3 notopodial capillaries from chaetiger 11 onwards. After the branchial region, capillaries suddenly become elongate, narrowly bilimbated (barely perceptible) and thinner. Notopodial posterior spines absent; notopodial chaetae elongated on last 3–4 chaetigers.

Hooks in neuropodia only, consistently starting from chaetiger 7, up to four per fascicle, accompanied by 1 –2 long non-limbate capillaries present only on the first 1–5 chaetigers after the appearance of the hooded hooks. Hooks bidentate, strongly curved near middle length of the shaft, lacking constriction (Fig 19E).

Four pairs of branchiae on chaetigers 7–10. Branchiae smooth and slightly flattened (Fig 18B), longest on chaetigers 8–9.

Pygidium disc-like, lacking lobes or gap.

Juvenile morphology: In juveniles, presence of 1–2 spines on chaetiger 5, which are embedded in the chaetiger, visible only through light microscopy. Additionally, non-limbate capillaries accompanying hooded hooks may be completely absent.

Methyl green staining pattern: Ventral side of peristomium and of chaetigers 7–10 intensely stained.

Remarks: Specimens examined in this work agree completely with Hartman’s (1945) original description and Blake’s (1971) redescription, based on specimens from North Carolina (Atlantic coast of the USA), which was originally described as Polydora tetrabranchia. Blake (1996) reestablished Dipolydora, transferred Polydora tetrabranchia to Dipolydora and divided species in five groups according to morphological characters, grouping D. tetrabranchia with D. giardi (Mesnil, 1896), D. bifurcata (Blake, 1981), D. magellanica (Blake, 1983), D. tridenticulata (Woodwick, 1964) and D. trilobata (Radashevsky, 1993).

465

Blake (1996) defined this group as lacking constriction on the hooded hooks (which is a character shared among all Dipolydora species), branchiae starting from chaetigers 7–8, presence of notopodial chaetae on chaetiger 1, spines from chaetiger 5 with one large accessory tooth and sometimes additional small teeth or flanges and posterior notopodial spines present or absent.

Among species in this group, D. giardi, D. bifurcata, D. magellanica, D. tridenticulata and D. trilobata possess spines from chaetiger 5 bearing one accessory tooth and an additional small teeth, spur or flange, while D. tetrabranchia possess a single large accessory tooth, almost as large as the spine in some specimens.

This species can be distinguished from the remaining Brazilian species by its very small size; short caruncle, reaching up to the end of chaetiger 1; spines from chaetiger 5 bearing a large accessory tooth and short branchial distribution – from chaetiger 7 to chaetiger 10. Dipolydora tetrabranchia is similar to D. socialis in lacking posterior notopodial spines and in having capillaries accompanying hood hooks for a limited number of chaetigers. However, both species can be separated based on caruncle length, to the end of chaetiger 1 in D. tetrabranchia and to the end of chaetigers 4–5 in D. socialis; morphology of spines from chaetiger 5, bifurcate in D. tetrabranchia and falcate, bearing subterminal protuberance in D. socialis and branchial distribution – on chaetigers 7–10 in D. tetrabranchia and from chaetiger 8, extending for most of the body in D. socialis.

Dipolydora tetrabranchia differs from all Brazilian Dipolydora species – and from most Dipolydora species in having only four pairs of branchiae, on chaetigers 7–10. This report represents the second report of D. tetrabranchia to the Brazilian Coast, as Assis et al. (2012) reported this species to Paraíba state, northeastern Brazil.

Habitat: medium sand to very fine sand, 38–40 m depth.

Distribution Atlantic Ocean: Brazil (Rio de Janeiro and Espírito Santo states), USA (North Carolina; type locality).

Identification key to Dipolydora species recorded in this work.

1. Branchiae starting from chaetiger 7………………………………………..……………….2 - Branchiae starting from chaetigers 8–10…………………………………………………….4 2(1) Only four pairs of branchiae. Lack of notopodial posterior spines...... Dipolydora tetrabranchia From 6 to 9 pairs of branchiae. Notopodial posterior spines present………………………….3

466

3(2) Falcate spines of chaetiger 5 with large lateral tooth; apical cowling-like structure absent on convex side of main fang. Notopodial posterior spines thin……………..Dipolydora blakei - Falcate spines of chaetiger 5 with large lateral tooth and apical structure appearing as wide cowling or third tooth on convex side of main fang. Notopodial posterior spines thick………………………………………………………………………...Dipolydora armata 4(1) Notopodial posterior spines present...... Dipolydora barbilla - Notopodial posterior spines absent……………………………………….Dipolydora socialis

Genus Boccardia Carazzi, 1893

Type species: Boccardia polybranchia Haswell, 1885

Synonym list: Perialla Kinberg, 1866 (senior subjective synonym)

Paraboccardia Rainer, 1973 (subjective synonym)

Neoboccardia Buzhinskaja, 1985 (subjective synonym)

Diagnosis (modified from Blake et al. 2017): Prostomium anteriorly rounded or incised, extending posteriorly as caruncle of variable length. Notopodial lamellae and notochaetae present or absent on chaetiger 1. Chaetiger 5 modified, with two types of major spines, one type simple, smooth, falcate; second type with expanded apex bearing collar or cloak of bristles; accompanied by few or no capillary notochaetae. Bidentate hooded hooks with conspicuous angle between teeth; hooks lacking constriction or manubrium on shaft; hooks first appear on chaetiger 7–11.

Posterior notopodial spines present or absent. Branchiae from chaetiger 2, absent on chaetiger 5, then present from chaetiger 6, continuing for variable number of chaetigers. Pygidium disclike, with or without separate lobes, or reduced to lobes or cuffs.

467

Boccardia capixaba sp. nov.

(Figs 20–22F)

Type material. Brazil. Espírito Santo Basin. Holotype: Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 02 Dec 2011 to 02 Fev 2012, 25m, MNRJP XXXX. Paratypes: Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 02 Dec 2011 to 02 Fev 2012, 25m MNRJP XXXX (3 ind), MZUSP XXXX (3 ind).

Additional material examined. Amb7 D1, 19°35'37,21" S 39°41'19,68" W, 25m (1 ind).

Comparative material examined. Boccardia polybranchia USNM 67602 (35 specimens, non- type).

Description: a medium-sized Boccardia species, largest complete specimen 5 mm long, 0.9 mm wide for 56 chaetigers, holotype complete, 4.8 mm long, 0.8 mm wide at the widest point for 53 chaetigers. Body dorsoventrally flattened throughout, especially on the postbranchial region (Fig 21B), tapering towards the pygidium. Pigmentation present as black stripes on the edges of prostomium and as a pair of brown stripes parallel to palp grooves. Body color light yellow to whitish in alcohol (Fig 20).

Prostomium narrow, bearing shallow median incision, extending posteriorly as a low caruncle to the end of chaetiger 3. Two pairs of eyes in trapezoidal arrangement. Peristomium partly fused to chaetiger 1, lacking lateral wings (Figs 20, 21A, C). Grooved palps reaching up to chaetiger 18, lost in most specimens (Fig 21A–C).

Chaetiger 1 bearing only a few chaetae on both rami – especially on the notopodium; shorter than chaetae on succeeding chaetigers. Postchaetal lamellae rounded on both rami, smaller than lamellae on succeeding chaetigers. Prechaetal lamellae absent (Fig 21C).

Chaetiger 5 modified, up to three times longer than chaetigers 4 or 6 (Figs 20, 21B–C). Presence of curved row of 5–7 spines of two types (Figs 22A, B) and ventral row of 4–8 bilimbate capillaries (Fig 22C). Geniculated chaetae and companion chaetae absent. Spines of two types: first type wide, with expanded apex covered in fine bristles (Fig 22A) and second type smooth and falcate (Figs 21D, 22B).

Notopodial postchaetal lamellae rounded on chaetigers 2–4, absent on chaetiger 5, rounded and much reduced on chaetigers 6–10, and absent from chaetiger 11 onwards. Notopodial prechaetal lamellae absent throughout.

468

Neuropodial postchaetal lamellae rounded on chaetigers 2–4, absent on chaetiger 5, rounded and much reduced on chaetigers 6, and absent from chaetiger 7 onwards. Neuropodial prechaetal lamellae absent throughout.

Chaetae from notopodia and neuropodia organized in two rows of non-granulated and bilimbate capillaries (Fig 22D). Notopodial chaetae up to 2.5 times longer than neuropodial chaetae. Towards the posterior region, capillaries progressively become elongate, narrowly bilimbated (barely perceptible), thinner, and less numerous. Notopodial posterior spines absent.

Hooks in neuropodia only, consistently starting from chaetiger 7 (Fig 21E), up to nine per fascicle, accompanied by 1–3 inferior long non-limbate capillaries accompanying the hooded hooks only on the first 2–4 chaetigers after the appearance of the hooded hooks (Fig 21F). Hooks bidentate, slightly curved near the distal end, lacking constriction (Fig 22E). Hoods covered in fine bristles.

Branchiae starting from chaetiger 2 (Fig 21A, C), absent on chaetiger 5; extending almost to the last chaetiger, absent on last 10–15 chaetigers. Branchiae smooth and cirriform (Fig 22F), basally fused to the notopodial lamellae, longest on chaetigers 10–35 and gradually reducing in length towards last branchial pair.

Pygidium unilobed, bearing median dorsal gap.

Methyl green staining pattern: Prostomium, peristomium, dorsal side of chaetigers 1–4 stained; dorsal region of anterior and median region, and region above and below notopodia intensely stained.

Remarks: Boccardia capixaba sp. nov. is similar to B. polybranchia from the Pacific Coast of Australia (New South Wales) and B. proboscidea, from the Pacific Coast of the USA (California), in having a caruncle extending to the posterior end of chaetiger 3; lack of occipital tentacle; lack of geniculate chaetae on chaetiger 5; neuropodial hooded hooks from chaetiger 7 and lack of posterior notopodial spines (Blake & Woodwick 1971; Kerckhof & Faasse 2014; Radashevsky et al. 2019).

However, B. capixaba sp. nov. differ from the aforementioned species in having curved falcate spines from chaetiger 5, only slightly curved in B. polybranchia and B. proboscidea; branchiae absent on last 10–15 chaetigers, absent on the posterior half in B. polybranchia and absent on last chaetigers in B. proboscidea; maximum number of neuropodial hooded hooks – up to nine per fascicle in B. capixaba sp. nov., instead of up to 13 hooks per fascicle, as in both B. polybranchia and B. proboscidea and pygidium morphology, disk-like, bearing small median gap, lacking a median gap in B. polybranchia and bearing four lobes in B. proboscidea.

469

Boccardia capixaba sp. nov. differs from B. polybranchia in the prostomium shape – narrow with slightly anterior incision in B. capixaba sp. nov. and incised in B. polybranchia; in the number of eyes – four in B. capixaba sp. nov. and 6–8 in B. polybranchia; lack of notopodial capillaries on chaetiger 1 in B. polybranchia and capillaries accompanying neuropodial hooded hooks in 2–4 chaetigers after the appearance of the hooks in B. capixaba sp. nov. and until middle region in B. polybranchia (Blake & Woodwick 1971; Blake & Kudenov 1978).

Boccardia capixaba sp. nov. also differs from B. proboscidea in the prostomium shape – narrow with slightly anterior incision in B. capixaba sp. nov. and rounded in B. proboscidea; in the number of eyes – four in B. capixaba sp. nov. and up to six in B. proboscidea; morphology of the bristled-topped spines from chaetiger 5 – straight in B. capixaba sp. nov. and slightly curved in B. proboscidea and capillaries accompanying neuropodial hooded hooks in 2–4 chaetigers after the appearance of the hooks in B. capixaba sp. nov. and present up to the posterior region in B. proboscidea (in inferior position in chaetigers 7–9 and alternating with hooded hooks on the posterior region) (Blake & Woodwick 1971; Radashevsky et al. 2019). The two species also seem to differ on the habitat, as B. proboscidea is a shell or stone borer (Radashevsky et al. 2019), while B. capixaba sp. nov. specimens were found inhabiting sandy tubes in the sediment.

Etymology. The species epithet, “capixaba” is the common denomination of natives from the state of Espírito Santo, southeastern Brazil, where the specimens were collected.

Habitat: Fine silt, 25 m depth.

Distribution: Southeastern Brazil (Espírito Santo state), South Atlantic Ocean.

Acknowledgments

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Legends of illustrations Figure 1. Map showing sampled area (blue polygon).

Figure 2. Dipolydora armata, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 3. SEM of Dipolydora armata. A. Whole fragmente, dorsal vew. B. Anterior region, dorsal view. C. Chaetigers 5 and 6, dorsolateral view. D. Posterior parapodia, lateral view. Abbreviations: br, branchia; gc, geniculated chaetae; ns, notopodial spine; pe, peristomium; pr, prostomium; sp, spine.

Figure 4. Dipolydora armata. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Ventral row of capillaries from chaetiger 5. D. Notopodial posterior spine. E. Hooded hook.

Figure 5. Dipolydora barbilla, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium.

Figure 6. SEM of Dipolydora barbilla. A. Whole specimen, dorsal view (prostomium) and ventral view (pygidium). B. Anterior region, dorsal view. C. Whole specimen, lateral view. D. Chaetigers 4 and 5, lateral view. E. Pygidium, ventral view. F. Notopodial spine, ventral view. Abbreviations: bc, bilimbate chaeta; br, branchia; car, caruncle; cha5, chaetiger 5, gc, geniculated chaetae; ns, notopodial spine; pc, pennoned chaetae; pr, prostomium; pyg, pygidum; sp, spine. Arrow on Fig A indicates where specimen is twisted and changes plane of view.

Figure 7. Dipolydora barbilla. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Companion chaetae from chaetiger 5. D. Ventral row of capillaries from chaetiger 5. E. Notopodial posterior spine. F. Hooded hook.

Figure 8. Dipolydora blakei, whole specimen, lateral view, showing characteristic pigmentation in ethanol. Abbreviations: pe, peristomium; pr, prostomium; pyg, pygidium.

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Figure 9. SEM of Dipolydora blakei. A. Anterior fragment, dorsal view. B. Anterior and middle region, lateral view. C. Whole specimen, lateral view. D. Anterior region, lateral view. E. Chaetigers 3–6, lateral view. F. Posterior region, lateral view. Abbreviations: bc, bilimbate chaetae; car, caruncle; cha5, chaetiger 5; gc, geniculated chaetae; hh, hooded hook; ns, notopodial spine; pr, prostomium; pyg, pygidium; sp, spine.

Figure 10. Dipolydora blakei. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Companion chaetae from chaetiger 5. D. Ventral row of capillaries from chaetiger 5. E. Notopodial posterior spine. F. Hooded hook.

Figure 11. Dipolydora cf. caulleryi, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pa, palp; pe, peristomium; pr, prostomium.

Figure 12. SEM of Dipolydora cf. caulleryi. A. Anterior fragment, dorsal view. B. Anterior fragment, lateral view. C. Chaetigers 3–6, lateral view. Abbreviations: bc, bilimbate chaetae; br, branchia; cha5, chaetiger 5; gc, geniculated chaetae; hh, hooded hook; pc, pennoned chaetae; pr, prostomium; pa, palp; sp, spine.

Figure 13. Dipolydora cf. caulleryi. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Companion chaetae from chaetiger 5. D. Ventral row of capillaries from chaetiger 5. E. Notopodial posterior spine. F. Hooded hook.

Figure 14. Dipolydora socialis, anterior chaetigers and mid-body, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pe, peristomium; pr, prostomium.

Figure 15. SEM of Dipolydora socialis. A. Anterior fragment, dorsal view. B. Anterior region, dorsal view. C. Middle region, dorsal view. D. Chaetigers 4 and 5, dorsal view. Abbreviations: car, caruncle; gc, geniculated chaetae; gi, gizzard-like structure; pc, pennoned chaetae; pe, peristomium; pr, prostomium; sp, spine.

Figure 16. Dipolydora socialis. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Companion chaetae from chaetiger 5. D. Ventral row of capillaries from chaetiger 5. E. Hooded hook.

Figure 17. Dipolydora tetrabranchia, whole specimen, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: pa, palp; pe, peristomium; pr, prostomium; pyg, pygidium.

Figure 18.SEM of Dipolydora tetrabranchia. A. Anterior fragment, dorsal view. B. Anterior and middle chaetigers, dorsal view. C. Chaetigers 3–5, dorsal view. D. Chaetigers 5 and 6, dorsolateral

480 view. Abbreviations: bc, bilimbate chaetae; br, branchia; cha5, chaetiger 5; gc, geniculated chaetae; pa, palp; pr, prostomium; sp, spine.

Figure 19. Dipolydora tetrabranchia. A. Dorsal geniculated chaetae. B. Modified spine from chaetiger 5. C. Companion chaetae from chaetiger 5. D. Ventral row of capillaries from chaetiger 5. E. Hooded hook.

Figure 20. Boccardia capixaba sp. nov., anterior chaetigers, dorsal view, showing characteristic pigmentation in ethanol. Abbreviations: br, branchiae; pa, palp; pe, peristomium; pr, prostomium.

Figure 21. SEM of Boccardia capixaba sp. nov. A. Anterior region, dorsal view. B. Anterior and middle fragment, lateral view. C. Anterior region, lateral view. D. Chaetiger 5, lateral view. E. Middle region, lateral view. F. Chaetigers 9–11, lateral view. Abbreviations: bc, bilimbated chaetae; br, branchia; bs, bristle-tipped spine; ca, capillary chaetae; cha5, chaetiger 5; hh, hooded hook; pa, palp; pe, peristomium; pr, prostomium; ss, smooth spine.

Figure 22. Boccardia capixaba sp. nov. A. Modified bristle-tipped spine from chaetiger 5. B. Modified falcate spine from chaetiger 5. C. Ventral row of capillaries from chaetiger 5. D. Hooded hooks. E. Branchia.

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Tables

Table 1 – Comparative diagnostic characters of Dipolydora species from this work.

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Table 1 (continued) – Comparative diagnostic characters of Dipolydora species from this work.

483

Table 2 – Comparative diagnostic characters of Boccardia species reported to Brazil.

484

Figure 1

485

Figure 2

Figure 3

486

Figure 4

Figure 5

487

Figure 6

488

Figure 7

Figure 8

489

Figure 9

490

Figure 10

Figure 11

491

Figure 12

492

Figure 13

Figure 14

493

Figure 15

Figure 16

494

Figure 17

Figure 18

495

Figure 19

Figure 20

496

Figure 21

497

Figure 22

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5. Considerações finais

Foram examinados 16.938 espécimes, pertencentes a 17 gêneros e 64 espécies, sendo duas espécies de Aonidella, duas espécies de Aonides, uma espécie de Aurospio, uma espécie de

Apoprionospio, uma espécie de Boccardia, seis espécies de Dipolydora, três espécies de Dispio, sete espécies de Laonice, uma espécie de Laubieriellus, uma espécie de Malacoceros, seis espécies de Microspio, uma espécie de Paraprionospio, 20 espécies de Prionospio, quatro espécies de Scolelepis, três espécies de Spio, uma espécie de Spiogalea e quatro espécies de

Spiophanes. Entre os gêneros citados acima, Spiogalea, Laubieriellus e Aonidella correspondem a novos registros para a costa Brasileira, enquanto as espécies Aonidella dayi, A. cf. cirrobranchiata, Dipolydora cf. caulleryi e Microspio profunda são novas ocorrências para a costa Brasileira.

Dentre as 64 espécies encontradas nessa Tese, foram registradas 12 espécies conhecidas pela ciência, oito espécies conhecidas pela ciência, mas que podem corresponder a táxons novos

(Aonidella cf. cirrobranchiata, Apoprionospio cf. dayi, Dipolydora cf. caulleryi, Dispio cf. brachychaeta, Laonice cf. antarcticae, L. cf. weddellia, Prionospio cf. delta e P. cf. ehlersi), sete espécies novas que não foram formalmente descritas devido ao baixo número de indivíduos e/ou mau estado de conservação do material (Dispio sp. 1, Microspio sp. 1, M. sp. 2, Scolelepis sp. 1,

S. sp. 2, S. sp. 3 e Spio sp. 1) e 37 espécies novas, sendo 4 dessas espécies já publicadas –

Spiogalea capixaba (Capítulo I), Laubieriellus decapitata, Prionospio solisi e P. nonatoi

(Capítulo II), enquanto as espécies novas dos Capítulos III a VIII serão submetidas para publicação no futuro.

Para a Bacia do Espírito Santo eram conhecidas apenas cinco espécies de Spionidae, nenhuma destas descrita para a costa Brasileira –Dipolydora socialis, Polydora cornuta, Prionospio heterobranchia, Apoprionospio pygmaeus e Spiophanes duplex. O número de espécies novas encontradas nesse trabalho supera em muito o número de espécies de Spionidae descritos na costa

Brasileira – apenas Laonice antarcticae, L. brevicornis, L. branchiata, L. parvabranchiata,

499

Pseudopolydora rosebelae e Polydora carinhosa. Todas essas espécies ocorrerem na plataforma continental (exceto por alguns registros de Laonice antarcticae no talude continental) e, no caso de espécies de Laonice, apresentam tamanho grande e em alguns casos possuem um carácter morfológico que facilita a identificação da espécie, como as brânquias acessórias de L. branchiata.

Embora fosse conhecido de antemão que o talude continental e cânions (Watu Norte e Doce) correspondessem a ambientes pouco conhecidos em relação à fauna (Paiva & Barroso, 2010;

Amaral et al., 2013), a plataforma continental e a foz do Rio Doce revelaram possuir um maior número de espécies novas para a ciência – 17 espécies novas exclusivas da plataforma continental e foz do Rio Doce e 14 espécies novas exclusivas do talude continental e cânions, além de 6 espécies novas ocorrendo tanto na plataforma continental quanto no talude continental. Esse trabalho foi ainda o primeiro a realizar um refinamento taxonômico em material do talude continental, uma vez que o material de Spionidae de projetos anteriores foi identificado apenas em nível de gênero ou morfotipado antes de ser depositado em coleções. Esse esforço taxonômico levou ao registro de uma espécie de Scolelepis de mar profundo, um gênero até então restrito à plataforma continental rasa.

A maior resolução taxonômica dessa Tese pode ter sido responsável pela ausência de registros de espécies “cosmopolitas” na área estudada. Por exemplo, nenhum exemplar de Prionospio steenstrupi, espécie descrita para a Islândia (e rotineiramente reportada na costa Brasileira), foi reportado nessa tese, enquanto três espécies com morfologia similar à P. steenstrupi foram reportados na área de estudo: Prionospio cristata, Prionospio triangularis sp. nov. e Prionospio posterocristata sp. nov. A ausência de espécies ditas “cosmopolitas” nesse trabalho levanta uma questão interessante: esses táxons simplesmente não ocorreram na área estudada ou uma maior resolução taxonômica foi capaz de distinguir um ou mais táxons que foram anteriormente identificados como uma espécie cosmopolita?

Em relação ao refinamento taxonômico, essa tese contribui para o conhecimento da família Spionidae no Brasil, sento útil como um trabalho introdutório para a diversidade do grupo –

500 mesmo tratando de uma região geográfica limitada e serve como literatura de referência para a identificação do grupo.

501

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Anexo 1 – Morfotipagem de espécies no projeto HABITATS.

Aonides sp. A Aonides sarsi sp. nov. Laonice sp. A Laonice kinbergi sp. nov. Microspio sp. B Microspio alexandrae sp. nov. Microspio sp. C Microspio sp. 1 Prionospio fauchaldi Prionospio corrugatus sp. nov. Prionospio sp. A Prionospio brevipinnulata sp. nov. Prionospio sp. B Prionospio capixaba sp. nov. Prionospio sp. C Prionospio mutatus sp. nov. Prionospio sp. D Prionospio absensi sp. nov. Prionospio sp. E Prionospio acutus sp. nov. Prionospio sp. F Prionospio quadrilamellata sp. nov. Prionospio sp. G Prionospio alexandrae sp. nov. Prionospio sp. L Prionospio triangularis sp. nov. Prionospio sp. M Prionospio posterocristata sp. nov. Prionospio sp. N (em parte) Prionospio kinbergi sp. nov. Prionospio sp. N (em parte) Prionospio hartmanae sp. nov. Prionospio sp. O Prionospio absensi sp. nov. Prionospio sp. P Laubieriellus decapitata Prionospio sp. R Prionospio fosterae sp. nov. Prionospio sp. S Prionospio cinthyae sp. nov. Prionospio sp. T Prionospio nonatoi Prionospio sp. U Prionospio solisi Scolelepis sp. A Scolelepis sp. 1 Spio sp. A Spio anaclaudiae sp. nov. Spio sp. B Spio capixaba sp. nov. Spiophanes sp. A Spiophanes grubei sp. nov. Spiophanes sp. B Spiophanes duplex Tabela 1 – Correspondência de espécies que ocorreram nos projetos HABITATS e AMBES, depositados no Museu de Zoologia Adão José Cardoso (UNICAMP). As espécies Aonidella cirrobranchiata, Aonides curvus sp. nov., Boccardia capixaba sp. nov., Dipolydora cf. caulleryi, Dispio levisi sp. nov., Dispio cf. brachychaeta, Dispio sp. 1, Laonice profunda sp. nov., Laonice brevicarunculata sp. nov., Malacoceros aureus sp. nov., Microspio tridentata sp. nov., Microspio quadridentata sp. nov., Microspio sp. 2, Paraprionospio

512 anterocristata sp. nov., Prionospio biancoi sp. nov., Scolelepis profunda sp. nov., Scolelepis sp. 2, Scolelepis sp. 3, Spio sp. 1, Spiogalea capixaba, Spiophanes aureus sp. nov. e S. profundus sp. nov. a princípio não foram reportadas no projeto HABITATS.

Entretanto, é importante notar que parte do material depositado no Museu de Zoologia Adão José Cardoso (UNICAMP) proveniente do projeto HABITATS foi identificado apenas até o nível de gênero. A insuficiência de resolução taxonômica impede que sejam feitas afirmações da ocorrência dessas espécies na Bacia de Campos. Apenas os táxons Microspio sp. A (que na realidade se trata de uma espécie do gênero Rhynchospio) e Polydora sp. (na realidade um Uncispionidae do gênero Rhamphispio Blake & Maciolek, 2018) foram exclusivos do projeto HABITATS.

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Anexo 2 – Chave para os gêneros de Spionidae reportados nesse trabalho.

1. Brânquias ausentes...... 2 - Brânquias presentes...... 3 2(1). Placa quitinosa recobrindo o prostômio; cerda em anzol ausente...... Spiogalea - Placa quitinosa ausente; cerda em anzol no neuropódio do setígero 1...... Spiophanes 3(1). Setígero 5 dotado de espinhos...... 4 - Setígero 5 dotados apenas de cerdas capilares...... 5 4(3). Dois tipos de espinhos no setígero 5...... Boccardia - Apenas um tipo de espinho no setígero 5...... Dipolydora 5. Brânquias presentes na maior parte do corpo...... 6 - Brânquias limitadas à região anterior do corpo...... 10 6(5). Prostômio pontiagudo ou cônico...... 7 - Prostômio largo, arredondado ou truncado a cônico (sem extremidade afilada)...... 8 7(6). Brânquias a partir do setígero 1. Brânquias acessórias presentes. Ausência de ganchos notopodiais...... Dispio - Brânquias a partir do setígero 2. Brânquias acessórias ausentes. Presenças de ganhos notopodiais...... Scolelepis 8(6). Brânquias a partir do setígero 2...... Microspio - Brânquias a partir do setígero 1...... 9 9(8). Prostômio dotado de cornos fronto-laterais. Pigídio com mais de 2 pares de cirros ...... Malacoceros - Ausência de cornos fronto-laterais. Pigídio com 2 pares de cirros...... Spio 10(5). Caruncula se estendendo por número variável de setígeros; antena e bolsas interneuropodiais geralmente presentes...... Laonice Órgãos nucais ausentes ou, quando presentes, não se estendendo além do setígero 2. Brânquias limitadas aos primeiros 20 setígeros...... 11 11(10). Ausência de ganchos notopodiais. Presença de cristas ventrais. 4 pares de brânquias lisas nos setígeros 2-5...... Laubieriellus - Presença de ganchos notopodiais...... 12 12(11). Prostômio cônico com extremidade afilada ou arredondada. Antena occipital presente ou ausente. Órgãos nucais ausentes...... Aonides - Prostômio alargado, arredondado ou truncado anteriormente. Antena occipital ausente. Órgãos nucais presentes...... 13 13(12) Ao menos um par de brânquias dotado de placas...... 14

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- Brânquias lisas ou dotadas de pínulas...... 15 14(13). Três pares de brânquias dotadas de placas a partir do setígero 1. Peristômio bem desenvolvido, envolvendo o prostômio...... Paraprionospio - Quatro pares de brânquias, primeiros três pares lisos e cirriformes e quarto par dotado de placas...... Apoprionospio 15(13). Dois a três pares de brânquias a partir do setígero 3, fundidas basalmente à lamela notopodial ou brânquias ausentes. Lamela notopodial do 3º setígero alargada...... Aurospio - Brânquias livres da lamela notopodial...... 16 16. Prostômio achatado. Lamelas parapodiais triangulares, sem alteração de formato ao longo do corpo. Ganchos com ângulo aberto (> 90º) entre o dente principal e o(s) dente(s) secundário(s)...... Aonidella - Lamelas parapodiais de formato variado, com alteração de formato e/ou tamanho ao longo do corpo. Ganchos com ângulo fechado (< 90º) entre o dente principal e os dentes secundários...... Prionospio

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