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UNIVERSIDADE ESTADUAL DE CAMPINAS Instituto De Biologia 1 UNIVERSIDADE ESTADUAL DE CAMPINAS Instituto de Biologia PAOLA VISNARDI FASSINA REDESCOBERTA DOS TARDÍGRADOS LIMNOTERRESTRES BRASILEIROS: REDESCRIÇÃO DE PSEUDECHINISCUS JUANITAE DE BARROS, 1939 E ​ ​ COMENTÁRIOS SOBRE PSEUDECHINISCUS NOVAEZEELANDIAE MARINAE ​ BARTOŠ, 1934 SENSU DE BARROS (1942) REDISCOVERY OF BRAZILIAN LIMNOTERRESTRIAL TARDIGRADES: ​ REDESCRIPTION OF PSEUDECHINISCUS JUANITAE DE BARROS, 1939 AND ​ ​ REMARKS OF PSEUDECHINISCUS NOVAEZEELANDIAE MARINAE BARTOŠ, ​ ​ 1934 SENSU DE BARROS (1942) CAMPINAS 2020 2 PAOLA VISNARDI FASSINA REDESCOBERTA DOS TARDÍGRADOS LIMNOTERRESTRES BRASILEIROS: REDESCRIÇÃO DE PSEUDECHINISCUS JUANITAE DE BARROS, 1939 E ​ ​ COMENTÁRIOS SOBRE PSEUDECHINISCUS NOVAEZEELANDIAE MARINAE ​ BARTOŠ, 1934 SENSU DE BARROS (1942) REDISCOVERY OF BRAZILIAN LIMNOTERRESTRIAL TARDIGRADES: ​ REDESCRIPTION OF PSEUDECHINISCUS JUANITAE DE BARROS, 1939 AND ​ ​ REMARKS OF PSEUDECHINISCUS NOVAEZEELANDIAE MARINAE BARTOŠ, ​ ​ 1934 SENSU DE BARROS (1942) Dissertation presented to the Institute of Biology of the University of Campinas in partial fulfillment of the requirements for the degree of Master in Animal Biology, in the area of Animal Biodiversity. Dissertação apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos requisitos exigidos para a obtenção do título de Mestra em Biologia Animal, na Área de Biodiversidade Animal. Orientador: PROF. DR. ANDRÉ RINALDO SENNA GARRAFFONI ESTE TRABALHO CORRESPONDE À VERSÃO FINAL DA DISSERTAÇÃO/TESE DEFENDIDA PELO ALUNA PAOLA VISNARDI FASSINA, E ORIENTADA PELO PROF. DR. ANDRÉ RINALDO SENNA GARRAFFONI. CAMPINAS 2020 3 4 Campinas, 8 de Setembro de 2020. COMISSÃO EXAMINADORA Prof. Dr. André Rinaldo Senna Garraffoni Profa. Dra. Clélia Márcia Cavalcanti da Rocha Prof. Dr. Flavio Dias Passos Os membros da Comissão Examinadora acima assinaram a Ata de Defesa, que se encontra no processo de vida acadêmica do aluno. A Ata da defesa com as respectivas assinaturas dos membros encontra-se no SIGA/Sistema de Fluxo de Dissertação/Tese e na Secretaria do Programa de Pós Graduação em Biologia Animal da Unidade Instituto de Biologia. 5 DEDICATÓRIA Dedico este trabalho primeiramente а Deus, ао mеυ pai Mauro, minha mãе Silvana, minha irmã e melhor amiga Roberta e ao meu noivo Victor que, cоm muito carinho е apoio, nãо mediram esforços para qυе еυ chegasse аté esta etapa dе minha vida. 6 AGRADECIMENTOS À Universidade Estadual de Campinas (UNICAMP) pela infraestrutura oferecida durante o desenvolvimento da pesquisa e durante meus anos de graduação. Ao Programa de Pós-Graduação em Biologia Animal da UNICAMP, por colaborar com a minha qualificação profissional e crescimento pessoal. À Fundação de Amparo à Pesquisa de São Paulo (FAPESP- 2014/23856-0; 2018/10313-0) pelo apoio financeiro ao projeto. Ao Prof. Dr. André R. S. Garraffoni pela orientação, apoio, confiança, paciência e todo o conhecimento passado para que eu pudesse realizar o meu projeto. Aos meus pai, Mauro Fassina, que apesar de seu silêncio sempre foi um grande incentivador de todas as minhas conquistas pessoais e profissionais. À minha mãe, Silvana Visnardi Fassina que me ensinou o amor pelo conhecimento e pelo ensinar. À minha irmã Roberta Visnardi Fassina por ser uma grande incentivadora do meu mestrado e das minhas decisões. Ao meu noivo, Victor de Backer Moura, que foi um grande apoiador na minha decisão de voltar a Unicamp depois de 3 anos de formada e seguir o meu sonho de ser mestre. Aos amigos do LEOM (Laboratório de Evolução de Organismos Meiofaunais) pelo companheirismo e apoio, e por me fazer sentir parte deste grupo mesmo estando longe fisicamente. Aos pesquisadores Dr. Rodolfo Corrêa de Barros, Dra. Cláudia M.L. Assunção e Érika Cavalcante Leite dos Santos por aceitarem o convite para serem a pré-banca desta dissertação e pelas valiosas contribuições ao texto. À amiga Ariane Campos que foi a maior incentivadora da minha volta à vida acadêmica e esteve presente em todos os momentos desta jornada. Aos pesquisadores in memoriam Ernest Gustav Gotthelf Marcus, Eveline du ​ Bois-Reymond Marcus e Rosina de Barros que, devido ao seu esforço e perseverança colocaram a pedra fundamental da tardigradologia brasileira e nos possibilitam "ver mais longe, por estar sobre os ombros de gigantes". À Deus, por me dar saúde e força para superar todas as dificuldades. 7 O presente trabalho foi realizado com apoio da Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Código de Financiamento 001. 8 I am among those who think that science has great beauty. (Marie Curie) If I have seen further it is by standing on the shoulders of Giants. (Isaac Newton) 9 RESUMO O filo Tardigrada é caracterizado por conter espécimes diminutos (70 a 1000 μm), possuírem um corpo robusto em forma de barril coberto por uma cutícula proteica, quatro pares de patas, garras no formato de gancho ou discos adesivos e um aparato buco-faringeano complexo. Os representantes desse grupo são comumente reconhecidos pela alta resistência em condições extremas de temperatura e pressão. O filo é composto por quase 1300 espécies, encontradas nos mais diversos ecossistemas terrestres, marinhos e de água doce no mundo todo. O estudo dos tardígrados está concentrado nas regiões Neártica e Paleoártica, pois, historicamente, foi onde estavam alocados os principais pesquisadores e em outras regiões biogeográficas os estudos são mais pontuais. Os tardígrados limnoterrestres brasileiros foram estudados de forma pioneira e intensa nas décadas de 30 a 50, por Rosina de Barros, Eveline Du Bois-Reymond Marcus e Ernest Marcus. Apesar desse pioneirismo, com mais de 10 artigos publicados em poucos anos, os pesquisadores não designaram nenhum material tipo e os trabalhos apresentam desenhos e descrições morfológicas pertinentes à época. A partir da utilização de técnicas modernas de obtenção de informação da morfologia externa e da anatomia (microscopia óptica de luz com contraste de interferência diferencial- DIC e microscopia eletrônica de varredura), moleculares (extração de genes 18S e COI) e coletas realizadas nas localidades tipo (Campos do Jordão, São Paulo) o presente estudo apresenta a redescrição da espécie Pseudechiniscus juanitae De Barros ​ 1939 e analisa a identidade da espécie Pseudechiniscus novaezeelandiae marinae ​ Bartoš, 1934 registrada e brevemente descrita por De Barros (1942). 10 ABSTRACT Tardigrada is characterized by small individuals (70 a 1000 μm) and a robust ​​ barrel-shaped body, covered by a proteic cuticle. Representatives of this group are famous for their resistance to extreme conditions of temperature and pressure. The phylum is composed by more than 1300 species, found in several terrestrial, marine and freshwater ecosystems worldwide. The study of the group is concentrated in the Nearctic and Palearctic regions, once it is where the main researchers are allocated. The limnoterrestrial brazilian tardigrades were extensively studied between the 1930s and 1950s by the researchers Rosina de Barros, Eveline Du Bois-Reymond Marcus and Ernest Marcus. Despite over 10 articles published in a few years, the researchers did not designate any type of material and the descriptions present only drawings and limited morphological information. Using modern techniques for obtaining information on external morphology and anatomy (DIC- differential interference contrast and SEM-Scanning Electron Microscopy), molecular studies (18S and COI) and sampling in the type location, this study presents a redescription of the species Pseudechiniscus juanitae De Barros 1939 and analyses the identity of ​ the species Pseudechiniscus novaezeelandiae Bartoš, 1934 registered and briefly ​ described by De Barros (1942). 11 SUMÁRIO I- INTRODUÇÃO 12 1.O filo Tardigrada 12 2. Resistência dos tardígrados 13 3. Biodiversidade e sistemática 14 4. Filogenética e registros fósseis 19 5. Dificuldades Taxonômicas 21 6. Taxonomia Integrativa e o conhecimento do grupo no Brasil 21 II- OBJETIVOS 23 III- REFERÊNCIAS BIBLIOGRÁFICAS 24 IV- CAPÍTULO 1 32 Redescription of Pseudechiniscus juanitae de Barros, 1939 and comments on the identity of Pseudechiniscus novaezeelandiae marinae Bartoš, 1934 sensu De Barros (1942) (Echiniscidae, Heterotardigrada)* 32 V- APÊNDICE 66 12 I- INTRODUÇÃO 1.O filo Tardigrada O filo Tardigrada é composto por microinvertebrados aquáticos (de 70 até aproximadamente que 1000 μm), distribuídos mundialmente e comumente encontrados em habitats terrestres e no mesobentos de ambientes aquáticos, de zonas tropicais a desertos áridos polares, de topos de montanhas a profundidades abissais nos oceanos (Kaczmarek et al. 2015; Nelson 2002). Apesar de serem encontrados no solo ou em musgos e liquens existentes na casca das árvores, esses organismos são considerados animais aquáticos independentemente de seu habitat, uma vez que necessitam de um filme d’água ao redor do corpo para realizar sua locomoção e respiração (Marley et al. 2011). O corpo diminuto e robusto dos tardígrados apresenta uma complexa morfologia, dividido em 5 segmentos com segmentação externamente indistinta: o segmento cefálico (cabeça), três segmentos torácicos e um segmento terminal. Cada segmento, com exceção do cefálico, possui um par de lobópodes (pernas sem articulação) com garras ou dedos com discos adesivos e o segmento caudal possui o quarto par de pernas voltadas posteriormente (Nelson 2002). A parede do corpo consiste em três camadas, sendo a mais externa uma cutícula acelular com α-quitina podendo formar escleritos articulados, seguida por ​ ​ uma
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