Crenicichla Ploegi, a New Species of Pike-Cichlid of the C. Saxatilis Group

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Crenicichla ploegi, a new species of pike-cichlid of the C. saxatilis group from the Rio Juruena and upper Rio Paraguai basins in Brazil, with an updated diagnosis and biogeographi...

Article in Zootaxa · February 2018 DOI: 10.11646/zootaxa.4377.3.3

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Henrique Varella Marina Loeb Universidade Federal da Bahia University of São Paulo

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Crenicichla ploegi, a new species of pike-cichlid of the C. saxatilis group from the Rio Juruena and upper Rio Paraguai basins in Brazil, with an updated diagnosis and biogeographical comments on the group (Teleostei: Cichlidae)

HENRIQUE R. VARELLA1, 2, 5, MARINA V. LOEB2, FLÁVIO C.T. LIMA3 & SVEN O. KULLANDER4 ¹Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Campus de Ondina, Rua Barão de Geremoabo s/n, 40170-115 Salvador, BA, Brazil. ²Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04218–970 São Paulo, SP, Brazil. 3Museu de Zoologia da Universidade Estadual de Campinas "Adão José Cardoso", Caixa Postal 6109, 13083-970 Campinas, SP, Bra- zil. 4Department of Zoology, Swedish Museum of Natural History, POB 50007, SE-104 05 Stockholm, Sweden. 5Corresponding author. E-mail: [email protected]

Abstract

Crenicichla ploegi, new species, is described based on material from the Rio Juruena (Rio Papagaio, Rio do Sangue, Rio Arinos, and Rio Juruena itself) and from tributaries of the upper Rio Paraguai (upper Rio Jauru, upper Rio Cabaçal and upper Rio Sepotuba) in Mato Grosso, Brazil. Crenicichla ploegi is the twenty-third species of the C. saxatilis group. It is distinguished from all other species of the group by the presence of dark spots and vermiculations on snout, interorbital area and dorsally on head in adults vs. dark markings absent or present only in juveniles but absent in adults, and by the presence of a deep and strongly pigmented lateral band present in both juveniles and adults, occupying a depth of 4–6 horizontal scale rows vs. lateral band narrower, occupying a depth of 2–3 horizontal scale rows, and conspicuous only in juveniles and smaller specimens, faded or absent in larger specimens. Diagnostic characteristics of the C. saxatilis group proposed in previous studies were discussed to update the diagnosis of the group, and morphological comparisons among the species included in the group, with biogeographical comments, are provided. The presence of conspicuous dark markings dorsally on the head and the presence of a prominent midlateral band are hypothesized to be paedomorphic characteristics, retained from juvenile conditions in the context of the species of the C. saxatilis group.

Key words: biogeography, coloration, paedomorphy, species groups

Resumo

Crenicichla ploegi, nova espécie, é descrita baseada em material de afluentes do Rio Juruena (Rio Papagaio, Rio do Sangue, Rio Arinos e do próprio Rio Juruena) e do alto Rio Paraguai (alto Rio Jauru, alto Rio Cabaçal e alto Rio Sepotuba) no estado de Mato Grosso, Brasil. Crenicichla ploegi é a vigésima-terceira espécie descrita do grupo C. saxatilis de espé- cies. Ela pode ser distinguida de todas as demais espécies do grupo pela presença de manchas escuras e vermiculações no focinho, região interorbital e dorsalmente na cabeça em adultos vs. marcas escuras ausentes ou presentes apenas em juvenis mas ausentes em adultos, e pela presença de uma larga faixa lateral fortemente pigmentada tanto em juvenis como adultos, ocupando uma altura de 4–6 séries horizontais de escamas vs. faixa lateral mais estreita, ocupando uma altura de 2–3 séries horizontais de escamas, e conspícuas apenas em juvenis e exemplares pequenos, apagada ou ausente em exemplares grandes. Características diagnósticas do grupo C. saxatilis propostas em estudos prévios foram discutidas a fim se atualizar a diagnose do grupo e comparações morfológicas entre as espécies incluídas no grupo, com comentários bio- geográficos, são oferecidas. Levanta-se a hipótese de que a presença de manchas escuras conspícuas na região dorsal da cabeça e a presença de uma banda médio-lateral proeminente no flanco de C. ploegi sejam características pedomórficas, retidas de condições juvenis no contexto do grupo C. saxatilis.

Palavras chaves: biogeografia, coloração, pedomorfia, grupos de espécies

Accepted by J. Armbruster: 4 Dec 2017; published: 2 Feb. 2018 361 Introduction

The South American cichlid genus Crenicichla Heckel, 1840, with 90 species (Kullander, 2003; Kullander & Varella, 2015), ranks as the most species-rich cichlid genus next to Apistogramma Regan, with 92 species (Kullander, 2003; Varella & Britzke, 2016; Römer et al. 2017). The division of Crenicichla into less inclusive, diagnosable groups has been a common procedure to make taxonomic revisions of the genus practicable. Kullander (1981, 1982) suggested the existence of at least two groups in the genus: C. lacustris and C. lepidota species groups, respectively. Kullander (1982: 654) compared the species placed in the Crenicichla lepidota species group with other forms related to Crenicichla saxatilis from the Amazon, Orinoco and Guyana basins, and tentatively separated the two assemblages by scale and fin counts. However, he highlighted the similarities between them in relation to color pattern, including the presence of a humeral blotch, and to sexual dimorphism, which led him to assume that they may be closely related. Ploeg (1991) proposed five groups of species within the genus, viz. C. saxatilis, C. lugubris, C. reticulata, C. lacustris and C. wallacii groups, the latter one also including the species of Teleocichla Kullander, 1988. The Crenicichla saxatilis group sensu Ploeg (1991) encompassed all species of Kullander´s C. lepidota group and C. saxatilis-like forms, plus C. guentheri Ploeg, 1991. In Ploeg’s (1991: 17) key to the species groups, the C. saxatilis group was diagnosed by the absence of a postorbital blotch (presence of a continuous postorbital stripe), frequent presence of a suborbital marking (instead of absent), number of scales below lateral lines usually less than 86, and presence of a humeral blotch (instead of absence). Ploeg (1991: 18) presents several other general characteristics that may be diagnostic for the group, but these have not been objectively compared among the species groups proposed by him. The Crenicichla saxatilis group has been corroborated as monophyletic by recent molecular phylogenies (Kullander et al., 2009; Piálek et al., 2012), although they had included only three and two specific-taxa belonging to the group, respectively. Varella et al. (2012) removed C. guentheri, which is junior synonym of C. hemera Kullander 1990, from the C. saxatilis group based on morphological characters. Both nominal species were absent from previous molecular analyses. The authors considered C. hemera and C. chicha Varella, Kullander & Lima, 2012 as sister-taxa that could not be promptly allocated in any of the other groups previously proposed in Crenicichla. With exception of these two species, the current composition of the C. saxatilis group has remained essentially that given by Ploeg (1991, Table 1), but its diagnosis has not been updated. Pike cichlids of the C. saxatilis group are widespread in almost all major rivers in cis-Andean South America, with exception of coastal river drainages of southern and southeastern Brazil (Kullander & Lucena, 2006), and are more commonly found in lacustrine environments or small streams. In this paper, we describe a new species of Crenicichla of the C. saxatilis group based on material from the Rio Juruena basin (a tributary of the upper Rio Tapajós) and the upper Rio Paraguai basin, Mato Grosso state, Brazil. It is the first species of this group described since Ploeg´s (1991) taxonomic revision of the genus and of interest both for being present in two different major river systems and for its distinctive, apparently paedomorphic colour pattern. Additionally, we provide an updated diagnosis for the Crenicichla saxatilis group and taxonomic and biogeographic comments on it.

Material and methods

Specimen lengths are given as Standard Length (SL), measured from the tip of the upper jaw to the middle of the base of the caudal fin. Measurements and counts were taken as described by Kullander (1986) and presented as percentage of SL. Statistical analysis of covariance (non-parametric robust ANCOVA) was performed using the software R v. 2.10.0 (R Development Core Team, 2009) and packages for statistical analyses (e.g. npsm; Kloke & McKean, 2014a, 2014b), with the raw morphometric data of all specimens and only adults specimens separately. Scales in a longitudinal row (E1 scales row) are counted in the row immediately dorsal to that containing the lower lateral line. Color marking terminology follows mainly Kullander (1986), Kullander & Nijssen (1989) and Ploeg (1991). Vertebral counts and osteological observations were based on three specimens (MZUSP 108380, 59.2–67.7 mm SL) cleared and stained (c&s) following the method of Taylor & Van Dyke (1985) and on x-rays of three other specimens (NRM 26098, 3, 54.1–84.6 mm SL). The lower pharyngeal tooth plate is measured as described by

362 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. Barel et al. (1977). The illustrations of the pharyngeal tooth plate and the infraorbitals were based on the cleared and stained specimens, photographed under a stereo microscope and further manipulated with image editing software. Nomenclature for individual ossicles of the infraorbital series follows López-Fernández et al. (2005: 645, fig. 11A).

TABLE 1. Valid and nominal species considered herein as belonging to the Crenicichla saxatilis species group. Valid species Nominal species 1. Crenicichla albopunctata Crenicichla saxatilis albopunctata Pellegrin, 1904 2. Crenicichla alta Crenicichla alta Eigenmann, 1912 Crenicichla vaillanti Pellegrin, 1903 Crenicichla pterogramma Fowler, 1914 Crenicichla cardiostigma Ploeg, 1991 3. Crenicichla anthurus Crenicichla anthurus Cope, 1872 4. Crenicichla brasiliensis Perca brasiliensis Bloch, 1792 Crenicichla menezesi Ploeg, 1991 5. Crenicichla britskii Crenicichla britskii Kullander, 1982 6. Crenicichla coppenamensis Crenicichla coppenamensis Ploeg, 1987 7. Crenicichla frenata Crenicichla frenata Gill, 1858 8. Crenicichla hummelincki Crenicichla hummelincki Ploeg, 1991 9. Crenicichla inpa Crenicichla inpa Ploeg, 1991 10. Crenicichla isbrueckeri Crenicichla isbrueckeri Ploeg, 1991 11. Crenicichla labrina Cychla labrina Spix & Agassiz, 1829 12. Crenicichla lepidota Crenicichla lepidota Heckel, 1840 Crenicichla edithae Ploeg, 1991 13. Crenicichla lucius Crenicichla lucius Cope, 1870 14. Crenicichla nickeriensis Crenicichla nickeriensis Ploeg, 1987 15. Crenicichla pellegrini Crenicichla pellegrini Ploeg, 1991 16. Crenicichla proteus Crenicichla proteus Cope, 1872 Crenicichla proteus argynnis Cope, 1872 Batrachops nemopterus Fowler, 1940 Crenicichla nijsseni Ploeg, 1991 17. Crenicichla pydanielae Crenicichla pydanieae Ploeg, 1991 18. Crenicichla santosi Crenicichla santosi Ploeg, 1991 19. Crenicichla saxatilis Sparus saxatilis Linnaeus, 1758 Scarus biocellatus Walbaum, 1792 Scarus pavo La Cepède, 1802 Scarus pavonius Gray, 1854 20. Crenicichla semicincta Crenicichla saxatilis semicincta Steindachner, 1892 Crenicichla clancularia Ploeg, 1991 21. Crenicichla sipaliwini Crenicichla sipaliwini Ploeg, 1987 22. Crenicichla sveni Crenicichla sveni Ploeg, 1991

Acronyms of ichthyological collections depository of the type specimens and comparative material: ANSP, The Academy of Natural Sciences, Philadelphia, USA; BMNH, Natural History Museum (former British Museum of Natural History), London, UK; CAS, California Academy of Sciences, San Fracisco, USA; FMNH, Field Museum of Natural History, Chicago, USA; INPA, Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil;

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 363 IRSNB, Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium; LBP, Laboratório de Biologia e Genética de Peixes, Universidade Estadual Paulista, Botucatu, Brazil; LIRP, Laboratório de Ictiologia, Universidade de São Paulo, Ribeirão Preto, Brazil; MCP, Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil; MHNCI, Museu de História Natural do Capão do Imbuia, Curitiba, Brazil; MHNG, Museum d'Histoire Naturelle, Ville de Genève, Genève, Switzerland; MNHN, Muséum National D´Histoire naturelle, Paris, France; MZUSP, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil; NMW, Naturhistorisches Museum, Wien, Austria; ZMA, Zoölogisch Museum, Amsterdam and RMNH, Rijksmuseum van Natuurlijke Histoire (RMNH), both housed in the Naturalis Biodiversity Center, Leiden; NRM, Swedish Museum of Natural History, Stockholm, Sweden; NUP, Núcleo de Pesquisa em Limnologia, Ictiologia e Aquicultura da Universidade Estadual de Maringá, Maringá, Brazil; ZUEC, Museu de Zoologia da Universidade Estadual de Campinas, Campinas, Brazil.

Results

Crenicichla ploegi, new species (Figs. 1, 2, 3a–c, 4, 5, 6a, 8 and 10b; Tab. 2)

Holotype. MZUSP 120714, male, 89.8 mm SL, Sapezal, Rio Papagaio, large river pool in the Fazenda Savocal (near road to Brasnorte), 12°46'04"S 58°23'05"W, 8–9 Oct 2006, F. A. Machado, F. C. T. Lima, C. M. C. Leite & N. E. Silva. Paratypes. Brazil. Mato Grosso. Rio Tapajós basin (168 specimens: 27.0–190.4 mm SL). LBP 7953, 4, 38.3–68.8 mm SL, Nova Mutum, Rio dos Patos, 13°48'03"S 056°01'38"W, 23 Jan 2009, C. Oliveira et al. LBP 7966, 2, 58.4–59.4 mm SL, Nova Mutum, tributary of Rio dos Patos, 13°48'13"S 56°09'25"W, 23 Jan 2009, C. Oliveira et al. LBP 7979, 12, 35.6–107.1 mm SL, Nova Mutum, tributary of Rio dos Patos, 13°49'08"S 56°10'53"W, 23 Jan 2009, C. Oliveira et al. LBP 7999, 1, 85.5 mm SL, Nova Mutum, Rio Igarapé, 14°08'14"S 56°05'48"W, 24 Jan 2009, C. Oliveira et al. LBP 13259, 5, 52.9–117.3 mm SL, Diamantino, riacho Alegre, 13°59'04"S 057°04'01"W, 10 Aug 2011, I. Camaragibe & J. Pacheco. LBP 19604, 4, 43.6–63.0 mm SL, Campo Novo dos Parecis, Rio Verde, 13°37'11"S 58°00'49"W, 21 Jun 2009, R. Devidé et al. MCP 30282, 2, 27.3–44.9 mm SL, São José do Rio Claro, stream near the road MT-010, Rio Arinos basin, 13°37'23"S 056°29'18"W, 17 Jan 2002, R. E. Reis et al. MCP 30295, 1, 59.0 mm SL, São José do Rio Claro, Rio Claro or Rio Água Verde, 12 km Southeast from São José do Rio Claro, road MT-010, 13°30'12"S 056°37'08"W, 17 Jan 2002, R. E. Reis et al. MCP 30311, 1, 87.5 mm SL (measured and counted), Tapurah, stream at road MT-338, 46 km North from Tapurah, 12°23'14"S 56°41'54"W, 19 Jan 2002, R. E. Reis et al. MCP 38400, 1, 41.2 mm SL, Sapezal, córrego Água Quente, road between Sapezal and Rio Papagaio, 13°32'25"S 058°43'32"W, 13 Jul 2004, R. E. Reis et al. MCP 38423, 3, 47.4–52.5 mm SL, Campos de Júlio, Rio Juruena, at road from Campos de Júlio to Sapezal, 13°33'05"S 059°02'03"W, 13 Jul 2004, R. E. Reis et al. MCP 39528, 1, 69.4 mm SL (measured and counted), Sapezal, upstream dam in Rio Sauê-Uiná, road BR-364 toward Sapezal, 13°32'22"S 058°50'48"W, 13 Jul 2004, P. Lehmann et al. MCP 39559, 1, 61.3 mm SL (measured and counted), Sapezal, Rio Sauê-Uiná dam, at road BR-364, 13°32'26"S 058°50'47"W, 13 Jul 2004, P. Lehmann et al. MCP 42678, 2, 32.6–32.8 mm SL, Sapezal, Rio Sacre, 13°36'52"S 058°05'38"W, 3 Aug 2007, V. Lampert & M. Teixeira. MCP 42680, 33, 19.4–23.6 mm SL, Sapezal, Rio Verde, 13°38'34"S 58°01'04"W, 8 Mar 2007, V. Lampert & M. Teixeira. MCP 42771, 1, 77.6 mm SL, Rio Sacre, 13°36'52"S 058°05'40"W, 7 Jun 2007, V. Lampert. MCP 42673, 2, 40.3–49.4 mm SL (1 measured and counted: 49.4 mm SL), Sapezal, Rio Sacre, 13°36'52"S 058°05'38"W, 3 Aug 2007, V. Lampert & M. Teixeira. MZUSP 61081, 2, 82.4–86.5 mm SL (measured and counted), Nova Mutum, Fazenda Buriti, unnamed stream of the Rio Arinos basin, 13°51'52"S 56°11'36"W, 16 Feb 2000, F.C.T. Lima et al. MZUSP 61099, 2, 37.1–53.6 mm SL (measured and counted), Nova Mutum, Rio Criquiri, tributary of Rio dos Patos, 13º48'10"S 56º09'29"W, 19 Feb 2000, F. C.T. Lima et al. MZUSP 82075, 1, 55.8 mm SL, Brasnorte, hydroelectric dam Bocaíva, Rio Cravari, tributary of Rio do Sangue, 3 Jan 2003, K. de Silimon. MZUSP 93548, 6, 39.3–94.8 mm SL (4 measured and counted: 34.7–94.8 mm SL), Sapezal, Rio Papagaio and at the mouth of the Rio Buriti, close of the bridge at the road between Sapezal and Brasnorte, 12°47'06"S 58°23'05"W, 7–9 Oct 2006, F. A. Machado et al. MZUSP 93697, 4, 65.5–118.7 mm SL (measured and counted), collected with holotype. MZUSP 108368, 1, 43.5 mm SL, Sapezal,

364 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. Rio Juruena, upstream hydroelectric dam Santa Lúcia, 13°39'18"S 59°01'12"W, 24 Sep 2006, K. de Silimon. MZUSP 108369, 1, 27.6 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Segredo, 13°14'54"S 59°01'43"W, 3 Sep 2006, K. de Silimon. MZUSP 108370, 2, 48.9–67.6 mm SL (1 measured and counted, 67.6 mm SL), Sapezal, Rio Juruena, downstream hydroelectric dam Santa Lúcia, 13°32'37"S 59°01'48"W, 25 Sep 2006, K. de Silimon. MZUSP 108371, 2, 29.7–66.2 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Telegráfica, 12°41'22"S 58°56'47"W, 15 Sep 2006, K. de Silimon. MZUSP 108372, 4, 24.6–62.5 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Rondon, 12°53'38"S 58°55'11"W, 20 Sep 2006, K. de Silimon. MZUSP 108373, 1, 41.6 mm SL, Campos de Júlio, Rio Juruena, Fazenda Tiroleza, 14°16'38"S 59°05'22"W, 19 Sep 2006, K. de Silimon. MZUSP 108374, 2, 49.8–61.4 mm SL, Campos de Júlio, Rio Juruena, Fazenda Tiroleza, 14°16'38"S 59°05'22"W, 19 Sep 2006, K. de Silimon. MZUSP 108375, 2, 45.9–46.0 mm SL, Campos de Júlio, pool between Rio Juruena and riacho Cabeceira, near road BR-364, 14°39'43"S 59°06'27"W, 18 Sep 2006, K. de Silimon. MZUSP 108376, 1, 47.2 mm SL, Campos de Júlio, Rio Juruena, near road BR-364, 14°39'40"S 59°06'27"W, 18 Sep 2006, K. de Silimon. MZUSP 108377, 3, 27.6–77.7 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Telegráfica, 12º37'32"S 58º55'36"W, 14 Sep 2007, K. de Silimon. MZUSP 108380, 4, 27.0–57.3 mm SL, 3 c&s, 59.2–67.7 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Ilha Comprida, 13°11'32"S 58°58'51"W, 27 Sep 2006, K. de Silimon. MZUSP 108381, 1, 69.4 mm SL (measured and counted), Campos de Júlio, Rio Juruena, downstream bridge at the road BR-364, 14°39'40"S 59°06'27"W, 18 Sep 2006, K. de Silimon. MZUSP 108382, 1, 43.8 mm SL, Sapezal, Rio Juruena, upstream hydroelectric dam Santa Lúcia, left margin, 13°37'55"S 59°03'03"W, 24 Sep 2006, K. de Silimon. MZUSP 108383, 1, 53.3 mm SL, Campos de Júlio, Rio Juruena, Fazenda Tiroleza, upstream airport, 14°15'47"S 59°05'25"W, 19 Sep 2006, K. de Silimon. MZUSP 108384, 1, 43.3 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Segredo, 13°13'47"S 59°01'59"W, 27 Sep 2006, K. de Silimon. MZUSP 108385, 1, 49.4 mm SL, Campos de Júlio, tributary of the Rio Juruena, near road BR-364, 14°39'40"S 59°06'27"W, 18 Sep 2006, K. de Silimon. MZUSP 108386, 1, 50.5 mm SL, Campos de Júlio, Rio Juruena, near road BR-364, 14°39'00"S 59°06'00"W, 17 Sep 2006, K. de Silimon. MZUSP 108387, 1, 56.4 mm SL, Sapezal, Rio Juruena, upstream hydroelectric dam Sapezal, right margin, 13°16'06"S 59°01'27"W, 22 Sep 2006, K. de Silimon. MZUSP 108388, 1, 50.3 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Ilha Comprida, 13°12'06"S 58°59'03"W, 26 Sep 2006, K. de Silimon. MZUSP 108389, 2, 37.2– 41.9 mm SL, Sapezal, Rio Juruena, upstream hydroelectric dam Santa Lúcia, 13°34'49"S 59°01'48"W, 24 Sep 2006, K. de Silimon. MZUSP 108390, 1, 30.5 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Sapezal, right margin, 13°14'47"S 59°00'52"W, 22 Sep 2006, K. de Silimon. MZUSP 108391, 2, 27.4–28.1 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Telegráfica, 12°41'05"S 58°56'29"W, 14 Sep 2006, K. de Silimon. MZUSP 108392, 1, 27.6 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Rondon, 12°53'38"S 58°55'11"W, 20 Sep 2006, K. de Silimon. MZUSP 108393, 1, 31.5 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Telegráfica, 12°39'51"S 58°55'53"W, 14 Sep 2006, K. de Silimon. MZUSP 108395, 1, 24.6 mm SL, Sapezal, Rio Juruena, downstream hydroelectric dam Telegráfica, 12°41'02"S 58°56'26"W, 14 Sep 2006, K. de Silimon. MZUSP 115689, 6, 45.8–62.3 mm SL, Campos de Júlio, Rio Juína, road between Comodoro and Campos de Júlio, 13°47'41.2"S 59°27'19.6"W, 29 Sep 2013, O. T. Oyakawa et al. NRM 16872, 1, 52.0 mm SL, Rio Capitao Elio, tributary to Rio do Sangue, where crossed by MT-170 road North of Brasnorte, 12 Oct 1989, S. O. Kullanderet al. NRM 26098. 3, 54.1–84.6 mm SL, Rio Juruena at ferry crossing 60 km East of Juína, 11°30'S 58°40'W, 14 Oct 1989, S. O. Kullander et al. NUP 11630, 2, 62.0–101.2 mm SL (measured and counted), Diamantino, Rio Preto, tributary of the Rio Arinos 14º07'46.6"S 56º42'15.3"W, 15 Sep 2010, C. H. Zawadzki. ZUEC 10053, 8, 43.2–146.1 mm SL (4 measured and counted: 96.9–146.1 mm SL), São José do Rio Claro, Rio Claro, above road MT-235, 13°52'S 56°41'W, 4–12 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10091, 1, 116.3 mm SL, São José do Rio Claro, Rio Claro, below road MT-235, 13°53'30"S 56°41'1"W, 4–5 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10103, 2, 124.3–164.4 mm SL (measured and counted), São José do Rio Claro, Rio Claro, below road MT-235, 13°47'38"S 56°42'10"W, 11 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10144, 4, 41.4–190.4 mm SL (3 measured and counted: 115.9–190.4 mm SL), São José do Rio Claro, Rio Claro, below road MT-235, 13°50'43"S 56°41'46"W, 8–11 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10179, 2, 34.1–123.7 mm SL (1 measured and counted: 123.7 mm SL), São José do Rio Claro, Rio Claro, above road MT- 235, 13°51'23"S 56°41'11"W, 4–6 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10181, 3, 55.6–142.0 mm SL, São José do Rio Claro, Rio Claro, below road MT-235, 13°48'20"S, 56°41'34"W, 4–10 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10244, 6, 41.7–97.3 mm SL, São José do Rio Claro, Rio Claro, below road MT-

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 365 235, 13°47'57"S 56°41'50"W, 10–15 May 2007, I. M. Fernandes & G. M. Alencar. ZUEC 10290, 1, 69.8 mm SL, São José do Rio Claro, córrego da Sucuri, tributary of the Rio Claro, 13°50'53"S 56°41'51"W, 8–9 May 2007, I. M. Fernandes & G. M. Alencar. Rio Paraguai basin (11 specimens: 48.4–127.2 mm SL). LBP 7540, 5, 52.7–86.7 mm SL (measured and counted), Campo Novo dos Parecis, Rio Verde, 14°33'24"S 57°48'43"W, 16 Jul 2008, W.P. Troy. MZUSP 78721, 1, 48.4 mm SL (measured and counted), Reserva do Cabaçal, Rio Juba, at the confluence with Rio São José, 14°48'16"S 58°17'56"W, 7 Mar 2002, F.C.T. Lima et al. MZUSP 78847, 1, 76.4 mm SL (measured and counted), Jauru, Rio Jauru, Fazenda do Sr. Salu, upstream of Salto da Fumaça, 15°11'04"S 58°43'43"W, 3–4 Mar 2002, F. A. Machado et al. MZUSP 78884, 3, 105.2–127.2 mm SL (measured and counted), Tangará da Serra, lake in the Rio Juba, 14°45'S 58°3'W, 18 May 1996, G. Borges. MZUSP 82067, 1, 88.3 mm SL (measured and counted), Arapatinga, Rio Jauru, downstream hydroelectric dam Alto Jauru, ca. 15°18'S 58°42'W, 11 Dec 2002, K. de Silimon.

FIGURE 1. Crenicichla ploegi: a) ZUEC 10053, paratype, adult male, 143.3 mm SL, Rio Claro, Rio Juruena basin; b) MZUSP 93697, holotype, adult male, 89.8 mm SL, Rio Papagaio, Rio Juruena basin; c) MCP 39528, adult female, 69.4 mm SL, Brazil, Rio Sauê-Uiná, Rio Juruena basin; d) MCP 42680, juvenile, 23.0 mm SL, Rio Verde, Rio Juruena basin.

366 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. FIGURE 2. Crenicichla ploegi: a) MZUSP 78884, paratype, 127.2 mm SL, Rio Juba, upper Rio Paraguai basin; b) LBP 7540, paratype, 85.1 mm SL; c) LBP 7540, paratype, 66.0 mm SL; and d) LBP 7540, 52.7 mm SL, Campo Novo dos Parecis, upper Rio Paraguai basin.

Diagnosis. Crenicichla ploegi is placed in the C. saxatilis species group as diagnosed here (see Discussion) by having a humeral blotch even in the smallest specimens observed, less than 80 scales in the E1 row (58–71), dimorphic males with many light dots scattered on the flank (Fig. 5), and infraorbital series with very slender lacrimal bone and five separate post-lacrimal tubular infraorbitals (Fig. 6a). It is distinguished from all other species of the C. saxatilis group by the presence of dark spots and vermiculations on snout, interorbital area and dorsally on head in many adults up to 146.6 mm SL (Fig. 3b, c) vs. dark markings absent or present only in juveniles but absent in adults (head uniformly dark gray in adults from 80 mm SL; Fig. 3e), and by the presence of a deep and strongly pigmented lateral band present in both juveniles and adults, occupying a depth of 4–6 horizontal scale rows vs. lateral band narrower, occupying a depth of 2–3 horizontal scale rows, and conspicuous only in juveniles and smaller specimens, faded or absent in larger specimens. In addition, Crenicichla ploegi is more elongate (body depth 17.8–21.7% SL) and has more scales in the E1 row (58–71 scales) than C. brasiliensis, C. britskii, C. frenata, C. isbrueckeri, C. labrina, C. lepidota, C.

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 367 proteus, C. santosi, C. saxatilis and C. semicincta (body depth more than 21% SL [Table 3, Fig. 9], and fewer than 56 scales in the E1 row). Crenicichla ploegi differs from C. alta, C. albopunctata, C. anthurus, C. coppenamensis, C. pydanielae and C. sipaliwini by having the humeral blotch situated almost entirely below the anterior branch of the lateral line (Fig. 10b) vs. humeral blotch with center on the level of the anterior branch of the lateral line (Fig. 10c).

FIGURE 3. Schematic drawing of dorsal view of the body in Crenicichla ploegi and Crenicichla aff. lepidota Tapajós, showing the dark markings in fishes with different sizes. a) Crenicichla ploegi, MCP 42680, 23.0 mm SL; b) Crenicichla ploegi, LBP 7540, 52.7 mm SL; c) Crenicichla ploegi, ZUEC 10053, 143.3 mm SL; d) Crenicichla aff. lepidota Tapajós, LBP 8008, juvenile, 21.6 mm SL; e) Crenicichla aff. lepidota Tapajós, LBP 8020, 92.4 mm SL.

368 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. FIGURE 4. Crenicichla ploegi, a male (left) and a female (right) photographed alive in the Rio Verde, tributary of the Rio Papagaio, Mato Grosso, Brazil. Photo by Marcelo Krause.

FIGURE 5. Crenicichla ploegi, uncatalogued, 60.0 mm SL, from the Rio Juba above the waterfall of Cachoeira do Juba, tributary of the Rio Sepotuba basin, upper Rio Paraguai, Tangará da Serra, Mato Grosso, Brazil (field number SACIB2016100103). Photographed alive in aquarium just after capture by Fernando Dagosta.

Description. Largest male 190.4 mm SL, largest female 115.9 mm SL. Refer to Figs. 1, 2, 4 and 5 for general aspect. Morphometric data in Table 2. Head slightly deeper than wide. Caudal peduncle longer than deep, or with approximately equal length and depth (caudal peduncle depth 61.0–98.2% caudal peduncle length). Snout short, rounded in dorsal view, pointed in lateral view. Lower jaw prognathous, its articulation with quadrate posterior to middle of orbit, reaching vertical through posterior margin of orbit in some specimens; posterior rear of maxilla almost reaching vertical through anterior margin of orbit. Ascending premaxillary processes reaching to about one-half the orbit diameter. Lower lip folds separate anteriorly, folds of upper lip discontinuous, with a symphyseal thickening. Postlabial skin fold margin truncate. Orbit supralateral, visible from below, entirely in anterior half of head. Interorbital area flat, narrower than mouth. Nostril dorsolateral, closer to anterior margin of orbit than to tip of postlabial skin fold,

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 369 with small tubular margin exposed, but no anterior marginal membranaceous skin flap. Posterior margin of preopercle regularly serrated or with projections on its superior portion in larger specimens. Posterior margin of supracleithrum smooth. Lateralis pores on head simple. Infraorbital series with one lacrimal (interpreted as lacrimals 1+2), with four pores, and five tubular post- lacrimal infraorbitals (3–7); lacrimal slender (length 26.4–28.9% depth); infraorbital 7 curved posteriorly (Fig. 6a). Flank scales ctenoid. Scales cycloid on head, on dorsum above anterior half of upper lateral line, along dorsal- fin base, on chest, and on belly below line from lower edge of pectoral-fin base to anal-fin origin. Predorsal scales small, embedded in skin, extending forward to transverse frontal lateralis canal; prepelvic scales smaller than predorsal ones. Cheek completely scaled, about 9–10 horizontal scale rows below orbit, superficially embedded in skin. Preopercle naked; interopercle naked or with a patch of 1–3 scales posteriorly, embedded in skin. Scales in E1 row 59 (1), 60*(2), 61 (3), 62 (2), 63 (3), 64 (2), 65 (3), 66 (7), 67 (2), 68 (6) or 71 (1) in specimens from Rio Juruena, 58 (1), 60 (4), 61 (1), 62 (3), 63 (1) or 65 (1) in specimens from upper Rio Paraguai. Transverse scale row 14–15+1+5. Circumpeduncular scale rows 11–13 dorsally, 11–13 ventrally (24–28 including lateral lines). Lateral-line scales 20/13 (2), 20/14 (2), 20/15 (1), 21/11 (2), 21/12 (2), 21/13 (3), 21/14 (3), 22/11 (2), 22/12 (2), 22/13 (4), 23/11* (4), 23/12 (4), 25/11 (1) in specimens from Rio Juruena, 21/12 (1), 22/10 (1), 22/11 (2), 22/ 12(1), 23/13 (1), 24/10 (2), 24/11 (3) or in specimens from upper Rio Paraguai, and 2–3 scales continuing lower line onto caudal fin. Upper and lower branches of lateral line not overlapping. Scales between upper lateral line and first dorsal-fin spine 9–11; scales between upper lateral line and last dorsal-fin spine 4–5; horizontal scale rows between branches of lateral line 3–4. Lateral-line scales larger and more elongate than scales of adjacent rows; each scale of upper branch of lateral line impinging 2–3 scales of ventrally-adjacent row, each scale of lower branch of lateral line impinging 2 (rarely 3) scales of dorsally-adjacent row. Dorsal, anal, pectoral and pelvic fins without scales. Caudal-fin squamation extending to about one third of fin length, posterior margin of scaled area concave. First dorsal-fin spine about one-fourth the length of last; spines subequal in length from 11th spine. Posterior edge of soft part of dorsal fin pointed, longest ray (10th) slightly beyond base of caudal fin, extending to approximately one-half the length of caudal fin in some specimens. Dorsal-fin count XVIII.13 (1), XVIII.14 (3), XVIII.15 (2), XVIII.16 (3), XIX.13 (1), XIX.14* (9), XIX.15 (6), XX.13 (2) or XX.14 (5) in specimens from rio Juruena, XVIII.14 (1), XVIII.15 (1), XIX.14 (5), XIX.15 (1), XX.14 (3) in specimens from upper rio Paraguai. Posterior edge of soft anal fin pointed, 7th or 8th ray longest, reaching caudal fin base or extending to approximately one-fifth the length of caudal fin. Anal fin count III.9 (2), III.10*(11), III.11 (17), III.12 (2) in specimens from rio Juruena, III.9 (3), III.10 (8) in specimens from upper rio Paraguai. Caudal fin rounded with 8 principal and 6–7 procurrent rays on each lobe. Pectoral fin rounded, 7th or 8th ray longest, not reaching halfway from its base to first anal-fin spine. Pectoral-fin rays 15 (7), 16 (23), 17*(2) in specimens from rio Juruena, 15 (1), 16 (8), 17 (2) in specimens from upper rio Paraguai. Pelvic fin rounded and nearly symmetrical, inserted slightly posterior to pectoral axilla at vertical through dorsal-fin spine III; 2th pelvic-fin ray longest, reaching halfway from its base to spinous anal fin, very far from genital papillae. All oral teeth conical, recurved. Teeth arranged in one outer row and 2–3 inner rows in dentary and premaxilla, with at least one of inner rows extending posteriorly almost as long as outer row and other rows more or less restricted to symphyseal portion. Teeth in outer row movable, but larger and slightly more fixed than teeth in inner rows, depressible; outer-row teeth near symphysis of dentary and premaxilla larger than posterior ones. Outer row with 22–30 teeth on right, 25–28 on left premaxilla; 27–33 on right, 25–30 on left dentary (counted in three c&s specimens, 59.2–67.7 mm SL). First gill arch of specimens from rio Juruena with 1 (6) or 2*(24) gill rakers externally on epibranchial and 8*(4), 9 (18), 10 (8), 11 (1) on ceratobranchial. Specimens from rio Paraguai with 1 (1), 2 (10) gill rakers on first epibranchial and 8 (2), 9 (5), 10 (4) on first ceratobranchial. Occasionally, one of the ceratobranchial gill rakers can be situated at angle between epi- and ceratobranchial instead of on ceratobranchial. Gill rakers on lower pharyngeal tooth plate 6 (1), 8*(4), 9 (8), 10 (7), 11 (1), 12 (5) in specimens from Rio Juruena and 8 (3), 9 (2),10 (2), 11 (1), 12 (1) in specimens from upper Rio Paraguai. No tooth plates on any ceratobranchials. Single concavity in frayed zone of posterior margin pharyngobranchial 4. Microbranchiospines present externally on second, third and fourth gill arches.

370 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. Crenicichla ploegi Crenicichla + 0 7M+) +3 ) + 6 / . +M+* +3 ( + ) , " , " p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

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 371 FIGURE 6. Infraorbital series: a) Crenicichla ploegi, MZUSP 108380, 67.7 mm SL; b) Crenicichla macrophthalma, MZUSP 105713, 117.0 mm SL, c) Crenicichla saxatilis MZUSP 101130, 102.8 mm SL, d) Crenicichla missioneira, MCP 18926, 129.9 mm SL; e) Crenicichla jupiaensis, MZUSP 16048, 62.0 mm SL; f) Teleocichla preta, NRM 65527, 63.3 mm SL; g) Crenicichla chicha, MZUSP 93683, 70.2 mm SL.

372 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. FIGURE 7. Crenicichla aff. lepidota Tapajós a) LBP 8020, adult male, 92.4 mm SL, tributary of Rio Arinos in Nova Mutum, Rio Juruena basin; b) LBP 7952, adult female, 75.1 mm SL, Rio dos Patos, Rio Juruena basin; c) LBP 8020, subadult, 41.3 mm SL, tributary of Rio Arinos in Nova Mutum, Rio Juruena basin; d) LBP 8008, juvenile, 21.6 mm SL, tributary of Rio Arinos in Nova Mutum, Rio Juruena basin.

Lower pharyngeal tooth-plate (dissected from three specimens: MZUSP 108380, 67.7 mm SL; ZUEC 10181, 142.0 mm SL; and ZUEC 10053, 146.1 mm SL, Fig. 8) strongly depressed in lateral view, with long posterior and anterior processes; tooth-plate length 76.7–82.7% of plate width; dentigerous-area length 42.0–63.5% of plate width, dentigerous-area width 72.0% plate width; 18 teeth in posterior row, 7–8 teeth in admedian rows. Medioposterior teeth largest and more robust, teeth gradually decreasing in height and becoming more slender rostrally and laterally. Teeth of posterior and admedian rows and on central dentigerous area bicuspid and compressed laterally, with main cusp antrorse; lateral and rostral teeth unicuspid, nearly conical.

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 373 FIGURE 8. Crenicichla ploegi, lower pharyngeal tooth plate in dorsal view, from ZUEC 10053, 146.1 mm SL. Posteromedial tooth, extracted during preparation of material, shown in lateral view.

Vertebrae (abdominal + caudal= total vertebrae): 18+13= 31 (1), 18+14=32 (1), 19+13=32 (1), 19+16=35 (2), 18+18=36 (1); with 6 vertebrae contained in caudal peduncle. Coloration in alcohol. Juveniles up to approximately 28 mm SL (Fig. 1d) with ground color yellowish pale, slightly darker and grayish dorsally than ventrally; dark markings brownish or blackish. Lateral-line scales uniformly pigmented with same color of adjacent ones. Preorbital stripe dark brown, from tip of lower jaw, continuing dorsally on upper lip, then running caudad below level of nostrils to anterior margin of orbit. Postorbital stripe aligned with preorbital one, from posterior margin of orbit to opercular angle. Dark brown, narrow stripe dorsally parallel to postorbital stripe, running from dorsoposterior margin of orbit to vertical through posterior preopercular margin; another dark, rounded blotch in extrascapular area. In dorsal view, dark brown, single median stripe runs from vertical through nostrils to vertical through middle of orbit on interorbital area, duplicating afterwards in two paired, irregular stripes running from area corresponding to frontal-supraoccipital crests posteriorly along dorsum close to base of dorsal fin. Paired stripes slightly fragmented into about five portions, first one close to origin of dorsal fin and last at end of fin. Dark blackish midlateral band very evident from just behind opercle to end of caudal fin, occupying depth of 4–5 horizontal scale rows up to caudal-fin flexure, becoming narrower and fainter from it to end of caudal fin. Humeral blotch rounded, appearing to be a darker portion of midlateral blotch ventrally but distinct from midlateral band dorsally, situated entirely below upper branch of lateral line. Pectoral and pelvic fins hyaline. Dorsal fin dusky, darker on distal half of spinous portion and on submarginal and marginal areas of its soft portion. Anal fin dusky, with darker margin. Caudal fin with blackish

374 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. rounded blotch tangent to fin flexure and between rays D3–V2, and some dark grayish pigmentation on its submarginal area of dorsal and ventral distal corners. Most of color pattern elements (dark markings) of subadults from 34 to around 65 mm SL (Fig. 2c, d) correspond to those of juveniles, but with some differences as highlighted below. Ground color on dorsal half of body is distinctively darker and more brownish, mainly due to posteroventral extension of dorsal blotches (from juveniles) forming 5–7 dark bars on dorsum; first bar oblique, running posteroventrally from dorsal fin origin to humeral area; last bar at end of dorsal fin, often extending more posteriorly onto caudal peduncle. Anterior to first vertical bar, viewed from above, a median dark stripe runs from snout to vertical through middle of orbit on interorbital area, duplicating afterwards into two irregularly-edged stripes caudally on posttemporal area. Bars on dorsum separated from midlateral band by a paler region following dorsally the midlateral band in many specimens and, in some of them, these bars also divided in two portions by paler pigmentation on scales of upper branch of lateral line. Specimens from 34 mm SL already showing some melanophores concentrated below eye, but dark marking which may be interpreted as a triangular suborbital blotch only distinguishable in specimens larger than 45 mm SL. A rounded conspicuous humeral blotch, its dorsal part distinct from midlateral band and surrounded by light, whitish pigmentation; in some specimens, light pigmentation extends posteriorly as series of whitish dots accompanying dorsally the midlateral band. Caudal blotch surrounded by lighter pigmentation, often with some whitish spots around it; posteriorly to caudal blotch, at least three vertical pale stripes contrast with the darker caudal-fin background and a very dark, blackish blotch at the end of fin. Dorsal fin more widely smoky and darker than in juveniles, with one or two rows of lighter spots on its spinous portion and at least three rows of lighter spots on its soft portion. Anal fin dusky as dorsal fin, with darker margin and three series of lighter dots on soft portion. Female with 69.4 mm SL (Fig. 1c) showing signals of sexual maturity and considered smallest adult specimen observed. Thus, specimens over 69 mm SL interpreted as possessing adult color pattern (Fig. 1a–c; Fig. 2a, b). Dark suborbital marking more evident and extended ventrally than in subadults, triangular shape distinct. With increasing of size, dark markings dorsally on snout and nape become more fragmented, forming coarse pattern of dark spots and vermiculations; this pattern observed in many male and female specimens, including specimens up to 146.6 mm SL. Bars on dorsum extending more ventrally, reaching midlateral area, often connecting with midlateral band. Rounded humeral blotch larger and very conspicuous, distinct from midlateral band, both dorsally and ventrally, by white ring surrounding entire blotch; black blotch on pectoral axilla present. Midlateral band still conspicuous and deep, occupying depth of five to six horizontal scale rows. Females showing more homogeneous color pattern, with dark markings (except humeral blotch) fainter than in males; in same way, dorsal, anal and caudal fins of females homogeneous dark pigmented, only with a darker blackish margin and lacking the lighter spots present in most subadults (and adult males); caudal-fin blotch in mature females lacking light ring. Males exhibiting more pronounced midlateral band and vertical bars. Besides a conspicuous white ring bordering humeral blotch (as in females), mature males showing numerous white dots scattered along midlateral area, on dorsum, caudal peduncle, eventually also onto basal portion of caudal fin and around caudal-fin blotch. Dorsal, anal and caudal fins smoky, without darker margin; numerous small, light (hyaline) spots scattered on spinous and soft portion of dorsal fin, on soft portion of anal fin and on caudal fin. Some specimens, mainly larger ones, with dark markings (head stripes and spots, midlateral band) much faded, except the humeral blotch. This variation may be due to preservation conditions or induced by stress during capture and/or subsequent fixation. Life colors. Description based on pictures of fishes in nature, as Fig. 4 taken in the Rio Verde, a tributary of the Rio Juruena, and on pictures of live fishes photographed in aquarium just after capture, as Fig. 5, taken of individual from the Rio Juba, upper Rio Paraguai basin. Ground color greenish-yellow dorsally and on dorsal, anal and caudal fins; ventral body pale to silvery; pectoral fin hyaline to yellowish. Melanin-based color pattern elements (e.g. head stripes and vermiculations, humeral and caudal blotches, and midlateral band) grayish to blackish, as described for preserved specimens. Iris red and two dark blotches laterally, forming what may be interpreted as a continuation from pre- to postorbital stripes. Suborbital triangular blotch brownish to yellowish. Iridescent regions around suborbital blotch and ventrally on postorbital area. White ring surrounding humeral blotch both in males and females. In males, same kind of white pigmentation forming dots scattered along midlateral area, on dorsum, and caudal peduncle; greenish and iridescent dots on caudal, dorsal and anal fins; dorsal-fin margin and margin of dorsal lobe of caudal fin red. In females, dorsal, anal and caudal fins uniform, dots absent; dorsal fin and dorsal margin of caudal fin with red submarginal stripe and blue margin.

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 375 Morphometric comparisons. The proportion body depth/standard length of Crenicichla ploegi is of 17.8– 21.7% SL, mean 19.7%. For the morphometric comparisons, the body depth was analyzed in function of the SL, firstly with data of the material encompassing juveniles and adults of all valid species of the Crenicichla saxatilis group. Graphically, there are two clouds of scattered points that corresponds to slender (Fig. 9 green and blue) and deep-bodied species (Fig. 9: red). ANCOVA analysis showed non-significant differences between the body depth of these two putative subsets (p-value 0.592). The hypothesis of homogeneity of slopes was rejected, indicating that body depth increases at different rates in the two subsets, with individuals becoming more different as standard length increases. To control for allometry, a second analysis was performed with adult specimens only (Fig. 9). Adults were defined as the specimens > 69 mm SL, as this is the standard length of the smallest specimen of Crenicichla ploegi with signals of sexual maturity. The second analysis showed significant difference between the body depth of the two subsets (p-value 0.039). The hypothesis of homogeneity of slopes was reject again i.e. body depth increases at different rates in the two subsets.

FIGURE 9. Bi-plots of body depth vs. SL with linear regressions of the 23 species of the Crenicichla saxatilis group, including C. ploegi, considering only adult specimens (> 69 cm SL). Species considered as deep-bodied taxa are shown in red; slender species are represented in green and blue.

Distribution (Fig. 11). Crenicichla ploegi is known both from the upper Rio Paraguai basin (upper reaches of Jauru, Cabaçal, and Sepotuba rivers), and from tributaries of the upper Rio Juruena (Papagaio, Sangue, Arinos rivers, and Rio Juruena itself), in Mato Grosso state, Brazil. Etymology. Crenicichla ploegi is named in honor of the Dutch ichthyologist Alex Ploeg, whose PhD thesis dealt with the taxonomic revision, biogeography and phylogeny of Crenicichla, and who published papers on systematics of the genus from 1986 to 1991, describing a total of 23 species, 18 of which still considered to be valid. Since then, he worked as interlocutor between the ornamental fish industry and other institutions worldwide.

376 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. Unfortunately, Dr. Ploeg perished with his wife and son in the Malaysia Airlines 17 (MH17) airplane attack in Ukraine on 17 July 2014. Habitat and ecological notes. Crenicichla ploegi has been collected in a variety of aquatic habitats, but mostly in streams and mid-sized rivers. The type locality is actually an exception, being a large river pool, more than 100 meters wide. Streams and rivers inhabited by C. ploegi possess clear water, a moderate to fast water flow, and rocky bottoms, usually with well-developed riparian forest, although the species has also been collected in rivers with highly disturbed or suppressed riparian forest.

FIGURE 10. Schematic drawings illustrating the correlation between general body shape and the position of the humeral blotch in relation to the anterior branch of lateral line: a) Crenicichla lepidota, lectotype NMW 33101, 157.8 mm SL—deep- bodied species with humeral blotch below anterior branch of lateral line; b) Crenicichla ploegi, holotype MZUSP 93697, 89.8 mm SL—slender species with humeral blotch below anterior branch of lateral line; and c) Crenicichla alta, SU 78954, 115.4 mm SL—slender species with humeral blotch displaced dorsally, with its center on the anterior branch of lateral line.

Discussion

Since Ploeg´s (1991) definition of the Crenicichla saxatilis group, no work has attempted to describe new species of this group. Many putative new species have been noticed, but the recognition of species described long ago is difficult, mainly because of inaccessible or unavailable type material or the condition of this material (e.g. Crenicichla brasiliensis, C. lepidota and C. saxatilis); this prevents meaningful comparisons with recently collected specimens. Moreover, in a sequence of three papers, Ploeg (1986, 1987, 1991) described fifteen nominal

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 377 species belonging to the C. saxatilis group, five of which have been subsequently considered junior synonyms of other species (Kullander, 2003; Table 1). Clearly, species of the C. saxatilis group still need a more in-depth taxonomic appraisal. To describe and compare Crenicichla ploegi, much of the type material and additional samples of almost all species of the C. saxatilis group were examined. As a critical reappraisal of the diagnostic characteristics of the group is lacking, we update the diagnosis of the group from a discussion of the morphological characters previously proposed, and provide comparisons among the included species as well as some biogeographical comments.

FIGURE 11. Distribution map of Crenicichla ploegi. Type locality as black star, collecting sites of paratypes as white circles.

Humeral blotch. The presence of a blotch on the humeral region of the body is indicated in the diagnoses proposed for this group by Kullander (1981, 1982), Ploeg (1991) and Kullander et al. (2009). A blotch in the humeral region is always present in all species of the Crenicichla saxatilis group sensu Ploeg (1991) except C. guentheri. The humeral blotch is present even in the smallest specimens, and the same kind of blotch is present in all specimens of C. ploegi. Crenicichla guentheri, recently considered a junior synonym of C. hemera by Varella et al. (2012), shows a similar but non-homologous dark blotch on the area dorsal to the base of pectoral fin. It was demonstrated that this blotch appears only late during ontogeny and that it is not present in all large specimens. Crenicichla hemera shares this kind of dark blotch with many species of the C. lugubris group (see Kulander & Varella, 2015). For this and other reasons, Varella et al. (2012) excluded C. hemera from the C. saxatilis group and suggested that it constitutes a monophyletic group with C. chicha. Thus, while the presence of a dark pigmentation patch dorsally to the pectoral fin is not exclusive for the C. saxatilis group, its presence in early ontogenetic stages yes. The position and shape of the humeral blotch vary widely and is useful to the interespecific comparisons in the group. Scales in E1 series. Previous authors characterized the Crenicichla saxatilis group as having a low number of scales in the longitudinal series. Ploeg (1991) followed Regan (1905, 1913) in counting the scales in the

378 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. longitudinal series below the lateral lines. Herein, we follow Kullander (1986), as do most recent taxonomic papers (e.g. Kullander & Nijssen, 1989; Lucena & Kullander, 1992; Kullander & Lucena, 2006; Kullander et al., 2009), and count the longitudinal scales along the E1 series. Species in the C. saxatilis group vary widely in E1 counts (31–71), with the lowest counts in C. britskii (31–43) and C. isbrueckeri (38–45) and the highest (up to 71 scales) in C. ploegi. These counts overlap with those of most species in other groups in Crenicichla and Teleocichla, being diagnostic only from to the C. lugubris and C. acutirostris groups sensu Kullander & Varella (2015), which have more than 89 and more than 74 scales, respectively. Color features. The presence of a continuous postorbital stripe instead of a postorbital blotch and the occasional presence (instead of absence) of a suborbital marking were used in Ploeg´s (1991) key to the species groups to distinguish species of the C. saxatilis, C. lacustris and C. lugubris groups from those of the C. reticulata and C. wallacii groups. As with many other Neotropical cichlids, all species of Crenicichla and Teleocichla possess a postorbital marking and there is ontogenetic variation of its shape. A postorbital stripe from the posterior margin of orbit to the posterior edge of the opercle is present in juveniles and subadults of most species of Crenicichla belonging to all groups recognized by Ploeg (1991). Most species of Crenicichla retain a continuous stripe in adults, including all species of the C. saxatilis group. Adults of other cichlids have this marking divided into a conspicuous, rounded blotch just behind the orbit and another elongate and fainter blotch in the opercle. The latter condition is less usual in Crenicichla, but is found in species of different groups, and may be correlated with the head relatively short and deep. Species of Teleocichla and of the C. wallacii and C. reticulata groups lack pigmentation on the cheek altogether. In the remaining species, the suborbital markings may have many distinct configurations, but at least three major patterns are observed:

i) a conspicuous, uniformly-pigmented dark blotch, which is short and triangular or elongate (stripe-like)— species of the C. saxatilis group, C. hemera and C. chicha; ii) an uniformly-pigmented suborbital marking, but much fainter and with a semicircular shape—Crenicichla cincta Regan, 1905, C. johanna Heckel, 1840, C. lugubris Heckel, 1840, C. rosemariae Kullander, 1997, C. strigata Günther, 1862 and C. tigrina Ploeg, Jégu & Ferreira, 1991, all belonging to the C. lugubris group sensu Kullander (1997); iii) the “spotted kind” (Kullander, 1982: fig. 12), the shape of the suborbital marking as a whole depends on the configuration of the small dark dots—species of the C. lacustris group sensu Piálek et al. (2012).

Sexual dimorphism. Kullander (1982) highlighted a strong sexual dimorphism of color pattern in species related to C. saxatilis. Mature females have the body almost uniformly dark, lacking most of the dark color pattern elements except the humeral and caudal blotches, preorbital, postorbital and suborbital markings. In most species of the C. saxatilis group, the mature females present the dorsal and anal fins with a dark margin and, occasionally, with a series of small dark spots (black in preserved, red in live specimens) along the submarginal area. Besides, the rounded belly is colored with reddish pigmentation in live mature females. Males retain the juvenile color pattern but large specimens usually become darker and with light—white or iridescent yellow—dots scattered on flanks. In Crenicichla albopunctata, the light dots occasionally extend anteriorly on lateral portion of the head. The females of Crenicichla ploegi examined herein present the dark-margined dorsal and anal fins but no series of small dark blotches on submarginal area, and the males show numerous white dots scattered on flanks. Sexual dimorphism in coloration is not exclusive of the Crenicichla saxatilis group. Reddish pigmentation on the belly is also found in females of some species of the C. wallacii and C. lugubris groups. However, the females of the C. lacustris, C. reticulata and C.wallacii groups show large, ocellated blotch(es) or a band on the dorsal fin but no dark margin and small dark spots on the submarginal area as in the C. saxatilis group. Besides, females of the C. lacustris and C. reticulata groups usually show an orange pigmentation on the lateral portion of the belly instead of a reddish pigmentation on its ventral portion. Features related to the sexually dimorphic coloration of males are more complex and constitute many different patterns, some of them being species-specific. Among the species of Crenicichla and Teleocichla, numerous light dots scattered on flanks similar to those of the species of the C. saxatilis group are only found in males of C. multispinosa, a member of the C. acutirostris group sensu Kullander (1997).

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 379 Extension of maxilla. Regan (1905, 1913) separated some species that are currently allocated in the C. saxatilis and C. reticulata groups by having the maxillary extending beyond the anterior margin of orbit. Kullander (1982) indicated a similar condition as one of the diagnostic characteristics of the C. lepidota species group. According to our data, species of the C. saxatilis group show the posterior border of the maxillary extending beyond the vertical through the anterior margin of orbit, as well as three species of the C. scottii complex within C. lacustris group sensu Piálek et al., (2012) and most species of the C. reticulata group. In the majority of species of Crenicichla, the posterior maxillary reaches the vertical through the anterior margin of orbit instead. A third condition, maxillary not reaching the anterior margin of orbit, is uncommon in Crenicichla (observed only in C. acutirostris, C. cyclostoma Ploeg, 1986, C. compressiceps Ploeg, 1986 and C. urosema Kullander, 1990a), but is showed by all species of Teleocichla (except T. gephyrogramma). Infraorbitals series. Cichoki (1976: character 46) and Kullander (1998: character 40) discussed the shape of lacrimal bone for South American Cichlidae. This character was reexamined in species of Crenicichla and Teleocichla, and in a number of other Neotropical cichlids. The lacrimal bone is very slender (Fig. 6a, c) in all species of the Crenicichla saxatilis group. This is an unusual characteristic among species of Crenicichla, being shared with those species of the C. reticulata group that have maxilla extending more posteriorly, and may correspond to Kullander´s state 2. Among other neotropical cichlids examined herein, this condition is found in some dwarf geophagines, viz. Apistogramma taeniata (Günther), Taeniacara candidi Myers, Biotoecus opercularis (Steindachner) and Dicrossus filamentosus (Ladiges), and in Acaronia nassa (Heckel). The most common condition in Crenicichla and Teleocichla is the lacrimal slightly longer than deeper (Fig. 6b, e, f, g), and in species of the C. missioneira group sensu Lucena & Kullander (1992) and C. tapii Piálek et al., 2015 the lacrimal is approximately square, as long as deep (Fig. 6d; Kullander, 1986: fig. 13 a, d)—these two conditions may correspond to a division of Kullander´s state 0. Varella et al. (2012) used the co-ossification of the second and third post-lacrimal infraorbitals (numbered 4 and 5; Fig. 6f, g) as one of the arguments to exclude Crenicichla hemera and C. chicha from the C. saxatilis group and to consider them as putative sister-taxa. In fact, all species of the C. saxatilis group (as considered herein) present separate five post-lacrimal infraorbital tubes (infraorbitals 3 to 7; Fig. 6a–d). This condition is shared with almost all congeners and with Cichla, Retroculus Eigenmann & Bray and Pterophyllum Heckel among the South American cichlids examined (Kullander, 1998; our data). The co-ossification and reduction in number of infraorbital bones are common among neotropical cichlids, being more intensified in geophagine taxa (Kullander, 1998; López-Fernández et al., 2005). Besides Crenicichla hemera and C. chicha, all species of Teleocichla have the apposed openings of the infraorbitals 4 and 5 co-ossified, leaving a middle pore (Fig. 6f, g). The unique examined specimen of Crenicichla igara Lucena & Kullander, 1992 examined also presents this condition. Crenicichla jupiaensis (Fig. 6e) show an unique condition among Crenicichla-Teleocichla: only four instead of five infraorbitals but no signal of co-ossification, as a middle pore, similar to the configuration found in Cleithracara (Kullander & Nijssen, 1989: fig. 103). Updated diagnosis. Species of the Crenicichla saxatilis group are diagnosed from all other species of Crenicichla and Teleocichla by the presence of a dark humeral blotch even in early ontogenetic stages and a dimorphic color pattern that consists in mature males with scattered light dots on the flanks and mature females with dark-margined dorsal and anal fins (occasionally also a series of small dark blotches on the submarginal area of the dorsal fin). They are also distinguished from all Crenicichla and Teleocichla except some species of the C. reticulata group sensu Ploeg (1991) and C. scottii complex sensu Lucena & Kullander (1992) by the posterior border of the maxilla extending beyond the vertical through the anterior margin of the orbit, and from all Crenicichla and Teleocichla except some few species of the C. reticulata group by the lacrimal bone very slender. In addition, species of the C. saxatilis group differ from those of the C. lugubris and C. acutirostris groups sensu Kullander & Varella (2015) by having 31–71 scales in the E1 series vs. more than 74 scales. From species of the C. reticulata and C. wallacii groups and of Teleocichla, they differ by the presence (vs. absence) of a suborbital marking and from species of the C. lacustris group sensu Piálek et al. (2012) by having a uniformly-pigmented suborbital marking vs. a composite suborbital marking formed by small dark dots. Finally, from the group formed by C. hemera and C. chicha (sensu Varella et al., 2012) and from Teleocichla, the species of the C. saxatilis group are distinguished by having five separate post-lacrimal infraorbital tubes vs. post-lacrimal infraorbitals 4 and 5 co- ossified, leaving a middle pore. Comparisons between species of the C. saxatilis group. Deep-bodied and slender taxa. The major shape

380 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. difference related to deep-bodied and slender taxa of the C. saxatilis group is not significant when all specimens (including juveniles) are analyzed, but significant if only adult specimens are included (see Morphometric comparisons section). Juveniles of the two subsets present functions of body depth vs. standard length more similar than those of the adults. Probably, the positive allometry in this proportion, common in cichlids, would be higher in the deep-bodied species than in the slender species. With a body-depth range of 17.8–21.7% SL (mean 19.7%), Crenicichla ploegi figures among the slender taxa of the Crenicichla saxatilis group (body depth 17.1–24.5% SL, Table 3), but with a common body depth compared with most other species of Crenicichla and Teleocichla. Color pattern and E1 counts. Deep-bodied taxa viz. Crenicichla brasiliensis, C. britski, C. frenata, C. isbrueckeri, C. labrina, C. lepidota, C. proteus, C. santosi, C. saxatilis and C. semicincta (body depth 22.1–32.1 % SL, means between 22.7–27.9 % SL, Table 3, Fig. 9) have comparatively few scales on E1 series (31–56 scales). All these species show the humeral blotch mostly or entirely below the upper branch of lateral line (Fig. 10a). The slender taxa (body depth 17.1–24.5 % SL, means 19.2–21.7 % SL) have comparatively more scales in the E1 series (54–71 scales). Some of these species (Crenicichla ploegi, C. hummelincki, C. inpa, C. lucius, C. nickeriensis and C. pellegrini) have the humeral blotch below anterior lateral line (Fig. 10b). In the others (C. albopunctata, C. alta, C. anthurus, C. coppenamensis, C. pydanielae and C. sipaliwini), the humeral blotch is displaced dorsally and with center on the anterior lateral line (Fig. 10c). Ploeg (1991: fig. 172) recovered a clade including all the seven species listed above but C. albopunctata, with the circular humeral blotch placed dorsally as synapomorphy. Ploeg (1991) also included C. lucius in this assemblage, but the neotype of Crenicichla lucius reexamined herein has a humeral blotch entirely below the anterior lateral line. Species of the C. saxatilis group have a conspicuous and uniformly-pigmented suborbital marking, but with a wide variation in relation to its shape. All deep-bodied taxa of the C. saxatilis group have the suborbital marking narrow and elongate, stripe-like (Kullander, 1986: fig.29A) except for an undescribed species from Northeast Brazil (Kullander, 1982) in which suborbital marking is lacking. Slender taxa possess suborbital marking shorter and broader, usually like a triangle (Kullander, 1986: 29B).

TABLE 3. Comparisons body depth (as percentage of SL) of the species of the Crenicichla saxatilis group (n, number of specimens measured; SD, standard deviation). n range mean SD n range mean SD C. ploegi 43 17.8–21.7 19.7 1.0 C. britskii 48 24.8–32.1 27.9 2.0 C. albopunctata 1 20.8 - - C. frenata 124.6 - - C. alta 30 18.3–23.8 20.7 1.1 C. labrina 8 22.3–25.2 23.3 1.2 C. anthurus 21 17.1–22.4 19.2 1.4 C. isbrueckeri 6 22.0–25.8 23.4 1.7 C. coppenamensis 14 17.9–22.9 20.1 1.5 C. lepidota 75 23.2–30.4 25.6 1.6 C. hummelincki 10 20.1–22.2 21.4 0.7 C. brasiliensis 11 21.3–27.1 24.0 1.8 C. inpa 17 18.8–23.4 21.0 1.6 C. proteus 9 22.2–25.4 24.3 0.9 C. lucius 8 21.0–22.4 21.7 0.4 C. santosi 10 21.4–25.4 23.5 1.5 C. nickeriensis 14 18.9–24.5 21.2 1.5 C. saxatilis 13 22.2–26.4 24.3 1.5 C. pellegrini 6 20.2–21.5 20.7 0.6 C. semicincta 7 22.1–23.6 22.7 0.6 C. pydanielae 5 20.5–21.0 20.7 0.2 C. sipaliwini 9 17.8–21.5 19.6 1.5 Crenicichla aff. lepidota 23 21.4–27.5 24.5 1.5 C. sveni 9 20.3–22.5 21.7 0.8 (Rio Tapajós basin)

Biogeographical comments. Species of Crenicichla and Teleocichla are distributed in almost all major river basins of the cis-Andean South America from the lower Río de La Plata in Argentina to Trinidad and coastal drainages of northern Venezuela. Members of the C. saxatilis group occur across the whole distribution of the genus Crenicichla, with exception of the coastal river drainages of south and southeastern Brazil from southern Bahia State to northern Rio Grande do Sul State (Kullander & Lucena, 2006). The distribution of the deep bodied-taxa overlaps that of the whole C. saxatilis group entirely, with highest diversity in the Amazon, Orinoco and in drainages of the Guyana Shield. Two species (Crenicichla lepidota and C. britskii) are distributed in the Río de La Plata basin and Lagoa dos Patos system (Kullander, 1982; Lucena &

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 381 Kullander, 1992; Varella, 2011); Crenicichla brasiliensis and C. menezesi are the only two nominal species from coastal rivers of Northeast Brazil (Kullander (1982; Ploeg, 1991) but they require further taxonomic investigation. Herein, C. brasiliensis is provisionally considered the only valid species of Crenicichla distributed in Northeast Brazil, with C. menezesi as a subjective junior synonym. The second northeastern Crenicichla is a still undescribed species from the Rio São Francisco basin (Kullander, 1982). The slender species of the C. saxatilis group are restricted to the Amazon basin, Rio Orinoco basin, coastal drainages of the Guyanas and Suriname and other smaller coastal drainages close to the mouth of the Rio Amazonas. Crenicichla ploegi is the only slender species distributed both in the Amazon basin (Rio Juruena basin) and in the Río de La Plata basin (upper Rio Paraguai basin). Among the slender species of the C. saxatilis group, those with humeral blotch displaced dorsally have a more restricted distribution. It encompasses drainages of the Western Amazon in Ecuador, Peru and Colombia, tributaries of the left margin of the Amazon basin (e.g. Rio Branco, Rio Trombetas and Rio Jari), and coastal drainages in the Guyanas and Suriname. They were not recorded for the major tributaries of the right margin of the Amazon river. Crenicichla ploegi occurs in the upper portion of the Rio Paraguai, where it is sympatric with Crenicichla lepidota—the only species of the C. saxatilis group distributed in the Rio Paraguai basin. In the Rio Juruena basin, C. ploegi is sympatric with a species preliminary identified herein as Crenicichla aff. lepidota Tapajós (Fig. 7). This upper Rio Paraguai-Rio Juruena distributional pattern was previously discussed by Lima et al. (2007) and Ribeiro et al. (2013). Lima et al. (2007) listed seven species of fish shared between these drainages, including C. ploegi as “an … apparently undescribed… species of Crenicichla”. The Rio Juruena and upper Rio Paraguai drainages possess a broad water divide. Thus, those authors inferred that the occurrence of shared taxa between these drainages points to hydrographic rearrangements caused by capture of headwaters (upper reaches of Jauru, Cabaçal, and Sepotuba rivers, in the case of C. ploegi) which would have formerly drained into the Amazon basin and that resulted in a northward extension of the upper Rio Paraguai basin (Lima & Ribeiro, 2011; Ribeiro et al., 2013). From field notes on the preserved material of the Museu de Zoologia da USP (MZUSP), Crenicichla ploegi is usually found in in streams and mid-sized rivers of the rio Juruena and the upper rio Paraguai basin, with clear water, moderate to fast water flow and rocky substrate. Crenicichla lepidota and C. aff. lepidota, which are sympatric with C. ploegi in the rio Juruena and upper rio Paraguai respectively, usually inhabits backwaters with slower water flow and substrate composed mainly of sand and clay. This kind of spatial partitioning occurs in other right-bank tributaries of the Amazon river, in which a pair of species of the C. saxatilis group are sympatric: one deep-bodied species occupying lentic environments and another slender species inhabiting the lotic (e.g. Crenicichla pellegrini and C. isbrueckeri in the Rio Aripuanã basin; C. lucius and C. proteus in Western Amazon river tributaries; Crenicichla cf. inpa and C. labrina in the Rio Tocantins). Putative paedomorphic characters. The smallest specimens (up to 30 mm SL) of all species of the C. saxatilis group show the same color pattern elements as described for juveniles of C. ploegi (Fig. 3a, d). Preorbital and postorbital stripes are conspicuous and almost continuous with a midlateral band running along the flank onto caudal fin. An humeral blotch can be distinguished on the anteriormost portion of the lateral band. Dark markings of the dorsal portion of head and body are slightly less conspicuous than lateral markings, and basically composed by a unique blotch dorsally on the snout, paired posttemporal markings, and paired stripes from the dorsal portion of the head, continuing backward on dorsum onto caudal peduncle. With increasing of size, in both males and females of species belonging the Crenicichla saxatilis group, those dorsal dark stripes on the head of juveniles become less conspicuous. In adult specimens over 80 mm SL, the dark markings are completely vanished and the dorsal portion of head is uniformly dark brown, with exception of the posttemporal that may be distinguishable or not (Fig. 3e). The dorsal stripes on dorsum are replaced by faint vertical bars. The lateral band of the juveniles is faded or modified to a series of irregular or rounded blotches (e.g. in males of C. brasiliensis, C. britskii, C. semicincta and C. nickeriensis). On the other hand, the humeral blotch remains conspicuous and, consequently, more prominent on the flank. From the signals of sexual maturity in coloration and examination of gonads of some specimens, it seems that Crenicichla ploegi reaches sexual maturity at similar length of other species of the C. saxatilis group i.e. around 65–80 mm SL. However, ontogenetic change of coloration of C. ploegi is different in some aspects. In subadults and smaller adults (ca. 69–80 mm SL), posttemporal markings, dorsal markings of head and dorsum are still conspicuous but become divided in smaller blotches or stripes (Fig. 3b). In most adults of C. ploegi up to 146.6 mm SL, these markings remain distinguishable, more fragmented, forming a coarse pattern of dark spots and

382 · Zootaxa 4377 (3) © 2018 Magnolia Press VARELLA ET AL. vermiculations on the dorsal portion of the snout and head (Fig. 3b, c). The conspicuousness of the midlateral band of subadults and of almost all adults (even larger specimens between 164.4–190.4 mm SL) remains similar to that of the juveniles (Fig. 3b, c). Therefore, considering that C. ploegi reaches sexual maturity at the same length of the other species of the C. saxatilis group, the presence of conspicuous dark markings dorsally on the head and the presence of a prominent midlateral band are here hypothesized to be paedomorphic characteristics, retained from juvenile conditions in the context of the species of the C. saxatilis group. Our results is in accordance with Lucena and Kullader (1992), who considered the retention of some color pattern elements (e.g. dark stripes and markings on head and the dark midlateral band on flanks) and absence of sexual dimorphism as indicative of juvenile coloration in species of the C. saxatilis group and in most small-scaled Crenicichla. Some of the sexually dimorphic features common to the species of C. saxatilis group are present in C. ploegi, as the numerous light dots on flanks in males and a dark margin of the dorsal fin in females. However, the females lacking the rounded belly with reddish pigmentation and a series of dark blotches in the submarginal area of the dorsal fin and the males not showing prolongation of dorsal and anal fins and with the light dots on flanks evidently smaller than in other species are suggestive of a weak sexual dimorphism.

Comparative material

Crenicichla albopunctata. MNHN 4898–59, lectotype, 144.9 mm SL, French Guiana. Crenicichla alta. Guyana, Essequibo River basin: FMNH 53777, holotype, 132.4 mm SL. BMNH 1911.10.31.11, 1, paratype, 60.8 mm SL. BMNH 1911.10.31.12, 1, paratype, 93.3 mm SL. BMNH 1911.10.31.13–14. 2, paratypes, 95.1–95.9 mm SL. CAS 66958, paratype, 1, 116.2 mm SL. CAS 78953, 2, paratypes, 68.1–71.1 mm SL. CAS 78954, 3, paratypes, 64.7–115.3 mm SL. CAS 78955, 1, paratype, 63.6 mm SL. CAS 78956, 4, 31.3– 84.1 mm SL. CAS 78958, 2, paratypes, 87.9–101.6 mm SL. CAS 78960, 4, paratypes, 63.9–119.4 mm SL. CAS 78961, 1, paratype, 98.6 mm SL. CAS 121780, 1, paratype, 68.1 mm SL. ANSP 39349, holotype of Crenicichla pterogramma, 125.1 mm SL. Brazil, Roraima, Rio Branco basin: INPA 2916, holotype of Crenicichla cardiostigma, 108.3 mm SL. Crenicichla anthurus. Peru, Departamento Loreto: CAS 36718, 9, 82.5–185.8 mm SL. MZUSP 26469, 2, 51.2– 55.1 mm SL. MZUSP 26100, 6, 57.5–92.0 mm SL. Crenicichla cf. anthurus. Colombia: CAS (SU) 53807, 10, 33.4–130.0 mm SL, Rio Orinoco-Guaviare. Crenicichla britskii. Brazil, São Paulo, upper Rio Paraná basin: MZUSP 16004, holotype, 1, 83.2 mm SL. MZUSP 16007–16021, 15, paratypes, 22.8–113.4 mm SL. MZUSP 41157, 2, 84.7–95.8 mm SL. MZUSP 41160, 2, 85.6–69.5 mm SL. MZUSP 41162, 1, 125.5 mm SL. MZUSP 41163, 1, 108.4 mm SL. MZUSP 41167, 1, 75.0 mm SL. MZUSP 83389, 1, 133.0 mm SL. MZUSP 83404, 4, 81.5–101.7 mm SL. MZUSP 88572, 1, 113.7 mm SL. MZUSP 102857, 2, 89.9–113.7 mm SL. MZUSP 100731, 5, 85.1–152.0 mm SL. MZUSP 100917, 1, 52.5–57.9 mm SL. Brazil, Paraná, upper Rio Paraná basin: MZUSP 16022, 1, 101.6 mm SL. MZUSP 42644, 7, 39.9–80.8 mm SL. NUP 2355, 1, 127.5 mm SL. NUP 6456, 3, 78.3–97.0 mm SL. NUP 6566, 2, 121.3– 126.1 mm SL. NUP 7880, 5, 46.9–120.5 mm SL. NUP 7895, 3, 46.6–96.9 mm SL. Crenicichla coppenamensis. Suriname, Saramaca district, Coppename River basin: ZMA 107841, holotype, 176.7 mm SL. IRSNB 764, 3, paratypes, 70.7–107.1 mm SL. ZMA 106512, 33, paratypes, 25.9–167.1 mm SL. ZMA 106513, 12, paratypes, 53.0–162.7 mm SL. Crenicichla frenata. CAS 77946, 2, ca. 150 mm SL (damaged) and 180.6 mm SL, West Indies, Trinidad (Trinidad and Tobago). Crenicichla hummelincki. Brazil, Pará, Rio Trombetas basin: MZUSP 39017, 10, paratypes, 30.1–91.3 mm SL; MZUSP 40315, 1, holotype, 104.5 mm SL. Crenicichla inpa. Brazil, Amazon River basin: INPA 1427, holotype, 135.6 mm SL. MZUSP 40140, 1, paratype, 77.3 mm SL. MZUSP 40161, 1, paratype, 86.3 mm SL. MZUSP 40147, 7, paratypes, 117.0–173.7 mm SL. MZUSP 40152, 20, paratypes, 25.2–134.8 mm SL. Crenicichla isbrueckeri. Brazil, Mato Grosso, Rio Madeira basin, Rio Aripuanã: INPA 2883, holotype, 92.8 mm SL. INPA 1492, 5 of 14 paratypes, 28.9–56.5 mm SL. Crenicichla labrina. Brazil, Rio Araguaia basin: MZUSP 89456, 3, 62.9–72.5 mm SL. MZUSP 97983, 5, 61.5– 101.1 mm SL. MZUSP 100006, 1, 121.0 mm SL.

CRENICICHLA PLOEGI NEW SPECIES OF PIKE CICHLID FROM BRAZIL Zootaxa 4377 (3) © 2018 Magnolia Press · 383 Crenicichla lepidota. Brazil, Mato Grosso, Rio Guaporé basin: NMW 33101, lectotype of C. lepidota, 158.0 mm SL. NMW 33102, 1, 159.3 mm SL. NMW 33109, 1, 74.5 mm SL. NMW 33114–17, 3, 31.6–35.7 mm SL. MCP 38281, 1, 60.5 mm SL. MCP 38384, 4, 41.1–82.5 mm SL. MCP 38405, 1, 46.5 mm SL. MCP 38438, 3, 41.1–90.1 mm SL. MCP 38446, 1, 88.7 mm SL. MCP 38608, 1, 46.7 mm SL. MCP 39610, 1, 144.0 mm SL. MZUSP 36939, 1, 137.7 mm SL. MZUSP 37501, 2, 43.8–46.5 mm SL. MZUSP 37625, 1, 64.4 mm SL. MZUSP 37678, 2, 54.3–56.2 mm SL. MZUSP 64952, 1, 155.6 mm SL. Brazil, Rio Paraná basin, Paraná: MHNCI 11779, 1, 89.0 mm SL. NUP 3676, 1, 80.4 mm SL. NUP 7331, 4, 82.3–96.9 mm SL. NUP 7359, 2, 88.8–91.4 mm SL. NUP 11365, 4, 69.4–109.1 mm SL. Brazil, Rio Grande do Sul, Lagoa dos Patos and Lagoa Mirim system: MZUSP 16035, 6, 51.3–78.3 mm SL. MZUSP 41246, 7, 58.3–75.6 mm SL. MZUSP 41249, 1, 74.9 mm SL. MZUSP 41265, 4, 55.7–118.6 mm SL. MZUSP 41269, 2, 125.4–136.3 mm SL. MZUSP 41270, 4, 137.8–189.0 mm SL. Argentina, Província de Entre Rios: MLP 8611, 1, 156.7 mm SL. Paraguay, Rio de La Plata basin: MHNG 2095.045, holotype of C. edithae, 76.9 mm SL. Paratypes of C. edithae: BMNH 1927.11.23.65, 1, 138.3 mm SL. BMNH 1935.6.4.416–19, 7, 38.9–111.4 mm SL. MHNG 2028.030, 1, 69.6 mm SL. MHNG 2028.031–039, 8, 44.6–87.2 mm SL. MHNG 2095.046–058, 10, 56.0–77.6 mm SL. MHNG 2095.040–047, 8, 49.5–111.0 mm SL. Crenicichla aff. lepidota Tapajós. Brazil. Rio Tapajós basin: LBP 7952, 1, 75.1 mm SL. LBP 8008, 24, 20.2–21.7 mm SL. LBP 8020, 4, 34.7–92.4 mm SL. LBP 8607, 2, 50.4–75.1 mm SL. LBP 19605, 1, 56.1 mm SL. LBP 19606, 1, 78.0 mm SL. LBP 8555, 5, 31.2–88.5 mm SL. Crenicichla lucius. Peru, Departamento Loreto: NRM 25662, neotype of C. lucius, 146.5 mm SL, Rio Napo drainage. CAS 36727, 4, 58.6–143.2 mm SL, Río Ampiyacu. Brazil, Amazonas, Rio Purus basin: MZUSP 41289, 3, 60.4–118.1 mm SL. Crenicichla brasiliensis. Brazil, Maranhão, Rio Mearim basin: MZUSP 43620, 3, 95.6–106.7 mm SL. MZUSP 98579, 1, 52.7 mm SL. MZUSP 98580, 1, 34.4 mm SL. MZUSP 98581, 1, 32.2 mm SL. Pernambuco, Rio São Francisco basin: MZUSP 108749, 3, 75.5–115.7 mm SL. Piauí, Rio Parnaíba basin: MZUSP 98563, 11, 10.6– 74.5 mm SL. MZUSP 98578, 12, 15.1–63.4 mm SL. Crenicichla nickeriensis. Suriname, district Nickerie, Nickerie River basin: ZMA 107843, holotype, 184.7 mm SL. IRSNB 765, 3, paratypes, 44.8–104.9 mm SL. ZMA 106505, 29, paratypes, 47.4–129.5 mm SL, 1 c&s. ZMA 106506, 36, paratypes, 34.5–109.69 mm SL. ZMA 116681, 5, paratypes, 86.4–102.2 mm SL. Crenicichla pellegrini. Brazil, Rio Aripuanã basin: INPA 2885, holotype, 112.6 mm SL. INPA 33479, 3, 61.1–77.4 mm SL. INPA 33546, 2, 37.8–60.1 mm SL. Crenicichla proteus. Peru, Departamento Loreto: ANSP 9050, lectotype of C. proteus, 118.4 mm SL. CAS 36717, 1, 105.0 mm SL. MZUSP 77825, 2, 41.8–76.5 mm SL. Brazil, Acre, Rio Japurá basin: MZUSP 40163, holotype of Crenicichla nijsseni, 78.0 mm SL. MZUSP 38532, 4, paratypes of C. nijsseni, 65.8–78.0 mm SL. Crenicichla pydanielae. Brazil, Pará, Rio Trombetas basin. INPA 3074, holotype, 114.3 mm SL. MZUSP 32864, 4, paratypes, 47.1–100.3 mm SL. Crenicichla santosi. Brasil, Rondônia, Rio Madeira basin, Rio Machado: INPA 2889, holotype, 117.1 mm SL. LIRP 10290, 1, 116.3 mm SL. LIRP 10519, 1, 133.8 mm SL. LIRP 10697, 1, 143.8 mm SL. LIRP 10885, 1, 108.9 mm SL. LIRP 11230, 4, 27.2–90.2 mm SL. LIRP 33555, 1, 100.9 mm SL. Crenicichla saxatilis. Suriname: BMNH 1853.11.12.24, holotype of Scarus pavoninus, 105.8 mm SL. NRM 5583, lectotype Sparus saxatilis, 111.5 mm SL; NRM 5585, paralectotype of Sparus saxatilis, 116.3 mm SL. Guyana: CAS 77935, 3, 86.5–118.9 mm SL. CAS 77936, 2, 98.5–108.8 mm SL. CAS 77942, 5, 43.3–94.4 mm SL. Crenicichla semicincta. Bolivia: NMW 33482, lectotype of C. semicincta, 132.2 mm SL. NMW 33483, paralectotype of C. semicincta, 78.6 mm SL. NMW 33484, paralectotype of C. semicincta, 76.2 mm SL. BMNH 1927.10.4.49, holotype of C. clancularia, 72,8 mm SL. BMNH 1927.10.4.50–56, 6, paratypes of C. clancularia, 42.3–76.0 mm SL. Crenicichla sipaliwini. Suriname, Corantijn river basin: ZMA 116683, 4, 62.3–71.4 mm SL. ZMA 107846, 8, 68.4–168.2 mm SL. Crenicichla sveni. Colombia, Rio Orinoco basin: MNHN 31622, holotype, 65.1 mm SL; ZMA 120388, paratype, 62.7 mm SL; ZMA 120651, paratype, 61.1 mm SL. Venezuela, Rio Orinoco basin: MZUSP 96506, 3, 30.7– 40.8 mm SL. MZUSP 96540, 4, 36.1–51.4 mm SL.

First author is grateful to the following curators of ichthyological collections for the loan of material and attention during visits: Georges Lenglet (IRSNB), Mark Sabaj Pérez (ANSP) Castro and Flávio Bockmann (LIRP-USP); Carla Pavanelli and Weferson da Graça (UEM); Vinicius Abilhoa (MHNCI); Roberto Reis e Carlos Lucena (MCP); Amalia Miquelarena e Hugo Lopez (ILPLA e MLP); Lúcia Rapp Py-Daniel and Jansen Zuanon (INPA), James Maclaine, Ralf Britz and Oliver Crimmen (BMNH), Sonia Fisch-Muller (MHNG), Ronald de Ruiter (Naturalis— ZMA, RMNH), Ernst Mikschi (NMW), Patrice Pruvost (MNHN), Kevin Swagel (FMNH), Luiz Rocha, David Catania and Jon Fong (CAS). Thanks to Gustavo Ballen (MZUSP) for the assistance in the statistical analysis. Thanks to Naércio Menezes (MZUSP) for continuous instruction in systematic work. Thanks to Marcelo Krause for the Fig. 4 and to Fernando Dagosta (MZUSP) for Fig. 5. HRV, MVL and FCTL were supported by FAPESP during the development of this paper (grants 2008/56130-1, 2011/14640-0 and 2013/17056-9 to HRV; 2011/06830- 0 to MVL; 2011/51532-7 and 2013/20936-0 to FCTL).

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