Five New Species of the Killifish Genus Anablepsoides from the Brazilian Amazon (Cyprinodontiformes: Rivulidae)

63 (3): 283 – 293

Five new species of the killifish genus Anablepsoides from the Brazilian Amazon (Cyprinodontiformes: Rivulidae)

Wilson J. E. M. Costa, Pedro H. N. Bragança & Pedro F. Amorim

Laboratório de Sistemática e Evolução de Peixes Teleósteos, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Caixa Postal 68049, CEP 21944-970, Rio de Janeiro, Brasil; wcosta(at)acd.ufrj.br, pedrobra88(at)gmail.com, pedro_f_a(at)hotmail.com

Accepted 01.xi.2013. Published online at www.senckenberg.de/vertebrate-zoology on 18.xii.2013.

Abstract Two new species of the Anablepsoides urophthalmus group and three new species of the A. ornatus group, collected in the Brazilian Ama- zon, are described. Anablepsoides jari, new species, and A. roraimaQHZVSHFLHVDUHWKH¿UVWUHFRUGVRIPHPEHUVRIWKHA. urophthalmus group for tributaries of the Amazonas river basin draining the Guiana Shield, but phylogenetic relationships of these species with other taxa are still uncertain. Anablepsoides jariIURPWKH-DULULYHUGUDLQDJHLVGLVWLQJXLVKHGIURPFRQJHQHUVE\WKHFRORXUSDWWHUQRIÀDQNDQG XQSDLUHG¿QVLQPDOHVZKHUHDVAnablepsoides roraimaIURPWKH%UDQFRULYHUGUDLQDJHLVGLVWLQJXLVKHGE\WKHVSRWWHGSDWWHUQRIÀDQNLQ males. Relationships of the three new species of the A. ornatus group, Anablepsoides henschelae, new species, Anablepsoides ottonii, new species, and Anablepsoides gamaeQHZVSHFLHVDUHWHQWDWLYHO\LQIHUUHGXSRQWKHEDVLVRIGHULYHGFRORXUSDWWHUQVRIFDXGDO¿QLQPDOHV which are also informative to diagnose each new species. Anablepsoides ottonii seems to be closely related to A. ornatus by both sharing DSRLQWHGFDXGDO¿QLQODUJHUVSHFLPHQVDQGWKHSUHVHQFHRIUHGGLVKEURZQVSRWVRQWKHGRUVDOSRUWLRQRIWKHFDXGDO¿QLQPDOHVZKHUHDV A. gamae and A. amanan VKDUHDFRORXUSDWWHUQRIFDXGDO¿QLQPDOHVXQLTXHDPRQJDSORFKHLORLGVLQZKLFKWKHGRUVDOSRUWLRQRIWKH¿Q LVRUDQJLVKUHGZLWKGRUVDOPDUJLQZKLWHDQGQDUURZEODFNRXWHUERUGHUAnablepsoides henschelae is considered as the sister group to all RWKHUVSHFLHVRILWVJURXSE\QRWVKDULQJDQDSRPRUSKLFDV\PPHWULFDOO\FRORXUHGFDXGDO¿QLQPDOHV

Key words Amazon Forest, Amazonas river, Biodiversity, Guiana Shield, Systematics.

Introduction

Anablepsoides HUBER, 1992 is a species-rich genus of cies presently assigned to Anablepsoides were described 1HRWURSLFDODSORFKHLORLGNLOOL¿VKHVRFFXUULQJLQQRUWK- from the Brazilian Amazon between 1863 and 1916 ern South America, including the Amazonas river basin, (STEINDACHNER, 1863, 1877; GÜNTHER, 1866; GARMAN, and in adjacent Caribbean islands (COSTA, 2011). Within 1895; HENN, 1916), but no new description for taxa oc- this geographical distribution, the Amazonas river ba- curring in this huge area was published between 1916 sin concentrates the highest diversity both in number DQGDOWKRXJKIUHTXHQWUHFHQWFROOHFWLRQVGHSRVLWHG of species and in specialized forms. However, whereas in Brazilian museums. Possibly this was a result of the taxonomical studies developed in the last decades fo- LQFRPSOHWH NQRZOHGJH DERXW VSHFLHV GHVFULEHG EHIRUH cused on taxa endemic to western Amazon, in Colombia, WKDW SHULRG PDNLQJ LGHQWL¿FDWLRQ RI UHFHQW FROOHFWLRQV Ecuador, and Peru, little advance has been recorded for uncertain. A new approach for the taxonomical state of the Brazilian Amazon (COSTA, 2013). Seven nominal spe- Anablepsoides from the Brazilian Amazon was provided

ISSN 1864-5755 283 W.J.E.M. Costa et al.: Five new Anablepsoides from the Brazilian Amazon

by COSTA (2006), when new morphological characters Table 1. Morphometric data of Anablepsoides jari. ZHUHLQFRUSRUDWHGWRULYXOLQHWD[RQRP\PDNLQJSRVVLEOH both to delimit taxa assigned to old names and to rec- holotype paratypes ognise new species (COSTA & LAZZAROTO, 2008; COSTA, male males (5) females (2) 2010, 2013). Standard length (mm) 25.6 22.5 – 31.0 20.8 – 24.5 Amazonian species of Anablepsoides have been Percent of standard length placed in different informal groups diagnosed by mor- Body depth 19.5 20.1 – 21.6 18.2 – 18.6 phological characters. COSTA (2010) recognised two groups of similar species sharing longitudinal rows Caudal peduncle depth 13.8 13.5 – 15.1 11.8 – 13.1 RI UHG VSRWV RQ WKH ÀDQN LQ PDOHV IURQWDO VTXDPDWLRQ Pre-dorsal length 80.3 78.0 – 81.9 77.9 – 79.9 (SDWWHUQHGDQGVKRUWXQSDLUHG¿QVGRUVDO¿QZLWK± Pre-pelvic length 53.6 54.2 – 57.8 54.4 – 55.2 Length of dorsal-fin base 6.8 6.2 – 8.8 7.3 – 7.5 UD\VDQGDQDO¿QZLWK±UD\V7KH¿UVWRQHWKHA. Length of anal-fin base 16.0 16.6 – 19.4 16.2 – 18.4 urophthalmus group, included species of central and Caudal-fin length 33.4 32.0 – 38.0 36.2 – 37.1 eastern Amazon, Guianas and eastern Venezuela, and the Pectoral-fin length 20.5 20.7 – 24.4 19.9 – 21.8 second one, the A. limoncochae group, included species Pelvic-fin length 10.0 8.8 – 11.8 8.0 – 9.3 from central and western Amazon. The former group was Head length 24.4 24.3 – 26.5 25.5 – 26.1 distinguished from the latter by the absence of contact or- JDQVRQERG\VFDOHVDQGDEVHQFHRIDQREOLTXHUHGVWULSH Percent of head length RQPLGGOHRIWKHGRUVDO¿QEHVLGHVVSHFLHVRIWKHIRUPHU Head depth 63.0 61.7 – 67.6 59.3 – 62.0 group often having yellow pigmentation concentrated on Head width 81.3 75.4 – 81.4 74.5 – 75.9 WKHGRUVDODQGYHQWUDOPDUJLQVRIWKHFDXGDO¿QLQPDOHV Snout length 14.3 14.8 – 17.1 12.8 – 14.0 Although species of both groups being morphologically Lower jaw length 21.2 18.6 – 21.3 18.4 – 19.0 similar, molecular evidence has indicated that species of Eye diameter 31.9 31.1 – 34.6 30.8 – 32.8 the A. urophthalmus group are more closely related to other species of Anablepsoides from the coastal rivers of northern South America than to species of the A. li- derived from distribution of melanophores were also di- moncochae group, whereas species of the latter group are rectly observed in all preserved specimens. Measurements more closely related to other species of Anablepsoides and counts follow COSTA (1995). Measure ments are pre- from the Amazon than to species of the A. urophthalmus sented as percentages of standard length (SL), except for group (e.g., MURPHY et al., 1999). those related to head morphology, which are expressed Another species group of Anablepsoides peculiar as percentages of head length. Fin-ray counts include all to the Amazon is that comprising A. ornatus (GARMAN, HOHPHQWV 1XPEHU RI YHUWHEUDH DQG JLOOUDNHUV ZHUH UH- 1895) and A. amanan (COSTA & LAZZAROTO, 2008), herein corded only from the cleared and stained specimen; the named as the A. ornatus group. Species of this group are compound caudal centrum was counted as a single ele- small (maximum size about 28 mm SL), have depressed ment. The osteological preparation was made according KHDGDQGORQJFDXGDO¿Q EHWZHHQDERXWDQG to TAYLOR & VAN DYKE (1985). Terminology for cephalic SL). The objective of this paper is to describe two new neuromast series follows COSTA (2001). The abbreviation species of the A. urophthalmus group and three new spe- C&S means specimens cleared and stained for bone and cies of the A. ornatus group collected in Amazonas river FDUWLODJH'HOLPLWDWLRQRIVSHFLHVLVDFFRUGLQJWRWKHPHWK- tributaries draining the Guiana Shield and adjacent areas odology of the Population Aggregation Analysis (DAVIS & of central Brazilian Amazon. NIXON LQZKLFKVSHFLHVDUHGHOLPLWHGE\DXQLTXH combination of morphological character states.

Material and methods Anablepsoides jari spec. nov.

Fig. 1, Table 1 Material is deposited in the ichthyological collections of ,(3$,QVWLWXWRGH3HVTXLVDV&LHQWt¿FDVH7HFQROyJLFDVGR Estado do Amapá, Macapá, INPA, Instituto Nacional de Holotype: 8)5- , male, 25.6 mm SL; Brazil: Estado do 3HVTXLVDVGD$PD]{QLD0DQDXVDQG8)5-,QVWLWXWRGH $PDSi0XQLFtSLRGH/DUDQMDOGR-DULVWUHDPWULEXWDU\RI-DULULY %LRORJLD8QLYHUVLGDGH)HGHUDOGR5LRGH-DQHLUR5LRGH HUORZHU$PD]RQDVULYHUEDVLQƍƎ6ƍƎ:P.H.N. BRAGANÇA & E. HENSCHEL, 25 July 2012. Janeiro. Morphological characters used in the description EHORZ ZHUH REWDLQHG IURP VSHFLPHQV ¿[HG LQ IRUPDOLQ Paratypes: $OO IURP 0XQLFtSLR GH /DUDQMDO GR -DUL (VWDGR GR just after collection, for a period of about ten days, and Amapá, Brazil. 8)5-, 1 female, 20.8 mm SL; 8)5-, 2 PDOHV±PP6/IHPDOHPP6/ & 6 FROOHFWHG WKHQWUDQVIHUUHGWRHWKDQRO'HVFULSWLRQVRIFRORXU ZLWKKRORW\SH±,13$PDOHV±PP6/IH patterns were based on photographs of live individuals PDOHV±PP6/VWUHDPWULEXWDU\WR-DULULYHUMXVWEHORZ WDNHQLQVPDOODTXDULDMXVWDIWHUFROOHFWLRQFRORXUSDWWHUQV 6DQWR$QW{QLRZDWHUIDOOM. JEGU & J. ZUANON, 21 June 1987.

284 VERTEBRATE ZOOLOGY — 63 (3) 2013

Fig. 1. Anablepsoides jari8)5-KRORW\SHPDOHPP6/%UD]LO$PDSi/DUDQMDOGR-DUL

Diagnosis. 'LVWLQJXLVKHG IURP DOO RWKHU VSHFLHV RI WKH GLDQRSHUFXODUYHQWUDORSHUFXODUSUHRSHUFXODU A. urophthalmus group by having longitudinal rows of PDQGLEXODU ODWHUDO PDQGLEXODU SDUDPDQGLEXODU red dots interrupted and replaced by short rows of golden /DWHUDOOLQHLQWHUUXSWHGDOWHUQDWLQJVHWVRI±VFDOHV GRWVRQWKHÀDQNLQPDOHV YVURZVRIUHGGRWVRIWHQFRQ- with one neuromast and without neuromasts. Two neuro- tinuous, when interrupted not replaced by golden dots), PDVWVRQFDXGDO¿QEDVH PDUJLQVRIXQSDLUHGGDUNJUH\LQPDOHV YVQHYHUGDUN %DVLK\DOVXEWULDQJXODUJUHDWHVWZLGWKDERXWRI SLJPHQW FRQFHQWUDWHG RQ ¿Q HGJH WR IRUP GLVWLQFW FRO- OHQJWKEDVLK\DOFDUWLODJHDERXWRIWRWDOOHQJWKRI oured margins), and a horizontal row of red dots on the basihyal. Six branchiostegal rays. Two teeth on second VXEEDVDOUHJLRQRIWKHDQDO¿QLQPDOHV QHYHUDVLPLODU SKDU\QJREUDQFKLDO *LOOUDNHUV RQ ¿UVW EUDQFKLDO DUFK colour pattern). 9RPHULQH WHHWK DEVHQW 'HUPRVSKHQRWLF SUHVHQW Ventral process of posttemporal rudimentary. Total ver- Description. 0RUSKRPHWULFGDWDDSSHDULQ7DEOH'RU WHEUDH± VDODQGYHQWUDOSUR¿OHVVOLJKWO\FRQYH[IURPVQRXWWRSRV- WHULRUHQGRIGRUVDODQGDQDO¿QEDVHVQHDUO\VWUDLJKWRQ Colouration. Males)ODQNOLJKWEOXHZLWKHLJKWORQJL caudal peduncle. Body slender, sub-cylindrical, slightly tudinal rows of red dots; rows often interrupted and re- depressed on head, compressed posteriorly. Greatest placed by short rows of golden dots; three rows poste- ERG\ GHSWK DW YHUWLFDO MXVW LQ IURQW WR SHOYLF¿Q EDVH ULRUO\ UHDFKLQJ FDXGDO ¿Q EDVH RWKHU ¿YH UHDFKLQJ -DZVVKRUWVQRXWZHDNO\SRLQWHGLQODWHUDOYLHZ YHUWLFDO EHWZHHQ DQWHULRU SRUWLRQ RI DQDO¿Q EDVH DQG 'RUVDODQGDQDO¿QVVPDOOVXEUHFWDQJXODUH[WUHPLW\ SRVWHULRUSRUWLRQRIGRUVDO¿QEDVH'RUVXPOLJKWEURZQ URXQGHG&DXGDO¿QHOOLSWLFDOVKRUW3HFWRUDO¿QURXQG- ODWHUDOO\ERUGHUHGE\LUUHJXODUGDUNEURZQLVKJUH\VWULSH HGSRVWHULRUPDUJLQUHDFKLQJYHUWLFDODWDERXWRI Venter white. Side of head and jaws light grey, opercular OHQJWKEHWZHHQSHFWRUDO¿QDQGSHOYLF¿QEDVHV3HOYLF region purplish grey, with greenish blue iridescence. Iris ¿QVPDOOVOLJKWO\SRLQWHGWLSUHDFKLQJDQXVSHOYLF¿Q SDOHJUHHQLVK\HOORZGRUVDOSRUWLRQGDUNJUH\LVK\HOORZ EDVHVPHGLDOO\LQFORVHSUR[LPLW\'RUVDO¿QRULJLQRQ 'RUVDO¿QOLJKW\HOORZZLWKVKRUWWUDQVYHUVHJUH\VWULSHV YHUWLFDOWKURXJKEDVHRIODVWWZRDQDO¿QUD\VRUVOLJKWO\ RQSRVWHULRUSRUWLRQGLVWDOPDUJLQEODFN$QDO¿QOLJKW SRVWHULRUWRDQDO¿QEDVHVHFRQGSUR[LPDOUDGLDORIGRU- \HOORZEDVDOSRUWLRQOLJKWEOXHGLVWDOPDUJLQGDUNJUH\ VDO¿QEHWZHHQQHXUDOVSLQHVRIndDQGth vertebrae, KRUL]RQWDO URZ RI UHG GRWV RQ VXEEDVDO UHJLRQ RI ¿Q ¿UVWSUR[LPDOUDGLDORIDQDO¿QEHWZHHQSOHXUDOULEVRI &DXGDO¿QJUHHQLVK\HOORZZLWKWUDQVYHUVHURZVRIVPDOO 15th and 17thYHUWHEUDH'RUVDO¿QUD\V±DQDO¿QUD\V JUH\VSRWV¿QPDUJLQJUH\WREODFN3HFWRUDO¿Q\HOORZ- ±FDXGDO¿QUD\V±SHFWRUDO¿QUD\V± LVKK\DOLQH3HOYLF¿QOLJKW\HOORZ SHOYLF¿QUD\V Females. Colour pattern of live females not recorded. Scales small, cycloid. Body and head entirely scaled, ,QSUHVHUYHGVSHFLPHQVÀDQNOLJKWEURZQZLWKLUUHJX- H[FHSW RQ MDZV %RG\ VTXDPDWLRQ H[WHQGLQJ RYHU DQ- larly arranged light grey dots. Side of head light brown. WHULRURIFDXGDO¿QEDVHQRVFDOHVRQGRUVDODQG 8QSDLUHG¿QVK\DOLQHZLWKWUDQVYHUVHURZVRIJUH\GRWV DQDO¿QEDVHV)URQWDOVTXDPDWLRQ(SDWWHUQHG(VFDOHV URXQGGDUNJUH\WREODFNVSRWRQGRUVDOSRUWLRQRIFDX- not overlapping medially; scales arranged in regular cir- GDO¿QEDVH3DLUHG¿QVK\DOLQH cular pattern around A-scale without exposed margins; transverse row of scales anterior to H-scale. Longitudinal Distribution. Known only from the type locality area VHULHVRIVFDOHV±WUDQVYHUVHVHULHVRIVFDOHVVFDOH in the lower Jari river drainage, Amazonas river basin, rows around caudal peduncle 16. No contact organs on northern Brazil. ÀDQNDQG¿QV Cephalic neuromasts: supraorbital 3 + 3, parietal 1, Etymology. The name jari is a reference to the main riv- anterior rostral 1, posterior rostral 1, infraorbital 1 + 15 + er at the type locality of the new species. 1, preorbital 2, otic 1, post-otic 2, supratemporal 1, me-

285 W.J.E.M. Costa et al.: Five new Anablepsoides from the Brazilian Amazon

Anablepsoides roraima spec. nov. Table 2. Morphometric data of Anablepsoides roraima.

Fig. 2, Table 2 holotype paratypes male males (3) females (5) Standard length (mm) 31.0 26.3 – 31.8 26.3 – 29.0 Holotype: 8)5- , male, 31.0 mm SL; Brazil: Estado de Percent of standard length 5RUDLPD0XQLFtSLRGH%RQ¿PVWUHDPNP1(RI%RD9LVWD URDG%57DFXWXULYHUGUDLQDJHSDUWRI%UDQFRULYHUGUDLQDJH Body depth 22.1 19.7 – 21.4 19.6 – 21.5 PLGGOH $PD]RQDV ULYHU EDVLQ ƍƎ1 ƍƎ: P.H.N. Caudal peduncle depth 14.3 13.4 – 14.4 13.6 – 14.2 BRAGANÇA, F. OTTONI & E. HENSCHEL, 10 September 2012. Pre-dorsal length 79.3 76.8 – 81.0 78.5 – 81.3 Paratypes: 8)5-PDOHV±PP6/IHPDOHV Pre-pelvic length 55.0 53.9 – 57.2 54.3 – 60.3 ±PP6/8)5-PDOHV±PP6/IH- Length of dorsal-fin base 7.2 7.0 – 8.7 6.7 – 8.1 PDOHV±PP6/FROOHFWHGZLWKKRORW\SH Length of anal-fin base 18.0 18.6 – 19.0 19.2 – 20.4 Caudal-fin length 33.8 32.8 – 34.7 34.3 – 35.6 Additional material (non-types): 8)5-, 8 ex.; 8)5-, Pectoral-fin length 18.3 18.6 – 20.7 18.9 – 21.2 H[ & 6 %UD]LO (VWDGR GH 5RUDLPD 0XQLFtSLR GH &DQWi Pelvic-fin length 7.5 7.1 – 9.5 7.3 – 8.7 ÀRRGSODLQV RI VWUHDP LQ WKH URDG %5 &DFKRUUR ULYHU GUDLQ- Head length 24.3 23.5 – 25.1 23.8 – 26.4 age, part of Branco river drainage, middle Amazonas river basin, Percent of head length ƍ1ƍ: P.H.N. BRAGANÇA, F. OTTONI & E. HEN- SCHEL, 13 September 2012. Head depth 72.4 65.8 – 71.9 65.7 – 70.4 Head width 85.8 81.9 – 85.6 78.8 – 86.4 Diagnosis. 'LVWLQJXLVKHG IURP DOO RWKHU VSHFLHV RI WKH Snout length 14.6 14.4 – 16.0 14.1 – 15.5 JHQXVE\SRVVHVVLQJÀDQNZLWKHLJKWORQJLWXGLQDOURZV Lower jaw length 22.8 23.0 – 23.5 20.0 – 24.5 of spots, among which the spots of the three dorsal-most Eye diameter 31.7 31.6 – 32.9 31.1 – 32.4 rows are reddish brown and separated by small interspace, DQGWKHVSRWVRIWKHRWKHU¿YHURZVDUHUHGDQGLQFRQ- tact to form long red stripes (vs. spots never forming that VHULHVRIVFDOHV±WUDQVYHUVHVHULHVRIVFDOHVVFDOH arrangement pattern). Also distinguished from all other rows around caudal peduncle 16. No contact organs on species of Anablepsoides from the Brazilian Amazon by ÀDQNDQG¿QV WKHSUHVHQFHRIWZRZHOOGH¿QHGEULJKW\HOORZVWULSHVRQ Cephalic neuromasts: supraorbital 3 + 3, parietal WKHGRUVDODQGYHQWUDOPDUJLQVRIWKHFDXGDO¿QLQPDOHV 2, anterior rostral 1, posterior rostral 1, infraorbital (vs. stripes diffuse or absent). ±SUHRUELWDORWLFSRVWRWLFVXSUDWHP- poral 1, median opercular 1, ventral opercular 1, preoper- Description. Morphometric data appear in Table 2. FXODUPDQGLEXODUODWHUDOPDQGLEXODUSDUD- 'RUVDODQGYHQWUDOSUR¿OHVJHQWO\FRQYH[IURPVQRXWWR mandibular 1. Lateral line interrupted, alternating sets of SRVWHULRUHQGRIGRUVDODQGDQDO¿QEDVHVQHDUO\VWUDLJKW ±VFDOHVZLWKRQHQHXURPDVWDQGZLWKRXWQHXURPDVWV on caudal peduncle. Body slender, sub-cylindrical an- 7ZRQHXURPDVWVRQFDXGDO¿QEDVH teriorly, slightly depressed on head, compressed pos- %DVLK\DOVXEWULDQJXODUJUHDWHVWZLGWKDERXWRI teriorly. Greatest body depth at vertical just in front to OHQJWKEDVLK\DOFDUWLODJHDERXWRIWRWDOOHQJWKRI SHOYLF¿QEDVH-DZVVKRUWVQRXWZHDNO\SRLQWHGLQODW- basihyal. Six branchiostegal rays. Two teeth on second eral view. SKDU\QJREUDQFKLDO *LOOUDNHUV RQ ¿UVW EUDQFKLDO DUFK 'RUVDODQGDQDO¿QVVPDOOVXEUHFWDQJXODUH[WUHPLW\ 9RPHULQHWHHWK'HUPRVSKHQRWLFSUHVHQW9HQWUDO URXQGHG&DXGDO¿QHOOLSWLFDOVKRUW3HFWRUDO¿QURXQG- process of posttemporal rudimentary. Total vertebrae HGSRVWHULRUPDUJLQUHDFKLQJYHUWLFDODWDERXWRI ± OHQJWKEHWZHHQSHFWRUDO¿QDQGSHOYLF¿QEDVHV3HOYLF ¿Q VPDOO VOLJKWO\ SRLQWHG WLS UHDFKLQJ DQXV LQ PDOHV Colouration. Males.)ODQNJURXQGFRORXUJROGHQWROLJKW VOLJKWO\VKRUWHULQIHPDOHVSHOYLF¿QEDVHVPHGLDOO\LQ EOXH RQ YHQWUDO SDUW )ODQN ZLWK HLJKW FRQWLQXRXV ORQ- FORVH SUR[LPLW\ 'RUVDO¿Q RULJLQ RQ YHUWLFDO WKURXJK JLWXGLQDOURZVRIGRWVVHYHQUHDFKLQJFDXGDO¿QEDVH base of 10th or 11thDQDO¿QUD\VHFRQGSUR[LPDOUDGLDO WKUHH GRUVDOPRVW URZV ZLWK VSRWV GDUN UHGGLVK EURZQ RIGRUVDO¿QEHWZHHQQHXUDOVSLQHVRIndDQGth ver- DQGVHSDUDWHGE\VPDOOLQWHUVSDFHRWKHU¿YHURZVZLWK WHEUDH¿UVWSUR[LPDOUDGLDORIDQDO¿QEHWZHHQSOHXUDO UHGVSRWVLQFRQWDFWIRUPLQJORQJVWULSHV'RUVXPOLJKW ULEVRIth and 16thYHUWHEUDH'RUVDO¿QUD\V±DQDO brown, venter white. Side of head light brown, opercular ¿QUD\V±FDXGDO¿QUD\V±SHFWRUDO¿QUD\V region greenish golden to purplish grey with blue irides- ±SHOYLF¿QUD\V cence on ventral portion. Iris pale greenish yellow, dor- Scales small, cycloid. Body and head entirely scaled, VDOSRUWLRQGDUNJUH\LVK\HOORZ'RUVDO¿QOLJKW\HOORZ H[FHSW RQ MDZV %RG\ VTXDPDWLRQ H[WHQGLQJ RYHU DQ- with short transverse grey stripes on posterior portion. WHULRURIFDXGDO¿QEDVHQRVFDOHVRQGRUVDODQG $QDO¿QOLJKW\HOORZEDVDOSRUWLRQOLJKWEOXHVRPHWLPHV DQDO¿QEDVHV)URQWDOVTXDPDWLRQ(SDWWHUQHG(VFDOHV ZLWKJUH\GRWVRQSRVWHULRUSRUWLRQ&DXGDO¿QOLJKWJUH\ not overlapping medially; scales arranged in regular cir- with minute brown dots, with yellow stripe on dorsal and cular pattern around A-scale without exposed margins; YHQWUDOPDUJLQV3HFWRUDO¿Q\HOORZLVKK\DOLQH3HOYLF¿Q transverse row of scales anterior to H-scale. Longitudinal light blue.

286 VERTEBRATE ZOOLOGY — 63 (3) 2013

Fig. 2. Anablepsoides roraima8)5-KRORW\SHPDOHPP6/%UD]LO5RUDLPD%RQ¿P

Fig. 3. Anablepsoides henschelae8)5-KRORW\SHPDOHPP6/%UD]LO5RUDLPD5RUDLQySROLV

Females.)ODQNOLJKWEURZQZLWKORQJLWXGLQDOURZVRI Diagnosis. 'LVWLQJXLVKHG IURP DOO RWKHU VSHFLHV RI WKH pale brown dots and irregularly arranged light blue to A. ornatus JURXSE\KDYLQJWKHFDXGDO¿QLQPDOHVZLWK pale golden dots. Side of head light brown, opercular re- similar colouration on its dorsal and ventral portions gion metallic green with purplish blue iridescence. Iris (vs. always distinct colouration) and the presence of a OLJKW\HOORZ8QSDLUHG¿QVK\DOLQHZLWKWUDQVYHUVHURZV ORQJLWXGLQDOURZRIGDUNUHGGRWVRQWKHYHQWUDOSRUWLRQ RIJUH\GRWVURXQGGDUNJUH\WREODFNVSRWRQGRUVDOSRU- RIWKHÀDQNLQPDOHV YVORQJLWXGLQDOURZRIGDUNUHG WLRQRIFDXGDO¿QEDVH3DLUHG¿QVK\DOLQH dots absent).

Distribution. Middle Branco river drainage, Amazonas Description. 0RUSKRPHWULFGDWDDSSHDULQ7DEOH'RU river basin, northern Brazil. VDOSUR¿OHVOLJKWO\FRQYH[IURPVQRXWWRHQGRIFDXGDO ¿QEDVH9HQWUDOSUR¿OHDSSUR[LPDWHO\VWUDLJKWVOLJKWO\ Etymology. The name roraima refers to the occurrence FRQYH[ RQ SHOYLF¿Q UHJLRQ %RG\ VOHQGHU GHSUHVVHG of the new species in the Brazilian State of Roraima. on head, gradually becoming compressed posteriorly. *UHDWHVWERG\GHSWKDWYHUWLFDOWKURXJKSHOYLF¿QEDVH Jaws short, snout slightly pointed in lateral view. 'RUVDO DQG DQDO ¿QV VPDOO H[WUHPLW\ URXQGHG GRUVDO VXEUHFWDQJXODU DQDO ¿Q VXEWULDQJXODU &DXGDO Anablepsoides henschelae spec. nov. ¿QRYDOVOHQGHUDQGORQJ3HFWRUDO¿QURXQGHGSRVWH- ULRUPDUJLQUHDFKLQJYHUWLFDOWKURXJKSHOYLF¿QEDVHLQ )LJVD7DEOH PDOHWKURXJKXURJHQLWDOSDSLOODLQIHPDOHV3HOYLF¿Q HOOLSWLFDO H[WUHPLW\ UHDFKLQJ DQDO¿Q RULJLQ LQ PDOHV UHDFKLQJXURJHQLWDOSDSLOODLQIHPDOHV3HOYLF¿QEDVHV Holotype: 8)5- , male, 22.5 mm SL; Brazil: Estado de PHGLDOO\ LQ FORVH SUR[LPLW\ 'RUVDO¿Q EDVH SRVWHULRU 5RUDLPD 0XQLFtSLR GH 5RUDLQySROLV VWUHDP DERXW NP: RI -XQGLiURDG%5-DXDSHULULYHUGUDLQDJHLWVHOIDSDUWRIWKH WR DQDO¿Q EDVH VHFRQG SUR[LPDO UDGLDO RI GRUVDO ¿Q th st lower section of the Negro river drainage, Amazonas river ba- between neural spines of 19 and 21 YHUWHEUDH ¿UVW th VLQ¶´6¶´:P.H.N. BRAGANÇA, F.P. OTTONI & E. SUR[LPDOUDGLDORIDQDO¿QEHWZHHQSOHXUDOULEVRI HENSCHEL, 15 September 2012. and 15thYHUWHEUDH'RUVDO¿QUD\VDQDO¿QUD\V± Paratypes: 8)5-IHPDOHV±PP6/8)5-, FDXGDO¿QUD\V±SHFWRUDO¿QUD\V±SHOYLF IHPDOHV±PP6/ F V FROOHFWHGZLWKWKHKRORW\SH ¿QUD\V

287 W.J.E.M. Costa et al.: Five new Anablepsoides from the Brazilian Amazon

otic 1, supratemporal 1, median opercular 1, ventral RSHUFXODUSUHRSHUFXODUPDQGLEXODUODWHUDO mandibular 2, paramandibular 1. Lateral line interrupted, DOWHUQDWLQJ VHWV RI ±VFDOHV ZLWK RQH QHXURPDVW DQG ZLWKRXWQHXURPDVWV7ZRQHXURPDVWVRQFDXGDO¿QEDVH %DVLK\DOVXEWULDQJXODUJUHDWHVWZLGWKDERXWRI OHQJWKEDVLK\DOFDUWLODJHDERXWRIWRWDOOHQJWKRI basihyal. Six branchiostegal rays. Second pharyngob- UDQFKLDOWHHWKDEVHQW*LOOUDNHUVRQ¿UVWEUDQFKLDODUFK 9RPHULQH WHHWK ± 'HUPRVSKHQRWLF SUHVHQW Ventral process of posttemporal absent. Total vertebrae ±

Colouration. Male.)ODQNSDOHSXUSOLVKJUH\ZLWKRE OLTXHGRXEOHURZVRIGDUNUHGGRWVFRQQHFWHGWRORQJLWX- GLQDOURZRIVLPLODUGRWVDORQJYHQWUDOSDUWRIÀDQNDQG other one on middle of caudal peduncle; golden irides- FHQFH DPRQJ REOLTXH GRW URZV 'RUVXP OLJKW \HOORZ- LVKJUH\ZLWKVHJPHQWHGORQJLWXGLQDOGDUNEURZQVWULSH RQODWHUDOSRUWLRQ2EOLTXHEODFNEDUEHWZHHQDUHDMXVW DERYH SHFWRUDO¿Q EDVH DQG PLGGOH RI RSHUFOH 9HQWHU light grey. Side of head purplish grey, opercular and sub- RUELWDO UHJLRQV ZLWK GHQVH FRQFHQWUDWLRQ RI VPDOO GDUN JUH\WREODFNVSRWVDQGGDUNEOXHLULGHVFHQFHRQYHQWUDO SRUWLRQRIRSHUFOHWUDQVYHUVHEODFNEDUEHWZHHQRUELWV through chin. Ventral portion of head with dense concen- WUDWLRQRIVPDOOEODFNVSRWVDQWHULRUO\ERUGHUHGE\QRW SLJPHQWHGDUHDMXVWSRVWHULRUWRFKLQEDU,ULVGDUNEURZQ ZLWKSRVWHURYHQWUDOVLOYHUVSRW'RUVDO¿QK\DOLQHZLWK WZRQDUURZGDUNJUH\WUDQVYHUVHVWULSHV$QDO¿QK\DOLQH WREOXLVKZKLWHRQEDVDOSRUWLRQZLWKWKUHHQDUURZGDUN JUH\ WUDQVYHUVH VWULSHV &DXGDO ¿Q K\DOLQH ZLWK VPDOO EODFNVSRWVDQGSDOHEOXHLULGHVFHQFHRQPLGGOHRIEDVDO SRUWLRQ3HFWRUDO¿Q\HOORZLVKK\DOLQH3HOYLF¿QK\DOLQH Females. )ODQN SDOH SXUSOLVK JUH\ ZLWK REOLTXH GRX- EOHURZVRIGDUNUHGGRWVDQGQDUURZORQJLWXGLQDOGDUN EURZQLVKJUH\VWULSHV'RUVXPOLJKW\HOORZLVKJUH\ZLWK VHJPHQWHGORQJLWXGLQDOGDUNEURZQVWULSHRQODWHUDOSRU- WLRQ2EOLTXHEODFNEDUEHWZHHQDUHDMXVWDERYHSHFWRUDO ¿QEDVHDQGPLGGOHRIRSHUFOH9HQWHUOLJKWJUH\6LGHRI Fig. 4. Colour pattern of the head in females, left side, of: a, head purplish grey, opercular and suborbital regions with Anablepsoides henschelae 8)5- SDUDW\SH PP 6/ GHQVH FRQFHQWUDWLRQ RI VPDOO GDUN JUH\ WR EODFN VSRWV b, Anablepsoides ottonii 8)5- SDUDW\SH PP 6/ F WUDQVYHUVHEODFNEDUEHWZHHQRUELWVWKURXJKFKLQ9HQWUDO Anablepsoides gamae8)5-SDUDW\SHPP6/ SRUWLRQRIKHDGZLWKORQJLWXGLQDOURZRIEODFNVSRWVDQ- teriorly bordered by not pigmented area just posterior WRFKLQEDU,ULVGDUNEURZQZLWKSRVWHURYHQWUDOVLOYHU Scales small, cycloid. Body and head entirely scaled, VSRW'RUVDO¿QK\DOLQHZLWKWZRQDUURZGDUNJUH\WUDQV- H[FHSW RQ MDZV %RG\ VTXDPDWLRQ H[WHQGLQJ RYHU DQ- YHUVH VWULSHV$QDO ¿Q K\DOLQH ZLWK WKUHH QDUURZ GDUN WHULRURIFDXGDO¿QEDVHQRVFDOHVRQGRUVDODQG JUH\WUDQVYHUVHVWULSHV&DXGDO¿QK\DOLQHZLWKQDUURZ DQDO¿QEDVHV)URQWDOVTXDPDWLRQ)SDWWHUQHG(VFDOHV GDUNJUH\WUDQVYHUVHEDUV3HFWRUDO¿Q\HOORZLVKK\DOLQH not overlapping medially; scales arranged in regular cir- 3HOYLF¿QK\DOLQH cular pattern around A-scale without exposed margins; transverse row of scales anterior to H-scale. Longitudinal Distribution. Known only from the type locality, a pool series of scales 27; transverse series of scales 7; scale DGMDFHQWWRVWUHDPDERXWNP:RI-XQGLiURDG%5 rows around caudal peduncle 12. No contact organs on -DXDSHULULYHUGUDLQDJH5RUDLPDVWDWH%UD]LO ÀDQNDQG¿QV Cephalic neuromasts: supraorbital 3 + 3, parietal 1, Etymology. Named after Elisabeth Henschel, for her rostral 1, infraorbital 2 + 11 + 1, preorbital 3, otic 1, post- valuable help during collecting trips in the Amazon.

288 VERTEBRATE ZOOLOGY — 63 (3) 2013

Fig. 5. Anablepsoides ottonii8)5-KRORW\SHPDOHPP6/%UD]LO$PD]RQDV%DUFHORV

Anablepsoides ottonii spec. nov. EDVH9HQWUDOSUR¿OHDSSUR[LPDWHO\VWUDLJKW%RG\VOHQ- der, depressed on head, gradually becoming compressed )LJVE7DEOH posteriorly. Greatest body depth at vertical through pel- YLF¿QEDVH-DZVVKRUWVQRXWVOLJKWO\SRLQWHGLQODWHUDO Holotype: 8)5-, male, 20.1 mm SL; Brazil: Estado do Ama- view. ]RQDV 0XQLFtSLRGH %DUFHORV,JDUDSp GR &DMDUD]LQKRD VWUHDP 'RUVDODQGDQDO¿QVVPDOOH[WUHPLW\URXQGHGGRU- QHDUWKHYLOODJHRI%DODLRDWULEXWDU\RIWKH&DXUpVULYHU1HJUR VDO VXEUHFWDQJXODU DQDO ¿Q VXEWULDQJXODU &DXGDO ¿Q ULYHU GUDLQDJH $PD]RQDV ULYHU EDVLQ ƍƎ6 ¶Ǝ: SRLQWHGVOHQGHUDQGORQJ3HFWRUDO¿QURXQGHGSRVWHULRU P.H.N. BRAGANÇA, P. AMORIM & F.P. OTTONI, 17 November 2012. PDUJLQUHDFKLQJRIGLVWDQFHEHWZHHQSHFWRUDO¿Q Paratypes: %UD]LO(VWDGRGR$PD]RQDV0XQLFtSLRGH%DUFHORV EDVHDQGSHOYLF¿QEDVH3HOYLF¿QHOOLSWLFDOH[WUHPLW\ Negro river drainage, Amazonas river basin. 8)5-, 3 males, UHDFKLQJXURJHQLWDOSDSLOODSHOYLF¿QEDVHVPHGLDOO\LQ ±PP6/IHPDOHV±PP6/DQGMXYHQLOHV FORVH SUR[LPLW\ 'RUVDO¿Q RULJLQ SRVWHULRU WR DQDO¿Q RIXQGHWHUPLQHGVH[±PP6/8)5-, 2 males, both PP6/IHPDOHV±PP6/DQGMXYHQLOHVRIXQ- EDVHVHFRQGSUR[LPDOUDGLDORIGRUVDO¿QEHWZHHQQHXUDO th st GHWHUPLQHGVH[±PP6/ F V FROOHFWHGZLWKWKHKROR- spines of 20 and 21 YHUWHEUDH¿UVWSUR[LPDOUDGLDORI W\SH±8)5-IHPDOHV±PP6/DQGMXYHQLOHV DQDO¿QEHWZHHQSOHXUDOULEVRIth and 15th vertebrae. RIXQGHWHUPLQHGVH[±PP6/,JDUDSp6DOJDGRDVWUHDP 'RUVDO¿QUD\V±DQDO¿QUD\V±FDXGDO¿QUD\V WULEXWDU\RIWKH&DXUpVULYHUƍƎ6ƍƎ:VDPHFRO- ±SHFWRUDO¿QUD\V±SHOYLF¿QUD\V lectors and date as holotype. Scales small, cycloid. Body and head entirely scaled, H[FHSW RQ MDZV %RG\ VTXDPDWLRQ H[WHQGLQJ RYHU DQ- Diagnosis. 'LVWLQJXLVKHG IURP DOO RWKHU VSHFLHV RI WKH WHULRURIFDXGDO¿QEDVHQRVFDOHVRQGRUVDODQG A. ornatus group E\ KDYLQJ D XQLTXH FRORXU SDWWHUQ LQ DQDO¿QEDVHV)URQWDOVTXDPDWLRQ)SDWWHUQHG(VFDOHV WKH FDXGDO ¿Q LQ PDOHV FRQVLVWLQJ RI GRUVDO SRUWLRQ RI not overlapping medially; scales arranged in regular cir- ¿QZKLWHZLWKGDUNUHGGLVKEURZQVSRWVFRQWUDVWLQJZLWK cular pattern around A-scale without exposed margins; WKHYHQWUDOSRUWLRQWKDWLVK\DOLQHZLWKGDUNJUH\GRWVDQG transverse row of scales anterior to H-scale. Longitudinal short lines (vs. never a similar colour pattern). Also dis- series of scales 28; transverse series of scales 7; scale tinguished from all species of the A. ornatus group, ex- rows around caudal peduncle 12. No contact organs on cept A. henschelae, by the absence of teeth on the second ÀDQNDQG¿QV pharyngobranchial bone (vs. presence); distinguished Cephalic neuromasts: supraorbital 3 + 3, parietal 1, ros- from A. henschelae by the absence of a longitudinal two tral 1, infraorbital 2 + 13 + 1, preorbital 2, otic 1, postotic RIUHGGRWVRQWKHYHQWUDOSDUWRIWKHÀDQNLQPDOHV YV 1, supratemporal 1, median opercular 1, ventral opercular presence). Also distinguished from all species of the SUHRSHUFXODUPDQGLEXODUODWHUDOPDQGLEXODU A. ornatus group, except A. ornatus, by having caudal 2, paramandibular 1. Lateral line interrupted, alter nating ¿QSRLQWHGLQODUJHUVSHFLPHQV YVHOOLSWLFDO DQGUHGGLVK VHWVRI±VFDOHVZLWKRQHQHXURPDVWDQGZLWKRXWQHXUR- EURZQVSRWVRQWKHGRUVDOSRUWLRQRIWKHFDXGDO¿Q also PDVWV7ZRQHXURPDVWVRQFDXGDO¿QEDVH distinguished from A. ornatus by the red pigmentation of %DVLK\DO VXEWULDQJXODU JUHDWHVW ZLGWK DERXW ventral portion of caudal peduncle not extending on cau- RIOHQJWKEDVLK\DOFDUWLODJHDERXWRIWRWDOOHQJWK GDO¿Q YVH[WHQGLQJRYHUZKROHYHQWUDOSRUWLRQRIWKH of ba sihyal. Six branchiostegal rays. Second pharyn- ¿Q DQGGRUVDOSRUWLRQRIFDXGDO¿QZKLWH YV\HOORZ JREUDQFKLDOWHHWK DEVHQW *LOOUDNHUV RQ ¿UVW EUDQFKLDO DUFK9RPHULQHWHHWK±'HUPRVSKHQRWLFSUHVHQW Description. 0RUSKRPHWULFGDWDDSSHDULQ7DEOH'RU Ventral process of posttemporal absent. Total vertebrae VDOSUR¿OHVOLJKWO\FRQYH[IURPVQRXWWRHQGRIFDXGDO¿Q ±

289 W.J.E.M. Costa et al.: Five new Anablepsoides from the Brazilian Amazon

Table 4. Morphometric data of Anablepsoides ottonii. EURZQZLWKSRVWHURYHQWUDOVLOYHUVSRW'RUVDO¿QK\D- OLQHZLWKWZRQDUURZGDUNJUH\WUDQVYHUVHVWULSHV$QDO holotype paratypes ¿Q K\DOLQH ZLWK WKUHH RU IRXU WUDQVYHUVH URZV RI GDUN male males (3) females (6) JUH\ GRWV &DXGDO ¿Q K\DOLQH ZLWK WUDQVYHUVH URZV RI Standard length (mm) 20.1 20.6 – 22.2 19.4 – 22.5 GDUNJUH\GRWVVRPHWLPHVGDUNJUH\EORWFKRQPLGGOHRI Percent of standard length FDXGDO¿QEDVH3HFWRUDO¿Q\HOORZLVKK\DOLQH3HOYLF¿Q Body depth 19.2 16.9 – 19.4 16.9 – 19.7 hyaline. Caudal peduncle depth 12.9 11.7 – 13.1 12.0 – 13.2 Pre-dorsal length 78.6 79.5 – 82.1 77.7 – 80.8 Distribution. .QRZQIURPWZRORFDOLWLHVLQWKH&DXUpV Pre-pelvic length 52.0 52.6 – 55.0 51.8 – 54.3 river drainage, a tributary of the Negro river, Amazonas Length of dorsal-fin base 6.7 6.5 – 8.6 6.7 – 8.1 river basin, Amazonas state, Brazil. Length of anal-fin base 16.3 15.9 – 17.8 14.1 – 16.2 Caudal-fin length 45.1 45.2 – 49.8 43.1 – 46.7 Etymology. Named after the ichthyologist Felipe Ottoni Pectoral-fin length 24.5 23.5 – 24.5 23.1 – 24.6 for his constant enthusiasm and friendship, and important Pelvic-fin length 11.9 10.9 – 13.6 10.1 – 12.5 participation during expeditions to northern Brazil. Head length 26.6 26.3 – 28.4 25.8 – 27.7 Percent of head length Head depth 58.7 52.9 – 55.1 51.8 – 56.9 Head width 79.3 75.4 – 77.3 77.8 – 80.5 Snout length 14.4 13.1 – 14.9 13.0 – 14.5 Anablepsoides gamae spec. nov. Lower jaw length 25.2 22.0 – 24.6 18.7 – 24.1 Eye diameter 35.7 30.0 – 33.0 30.8 – 33.5 )LJVF7DEOH

Holotype: 8)5- , male, 21.1 mm SL; Brazil: Estado do $PDSi 0XQLFtSLR GH /DUDQMDO GR -DUL VWUHDP WULEXWDU\ WR -DUL ULYHU EHWZHHQ /DUDQMDO GR -DUL DQG 6DQWR $QW{QLR ZDWHUIDOO Colouration. Males.)ODQNSDOHJUHHQZLWKREOLTXHURZV $PD]RQDVULYHUEDVLQ¶´6¶´:P.H.N. BRAGANÇA RIGDUNEURZQLVKUHGGRWVORQJLWXGLQDOURZRIUHGGLVK & E. HENSCHEL, 25 July 2012. brown dots on caudal peduncle, separated from reddish Paratypes: Brazil: Estado do Amazonas: Estado do Amapá: Mu ni- orange on postero-ventral part of caudal peduncle by FtSLRGH/DUDQMDOGR-DUL-DULULYHUGUDLQDJH$PD]RQDVULYHUEDVLQ EULJKW EOXH ]RQH VOLJKWO\ H[WHQGLQJ RQ FDXGDO¿Q EDVH 8)5-PDOHV±PP6/IHPDOHV±PP 'RUVXP OLJKW EURZQ ZLWK VHJPHQWHG ORQJLWXGLQDO GDUN SL; 8)5-PDOHV±PP6/IHPDOHPP6/ EURZQ VWULSH RQ ODWHUDO SRUWLRQ 2EOLTXH GDUN JUH\ EDU (c&s); 8)5-IHPDOHV±PP6/FROOHFWHGZLWK EHWZHHQ DUHD MXVW DERYH SHFWRUDO¿Q EDVH DQG PLGGOH WKHKRORW\SH±IEPA 2755, 1 male, 22.2 mm SL; Laranjal do Jari; C. GAMA, 27 Oct. 2007. of opercle. Venter light grey. Side of head light brown, opercular and suborbital regions with dense concentra- WLRQRIVPDOOGDUNJUH\WREODFNVSRWVDQGGDUNEOXHLUL- Diagnosis. 'LVWLQJXLVKHG from all other congeners, ex- GHVFHQFHRQYHQWUDOSRUWLRQRIRSHUFOHWUDQVYHUVHEODFN cept A. amananE\WKHFRORXUSDWWHUQRIWKHFDXGDO¿Q bar between orbits through chin. Ventral portion of head DQGDQDO¿QLQPDOHVFRQVLVWLQJRIGRUVDOSRUWLRQRIFDX- ZLWKGHQVHFRQFHQWUDWLRQRIVPDOOEODFNVSRWVDQWHULRUO\ GDO¿QRUDQJLVKUHGZLWKGRUVDOPDUJLQZKLWHDQGQDUURZ bordered by not pigmented area just posterior to chin bar. EODFNRXWHUERUGHU YVQHYHUDVLPLODUFRORXUSDWWHUQ ,ULVGDUNEURZQZLWKSRVWHURYHQWUDOVLOYHUVSRW'RUVDO PLGGOHSRUWLRQRIFDXGDO¿QEODFNZLWKEULJKWJUHHQLVK ¿QZKLWHZLWKWZRQDUURZGDUNEURZQWUDQVYHUVHVWULSHV blue stripe extending from caudal peduncle to posterior $QDO¿QK\DOLQHZLWKWKUHHWUDQVYHUVHURZVRIGDUNJUH\ SRUWLRQRI¿Q YVQHYHUDVLPLODUFRORXUSDWWHUQ DQDO¿Q GRWVDQGGDUNJUH\YHUPLFXODWHPDUNV&DXGDO¿QZKLWH yellow with transverse red stripes on the posterior por- ZLWKGDUNUHGGLVKEURZQVSRWVRQGRUVDOSRUWLRQK\DOLQH tion (vs. never a similar colour pattern) and by having a ZLWK GDUN JUH\ GRWV DQG VKRUW OLQHV RQ YHQWUDO SRUWLRQ continuous row of 6 neuromasts on the mandibular se- 3HFWRUDODQGSHOYLF¿QV\HOORZLVKK\DOLQH ULHV YVRUQHXURPDVWVRQDQDQWHULRUVHFWLRQDQGD Females.)ODQNSDOH\HOORZLVKJUH\ZLWKREOLTXHGDUN VLQJOHLVRODWHGSRVWHULRUQHXURPDVW 'LVWLQJXLVKHGIURP EURZQ EDUV RIWHQ YHQWUDOO\ WHUPLQDWLQJ LQ EODFN VSRW A. amanan and all other species of the A. ornatus group DQG QDUURZ ORQJLWXGLQDO GDUN EURZQLVK JUH\ VWULSHV E\KDYLQJDXQLTXHFRORXUSDWWHUQRIÀDQNLQDGXOWPDOHV 'RUVXPOLJKWEURZQZLWKVHJPHQWHGORQJLWXGLQDOGDUN LQZKLFKWKHUHDUHSDLUVRIREOLTXHURZVRIGDUNUHGGRWV EURZQ VWULSH RQ ODWHUDO SRUWLRQ 2EOLTXH EODFN EDU EH- the dots of each posterior pair row being connected to WZHHQ DUHD MXVW DERYH SHFWRUDO¿Q EDVH DQG PLGGOH RI IRUPGDUNUHGREOLTXHEDUVDQGHDFKGRWRIWKHDQWHULRU opercle. Venter light grey. Side of head light grey, oper- pair row being usually connected to the posterior adja- cular and suborbital regions with dense concentration of cent row (vs. never a similar colour pattern) and by a VPDOOGDUNJUH\WREODFNVSRWVWUDQVYHUVHEODFNEDUEH- XQLTXHFRORXUSDWWHUQRQWKHKHDGLQIHPDOHVFRQVLVWLQJ tween orbits through chin. Ventral portion of head with of a distinctive not pigmented area between a zone of ORQJLWXGLQDOURZRIEODFNVSRWVDQWHULRUO\ERUGHUHGE\ small brownish grey spots on the post-orbital region and QRWSLJPHQWHGDUHDMXVWSRVWHULRUWRFKLQEDU,ULVGDUN DQREOLTXHEODFNEDUFURVVLQJWKHPLGGOHRIRSHUFOH )LJ

290 VERTEBRATE ZOOLOGY — 63 (3) 2013

Fig. 6. Anablepsoides gamae8)5-KRORW\SHPDOHPP6/%UD]LO$PDSi/DUDQMDOGR-DUL

F YVVLGHRIKHDGSLJPHQWHGZLWKVPDOOGDUNJUH\WR %DVLK\DO VXEWULDQJXODU JUHDWHVW ZLGWK DERXW EODFNVSRWVRYHUODSSHGRYHURSHUFXODUSRVWRUELWDODQG RIOHQJWKEDVLK\DOFDUWLODJHDERXWRIWRWDOOHQJWK LQIUDRUELWDOUHJLRQ)LJVD±E $OVRGLVWLQJXLVKHGIURP of basihyal. Branchiostegal rays 6. Second pharyngob- A. amanan E\DORQJHUSUHGRUVDOGLVWDQFHLQPDOHV ± UDQFKLDOZLWKWHHWK*LOOUDNHUVRQ¿UVWEUDQFKLDODUFK 6/YV±6/ 9RPHULQH WHHWK ± 'HUPRVSKHQRWLF SUHVHQW Ventral process of posttemporal absent. Total vertebrae Description. Morphometric data appear in Table 5. ± 'RUVDO SUR¿OH VOLJKWO\ FRQYH[ IURP VQRXW WR HQG RI FDXGDO¿Q EDVH 9HQWUDO SUR¿OH DSSUR[LPDWHO\ VWUDLJKW Colouration. Males. )ODQN JUHHQLVK JROGHQ ZLWK VOLJKWO\FRQYH[RQSHOYLF¿QUHJLRQ%RG\VOHQGHUGH- REOLTXHURZSDLUVRIGDUNUHGGRWVGRWVRIHDFKSRVWHULRU pressed on head, gradually becoming compressed poste- SDLUURZFRQQHFWHGWRIRUPGDUNUHGREOLTXHEDUVHDFK ULRUO\*UHDWHVWERG\GHSWKDWYHUWLFDOWKURXJKSHOYLF¿Q dot of anterior pair row usually connected to posteriorly base. Jaws short, snout slightly pointed in lateral view. adjacent row; red dots concentrated on posterior portion 'RUVDODQGDQDO¿QVVPDOOH[WUHPLW\URXQGHGGRU- RIFDXGDOSHGXQFOH'RUVXPOLJKWEURZQZLWKVHJPHQWHG VDO VXEUHFWDQJXODU DQDO ¿Q VXEWULDQJXODU &DXGDO ¿Q ORQJLWXGLQDOGDUNEURZQVWULSHRQODWHUDOSRUWLRQ2EOLTXH RYDO VOHQGHU DQG ORQJ 3HFWRUDO ¿Q URXQGHG SRVWHULRU GDUNJUH\EDUEHWZHHQDUHDMXVWDERYHSHFWRUDO¿QEDVH PDUJLQUHDFKLQJYHUWLFDOWKURXJKSHOYLF¿QEDVHLQPDOH and middle of opercle. Venter light grey. Side of head WKURXJKXURJHQLWDOSDSLOODLQIHPDOHV3HOYLF¿QHOOLSWL- light grey, opercular and suborbital regions with dense FDOH[WUHPLW\UHDFKLQJDQDO¿QRULJLQLQPDOHVUHDFKLQJ FRQFHQWUDWLRQRIVPDOOGDUNJUH\VSRWVDQGEOXHLULGHV- XURJHQLWDOSDSLOODLQIHPDOHV3HOYLF¿QEDVHVPHGLDOO\ FHQFHRQYHQWUDOSRUWLRQRIRSHUFOHWUDQVYHUVHEODFNEDU LQFORVHSUR[LPLW\'RUVDO¿QEDVHSRVWHULRUWRDQDO¿Q between orbits through chin. Ventral portion of head with EDVHVHFRQGSUR[LPDOUDGLDORIGRUVDO¿QEHWZHHQQHXUDO GHQVHFRQFHQWUDWLRQRIVPDOOEODFNVSRWVDQWHULRUO\ERU- spines of 19th and 21stYHUWHEUDH¿UVWSUR[LPDOUDGLDORI dered by not pigmented area just posterior to chin bar. DQDO¿QEHWZHHQSOHXUDOULEVRIthDQGth vertebrae. ,ULVGDUNEURZQZLWKSRVWHURYHQWUDOVLOYHUVSRW'RUVDO 'RUVDO¿QUD\V±DQDO¿QUD\V±FDXGDO¿QUD\V ¿QZKLWHZLWKWZRGDUNUHGGLVKEURZQWUDQVYHUVHVWULSHV ±SHFWRUDO¿QUD\V±SHOYLF¿QUD\V $QDO¿QSDOH\HOORZEDVDOSRUWLRQOLJKWEOXHZLWKGDUN Scales small, cycloid. Body and head entirely scaled, red dots, posterior portion light blue with two or three H[FHSW RQ MDZV %RG\ VTXDPDWLRQ H[WHQGLQJ RYHU DQ- GDUNUHGWUDQVYHUVHVWULSHV'RUVDOSRUWLRQRIFDXGDO¿Q WHULRURIFDXGDO¿QEDVHQRVFDOHVRQGRUVDODQG RUDQJLVKUHGGRUVDOPDUJLQZKLWHZLWKQDUURZEODFNRXW- DQDO¿QEDVHV)URQWDOVTXDPDWLRQ)SDWWHUQHG(VFDOHV HUERUGHUPLGGOHSRUWLRQRIFDXGDO¿QEODFNZLWKEULJKW not overlapping medially; scales arranged in regular cir- greenish blue stripe extending from caudal peduncle to cular pattern around A-scale without exposed margins; SRVWHULRUIRXUWKRI¿QYHQWUDOSRUWLRQRIFDXGDO¿QGDUN transverse row of scales anterior to H-scale. Longitudinal UHGWRJUDGXDOO\EODFNSRVWHULRUO\3HFWRUDO¿QK\DOLQH series of scales 28; transverse series of scales 7; scale 3HOYLF¿Q\HOORZLVKK\DOLQH rows around caudal peduncle 12. No contact organs on Females.)ODQNOLJKWJUH\ZLWKREOLTXHGDUNJUH\EDUV ÀDQNDQG¿QV RIWHQ YHQWUDOO\ WHUPLQDWLQJ LQ EODFN VSRW DQG QDUURZ Cephalic neuromasts: supraorbital 3 + 3, parietal ORQJLWXGLQDO GDUN EURZQLVK JUH\ VWULSHV 'RUVXP OLJKW 1, rostral 1, infraorbital 2 + 12 + 1, preorbital 2, otic 1, JUH\ZLWKVHJPHQWHGORQJLWXGLQDOGDUNEURZQVWULSHRQ post-otic 1, supratemporal 1, median opercular 1, ven- ODWHUDOSRUWLRQ9HQWHUOLJKWJUH\2EOLTXHEODFNEDUIURP WUDORSHUFXODUSUHRSHUFXODUPDQGLEXODUODWHUDO DUHD MXVW DERYH SHFWRUDO¿Q EDVH WR LQIUDRUELWDO UHJLRQ PDQGLEXODU±SDUDPDQGLEXODU7ZRQHXURPDVWVRQ through middle of opercle. Side of head light grey, with FDXGDO¿QEDVH zone of small brownish grey spots on post-orbital region,

291 W.J.E.M. Costa et al.: Five new Anablepsoides from the Brazilian Amazon

Table 5. Morphometric data of Anablepsoides gamae. longitudinal rows of red dots interrupted and replaced E\ VKRUW URZV RI JROGHQ GRWV RQ WKH ÀDQN LQ PDOHV holotype paratypes PDUJLQVRIXQSDLUHGGDUNJUH\LQPDOHVDQGDKRUL]RQ- male males (4) females (2) tal row of red dots on the sub-basal region of the anal Standard length (mm) 21.1 17.5 – 22.2 18.3 – 21.8 ¿QLQPDOHVWKUHHFKDUDFWHUVWDWHVRIWKHFRORXUSDWWHUQ Percent of standard length not found in any other species of the group. In addition, Body depth 19.6 19.2 – 20.5 19.3 – 19.5 vomerine teeth are absent in A. jari, a condition not re- Caudal peduncle depth 12.2 12.3 – 13.3 11.4 – 11.6 ported previously for species of that group. Contrasting Pre-dorsal length 82.3 81.3 – 84.6 82.6 – 83.3 ZLWKWKRVHXQLTXHGLDJQRVWLFIHDWXUHVLWZDVQRWSRVVL- Pre-pelvic length 54.7 52.2 – 54.3 54.1 – 54.5 ble to identify any morphological trait suggesting close Length of dorsal-fin base 6.6 5.4 – 7.6 4.8 – 5.5 relationships between A. jari and any member of the A. Length of anal-fin base 15.9 14.9 – 17.8 13.5 – 14.9 urophthalmus group. However, a similar horizontal row Caudal-fin length 43.9 46.1 – 48.1 40.6 – 42.3 RIUHGGRWVRQWKHVXEEDVDOUHJLRQRIWKHDQDO¿QLQPDOHV Pectoral-fin length 24.9 23.7 – 25.8 23.2 – 23.6 is also found in A. beniensis (MYERS, 1927), a species Pelvic-fin length 12.9 11.8 – 14.3 10.4 – 11.4 geographically widespread in the Madeira river drain- Head length 26.9 27.1 – 29.4 26.6 – 26.9 age (COSTA, 2006), not belonging to the A. urophthal- Percent of head length mus group. Anablepsoides jari is readily distinguished Head depth 57.1 52.9 – 54.7 54.0 – 55.6 from A. beniensis by the former having more vertebrae Head width 80.4 73.9 – 78.1 80.7 – 83.5 ± YV ± PRUH FDXGDO¿Q UD\V ± YV Snout length 15.2 13.8 – 15.0 14.2 – 15.6 ± PRUHVFDOHVRQWKHORQJLWXGLQDOVHULHV ± Lower jaw length 23.5 21.0 – 22.3 20.3 – 21.2 YV± YRPHULQHWHHWKDEVHQW YVYRPHULQHWHHWK Eye diameter 33.2 31.8 – 35.0 31.8 – 33.1 DQGORZHUMDZQRWGLVWLQFWLYHO\SLJPHQWHG YVEODFN Anablepsoides roraima is distinguished from all RWKHU VSHFLHV RI WKH JHQXV E\ SRVVHVVLQJ ÀDQN JROGHQ VHSDUDWHGIURPREOLTXHRSHUFXODUEDUE\QRWSLJPHQWHG in males, with eight longitudinal rows of spots, among DUHDWUDQVYHUVHEODFNEDUEHWZHHQRUELWVWKURXJKFKLQ which the spots of the three dorsal-most rows are red- 9HQWUDOSRUWLRQRIKHDGZLWKJUHDWFRQFHQWUDWLRQRIEODFN dish brown and separated by small interspace, and the spots anteriorly bordered by not pigmented area just pos- VSRWV RI WKH RWKHU ¿YH URZV DUH UHG DQG LQ FRQWDFW WR WHULRU WR FKLQ EDU ,ULV GDUN EURZQ ZLWK SRVWHURYHQWUDO form long red stripes, a colour pattern not occurring in VLOYHUVSRW'RUVDO¿QK\DOLQHZLWKWZRQDUURZGDUNJUH\ RWKHUVSHFLHV7KHWZRZHOOGH¿QHGEULJKW\HOORZVWULSHV WUDQVYHUVHVWULSHV$QDO¿QK\DOLQHZLWKWKUHHWUDQVYHUVH RQ WKH GRUVDO DQG YHQWUDO PDUJLQV RI WKH FDXGDO ¿Q LQ URZV RI GDUN JUH\ GRWV &DXGDO ¿Q K\DOLQH ZLWK WUDQV- males of A. roraima suggest a close relationship with YHUVHURZVRIGDUNJUH\GRWV3DLUHG¿QVK\DOLQH A. deltaphilus (SEEGERS, 1983), A. lungi (BERKENKAMP, A. mazaruni (MYERS and A. stagnatus Distribution. Known only from the Jari river drainage, (EIGENMANN, 1909), all endemic from the Guianas and an lower Amazonas river basin, Amapá state, Brazil. adjacent region in Venezuela. The stripes are pale yellow or pale orange in A. urophthalmus (GÜNTHER, 1866), dif- Etymology. Named after Cecile Gama in recognition of fuse and inconspicuous in A. tocantinensis (COSTA, 2010) KHUSLRQHHULQJLFKWK\RORJLFDO¿HOGVWXGLHVLQWKH$PDSi and A. xinguensis (COSTA, 2010), and absent in A. jari. state, Brazilian Amazon. Among the species of the A. ornatus group herein described, A. ottonii is considered closely related to A. or- natusDVSHFLHVRFFXUULQJLQWKHÀRRGSODLQVRIWKHORZHU Negro river and adjacent rivers of central Amazon, and Discussion A. gamae to A. amanan, endemic to the Japurá-Solimões drainage. Anablepsoides ottonii and A. ornatus share a SRLQWHGFDXGDO¿QLQODUJHUVSHFLPHQV YVHOOLSWLFDO DQG the presence of reddish brown spots on the dorsal portion Anablepsoides jari and A. roraima DUHWKH¿UVWUHFRUGV RIWKHFDXGDO¿QLQPDOHV YVUHGGLVKEURZQPDUNVDE- of members of the A. urophthalmus group for tributaries VHQW EHVLGHVWKHFDXGDO¿QXVXDOO\EHLQJORQJHULQPDOHV of the Amazonas river basin draining the Guiana Shield. ±6/YV±6/ EXWERWKVSHFLHV Phylogenetic relationships of these species with other KDYHGLVWLQFWFRORXUSDWWHUQVRIWKHFDXGDO¿QLQPDOHV taxa included in that group are still uncertain, since both (see diagnosis above). On the other hand, A. gamae and GHHSPRUSKRORJLFDOGDWDDQG'1$VHTXHQFHVDUHVWLOOXQ- A. amanan VKDUHDQLGHQWLFDODQGXQLTXHFRORXUSDWWHUQ available for most taxa included in that group. However, RIFDXGDO¿QLQPDOHVPRUHVWULNLQJLQWKHGRUVDOSRUWLRQ PRUSKRORJLFDOIHDWXUHVH[KLELWHGE\WKHQHZWD[DPDNH RIWKH¿QZKLFKLVRUDQJLVKUHGZLWKGRUVDOPDUJLQZKLWH possible to unambiguously distinguish them from all DQGQDUURZEODFNRXWHUERUGHUDFRORXUSDWWHUQQRWIRXQG other congeners. HOVHZKHUHDPRQJDSORFKHLORLGNLOOL¿VKHVAnablepsoides Anablepsoides jari is easily distinguished from all gamae and A. amanan can be distinguished by the former other species of the A. urophthalmus group by having KDYLQJLQPDOHVSDLUVRIREOLTXHURZVRIGDUNUHGGRWV

292 VERTEBRATE ZOOLOGY — 63 (3) 2013

the dots of each posterior pair row being connected to VKLSVWD[RQRPLFUHYLVLRQDQGELRJHRJUDSK\±,FKWK\RORJLFDO IRUPGDUNUHGREOLTXHEDUVDQGHDFKGRWRIWKHDQWHULRU Exploration of Freshwaters, 12± pair row being usually connected to the posterior adja- COSTA, W.J.E.M. 5HODWLRQVKLSVDQGWD[RQRP\RIWKHNLOOL FHQWURZ YVVLQJOHREOLTXHURZVDUUDQJHGDVFKHYURQVLQ ¿VKJHQXVRivulus (Cyprinodontiformes: Aplocheiloidei: Ri vu- A. amanan), the presence in females of a not pigmented lidae) from the Brazilian Amazonas river basin, with notes on area between a zone of small brownish grey spots on the historical ecology. ±$TXD±-RXUQDORI,FKWK\RORJ\DQG$TXDWLF SRVWRUELWDOUHJLRQDQGDQREOLTXHEODFNEDUFURVVLQJWKH Biology, 11± middle of opercle (vs. not pigmented area absent), and a COSTA, W.J.E.M. (2011): Phylogenetic position and taxonomic sta- ORQJHUSUHGRUVDOGLVWDQFHLQPDOHV ±6/YV tus of Anablepsoides, Atlantirivulus, Cynodonichthys, Lai mo- ±6/ )LQDOO\A. henschelae possibly is the se mion and Melanorivulus (Cyprinodontiformes: Rivulidae). ± sister group to the remaining species of the group, since Ichthyological Exploration of Freshwaters, 22± it does not share an apomorphic asymmetrically coloured GARMAN6 7KHF\SULQRGRQWV±0HPRLUVRIWKH0XVHXP FDXGDO¿QLQPDOHVLQZKLFKWKHGRUVDODQGYHQWUDOSRU- of Comparative Zoology, 19± WLRQRIWKH¿QH[KLELWGLIIHUHQWFRORXUSDWWHUQV GÜNTHER$ &DWDORJXH RI ¿VKHV LQ WKH %ULWLVK 0XVHXP Volume 6. Catalogue of the Physostomi, containing the fam- ilies Salmonidae, Percopsidae, Galaxidae, Mormyridae, Gym- QDUFKLGDH(VRFLGDH8PEULGDH6FRPEUHVRFLGDH&\SULQRGRQ Acknowledgements tidae, in the collection of the British Museum. London: British Museum, xv + 368p. HENN, A.W. ( 2QYDULRXV6RXWK$PHULFDQSRHFLOLLG¿VKHV± 7KDQNVDUHGXHWR)2WWRQLDQG(+HQVFKHOIRUKHOSLQWKH¿HOG Annals of the Carnegie Museum, 10±SO WR/53\'DQLHODQG&*DPDIRUWKHORDQRIPDWHULDODQGWR& HOEDEMAN, J.J. (1958): The frontal scalation pattern in some groups 0HOORIRUWHFKQLFDODVVLVWDQFH7KLVVWXG\ZDVVXSSRUWHGE\&13T RIWRRWKFDUSV 3LVFHV&\SULQRGRQWLIRUPHV ±%XOOHWLQRI$TXD &RQVHOKR1DFLRQDOGH'HVHQYROYLPHQWR&LHQWt¿FRH7HFQROyJLFR tic Biology, 1± 0LQLVWpULRGH&LrQFLDH7HFQRORJLD DQG&$3(6 &RRUGHQDomRGH MURPHY, W.J.; THOMERSON, J.E. & COLLIER, G.E. (1999): Phylo- $SHUIHLoRDPHQWRGH3HVVRDOGH1tYHO6XSHULRU &ROOHFWLRQVZHUH JHQ\RIWKHQHRWURSLFDONLOOL¿VKIDPLO\5LYXOLGDH &\SULQRGRQ made with permits provided by ICMBio (Instituto Chico Mendes WLIRUPHV$SORFKHLORLGHL LQIHUUHG IURP PLWRFKRQGULDO '1$ GH&RQVHUYDomRGD%LRGLYHUVLGDGH VHTXHQFHV±0ROHFXODUDQG3K\ORJHQHWLF(YROXWLRQ13± 301. STEINDACHNER, F. (1863): Beiträge zur Kenntniss der Sciaenoiden %UDVLOLHQVXQGGHU&\SULQRGRQWHQ0HMLFRV±6LW]XQJVEHULFKWH References GHU.DLVHUOLFKHQ$NDGHPLHGHU:LVVHVFKDIWHQ0DWKHPDWLVFK Naturwissenschaftliche Klasse, 48±SO STEINDACHNER,) ,FKWK\RORJLVFKH %HLWUlJH ± 6LW]XQJV COSTA, W.J.E.M. 3HDUO NLOOL¿VKHV ± WKH &\QROHELDWLQDH EHULFKWHGHU.DLVHUOLFKHQ$NDGHPLHGHU:LVVHVFKDIWHQ0DWKH V\VWHPDWLFV DQG ELRJHRJUDSK\ RI WKH QHRWURSLFDO DQQXDO ¿VK matisch-Naturwissenschaftliche Klasse, 74±SO subfamily. Neptune City: TFH, 128p. TAYLOR, W.R. & VAN DYKE, G.C. (1985): Revised procedures for COSTA, W.J.E.M. (2001):7KHQHRWURSLFDODQQXDO¿VKJHQXVCy no- VWDLQLQJ DQG FOHDULQJ VPDOO ¿VKHV DQG RWKHU YHUWHEUDWHV IRU lebias (Cyprinodontiformes: Rivulidae): phylogenetic relation- ERQHDQGFDUWLODJHVWXG\±&\ELXP9±

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