Two new species of seasonal killifishes of the

Two new species of seasonal killifishes of the

Nothobranchius melanospilus species complex from

the East Africa biodiversity hotspot (Cyprinodontiformes:

Aplocheilidae*)

Wilson J. E. M. Costa

Laboratory of Systematics and Evolution of Teleost Fishes, Institute of Biology, Federal University of Rio de Janeiro, Caixa Postal 68049, CEP 21944-970, Rio de Janeiro, Brasil; wcosta@acd.ufrj.br

Submitted February 16, 2018.

Accepted October 30, 2018.

Published online at www.senckenberg.de/vertebrate-zoology on February 15, 2019.

Published in print on February 27, 2019.

Editor in charge: Uwe Fritz

Abstract

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Key words

.HQ\D0NDWD5LYHU5DPLVL5LYHU7DQ]DQLD7HQGLJR6ZDPS6\VWHPDWLFV

Introduction

7KH $IULFDQ JHQXV Nothobranchius Peters, 1868 is DPRQJ WKH PRVW GLYHUVL¿HG JHQHUD RI WKH NLOOL¿VK VXE RUGHU$SORFKHLORLGHLH[KLELWLQJLQWHUHVWLQJWURSKLFVSH FLDOL]DWLRQVC^OSTA, 2011, 2018$OOWKHDERXWVSHFLHV LQFOXGHG LQ WKLV JHQXV LQKDELW WHPSRUDU\ SRROV IRUPHG GXULQJUDLQ\VHDVRQVWKDWDUHDOWHUQDWHGE\LQWHQVLYHO\GU\

SHULRGVZKHQSRROVGLVDSSHDUDQGUHVLVWDQWHJJVVXUYLYH ZLWKLQ WKH ERWWRP VXEVWUDWH HJJ^UBB, 1969; B^AILEY,

1972; W^OURMS,WVJUHDWHVWGLYHUVLW\LVIRXQGLQ

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M^YERS et alWRZKHUHDERXWRQHKDOIRIDOOVSH

FLHVRINothobranchius DUHHQGHPLFHJC^OSTA, 2008, 2017a).

Nothobranchius melanospilus 3IHIIHU KDV EHHQ FRQVLGHUHG D JHRJUDSKLFDOO\ ZLGHVSUHDG VSHFLHV RFFXUULQJLQWKHFRDVWDOIRUHVWVRIHDVWHUQ7DQ]DQLDDQG VRXWKHDVWHUQ.HQ\DFRPPRQO\GLDJQRVHGE\WKHSUHV HQFH RI EODFN GRWV RQ WKH ÀDQN LQ IHPDOHV W^ILDEKAMP et al +RZHYHU WKH VWXG\ RI DSORFKHLOLG FROOHF WLRQV GHSRVLWHG LQ 0XVpH 5R\DO GH O¶$IULTXH &HQWUDO 7HUYXUHQ05$&UHYHDOHGWKDWN. melanospilus sense W^ILDEKAMP et alLVDVSHFLHVFRPSOH[5HFHQWO\

$SORFKHLOLGDHLVKHUHDGRSWHGDVDVLQJOH$SORFKHLORLGHLIDPLO\IROORZLQJC^OSTA

N. melanospilusZDVUHGHVFULEHGDQGJHRJUDSKLFDOO\UH VWULFWHGWR=DQ]LEDU,VODQGDQGDGMDFHQWFRQWLQHQWDOULYHU EDVLQVRI7DQ]DQLDCOSTAE7KHSRSXODWLRQVRF FXUULQJLQWKHQRUWKHUQDQGQRUWKZHVWHUQSDUWRIWKHGLV WULEXWLRQRIWKLVFRPSOH[FRUUHVSRQGWRWZRQHZVSHFLHV WKDWDUHKHUHLQGHVFULEHG

Material and methods

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Nothobranchius kwalensis, spec. nov.

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Diagnosis. Nothobranchius kwalensis LV GLVWLQJXLVKHG IURP DOO RWKHU FRQJHQHUV RI WKHN. melanospilus FRP SOH[ E\ WKH FRPELQDWLRQ RI WKH IROORZLQJ PRUSKRORJL FDOFKDUDFWHUVWDWHVORZHUMDZQRWDQWHULRUO\SURMHFWLQJ EH\RQG XSSHU MDZ vs DQWHULRUO\ SURMHFWLQJ LQN. prog- nathusGRUVDO¿QRULJLQDQWHULRUWRDQDO¿QRULJLQLQIH PDOHV vs SRVWHULRU LQ ERWK VH[HV LQN. melanospilus);

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17 – 21 in N. melanospilus); in females, few rounded

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Fig. 2. *HRJUDSKLFDOGLVWULEXWLRQRINothobranchius kwalensisEOXHFLUFOHN. prognathusJUHHQOR]HQJHVDQGN. melanospilusUHGVTXDUHV

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dense vivid red).

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Holotype: 05$& 3 PDOH PP 6/ 7DQ]DQLD 0RURJRURGLVWULFWVZDPSSRRODERXWNPIURP.LPDPEDLQWKHWR .LORVDƒ¶´6ƒ¶´(DERXWPDVO5+W^ILDEKAMP, -XQ

Paratypes: 05$& 3 PDOHV ± PP 6/

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Diagnosis. Nothobranchius prognathus differs from all RWKHU VSHFLHV RI WKHN. melanospilus FRPSOH[ LQ SRV VHVVLQJ ORZHU MDZ DQWHULRUO\ SURMHFWLQJ EH\RQG XSSHU MDZ vs QRW SURMHFWLQJ SHOYLF ¿QV PHGLDOO\ XQLWHG LQ PDOHVvsEDVHVMXVWLQFRQWDFWEXWQRWXQLWHGDQGSUHV HQFHRIWKUHHORQJLWXGLQDOVHULHVRIVFDOHVRQIURQWDOUH JLRQvs. WZR$OVRGLVWLQJXLVKHGIURPDOORWKHUVSHFLHV RI WKHN. melanospilus FRPSOH[ E\ WKH FRPELQDWLRQ RI /HQJWKRIGRUVDO¿QEDVH ± 22.9

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+HDGOHQJWK ±

Percent of head length

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/RZHUMDZOHQJWK 29.8 26.7 – 29.9 (\HGLDPHWHU ± 27.1

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in N. melanospilus DQG ± DQG 6/ UH

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Fig. 3. Nothobranchius prognathusA, 05$& 3 KRORW\SH PDOH PP 6/B, 05$& 3 SDUDW\SH PDOH PP6/C, 05$&3SDUDW\SHIHPDOHPP6/7DQ]DQLD0RURJRUR.LPDPED

A

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Etymology. )URPWKH/DWLQprognathus SURJQDWKRXV UHIHUULQJWRWKHGLVWLQFWLYHORZHUMDZRIWKHQHZVSHFLHV WKDWLVDQWHULRUO\SURMHFWHGVOLJKWO\EH\RQGWKHXSSHUMDZ

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Distribution. 7HQGLJR6ZDPSDUHD0NDWD5LYHUÀRRG SODLQVXSSHU:DPL5LYHUEDVLQDOWLWXGHDERXW±

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Remarks. 7KH FRORXUDWLRQ LQ OLIH RIN. prognathus is GRFXPHQWHG LQ WKH DTXDULXP ¿VK OLWHUDWXUH LQ SKRWR JUDSKVWDNHQIURPOLYHLQGLYLGXDOVFROOHFWHGLQWKHW\SH ORFDOLW\UHJLRQHJ SEEGERS, 1997).

Discussion

WILDEKAMP et alSURYLGHGDVKRUWUHGHVFULSWLRQRI N. melanospilusZKLFKZDVFRQVLGHUHGWRRFFXULQDYDVW UHJLRQRIVRXWKHDVWHUQ.HQ\DDQGHDVWHUQ7DQ]DQLD7KH GLVWULEXWLRQ DUHD DWWULEXWHG WRN. melanospilus overlaps WKH JHRJUDSKLFDO UDQJH RI DOO RWKHU FRQJHQHUV IRXQG LQ WKH ($%+ ZKLFK DUH DOZD\V HQGHPLF RI VPDOOHU DUHDV DVRFFXUULQJZLWKPRVWVHDVRQDONLOOL¿VKHVHJCOSTA, 2QWKHRWKHUKDQGWKHFRORXUSDWWHUQIRXQGLQDOO /HQJWKRIGRUVDO¿QEDVH 28.9 27.9 – 29.9±

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A

B

Fig. 4. Nothobranchius melanospilusA, 8)5-PDOHPP6/B, 8)5-IHPDOHPP6/7DQ]DQLD3ZDQL0EH]L 5LYHUÀRRGSODLQV

EXWC^OSTA E SURYLGHG D KLVWRULFDO UHYLHZ RI WKH WD[RQRP\ RIN. melanospilus VKRZLQJ WKDW WKH

=DQ]LEDU LVODQG VKRXOG EH FRQVLGHUHG DV LWV W\SH ORFDO LW\,QIDFWP^FEFFEREDVHGLWVRULJLQDOGHVFULSWLRQ of N. melanospilus RQ IRXU IHPDOH VSHFLPHQV FROOHFWHG LQ /RQJR%D\ SRVVLEO\ SUHVHQW ,ORQJD E\P^ASHA and S^TUHLMANNLQDQGVSHFLPHQVUHSRUWHGLQP^LAYFAIR

& G^ÜNTHERFROOHFWHGLQ=DQ]LEDUC^OSTA E 7KH RQO\ LOOXVWUDWLRQ RIN. melanospilus ap pearing in P^FEFFER ZDV D UHSURGXFWLRQ RI D ¿J XUHRIDIHPDOHVSHFLPHQIURP=DQ]LEDU¿UVWLOOXVWUDWHG in P^LAYFAIR & G^ÜNTHER+RZHYHUQRLQGLYLGXDO W\SH VSHFLPHQ ZDV WKHQ GHVLJQDWHGB^OULENGER DWWULEXWHG W\SH VWDWXV IRU WKRVH VSHFLPHQV FROOHFWHG LQ =DQ]LEDU XVLQJ WKH VDPH LOOXVWUDWLRQ SXEOLVKHG LQ P^LAYFAIR & G^ÜNTHER DQGP^FEFFER RI D

IHPDOH IURP =DQ]LEDU %DVHG RQ UHFRPPHQGDWLRQ % RI WKH ,QWHUQDWLRQDO &RGH RI =RRORJLFDO 1RPHQFODWXUH UHIHUULQJWROHFWRW\SHVHOHFWLRQLHSUHIHUHQFHWR D V\QW\SH RI ZKLFK DQ LOOXVWUDWLRQ KDV EHHQ SXEOLVKHG C^OSTAEGHVLJQDWHGWKDWVSHFLPHQIURP=DQ]LEDU illustrated in P^LAYFAIR & G^ÜNTHERP^FEFFER

Fig. 5. 'LDJUDPPDWLFUHSUHVHQWDWLRQRIWKHODWHURVHQVRU\V\VWHPRQWKHGRUVDOVXUIDFHRIWKHKHDGLQA, Nothobranchius prognathus sp. n., 05$&3PDOHSDUDW\SHPP6/B, Nothobranchius melanospilus, 05$&3PDOHPP6/DUQ±

DQWHULRUURVWUDOQHXURPDVWDVDVV±DQWHULRUVHFWLRQRIWKHDQWHULRUVXSUDRUELWDOVHULHVSUQ±SRVWHULRUURVWUDOQHXURPDVWSVDVV±SRVWHULRU VHFWLRQRIWKHDQWHULRUVXSUDRUELWDOVHULHVSVV±SRVWHULRUVXSUDRUELWDOVHULHV

Fig. 6. 'LDJUDPPDWLFUHSUHVHQWDWLRQRIWKHSHOYLF¿QVYHQWUDOYLHZ RIA, Nothobranchius prognathus sp. n., 05$&3, PDOH SDUDW\SH PP 6/B, Nothobranchius melanospilus, 05$&3PDOHPP6/

Fig. 7. +HDG ODWHUDO YLHZ RIA KRORW\SH RINothobranchius kwalensis, 05$& 3 PDOH PP 6/B KROR W\SH RINotho branchius prognathous, 05$& 3, male, PP6/

and BOULENGERDVWKHOHFWRW\SHRIN. melanospi- lus5HFHQWWD[RQRPLFDODQDO\VHVCOSTA,EWKHSUH VHQW VWXG\ KDYH FRQVLVWHQWO\ LQGLFDWHG WKDWN. melano- spilus LVUHVWULFWHGWR=DQ]LEDUDQGDGMDFHQWFRQWLQHQWDO ULYHUEDVLQVRI7DQ]DQLDVRXWKRIWKH:DPL5LYHU)LJ DVZHOODVVSHFLPHQVFROOHFWHGLQWKHXSSHU:DPL5LYHU EDVLQLQFOXGLQJDSRSXODWLRQIRXQGLQWKHURDGEHWZHHQ ,ORQJD DQG .LORVD DUH KHUH LGHQWL¿HG DV EHORQJLQJ WR N. prognathus, not N. melanospilus.

Key for identification of species of the N. melanospilus complex

1 /RZHU MDZ QRW DQWHULRUO\ SURMHFWLQJ EH\RQG XS SHUMDZ)LJ$SHOYLF¿QVQHYHUPHGLDOO\XQLWHG )LJ$WZRORQJLWXGLQDOVHULHVRIVFDOHVRQIURQWDO UHJLRQ)LJ% ... 2 1’ /RZHU MDZ DQWHULRUO\ SURMHFWLQJ EH\RQG XSSHU

MDZ )LJ% SHOYLF ¿QV PHGLDOO\ XQLWHG LQ PDOHV )LJ%WKUHHORQJLWXGLQDOVHULHVRIVFDOHVRQIURQ WDOUHJLRQ)LJ$ ... N. prognathus 2 'RUVDO¿QRULJLQDQWHULRUWRDQDO¿QRULJLQLQIHPDOHV

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... N. melanospilus

Acknowledgements

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W^ILDEKAMP, R+S^HIDLOVSKIY, K. M. & W^ATTERS,%5 6\VWHPDWLFV RI WKHNothobranchius melanospilus VSHFLHV JURXS &\SULQRGRQWLIRUPHV 1RWKREUDQFKLLGDH ZLWK GHVFULS

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