Upeneus)DPLO\0XOOLGDHZLWKGHVFULSWLRQVRI IRXUQHZVSHFLHV
Franz Uiblein1 and Phillip C. Heemstra2
1Institute of Marine Research, Nordnesgaten 33, 5817 Bergen, Norway; e-mail: franz@imr.no
1 & 2South African Institute of Aquatic Biodiversity,
Private Bag 1015, Grahamstown, South Africa;
e-mail: p.heemstra@saiab.ac.za
Received 17 June 2009; accepted 30 September 2009
ABSTRACT.7KH WD[RQRP\ RI WKH JRDWÀVK VSHFLHV RI WKH JHQXVUpeneus from the Western Indian Ocean is reviewed. Sixteen species are recognized: U. davidaromi, U. doriae, U. guttatus, U. indicus sp. nov., U.
margarethaesp. nov., U. mascareinsis, U. moluccensis, U. oligospilus, U. pori, U. suahelicus sp. nov., U. sulphureus, U. sundaicus, U. supravittatus sp. nov., U. taeniopterus,U. tragula, and U. vittatus. Upeneus oligospilus, which had been synonymized with U. tragula, is resurrected as a valid species, and U. taeniopterus is shown to be a senior synonym of U. arge. A neotype is designated for U. vittatus, the type species of the genus, and described.
Several new occurrences of species in distinct areas of the Western Indian Ocean are documented. Four VSHFLHVFRPSOH[HVDUHGLVWLQJXLVKHGEDVHGRQQXPEHUVRIGRUVDOÀQVSLQHVJLOOUDNHUVDQGSHFWRUDOÀQUD\V DQGFDXGDOÀQFRORXU7KHVHFRPSOH[HVLQFOXGHRIWKHWRWDOYDOLGUpeneus species. Based on examination RIVSHFLHVIURPWKH,QGR3DFLÀFWKH$WODQWLFDQGWKH0HGLWHUUDQHDQZLWKLQJURXSFRPSDULVRQVDUHPDGH XVLQJWKHEHVWGLVWLQJXLVKLQJFKDUDFWHUV)UHVKFRORXUDWLRQRIWKHKHDGERG\DQGÀQVLVLPSRUWDQWIRUVSHFLHV GLDJQRVLV7KHQXPEHUDQGWKHFRQÀJXUDWLRQRIEDUVRQWKHFDXGDOÀQDUHDOVRXVHIXOWRGLVWLQJXLVKVHYHUDO VSHFLHV'DUNEDUVRIWHQSHUVLVWLQSUHVHUYHGÀVKDQGFDQEHXVHGWRLGHQWLI\ERWKIUHVKDQGSUHVHUYHGUpeneus specimens of several species. Morphometric characters are shown to clearly distinguish many species and provide valuable information on differentiation in body shape among populations and life-history stages.
Implications for further research priorities on the taxonomy of Upeneus are discussed.
KEYWORDS: Morphology, colour patterns, new species, Mullidae, Upeneus, Western Indian Ocean
INTRODUCTION
The Western Indian Ocean region extends from South Africa to the Northern Red Sea, the Persian Gulf, Sri /DQNDDQGDOLQHVRXWKIURPWKHWLSRI,QGLD7KHÀVK fauna of this large area that comprises many different coastal, island, and deep-water habitats is not well NQRZQ2QO\DIHZSDUWVRIWKLVDUHDKDYHEHHQVWXGLHG and a comprehensive inventory of the entire region LV ODFNLQJ 3UHSDUDWLRQ RI D ERRN RQ WKH FRDVWDO ÀVKHV of the Western Indian Ocean has motivated detailed WD[RQRPLFVWXGLHVRIWKHÀVKHVRIWKLVDUHD2QHRIWKH JURXSVXUJHQWO\UHTXLULQJUHYLVLRQLVWKHJRDWÀVKJHQXV Upeneus.
The genus Upeneus Cuvier 1829 was revised by Lachner (1954), who recognized 10 species.
Subsequently an additional seven species were described: U. mascareinsis )RXUPDQRLU *Xp]p from Réunion Island, U. crosnieri Fourmanoir & Guézé IURP0DGDJDVFDUU. pori Ben-Tuvia & Golani 1989
from the Red Sea, U. francisi Randall & Guézé 1992 from 1RUIRON ,VODQG 1HZ =HDODQGU. davidaromi Golani 2001 from the Red Sea, U. australiae .LP 1DND\D 2002 from Queensland, NE Australia, and U. mouthami 5DQGDOO .XOELFNLIURPWKH&KHVWHUÀHOG%DQNLQ the Coral Sea. One species, Upeneus crosnieri, was later synonymized with U. guttatus 'D\ %DXFKRW HW al. 1985). Another species, U. oligospilus Lachner 1954, described from the Persian Gulf, was assumed to have only subspecies status by Randall (1995) and was more recently synonymized with U. tragula by Randall &
.XOELFNL
The only previous review of the Western Indian OceanUpeneusVSHFLHVGDWHVEDFNFD\HDUV.XPDUDQ
& Randall 1984) and lists nine species: Upeneus asymmetricus Lachner 1954, Upeneus bensasi7HPPLQFN
& Schlegel 1843), Upeneus luzonius Jordan & Seale 1907, Upeneus moluccensis%OHHNHUUpeneus sulphureus Cuvier 1829, Upeneus sundaicus%OHHNHUUpeneus taeniopterus Cuvier 1829, Upeneus tragula Richardson
DQGUpeneus vittatus )RUVVNnO 2I WKHVH Randall et al. (1993) found that U. bensasi was a junior synonym of U. japonicus (Houttuyn 1782). In a review RIWKHJRDWÀVKHVRIWKHZHVWHUQ&HQWUDO3DFLÀF5DQGDOO (2001) reported that U. arge Jordan & Evermann 1903 occurred at several Western Indian Ocean localities. The need for a review of the Western Indian Ocean species became evident when we participated in a cruise of the Dr. Fridtjof Nansen in the southWestern Indian Ocean from September to November 2007. Upeneus species WKDW ZHUH FROOHFWHG FRXOG QRW EH LGHQWLÀHG XVLQJ WKH DYDLODEOHOLWHUDWXUHRQWKHFRDVWDOÀVKHVRIWKLVDUHD
A particular problem in the taxonomy of the Mullidae is the dearth of diagnostic meristic characters.
One of the few useful meristic characters is the number
RIGRUVDOÀQVSLQHVZKLFKUHTXLUHVFDUHIXOH[DPLQDWLRQ LQRUGHUWRGHWHFWWKHPLQXWHUHFXPEHQWÀUVWVSLQHLQ the eight-spine species group that distinguishes it from WKH VHYHQVSLQH JURXS /DFKQHU .LP 1DND\D 2002). Another important character is the number of SHFWRUDOÀQ UD\V ZKLFK PD\ GLIIHU DPRQJ VSHFLHV E\
only one ray. A third useful character is the number RI JLOO UDNHUV 7KHVH FDQ EH VHSDUDWHG LQWR VHYHUDO sub-characters according to position on the lower or upper limb and whether they are rudimentary or well-developed. The number of lateral-line scales is often used, but this is a less reliable character, as scales are frequently lost during collecting, transport or preservation. Also, in many species the scales extend IURPWKHERG\RQWRWKHEDVHRIWKHFDXGDOÀQDQGKHQFH Table 1. Abbreviations and description of characters.
LWPD\EHGLIÀFXOWWRFRXQWWKHH[DFWQXPEHURIVFDOHV There has consequently been considerable variation in VFDOHFRXQWV7KRPDV
In this account we examined 252 specimens of 24 species from the Western Indian Ocean from recent ÀHOG FROOHFWLRQV DQG VHYHUDO PXVHXPV DV ZHOO DV comparative material from the Eastern Indian Ocean, WKH3DFLÀFWKH0HGLWHUUDQHDQDQGWKH$WODQWLF2QHRI WKHJRDOVZDVWRSUHSDUHDNH\WRWKH:HVWHUQ,QGLDQ Ocean species that could be used for both fresh and SUHVHUYHGÀVK6SHFLDOHPSKDVLVLVSODFHGRQWKHXVHRI UHODWLYHO\ TXLFNO\ LGHQWLÀDEOH PHULVWLF PRUSKRPHWULF and colour characters. Forty morphometric characters were recorded to compare body shape variation among individuals and species. A further goal of this study was to designate a neotype for Upeneus vittatus, as no original type material exists (Randall 1974 and pers.
comm.).
MATERIALS AND METHODS
Abbreviations and descriptions of 51 characters are provided in Table 1. An overview of all Upeneus species currently recognized as valid is given in Table 2. Morphometric characters were measured with an electronic caliper and expressed as % SL. For comparison with earlier studies—and in order to facilitate the DSSOLFDWLRQ LQ WKH ÀHOG³WKH GLDJQRVWLFDOO\ PRVW important morphometric characters are provided as UDWLRVRIWKH6/LQWKHNH\DQGLQ7DEOHF0RUSKRPHWULF UDWLRVOHVVWKDQDUHJLYHQWRWZRVLJQLÀFDQWGLJLWV
Only meristic characters that vary among species are referred to in the diagnoses and comparisons:
WKH QXPEHU RI GRUVDOÀQ VSLQHV SHFWRUDOÀQ UD\V UXGLPHQWDU\ DQG GHYHORSHG JLOO UDNHUV RQ ORZHU DQG upper limb, and lateral-line scales. Rare counts are given LQSDUHQWKHVHV,QRUGHUWRVHHWKHÀUVWPLQXWHGRUVDO ÀQVSLQHLQHLJKWVSLQHVSHFLHVDVWHUHRPLFURVFRSHZDV used. Sometimes it was also necessary to move the VFDOHV DW WKH EDVH RI WKH VHFRQG VSLQH WR VHH WKH ÀUVW VSLQH*LOOUDNHUVZHUHLGHQWLÀHGDVUXGLPHQWDU\LIWKHLU OHQJWKZDVOHVVWKDQWKHLUZLGWK7KHJLOOUDNHULQWKH DQJOH EHWZHHQ WKH XSSHU DQG ORZHU OLPEV RI WKH ÀUVW gill arch was included in the count for the lower limb.
/DWHUDOOLQHVFDOHFRXQWVDUHGLIÀFXOWRQVSHFLPHQVZLWK missing scales and do not include scales on the caudal ÀQ
0HDVXUHPHQWV VKRZLQJ KLJK RYHUDOO LQWUDVSHFLÀF YDULDWLRQ HJ ÀQ GLVWDQFHV IURP WKH VQRXW ZHUH QRW included in the diagnoses and only rarely in species comparisons. Body depth measurements were only considered when there was consistent co-variation with other closely correlated measurements.
&RORXU FKDUDFWHUV WKDW IDGH LQ SUHVHUYHG ÀVK particularly yellowish, pale brown or reddish patterns, were only included as supplementary information along with comments on their restricted applicability.
%DQGV RQ WKH GRUVDO ÀQV DUH RIWHQ GLIÀFXOW WR GHWHFW RQ SUHVHUYHG ÀVK DQG ZHUH QRW FRQVLGHUHG LQ WKH
NH\ZLWKWKHH[FHSWLRQRIWKHEODFNÀUVWGRUVDOÀQWLS which is usually retained. Because the latter occurs LQGHSHQGHQWO\ RI RWKHU PDUNLQJV LW ZDV WUHDWHG separately in all cases and was not included in counts RIGRUVDOÀQEDQGV&RXQWVRIFDXGDOÀQEDUVLQFOXGH all bars from the base of each lobe to the lobe tip. The FRXQWRIEDUVRQWKHXSSHUFDXGDOÀQOREHDOVRLQFOXGHV the bar close above or behind the end of lateral line.
Colour photographs for each species were selected from the Western Indian Ocean material that we examined in the current study, or material from the DUHD WKDW ZDV LGHQWLÀDEOH 3ODWHV &DXGDO ÀQ photographs of selected preserved specimens were XVHGWRSUHSDUHGUDZLQJVRIFDXGDOÀQFRORXUSDWWHUQV (Plate 3).
In cases where species showed no clear separation in any one character, but where a combination of several characters showed only minor overlap, Principal Component Analysis (PCA), with size-adjustment based on the residuals gained from log-log regressions of the morphometric variables with standard length, was used to obtain information on the optimal distinction HJ8LEOHLQ :LQNOHU1LHOVHQ
Complementary information on species distributions was obtained from Fischer et al. (1990), Randall (2001), 5DQGDOO .XOELFNLDQG)ULFNHHWDO
TAXONOMY
*HQXVUpeneus&XYLHU UpeneusCuvier 1829: 157. Type species Mullus vittatus)RUVVNnOE\VXEVHTXHQW
GHVLJQDWLRQRI'HVPDUHVW
DIAGNOSIS'RUVDOÀQV9,,RU9,,,DQDOÀQ,SHOYLF ÀQV,SHFWRUDOÀQUD\V²SULQFLSDOFDXGDOÀQ UD\VPHGLDQEUDQFKHGJLOOUDNHUV²² ²ODWHUDOOLQHVFDOHV²ODWHUDOOLQHFRPSOHWH small scales present basally on second dorsal, anal and FDXGDO ÀQV VPDOO WHHWK SUHVHQW RQ YRPHU SDODWLQHV and both jaws, multiserial and villiform on jaws; body REORQJVOLJKWO\FRPSUHVVHGEDUEHOOHQJWKLQDGXOWV² WLPHVLQ6/VQRXWOHQJWK²WLPHVLQ6/VXEHTXDOWR SRVWRUELWDOOHQJWK²WLPHVLQ6/LQIUHVKÀVKODWHUDO ERG\VWULSHVDQGRUFDXGDOÀQEDUVRIGLIIHULQJFRORXU GDUNFDXGDOÀQEDUVIUHTXHQWO\UHWDLQHGRQSUHVHUYHG ÀVK
DISTRIBUTION (Table 2). In all major oceans, tropical to subtropical, only a single species in the Atlantic and two in the Mediterranean, both so-called “Lessepsian”
immigrants from the Red Sea after the opening of the 6XH]&DQDO%HQ7XYLD
REMARKS.:HUHFRJQL]HVSHFLHVDVYDOLGVSHFLHV IURPWKH:HVWHUQ,QGLDQ2FHDQ7DEOH6PLWK consideredUpeneus queketti Gilchrist & Thompson 1908,
described from Durban, as a synonym of Mulloidichthys
DXULÁDPPD )RUVVNnO %DXFKRW HW DO
consideredU. niebuhri*Xp]pGHVFULEHGIURPWKH Gulf of Suez, a synonym of U. tragula.
SPECIES ACCOUNTS
%DVHG RQ D FRPELQDWLRQ RI WKH QXPEHU RI GRUVDOÀQ VSLQHVJLOOUDNHUVQXPEHURISHFWRUDOÀQUD\VDQGEDUV RQFDXGDOÀQDOOXVHGLQHDUOLHUDFFRXQWVHJ/DFKQHU 7KRPDV6DLQVEXU\HWDO*RODQL 5DQGDOO.LP 1DND\DIRXUPDMRUVSHFLHV complexes can be distinguished (Table 2).
6SHFLHVFRPSOH[, the “japonicusJURXSµGRUVDOÀQ spines distinguish this group clearly from all others; in DGGLWLRQWKHVHVSHFLHVKDYH²WRWDOJLOOUDNHUVDQG
²SHFWRUDOÀQUD\VIUHVKÀVKRIWKLVJURXSDOOKDYH EDUV RQ WKH XSSHU FDXGDOÀQ OREH DQG VHYHUDO VSHFLHV DOVRRQWKHORZHUFDXGDOÀQOREH7KLVFRPSOH[LQFOXGHV Upeneus guttatus and U. pori and the non-Western Indian Ocean species, U. asymmetricus,U. australiae, U.
francisi,U. japonicus, and U. parvus, the last being the only Atlantic representative of the genus.
6SHFLHV FRPSOH[ , the “tragula JURXSµ GRUVDOÀQ VSLQHV ²UDUHO\ WRWDO JLOO UDNHUV DQG ² UDUHO\SHFWRUDOÀQUD\VDOOVSHFLHVZLWKEDUVRQWKH
XSSHUFDXGDOÀQOREHDQGVHYHUDOZLWKEDUVRQWKHORZHU FDXGDOÀQOREH7KLVFRPSOH[LQFOXGHVU. margarethae sp. nov., U. oligospilus,U. sundaicus,U. taeniopterus,U.
tragula, and the non-Western Indian Ocean species U.
luzonius, and U. mouthami.
6SHFLHVFRPSOH[, the “moluccensisJURXSµGRUVDOÀQ VSLQHVUDUHO\²WRWDOJLOOUDNHUVUDUHO\² SHFWRUDOÀQUD\VDQGQREDUVRQWKHORZHUFDXGDOÀQ OREHEXWVRPHVSHFLHVKDYHEDUVRQWKHXSSHUFDXGDOÀQ lobe. This complex includes U. doriae,U. moluccensis,U.
sulphureus, and the non-Western Indian Ocean species U. quadrilineatus.
6SHFLHV FRPSOH[ , the “vittatus group”: 8 dorsal- ÀQ VSLQHV ² WRWDO JLOO UDNHUV ² SHFWRUDOÀQ UD\V DQG EDUV RQ ERWK FDXGDOÀQ OREHV LQ IUHVK DQG SUHVHUYHGÀVK7KLVFRPSOH[LQFOXGHVU. davidaromi,U.
indicus sp. nov., U. mascareinsis,U. suahelicus sp. nov., U.
supravittatussp. nov., U. vittatus, and the non-Western Indian Ocean species U. subvittatus.
7KH 3DFLÀF VSHFLHV8SHQHXV ÀOLIHU KDV GRUVDOÀQ spines with a unique, extremely long second spine WKDWH[WHQGVEH\RQGWKHEDVHRIWKHVHFRQGGRUVDOÀQ This species is not included in any of the four species FRPSOH[HVLWKDV²JLOOUDNHUVRUSHFWRUDOÀQ UD\VDQGQREDUVRQWKHFDXGDOÀQ5DQGDOO .XOELFNL 8LEOHLQXQSXEOLVKHGGDWD
Table 2. Distribution and taxonomic grouping of all 26 currently recognized Upeneus species. The species occurring in the Western Indian Ocean are emphasized in bold.
KEY TO THEWESTERNINDIANOCEANSPECIES OFUPENEUS
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D 'RUVDOÀQVSLQHVÀUVWVSLQHVXVXDOO\WKHORQJHVWSHFWRUDOÀQUD\V²WRWDOJLOOUDNHUV²ODWHUDOOLQH VFDOHV²SHFWRUDODQGSHOYLFÀQVVXEHTXDOLQOHQJWK E 'RUVDOÀQVSLQHVÀUVWVSLQHPLQXWHUHFXPEHQWSDUWO\KLGGHQE\VNLQDQGVFDOHVDWÀQRULJLQSHFWRUDO ÀQUD\V²WRWDOJLOOUDNHUV²ODWHUDOOLQHVFDOHV²SHFWRUDODQGSHOYLFÀQVVXEHTXDOLQOHQJWKRU SHFWRUDOÀQVORQJHUWKDQSHOYLFÀQV D 3HFWRUDOÀQUD\VPRVWO\UDUHO\WRWDOJLOOUDNHUV²ERG\UHGGRUVDOO\SUHVHUYHGÀVKSDOHEURZQ QRWGDUNHUGRUVDOO\ZLWKFDXGDOÀQEDUVQRWUHWDLQHG,QGR:HVW3DFLÀFU. guttatus E 3HFWRUDOÀQ UD\V WRWDO JLOO UDNHUV ² ERG\ JUH\ WR UHGGLVKEURZQ GRUVDOO\ SUHVHUYHG ÀVK GDUNHU
GRUVDOO\ZLWKFDXGDOÀQEDUVFRPPRQO\UHWDLQHG5HG6HDWR2PDQ0DGDJDVFDU0HGLWHUUDQHDQU. pori D 7RWDOJLOOUDNHUV²UDNHUVRQORZHUOLPE²SHFWRUDOÀQUD\V²SHFWRUDODQGSHOYLFÀQVVXEHTXDO
in length ... 4 E 7RWDOJLOOUDNHUV²UDNHUVRQORZHUOLPE²SHFWRUDOÀQUD\V²SHFWRUDOÀQVORQJHUWKDQSHOYLFÀQV ... 8 D 'DUNGRWVRUEORWFKHVRQERG\DQGSDLUHGÀQVÀUVWGRUVDOÀQWLSGDUNRQHEURZQWREODFNPLGODWHUDOERG\
VWULSHLQIUHVKDQGSUHVHUYHGÀVKXSSHUMDZOHQJWK²WLPHVLQ6/
E 1RGDUNGRWVRUEORWFKHVRQERG\DQGSDLUHGÀQVQRGDUNGRUVDOÀQWLSRQHRUWZR\HOORZLVKRUSDOHEURZQ ODWHUDOERG\VWULSHVRQIUHVKÀVKXSSHUMDZOHQJWK²WLPHVLQ6/
D 7RWDOQXPEHURIFDXGDOÀQEDUV²RUIHZHULQMXYHQLOHVFP6/FDXGDOÀQOHQJWK²WLPHVLQ6/
DQG²WLPHVLQKHDGOHQJWKSHOYLFÀQOHQJWK²WLPHVLQ6/3HUVLDQ*XOIU. oligospilus E 7RWDOQXPEHUFDXGDOÀQEDUVRUPRUH²LQMXYHQLOHVFP6/FDXGDOÀQOHQJWK²WLPHVLQ6/
DQG²WLPHVLQKHDGOHQJWKSHOYLFÀQOHQJWK²WLPHVLQ6/,QGR:HVW3DFLÀFU. tragula D 3HFWRUDOÀQOHQJWK²WLPHVLQ6/ODWHUDOOLQHVFDOHV²XSSHUFDXGDOÀQOREHZLWKGDUNEDUVZHOO
UHWDLQHGRQSUHVHUYHGÀVKIUHVKÀVKZLWKDSDOHEURZQPLGODWHUDOERG\VWULSHDQGDZHDNHUPRUH\HOORZLVK VWULSHEHORZ,QGR&HQWUDO3DFLÀFU. taeniopterus E 3HFWRUDOÀQOHQJWK²WLPHVLQ6/ODWHUDOOLQHVFDOHV²XSSHUFDXGDOÀQOREHZLWKUHGRUJUH\
EDUVQRWRURQO\WUDFHVUHWDLQHGRQSUHVHUYHGÀVKIUHVKÀVKZLWKD\HOORZRUSDOHEURZQPLGODWHUDOERG\
stripe ... 7 D 7RWDOJLOOUDNHUV²ODWHUDOOLQHVFDOHV²ÀUVWGRUVDOÀQKHLJKW²WLPHVLQ6/FDXGDOSHGXQFOH
GHSWK²WLPHVLQ6/EDUEHOVZKLWHIRXUUHGEDUVRQXSSHUFDXGDOÀQOREHLQIUHVKÀVKWKUHHGLVWDOO\IURP IRUN,QGLDQ2FHDQDQG$UDIXUD6HDU. margarethae sp. nov.
E 7RWDOJLOOUDNHUV²ODWHUDOOLQHVFDOHV²ÀUVWGRUVDOÀQKHLJKW²WLPHVLQ6/FDXGDOSHGXQFOH GHSWK²WLPHVLQ6/EDUEHOVIUHTXHQWO\\HOORZLQIUHVKÀVK²UHGRUJUH\EDUVRQXSSHUFDXGDOÀQOREH LQIUHVKÀVK,QGR3DFLÀFU. sundaicus D 1REDUVRQFDXGDOÀQDOVRQRWLQOLYHRUIUHVKÀVKÀUVWGRUVDOÀQWLSEODFNRUSDOHEURZQWR\HOORZLVKFRORXU
PD\GLVDSSHDURQSUHVHUYHGÀVKERG\GHSWKDWÀUVWGRUVDOÀQRULJLQ²WLPHVLQ6/LQDGXOWÀVK!FP 6/SHFWRUDOÀQOHQJWK²WLPHVLQKHDGOHQJWK E &DXGDOÀQZLWKHLWKHU²UHGEDUVRQXSSHUOREHRQO\RU²SDOHEURZQWREODFNEDUVRQHQWLUHÀQ²EDUV
RQXSSHUDQG²RQORZHUOREHEDUVFRQVSLFXRXVLQOLYHÀVKDQGPRVWO\ZHOOUHWDLQHGRQSUHVHUYHGÀVK EURZQWREODFNÀUVWGRUVDOÀQWLSUHWDLQHGRQSUHVHUYHGÀVKERG\GHSWKDWÀUVWGRUVDOÀQRULJLQ²WLPHV LQ6/LQDGXOWÀVK!FP6/SHFWRUDOÀQOHQJWK²WLPHVLQKHDGOHQJWK D 7RWDOJLOOUDNHUV²UDNHUVRQORZHUOLPEWLSRIÀUVWGRUVDOÀQSDOHEURZQWR\HOORZLVKLQIUHVKÀVK
QRWRURQO\IDLQWO\UHWDLQHGRQSUHVHUYHGÀVKIUHVKÀVKZLWKDQDUURZ\HOORZPLGODWHUDOERG\VWULSHDQDO ÀQKHLJKW²WLPHVLQ6/ÀUVWGRUVDOÀQKHLJKW²WLPHVLQ6/VHFRQGGRUVDOÀQKHLJKW²WLPHV in SL (Persian Gulf and Gulf of Oman) ... U. doriae E 7RWDOJLOOUDNHUV²UDNHUVRQORZHUOLPE²WLSRIÀUVWGRUVDOÀQEODFNLQIUHVKDQGSUHVHUYHGÀVK
IUHVKÀVKZLWKWZRFRQVSLFXRXV\HOORZODWHUDOERG\VWULSHVDQDOÀQKHLJKW²WLPHVLQ6/ÀUVWGRUVDOÀQ KHLJKW²WLPHVLQ6/VHFRQGGRUVDOÀQKHLJKW²WLPHVLQ6/,QGR:HVW3DFLÀFU. sulphureus
D 1R EDUV RQ ORZHU FDXGDOÀQ OREH XSSHU OREH ZLWK ² UHG EDUV IDLQWO\ UHWDLQHG RQ SUHVHUYHG ÀVK RQH FRQVSLFXRXV\HOORZRUJROGPLGODWHUDOERG\VWULSHLQIUHVKÀVKKHDGOHQJWK²WLPHVLQ6/KHDGGHSWK WKURXJKH\H²WLPHVLQ6/,QGR:HVW3DFLÀFHDVWHUQ0HGLWHUUDQHDQU. moluccensis E /RZHUFDXGDOÀQOREHZLWK²EURZQRUEODFNEDUVXSSHUFDXGDOÀQOREHZLWK²EURZQRUEODFNEDUVQRRU
DWOHDVWWZRODWHUDOERG\VWULSHVLQIUHVKÀVKKHDGOHQJWK²WLPHVLQ6/KHDGGHSWKWKURXJKH\H² times in SL ... 11 D &DXGDOÀQ EDUV SDOH EURZQ WR GDUN EURZQ XQLIRUPO\ FRORXUHG SDOH VSDFHV EHWZHHQ EDUV RI QHDUO\ HTXDO ZLGWKEDUVRQXSSHUFDXGDOÀQOREHFXUYHGWZR\HOORZRUSDOHEURZQODWHUDOERG\VWULSHVRQIUHVKÀVKERG\
GHSWKDWDQDOÀQRULJLQ²WLPHVLQ6/SHFWRUDOÀQOHQJWK²WLPHVLQ6/DQG²LQKHDGOHQJWK WRWDOJLOOUDNHUV² E &DXGDOÀQEDUVDWOHDVWSDUWO\EODFNRUGDUNEURZQYDU\LQFRORXULQWHQVLW\EDUVRUWKHVSDFHVEHWZHHQWKHP
YDU\RIWHQLQZLGWKFDXGDOÀQEDUVRQERWKOREHVPRVWO\VWUDLJKWDQGQRWFXUYHGQRRUPRUHWKDQWZRODWHUDO ERG\VWULSHVRQIUHVKÀVKERG\GHSWKDWDQDOÀQRULJLQ²WLPHVLQ6/SHFWRUDOÀQOHQJWK²WLPHVLQ 6/DQG²LQKHDGOHQJWKWRWDOJLOOUDNHUV² D 3HFWRUDOÀQOHQJWK²WLPHVLQ6/DQGWLPHVLQKHDGOHQJWKVKRUWHUWKDQERG\GHSWKDWDQDOÀQRULJLQ WKHODWWHU²WLPHVLQ6/6:,QGLDU. indicus sp. nov.
E 3HFWRUDOÀQOHQJWK²WLPHVLQ6/DQG²WLPHVLQKHDGOHQJWKVXEHTXDOWRRUORQJHUWKDQERG\GHSWK DWDQDOÀQRULJLQWKHODWWHU²WLPHVLQ6/
D 7RWDOQXPEHURIJLOOUDNHUV²UDNHUVRQORZHUOLPE²EDUEHOOHQJWK²WLPHVLQ6/SHFWRUDOÀQ OHQJWK ² WLPHV LQ 6/ DQG ² WLPHV LQ VXEHTXDO WR ERG\ GHSWK DW DQDOÀQ RULJLQ KHDG OHQJWK
²WLPHVLQ6/(DVW$IULFDDQGVRXWKHUQ5HG6HDU. suahelicus sp. nov.
E 7RWDOQXPEHURIJLOOUDNHUVUDUHO\²UDNHUVRQORZHUOLPE²EDUEHOOHQJWK²WLPHVLQ6/
SHFWRUDOÀQOHQJWK²WLPHVLQ6/DQG²WLPHVLQ JUHDWHUWKDQERG\GHSWKDWDQDOÀQRULJLQKHDG OHQJWK²WLPHVLQ6/6RXWK,QGLDDQG6UL/DQNDU. supravittatus sp. nov.
D +HDGGHSWKWKURXJKH\H²WLPHVLQ6/DQDOÀQKHLJKW²VHFRQGGRUVDOÀQKHLJKW²WLPHVLQ 6/QRODWHUDOERG\VWULSHLQIUHVKÀVK5HG6HDU. davidaromi E +HDGGHSWKWKURXJKH\H²WLPHVLQ6/DQDOÀQKHLJKW²VHFRQGGRUVDOÀQKHLJKW²WLPHVLQ
6/QRRUPRUHWKDQWZRODWHUDOERG\VWULSHVRQIUHVKÀVK D %RG\GHSWKDWDQDOÀQRULJLQ²WLPHVLQ6/FDXGDOSHGXQFOHGHSWK²WLPHVLQ6/ÀUVWGRUVDOÀQ
KHLJKW ² WLPHV LQ 6/ YHUWLFDO OHQJWK RI EODFN WLS RQ ÀUVW GRUVDO ÀQ DQG ZLGWK RI ODUJHVW EDU DQGRU LQWHUVSDFHEHWZHHQGLVWDOPRVWEDUVRIORZHUFDXGDOÀQOREHQDUURZHUWKDQRUELWQRODWHUDOERG\VWULSHVRQ IUHVKÀVK,QGLDQ2FHDQU. mascareinsis E %RG\GHSWKDWDQDOÀQRULJLQ²WLPHVLQ6/FDXGDOSHGXQFOHGHSWK²WLPHVLQ6/ÀUVWGRUVDOÀQ
KHLJKW ² WLPHV LQ 6/ YHUWLFDO OHQJWK RI EODFN WLS RQ ÀUVW GRUVDO ÀQ DQG ZLGWK RI ODUJHVW EDU DQGRU LQWHUVSDFHEHWZHHQGLVWDOPRVWEDUVRIORZHUFDXGDOÀQOREHVXEHTXDOWRRUZLGHUWKDQRUELWRUODWHUDO ERG\VWULSHVRQIUHVKÀVKWZR\HOORZRUFRSSHU\VWULSHVPLGODWHUDOO\DQGEHORZDQGRQHRUWZREURZQRU SDOHEURZQVWULSHVGRUVDOO\,QGR:HVW3DFLÀFU. vittatus
Upeneus davidaromi *RODQL
(Tables 3, 8; Plates 1, 3)
Upeneus subvittatus (non 7HPPLQFN 6FKOHJHO Ben-Tuvia & Kissil 1988: 12, Fig. 11 (type locality:
(ODW*XOIRI$TDED5HG6HDGHSWKUDQJH² m; Baranes & Golani 1993: 310.
Upeneus vitatus [sic] (non )RUVVNnO.KDODI 'LVL 120 (misspelling of name).
Upeneus davidaromi Golani 2001: 112, Fig. 1a (type locality Eilat, Gulf of Aqaba).
DIAGNOSIS'RUVDOÀQV9,,,SHFWRUDOÀQVRU JLOOUDNHUV² ²ODWHUDOOLQHVFDOHV² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ 6/ ERG\
GHSWK DW DQXV ² FDXGDOSHGXQFOH GHSWK ² PD[LPXPKHDGGHSWKKHDGGHSWKWKURXJKH\H² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWK²EDUEHOOHQJWK²FDXGDOÀQOHQJWK² DQDOÀQKHLJKW²SHOYLFÀQOHQJWKSHFWRUDO ÀQ OHQJWK ² ÀUVW GRUVDOÀQ KHLJKW ² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO QXPEHU RI FDXGDO ÀQ EDUV ² ² GDUN EDUV RQ XSSHU FDXGDOÀQ OREH (including one bar close to rear end of lateral line), colour of bars changing from pale brown proximally WRGDUNEURZQRUEODFNDWUHDUFDXGDOÀQPDUJLQDQG WLS LQ IUHVK ÀVK ² EDUV RQ ORZHU FDXGDOÀQ OREH slightly increasing in width distally, one usually at tip and smaller, the distal-most two or three bars almost HQWLUHO\GDUNEURZQRUEODFNÀUVWGRUVDOÀQWLSEODFN
FDXGDOÀQ EDUV DQG EODFN GRUVDOÀQ WLS UHWDLQHG RQ SUHVHUYHG ÀVK EDUEHOV ZKLWH QR ODWHUDO ERG\ VWULSHV head and body silvery laterally, the snout and dorsal part of head and body reddish, and belly white in fresh ÀVKZKLWHEHOO\FRORXUDWLRQSDUWO\UHWDLQHGDQGERG\
GRUVDOO\GDUNHQHGLQSUHVHUYHGÀVK DISTRIBUTION. Red Sea
COMPARISONS. A Western Indian Ocean species of
the vittatus JURXS 7DEOHV 3ODWHV ²Upeneus davidaromi differs from U. indicus in fewer gill UDNHUV ERG\ DQG FDXGDOSHGXQFOH OHVV GHHS GHHSHU head through eye, larger suborbital depth, longer KHDG ODUJHU RUELW ORQJHU XSSHU MDZ KLJKHU DQDO ÀQ ORZHU ÀUVW GRUVDOÀQ FDXGDOÀQ EDUV OHVV XQLIRUPO\
coloured, and lateral body stripes absent; from U.
mascareinsis it differs in deeper head through eye, greater suborbital depth, longer postorbital and head, and body colouration silvery instead of rose-red; it Table 3. Meristic and colour characters (a), morphometric characters as %SL (b), and morphometric characters as
ratios of SL (c) for the Western Indian Ocean Upeneusspecies.
differs from U. suahelicus in body and caudal-peduncle less deep, deeper head through eye, larger suborbital GHSWK ORQJHU KHDG DQG VQRXW VKRUWHU SHFWRUDO ÀQV ORZHU ÀUVW GRUVDOÀQ ODWHUDO ERG\ VWULSHV DEVHQW DQG FDXGDOÀQEDUVOHVVXQLIRUPO\FRORXUHGLWGLIIHUVIURP U. supravittatusLQIHZHUJLOOUDNHUVERG\DQGFDXGDO peduncle less deep, deeper head through eye, larger VXERUELWDO GHSWK ODUJHU RUELW VKRUWHU SHFWRUDO ÀQV ORZHU ÀUVW GRUVDOÀQ ODWHUDO ERG\ VWULSHV DEVHQW DQG FDXGDOÀQEDUVOHVVXQLIRUPO\FRORXUHGDQGLWGLIIHUV from U. vittatus LQ ERG\ OHVV GHHS DW GRUVDOÀQ RULJLQ and caudal-peduncle less deep, deeper head through eye, larger suborbital depth, longer head, larger orbit, ORQJHUXSSHUMDZVKRUWHUGLVWDQFHEHWZHHQGRUVDOÀQV VKRUWHUDQDOÀQEDVHORZHUÀUVWGRUVDOÀQODWHUDOERG\
VWULSHVDEVHQWDQGZLGHVWEDURQORZHUFDXGDOÀQOREH smaller.
REMARKS. $PRQJ WKH WZR NQRZQ GHHSZDWHU VSHFLHV ofUpeneus from the Indian Ocean region, U. davidaromi occurs deepest, with a reported depth range of 150 to PGHSWKZKLOHWKHPD[LPXPGHSWKUHSRUWHGIRUU.
mascareinsisLVP)RXUPDQRLU *Xp]p7KH photographed specimen (Plate 1) differs from another individual of Upeneus davidaromi illustrated in the species description by Golani (2001) in a more regular EDUDQGLQWHUVSDFHZLGWKRQWKHORZHUFDXGDOÀQOREH
Upeneus davidaromi attains 24 cm SL.
Upeneus doriae*QWKHU
(Tables 3, 7; Plates 1, 3)
Upeneoides doriae *QWKHU W\SH ORFDOLW\
Bender Abassi, Persian Gulf).
0XOORLFKWK\V DXULÁDPPD (non )RUVVNnO %OHJYDG Løppenthin 1944: 133, Plate 7, Fig. 1 (Persian Gulf).
Upeneus doriae 5DQGDOO )LJ SKRWR RI FP ÀVK IURP 2PDQ GLDJQRVLV DQG FRORXU photograph; Carpenter et al. 1997: 193, Pl. 13, Fig. 88 3HUVLDQ*XOI.LP 1DND\DII)LJUH GHVFULSWLRQDQGSKRWRJUDSKRISUHVHUYHGÀVK DIAGNOSIS 'RUVDO ÀQV 9,,, SHFWRUDO ÀQV ² JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/
ERG\GHSWKDWDQXVFDXGDOSHGXQFOHGHSWK² PD[LPXPKHDGGHSWK²KHDGGHSWKWKURXJKH\H
² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWK EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK
² DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK ² SHFWRUDOÀQOHQJWK²ÀUVWGRUVDOÀQKHLJKW² VHFRQGGRUVDOÀQKHLJKW²6/QREDUVRQFDXGDO ÀQRQHQDUURZ\HOORZPLGODWHUDOERG\VWULSHXVXDOO\
QRWUHWDLQHGRQSUHVHUYHGÀVKÀUVWGRUVDOÀQWLSSDOH EURZQWR\HOORZLVKLQIUHVKÀVKSLJPHQWDWLRQIDLQWO\
UHWDLQHGRUORVWLQSUHVHUYHGÀVKEDUEHOVZKLWHERG\
VLOYHU\GRUVDOO\GDUNHQHGLQIUHVKÀVKDQGSDOHWRGDUN EURZQVRPHWLPHVGRUDOO\GDUNHULQSUHVHUYHGÀVK
DISTRIBUTION. Persian Gulf and Gulf of Oman
COMPARISONS. Western Indian Ocean species of the moluccensis JURXS 7DEOHV 3ODWHV ²Upeneus doriae differs from U. moluccensis LQ PRUH JLOO UDNHUV GHHSHUERG\DQGKHDGQREDUVRQFDXGDOÀQDQGERG\
FRORXUDWLRQ VLOYHU\ YV UHG LQ IUHVK ÀVK DQG LW GLIIHUV from U. sulphureus LQ PRUH WRWDO JLOO UDNHUV DQG JLOO UDNHUVRQORZHUOLPEERG\OHVVGHHSVKRUWHUSHOYLFÀQV DQGORZHUGRUVDOÀQV
REMARKS. In their redescription of Upeneus doriae, Kim DQG1DND\DHPSKDVL]HWKHKLJKHUQXPEHURIJLOO UDNHUVRQWKHORZHUOLPEDVLPSRUWDQWIRUWKHGLVWLQFWLRQ with U. sulphureus. Comparison of material from the two Gulf areas revealed geographic differences in orbit VL]H DQG FDXGDOÀQ OHQJWK LQ DGXOW VSHFLPHQV 7DEOH 7). However, because the Gulf of Oman specimens are clearly larger in size than those of the Persian Gulf, the possibility of allometric changes during ontogeny FDQQRWEHUXOHGRXW2QHMXYHQLOHPP6/IURP the Persian Gulf, which was examined in detail, clearly deviated in several morphometric measurements from adult specimens and hence, similar to U. mascareinsis, U. tragula, and U. oligospilus, was treated separately, WRJHWKHUZLWKWKHVHFRQGMXYHQLOHVSHFLPHQPP6/
Table 7).
Meristic and selected morphometric data, and SKRWRJUDSKV RI IUHVK DQG SUHVHUYHG ÀVK IURP WKH HDVWHUQ *XOI RI 2PDQ %3%0 DQG ZHUH PDGH DYDLODEOH E\ -DFN 5DQGDOO $UQROG Suzumoto, Jerry Finan and Jeff Williams, indicating WKDWLQWKH*XOIRI2PDQWKHÀUVWGRUVDOÀQWLSRIIUHVK ÀVKLVSDOH\HOORZLVKUDWKHUWKDQSDOHEURZQDQGWKLV pigmentation can be completely lost after preservation.
The same is true for the yellow mid-lateral body stripe;
WKHERG\FRORULQSUHVHUYHGÀVKYDULHVIURPSDOHEURZQ WREURZQDQGLVLQVRPHLQVWDQFHVGRUVDOO\GDUNHURU D EURDG GDUNHU VWULSH DSSHDUV DORQJ WKH PLG ERG\
There is a tight overlap in counts, the BPBM material KDYLQJ²WRWDOJLOOUDNHUV²RQORZHUOLPE,I WKHPRUSKRPHWULFPHDVXUHPHQWVRIDGXOWÀVK!FP SL) examined in the current study are combined with data from the BPBM specimens, important diagnostic FKDUDFWHUVVXFKDVERG\GHSWKDWÀUVWGRUVDOÀQRULJLQ
² 6/ ERG\ GHSWK DW DQDOÀQ RULJLQ ² 6/DQGKHDGOHQJWK²6/UHWDLQUDWKHUQDUURZ ranges and overlap. The BPBM specimens are included as “additional material” in the material list.
Upeneus doriae attains 20 cm SL; it occurs in shallow water to 45 m depth.
Upeneus guttatus'D\
(Tables 3, 4; Plates 1, 3)
Upenoides guttatus 'D\W\SHORFDOLW\0DGUDV India).
Upeneus crosnieri )RXUPDQRLU *Xp]p )LJ
,FW\SHORFDOLW\0LWVLR3UDFHO%DQN0DGDJDVFDU Bauchot et al. 1985: 7 (synonym of Upeneoides guttatus'D\
Upeneus guttatus5DQGDOO .XOELFNLSS)LJV
²GLDJQRVLVDQGWZRFRORXUSKRWRJUDSKV Upeneus bensasi (non7HPPLQFN 6FKOHJHO'H%UXLQHW
DO3O)LJ6UL/DQND
DIAGNOSIS 'RUVDO ÀQV 9,, SHFWRUDO ÀQV ² PRVWO\ LQ RI VSHFLPHQV JLOO UDNHUV ²
² ²ODWHUDOOLQHVFDOHV²ERG\GHSWKDW ÀUVWGRUVDOÀQRULJLQ²6/ERG\GHSWKDWDQXV
² FDXGDOSHGXQFOH GHSWK ² PD[LPXP KHDG GHSWK ² KHDG GHSWK WKURXJK H\H ² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWK
² EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK ² DQDOÀQKHLJKWSHOYLFÀQOHQJWK²SHFWRUDO ÀQ OHQJWK ² ÀUVW GRUVDOÀQ KHLJKW ² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO EDUV RQ FDXGDO ÀQ
² XSSHU FDXGDOÀQ OREH ZLWK UHGGLVK EDUV RI similar width or narrower than the pale interspaces EHWZHHQ WKH EDUV QRW UHWDLQHG RQ SUHVHUYHG ÀVK ² faint, irregular red bars on exterior margin of lower FDXGDOÀQOREHVRPHWLPHVH[WHQGLQJRYHUHQWLUHOREH RUDEVHQWQRWRURQO\WUDFHVUHWDLQHGRQSUHVHUYHGÀVK QRODWHUDOERG\VWULSHVÀUVWGRUVDOÀQWLSSDOHEDUEHOV
\HOORZRUZKLWHLQIUHVKÀVKERG\ZKLWHEHORZODWHUDO line, covered by red pigmentation above lateral line which may also reach down ventrally, forming red patches or blotches; belly white; body pale brown and QRWGRUVDOO\GDUNHQHGLQSUHVHUYHGÀVK
DISTRIBUTION. Red Sea to Somalia, Kenya, Mozambique, 6RXWK$IULFD.ZD=XOX1DWDO0DGDJDVFDU5pXQLRQ 6H\FKHOOHV,QGLD6UL/DQND%D\RI%HQJDO$QGDPDQ Sea, Malaysia, Singapore, northern Australia and Philippines.
COMPARISONS. Western Indian Ocean species of the japonicus group (Tables 3, 4, Plates 1, 3): Upeneus guttatus differs from U. pori in mostly 13 vs 14 pectoral- ÀQUD\VIHZHUJLOOUDNHUVQRRURQO\WUDFHVRIEDUVYV FRQVSLFXRXV EDU SDWWHUQV RQ ERWK FDXGDOÀQ OREHV RI SUHVHUYHGÀVKDQGSDOHEURZQYVGRUVDOO\GDUNERG\
RQSUHVHUYHGÀVK
Non-Western Indian Ocean species of the japonicus group (Table 4): Upeneus guttatus differs from U.
asymmetricus LQIHZHUJLOOUDNHUVKDOIERG\DWDQDOÀQ origin less deep, bars not crossing entire lower caudal- ÀQOREHLQIUHVKÀVKQRODWHUDOERG\VWULSHLQIUHVKÀVK DQGEDUVLQPRVWFDVHVQRWUHWDLQHGRQSUHVHUYHGÀVK fromU. australiae it differs in mostly 13 vs 14 pectoral- ÀQUD\VODWHUDOERG\VWULSHDEVHQWLQIUHVKÀVKORZHU FDXGDOÀQOREHQRWHQWLUHO\FURVVHGE\EDUVLQIUHVKÀVK FDXGDOÀQEDUVLQPRVWFDVHVQRWUHWDLQHGRQSUHVHUYHG ÀVK DQG ERG\ GRUVDOO\ SDOHU LQ SUHVHUYHG ÀVK IURP U. francisiLWGLIIHUVLQPRVWO\YVSHFWRUDOÀQUD\V IHZHU JLOO UDNHUV VKRUWHU KHDG VKRUWHU SRVWRUELWDO OHQJWK VKRUWHU SDLUHG ÀQV DQG ERG\ FRORXU UHG YV
VLOYHU\LQIUHVKÀVKIURPU. japonicus it differs in fewer JLOO UDNHUV VKRUWHU EDUEHOV VKRUWHU SHFWRUDO ÀQV DQG EDUVSUHVHQWDORQJYHQWUDOPDUJLQRIORZHUFDXGDOÀQ OREH LQ IUHVK ÀVK DQG IURP WKH $WODQWLFU. parvus it GLIIHUV PRVWO\ LQ YV SHFWRUDOÀQ UD\V IHZHU JLOO UDNHUV DQG ODWHUDOOLQH VFDOHV VKRUWHU EDUEHOV ORQJHU DQGKLJKHUDQDOÀQVKRUWHUSHFWRUDOÀQDQGEDUVQRW FURVVLQJORZHUFDXGDOÀQOREHHQWLUHO\RQIUHVKÀVK REMARKS. Bauchot et al. (1985) synonymized Upeneus crosnieri)RXUPDQRLU *Xp]pZLWKU. guttatus (Day :HIROORZWKLVFRQFOXVLRQEDVHGRQH[DPLQDWLRQ of the holotype of U. crosnieri which overlaps U. guttatus in all important characters.
New record for the Red Sea. Upeneus guttatus attains FP6/GHSWKUDQJH²P
Upeneus indicusVSQRY
(Tables 3, 8; Plates 1, 3)
Holotype: BPBM 27524, 137 mm, Cochin, western India, FROOHFWHG E\ - ( 5DQGDOO YLD ORFDO ÀVKHUPHQ 1980).
Paratype: BPBM 40987, 132 mm, Cochin, collected together with holotype.
DIAGNOSIS'RUVDOÀQ9,,,SHFWRUDOÀQV²JLOO UDNHUV ² ² ODWHUDOOLQH VFDOHV ERG\
GHSWKDWÀUVWGRUVDOÀQRULJLQ²6/ERG\GHSWK DW DQDOÀQ RULJLQ ² FDXGDOSHGXQFOH GHSWK PD[LPXPKHDGGHSWKKHDGGHSWKWKURXJKH\H² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWKEDUEHOOHQJWK²FDXGDOÀQOHQJWK² DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK SHFWRUDO ÀQ OHQJWK ² ÀUVW GRUVDOÀQ KHLJKW ² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO EDUV RQ FDXGDO ÀQ
² ² SDOH EURZQ EDUV RQ XSSHU FDXGDOÀQ OREH RI VDPH FRORXU RQ ORZHU FDXGDOÀQ OREH WKH WKUHH SUR[LPDOEDUVRQXSSHUFDXGDOÀQOREHVOLJKWO\FXUYHG FDXGDOÀQ EDUV ZLGHU WKDQ SDOH LQWHUVSDFHV EHWZHHQ WKHPEDUVIDLQWO\UHWDLQHGRQSUHVHUYHGÀVKWKUHHSDOH EURZQODWHUDOERG\VWULSHVRQIUHVKÀVKRQHPLGODWHUDO VWULSH UHDFKLQJ IURP EHKLQG KHDG WR FDXGDOÀQ EDVH RQHVWULSHZHOOEHORZVWDUWLQJEHKLQGSHFWRUDOÀQEDVH and a third, short stripe just above lateral line below GRUVDO ÀQV OHQJWK RI GDUN ÀUVW GRUVDOÀQ WLS FOHDUO\
smaller than orbit; head and body silvery-rose laterally, GRUVDOO\GDUNUHGGLVKDQGEHOO\VLOYHU\ZKLWHLQIUHVK ÀVKERG\LQSUHVHUYHGÀVKXQLIRUPO\EURZQ
DESCRIPTION. Measurements in % SL and counts are given in Table 8; morphometric data as ratios of SL IRUKRORW\SHGDWDIRUSDUDW\SHLIGLIIHUHQWLQEUDFNHWV ERG\GHHSLWVGHSWKDWÀUVWGRUVDOÀQRULJLQ>@
ERG\GHSWKDWDQDOÀQRULJLQ>@KHDGOHQJWK [3.3], similar to maximum depth of body and longer
WKDQFDXGDOÀQOHQJWK>@KHLJKWRIVHFRQGGRUVDO ÀQ>@OHVVWKDQEDUEHOOHQJWK>@SHFWRUDO ÀQOHQJWK>@FOHDUO\ORQJHUWKDQSHOYLFÀQOHQJWK (4.8); caudal-peduncle depth 9.1, clearly larger than orbit (14).
Fresh colour (Plate 1): Head and body silvery-rose, belly silvery-white, dorsal surface of body and head reddish;
body with three narrow, pale brown lateral stripes, one PLGODWHUDOIURPKHDGWRFDXGDOÀQEDVHDQWHULRUO\ZHOO EHORZODWHUDOOLQHFURVVLQJOLQHEHORZVHFRQGGRUVDOÀQ and connecting to proximal-most bar of upper caudal- ÀQ OREH RQH VWULSH VWDUWLQJ EHKLQG SHFWRUDOÀQ EDVH and connecting to proximal-most bar of lower caudal- ÀQOREHDQGDWKLUGVKRUWVWULSHMXVWDERYHODWHUDOOLQH VWDUWLQJ EHORZ ÀUVW GRUVDO DQG HQGLQJ EHORZ VHFRQG GRUVDO ÀQ XSSHU OREH RI FDXGDO ÀQ ZKLWH UD\V SDOH URVHJUD\SUR[LPDOO\DQGK\DOLQHGLVWDOO\ZLWKRU SDOHEURZQEDUVWKHÀQOREHWLSGDUNWKHSUR[LPDO RUEDUVVOLJKWO\FXUYHGORZHUFDXGDOÀQOREHZKLWH rays mostly hyaline with 4 straight pale brown bars, the lobe tip pale; bars and white interspaces on both lobes uniform in width, the bars slightly wider than WKH LQWHUVSDFHV VSLQRXV GRUVDO ÀQ ZKLWH ZLWK WZR pale brown horizontal stripes, the lower one thin and considerably separated by a pale white interspace from EDVHWKHXSSHURQHDVZLGHDVWKHEODFNÀQWLSDERYH YHUWLFDO OHQJWK RI EODFN ÀQ WLS VPDOOHU WKDQ RUELW DQG QHDUO\ HTXDO WR FDXGDOÀQ EDU ZLGWK DOO WKUHH SDOH LQWHUVSDFHV EHWZHHQ VWULSHV ZLGHU WKDQ VWULSHV RU ÀQ WLS VRIW GRUVDO ÀQ SDOH ZKLWH ZLWK IRXU SDOH EURZQ horizontal stripes, the uppermost stripe covering the ÀQ WLS DOPRVW FRPSOHWHO\ H[FHSW IRU D WLQ\ SDOH VSRW at the very tip itself; the two lower stripes narrow, the ORZHVWRQHRQO\SUHVHQWRQGLVWDOSDUWRIWKHÀQWKHWZR uppermost stripes of similar width as the middle stripe RQWKHVSLQRXVGRUVDOÀQDQDODQGSHOYLFÀQVK\DOLEDQH ZLWKZKLWHUD\VURVHDWEDVHSHFWRUDOÀQVK\DOLQH Preserved colour (Plate 3): Head and body uniformly EURZQ EODFN ÀUVW GRUVDOÀQ WLS GLVWLQFW ODWHUDO ERG\
VWULSHVDEVHQWEDUVRQFDXGDOÀQOREHVIDLQWDQGUDWKHU LQGLVWLQFW QRW UHWDLQHG RQ XSSHU FDXGDOÀQ OREH RI SDUDW\SHSHFWRUDOSHOYLFDQGDQDOÀQVWUDQVSDUHQW DISTRIBUTION 2QO\ NQRZQ IURP WKH W\SH ORFDOLW\
Cochin, Western India
ETYMOLOGY. The name “indicus” derives from the type locality and is treated as a noun in apposition.
COMPARISONS. Western Indian Ocean species of the vittatusJURXS7DEOHV3ODWHV²Upeneus indicus differs from U. davidaromi LQ PRUH JLOO UDNHUV GHHSHU body and caudal peduncle, head through eye less deep, smaller suborbital depth, shorter head, smaller RUELW VKRUWHU XSSHU MDZ ORZHU DQDO ÀQ KLJKHU ÀUVW
GRUVDO ÀQ FDXGDOÀQ EDUV PRUH XQLIRUPO\ FRORXUHG DQGODWHUDOERG\VWULSHVSUHVHQWLQIUHVKÀVKU. indicus differs from U. mascareinsisLQPRUHJLOOUDNHUVRQXSSHU limb, deeper body and caudal peduncle, larger head GHSWK VPDOOHU RUELW FDXGDOÀQ EDUV PRUH XQLIRUPO\
coloured and more regularly sized on lower lobe, and ODWHUDOERG\VWULSHVSUHVHQWLQIUHVKÀVKLWGLIIHUVIURP U. suahelicusLQPRUHJLOOUDNHUVDQGODWHUDOOLQHVFDOHV GHHSHUKDOIERG\DWÀUVWGRUVDOÀQRULJLQRUELWORZHU DQDO ÀQ VKRUWHU SHFWRUDO ÀQV DQG ODWHUDO KHDG DQG body colour reddish vs silvery-white; it differs from U. supravittatusLQDGHHSHUERG\VKRUWHUSHFWRUDOÀQV and lateral head and body colour reddish vs brassy in IUHVK ÀVK DQG LW GLIIHUV IURPU. vittatus in more gill UDNHUV RQ XSSHU OLPE GHHSHU ERG\ DW DQDOÀQ RULJLQ VPDOOHU RUELW FDXGDOÀQ EDUV OHVV LQWHQVHO\ DQG PRUH uniformly coloured, more regularly-sized on lower OREHEODFNÀUVWGRUVDOÀQWLSVPDOOHUDQGODWHUDOERG\
and head colour reddish vs silvery white.
REMARKS. Upeneus indicus attains 14 cm SL; no depth information is available.
Upeneus margarethaeVSQRY
(Tables 3, 5; Plates 1, 3)
Upeneus bensasi 7HPPLQFN 6FKOHJHO%DXFKRW
& Bianchi. 1984, in part; Fischer et al. 1990, 155; in part.
Holotype: SAIAB 82217, male 82 mm, Mozambique, 59Dr. F. Nansen0Ý·6Ý·(
m, bottom trawl.
Paratypes: Somalia 8610 PP ·
1 · (Kenya: 6$,$% ² PP RII .LSLQL · 6 · ( ² P0R]DPELTXH 6$,$% PP 59. Dr. F. Nansen, M07-77,
· 6 · ( P ERWWRP WUDZO 6$,$%
²PP:HVWHUQ,QGLDQ2FHDQ0R]DPELTXH 59Dr. F. Nansen0·6·(
m, bottom trawl; Madagascar6$,$%² PP7VLPLSDLND%D\0$'²P
Non-types01+1²PP·1·
( (ULWUHD =08& 3 PP 3HUVLDQ *XOI 8610²PP&RORPERÀVKPDUNHWODQGHG DW(UDYXU6UL/DQND%01+² PP·6·(1:$XVWUDOLD$06, PP·6·($UDIXUD6HD59Soela, 100-105 m, engel trawl (N Australia).
DIAGNOSIS 'RUVDO ÀQV 9,,, SHFWRUDO ÀQV ² PRVWO\LQRILQGLYLGXDOVJLOOUDNHUV²² ²ODWHUDOOLQHVFDOHV²ERG\GHSWKDWÀUVW GRUVDOÀQRULJLQ²6/ERG\GHSWKDWDQXV² FDXGDOSHGXQFOH GHSWK ² PD[LPXP KHDG GHSWK
²KHDGGHSWKWKURXJKH\H²KHDGOHQJWK² RUELWOHQJWK²XSSHUMDZOHQJWK²EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK ² SHFWRUDOÀQ OHQJWK
² ÀUVW GRUVDOÀQ KHLJKW ² VHFRQG GRUVDO ÀQ KHLJKW ² 6/ WRWDO EDUV RQ FDXGDO ÀQ ² EDUVRQXSSHUFDXGDOÀQOREHRQOREHLWVHOIDQGRQH DWOREHEDVHEURDGUHGEDQGRQORZHUFDXGDOÀQOREH FRYHULQJ XS WR RU UHG EDUV ZKLFK DUH RQO\ SDUWO\
YLVLEOHDORQJWKHRXWHUYHQWUDOÀQPDUJLQLQIUHVKÀVK EDUVPD\EHFRPHPRUHYLVLEOHLQSUHVHUYHGÀVKEXWDUH XVXDOO\QRWUHWDLQHGRQSUHVHUYHGÀVKDSDUWIURPVRPH WUDFHVRIEDUVDQGGDUNSLJPHQWDWLRQDWWKHORZHUOREH ÀQWLSDVLQJOHPLGODWHUDOERG\VWULSHUXQQLQJWKURXJK eye, red from snout tip to eye and yellow from behind H\HWRFDXGDOÀQEDVHQRWUHWDLQHGRQSUHVHUYHGÀVK ÀUVWGRUVDOÀQWLSSDOHEDUEHOVZKLWHERG\UHGDERYH lateral line, body and head white laterally, with red or rose dots or blotches, ventral side of head and belly ZKLWH ERG\ XQLIRUPO\ SDOH EURZQ LQ SUHVHUYHG ÀVK ÀUVWPLQXWHGRUVDOÀQVSLQHIUHTXHQWO\GDUNSLJPHQWHG IRUPLQJDVPDOOVSRWDWÀQRULJLQWKDWLVRIWHQGLVWLQFW IURPDERYHLQSUHVHUYHGÀVK
DESCRIPTION. Measurements in % SL and counts are given in Table 5; morphometric data as ratios of SL for KRORW\SHGDWDIRUSDUDW\SHVLQEUDFNHWVERG\HORQJDWH ERG\GHSWKDWÀUVWGRUVDOÀQRULJLQ>²@ERG\
GHSWK DW DQDOÀQ RULJLQ >²@ KHDG OHQJWK
>²@ODUJHUWKDQPD[LPXPGHSWKRIERG\DQGVXE HTXDOWRFDXGDOÀQOHQJWK>²@VHFRQGGRUVDO ÀQOHQJWK>²@VLPLODUWREDUEHOOHQJWK>²
@SHOYLFÀQOHQJWK>²@VLPLODUWROHQJWKRI SHFWRUDOÀQV>²@FDXGDOSHGXQFOHGHSWK
>²@ODUJHUWKDQRUELWOHQJWK>²@
Fresh colour (Plate 1): Head and body white laterally, ZLWK VRPH LUUHJXODU IDLQW UHG PDUNLQJV UHG GRUVDOO\
(above lateral line), belly white; a red mid-lateral stripe RQ VQRXW WKURXJK H\H \HOORZ IURP H\H WR FDXGDOÀQ base, stripe anteriorly below lateral line, posteriorly along or slightly above lateral line; the stripe sometimes RYHUODLQZLWKUHG,ULVFRPSOHWHO\UHG'RUVDOÀQVSDOH ZLWK WZR VXEKRUL]RQWDO UHG VWULSHV SHFWRUDO ÀQV K\DOLQH WKH UD\V SDOH UHGGLVK SHOYLF ÀQV SDOH ZLWK RU IDLQW GXVN\UHG VWULSHV FDXGDOÀQ XSSHU OREH K\DOLQH ZLWK WKUHH UHG EDUV RQ ÀQ OREH LWVHOI DQG D fourth bar at the base of lobe; bar width equal to width of pale interspaces between bars; lower lobe with broad UHGEDQGFRYHULQJRUUHGEDUVYLVLEOHDORQJYHQWUDO ÀQPDUJLQPPSDUDW\SHZLWKLQQHUPDUJLQRIORZHU FDXGDOÀQ OREH GDUN DQG D VHULHV RI WKUHH VPDOO SDOH VSRWVDQDOÀQK\DOLQHZLWKZKLWHUD\VEDUEHOVZKLWH Preserved colour (Plate 3): head and body pale beige to SDOHEURZQVRPHLVRODWHGGDUNSDWFKHVRQRSHUFXOXP RQGRUVDOVLGHRIERG\DIHZW\SHVVKRZDGDUNVDGGOH EHKLQGWKHVHFRQGGRUVDOÀQWKDWUHDFKHVWRODWHUDOOLQH DOOÀQVWUDQVSDUHQWDQGZLWKRXWSLJPHQWH[FHSWIRUD
GDUNLQWHULRUPDUJLQRQORZHUFDXGDOÀQOREHWRZDUGV WLSRIOREHDQGVRPHVPDOOGDUNVSRWVRUVWULSHVRQERWK OREHV VRPHWLPHV DOVR UHPDLQV RI EDUV RQ FDXGDOÀQ OREHVRQORZHUOREHXSWRVL[GDUNQDUURZEDUVUHDFKLQJ from close to ventral margin to about two-thirds of lobe width, becoming most conspicuous at mid of lobes. In holotype and several paratypes the minute spine in IURQWRIWKHÀUVWGRUVDOÀQLVLQGLFDWHGE\DGDUNGRW,Q specimens with intact scales, the lateral line is distinct, H[WHQGLQJWRRUVOLJKWO\EH\RQGFDXGDOÀQEDVH
DISTRIBUTION. Red Sea to Somalia, Kenya, Mozambique, 0DGDJDVFDU3HUVLDQ*XOI6UL/DQND1:$XVWUDOLD1 Australia (Arafura Sea).
ETYMOLOGY The name of this species “margarethae”
LVLQKRQRXURIWKHPRWKHURIWKHÀUVWDXWKRUWKHODWH Margaretha Uiblein (née Feichtinger).
COMPARISONS. Western Indian Ocean species of the tragula JURXS 7DEOHV 3ODWHV ² Upeneus margarethae differs from U. oligospilus in shorter snout DQGSRVWRUELWDOOHQJWKORQJHUFDXGDOÀQORQJHUSHOYLF DQG SHFWRUDO ÀQV UHG YV ² EURZQ RU EODFN EDUV RQ XSSHU FDXGDOÀQ OREH QR FRQVSLFXRXV GDUN EDUV RQ ORZHU FDXGDOÀQ OREH ERG\ DQG ÀQV ODFNLQJ GDUN EORWFKHVRUGRWVÀUVWDQGVHFRQGGRUVDOÀQWLSVSDOHLQ ERWKIUHVKDQGSUHVHUYHGÀVKFRORXURIPLGODWHUDOERG\
VWULSH\HOORZYVEURZQRUGDUNEURZQDQGPLGODWHUDO ERG\ VWULSH QRW UHWDLQHG RQ SUHVHUYHG ÀVK LW GLIIHUV from U. sundaicus LQ PRUH JLOO UDNHUV IHZHU ODWHUDO line scales, caudal peduncle and suborbital less deep, ORZHU ÀUVW GRUVDO ÀQ YV ² EDUV RQ XSSHU FDXGDO ÀQ OREH WKH PLGODWHUDO ERG\ VWULSH VWDUWLQJ DW VQRXW vs behind eye, and barbels white vs yellow (colour SDWWHUQVDSSO\WRIUHVKÀVKRQO\IURPU. taeniopterus it differs in fewer lateral-line scales, larger orbit, longer SHOYLFDQGSHFWRUDOÀQVKLJKHUVHFRQGGRUVDOÀQRQH YVWZRODWHUDOERG\VWULSHVLQIUHVKÀVKDQGFDXGDOÀQ EDUVLQPRVWFDVHVQRWUHWDLQHGRQSUHVHUYHGÀVKDQG fromU. tragulaLWGLIIHUVLQPRVWO\LQRIÀVK YVPRVWO\LQRIÀVKSHFWRUDOÀQUD\VVKRUWHU MDZVORQJHUSHFWRUDOÀQVUHGYV²EURZQRUEODFN EDUV RQ XSSHU FDXGDOÀQ OREH LQ DGXOWV ! FP 6/
GDUNEDUVRQORZHUFDXGDOÀQOREHDEVHQWWKHERG\DQG ÀQVODFNLQJGDUNEORWFKHVRUGRWVWKHÀUVWDQGVHFRQG GRUVDOÀQWLSVSDOHPLGODWHUDOERG\VWULSH\HOORZYV EURZQWREODFNDQGPLGODWHUDOERG\VWULSHFRORXUQRW UHWDLQHGRQSUHVHUYHGÀVK
Non-Western Indian Ocean species of the tragula JURXS 7DEOHV Upeneus margarethae differs from U. luzoniusLQPRUHJLOOUDNHUVIHZHUODWHUDOOLQHVFDOHV KDOIERG\DWDQDOÀQRULJLQDQGFDXGDOSHGXQFOHOHVV GHHSKHDGDQGERG\QRWGDUNHUGRUVDOO\LQSUHVHUYHG ÀVK DQG SDOH LQWHUVSDFHV EHWZHHQ XSSHU OREH EDUV wider; from U. mouthami it differs in mostly 14 vs 13 SHFWRUDOÀQUD\VVKRUWHUEDUEHOVVKRUWHUÀUVWGRUVDOÀQ base, broad red band vs brown to red bars on lower FDXGDOÀQ OREH LQ IUHVK ÀVK QR EDU UHPDLQV RU RQO\
ZHDNOLQHVYVGDUNEORWFKHVDVUHPDLQVRIEDUVRQORZHU FDXGDOÀQOREHLQSUHVHUYHGÀVKDQGZKLWHYV\HOORZ EDUEHOVLQIUHVKÀVK
REMARKS. Since Lachner (1954) there has been speculation that there may be a different form of Upeneus bensasi (Houttyn, 1782) in the Western Indian Ocean, a name that he applied erroneously to Upeneus guttatus, which was described from off Madras, India.
Since the synonymization of U. bensasi with the earlier described, seven dorsal-spine U. japonicus by Randall et al. (1993), it is clear that this eight-spine form cannot be ascribed to U. bensasi(DUOLHUZRUNHUVVXFKDV7KRPDV PD\ KDYH DOUHDG\ HQFRXQWHUHG WKLV PDWHULDO but probably missed the minute spine in front of the dorsal, as he did when examining the holotype of U.
taeniopterus at MNHN in Paris (see also the species account of U. taeniopterus below).
1R LQIRUPDWLRQ RQ FRORXUDWLRQ RI IUHVK ÀVK IURP areas other than the southern East African coast is currently available. Also, the specimens from northern SDUWVRIWKH:HVWHUQ,QGLDQ2FHDQUHJLRQ6UL/DQNDDQG NW Australia showed some deviation in a PCA based on morphometric characters (Uiblein, unpublished data) and hence were not given a type status.
Upeneus margarethae attains 11 cm SL; depth range 8—50 m.
Upeneus mascareinsis )RXUPDQRLU
*Xp]p
(Tables 3, 8; Plates 1, 3)
Upeneus mascareinsis )RXUPDQRLU *Xp]p )LJ,ESKRWRJUDSKRISUHVHUYHGÀVKW\SHORFDOLW\
5pXQLRQ /HWRXUQHXU HW DO )ULFNH HW DO 2009.
DIAGNOSIS'RUVDOÀQV9,,,SHFWRUDOÀQV²JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/ ERG\
GHSWK DW DQXV ² FDXGDOSHGXQFOH GHSWK ² PD[LPXPKHDGGHSWK²KHDGGHSWKWKURXJKH\H
² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWK ² EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK ² DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK
²SHFWRUDOÀQOHQJWK²ÀUVWGRUVDOÀQKHLJKW
² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO EDUV RQFDXGDOÀQ²²EDUVRQXSSHUFDXGDOÀQOREH colour of bars changing from pale brown proximally WR GDUN EURZQ RU EODFN DW GLVWDO PDUJLQ RI OREH ² EURZQEDUVRQORZHUFDXGDOÀQOREHEHFRPLQJGDUNHU GLVWDOO\ODVWWZRFRPSOHWHO\GDUNEURZQRUEODFNEDUV slightly increasing in width distally, bar on tip small;
ZLGWKRIODUJHVWORZHUFDXGDOÀQOREHEDUDQGRUVSDFH between distal-most bars less than orbit; no lateral ERG\ VWULSHV ÀUVW GRUVDOÀQ WLS GDUN EDUEHOV ZKLWH body white ventrally and rose-red laterally, becoming GRUVDOO\GDUNHUZLWKVRPHJROGLULGHVFHQFHDORQJGRUVDO
ÁDQNVDQGDGDUNHQHGVDGGOHEHKLQGVHFRQGGRUVDOÀQ SUHVHUYHG ÀVK ERG\ XQLIRUPO\ EURZQ RU D ELW GDUNHU GRUVDOO\EDUVDQGEODFNÀUVWGRUVDOÀQWLSUHWDLQHG DISTRIBUTION. Madagascar, Mozambique, Réunion, Western Indonesia
COMPARISONS. Western Indian Ocean species of the vittatus JURXS 7DEOHV 3ODWHV ²Upeneus mascareinsis differs from U. davidaromi in head through eye less deep, smaller suborbital depth, shorter postorbital distance, and head and body colouration rose-red laterally instead of silvery; from U. indicus it GLIIHUV LQ IHZHU JLOO UDNHUV RQ XSSHU OLPE ERG\ DQG caudal peduncle less deep, smaller maximum head GHSWK ODUJHU RUELW FDXGDOÀQ EDUV OHVV XQLIRUPO\
coloured and less regularly sized on lower lobe, lateral ERG\ VWULSHV DEVHQW LQ IUHVK ÀVK IURPU. suahelicus it differs in body and caudal-peduncle less deep, shorter SHFWRUDO ÀQ FDXGDOÀQ EDUV OHVV XQLIRUPO\ FRORXUHG and less regularly sized on lower lobe, lateral body stripes absent, and head and body colour reddish vs pale brown laterally; from U. supravittatus it differs in body and caudal peduncle less deep, shorter pectoral ÀQORZHUÀUVWGRUVDOÀQFDXGDOÀQEDUVOHVVXQLIRUPO\
coloured and less regularly sized on lower lobe, bar interspace width on both lobes not generally narrower than bars; lateral body stripes absent, and head and body colour more reddish than brassy; and it differs fromU. vittatus in caudal peduncle less deep, smaller suborbital depth, the largest bar and/or pale interspace EHWZHHQ GLVWDOPRVW EDUV RQ ORZHU FDXGDO ÀQOREH QDUURZHUÀUVWGRUVDOÀQEODFNWLSVPDOOHUODWHUDOERG\
stripes absent, and body and head colour laterally reddish instead of silvery-white.
Non-Indian Ocean species of vittatus group (Table 8, Fig. 1): U. mascareinsis differs from U. subvittatus in more lateral-line scales, body and caudal peduncle less deep, VKRUWHUVQRXWDQGSRVWRUELWDOOHQJWKORZHUDQDOÀQDQG VKRUWHUSHFWRUDOÀQVWKHWZRVSHFLHVODUJHO\UHVHPEOH each other in colour, U. subvittatus laterally silvery-grey LQFRQWUDVWWRUHGGLVKWKHGDUNVDGGOHEHKLQGWKHVHFRQG GRUVDOÀQORQJHUH[WHQGLQJIURPLPPHGLDWHO\EHKLQG VHFRQGGRUVDOÀQEDVHWREDVHRIXSSHUFDXGDOÀQOREH Because of considerable overlap of morphometric measurements, a PCA based on 37 variables gathered from 9 U. mascareinsis and 5 U. subvittatus was carried out (Fig. 1). A clear separation was found between WKH WZR VSHFLHV %RG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ caudal-peduncle depth, maximum head depth, snout OHQJWKDQGSHFWRUDOÀQOHQJWKDUHDPRQJWKHYDULDEOHV contributing most to the 31% of total variance explained E\WKHÀUVWSULQFLSDOFRPSRQHQW
REMARKS. Our data agree well with several details given in the original species description of U. mascareinsis by )RXUPDQRLU *Xp]p 6LPLODU WRU. davidaromi, FRQVLGHUDEOH LQWUDVSHFLÀF YDULDWLRQ LQ ZLGWK RI ORZHU
FDXGDOÀQOREHEDUVDQGSDOHLQWHUVSDFHVZHUHIRXQG These differences occur also among individuals of similar size and overlapping distribution and hence may indicate a colour polymorphism.
One silvery juvenile specimen with an elongated ERG\DQGVKRUWSHFWRUDOÀQVFROOHFWHGRII0R]DPELTXH was found to be closely related to Upeneus mascareinsis based on meristic characters, some measurements of body form, and collecting depth (100 m; Fig. 2). This ÀVKGLIIHUHGKRZHYHUFRQVLGHUDEO\IURPDGXOWVRIWKH VSHFLHV LQ RUELW OHQJWK SHFWRUDOÀQ OHQJWK GLVWDQFH RI SHFWRUDOÀQ RULJLQ IURP VQRXW WLS PD[LPXP KHDG depth, and the body colour dorsally silvery-bluish instead of reddish (Table 8, Plate 1, Fig. 2). Some of these GHYLDWLRQV PD\ UHÁHFW DQ HDUO\ RQWRJHQHWLF WUDQVLWLRQ in lifestyle, as similar differences have been reported IRURWKHUJRDWÀVKVSHFLHVHJ&DOGZHOO8LEOHLQ 'XHWRWKHVHFRQVLGHUDEOHGLIIHUHQFHVDQGODFNRI additional comparative material this specimen is only referred to as cf. mascareinsis.
During the course of comparing U. mascareinsis with U. subvittatus we found indications for considerable variation in body colour among U. subvittatus specimens IURP GLIIHUHQW DUHDV LQ WKH 3DFLÀF &RPSDULVRQ RI D specimen photographed by J. E. Randall from the
0ROXFFD ,VODQGV ,QGRQHVLD %3%0 FP 6/
with photographs of two specimens from Japan, one shown in Masuda et al. (1984) and another one in 2NDPXUD $PDRND VXJJHVWV WKH H[LVWHQFH RI additional colour variants with either a ventrally pale grey and dorsally brown body or an overall reddish ERG\ %RWK VSHFLPHQV IURP -DSDQ ODFN D GDUN VDGGOH EHKLQG WKH VHFRQG GRUVDO ÀQ 'HWDLOHG WD[RQRPLF studies of these different forms are needed.
To date Upeneus mascareinsis had only been recorded from Réunion and was assumed to be endemic there /HWRXUQHXUHWDO)ULFNHHWDO2XUUHFRUGV for Madagascar, Mozambique and Western Indonesia are new for this species. It has been collected between 100 and 400 m depth and thus inhabits the lower shelf and upper slope in the Indian Ocean proper, as does U.
davidaromi in the Red Sea. Upeneus mascareinsis attains 18 cm SL.
Upeneus moluccensis%OHHNHU
(Tables 3, 7; Plates 1, 3)
Upeneoides moluccensis %OHHNHUW\SHORFDOLW\
Ambon, Moluccan Islands).
Upeneus moluccensis: Kumaran & Randall 1984, no page number, Plate III; colour photograph; Ben-Tuvia &
Kissil 1988: 11, Fig. 10 (Red Sea: Eritrea and Elat, Gulf of Aqaba); De Bruin et al. 1994: 270, Pl. 20, Fig. 137 FRORXUSKRWR6UL/DQND5DQGDOO .XOELFNL 301-302, Fig. 5 (diagnosis and colour photograph, New Caledonia).
DIAGNOSIS'RUVDOÀQV9,,,SHFWRUDOÀQV²JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/ ERG\
GHSWK DW DQXV ² FDXGDOSHGXQFOH GHSWK ² PD[LPXPKHDGGHSWK²KHDGGHSWKWKURXJKH\H
² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZOHQJWK²EDUEHOOHQJWK²FDXGDOÀQOHQJWK
² DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK ² SHFWRUDOÀQOHQJWK²ÀUVWGRUVDOÀQKHLJKW² VHFRQGGRUVDOÀQKHLJKW²6/²WKLQUHGEDUV RQXSSHUFDXGDOÀQOREHIDLQWO\UHWDLQHGRQSUHVHUYHG ÀVKQREDUVRQORZHUFDXGDOÀQOREHEXWDUHGEURDG band covering the entire lower lobe apart from distal, LQQHUPDUJLQWKHODWWHUVRPHZKDWGDUNHUPRVWRIORZHU FDXGDOÀQOREHSLJPHQWDWLRQORVWLQSUHVHUYHGÀVKRQH conspicuous yellow or gold mid-lateral body stripe IURPH\HWRXSSHUFDXGDOÀQEDVHQRWRURQO\IDLQWO\
UHWDLQHG RQ SUHVHUYHG ÀVK ÀUVW GRUVDOÀQ WLS GDUN DOVRLQSUHVHUYHGÀVKEDUEHOVZKLWHERG\VLOYHU\URVH GDUNHQHG DERYH ODWHUDO VWULSH LQ SUHVHUYHG ÀVK ERG\
SDOHEURZQVOLJKWO\GDUNHQHGGRUVDOO\
DISTRIBUTION. Eastern Mediterranean, Red Sea to Mozambique, Arabian Sea, Madagascar, India, Andaman Sea, Indonesia, New Caledonia, Philippines, Taiwan and southern Japan.
Fig. 2. Photograph of Upeneuscf.mascareinsis, SAIAB 82328, 6.9 cm SL, collected off Mozambique (O. Alvheim, IMR)
Fig. 1. First principal component (PC1) based on 37 morphometric characters against number of lateral- line scales for 9 Upeneus mascareinsis (Indian Ocean) and 5 U. subvittatus :HVW3DFL¿F:,2
= Western Indian Ocean, EIO = Eastern Indian Ocean)
COMPARISONS. Western Indian Ocean species of the moluccensis JURXS 7DEOHV 3ODWHV ² )LJ Upeneus moluccensis differs from U. doriae in fewer gill UDNHUV ERG\ DQG KHDG OHVV GHHS EDUV RQ FDXGDO ÀQ present, and body colouration red vs silvery in fresh ÀVKDQGLWGLIIHUVIURPU. sulphureus in body, head and caudal peduncle less deep, shorter and lower anal and GRUVDOÀQVEDUVRQWKHXSSHUFDXGDOÀQOREHSUHVHQW one vs two lateral stripes and red vs white to silvery ERG\FRORXUDWLRQLQIUHVKÀVK
REMARKS.Upeneus moluccensis attains 18 cm SL; depth UDQJH²P
Upeneus oligospilus /DFKQHU
7DEOHV3ODWHV
Upeneus oligospilus Lachner 1954 (type locality: Persian Gulf).
Upeneus tragula 5LFKDUGVRQ .XURQXPD $EH 1972, p. 88, Fig. 29 (drawing of 13.8 cm specimen from Kuwait with typical features of U. oligospilus).
Synonym of Upeneus tragula5DQGDOO .XOELFNL p. 305.
DIAGNOSIS'RUVDOÀQV9,,,SHFWRUDOÀQV²JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/ ERG\
GHSWK DW DQXV ² FDXGDOSHGXQFOH GHSWK ² PD[LPXPKHDGGHSWK²KHDGGHSWKWKURXJKH\H
² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZOHQJWK²EDUEHOOHQJWK²FDXGDOÀQOHQJWK
² DQDOÀQ KHLJKW ² SHOYLFÀQ OHQJWK ² SHFWRUDOÀQOHQJWK²ÀUVWGRUVDOÀQKHLJKW² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO QXPEHU RI EDUVRQFDXGDOÀQ²LQDGXOWVRUOHVVLQMXYHQLOHV!
FP6/XSSHUFDXGDOÀQOREHZLWK²EURZQGDUN EURZQRUEODFNEDUV²EURZQGDUNEURZQRUEODFN EDUVRQORZHUOREHDEURZQRUGDUNEURZQPLGODWHUDO ERG\VWULSHIURPWLSRIVQRXWWRFDXGDOÀQEDVHERG\
above stripe greenish grey, body below stripe white RUEHLJHZLWKLUUHJXODUGDUNUHGGLVKEURZQVSRWVDQG EORWFKHV DOVR RQ SDLUHG ÀQV GDUN PDUNV RQ RU YHU\
FORVHWRÀUVWDQGVHFRQGGRUVDOÀQWLSVEDUEHOV\HOORZ LQIUHVKÀVKPRVWRIERG\DQGÀQSLJPHQWDWLRQUHWDLQHG RQSUHVHUYHGÀVK
DISTRIBUTION. Persian Gulf
COMPARISONS. Western Indian Ocean species of the tragula JURXS 7DEOHV 3ODWHV ² )LJ Upeneus oligospilus differs from U. margarethae in longer snout DQG SRVWRUELWDO OHQJWK VKRUWHU FDXGDO ÀQ VKRUWHU SHOYLF DQG SHFWRUDO ÀQV ² EURZQ RU EODFN YV UHG EDUV RQ XSSHU FDXGDOÀQ OREH FRQVSLFXRXV GDUN EDUV RQORZHUFDXGDOÀQOREHSUHVHQWLQIUHVKÀVKERG\DQG ÀQV VKRZLQJ GDUN EORWFKHV RU GRWV ÀUVW DQG VHFRQG GRUVDOÀQ WLSV GDUN LQ ERWK IUHVK DQG SUHVHUYHG ÀVK
FRORXURIPLGODWHUDOERG\VWULSHEURZQRUGDUNEURZQ vs yellow, and mid-lateral body stripe retained on SUHVHUYHGÀVKIURPU. sundaicusLQPRUHJLOOUDNHUV IHZHU ODWHUDOOLQH VFDOHV ERG\ DW DQDOÀQ RULJLQ DQG FDXGDOSHGXQFOHOHVVGHHSSDLUHGÀQVPRVWO\VKRUWHU ÀUVWGRUVDOÀQFRQVLGHUDEO\ORZHUORZHUFDXGDOÀQOREH ZLWKRUGDUNEDUVGDUNEORWFKHVDQGGRWVRQERG\
DQG SDLUHG ÀQV SUHVHQW EODFN GRUVDOÀQ WLSV LQ ERWK IUHVKDQGSUHVHUYHGÀVKSUHVHQWDQGPLGODWHUDOERG\
VWULSH EURZQ RU GDUN EURZQ YV SDOH EURZQ UHWDLQHG RQ SUHVHUYHG ÀVK IURPU. taeniopterus it differs in fewer lateral-line scales, greater maximum head depth, ORQJHU KHDG DQG VQRXW VKRUWHU FDXGDO ÀQ KLJKHU VHFRQGGRUVDOÀQGDUNEORWFKHVDQGGRWVRQERG\DQG SDLUHG ÀQV SUHVHQW EODFN GRUVDOÀQ WLSV LQ ERWK IUHVK DQGSUHVHUYHGÀVKSUHVHQWDQGPLGODWHUDOERG\VWULSH EURZQRUGDUNEURZQYVSDOHEURZQRQO\WKHIRUPHU UHWDLQHGRQSUHVHUYHGÀVKDQGLWGLIIHUVIURPU. tragula LQVKRUWHUFDXGDOÀQDQGIHZHUFDXGDOÀQEDUVERWKLQ DGXOWV²YV²DQGMXYHQLOHVYV²)LJV$
B, D) and in greater postorbital length, longer distance EHWZHHQ WLS RI VQRXW WR SHFWRUDOÀQ RULJLQ VKRUWHU SHOYLFÀQDQGORZHUVHFRQGGRUVDOÀQLQDGXOWÀVKLI size is referred to (e.g. Fig. 3C). [FIG. 3]
Non-Western Indian Ocean species of the tragula JURXS 7DEOH Upeneus oligospilus differs from U.
luzoniusLQPRUHJLOOUDNHUVFDXGDOSHGXQFOHOHVVGHHS ORQJHU KHDG DQG VQRXW VKRUWHU SHOYLF ÀQ ORZHU ÀUVW GRUVDOÀQGDUNGRWVDQGEORWFKHVRQERG\DQGSDLUHG ÀQVSUHVHQWDQGGDUNGRUVDOÀQWLSVSUHVHQWLWGLIIHUV from U. mouthami in longer snout and postorbital length, smaller orbit, longer upper jaw, shorter barbels, ORQJHUÀUVWGRUVDOÀQEDVHVKRUWHUFDXGDOÀQVKRUWHU SDLUHG ÀQV EURZQ RU GDUN EURZQ YV \HOORZ ODWHUDO ERG\VWULSHWKHIRUPHUUHWDLQHGRQSUHVHUYHGÀVKGDUN GRWVDQGEORWFKHVRQERG\DQGSDLUHGÀQVSUHVHQWDQG GDUNGRUVDOÀQWLSVSUHVHQW
REMARKS. Upeneus oligospilus is resurrected from synonymy with U. tragula 5DQGDOO .XOELFNL because these two species can be clearly distinguished by the combination of two characters, the total number RI GDUN EDUV RQ WKH FDXGDO ÀQ DQG FDXGDOÀQ OHQJWK (Fig. 3D). This distinction did not become apparent in the earlier taxonomic studies of Lachner (1954) and 7KRPDVZKRSURYLGHGFDXGDOÀQEDUFRXQWVEXW QRGHWDLOHGFRPSDULVRQVRIFDXGDOÀQOHQJWK
$SKRWRJUDSKRIDIUHVKO\FROOHFWHGFPÀVKIURP WKH*XOIRI6XH]ZLWKVKRUWXQSDLUHGÀQVDQGEDUVRQ WKHFDXGDOÀQWKDWEHFDPHUHFHQWO\DYDLODEOHWRWKHÀUVW DXWKRU6HUJH\%RJRURGVN\SHUVFRPPPD\LQGLFDWH thatU. oligospilus is distributed more widely. Thomas DWWULEXWHG RQH VSHFLPHQ IURP QRUWKHUQ 6UL /DQNDWRWKLVVSHFLHV7KHWZRMXYHQLOHVSHFLPHQVIURP 6UL/DQNDH[DPLQHGLQWKLVVWXG\FOHDUO\IDOOZLWKLQWKH FDXGDOÀQOHQJWKUDQJHRIUpeneus tragula.
Upeneus oligospilus attains 17 cm SL; maximum reported depth is 13 m.
Upeneus pori %HQ7XYLD *RODQL
(Tables 3, 4; Plates 1, 3)
Upeneus pori Ben-Tuvia & Golani 1989: 13 ff., Figs.
² SKRWRJUDSKV RI IUHVK ÀVK ex-situ and in-situ) (type locality: Elat, Gulf of Aqaba, Red Sea); Kim &
1DND\D)LJGLDJQRVWLFFKDUDFWHUVDQG SKRWRJUDSKVKRZLQJFDXGDOÀQSDWWHUQ
DIAGNOSIS 'RUVDO ÀQV 9,, SHFWRUDO ÀQV JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/ ERG\
GHSWK DW DQXV ² FDXGDOSHGXQFOH GHSWK ² PD[LPXPGHDGGHSWK²KHDGGHSWKWKURXJKH\H
² KHDG OHQJWK ² RUELW OHQJWK ² XSSHU MDZ OHQJWK ² EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK ² DQDOÀQ KHLJKW SHOYLFÀQ OHQJWK
²SHFWRUDOÀQOHQJWK²ÀUVWGRUVDOÀQKHLJKW
² VHFRQG GRUVDOÀQ KHLJKW ² 6/ WRWDO EDUV RQFDXGDOÀQ²XSSHUFDXGDOÀQOREHZLWK²UHG EURZQEDUV²UHGEURZQRUJUH\EDUVRQYHQWUDOKDOI VLGHRIORZHUOREHH[WHQGLQJWRDEURDGEURZQRUGDUN grey stripe along middle of lobe; 3 or 4 brownish-red or JUH\EDUVRQLQQHUGRUVDOKDOIRIORZHUFDXGDOÀQOREH LQSUHVHUYHGÀVKEDUSDWWHUQVRIWHQUHWDLQHGVKRZLQJ GDUNEURZQFRORXUDWLRQQRRURQO\IDLQWLQGLFDWLRQRI D PLGODWHUDO ERG\ VWULSH LQ IUHVK ÀVK ÀUVW GRUVDO ÀQ
not pigmented; barbels white or yellow; body greyish RU UHGEURZQ GRUVDOO\ GDUNHQHG DQG FRYHUHG ZLWK grey to reddish-brown spots extending to lateral side;
OLYHÀVKPD\VKRZDGDUNEURZQRUUHGEDUIURPH\H YHUWLFDOO\ GRZQZDUGV ERG\ GRUVDOO\ GDUNHQHG LQ SUHVHUYHGÀVK
DISTRIBUTION. Mediterranean, Red Sea to south coast of Oman, Madagascar.
COMPARISONS. Western Indian Ocean species of the japonicus group (Tables 3, 4, Plates 1, 3): Upeneus pori differs from U. guttatus LQYVPRVWO\SHFWRUDOÀQ UD\VPRUHJLOOUDNHUV²EURZQRUJUH\EDUVRQYHQWUDO KDOI VLGH RI ORZHU FDXGDOÀQ OREH YV ² UHG SDUWO\
KLGGHQ EDUV DORQJ YHQWUDO PDUJLQ RI ORZHU FDXGDOÀQ OREHRIIUHVKÀVKLQFRQVSLFXRXVEDUSDWWHUQVYVQRRU RQO\WUDFHVRIEDUVRQERWKFDXGDOÀQOREHVRISUHVHUYHG ÀVK DQG GRUVDOO\ GDUNHQHG YV SDOH EURZQ ERG\ LQ SUHVHUYHGÀVK
Non-Western Indian Ocean species of the japonicus group (Table 4): Upeneus pori differs from U. asymmetricus LQ YV SHFWRUDOÀQ UD\V VPDOOHU interorbital distance, shorter and thinner barbels, longer DQGKLJKHUDQDOÀQDQGGDUNEDUVRQYHQWUDOPDUJLQRI ORZHUFDXGDOÀQOREHYVEDUVRQO\RQLQQHUGRUVDOVLGH of lower lobe; from U. australiae it differs in more gill UDNHUVERG\DWÀUVWGRUVDORULJLQVPDOOHUVPDOOHUKDOI Fig. 3.1XPEHURIEDUVRQFDXGDO¿Q$FDXGDO¿QOHQJWKUDWLR%DQGVHFRQGGRUVDO¿QKHLJKWUDWLR&DJDLQVW
SL in 16 Upeneus oligospilus and 16 U. tragula; WKHFRPELQDWLRQRIFDXGDO¿QOHQJWKUDWLRDQGFDXGDO¿QEDU number (D) allows size-independent distinction between the two species.
ERG\ GHSWK DW DQDOÀQ RULJLQ FDXGDO SHGXQFOH PRUH compressed and less deep, smaller head (both vertically and horizontally, with smaller interorbital distance), VKRUWHU DQGWKLQQHU EDUEHOV DQGGDUN EDUV RQYHQWUDO PDUJLQRIORZHUFDXGDOÀQOREHYVEDUVFURVVLQJHQWLUH lower lobe; it differs from U. francisiLQIHZHUJLOOUDNHUV GHHSHU ERG\ DW DQDOÀQ RULJLQ KHDG WKURXJK H\H OHVV deep, shorter head and postorbital length, larger orbit, VKRUWHU MDZV ORQJHU VHFRQG GRUVDOÀQ EDVH VKRUWHU SDLUHGÀQVDQGFDXGDOEDUVLQPRVWFDVHVZHOOUHWDLQHG on preserved material; it differs from U. japonicus in shorter head length, shorter barbels and shorter caudal SHGXQFOHORQJHUFDXGDOÀQVKRUWHUSHFWRUDOÀQORZHU GRUVDOÀQVDQGEDUVRQORZHUFDXGDOÀQOREHSUHVHQW and it differs from U. parvus LQ YV SHFWRUDOÀQ UD\V IHZHU ODWHUDOOLQH VFDOHV ERG\ DW DQDOÀQ RULJLQ and head less deep, shorter jaws, shorter barbels and shorter interdorsal distance, longer caudal peduncle, ORQJHU DQG KLJKHU DQDO ÀQ VKRUWHU SHFWRUDO ÀQV DQG EDUV UHVWULFWHG WR YHQWUDO PDUJLQ RI ORZHU FDXGDOÀQ lobe vs bars crossing entire lobe.
REMARKS. Comparison of colour patterns with the Red 6HDSRSXODWLRQLVUHVWULFWHGWRSUHVHUYHGÀVKGXHWRWKH DEVHQFH RI SKRWRJUDSKV RI OLYH RU IUHVK ÀVK IURP WKH Western Indian Ocean proper.
7KLV LV WKH ÀUVW UHFRUG RI Upeneus pori in the southWestern Indian Ocean. The species attains 19 cm SL; maximum reported depth is 45 m.
Upeneus suahelicusVSQRY
(Tables 3, 8; Plates 2, 3)
Holotype: SAIAB 13948, male, 101 mm, Kenya, off 0DOLQGL59Dr. F. Nansen3&+·6·
E, collected by P.C. Heemstra, December 1980.
Paratypes:Kenya: 6$,$% PP RII 0DOLQGL
59 Dr. F. Nansen 3&+ · 6 · (
0R]DPELTXH: SAIAB 82805, 135 mm, Lipobane HVWXDU\ ··· 6 ··· ( 6$,$%
PP/LSREDQHHVWXDU\···6···(6$,$%
² PP RII 4XHOLPDQH P 6$,$%
PP59Dr. F. Nansen0·6
·(PGHSWKERWWRPWUDZO6$,$%
PP 59Dr. F. Nansen 0 ¶6 ·
( P ERWWRP WUDZO 8610 PP 3RQWD Mahone-Inhaca; 6RXWK $IULFD: SAIAB 11478, 101 PP'XUEDQ6(6$,$%²PP .ZD=XOX1DWDO5LFKDUGV%D\···6·µ(
45 m depth.
Non-types: HUJ 8555, 124 mm, Massawa, Eritrea (Red 6HD 6$,$% PP 59Dr. F. Nansen M07-
·6·(ERWWRPWUDZOGHSWKP 0R]DPELTXH6$,$%PP.ZD=XOX1DWDO 5LFKDUGV %D\ ··· 6 ··· ( P GHSWK (South Africa).
DIAGNOSIS 'RUVDO ÀQV 9,,, SHFWRUDO ÀQV ² JLOO UDNHUV ² ² ² ODWHUDOOLQH VFDOHV ² ERG\ GHSWK DW ÀUVW GRUVDOÀQ RULJLQ ² 6/
ERG\ GHSWK DW DQDOÀQ RULJLQ ² FDXGDOSHGXQFOH GHSWK²PD[LPXPKHDGGHSWK²KHDGGHSWK WKURXJK H\H ² KHDG OHQJWK ² RUELW OHQJWK
² XSSHU MDZ OHQJWK ² EDUEHO OHQJWK ² FDXGDOÀQ OHQJWK ² DQDOÀQ KHLJKW SHOYLF ÀQOHQJWK²SHFWRUDOÀQOHQJWK²ÀUVWGRUVDO ÀQ KHLJKW ² VHFRQG GRUVDOÀQ KHLJKW ² 6/
WRWDOQXPEHURIEDUVRQFDXGDOÀQ²²SDOHEURZQ EDUVRQXSSHUFDXGDOÀQOREHDQG²RIVDPHFRORXU RQ ORZHU FDXGDOÀQ OREH WKH WKUHH SUR[LPDO EDUV RQ XSSHU FDXGDOÀQ OREH VOLJKWO\ FXUYHG FDXGDOÀQ EDUV and white interspaces nearly equal in width; all bars HQWLUHO\UHWDLQHGRQSUHVHUYHGÀVKWZR\HOORZRUSDOH EURZQ ODWHUDO ERG\ VWULSHV RQ IUHVK ÀVK RQH FORVH WR ODWHUDO OLQH UHDFKLQJ IURP EHKLQG KHDG WR FDXGDOÀQ base; the other stripe below, starting behind pectoral- ÀQEDVHÀUVWGRUVDOÀQWLSGDUNDOVRLQSUHVHUYHGÀVK barbels white; head and body silvery white laterally DQGUHGGLVKEURZQGDUNHUGRUVDOO\LQIUHVKÀVKEHOO\
ZKLWH LQ SUHVHUYHG ÀVK ERG\ SDOH EURZQ VRPHWLPHV GDUNHUGRUVDOO\
DESCRIPTION. Measurements as % SL and counts are given in Table 8; morphometric data as ratios of SL IRU KRORW\SH GDWD IRU SDUDW\SHV LQ EUDFNHWV ERG\
PRGHUDWHO\GHHSERG\GHSWKDWÀUVWGRUVDOÀQRULJLQ >²@ DQG DW DQDOÀQ RULJLQ >²@ KHDG OHQJWK>²@VLPLODUWRPD[LPXPGHSWKRIERG\
DQG FDXGDOÀQ OHQJWK >²@ KHLJKW RI VHFRQG GRUVDOÀQ>²@VLPLODUWREDUEHOOHQJWK>²
@ SHFWRUDOÀQ OHQJWK >²@ FOHDUO\ ORQJHU WKDQ SHOYLFÀQ OHQJWK >²@ FDXGDOSHGXQFOH GHSWK>²@ODUJHUWKDQRUELWOHQJWK>²@
Fresh colour (Plate 2): Head and body silvery white ODWHUDOO\GRUVDOVXUIDFHGDUNUHGGLVKRUUHGGLVKJUH\
belly white; with two pale brown or yellow lateral body VWULSHVWKHXSSHUVWULSHIURPKHDGWRFDXGDOÀQEDVH anteriorly well below lateral line, crossing line below VHFRQGGRUVDOÀQDQGFRQQHFWLQJWRSUR[LPDOPRVWEDU RI XSSHU FDXGDOÀQ OREH WKH ORZHU VWULSH QDUURZHU VWDUWLQJ EHKLQG SHFWRUDOÀQ EDVH DQG FRQQHFWLQJ WR SUR[LPDOPRVW EDU RI ORZHU FDXGDOÀQ OREH LQ RQH RI the two photographed specimens a third, indistinct, reddish-brown stripe, separated dorsally by a series of pale spots can be seen, extending dorsolaterally from KHDG WR EHKLQG VHFRQG GRUVDOÀQ EDVH XSSHU OREH RI FDXGDO ÀQ ZKLWH RU SDOH JUH\ ZLWK RU SDOH EURZQ EDUV WKH ÀQOREH WLS GDUN WKH SUR[LPDO WR EDUV VOLJKWO\ FXUYHG ORZHU FDXGDOÀQ OREH ZKLWH RU SDOH JUH\ZLWKIRXUSDOHEURZQEDUVWKHÀUVWWZRSUR[LPDO ones slightly curved, mostly uniform in width, and the OREHWLSSDOHRUGDUNSDOHLQWHUVSDFHEHWZHHQEDUVRI similar width or wider than bars on upper lobe, and RIWHQQDUURZHURQORZHUOREHVSLQRXVGRUVDOÀQZKLWH to pale grey, with two pale brown horizontal stripes,
WKH ORZHU VHSDUDWHG IURP ÀQ EDVH E\ D QDUURZ SDOH interspace, the upper stripe separated from the lower RQHDQGWKHEODFNÀQWLSE\DSDOHLQWHUVSDFHRIVLPLODU width to the stripe and the tip, width about or less than KDOIRUELWGLDPHWHUVRIWGRUVDOÀQVLPLODUEXWZLWKWKUHH pale brown horizontal stripes, the lower one connected WRWKHÀQEDVHDQGWKHXSSHUPRVWRQHFRYHULQJWKHWLS DQDO DQG SHOYLF ÀQV K\DOLQH ZLWK UD\V SDOH \HOORZ RU ZKLWHDWWKHÀQEDVHSHFWRUDOÀQVK\DOLQH
Preserved colour (Plate 3): Head and body uniformly pale brown, with white silvery or pale yellow pigmentation on ventral side and on head, with large white colour SDWFKHV RQ RSHUFXODU PDUJLQ DQG EHORZ H\HV %ODFN ÀUVW GRUVDOÀQ WLS GLVWLQFW EDUV RQ ERWK FDXGDOÀQ OREHVSDOHRUGDUNEURZQSHFWRUDOSHOYLFDQGDQDOÀQV transparent.
DISTRIBUTION. Kenya to South Africa, Eritrea (Red Sea) ETYMOLOGY: The name suahelicus is derived from the
$UDELF ¶VDZDKLO· PHDQLQJ ¶FRDVW· DQG UHIHUV WR WKH occurrence of this species off the East African coast.
In a large part of this area the lingua franca is Swahili, which has the same derivation.
COMPARISONS. Western Indian Ocean species of the vittatus group (Tables 3, 8, Plates 1-3, Fig. 4): Upeneus suahelicus differs from U. davidaromi in deeper body and caudal peduncle, head through eye less deep and smaller suborbital depth, shorter head and snout, ORQJHUSHFWRUDOÀQVWDOOHUÀUVWGRUVDOÀQODWHUDOERG\
VWULSHV SUHVHQW DQG FDXGDOÀQ EDUV PRUH XQLIRUPO\
coloured; it differs from U. indicusLQIHZHUJLOOUDNHUV DQGODWHUDOOLQHVFDOHVKDOIERG\GHSWKDWÀUVWGRUVDOÀQ origin less deep, larger orbit, longer anal and pectoral ÀQVDQGODWHUDOKHDGDQGERG\FRORXUVLOYHU\ZKLWHYV reddish; from U. mascareinsis it differs in deeper body DQG FDXGDO SHGXQFOH ORQJHU SHFWRUDO ÀQV FDXGDOÀQ bars more uniformly coloured and more regularly sized on lower lobe, lateral body stripes present, and head and body colour pale brown vs reddish laterally;
from U. supravittatus LW GLIIHUV LQ IHZHU JLOO UDNHUV shorter barbels and head, shorter distances from tip RI VQRXW WR SDLUHG ÀQ RULJLQV VKRUWHU SHFWRUDO ÀQV KLJKHUVHFRQGGRUVDOÀQDQGERG\FRORXUVLOYHU\ZKLWH vs brassy (for further results of comparisons, see also next paragraph); and from U. vittatus it differs in fewer ODWHUDOOLQH VFDOHV ORQJHU SHFWRUDO ÀQV KLJKHU VHFRQG GRUVDO ÀQ FDXGDOÀQ EDUV PRUH XQLIRUPO\ FRORXUHG DQGEODFNÀUVWGRUVDOÀQWLSVPDOOHU
BecauseUpeneus suahelicus and U. supravittatus are rather similar in body shape and overlap slightly in all characters, a PCA of 40 morphometric variables from all specimens studied (n=37) was conducted, which VKRZHG D FOHDU GLVWLQFWLRQ ZKHQ WKH ÀUVW SULQFLSDO component was plotted against total number of gill UDNHUV )LJ 0D[LPXP KHDG GHSWK KHDG OHQJWK
GLVWDQFHVIURPVQRXWWRSDLUHGÀQDQGERWKGRUVDOÀQ origins, barbel length, orbit depth, postorbital length, ORZHUMDZOHQJWKDQGSHFWRUDOÀQOHQJWKZHUHDPRQJ the variables contributing most to the 22 % of total YDULDQFHH[SODLQHGE\WKHÀUVWSULQFLSDOFRPSRQHQW
Non-Western Indian Ocean species of the vittatus group (Table 8): Upeneus suahelicus differs from U.
subvittatus in less deep caudal peduncle, shorter head, VQRXW ORZHU MDZ DQG EDUEHOV KLJKHU ÀUVW GRUVDO ÀQ ODWHUDO ERG\ VWULSHV SUHVHQW LQ IUHVK ÀVK DQG PRUH XQLIRUPFDXGDOÀQEDUSDWWHUQ
REMARKS. The specimen of U. suahelicus from the
southern Red Sea (HUJ 8555) has smaller eyes (see 7DEOHDQGWKHFDXGDOÀQEDUFRORXUDWLRQLVPRUHIDLQW compared to the other specimens examined, which all originate from further south along the east African coast.
This specimen was listed among the material examined in an earlier account of Upeneus vittatus by Ben-Tuvia and Kissil (1988). These authors also provided a live colour description based on a photograph of a specimen collected in the Red Sea which, however, was not SXEOLVKHGVHHDOVRWKHUHPDUNVVHFWLRQRIWKHDFFRXQW ofU. vittatus further below).
Upeneus suahelicus attains 13.5 cm; depth range is
²P
Upeneus sulphureus&XYLHU
7DEOHV3ODWHV²
Upeneus sulphureus &XYLHU LQ &XYLHU 9DOHQFLHQQHV 1829: 450 (type locality: Antjer, western Java);
Kumaran & Randall 1984: no page number, Plate III (colour photograph); Ben-Tuvia & Kissil 1988:
)LJEODFNDQGZKLWHSKRWRJUDSK5HG6HD Fig. 4. First principal component based on 40
morphometric characters against total number of gill rakers for Upeneus suahelicus andU.
supravittatus.
