Journal of the
Ocean Science Foundation
2015 Volume 18
Randall, J.E. & Victor, B.C.
Descriptions of thirty-four new species of the ish genus Pempheris (Perciformes:
Pempheridae), with a key to the species of the western Indian Ocean
Published December 24, 2015 by the Ocean Science Foundation © 2015 OSF, Inc.
ISSN: 1937-7835
Online edition at http://oceansciencefoundation.org/josf.html
Printed copies available at UC Libraries and free from OSF, 4051 Glenwood, Irvine CA
Cover photo: Pempheris sp., Watamu, Kenya; © Dawn Goebbels
Printing sponsored by Walsh Paper Distribution, Inc. of Westminster, CA
p. 1
Editor: Benjamin Victor
Editorial Advisory Board
John E. Randall
Gerald R. Allen
Sergey V. Bogorodsky
Published by
the Ocean Science Foundation, Inc.
George Walsh, President
OSF Board of Directors
Chris Caldow
Michael Orencia
Fernando Rivera
Benjamin Victor
Anjalee Gonyea, Ofice Manager
Jessica Root, Senior Project Manager
Autumn Gries, Project Manager
Leah Smith, Project Manager
© 2015 OSF, Inc.
Descriptions of thirty-four new species of the ish genus Pempheris
(Perciformes: Pempheridae), with a key to the species
of the western Indian Ocean
JOHN E. RANDALL
Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA
E-mail: jackr@hawaii.rr.com
BENJAMIN C. VICTOR
Ocean Science Foundation, 4051 Glenwood, Irvine, CA 92604
and Guy Harvey Research Institute, Nova Southeastern University,
8000 North Ocean Drive, Dania Beach, FL 33004
E-mail: ben@coralreefish.com
Abstract
The following thirty-four new species of the pempherid ish genus Pempheris are described from the western
Indian Ocean, raising the total number for the region to 47 species: P. andilana from northwest Madagascar; P.
argyrea from the Seychelles; P. bineeshi from southeast India; P. bruggemanni from Réunion and Mauritius; P.
connelli from KwaZulu-Natal, South Africa; P. ellipse from the south coast of Oman; P. hadra from the Republic
of the Maldives; P. heemstraorum from Rodrigues, Mascarene Islands; P. hollemani from southeast Madagascar;
P. ibo from Mozambique and South Africa; P. kruppi from the Gulf of Aden and Socotra; P. kuriamuria from the
south coast of Oman; P. leiolepis from Chumbe Island, Zanzibar; P. megalops from the Seychelles; P. micromma
from the south coast of Oman; P. muscat from the Gulf of Oman; P. orbis from the Red Sea; P. pathirana from Sri
Lanka; P. peza from southern Mozambique; P. rochai from the south coast of Oman; P. rubricauda from northwest
Madagascar; P. sergey from the Red Sea; P. shimoni from Kenya; P. shirleen from the Red Sea; P. smithorum from
Zanzibar; P. tau from the Red Sea; P. ternay from the Seychelles; P. tilman from the Gulf of Aqaba, Red Sea; P. tiran
from the Strait of Tiran, Red Sea; P. trinco from Sri Lanka; P. viridis from the Red Sea; P. wilsoni from the Gulf of
Oman; P. xanthomma from the Gulf of Aden (Yemen); and P. zajonzi from the Socotra Archipelago. Many of the
new species are described from single specimens or one location, indicating a need for more collecting to resolve
the taxonomy of this group: we emphasize that some decisions are tentative and apply to the state of collections
at present. The barcode mtDNA COI sequences for 21 of the 47 WIO species are analyzed, showing some deep
divergences between species, but also several sibling species or species complexes that apparently share COI
sequences. The genetic analysis is also limited by small sample sizes; more extensive genetic surveys are required
to conirm the results. An appendix includes the description of the hybrid P. malabarica × P. russellii, a discussion
of a complex of Pempheris in St. Brandon’s Shoals in need of further study, and underwater photographs of
potential new species of Pempheris for which we have no specimens.
Key words: taxonomy, ichthyology, systematics, coral-reef ishes, sweepers, DNA barcoding.
1
Introduction
The ish family Pempheridae, with the common name sweepers, consists of two genera, Pempheris Cuvier
1829, with 35 previously described valid species, and Parapriacanthus Steindachner 1870, with ive species
(Eschmeyer & Fricke 2015). These ishes are found in all tropical to warm temperate seas, except the eastern
Paciic Ocean. They are characterized by a compressed, rhomboid-shaped body; very large eyes; short snout; the
dorsal in near the middle of the body, with VI spines and 9 (rarely 10) rays; and a very long, low anal in with III
spines and 35–48 rays. They are unusual in having two layers of scales, those of the outer layer larger and very
deciduous. These ishes typically form aggregations in caves or below ledges on reefs by day, and emerge from
shelter at night to feed near the surface on zooplankton.
The South African Institute for Aquatic Biodiversity (SAIAB) will soon publish a series of volumes on all the
coastal ishes of the western Indian Ocean (WIO), including the Red Sea and the Persian Gulf. Most of the family
accounts have been written by author(s) recognized as authorities of the group. When it was determined that the
family Pempheridae needed a more comprehensive review, the irst author, as one of the editors of the SAIAB
volumes, volunteered to prepare it. He enlisted the collaboration of the second author for the latter’s expertise on
the molecular taxonomy of coral-reef ishes. The family is now the last needed before the volumes can be published.
Pempheris is a particularly troublesome genus for taxonomists, since the species’ morphology varies little, their
few distinctive markings are frequently shared, their meristics broadly overlap, their outer scales are deciduous,
and they are not frequently collected or targeted in surveys. As a result, most of the literature prior to 2013 on the
genus for the western Indian Ocean is based on misidentiications. Of the checklists and guidebooks that have
been published in the last 30 years that list species of Pempheris of the western Indian Ocean, none have any
species of Pempheris correctly identiied. The irst author’s book Coastal Fishes of Oman, published in 1995,
will serve as an example. One species of Pempheris is illustrated on page 244 by two color igures identiied as
P. vanicolensis. The irst photograph (Fig. 637) is P. convexa, described by the authors in 2014. The second (Fig.
638) is an underwater photograph of a species of Pempheris in the schwenkii complex, clearly not P. vanicolensis
(a species known from the Andaman Sea and Paciic Ocean). An additional species is illustrated by Fig. 639, and
identiied only as Pempheris sp. It should have been labeled P. malabarica (however, we suspect that more than
one species may be presently included in nominal P. malabarica).
Our rsearch on the genus began in 2013 with the description of the new Indian Ocean species Pempheris
lavicycla, which had long been misidentiied as P. vanicolensis (Randall et al. 2013). For the irst time, mtDNA
sequencing was used in Pempheris to assist in differentiating similar-appearing specimens. Subsequently, Mooi
& Randall (2014) described P. bexillon from the southwestern Indian Ocean, Randall & Bineesh (2014) clariied
the species from India (with one new species), and Randall & Victor (2014) described four new species from
Oman and the African coast. The Pempheris species of the Red Sea have also been recently revised: Koeda et al.
(2014) described the new species P. tominagai, but then mistakenly listed three other species which do not occur
in the Red Sea. Randall et al. (2014) corrected Koeda et al. (2014) and concluded that there were three Red Sea
species. This was conirmed by Azzurro et al. (2015), who documented the invasion of the Mediterranean by P.
rhomboidea and recorded P. lavicycla and P. tominagai as the two other Red Sea species. Only ive of the now
47 species of Pempheris in the western Indian Ocean (including India and Sri Lanka) were described before 2013
(all before 1889), one was added in 2013, seven were added in 2014, and 34 more are described herein.
An appendix includes the description of the hybrid Pempheris malabarica × P. russellii from Pakistan;
underwater photographs of unidentiied species of Pempheris from the western Indian Ocean are also shown
in the hope that specimens can be collected, including at least eight probable undescribed species of Pempheris
represented by underwater photographs taken by Dawn Goebbels in the Watamu Marine National Park, Kenya;
and a discussion of a complex of Pempheris in St. Brandon’s Shoals in need of further study.
Little is known of the early life history of Pempheris, but their larvae are known to settle at a particularly small
size for reef ishes, as small as 6 mm SL (Leis & Carson-Ewart 2000). This correlates with a relatively short larval
life and reduced dispersal, which would promote local adaptation and the development of species complexes in
areas with widely dispersed reef habitats such as the broad western Indian Ocean region. While checking the
sex of specimens of Pempheris, we have noticed mature gonads in ish of surprisingly small size for the species,
as well as a very high percentage of ish in spawning condition, indicating year-round spawning. Underwater
2
photographs and our observations of aggregations of Pempheris often reveal more than one species. Pempheris
are broadcast spawners, which could facilitate hybridization and a degree of introgression that may slow the
divergence of mitochondrial DNA lineages as species complexes evolve. These speculations require additional
evidence, but may account for the high diversity in the genus in the western Indian Ocean and the frequent sharing
of mitochondrial haplotypes within some species complexes.
Many of the species of Pempheris described below are without enough specimens for adequate description
(nine from single specimens!), and most without knowledge of the life color. We hope that this publication,
admittedly preliminary, will aid in acquiring more material of this troublesome genus.
Materials and Methods
Few species of ishes have proven more dificult to classify than those of the genus Pempheris. Most are
morphologically very similar, the color is often a near-uniform copper, bronze or silver, and the dorsal in is either
broadly black-tipped, or the anterior margin black, or both. Of the ins, only the number of anal-in soft rays is of
diagnostic value, but the range is broad (Table 1); therefore, many specimens would be needed for this count to
have value. The last segmented ray in the dorsal and anal ins is divided at its base: it is counted as a single ray.
Counts of pectoral-in rays include the rudimentary upper ray and vary from 16 to 19, usually 17 or 18; they can
be of modal value only when many specimens are available. The pectoral-in rays of both sides are counted but
presented as separate counts (Table 1). The ins of museum specimens of Pempheris are often damaged, especially
the dorsal and caudal ins.
Because the large scales of the outer layer are so easily shed, the only viable scale count is that of the more
adherent small scales of the lateral line (Table 2); again, many specimens are needed to obtain the deinitive range.
The lateral-line of species of Pempheris continues to the end of the caudal in; the lateral-line scales are counted
to the base of the caudal in. Occasional lateral-line scales on some specimens were found with two sensory pores.
In such cases, the number of scales is counted, not the pores. Scales with two pores usually have two narrow
columns of scales above. These double columns are more easily seen than the pores. The species of Pempheris
have both ctenoid and cycloid scales. Whether the scales are ctenoid or cycloid, and on what parts of the body
they are found, are also of importance for species identiication. All species of the genus have a midventral row of
small scales that bear a median ridge, and all have two rows of small, strongly ctenoid scales to each side of the
median ventral row. The scales on the side of the chest (deined as above the ventral two rows of scales, below
the base of the pectoral in, posterior to the margin of the operculum, and anterior to a line from the rear base of
the pectoral ins to the origin of the pelvic ins) may be ctenoid or cycloid, or divided into a ventroanterior ctenoid
part and a dorsoposterior cycloid part. This is often of diagnostic importance when all of the scales of the outer
layer are present on the side of the chest (too often they are not). All species of Pempheris have ive longitudinal
rows of ctenoid scales dorsally on the body posterior to the dorsal in; the lower row on each side is only about
half ctenoid. If any part of a scale is ctenoid, it is regarded here as ctenoid. Depending on the species, the scales
may be either ctenoid or cycloid on the nape, below the dorsal in, on the mid-side of the caudal peduncle, and on
the chest. The caudal peduncle has the highest loss of scales of any part of the body, so one can rarely determine
whether the scales on the side of the peduncle are cycloid or ctenoid.
Gill-raker counts are important meristic data, in general, but the range is also broad. The counts are divided
into the dorsal and ventral parts, with the raker at the angle included in the ventral part. However, only the total
count is recorded in the species accounts and Table 3. The low, rounded ventroposterior margin of the preopercle
usually has one or two lat triangular spines and thin, lexible, translucent projections of variable shape, some with
small holes. There is so much intraspeciic variation with this ridge that we have not used it for species identiication; also it is normally covered by scales.
The species of Pempheris have steeply oblique jaws, the edge of the maxilla varying from 59–75° to the
horizontal axis of the body. The angle is a useful diagnostic character. It is measured by protractor at the ventral
edge of the maxilla with the jaws slightly open. The horizontal axis of the body is taken from the tip of the snout
through the center of the eye to a point at the middle of the ventral half of the caudal peduncle. The teeth in the
jaws are small, but their features are often important for species identiication. Very small teeth are consistently
3
4
present on the vomer and palatines, but they are of little diagnostic value. All specimens of Pempheris we have
examined have a close-set pair of vertically oval nostrils directly anterior to the orbit.
Most of the specimens of Pempheris examined for this study are from the ish collections of the Bishop
Museum, Honolulu (BPBM); South African Institute for Aquatic Biodiversity, Grahamstown, South Africa
(SAIAB); and the United States National Museum of Natural History, Washington, D.C. (USNM). We have
also obtained loans from the Natural History Museum, London (BMNH); California Academy of Sciences, San
Francisco (CAS); Australian National Fish Collection at Hobart, Tasmania (CSIRO); Hebrew University of
Jerusalem (HUJ); King Abdulaziz University Marine Museum, Jeddah (KAUMM); Muséum national d’Histoire
naturelle, Paris (MNHN); National Bureau of Fish Genetic Resources, Kochi, India (NBFGR); Natural History
Collection of Yemen (NHCY; now curated at SMF); Phuket Marine Biological Centre, Phuket, Thailand
(PMBC); Senckenberg Museum, Frankfurt (SMF); Steinhardt Museum of Natural History, Tel Aviv University
(SMNHTAU); the Université de La Réunion, St. Denis (URUN); and the Zoological Survey of India, Kozhikod
(formerly Calicut), India (ZSI/CLT).
Data in parentheses in the descriptions refer to paratypes. Lengths of specimens are given as standard length
(SL), measured from the median anterior end of the upper lip to the base of the caudal in (posterior end of
hypural plate); head length is measured from the same anterior point to the posterior end of the opercular lap, and
upper-jaw length from the same anterior point to the posterior end of the maxilla; body depth is taken from the
origin of the anal in vertically to the internal base of the dorsal in (not always the maximum depth); body width
is measured just posterior to the opercular lap (used only for adults; juveniles are much thinner); predorsal and
prepelvic lengths have proven to be of value as identiication characters, often more important than the length of
rays of the ins, because the latter are so often broken on museum specimens, the measurements are taken from
the origin of the irst spine to the tip of the upper lip; eye diameter is the horizontal length of the bony orbit;
interorbital width the least bony width; lengths of spines and rays of median ins are measured from their extreme
base to tip of longest ray; caudal-in length is taken horizontally from the base to a perpendicular at the end of the
longest ray, however, most museum specimens have lost enough of this in to prevent taking this measurement,
The same is true to a lesser extent for the dorsal in. Pectoral- and pelvic-in lengths are the lengths of the longest
ray, taken to the extreme base. The head of ishes of the genus Pempheris is much broader (thickness) than the
strongly tapering body. When photographing a specimen, it is necessary to elevate the tail end to have the ish
in optimal focus. This results in a shorter snout in the photograph. Measurements of specimens were taken from
specimens, not photographs. Proportional measurements are rounded to the nearest 0.05.
A small number of the more recently collected specimens have been preserved for DNA sequencing studies, a
number clearly inadequate to assess the degree of genetic variation in the genus. DNA extractions were performed
with the NucleoSpin96 (Machery-Nagel) kit according to manufacturer speciications under automation with a
Biomek NX liquid-handling station (Beckman-Coulter) equipped with a iltration manifold. A 652-bp segment
was ampliied from the 5′ region of the mitochondrial COI gene using a variety of primers (Ivanova et al. 2007).
PCR ampliications were performed in 12.5 µL volume including 6.25 µL of 10% trehalose, 2 µL of ultra pure
water, 1.25 µL of 10 × PCR buffer (10 mmol/L KCl, 10 mmol/L (NH4)2SO4, 20 mmol/L Tris-HCl (pH 8.8), 2
mmol/L MgSO4, 0.1% Triton X-100), 0.625 µL of MgCl2 (50 mmol/L), 0.125 µL of each primer (0.01 mmol/L),
0.0625 µL of each dNTP (10 mmol/L), 0.0625 µL of Taq DNA polymerase (New England Biolabs), and 2 µL of
template DNA. The PCR conditions consisted of 94°C for 2 min, 35 cycles of 94°C for 30 s., 52°C for 40 s., and
72°C for 1 min, with a inal extension at 72°C for 10 min. Specimen information and barcode sequence data from
this study were compiled using the Barcode of Life Data Systems (Ratnasingham & Hebert 2007). The sequence
data is publicly accessible on BOLD and GenBank. Additional sets of comparison COI sequences were obtained
from prior studies compiled in GenBank. Sequence divergences were calculated using BOLD with the Kimura
2-parameter (K2P) model generating a mid-point rooted neighbor-joining (NJ) phenogram to provide a graphic
representation of the species’ sequence divergence.
Mitochondrial DNA lineages obtained in the BOLD database have been assigned code numbers, or BINs
(Barcode Index Numbers), to permanently name lineages. BINs are clusters calculated by an algorithm taking
into account similarities and connectivity and assessing cluster boundaries. The BINs occupied by the mtDNA
lineages for the species that have been sequenced for this paper are presented under the species title after the
Figures and Tables.
Key to the Species of the Genus Pempheris of the western Indian Ocean
1a.
Band of small teeth anteriorly in lower jaw expanded medially to a semi-circular patch,
partly exposed when mouth fully closed; the outer row of teeth in patch erect in females,
enlarged and forward-projecting in males ...................................................................................................2
1b.
No semicircular patch of teeth anteriorly in lower jaw partly exposed when mouth fully closed ...............3
2a.
Anal in with a prominent jet-black margin; gill rakers 31 or 32; lateral-line scales 62–64 (two specimens,
121 & 122 mm SL, Strait of Tiran, Gulf of Aqaba, Red Sea) .................................................P. tiran, n. sp.
2b.
Anal in without a black margin (tips of a few anterior rays blackish on some specimens); gill rakers
24–28; lateral-line scales 65–75 (Oman, India to Indonesia, Thailand, and Cambodia) ........P. malabarica
3a.
Eyes extremely large, bony diameter of orbit 6.2 in SL; head length 2.95 in SL; gill rakers 31; teeth in
upper jaw extremely small and slightly recurved, in ive or six close-set rows to each side of symphyseal
gap (one specimen, 121 mm SL, Seychelles) ...................................................................P. megalops, n. sp.
3b.
Eyes not extremely large, bony diameter of orbit of adults 6.7–9.35 in SL; head length 3.0–3.6 in SL;
gill rakers 24–31; teeth in upper jaw not extremely small, mainly in two rows ....................................4
4a.
A distinct dark-brown to black band at base of anal in, narrowing posteriorly as in narrows; dorsal in
with an anterior blackish margin which may darken distally, sometimes broadening to a large apical spot;
no large, oval, dark brown to black spot at base of pectoral in; no dark stripes along lanks (i.e. the 25
species of the schwenkii complex in the western Indian Ocean) ...............................................................23
4b.
Usually no distinct dark-brown to black band at base of anal in (exceptions either have a prominent
dark edge to the anal in [in P. bexillon below] or dark stripes along the lanks in P. kuriamuria, P.
russellii, & P. wilsoni); no blackish band on anterior margin of dorsal in; a large, oval, dark brown
to black spot at base of pectoral ins present or absent .................................................................................5
5a.
Anal in with a complete black border (broad anteriorly, progressively narrower posteriorly; can be dark
reddish in P. mangula) ….............................................................................................................................6
5b.
Anal in without a complete black border; usually dark stripes along lanks made up of darkened centers
of lateral scale rows in life (i.e. the rhomboidea complex) .....................................................................8
6a.
Dorsal in all yellow with a full-depth black posterior margin, progressively broader dorsally to in tip;
lateral-line scales 57–64 (Chagos Archipelago, Mascarene Islands, Comoro Islands, and Mozambique
Channel) …...................................................................................................................................P. bexillon
6b.
Dorsal in not all yellow, usually without a complete black posterior margin (if present, in some
P. lavicycla, iris bright yellow); lateral-line scales 53–60 ......................................................................7
5
6
7a.
A bright yellow ring around pupil of eye; an oval dark spot at base of pectoral ins; predorsal length 2.55–
2.6 in SL; gill rakers 29–33 (Red Sea, Oman, and islands of Indian Ocean south to Kenya, Seychelles and
Chagos Archipelago, east to Andaman Islands) .........................................................................P. lavicycla
7b.
No bright yellow ring around pupil of eye (can be dusky yellow); a narrow dark brown band
across base of pectoral ins; predorsal length 2.65–2.75 in SL; gill rakers 27–30 (east and SW coasts
of India, and eastward) ...............................................................................................................P. mangula
8a.
Eyes very small, the orbit diameter in adults 8.0–9.45 in SL …..................................................................9
8b.
Eyes not very small, the orbit diameter in adults 7.0–8.2 in SL ...............................................…...............10
9a.
Body not elongate, the depth 2.0–2.1 in SL; silvery gray when fresh, scales below lateral line rimmed with
yellowish brown (south coast Oman to Strait of Hormuz, Republic of Maldives) ......................P. convexa
9b.
Body elongate, the depth 2.35 in SL; gray dorsally, often with iridescence, pale green or yellow below
lateral line, scales rimmed in gray (one specimen, 116 mm SL, Oman, south coast) .....P. micromma, n. sp.
10a.
Pectoral-in rays 16 or 17; gill rakers 26–28 (3 specimens, 115–136 mm SL, northern Mozambique) …...
........................................................................................................................................................P. cuprea
10b.
Pectoral-in rays 17–19 (modally 18, except P. nesogallica with mode of 17); gill rakers 28–34 ...............11
11a.
Pectoral ins very short, the longest ray 3.65 in SL; body depth 2.05 in SL; pectoral-in rays 18 or 19
(mainly 19) (southeast coast of India) ............................................................................................P. sarayu
11b.
Pectoral ins not very short, the longest ray 2.85–3.15 in SL; body depth 2.1–2.35 in SL; pectoral-in rays
17–19 (rarely 19) ......................................................................................................................................12
12a.
Pectoral ins very long, the in length 2.9–3.0 in SL …............................................................................13
12b.
Pectoral ins not very long, the in length 3.0–3.55 in SL .................................................…...............14
13a.
Scales on chest one-half cycloid and one-half ctenoid; longest dorsal-in ray 4.05–4.25 in SL; lateral-line
scales 52–58 (Red Sea, and as an immigrant to the Mediterranean Sea) .................................P. rhomboidea
13b.
Scales on chest all ctenoid; longest dorsal-in ray 3.65–3.75 in SL; lateral-line scales 59–61 (two specimens,
95 & 112.5 mm SL, Oman) ..........................................................................................P. kuriamuria, n. sp.
14a.
Head length 3.0–3.05 in SL (Gulf of Oman) .................................................................................P. darvelli
14b.
Head length 3.1–3.45 in SL .......................................................................................................................15
15a.
Prepelvic length 2.85 in SL; head length 3.45 in SL; anal-in soft rays 45 (one specimen, 125 mm SL,
Fahal Island, Gulf of Oman) ................................................................................................P. wilsoni, n. sp.
15b.
Prepelvic length 2.45–2.7 in SL; head length 3.1–3.4 in SL; anal-in soft rays 35–44 ...................................16
16a.
Scales on side of chest entirely ctenoid; pectoral-in length 3.1–3.55 in SL .............................................17
16b.
Scales on chest divided into ventroanterior ctenoid and dorsoposterior cycloid parts; pectoral-in
length 3.0–3.15 in SL ..............................................................................................................................21
17a.
Tongue triangular basally, narrowing to a slender rectangle with rounded tip; head length 3.25–3.5 in SL
(Sind coast of Pakistan) ................................................................................................................P. russellii
17b.
Tongue entirely triangular, without a slender rectangular tip; head length 3.1–3.4 in SL .....................18
18a.
Pectoral-in length 3.3–3.55 in SL; predorsal length 2.55–2.7 in SL; lateral-line scales 52–60 ...............19
18b.
Pectoral-in length 3.0–3.4 in SL; predorsal length 2.7–2.8 in SL; lateral-line scales 55–64 ...................20
19a.
Predorsal length 2.55–2.6 in SL; prepelvic length 2.55–2.6 in SL; angle of maxilla to horizontal axis of
body 65–70°; lower jaw not or only slightly protruding when mouth irmly closed (KwaZulu-Natal to
Mozambique) ..................................................................................................................................P. eatoni
19b.
Predorsal length 2.7 in SL; prepelvic length 2.45–2.5 in SL; angle of maxilla to horizontal axis of body
61–66°; lower jaw strongly protruding when mouth irmly closed (two specimens, 126.5 & 139 mm SL,
Zanzibar) .......................................................................................................................P. smithorum, n. sp.
20a.
Front of upper jaw with an outer row of sharply conical, recurved teeth, followed inwardly by a broad zone
of progressively smaller teeth; upper lip with scattered, very small papillae; anal-in soft rays 38–43; gill
rakers 29 or 30 (14 specimens, 113–139 mm SL, Sri Lanka) ..........................................P. pathirana, n. sp.
,
20b.
Front of upper jaw with two irregular rows of sharply conical, recurved teeth, not followed inwardly by a
broad zone of small teeth; upper lip densely covered with papillae of moderate size; anal-in soft rays 37–
41; gill rakers 29–32 (16 specimens, 106.5–124 mm SL, Gulf of Aden and Socotra) .......P. kruppi, n. sp.
21a.
Pectoral-in length 3.0–3.15 in SL; anal-in soft rays 38–44; gill rakers 30–34 (Mascarene Islands,
Madagascar, and KwaZulu-Natal) ...........................................................................................P. nesogallica
21b.
Pectoral-in length 3.15 in SL; anal-in soft rays 35–41; gill rakers 28–32 ............................................22
22a.
Head length 3.3 in SL; tongue triangular, straight-sided, with a pointed tip; anal-in soft rays 35–40; gill
rakers 30–32 (5 specimens, 124–132 mm SL, east coast of Oman) ...................................P. muscat, n. sp.
22b.
Head length 3.15 in SL; tongue triangular, concave-sided, ending in a thin, rounded, lexible lap; anal-in
soft rays 37–41; gill rakers 28–31 (12 specimens, 31–133 mm SL, Shimoni, Kenya) ........P. shimoni, n. sp.
7
P. schwenkii complex below: (note P. bexillon above is genetically also in the schwenkii complex)
8
23a.
Band at base of anal in consisting of discrete black dots of variable size; head and body below lateral line
light tan when fresh, near-white ventrally and posteriorly (two specimens, 117 & 118 mm SL, southeast
Madagascar) ...................................................................................................................P. hollemani, n. sp.
23b.
Band at base of anal in uniformly dark brown to black; color not as in 23a .....................................24
24a.
Body strongly compressed, maximum body width 3.35 in body depth; pelvic ins long, 5.3 in SL (one
specimen, 127 mm SL, Rodrigues, Mascarene Islands; largest specimen of the schwenkii complex
examined) .............................................................................................................. P. heemstraorum, n. sp.
24b.
Body not strongly compressed, maximum width 2.2–3.1 in body depth; pelvic ins 5.3–7.45 in SL .......25
25a.
Body thick, maximum width 2.2 in body depth; scales on caudal peduncle ctenoid (except those of lateral
line); color when fresh bronze, except for broad central area of body enclosing pectoral ins with metallic
green scale centers; caudal in orange-red without black margins (one specimen, 118 mm SL, Republic of
Maldives) ..............................................................................................................................P. hadra, n. sp.
25b.
Body not thick, maximum width 2.35–3.1 in body depth; scales on caudal peduncle both ctenoid
and cycloid; color not as in 25a ................................................................................................................26
26a.
Lateral-line scales 43–49; straight ascending anterior part of lateral line abruptly angular below dorsal in
to remainder of lateral line that parallels dorsal body contour; color in life variable, often greenish yellow
with copper iridescence dorsally, others iridescent silvery (largest of 14 type specimens examined, 93 mm
SL, KwaZulu-Natal to Mozambique) ....................................................................................... P. ibo, n. sp.
26b.
Lateral-line scales 46–59 (only P. tominagai with fewer than 49); ascending anterior part of lateral line
bends in a gradual curve into posterior portion of lateral line; color not as in 26a ..........................27
27a.
Dorsal-in soft rays 10; a single row of projecting recurved conical teeth anteriorly in upper jaw (one
specimen, 116.5 mm SL, south coast of Oman) ...................................................................P. rochai, n. sp.
27b.
Dorsal-in soft rays 9; two or more rows of recurved conical teeth anteriorly in upper jaw .................28
28a.
Lateral line very strongly arched anteriorly, rising 3/4 orbit diameter above level of origin of lateral
line; orbit of eye forming an ellipse, the vertical height 8.8% higher than width; body depth 2.1 in SL (one
adult, 93.5 mm SL, and 15 juveniles, south coast of Oman) ............................................... P. ellipse, n. sp.
28b.
Lateral line not strongly arched anteriorly, rising at most 1/2 orbit diameter above level of
origin of lateral line; orbit of eye circular or only slightly higher than wide; body depth 2.15–2.85 in SL
(except P. connelli with body depth 2.0–2.25 in SL) ............................................................................29
29a.
Caudal in bright red-orange with broad black dorsal, ventral, and posterior borders …..........................30
29b.
Caudal in not bright red-orange with broad black borders ..................................................................31
30a.
Pectoral-in length 3.0 in SL; lateral-line scales 51; maxilla forming an angle of 60° to horizontal
axis of body; dark brown to black band at base of anal in narrow, one-third pupil diameter in width
at mid-length; anal in with blackish margin; pectoral-in rays 17 (one specimen, 92 mm SL, Mahé,
Seychelles) ...........................................................................................................................P. ternay, n. sp.
30b.
Pectoral-in length 3.4–3.6 in SL; lateral-line scales 55–57; maxilla forming an angle of 69–73°
to horizontal axis of body; dark brown to black band at base of anal in broad; anal in without a
blackish margin; pectoral-in rays 17 or 18 (modally 18)(5 specimens, 68.5–77 mm SL, northwestern
Madagascar) ...............................................................................................................P. rubricauda, n. sp.
31a.
Body depth 2.0–2.25 in SL; gill rakers 29–32; color in life olivaceous dorsally, brassy yellow below lateral
line, the scales with indistinct brown edges; largest specimen examined, 126.5 mm SL (39 specimens, 53–
126.5 mm SL, KwaZulu-Natal, South Africa) ..................................................................P. connelli, n. sp.
31b.
Body depth 2.2–2.7 in SL (except range down to 2.15 in P. xanthomma); gill rakers 25–30 (except
P. orbis and P. tau); largest specimens usually less than 110 mm SL; color not as in 31a .............32
32a.
Lateral-line scales 58–63; pectoral-in length 3.0–3.1 in SL; color of body of adults when fresh
dusky silver–gray, the scales below lateral line rimmed with brownish orange; caudal in with
dark brown band over lateral line, on upper and lower margins, but none on posterior margin
(6 specimens, 83–118 mm SL, Réunion and Mauritius) ......................................P. bruggemanni, n. sp.
32b.
Lateral-line scales 48–59; pectoral-in length 3.0–3.9 in SL; color not as in 32a .................................33
33a.
Teeth at front of upper jaw in three or more irregular, close-set rows, more recurved inwardly ..............34
33b.
Teeth at front of upper jaw in two rows, the outer slightly recurved, the inner larger and more
strongly recurved ......................................................................................................................................42
34a.
Head length 3.0 in SL; dorsal proile of head straight (one specimen, 100 mm SL, Chumbe Island, Zanzibar)
............................................................................................................................................P. leiolepis, n. sp.
34b.
Head length 3.05–3.35 in SL; dorsal proile of head convex (except for P. viridis where proile
nearly straight) ........................................................................................................................................35
35a.
Body thin, the width 2.75–2.9 in body depth ….........................................................................................36
35b.
Body not thin, the width 2.4–2.7 in body depth ............................................................................................37
9
10
36a.
Body very deep, the depth 2.2–2.25 in SL; predorsal length 2.4–2.55 in SL; pelvic fins not short,
reaching anus, their length 5.7–6.8 in SL (10 specimens, 91–103 mm SL, northwestern Madagascar) ..
..........................................................................................................................................P. andilana, n. sp.
36b.
Body not very deep, the depth 2.4 in SL; predorsal length 2.6–2.7 in SL; pelvic ins short, not
reaching anus, their length 7.45 in SL (two specimens, 103 & 105 mm SL, Hurghada, Egypt,
Red Sea) ..............................................................................................................................P. shirleen, n. sp.
37a.
Body depth 2.65 in SL; dorsal proile of head nearly straight; gill rakers 28; color when fresh metallic green,
scale edges yellowish brown (one specimen, 79.5 mm SL, Al Lith, Saudi Arabia, Red Sea) ....P. viridis, n. sp.
37b.
Body depth 2.3–2.7 in SL; dorsal proile of head convex; gill rakers 26–33; color not as in 37a ...................38
38a.
A copper-colored ring around pupil of eye in alcohol; lateral-line scales 58–64; dark brown band at base
of anal in equal in width to pale outer part of in (fully pigmented width measured at mid-length of in) (3
specimens, 74–85 mm SL, Gulf of Aqaba) ............................................................................P. orbis, n. sp.
38b.
No copper ring around pupil of eye in alcohol; lateral-line scales 46–59; dark brown band at
base of anal in not equal in width to pale outer part of in .....................................................39
39a.
Dark brown band at base of anal in broader than pale outer part of in (measured at mid-length of in);
caudal in without a broad black posterior border (dark border present only on dorsal and ventral margins
of in); scales on side of chest entirely cycloid; lateral-line scales 46–51 (Red Sea) ...............P. tominagai
39b.
Dark brown band at base of anal in narrower than pale outer part of in; caudal in with a broad
black posterior margin (also present on dorsal and ventral margins of in, but half as broad);
scales on side of chest cycloid, except for a few ctenoid scales ventrally at edge of opercle
(chest scales missing on P. tau); lateral-line scales 49–59 .....................................................................40
40a.
Prepelvic length 2.4–2.45 in SL; eyes large, orbit diameter less than 7.1 in SL in large adults; color of
body when fresh silvery below lateral line, the scale centers broadly pale blue-green in pectoral region (17
specimens, 61–102 mm SL, Seychelles) ............................................................................P. argyrea, n. sp.
40b.
Prepelvic length 2.55–2.7 in SL; eyes small, orbit diameter 7.9–8.65 in SL in adults; color not as in
40a .............................................................................................................................................................41
41a.
Prepelvic length 2.65–2.7 in SL; head length 3.25–3.3 in SL; gill rakers 30–33 (3 specimens, 95–116.5
mm SL, Gulf of Aqaba and southern tip of Sinai Peninsula) ......................................................P. tau, n. sp.
41b.
Prepelvic length 2.55 in SL; head length 3.15 in SL; gill rakers 28–30 (one 89-mm SL specimen and four
juveniles, Socotra Archipelago) ...........................................................................................P. zajonzi, n. sp.
42a.
Scales all ctenoid on nape and partly ctenoid below dorsal in; lateral line scales 50–59 (16 specimens,
75–114 mm SL, southern Mozambique) .................................................................................P. peza, n. sp.
42b.
Scales cycloid on nape and below dorsal in; lateral-line scales 46–58 .....................................................43
43a.
Prepelvic length 2.8–2.85 in SL; body depth 2.5–2.7 in SL; gill rakers 25–27; caudal in with a black
border only on posterior margin (7 specimens, 95–108 mm SL, southern India) ..............P. bineeshi, n. sp.
43b.
Prepelvic length 2.45–2.8 in SL; body depth 2.15–2.55 in SL; gill rakers 26–30; caudal in with a black
border on all outer margins (may be narrower and only blackish on upper and lower margins) ...................44
44a.
Prepelvic length 2.7–2.8 in SL; gill rakers 29 or 30; a dark brown to blackish line at base of rays
and in axil of pectoral in (3 specimens, 75–97 mm SL, Gulf of Aqaba) .............................P. tilman, n. sp.
44b.
Prepelvic length 2.5–2.65 in SL; gill rakers 26–29; a black band across base of pectoral in
below base of rays (straight ventrally and rounded dorsally) ...............................................................45
45a.
Dark brown band at base of anal in broader than pale outer part of in; predorsal length 2.7–2.8 in SL;
margin of upper jaw forming an angle of 67–70° to horizontal axis of body (11 specimens, 57–100 mm
SL, Yemen, Gulf of Aden) .............................................................................................P. xanthomma, n. sp.
45b.
Dark brown band at base of anal in narrower than pale outer part of in; predorsal length 2.6–2.75 in
SL; margin of upper jaw forming an angle of 60–68° to horizontal axis ............................................46
46a.
Body depth 2.5–2.55 in SL; head length 3.3–3.4 in SL; pectoral-in rays modally 17; ground color when
fresh greenish yellow, the scales rimmed with brown (5 specimens, 88–105 mm SL, Sharm Obhur,
Saudi Arabia, Red Sea) ........................................................................................................P. sergey, n. sp.
46b.
Body depth 2.3–2.5 in SL; head length 3.2–3.3 in SL; pectoral-in rays modally 18; ground color when
fresh silver-gray, the scales rimmed in copper (22 specimens, 75–108 mm SL, Sri Lanka) ..P. trinco, n. sp.
11
0.02
...............
P. eatoni South Africa
P. kruppi Yemen
P. kruppi Yemen
P. kruppi Yemen
P. eatoni South Africa
P. eatoni South Africa
P. sp. Madagascar
P. eatoni South Africa
P. eatoni South Africa
P. sp. South Africa
P. kruppi Yemen
P. russellii Pakistan
P. kuriamuria Oman
P. kuriamuria Oman
P. kruppi Yemen
P. eatoni South Africa
P. wilsoni Oman
P. sp. South Africa
}
P. rhomboidea Red Sea
}Bali, Indonesia
}
SE India
P. mangula
.....
.........
...... ... ... ..............
.
..
..........
}
}
}
P. bineeshi SE India
P. schwenkii Bali, Indonesia
P. hadra Maldives
}
P. tominagai Red Sea
}
P. sp. Chagos Islands
P. xanthomma Yemen
P. ternay Seychelles
}
}
P. rubricauda Madagascar
P. bexillon Réunion
}
P. ibo South Africa
P. hollemani Madagascar
}
P. connelli South Africa
P. rochai Oman
P. malabarica Pakistan
Djibouti
}
}
Red Sea
}Seychelles
P. sp. 1 South Africa
P. bruggemanni Réunion
}
P. lavicycla
Parapriacanthus ransonneti Tonga
12
DNA Barcoding Results. Mitochondrial DNA sequences can assist in decisions on the taxonomic status
of populations, but cannot be used in isolation to decide species status (Victor 2015). Although most reef ish
species show monophyletic mtDNA COI lineages more than 2% different from their nearest relatives (Steinke
et al. 2009, Ward et al. 2009), there is no clear threshold degree of divergence between the mtDNA lineages
of established species. Indeed, many cases of species that share COI sequences (the “barcode” mtDNA marker
used in the Barcode of Life project) have been documented, i.e. “phenovariant” species sensu Victor (2015).
Two species that are on their own evolutionary trajectories can share mtDNA sequences due to insuficient time
since speciation events for mitochondrial mutations to accumulate (incomplete lineage sorting) or some degree
of hybridization allowing mitochondrial haplotypes to be traded (via maternal descent, and then taking extensive
time to be eliminated by chance). In cases of phenovariant populations, a close look at the morphological,
meristic, and marking differences is required to make taxonomic decisions about species status.
Where we ind non-overlapping monophyletic sets of haplotypes (i.e. divergent lineages), it conirms that
there is some degree of reproductive isolation between populations. This alone does not deine species, and
when there are no corresponding morphological, meristic, and marking differences of a “suficient” degree,
the populations would be considered “genovariant” populations of the same species (Victor 2015). The inal
decision on species status is essentially a taxonomist’s subjective determination.
In the case of the western Indian Ocean Pempheris, there is limited information on the genetic structure of
populations and species. At present, 21 of the 47 regional species have been barcoded (sequenced for the COI
marker), however many are based on single specimens. An additional lineage occurs in South Africa based on a
sequence from an egg (P. sp. 1); it may represent one of the many unsequenced species or an additional species.
The phenetic tree based on the COI sequences of Indian Ocean populations of Pempheris is presented in
Fig. 1. Many species form deeply divergent monophyletic lineages well isolated from their nearest relatives, but
some are only minimally divergent lineages, although clearly distinct (e.g. the 1% divergence between the P.
rhomboidea complex vs. P. mangula). Several new species described here have sequences only slightly different
from their nearest relatives, but, since they are based on single sequences, the consistency of the divergence is
untested. Several more new species share identical haplotypes with their closest relatives, but those are also
mostly based on single sequences. In general, most phenovariant species documented in this study are allopatric,
suggesting that regional species complexes are in the process of diverging. In one case the phenovariants are
both from Oman (P. kuriamuria from the south coast of Oman and P. wilsoni from the northern coast in the Gulf
of Oman), but this is also based on single sequences and clearly needs to be conirmed.
The P. rhomboidea species complex appears to be the largest complex of phenovariant species documented
to date, with six putative species in the region mostly sharing haplotypes (and many more unsequenced) and
ranging from the Red Sea (and invasive in the Mediterranean), south to South Africa, and eastward to Pakistan.
The six sequenced phenovariants are mostly allopatric, but several species in the complex occupy the coastline
from Yemen to Oman; unfortunately these species are among the least well documented and this area is now one
of the more dificult places to work on the globe. The true P. schwenkii complex (excluding the distantly related
“schwenkioid” species, most of which are from the southwestern Indian Ocean) is another group that contains
sets of phenovariant species, falling into two distinctly divergent sets of species that either share haplotypes (P.
bineeshi/hadra/schwenkii) or are fractionally different (P. tominagai/xanthomma/ternay/rubricauda). The true P.
schwenkii complex in the Indian Ocean is entirely allopatric, with the larger genetic split between the far western
Indian Ocean species and the central/eastern species, i.e. from Maldives, India, to Indonesia.
It is unclear what exactly is the origin and signiicance of these shared lineages in Pempheris; it is an unusual
inding among tropical marine reef ishes that needs to be further explored. One could speculate that it may
be a result of an increased opportunity for accidental hybridization among crepuscular or nocturnal groupspawning ishes, or something to do with the presumably short larval duration of Pempheris facilitating regional
differentiation, but not complete enough to develop corresponding genetic structure. Clearly, much more
information on the genetics of this particularly dificult genus is needed.
Figure 1 (opposite). The neighbor-joining phenetic tree of COI sequences for Indian Ocean populations of Pempheris
following the Kimura two-parameter model (K2P) generated by BOLD (Barcode of Life Database). The scale bar at left
represents a 2% sequence difference. Collection locations for specimens are indicated, and the pempherid Parapriacanthus
ransonneti is used as the outgroup. GenBank accession numbers and collection data for the sequences are listed in Appendix A.
13
TABLE 1
Anal-in and pectoral-in ray counts of specimens of Pempheris of the western Indian Ocean
(pectoral-in rays counted separately on both sides)
Anal-in soft rays
35
36
37
38
39
40
41
3
5
1
4
3
2
1
1
2
5
1
3
5
16
1
1
2
3
4
2
1
11
3
3
15
1
6
P. andilana
P. argyrea
P. bexillon
P. bineeshi
P. bruggemanni
1
P. connelli
P. convexa
P. cuprea
P. darvelli
P. eatoni
1
P. ellipse
5
P. lavicycla
P. hadra
7
1
2
P. hollemani
1
5
P. kruppi
4
3
P. kuriamuria
1
2
1
6
5
3
2
1
2
4
1
4
1
P. pathirana
1
3
6
5
1
1
2
1
4
6
P. rochai
1
1
P. russellii
1
P. sarayu
P. sergey
1
1
P. shimoni
P. shirleen
2
4
4
1
2
1
1
4
1
3
3
7
1
1
1
1
2
2
P. smithorum
2
P. tau
3
1
P. xanthomma
P. zajonzi
1
2
17
18
7
21
16
4
1
16
1
5
13
12
16
10
11
49
5
3
2
4
51
26
22
12
1
2
30
5
26
2
2
6
3
2
7
10
13
9
6
1
2
2
2
1
2
4
1
2
8
5
1
4
3
3
3
2
1
2
1
2
6
1
1
1
1
5
2
4
1
1
1
1
1
1
1
2
2
7
6
1
2
1
1
5
2
8
1
6
2
3
2
1
21
2
1
13
15
1
4
3
1
1
2
P. wilsoni
14
1
2
1
1
1
P. trinco
P. viridis
16
1
1
P. orbis
P. tominagai
48
1
P. nesogallica
P. tilman
P. tiran
47
1
P, micromma
P. ternay
46
1
1
6
P. megalops
P. rubricauda
45
1
2
P. mangula
P. rhomboidea
1
1
1
10
P. malabarica
P. peza
3
44
1
P. leiolepis
P. muscat
43
1
P. heemstraorum
P. ibo
3
6
1
1
6
2
2
5
42
Pectoral-in rays
2
4
8
7
3
2
4
2
6
1
8
17
1
2
39
10
2
2
22
29
3
2
42
25
2
10
18
4
18
19
45
1
6
19
2
2
13
1
2
2
3
2
23
9
19
2
1
38
5
1
1
3
4
4
6
2
TABLE 2
Lateral-line scale counts of specimens of Pempheris of the western Indian Ocean
43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
1 1 2 1 1 2 2
2 2 3 3 3 1 1 1
2 3 4
1 1 2 2 2
2 1
1 4 7 9 5 2 5
1
1 1
1
1
1 1
1 1
1 5 4 7 9 10 7 7
1 1 2 4 3 3 1 1
1 1 2 8 8 4 4
P. andilana
P. argyrea
P. bexillon
P. bineeshi
P. bruggemanni
P. connelli
P. convexa
P. cuprea
P. darvelli
P. eatoni
P. ellipse
P. lavicycla
P. hadra
1
3
3
1
1
P. hollemani
1
1 1 2 2 4 3 1
1 1 3 2 1 2 1 2 2 1
1
1
P. kruppi
P. kuriamuria
P. leiolepis
2
1
P. heemstraorum
P. ibo
3 2 1 2 1
1
2 3 4 7 8 5 5 2 3 1 1
P. malabarica
P. mangula
P. megalops
P, micromma
P. muscat
P. nesogallica
P. orbis
P. pathirana
P. peza
P. rhomboidea
P. rochai
P. rubricauda
P. russellii
P. sarayu
P. sergey
P. shimoni
P. shirleen
P. smithorum
P. tau
P. ternay
P. tilman
P. tiran
P. tominagai
P. trinco
P. viridis
3 4 5 2 1
1
1
1
1 2 1
1 2 1 3 6 4
1
2 2 1
1 2 1
2 1 3 3 2
2 2 5 6 6 5 5
P. zajonzi
1
2 1 1
1
1 2 2 1
1
1
1
3 1 1
1 5 4 1 2
2
1
1
1
2 1 1
2 1 2 3 2 1
1
1
1
1
1
1
1
1
1 1
1
1
1 2 1 1
1
1 1 2 2 4
1
1
4 4 1 1
1
P. wilsoni
P. xanthomma
1
1 3 5 4 5 1 3 1 3
1 1
2 1
15
TABLE 3
Gill-raker counts of specimens of Pempheris of the western Indian Ocean
24
25
P. andilana
P. argyrea
26
1
1
27
4
1
3
3
28
4
9
29
1
5
P. bexillon
2
P. bineeshi
1
P. bruggemanni
P. connelli
P. convexa
1
P. cuprea
1
5
11
1
30
31
32
33
34
1
1
3
6
6
2
14
1
7
1
2
1
14
1
11
1
10
7
3
1
P. darvelli
P. eatoni
2
P. ellipse
3
1
6
P. lavicycla
6
4
1
15
1
5
1
P. hadra
1
P. heemstraorum
1
P. hollemani
P. ibo
1
2
4
P. kruppi
7
2
1
5
2
6
1
P. kuriamuria
P. leiolepis
P. malabarica
1
11
18
P. mangula
10
3
1
1
6
P. micromma
P. muscat
P. nesogallica
P. orbis
P. pathirana
3
P. peza
P. rhomboidea
1
3
P. rochai
P. rubricauda
5
4
5
2
P. sarayu
2
P. sergey
2
9
1
2
3
4
1
9
3
3
1
10
1
3
1
4
1
1
1
1
1
3
P. shimoni
1
P. shirleen
1
2
4
2
6
9
11
2
P. russellii
2
1
4
P. smithorum
P. tau
1
P. tilman
2
1
P. tiran
P. tominagai
12
P. trinco
5
4
3
16
1
1
P. wilsoni
P. zajonzi
1
1
P. ternay
P. xanthomma
1
1
P. megalops
P. viridis
8
1
8
14
3
1
3
1
1
1
Pempheris andilana Randall & Victor, n. sp.
Figure 2, Tables 1–3.
Holotype. USNM 307628, male, 102.5 mm SL, Madagascar, Nosy Be, off Andilana Beach, near hotel at
beach, Cruise 17 of R/V Vityaz, J.R. Paxton, D.M. Cohen, & B.B. Collette, rotenone, Nov. 9, 1988.
Paratypes. USNM 435863, 5: 93–103 mm SL, & BPBM 41193, 96 mm SL, same data as holotype; SAIAB
31327, 2: 91.5–102.5 mm SL, Madagascar, off Andilana Beach Hotel, rotenone, M. Anderson, Nov. 9, 1988;
SAIAB 52862, 100.5 mm SL, Madagascar, Ambariotelo, 13.416° S, 48.366° E, rotenone, P.C. Heemstra & D.A.
Hensley, Sep. 5, 1995.
Description. Dorsal-in rays VI,9; anal-in rays III,38 (37–41); pectoral-in rays 17 or 18 (modally 18); lateralline scales 56 (52–58); gill rakers 27 (26–29); body depth 2.2 (2.2–2.25) in SL; body width 2.75 (2.75–2.9) in body
depth; head length 3.35 (3.15–3.35) in SL; eye moderate in size for the genus, orbit diameter 7.35 (7.3–8.0) in
SL; interorbital width 10.9 (11.2–11.8) in SL; mouth oblique, forming an angle of 63°(62–65°) to horizontal axis
of body; maxilla reaching posterior to a vertical at center of eye; tip of lower jaw projecting anterior to upper jaw
when mouth fully closed; teeth in upper jaw in three irregular rows medially, soon narrowing laterally to a double
row, and inally to a single row of very small but still pointed teeth in posterior ifth of jaw; teeth in lower jaw
very small and densely spaced, in a band of about four to six rows medially, where broadest, progressively more
retrorse and more pointed inwardly, the innermost largest and nearly horizontal (larger in males than females);
tongue triangular, narrowing to a small, rounded, rectangular tip; lips smooth (densely and inely papillate by
microscope); scales on nape and below dorsal in cycloid; scales on ventroanterior fourth of side of chest ctenoid,
the rest cycloid; scales dorsally on caudal peduncle ctenoid to one scale row above lateral line, cycloid below
lateral line except midventrally; origin of dorsal in at greatest body depth, the predorsal length 2.5 (2.4–2.55) in
SL; longest dorsal-in ray 3.4–3.8 in SL (broken in holotype); caudal in forked, the caudal concavity about 4 in
head length; pectoral-in length 3.2 (3.0–3.2) in SL; prepelvic length 2.45 (2.45–2.5) in SL; pelvic ins reaching
anus or extending slightly posterior, their length 6.8 (5.7–6.8) in SL. Color of body in alcohol light brown, many
scales with very narrow pale yellowish edges, becoming darker above lateral line, and much darker on nape;
chest notably lighter brown; ins yellowish, the dorsal in with a black anterior band, not becoming broader at tip;
caudal in with wide blackish margins, broadest posteriorly; dark brown band at base of anal in almost as broad
anteriorly as outer yellowish part of in, but narrower posteriorly; and pectoral ins with a blackish line centered
on upper part of in base. Color in life unknown.
Figure 2. Pempheris andilana, holotype, USNM 307628, 102.5 mm SL, Andilana Beach, Nosy Be, Madagascar (H.A.
Randall).
17
Etymology. This species is named Pempheris andilana for the beach on the island of Nosy Be, NW Madagascar,
adjacent to the reef from which the type specimens were collected. The speciic epithet is a noun in apposition.
Remarks. This new species is one of the large group of species of Pempheris that we term the schwenkii
complex, named for the irst species, P. schwenkii Bleeker, that was described from the Batu Islands off the SW
coast of Sumatra in 1855. The last twenty-ive species in the above key are from this group. They tend to be more
slender, on average, than the remaining species of Pempheris, and all have a broad, dark brown to black band at
the base of the anal in, as well as a dark brown to black anterior margin on the dorsal in that, on some species,
expands to a dark distal spot. Also they lack the lateral dark stripes on the lanks made up of darkened scale
centers characteristic of the rhomboidea complex of species. Two other members of the schwenkii complex are
known from Madagascar: P. rubricauda from the same region as P. andilana and P. hollemani from the SE of the
island: both differ in having a shorter predorsal length. No DNA sequences are available for P. andilana.
Pempheris argyrea Randall & Victor, n. sp.
Figures 3 & 4, Tables 1–3.
Pempheris schwenkii [non Bleeker] Randall & Bineesh 2014: 37.
Holotype. BPBM 21599, female, 102 mm SL, Seychelles, La Digue, North Point, rocky shore, 0–1 m,
rotenone, J.E. Randall, H.A. Randall, & D.J. Woodland, June 1, 1977.
Paratypes. SAIAB 51264, 90 mm SL, Seychelles, Mahé, 4.616° S, 55.450° E, J.L.B. Smith & M.M. Smith,
Sept. 1954; SAIAB 51265, 90.5 mm SL, Praslin, 4.316° S, 55.733° E, J.L.B. Smith & M.M. Smith, Oct. 11, 1954;
BPBM 22980, 5: 61–86 mm SL, CAS 237995, 2: 77–79 mm SL, & USNM 434841, 6: 65–91 mm SL, all with
same data as holotype; BPBM 22937, 90 mm SL, Mahé, W coast, off Residence Danzilles, 2–3 m, speared from
aggregation, J.E. Randall, April 6, 1980 (specimen in poor condition).
Figure 3. Pempheris argyrea, holotype, BPBM 21599, 102 mm, La Digue, Seychelles (J.E. Randall).
18
Description. Dorsal-in rays VI,9; anal-in rays III,38 (36–40); pectoral-in rays 17 (16–18, modally 17);
lateral-line scales 53 (50–57); gill rakers 28 (26–30); body depth 2.4 (2.3–2.5) in SL; body width 2.65 (2.55–2.7)
in body depth; head length 3.15 (3.05–3.25) in SL; dorsal proile of head strongly convex; orbit diameter 7.35
(6.9–7.75) in SL; interorbital width 11.3 (10.9–11.5) in SL; mouth oblique, forming an angle of 62°(62–65°) to
horizontal axis of body; tip of lower jaw projecting anterior to upper jaw when mouth fully closed; teeth in upper
jaw in three irregular, close-set rows medially, soon narrowing to two well-separated rows, and inally to a single
row of very small teeth on about posterior ifth of jaw; teeth in lower jaw very small and densely spaced, in a band
of four to ive irregular rows medially, where broadest, progressively more retrorse and more pointed posteriorly,
the innermost teeth largest and nearly horizontal; tongue narrowly triangular and pointed; lips smooth (dense low
papillae by microscope); scales on nape and below dorsal in cycloid; scales on ventroanterior fourth of side of
chest ctenoid, the rest cycloid; scales ctenoid on caudal peduncle only dorsally and ventrally; predorsal length
2.65 (2.55–2.7) in SL; dorsal in broken (longest ray 4.3 in SL from photograph); caudal in slightly forked, the
caudal concavity about 4–5 in head length; pectoral-in length 3.1 (3.1–3.3) in SL; prepelvic length 2.5 (2.4–2.45)
in SL; pelvic ins varying from just reaching anus to reaching anterior end of anal-in base, their length 6.0 (5.8–
6.55) in SL. Color in alcohol light brown, the scales dorsally on head and nape a little darker; lateral line light
brown with a slightly darker brown margin; ins pale yellowish, the dorsal with a broad brown anterior margin,
expanding to a spot distally; anal in with a broad dark brown band that narrows posteriorly; no dark spot at base
or axil of pectoral ins; lips dusky. Color when fresh as in Fig. 3.
Etymology. This species is named Pempheris argyrea, from Greek for silvery, in reference to its overall
silvery coloration, as noted when irst collected and photographed as a specimen.
Remarks. This species has been observed and collected only from the three main islands of the Seychelles,
Figure 4. Pempheris argyrea, aggregation, La Digue, Seychelles (J.E. Randall).
19
with La Digue selected as the type locality. It is typically seen by day in aggregations in the shelter of coral reefs,
at times in less than 1 m. One of 25 species of the broad schwenkii complex presently conirmed for the western
Indian Ocean, distinguished by relatively large eyes, low count of 36–40 anal-in soft rays, modally 17 pectoralin rays, 26–30 gill rakers, small size (largest specimen, the holotype, 102 mm SL), and silvery coloration. P.
ternay, the other member of the schwenkii complex from the Seychelles, is distinguished by a black-bordered
bright red-orange caudal in. Two non-schwenkii complex species also occur in Seychelles: P. lavicycla and P.
megalops. No DNA sequences are available for P. argyrea.
Pempheris bineeshi Randall & Victor, n. sp.
Figures 5 & 6, Tables 1–3. BIN AAM9396 (in part).
Pempheris schwenkii [non Bleeker] Randall & Bineesh 2014: 36 (Tuticorin, India).
Holotype. NBFGR-CH-1143, female, 101 mm SL, India, Tamil Nadu, Tuticorin, trawler, 10–30 m, K.K.
Bineesh, March 14, 2013.
Paratypes. BPBM 41096, 100 mm SL, India, Tamil Nadu, Tuticorin, from isherman, K. Rama Rao & J.E.
Randall, March 1, 1975; ZSI/CLT 2492, 5: 95–108 mm SL, same locality and date as holotype.
Description. Dorsal-in rays VI,9; anal-in rays 39 (38–41); pectoral-in rays 17 or 18 (modally 18); lateralline scales 49 (50–54); gill rakers 26 (25–27); body depth 2.5 (2.5–2.7) in SL; body width 2.6 (2.55–2.65) in body
depth; head length 3.3 (3.2–3.35) in SL; dorsal proile of nape nearly straight, curving slightly to tip of snout;
eye moderately large, 7.75 (7.5–7.85) in SL; interorbital width 11.9 (12.0–12.2) in SL; mouth oblique, forming
an angle of 63°(60–64°) to horizontal axis of body; tip of lower jaw projecting slightly anterior to upper jaw
when mouth irmly closed; small, strongly recurved, sharp teeth in two rows in upper jaw, those of upper row
more forward-projecting, of lower row larger and more strongly recurved; teeth progressively smaller posteriorly
in jaw, narrowing to a single row on about posterior ifth, and disappearing as maxilla expands; teeth in lower
Figure 5. Pempheris bineeshi, holotype, NBFGR-CH-1143, female, 101 mm SL, Tuticorin, India (K.K. Bineesh).
20
Figure 6. Pempheris bineeshi, paratype, ZSI/CLT 2492, 104 mm SL, Tuticorin, India (K.K. Bineesh).
jaw very small and densely spaced, in a band of four to ive irregular rows medially, where band broadest,
progressively larger, more retrorse and more pointed inwardly, the innermost teeth much the largest and nearly
horizontal; lips smooth; tongue nearly forming an equilateral triangle; outer layer of scales, nearly all of inner
layer, and even some of the strongly adhering lateral-line scales missing; predorsal length 2.8 (2.8–2.85) in SL;
longest dorsal-in ray 3.9 (3.95–4.05) in SL; caudal-in length 3.55 in SL (broken on paratypes); caudal concavity
2.35 in head length of holotype; pectoral-in length 3.15 (3.1–3.3) in SL; prepelvic length 2.8 (2.8–2.85) in SL;
pelvic ins just reaching anus, their length 5.6 (5.75–5.9) in SL. Color in alcohol (Fig. 5) wth the dorsal in broadly
bordered anteriorly and distally with black, the caudal in with a blackish posterior border and no dark pigment
on upper and lower margins, and two large dark areas forming an indistinct bar across nearly the basal half of in;
dark band at base of anal in broad; only a faint narrow dark band across upper base of pectoral ins. Color when
fresh as in Fig. 6.
Etymology. This species is named for K.K. Bineesh of the ICAR-National Bureau of Fish Genetic Resources
(NBFGR) in Kochi, Kerala, in recognition of the extensive efforts he has made to collect, photograph, and DNAbarcode species of Pempheris in India.
Remarks. Presently known only from the type location, but to be expected elsewhere in SE India, and maybe
well beyond. All of our specimens have been obtained from ishermen in Tuticorin, one of the oldest cities of
India. The sweepers are typically collected as bycatch on small trawlers that ish in the evening and at night off
the coast; the objective is shrimp and ishes of far more value than the small, thin-bodied species of Pempheris.
Specimens collected by trawling have always lost nearly all of their scales. P. bineeshi is the only representative
of the schwenkii complex from Indian waters, although P. trinco is described from nearby Sri Lanka. Other
congeners from south India include P. malabarica, P. mangula, and P. sarayu. The barcode mtDNA COI sequences
of P. bineeshi are the same as the sequence from P. hadra from the Maldives well to the south and specimens of
presumed true P. schwenkii from Bali, Indonesia (the type location of P. schwenkii is Sumatra, midway between
the Indian subcontinent and Bali). The three species form a lineage 3.4% divergent from a set of schwenkiicomplex species from the western Indian Ocean, i.e. P. tominagai from the Red Sea, P. xanthomma from Yemen,
P. ternay from Seychelles, and P. rubricauda from Madagascar (Fig. 1).
21
Pempheris bruggemanni Randall & Victor, n. sp.
Figures 7–9, Tables 1–3. BIN AAT9853.
Holotype. MNHN 2015-0002, male, 108 mm SL, Réunion, W side, S of Saint-Leu, 21°11.600 S; 55°16.946
E, 18 m, rotenone, H. Bruggemann, N. Hubert, & S. Planes, 13 Aug. 2007.
Paratypes. SMF 1537, 118 mm SL, Mauritius, 1885 (no further data); URUN 2007-1208, 83 mm SL &
URUN 2007-1209, 86.5 mm SL, fore reef slope in front of Saint-Gilles, Réunion, 21°3.460 S, 55°13.280 E, 18
m, rotenone, H. Bruggemann, N. Hubert, & S. Planes, Aug. 12, 2007; URUN 2007-1419, female, 97 mm SL, &
BPBM 41198, female, 103 mm SL (eyes missing), same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,41 (40–42); pectoral-in rays 18 (17–18); lateral-line scales
60 (58–63); gill rakers 28 (28–29); body depth 2.3 (2.3–2.35) in SL; body width 3.1 (2.85–2.95) in body depth;
head length 3.1 (3.1–3.2) in SL; nape slightly convex; orbit diameter 7.2 (7.2–7.3) in SL; interorbital width 10.8
(10.8–11.4) in SL; maxilla forming an angle of 68° (65–70°) to horizontal axis of body, reaching to or slightly
posterior to a vertical at center of eye; tip of lower jaw projecting slightly anterior to upper jaw when mouth
fully closed; teeth in upper jaw typical of genus, in two rows medially, soon narrowing to a single row; teeth in
lower jaw very small, in 3 or 4 irregular rows medially, progressively more retrorse and more pointed medially,
the innermost largest and nearly horizontal; lips smooth; scales on nape and below dorsal in ctenoid; scales on
ventroanterior two-thirds of side of chest ctenoid, the rest cycloid; scales ctenoid on caudal peduncle only dorsally
and ventrally; predorsal length 2.5 (2.5–2.6) in SL; caudal and dorsal ins broken; pectoral-in length 3.1 (3.0–
3.1) in SL; prepelvic length 2.4 (2.35–2.4) in SL; pelvic ins reaching or extending slightly beyond anus, their
length 5.7 (5.3–6.25) in SL. Color in alcohol gray, each scale of body with a vertically elongate brown bar; ins
yellowish, the dorsal with a blackish anterior margin that expands distally; anal in with a broad dark brown band
at base that narrows posteriorly, and a broad blackish distal margin, also narrower posteriorly; caudal in with dark
upper and lower margins and a dark band enclosing the lateral line, but no black posterior margin; paired ins pale
yellowish, the pectoral ins with a narrow, curved, dark brown band at extreme base of rays; no dark axillary spot.
Color when fresh as in Figs. 7 & 8; color in life as in Fig. 9.
Etymology. This species is named for Professor Henrich Bruggemann of the Laboratoire d’Ecologie Marine,
Université de la Réunion, who collected the type specimens with N. Hubert and S. Planes and provided the
photographs. He was supported by the BIOTAS program of the French Agence Nationale de Recherche (ANR06-BDIV), with the collaboration of Gustav Paulay (Florida Museum of Natural History).
Figure 7. Pempheris bruggemanni, holotype, MNHN 2015-002, male, 108 mm SL, Réunion (H. Bruggemann).
22
Figure 8. Pempheris bruggemanni, paratype, URUN 2007-1419, female, 97 mm SL, Réunion (H. Bruggemann).
Remarks. Presently known only from Réunion and Mauritius. The head of the holotype of P. bruggemanni
must have been bent inwardly when photographed, resulting in an unnaturally short snout (a distortion also
noted in some photographs of other species of the genus). The 83-mm male and 86.5-mm female paratypes are
sexually mature. The paratype from Mauritius is in poor condition and was not used for measurements. Prof.
Bruggemann conirms our identiication of Alain Diringer’s underwater photograph of Fig. 9 as the subadult of
the present species. One other member of the schwenkii complex, P. heemstraorum, has also been collected in
the Mascarene Islands, a single specimen from Rodrigues: it is a larger ish with a deeper body than the present
species. The barcode mtDNA COI sequences of P. bruggemanni form a lineage 6.5% divergent (K2P; 6.3%
uncorrected pairwise) from their nearest relatives, the clade including the common P. connelli of South Africa as
well as P. hollemani from Madagascar and P. rochai from Oman (Fig. 1).
Figure 9. Pempheris bruggemanni, underwater photograph, Réunion (A. Diringer).
23
Pempheris connelli Randall & Victor, n. sp.
Figures 10 & 11, Tables 1–3. BIN AAC6084 (in part).
Pempheris tominagai [non Koeda, Yoshino, Imai & Tachihara] Koeda et al. 2014: 317, in part.
Holotype. SAIAB 58560, female, 117 mm SL, South Africa, KwaZulu-Natal, Aliwal Shoal, 30.293° S,
30.817° E, ichthyocide, A. Bentley, P. Heemstra, E. Heemstra, & W. Holleman, June 14, 1998.
Paratypes. SAIAB 40347, 9: 60–111 mm SL, South Africa, KwaZulu-Natal, Park Rynie, Cowrie Reef,
30.3166° S, 30.733° E, C. Buxton, Aug. 4, 1992; SAIAB 43263, 2: 85–93 mm SL, KwaZulu-Natal, Horn’s
Reef, down beach from Mile 13, C. Buxton, June 24, 1993; SAIAB 46185, 2: 100–100.5 mm SL, South Africa,
KwaZulu-Natal, Anchor Reef, 30° 25.00’ S, 30.816° E, ichthyocide, C. Buxton, P. Heemstra, R. Tilney, & L.
Berger, June 17, 1994; BPBM 41202, 106 mm SL; same data as SAIAB 46185; USNM 435889, 2: 116–119 mm
SL, same data as holotype; SAIAB 61199, 3: 105–126.5 mm SL, Aliwal Shoal, 30° 25’ S, 30.817° E. ichthyocide,
P. Buchal, E. Heemstra, P. Heemstra, H. Larson, M. Smale, & A. Bentley, Nov. 25, 1999; SAIAB 62965, 2:
98–102 mm SL, Aliwal Shoal, ichthyocide, P. Buchal, P. Heemstra, E. Heemstra, H. Larson, M. Smale, & A.
Bentley, Aug. 3, 2000; SAIAB 64662, 3: 91–99.5 mm SL, KwaZulu-Natal, 8-Mile Reef, 27.441° S, 32.717° E,
ichthyocide, P. Heemstra, H. Larson, M. Smale, & A. Bentley, May 14, 2001; SAIAB 75776, 120 mm SL, female,
KwaZulu-Natal, Aliwal Shoal, Cathedral Reef, 30.250° S, 30.81° E, spear, A. Connell, July 29, 2003; SAIAB
75850, 2: 78–103 mm SL, Aliwal Shoal, 30.300° S, 30.816° E, ichthyocide, P. Heemstra, E. Heemstra, J. Stapley,
S. Warren, K. Hutchings, J. Dives, & R. Reaugh, Sept. 25, 2003; SAIAB 194781, 2: 96.5–98 mm SL, KwaZuluNatal, Brighton Beach, 29.937° S, 31° 0.533’ E, S. Chater et al., Mar. 1, 2012; SAIAB 194782, 6: 53–63 mm SL,
KwaZulu-Natal, Blood Reef (30 km S of Durban), 29° 53.041’ S, 31° 3.606’ E, 12 m, ichthyocide, A. Connell,
Jan. 27, 2014; SAIAB 194783, 2: 102–113.5 mm SL, & BPBM 41204, 106 mm SL, Aliwal Shoal, 30° 16.682’ S,
30° 48.555’ E, ichthyocide and spear, A. Connell, July 5, 2014.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (37–43); pectoral-in rays 18 (17–19); lateral-line scales
Figure 10. Pempheris connelli, holotype, SAIAB 58560, female, 117 mm, Aliwal Shoal, KwaZulu-Natal; pale blue areas
on the body are where both layers of scales are missing (P.C. Heemstra).
24
53 (50–58); gill rakers 30 (29–32); body depth 2.2 (2.0–2.25) in SL; body width 2.95 (2.8–2.95) in body depth;
head length 3.25 (3.2–3.3) in SL; eye moderately large, the orbit diameter 7.2 (7.05–7.55) in SL (in specimens
95–117 mm SL); interorbital width 10.6 (10.6–11.2) in SL; mouth oblique, forming an angle of 72° (67–72°) to
horizontal axis of body; tip of lower jaw expanded to a small knob that projects slightly anterior to upper jaw
when mouth fully closed; small, recurved, inwardly depressible teeth in two irregular rows anteriorly in upper
jaw, soon narrowing to a single row posteriorly; small, recurved, depressible teeth in lower jaw in two to three
rows medially, interspersed with tiny blackish nodules; lips with numerous, close-set papillae; tongue triangular,
the sides indented near tip to form a small rectangle with rounded tip; scales on nape and below dorsal in ctenoid;
scales of side of chest about 3/4 cycloid, the ctenoid scales ventroanteriorly and adjacent to operculum; scales on
side of caudal peduncle cycloid; predorsal length 2.6 (2.55–2.65) in SL; longest dorsal-in ray 4.5 (4.1–4.9) in SL;
pectoral-in length 3.2 (3.0–3.2) in SL; prepelvic length 2.45 (2.4–2.45) in SL; pelvic ins not reaching anus, 6.5
(6.1–6.45) in SL. Color when fresh as in Fig. 10; no black spot at base of pectoral ins, but a narrow, curved dark
brown line at base of pectoral-in rays (not crossing entire base); axil of pectoral ins dark brown to black; no black
posterior border on caudal in; color in life as in Fig. 11. Largest specimen examined, 126.5 mm SL.
Etymology. This species is named for Dr. Allan D. Connell, whose untiring efforts to document the ishes of
KwaZulu-Natal have resulted in the discovery of many new species. He collected several of the recent lots of the
types of this species, as well as numerous specimens, photographs, and tissue samples of other species of ishes
from southern Africa.
Remarks. Presently known only from KwaZulu-Natal, South Africa. A second member of the schwenkii
complex, P. ibo, occurs in South Africa and Mozambique and can be found on the same reef and in the same
collections as P. connelli. Dr. Connell speared one of each species in 24 m at Aliwal Shoal. The two species are
easily confused as preserved specimens, but can be fully separated by the lateral-line scale count: 50–58 for P.
connelli vs. 43–49 for P. ibo. Also P. connelli appears to attain larger size; the largest of 14 specimens of P. ibo
examined by us measures 93 mm SL. Fortunately, despite their outward similarity, the barcode mtDNA COI
sequences of the two species are very different: P. connelli sequences are 13.3% divergent (K2P; 12% uncorrected
pairwise) from P. ibo. In fact, P. bruggemanni from the Mascarene Islands are actually closer genetically to P.
connelli (6.3% divergent) and are the nearest-neighbor lineage (Fig. 1). The barcode sequence from the type
specimens of P. hollemani from Madagascar and P. rochai from Oman have the same COI sequence as P. connelli,
indicating either a recent split of closely related species or occasional hybridization may occur between the three
species across the western Indian Ocean.
Figure 11. Pempheris connelli, underwater aggregation, Aliwal Shoal, KwaZulu-Natal (D. King).
25
Pempheris ellipse Randall & Victor, n. sp.
Figures 12 & 13, Tables 1–3.
Holotype. BPBM 35867, female, 93 mm SL, Oman, south coast, Rahah Bay, 16°57’4” N, 54°49’4” E, 6–9 m,
spear, J.L. Earle, Oct. 10, 1993.
Paratypes. BPBM 41082, 15: 31–41 mm SL, Oman, south coast, Rahah Bay, tide pools at low tide, 0–2 m,
rotenone, J.E. Randall, J.P. Hoover, & I. McLeish, Feb. 8, 1993.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (36–40); pectoral-in rays 18 (17–19, strongly modal 18;
only one count of 19); lateral-line scales 53 (50–57); lateral line very strongly arched anteriorly, rising 3/4 orbit
diameter above level of origin of lateral line; scales on nape and below dorsal in ctenoid; scales on ventroanterior
third of side of chest ctenoid, remaining chest scales cycloid; most scales missing from caudal peduncle, but
the few intact scales are cycloid; gill rakers 27 (26–30); body deep, the depth 2.1 in SL, and moderately thick,
the width 2.95 in depth; head length 3.35 in SL; eye forming a vertical ellipse (bony orbit width 12.2 mm; bony
orbit height 13.9 mm); orbit diameter 7.45 in SL; interorbital width 10.9 in SL; jaws steeply oblique, the edge
of maxilla forming an angle of 70° (69–75° in juveniles) to horizontal axis of body; upper jaw with a close-set,
double row of small, recurved, sharply conical teeth that are progressively smaller laterally, ending in a single
row of very small nodular teeth; band of teeth medially in lower jaw crossed by four to six irregular rows of very
small recurved teeth, not clearly larger inwardly; tongue narrowly triangular with pointed tip; predorsal length 2.8
in SL; longest dorsal-in ray 3.75 in SL; caudal in broken; pectoral-in length 3.3 in SL; prepelvic length 2.65 in
SL; pelvic ins reaching origin of anal in, their length 5.65 in SL. Color of holotype in alcohol as shown in Fig.
12; the body is uniform brown, only the scale edges a little lighter, except for a reddish hue on the chest and a faint
greenish zone beneath the pectoral in and continuing ventrally to the pelvic ins. Color of a juvenile in alcohol as
in Fig. 13. Life color not reported.
Etymology. This species is named Pempheris ellipse in reference to the longer vertical axis of the orbit, giving
the eye an elliptical shape. The speciic epithet is a noun in apposition.
Figure 12. Pempheris ellipse, holotype, BPBM 35867, female, 93 mm SL, Rahah Bay, south coast of Oman, with two spear
wounds on side below dorsal in (H.A. Randall).
26
Figure 13. Pempheris ellipse, paratype, juvenile, 41 mm, Rahah Bay, southern Oman (H.A. Randall).
Remarks. Fifteen juveniles collected from a tidepool at Rahah Bay were identiied as this species by the shape
of the eye, the highly arched anterior end of the lateral line, the very steep angle of the upper jaw, and meristic
characters. Three other species of Pempheris were collected in the bay: P. convexa Randall & Victor 2014, P.
micromma, described in the present paper, and a subadult of an unidentiied schwenkioid species collected from a
cave in 14 m (BPBM 36325, 72 mm SL), for which no photograph or color note was taken. It has a very steep jaw
angle, straight dorsal proile of the head and nape, ctenoid scales on the nape and below the dorsal in (all scales
missing from chest), but differs from P. ellipse in that the eye is round, the body more slender, and the anal in has
a blackish margin. Pempheris rochai, described herein from the nearby coast of Oman, might also be expected in
the bay. A photograph of the tidal channels of Rahah Bay was published by Randall & Victor (2014: Fig. 2). No
DNA sequences are available for P. ellipse.
Pempheris hadra Randall & Victor, n. sp.
Figure 14, Tables 1–3. BIN AAM9396 (in part).
Holotype. SAIAB 187537, female, 118 mm SL, Republic of Maldives, Kaafu Atoll, 4°10.180’ N; 73°30.800’
E, rocky wall of swimming pool opposite power station, rotenone, O. Gon & G. Gouws, Dec. 6, 2010.
Description. Dorsal-in rays VI,9; anal-in rays III,38; pectoral-in rays 17; lateral-line scales 49; gill rakers
26; body depth 2.45 in SL; body very stout, the width (taken just behind gill opening) 2.2 in body depth; head
length 3.5 in SL; eye relatively small, orbit diameter 8.4 in SL; interorbital width 12.5 in SL; mouth oblique,
forming an angle of 64° to horizontal axis of body; tip of lower jaw slightly anterior to upper jaw when mouth
fully closed; teeth in upper jaw in two well-separated rows medially, narrowing to a single row posteriorly; conical
teeth medially in lower jaw in four or ive densely packed rows, those of outer row nearly erect, the remaining
teeth progressively more recurved inwardly; scales on nape and below dorsal in cycloid; scales on ventroanterior
two-ifths of chest and along edge of operculum ctenoid, cycloid on the remaining three-ifths; scales on caudal
peduncle ctenoid; predorsal length 2.7 in SL; longest dorsal-in ray 3.95 in SL; caudal in emarginate, the caudal
27
Figure 14. Pempheris hadra, holotype, SAIAB 187537, female, 118 mm SL, Kaafu Atoll, Republic of Maldives (O. Gon).
concavity 3.8 in head length; pectoral-in length 3.25 in SL; prepelvic length 2.55 in SL; pelvic ins nearly
reaching origin of anal in, 6.2 in SL. Color in alcohol dark orangish brown, the scale edges narrowly pale yellow;
ins yellowish, the leading edge of dorsal in blackish, expanding into a large black apical spot; dark reddish
streaks following longitudinal scale rows below lateral line: anal in with a very dark brown band at base, 3 times
broader anteriorly (comparable to the width of the in); a middorsal V-shaped blackish mark on nape before dorsal
in; caudal in with a broad blackish posterior margin, the upper and lower margins only slightly darker; no black
spot on base or axil of pectoral ins, but a curving dark brown line following base of rays. Color when fresh as
shown by Fig. 14.
Etymology. This species is named Pempheris hadra, from Greek for thick or stout, in reference to the unusual
stocky body.
Remarks. We are aware of only a single specimen of Pempheris hadra. It is distinct in being more stoutbodied than any schwenkii-complex species we have examined, and unusual in having cycloid scales on the nape,
but ctenoid scales on the side of the caudal peduncle. The unusual habitat is a pool constructed in the lagoon of
Kaafu Atoll from which water is taken for cooling the electric power plant. The pool is also used for recreational
swimming by the locals. We suspect that this ish found shelter in the incurrent cooling pipe (as have two other
species of Pempheris we have documented in Guam). This species is the only schwenkii-complex species collected
in Maldives; two unrelated species, P. convexa and P. lavicycla, are the other species known from the archipelago.
The barcode mtDNA COI sequence of P. hadra is the same as a lineage of Pempheris that includes P. bineeshi
from India and the probable true P. schwenkii (type location Sumatra) from Bali, Indonesia (Fig. 1). The lineage
is 3.4% divergent from the nearest lineage in the neighbor-joining tree, which contains the western Indian Ocean
complex of P. tominagai from the Red Sea, P. xanthomma from Yemen, P. rubricauda from Madagascar, and P.
ternay from Seychelles.
28
Pempheris heemstraorum Randall & Victor, n. sp.
Figure 15, Tables 1–3.
Holotype. SAIAB 69487, female, 127 mm SL, Mascarene Islands, Rodrigues, 19.6502° S, 63.4297° E, reef
in 4–7 m, ichthyocide, E. Heemstra, P. Heemstra, M. Smale, & J. Stapely, Oct. 8, 200l.
Description. Dorsal-in rays VI,9; anal-in rays III,37; pectoral-in rays 18; lateral-line scales 59; gill rakers
31; body depth 2.2 in SL; body width 3.35 in body depth; head length 3.05 in SL; eye relatively small, the orbit
diameter 8.45 in SL; interorbital width 10.7 in SL; mouth oblique, forming an angle of 63° to horizontal axis of
body; tip of lower jaw slightly projecting when mouth fully closed; teeth anteriorly in upper jaw recurved, in two
rows medially, reduced to an irregular single row of progressively smaller teeth about one-third back in jaw; teeth
of lower jaw very small, conical and recurved, in a dense band of ive to six rows medially, progressively more
retrorse inwardly; upper lip with dense papillae; lower lip with small well-separated papillae; tongue narrowly
triangular; outer layer of scales missing from nape, side of chest, and caudal peduncle; predorsal length 2.45 in
SL; longest dorsal-in ray 3.85 in SL; caudal in slightly forked, caudal concavity about 5 in head length; pectoral
ins long, 3.0 in SL; prepelvic length 2.5 in SL; pelvic ins long, reaching origin of anal in, 5.3 in SL. Color in
alcohol of body below lateral line nearly uniform light brown, the scale edges a little paler; head and body above
lateral line darker brown; dorsal in translucent with dusky rays, the spinous portion and irst ray dark brown, with
a little dark pigment extending onto distal end of next few rays; anal in with a dark brown band at base, half depth
of in anteriorly, but narrowing posteriorly to only one-ifth height of in; caudal in with translucent yellowish
membranes and light brown rays, the upper and lower margins dark brown, a narrow brown band following lateral
line to end of in, no posterior dark margin; paired ins pale yellowish, the pectoral ins without a dark brown spot
on base or axil. Color when fresh as shown in Fig. 15.
Etymology. This species is named for Dr. Phillip C. Heemstra and Elaine Heemstra of the South African
Institute for Aquatic Biodiversity, in recognition of their extensive body of work in ichthyology. They collected
and photographed the holotype of this species, along with providing many other specimens and photographs of
western Indian Ocean ishes.
Remarks. This species is unusually large for the schwenkii complex, although the markings are typical for the
complex. Three other species are found in the Mascarenes: another schwenkii-complex species, P. bruggemanni,
which has more anal-in rays and fewer gill-rakers, and the rhomboidea-complex species P. nesogallica, which
was described from Mauritius, and the unrelated P. bexillon. No DNA sequences are available for P. heemstraorum.
Figure 15. Pempheris heemstraorum, holotype, SAIAB 69487, 127 mm SL, Rodrigues, Mascarene Islands (P.C. Heemstra).
29
Pempheris hollemani Randall & Victor, n. sp.
Figure 16, Tables 1–3. BIN AAC6084 (in part).
Pempheris tominagai [non Koeda, Yoshino, Imai & Tachihara] Koeda et al. 2014: 317, in part.
Holotype. SAIAB 200564, 118 mm SL, Madagascar, Anosy Region, Fort Dauphin, Libanona Beach, 25.5166°
S, 32.6833° E, gill net, P.C. Heemstra, E. Heemstra, et al., April 30, 2010.
Paratype. SAIAB 97406, 117 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,38 and 40; pectoral-in rays 17 or 18; lateral-line scales 58
(55); gill rakers 27 (26); body depth 2.3 (2.2) in SL; body width 2.65 (2.6) in body depth, head length 3.6 (3.5)
in SL; eye relatively small, the orbit 7.7 (7.85) in SL; interorbital width 12.1 in SL; mouth oblique, forming an
angle of 70° to horizontal axis of body; tip of lower jaw projecting when mouth fully closed; small slender teeth
in two rows anteriorly in upper jaw, soon narrowing to a single row; very small teeth in a dense band in lower
jaw, also forming a single row posteriorly; tiny black nodules scattered within band of teeth of lower jaw; tongue
triangular, the dorsal surface with scattered small black spots; scales cycloid, except ctenoid on nape, below
and posterior to dorsal in, and below a line from midway along edge of opercle to base of pelvic ins; predorsal
length 2.7 in SL; longest dorsal-in ray 4.1 in SL; caudal in forked, length 4.0 (4.3) in SL, the caudal concavity
4.3 (4.0) in head length; pectoral-in length 3.2 in SL; prepelvic length 2.5 in SL; pelvic ins not reaching anus,
6.9 (6.75) in SL. Color in alcohol pale brown, the head and body above lateral line a little darker; dorsal in with
dense melanophores anteriorly and distally; caudal in with ine melanophores on rays, fewer on membranes, the
margins of in a little darker; a broad dark band at base of anal in, as if stippled with large melanophores; a curved
blackish line at base of pectoral-in rays, and a large black spot in axil of ins. Color when fresh as in Fig. 16.
Etymology. This species is named for Dr. Wouter Holleman of the South African Institute for Aquatic
Biodiversity, in recognition of his extensive research on western Indian Ocean ichthyology. He provided the data
on this species from the two type specimens when they could not be sent on loan.
Remarks. The two specimens of P. hollemani were collected during the Atimo Vatae Expedition to southern
Madagascar (Philippe Bouchet of the Museum national d’Histoire naturelle, Principal Investigator). Koeda et al.
(2014: 317) erred by including this species as a paratype of their new species P. tominagai, type locality Sharm
Figure 16. Pempheris hollemani, holotype, SAIAB 200564, 118 mm, SE Madagascar (E. Heemstra).
30
el-Sheikh, northern Red Sea. The specimens are clearly not P. tominagai, a Red Sea species with a different color
pattern and more than 15% different mtDNA sequences (Randall et al. 2014). All species of Pempheris of the
schwenkii complex share a broad dark brown to black band at the base of the anal in. Pempheris hollemani has
this dark anal-in band, but instead of being uniformly dark brown or black, it is unique in consisting of discrete
black dots of variable size. The barcode mtDNA COI sequence of P. hollemani is the same as P. connelli from
South Africa and P. rochai from Oman, indicating either a recent split of closely related species or some degree
of gene low across the western Indian Ocean between the three species (Fig. 1).
Pempheris ibo Randall & Victor, n. sp.
Figures 17 & 18, Tables 1–3. BIN AAF8820.
Holotype. SAIAB 191483 (out of SAIAB 3874), female, 93 mm SL, Mozambique, Ibo Island, lighthouse,
12.3333° S, 40.6166° E, M.M. Smith & R. Stobbs, Oct. 1, 1973.
Paratypes. SAIAB 3759, 8: 35–45 mm SL, Mozambique, Quilalea Island, 12.4891° S, 40.5994° E, M.M.
Smith & R. Stobbs, Oct. 12, 1973; BPBM 41201, 80 mm SL, male, Mozambique, 15°32.91’ S, 40°38.57’ E, R/V
Fridtjoff Nansen, M. Lisher, Aug. 13, 2009; SAIAB 189307, 2: 54.5–65 mm SL, South Africa, KwaZulu-Natal,
sunken barge DAR 1, 28°09.682’ S, 32°33.649’ E, 25 m, A.D. Connell, spear, Apr. 13, 2010; SAIAB 189165, 2:
41.5–60 mm SL, South Africa, KwaZulu-Natal, sunken barge DAR 1, 25 m, A.D. Connell, Aug. 1, 2011.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (35–39); pectoral-in rays 18 (17 or 18, modally 18);
lateral-line scales 48 (43–49); lateral line rising steeply and straight from origin to a level about 2/3 orbit diameter
above origin, then forming a distinct sharp angle with remaining lateral line; scales on nape and below dorsal
in ctenoid; scales on ventroanterior three-quarters of side of chest ctenoid, remaining scales cycloid; scales on
caudal peduncle ctenoid, except for scale row above and below lateral line; gill rakers 30 (27–30); body depth
2.35 (2.2–2.45) in SL, moderately compressed, width 2.8 (2.75–3.0) in body depth; head length 3.3 (3.25–3.3) in
SL; orbit diameter 7.8 (7.25–7.95) in SL; interorbital width 11.6 (11.3–12.4) in SL; mouth oblique, forming an
angle of 68° (66–69°) to horizontal axis of body; dentition typical of most species of the genus; tongue slender,
Figure 17. Pempheris ibo, holotype, SAIAB 191483, female, 93 mm, Mozambique, Ibo Island (H.A. Randall).
31
triangular, and sharply pointed; predorsal length 2.7 (2.7–2.75) in SL; longest dorsal-in ray 3.9 (3.9–4.15) in
SL; caudal in forked, the caudal concavity about 2.6 in SL; pectoral-in length 3.0 (2.95–3.1) in SL; prepelvic
length 2.6 (2.55–2.6) in SL; pelvic ins just reaching anus, in length 6.45 (5.75–6.65) in SL. Color in alcohol of
holotype shown in Fig. 17. The faint blackish border on the anal in is not shared by all specimens. The life color
is variable, as shown in Fig. 18.
Etymology. This species is named for the type locality, Ibo Island, Mozambique. The speciic epithet is a noun
in apposition.
Remarks. The 34-m lighthouse at Ibo Island, Pinda, Mozambique was erected in 1923 to warn mariners of
nearby shoal waters. Fifty years later, Margaret M. Smith and Robin Stobbs made a small collection of ishes off
the lighthouse that included 11 specimens of Pempheris. Ten were identiied as P. adusta Bleeker, later corrected
by us to P. eatoni Randall & Victor. The remaining 93-mm specimen is the holotype of the present species. Ten
days later, Smith & Stobbs made an inshore ish collection at Quilalea Island, Mozambique that included eight
juveniles of P. ibo. Recent collections have extended the range to KwaZulu-Natal, the most southern at Aliwal
Shoal, where P. ibo can be found in the same collections as their similar-appearing and more abundant relative,
P. connelli.
The small size, slender body, meristic data, broad blackish band at base of anal in, and dark leading edge of
the dorsal in identify this species as one of the schwenkii complex. It is readily identiied as a distinct species by
having 43–49 lateral-line scales (lowest for any species of the genus in the western Indian Ocean– see Table 2),
steeply angular anterior lateral line, relatively low count of anal-in soft rays, long pectoral ins, and short predorsal
length. The ish of Fig. 18 are readily identiied as P. ibo by the slender body, relatively small eye, angular anterior
lateral-line, long pectoral ins, and especially by being able to count the low number of anal-in soft rays and
lateral-line scales. One other species of the schwenkii complex, P. peza, has been collected in Mozambique and
one other in South Africa, the common P. connelli; both have many more lateral-line scales. Despite the similarity
in appearance, the barcode mtDNA COI sequences of P. ibo are very different from the P. connelli lineage, 13.3%
divergent (K2P; 12% uncorrected pairwise). The nearest-neighbor lineage to P. ibo is P. bruggemanni from the
Mascarenes, but still far distant with 11.8% divergence (K2P; 11% uncorrected pairwise)(Fig. 1).
Figure 18. Pempheris ibo, underwater aggregation, note color variation, Mozambique (D. Polack).
32
Pempheris kruppi Randall, Victor & Aideed, n. sp.
Figures 19 & 20, Tables 1–3. BIN AAD1777 (in part).
Pempheris sp. Zajonz et al. 2000: 156 (Socotra Archipelago).
Holotype. SMF 35711, female, 123.5 mm SL, Gulf of Aden, Yemen, Ras Belhaf, 13°58.361’ N, 48°11.743’
E, 8 m, moderately steep reef that shelves southward, high live coral cover of massive Porites, abundant foliose
Montipora, and encrusting Echinopora, rotenone, E. Lavergne, M. Schneider, & U. Zajonz, June 3, 2006.
Paratypes. SMF 35528, 6: 106.5–123 mm SL, & NHCY-P-12, 6: 108–118.5 mm SL, Gulf of Aden, Yemen,
Ras Majdaha, 14°00.868’ N, 48°25.572’ E, steep rocky slope to 6 m, collected with rotenone from 8–11 m, live
coral cover less than 20%, isolated corals, mainly Porites, some Pocillopora, rotenone, F. Krupp, M. Schneider,
& E. Lavergne, May 20, 2005; BPBM 41214, 121 mm SL, USNM 435864, 124 mm SL, & NHCY-P-13, 123 mm
SL, all with same data as holotype.
Description. Dorsal-in rays VI,9 (rarely VI,10); anal-in rays III,39 (37–41); pectoral-in rays 18 (17 or 18,
rarely 17); lateral-line scales 60 (55–64); scales on nape and chest ctenoid, mixed cycloid and ctenoid below
dorsal in, and cycloid on side of caudal peduncle; gill rakers 30 (29–32); body depth 2.15 (2.15–2.25) in SL;
body width 2.7 (2.6–2.7) in body depth; head length 3.2 (3.1–3.35) in SL; eye relatively small, the orbit diameter
7.4 (7.15–7.45) in SL; interorbital width 10.9 (10.7–11.0) in SL; mouth strongly oblique, forming an angle of 69°
(67–70°) to horizontal axis of body; very small, slender, recurved, conical teeth in a double row in medial third of
upper jaw, continuing as a single row to end of jaw; slender, conical, recurved teeth in lower jaw in a narrow band
of four to ive rows medially, narrowing to a single row at nearly the same point as upper jaw; tongue triangular,
straight-sided and pointed; predorsal length 2.7 (2.6–2.7) in SL; longest dorsal-in ray 4.0 (3.8–4.0) in SL; caudal
in moderately forked, the caudal concavity about 4–5 in head length; pectoral-in length 3.35 (3.15–3.4) in SL;
prepelvic length 2.6 (2.5–2.6) in SL; pelvic ins reaching or extending slightly posterior to anus, 5.85 (5.1–6.05) in
SL. Color in alcohol as in Fig. 19; dark brown, the scale edges silvery gray; dorsal in yellowish gray with a distal
black spot on irst six soft rays, its length two-thirds eye diameter; caudal in gray-brown, progressively lighter
Figure 19. Pempheris kruppi, holotype, SMF 35711, female, 123.5 mm, Gulf of Aden, Yemen, off Ras Belhaf (H.A. Randall).
33
Figure 20. Pempheris kruppi, Gulf of Aden off Majdaha, south coast of Yemen; note the variation in the color of the
posterior margin of the caudal in (M. Shaikh Aideed).
posteriorly, except for a blackish posterior border (when in intact); anal in reddish brown on basal half, followed
by a middle zone of brownish yellow, inely lecked with black dots, and a narrow translucent outer zone about
one-third pupil diameter in average width; paired ins yellowish, the pectoral-in base with a purplish black band
across base, one-half pupil diameter in depth; axil of pectoral ins black; iris brassy yellow with a black circle
that expands dorsally. Color in life dark brown with prominent lateral dark stripes; yellowish to orange ins and a
distinct oval apical spot on the dorsal in; anal in with dark-tipped membranes on irst few rays (Fig. 20). Largest
specimen examined, the holotype.
Etymology. This species is named for Dr. Fareed Krupp of the Forschungsinstitut Senckenberg, authority
on the ishes of the Gulf of Aden and Red Sea, and principal collector of the irst series of type specimens of the
species.
Remarks. Presently known only from the Gulf of Aden off Yemen and from the islands of the Socotra
Archipelago. It is one of several species in the rhomboidea complex in the region, and is closely related to P.
pathirana from Sri Lanka, from which it primarily differs by tooth arrangement. Pempheris rhomboidea from
the adjacent Red Sea and P. kuriamuria from Oman have longer pectoral ins; P. darvelli from Oman has a longer
head length; P. wilsoni from Oman has a shorter head and prepelvic length; P. muscat, also from Oman, has
divided scale types on the chest (vs. entirely ctenoid); and P. russellii from Pakistan has a different shape of the
tongue. Moteah Shaikh Aideed earned a coauthor role for the description of this species by providing specimens,
collection data, tissue samples, and photographs. The barcode mtDNA COI sequences of P. kruppi from Yemen
are the same as others of the rhomboidea complex, including two species described from specimens from nearby
Oman: P. kuriamuria and P. wilsoni (Fig. 1).
34
Pempheris kuriamuria Randall & Victor, n. sp.
Figures 21 & 22, Tables 1–3. BIN AAD1777 (in part).
Holotype. CAS 236521, female, 112.5 mm SL, Oman, south coast, Ras Hamar, 16°54’44.3” N, 53°47’25.9”
E, reef, ichthyocide, L.A. Rocha, J.D. DiBattista, M. Priest, T. Sinclair, & J.H. Choat, March 17, 2013.
Paratype. BPBM 41239, gender unknown, 95 mm SL, Oman, Gulf of Oman, Jissah Island, 23°33.513’ N,
58°39.144’ E, spear, B.W. Darvell & K.D.P. Wilson, Nov. 27, 2013.
Description. Dorsal-in rays VI,9; anal-in rays III,40 (38); pectoral-in rays 18 (one side 19); lateral-line
scales 61 (59); scales on nape and side of chest ctenoid, mixed cycloid and ctenoid below dorsal in, and cycloid on
side of caudal peduncle; gill rakers 30; body depth 2.15 in SL; body width 2.95 (2.85) in body depth; head length
3.3 (3.35) in SL; eye relatively small, equal to postorbital length, the orbit diameter 7.45 (7.3) in SL; interorbital
width 12.4 (10.4) in SL; mouth forming an angle of 66° (69°) to horizontal axis of body; upper jaw with small,
slender, recurved, conical teeth in two close-set rows to below nostrils, continuing as a single row to end of jaw;
teeth in lower jaw in a narrow band, crossed where broadest by six or seven rows of extremely small, recurved,
conical teeth that are progressively more retrorse posteriorly; tongue narrowly triangular with slightly incurved
sides; predorsal length 2.45 (2.55) in SL; longest dorsal-in ray 3.75 (3.65) in SL; caudal in slightly forked, 3.6 in
SL in paratype, caudal concavity about 3.5 in HL (estimated, longest rays broken); pectoral-in length 2.9 (2.95)
in SL; prepelvic length 2.5 in SL; pelvic ins just reaching origin of anal in, 5.65 (5.2) in SL. Color in alcohol
(Fig. 21) with dark band at base of anal in (very broad anteriorly and progressively narrower posteriorly), broad
translucent light middle zone of in, densely stippled with dark brown, and a narrow transparent outer margin; a
curved dark purplish brown band at base of pectoral ins; and dorsal in with a black apical spot, no dark leading
edge, and a narrow black band at base. Color in life as shown in Fig. 22.
Etymology. This species is named for the type locality, the Kuriamuria Islands off the south coast of Oman.
The speciic epithet is a noun in apposition.
Remarks. The underwater photographs of Fig. 22 are identiied as P. kuriamuria by the long pectoral ins,
eye size (equal to postorbital head), body and in proportions, lateral-line scale count, and color pattern, especially
the dark brown band at the base of the anal in that narrows posteriorly. Although this band is a typical feature
of the schwenkii complex, P. kuriamuria has the stripes along the lank following the scale rows typical of the
rhomboidea-complex. The holotype is notable for the relatively long pectoral ins distinguishing it from the other
species in the complex (shared only with P. rhomboidea of the Red Sea which differs by having two scale types
on the chest vs. all ctenoid). The barcode mtDNA COI sequence of P. kuriamuria is the same as others of the
rhomboidea complex, including P. wilsoni, also from Oman (Fig. 1).
Figure 21. Pempheris kuriamuria, holotype, CAS 236521, female, 112.5 mm SL, Oman, Ras Hamar (H.A. Randall). Caudal
in damaged (posterior margin of in has a uniform dark edge when intact).
35
Figure 22. Pempheris kuriamuria, underwater photographs; upper: Raysut, south coast of Oman; lower: Kuriamuria
Islands, southern Oman (J.E. Randall).
36
Pempheris leiolepis Randall & Victor, n. sp.
Figure 23, Tables 1–3.
Holotype. SAIAB 200715 (out of SAIAB 51259), male, 100 mm SL, Zanzibar, Chumbe Island, 6.1666° S,
39.1833° E, J.L.B. & M.M. Smith, Aug. 5, 1952.
Description. Dorsal-in rays VI,9; anal-in rays III,37; pectoral-in rays 18; lateral-line scales 53; scales on
nape, below dorsal in, and side of caudal peduncle cycloid; scales on side of chest cycloid, except for two or three
ventral ctenoid scales adjacent to operculum; gill rakers 28; body depth 2.4 in SL; body moderately compressed,
the width 2.6 in body depth; head length 3.0 in SL; dorsal proile of head nearly straight; eye diameter 7.75 in
SL; interorbital width 11.6 in SL; mouth oblique, forming an angle of about 63° to horizontal axis of body; teeth
in upper jaw typical of those of lower jaw of the genus, with three or four irregular rows of small conical teeth
that are progressively larger and more recurved posteriorly; teeth of lower jaw much smaller and more densely
packed, in a maximum of approximately six rows; tip of lower jaw expanded to a small knob that projects
slightly forward when mouth irmly closed; tongue triangular with slightly indented sides, the tip rounded and
very lexible; predorsal length 2.5 in SL; outer part of dorsal and caudal ins broken; pectoral-in length 3.15 in
SL; prepelvic length 2.4 in SL; pelvic ins just reaching origin of anal in, their length 6.2 in SL. Color in alcohol
as shown in Fig. 23; the body is a nearly uniform orangish brown, except for a reddish hue to the chest and a
greenish zone beneath the basal part of the pectoral in, continuing to the base of pelvic ins; note the absence of
a prominent dark spot at the base of the pectoral ins (instead there is a blackish line that nearly crosses the upper
part of the in base). Color in life unknown.
Etymology. This species is named Pempheris leiolepis, from the Greek meaning smooth scales, in reference
to its having the greatest cover of the body in cycloid scales.
Remarks. The type locality, Chumbe, is a 16-acre, privately-owned island 8 miles SW of Zanzibar, famous
for its spectacular coral reefs. Details are lacking for the Smiths’ collection of our single specimen. The species
is a member of the schwenkii complex; it is the only species of the complex we have examined along the African
coast north of Mozambique, where there are two other species, P. ibo and P. peza. The upper-jaw dentition is very
similar to that of P. andilana and P. argyrea, the basis for grouping them in the same part of the identiication key.
No DNA sequences are available for P. leiolepis.
Figure 23. Pempheris leiolepis, holotype, SAIAB 200715, male, 100 mm SL, Chumbe Island, Zanzibar (H.A. Randall).
37
Pempheris megalops Randall & Victor, n. sp.
Figure 24, Tables 1–3.
Holotype. BPBM 41063, female, 121 mm SL, Seychelles, La Digue, North Point, rocky shore, 0–l m, rotenone,
J.E. Randall, H.A. Randall, & D.J. Woodland, June 1, 1977.
Description. Dorsal-in rays VI,9; anal-in rays III,42; pectoral-in rays 17; lateral-line scales 58; outer layer
of scales on nape and side of caudal peduncle missing; scales on about ventroanterior half of side of chest ctenoid,
those on remaining half cycloid; gill rakers 31; body depth 2.35 in SL; body width 2.9 in body depth; head length
2.95 in SL; eye extremely large, the orbit diameter 6.2 in SL; interorbital width 11.0 in SL; upper jaw with a
narrow symphyseal gap, the larger teeth sharply conical and recurved, mainly in two irregular rows, those of the
upper row more jutting, narrowing to a single irregular row nearly half way to end of jaw; very small recurved
teeth above and interspersed among larger teeth; lower jaw with a band of very small, nodular, recurved teeth in
four or ive irregular rows where broadest medially; predorsal and prepelvic lengths 2.4 in SL; longest dorsal-in
ray 3.35 in SL; caudal in damaged; pectoral-in length 3.0 in SL; pelvic ins just reaching origin of anal in, 5.8
in SL. Color in alcohol as in Fig. 24; the caudal in had a broad blackish posterior margin in life, as well as one
on the outer margin of anal in, though not as strongly developed. There is no dark spot at the base or axil of the
pectoral ins. Color in life unknown.
Etymology. The species is named Pempheris megalops from the Greek meaning large eye, in reference to the
type having the largest eye of any species of the genus that we have examined.
Remarks. This species is apparently not a member of the schwenkii complex, but its placement is uncertain.
Three other Pempheris species are found in Seychelles: two in the schwenkii complex, P. ternay and P. argyrea
(type specimens of the latter in the same collection as this holotype), and the unrelated P. lavicycla. Pempheris
species typically occur in aggregations, so more than one specimen was probably collected in the rotenone station
that resulted in the holotype of P. megalops. Not realizing the species was undescribed, only one specimen was
preserved, and no photograph was taken. It is odd that this species, with such a large eye, was found by day at a
depth of less than one meter. No DNA sequences are available for P. megalops.
Figure 24. Pempheris megalops, holotype, BPBM 41063, female, 121 mm, La Digue, Seychelles (H.A. Randall).
38
Pempheris micromma Randall & Victor, n. sp.
Figures 25 & 26, Tables 1–3.
Holotype. BPBM 41157, female, 117 mm SL, Oman, south coast, Rahah Bay, west side, rocky point, 16°57’
N, 54°49’12” E, tide pools, 0–1.5 m, rotenone, J.E. Randall & I. McLeish, Feb. 6, 1993.
Description. Dorsal-in rays VI,9; anal-in rays III,43; pectoral-in rays 18; lateral-line scales 56; scales on
nape and below dorsal in cycloid; ctenoid scales on at least ventroanterior two-thirds of chest (missing from rest
of chest); scales on caudal peduncle ctenoid; gill rakers 30; body depth 2.35 in SL; body strongly compressed, the
width 3.1 in depth; head length 3.6 in SL; eye very small, the orbit diameter 9.45 in SL; interorbital width 13.1
in SL; jaws steeply oblique, the edge of maxilla forming an angle of 67° to horizontal axis of body; dentition of
upper jaw typical of the genus; band of teeth in lower jaw broadest medially where crossed by four or ive rows
of very small conical teeth, progressively more recurved inwardly; predorsal and prepelvic lengths 2.75 in SL;
longest dorsal-in ray 3.65 in SL; caudal in damaged; pectoral-in length 3.3 in SL; pelvic ins short, just reaching
anus, 6.65 in SL. Color in alcohol as in Fig. 25, the body brown, except for relection from scale edges, with a
greenish gray zone from beneath basal half of pectoral in to ventral abdomen; each scale with a darker vertical
zone from a stippling of dark brown dots; dorsal in with only a dusky band on the leading edge, but with a typical
distal black spot and a narrow blackish band at base; caudal in with a broad blackish posterior border, but without
dark pigment on upper and lower margins; anal in with three color zones, a very broad basal band a little darker
brown than body, strongly narrowing posteriorly, a middle pale yellow zone with brown dots, and a translucent
whitish margin; paired ins pale yellowish, the pectoral ins with a narrow dark brown band across most of base
where rays end; color in life as shown by the photograph of an aggregation in Fig. 26.
Etymology. This species is named Pempheris micromma from the Greek meaning small eye, in reference to
the type having the smallest eye for the genus.
Remarks. The holotype was collected from a large tidal channel in Rahah Bay on the south coast of Oman,
along with 15 juveniles of what was believed to be the same species. However, the juveniles were later identiied,
chiely by meristics, to be the same as an adult specimen of P. ellipse speared in the bay at a depth of 8 m. The
species of the underwater photograph of Fig. 26 is identiied as P. micromma by the very small eye, steep angle
of the upper jaw, predorsal and prepelvic lengths, and by the locality of the south coast of Oman near Rahah Bay.
The eye of the holotype in Fig. 25 appears vertically elliptical, but this is a result of the head not being in the same
plane as the body for the photograph. The left eye measures only 0.5 mm higher than broad, and the right eye is a
Figure 25. Pempheris micromma, holotype, BPBM 41157, female, 117 mm, Rahah Bay, southern Oman (H.A. Randall).
39
Figure 26. Pempheris micromma, underwater aggregation, south coast of Oman (J.E. Randall).
perfect circle, clearly distinguishing this species from P. ellipse. The holotype does not have the dark band at the
base of the anal in characteristic of the schwenkii complex, but the appearance in the underwater photograph is
consistent with schwenkii-complex sweepers. Additional material will resolve the status of this species.
Two other species co-occur in Rahah Bay with P. micromma: P. ellipse, distinguished by elliptical eyes and a
highly arched anterior lateral line, and P. convexa, diagnosed chiely by having a pronounced convex head proile
and by lacking a dark brown band at the base of the anal in. Pempheris rochai, described from one specimen
from Ras Hamar, nearby on the southern coastline of Oman, might be expected from the bay. It differs markedly
by having relatively large eyes, only a single row of teeth medially in the upper jaw, and the unusual count of 10
dorsal-in soft rays. A photograph of the tidal channels of Rahah Bay was published by Randall & Victor (2014:
Fig. 2). No DNA sequences are available for P. micromma.
Pempheris muscat Randall & Victor, n. sp.
Figure 27, Tables 1–3.
Holotype. BPBM 35845, male, 127 mm SL, Gulf of Oman, Oman, Cat Island, 23°35’8” N, 58°36’34” E, 5–6
m, spear, J.E. Randall, April 28, 1990.
Paratypes. BMNH 1889.4.15.35-38, 3: 124–132 mm SL (a fourth specimen in poor condition, not measured),
Gulf of Oman, Oman, Muscat, A. Jayakar, 1889; BPBM 21359, 129 mm SL, Oman, 2–3 miles south of Bandar
Sidab, islet about 300 m offshore, depth 3–5 m, spear, J.E. Randall, March 5, 1977.
40
Figure 27. Pempheris muscat, holotype, BPBM 35845, male, 127 mm SL, Cat Island, Gulf of Oman (H.A. Randall).
Description. Dorsal-in rays VI,9 (one of six with VI,10); anal-in rays III,40 (35–40); pectoral-in rays 18
(17–19); lateral-line scales 59 (54–59); outer layer of scales on nape and side of caudal peduncle missing; scales
ctenoid on about ventroanterior half of side of chest, those remaining cycloid; gill rakers 31 (30–32); body depth
2.35 (2.2–2.35) in SL; body width 2.95 (2.9–3.5) in body depth; head length 3.15 (3.1–3.25) in SL; eye large,
orbit diameter 6.85 (6.8–7.4) in SL; interorbital width 11.7 (11.1–12.4) in SL; jaws moderately oblique, the edge
of maxilla an angle of 65° (63–68°) to horizontal axis of body; upper jaw dentition similar to that of the generic
lower jaw, with a band of 3 to 5 irregular rows of very small, recurved, sharply conical teeth, progressively more
sessile inwardly, gradually narrowing posteriorly to side of jaw to a short single row of very small nodules; lower
jaw dentition similar to that of upper jaw, the teeth more widely spaced; predorsal and prepelvic lengths 2.5
(2.45–2.55) in SL; longest dorsal and caudal rays broken; longest pectoral ray in paratype with intact in 3.15 in
SL; pelvic ins just reaching anus, 6.8 (6.0) in SL. Color of holotype in alcohol as in Fig. 27, the scales of body
gray-brown, each with a vertically elongate orange-brown bar stippled with dark brown dots; dorsal in with
apical black spot, a narrow dark brown band at base, but no darker leading margin; caudal in with a blackish
posterior margin, judging from black pigmentation midposteriorly on remaining in; anal in densely stippled with
dark brown, darkest anteriorly, except for translucent whitish margin; paired ins pale orangish, the pectoral ins
with a faint narrow dark bar across base of rays. Color in life unknown.
Etymology. This species is named for the port city of Muscat, the capital of Oman, the only locality given for
the irst museum collection of the species. The speciic epithet is a noun in apposition.
Remarks. Pempheris muscat has as few as 35 anal-in soft rays, the lowest count for the genus in the region,
shared only with two species in the schwenkii complex, of which the new species is apparently not a member.
The appearance broadly overlaps other rhomboidea-complex species, including P. russellii Day, a species from
the Sind coast of Pakistan, which was our initial identiication for these specimens until the dentition was closely
examined. The teeth at the front of the lower jaw of P. muscat are twice as large as those of P. russellii, well
spaced, and in regular rows; those of the upper jaw are unique in the genus in being essentially the same as the
teeth of the lower jaw. Further comparison revealed that the new species has a larger head, less body depth, and,
if specimens of equal size are compared, a larger eye than P. russellii. No DNA sequences are available for P.
muscat.
41
Pempheris orbis Randall & Victor, n. sp.
Figure 28, Tables 1–3.
Holotype. SMNHTAU P.5284, female, 85 mm SL, Red Sea, Gulf of Aqaba, Dahab, L. Fishelson, Oct. 13, 1968.
Paratypes. HUJ 4727, females, 2: 77–77 mm SL, Red Sea, Ras Muhammad, near south tip of Sinai Peninsula,
rotenone, E. Clark, Aug. 7, 1968.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (38–39); pectoral-in rays 18 (17–18); lateral-line scales
58 (61–64); scales on nape, below dorsal in, and on side of chest cycloid; gill rakers 30–31; body depth 2.7 (2.3–
2.5) in SL; body width 2.6 (2.55–2.7) in body depth; head length 3.3 (3.2) in SL; dorsal proile of head slightly
convex; orbit diameter 7.7 (7.35–7.45) in SL; interorbital width 11.5 (11.3–11.8) in SL; jaws oblique, the edge
of maxilla an angle of 63° (62–63°) to horizontal axis of body; tip of lower jaw projecting anterior to upper jaw
when mouth fully closed; teeth small, conical and recurved, those medially in upper jaw in three to four irregular
rows, soon narrowing to two rows of progressively smaller teeth, and inally to a single row of very small nodules
on about posterior ifth of jaw, ending just 2 mm before posterior end of maxilla in holotype; teeth in lower jaw
in a band of four to ive, irregular, close-set rows (medially where band is broadest) comprising distinctly smaller,
more nodular, but still progressively more recurved teeth inwardly; innermost teeth largest and nearly horizontal;
lips dark purplish brown with distinct, well-spaced papillae; tongue triangular and straight-sided, except slightly
indented near tip; predorsal and prepelvic lengths 2.5 (2.5–2.6) in SL; longest dorsal-in ray 3.95 (4.1) in SL
(longest ray broken in largest paratype); all caudal-in rays broken; pectoral-in rays broken on holotype and
largest paratype, in length in smallest paratype 3.5 in SL; pelvic ins reaching or extending slightly posterior to
anus, their length 6.3 (6.4–6.8) in SL. Color of holotype in alcohol as in Fig. 28. Scales below the lateral line of the
largest paratype with light golden centers and orange margins, brightest in pectoral region; dorsal in with black
membranes distally between last three spines and irst four soft rays; broad anterior margin of dorsal in darker
than rest of in, except distal black part of in. Color in life unknown.
Etymology. This species is named orbis, Latin for circle or ring, for the most striking color feature of this
species in alcohol (and presumably also in life), i.e. the bright copper ring around the pupil of the eye. The speciic
epithet is a noun in apposition.
Remarks. No information is available on the habitat of this species; the collectors of the holotype and paratypes
are deceased. Evidently a small species; the smallest paratype is a fully mature female. One of seven schwenkii
complex species in the Red Sea, distinguished by high lateral-line scale counts, along with more than two rows of
teeth medially in the upper jaw, a wide dark band at the base of the anal in, and the copper ring around the pupil.
No DNA sequences are available for P. orbis.
Figure 28. Pempheris orbis, holotype, SMNHTAU P.5284, female, 85 mm SL, Red Sea, Gulf of Aqaba (H.A. Randall).
42
Pempheris pathirana Randall & Victor, n. sp.
Figure 29, Tables 1–3.
Holotype. USNM 432500, male, 129 mm SL, Sri Lanka, Trincomalee District, Koddiyar Pattu, Foul Point (on
E side of lighthouse base), 8°3l’36” N, 81°l9’ E, 0–2 m, Ceylon – University of Miami & Smithsonian Institution
Project 3818, rotenone, T. Iwamoto et al., May 15, 1970.
Paratypes. USNM 402255, female, 125 mm SL, Sri Lanka, North Eastern Province, Trincomalee, outside
harbor, large rock outcrop, about 50 yards offshore, 1–5 m, rotenone, W.F. Smith-Vaniz, June 26, 1969; USNM
402235, 9: 104–140 mm SL, same data as holotype; BPBM 41225, 121 mm SL, & SAIAB 200844, 2: 126–127
mm SL (formerly three specimens of USNM 379412), same collection data as USNM 402255 except date is July
1, 1969.
Description. Dorsal-in rays VI,9 (one paratype with VI,10); anal-in rays III,43 (38–43); pectoral-in rays
18 (17–19, strongly modal 18); lateral-line scales 57 (56–62); scales on nape, all of chest, and caudal peduncle
ctenoid, those below dorsal in cycloid; gill rakers 30 (29 or 30); body depth 2.3 (2.1–2.3) in SL; body width 2.8
(2.7–2.95) in depth; head length 3.2 (3.1–3.3) in SL; eye moderate in size, orbit diameter 7.2 (7.0 in 140-mm
paratype to 7.9 in 104-mm paratype); interorbital width 11.5 (11.3–12.5) in SL; jaws steeply oblique, the edge of
maxilla forming an angle of 63–68° to horizontal axis of body; upper jaw to each side of symphyseal gap with a
single row of slender recurved teeth within a narrow zone of very small nodular teeth; main row of teeth smaller
laterally, in an irregular row (nearly two rows in places); small slender teeth at front of lower jaw in three to ive
irregular rows, progressively more recurved inwardly; predorsal length 2.7 (2.65–2.7) in SL; longest dorsal-in
ray 3.7 (3.75–4.15) in SL; pectoral–in length 3.2 (3.15–3.3) in SL; prepelvic length 2.65 (2.6–2.7) in SL; pelvic
ins varying from reaching anus to just reaching origin of anal in, 5.75 (5.2–6.1) in SL, relatively longest in
smallest paratype. Color in alcohol as in Fig. 29; note the absence of black pigment at the base of the pectoral ins
(also lacking in axil), no black on base or margin of anal in, and black pigmentation forming a spot only distally
on the dorsal in. Color in life unknown.
Etymology. This species is named for Yohan Pathirana of Aquamarines International in Sri Lanka, who has
provided specimens and photographs of Pempheris from the island nation. He also had his divers search for P.
pathirana at Trincomalee, where it was collected in 1969 and 1970, but they failed to ind it.
Figure 29. Pempheris pathirana, holotype, USNM 432500, male, 129 mm, Trincomalee District, Sri Lanka (S. O’Hara).
43
Remarks. Pempheris pathirana is presently known only from Sri Lanka, but it might be expected in suitable
shallow reef habitat in southern India. Specimens from the 1969 and 1970 collections of this species from Sri
Lanka in the U.S. National Museum of Natural History were irst identiied as Pempheris nesogallica Cuvier in
C. & V. The two species appear very similar as preserved specimens. Pempheris nesogallica was described from
the Mascarene Islands and was recently found in KwaZulu-Natal. It has slightly longer predorsal and prepelvic
lengths (2.55–2.65 vs. 2.6–2.7 in P. pathirana); slightly longer pectoral ins (3.05–3.2 in SL vs. 3.15–3.3 in P.
pathirana); ctenoid vs. cycloid scales on side of caudal peduncle; a few cycloid scales above ctenoid scales on
side of chest of P. nesogallica (vs. all ctenoid on P. pathirana); a large black spot on base and axil of pectoral ins
of P. nesogallica (at most dark brown on P. pathirana); and 30–34 gill rakers for P. nesogallica (vs. 29 or 30 for
P. pathirana).
Pempheris pathirana is also a close relative of P. kruppi, found on the coast of Yemen and Socotra. Pempheris
kruppi differs in the steeper angle of the jaws, 37–41 anal-in soft rays (vs. 38–43 in P. pathirana), and in dentition.
Specimens of P. pathirana have a single row of recurved teeth anteriorly in the upper jaw, compared to two in
most other species in the rhomboidea species complex, followed in P. pathirana by many irregular rows of very
small teeth. Two related species are found in India: P. sarayu and P. mangula. Pempheris sarayu differs in having
a shorter pelvic in, 6.15 in SL, and a mode of 19 pectoral-in rays. Pempheris mangula has a characteristic dark
brown or dark red margin on the anal in. The representative of the schwenkii complex in India is P. bineeshi, and
in Sri Lanka the local representative is P. trinco. The unrelated P. malabarica is also found in the region. No DNA
sequences are available for P. pathirana.
Pempheris peza Randall & Victor, n. sp.
Figure 30, Tables 1–3.
Holotype. SAIAB 50637, male, 107 mm SL, Mozambique, Ponta do Ouro, 26.833° S, 32.8833° E, rotenone,
P.C. Heemstra, A. Wood, & A. Paterson, Nov. 9, 1995.
Paratypes. SAIAB 200761, 9: 75–114 mm SL, & BPBM 41192, 6: 91–110 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (36–40); pectoral-in rays 17 or 18; lateral-line scales
51 (50–59); lateral line moderately arched anteriorly, rising slightly more than 1/2 orbit diameter above level of
Figure 30. Pempheris peza, holotype, SAIAB 50637, male, 107 mm SL, Mozambique (H.A. Randall).
44
origin of lateral line; scales ctenoid on nape and below dorsal in, but cycloid on side of caudal peducle and on
chest, except for two or three ventroanterior ctenoid scales; gill rakers 28 (28–30); body depth 2.35 (2.2–2.35) in
SL, and compressed, the width 2.9 (2.9–3.2) in body depth; head length 3.2 (3.2–3.35) in SL; orbit diameter 7.5
(7.2–8.1) in SL; interorbital width 10.7 (10.2–12.5) in SL; jaws moderately oblique, angle of maxilla to horizontal
axis of body 68° (66–69°); no prominent knob at tip of lower jaw; lower jaw slightly to moderately projecting
when mouth irmly closed; a single row of sharply conical, recurved teeth anteriorly in upper jaw; a second more
anterior and more projecting row of teeth present laterally, reduced to a single row of progressively smaller teeth
half way back in jaw, and ending posteriorly as a row of small nodules; a narrow band of small teeth in three or
four rows medially in lower jaw, progressively more recurved inwardly, and ending laterally in jaw as a single
row of small nodules; tongue triangular with pointed tip; predorsal length 2.7 (2.65–2.7) in SL; no dorsal-in rays
fully intact; caudal concavity about 4 in head length; pectoral-in length 3.25 (3.0–3.25) in SL; prepelvic length
2.45 (2.4–2.5) in SL; pelvic ins nearly or just reaching anus, 6.3 (6.0–6.8) in SL. Color in alcohol as in Fig. 30;
scales brown with whitish edges; dorsal in with a broad anterior black margin expanding slightly to a distal spot;
caudal in with broad blackish border posteriorly and on dorsal and ventral margins; anal in with a broad, very
dark brown band at base, the outer part of in pale yellowish and nearly as broad; three blackish spots at margin
of in (one to three such spots on a few paratypes); these are not remnants of a black border, nor are they parasites
(though they could be of parasitic origin); iris light yellow, the upper half of the yellow dorsal part of the iris
abruptly black (irregular on holotype, but a straight margin on most paratypes). Color in life unknown.
Etymology. This species is named Pempheris peza from the Greek meaning border, in reference to the very
broad blackish border on all of the outer edge of the caudal in.
Remarks. The 16 specimens of the new species are all from a single collecting station at Ponta do Ouro, a
small coastal town 10 km north of the border of South Africa and Mozambique. Two of the collectors were asked
for details of the collection. They only recall that rotenone was used in relatively shallow water. Two other species
of the schwenkii complex occur along the southern Mozambique coast, P. ibo and P. connelli: P. ibo has many
fewer lateral-line scales (43–49), and P. connelli has a wider body and less marked caudal in. No DNA sequences
are available for P. peza.
Pempheris rochai Randall & Victor, n. sp.
Figure 31, Tables 1–3. BIN AAC6084 (in part).
Holotype. CAS 236522, female, 116.5 mm SL, Oman, south coast, Ras Hamar, 16°54’ N, 53°57’ E, spear,
L.A. Rocha, March 18, 2013.
Description. Dorsal-in rays VI,10; anal-in rays III,39; pectoral-in rays 17 or 18; lateral-line scales 59;
lateral line moderately arched anteriorly, rising 1/2 orbit diameter above level of origin of lateral line; outer layer
of scales almost totally missing (no cycloid vs. ctenoid scale characters possible); gill rakers 28; body moderately
deep, depth 2.25 in SL, and compressed, the width 3.1 in depth; head length 3.15 in SL; eye moderately large, orbit
diameter 6.8 in SL; interorbital width 11.6 in SL; jaws moderately oblique, the edge of maxilla forming an angle
of about 68° to horizontal axis of body; knob at tip of lower jaw bulges more posteriorly than anteriorly; upper
jaw with a single row of sharply conical, recurved teeth that are progressively smaller posteriorly to side of jaw,
ending as small nodules; lower jaw with a narrow band of very small conical teeth in three to ive rows medially,
progressively more recurved inwardly; predorsal length 2.65 in SL; longest dorsal-in rays broken; pectoral-in
length 3.0 in SL; prepelvic length 2.5 in SL; pelvic ins just reaching anus, 6.15 in SL. Color in alcohol as shown
in Fig. 31; the scales silvery with thin brown margin (when wet, they show lavender iridescence; however these
are the scales of the inner layer); leading edge of dorsal in dark brown, broadening to a dark spot distally; dark
brown band at base of anal in broad; tips of heavily abraded anal rays with traces of black pigment, suggesting a
blackish margin when intact; caudal in probably fully bordered in dark brown or black in life; pectoral ins with
45
Figure 31. Pempheris rochai, holotype, CAS 236522, female, 116.5 mm, Ras Hamar, Oman (H.A. Randall). Spear wound
below pectoral in.
a narrow blackish band at base of rays, except lateral two or three rays. Color in life unknown.
Etymology. This species is named for the collector, Dr. Luiz A. Rocha, Curator of Ichthyology at the California
Academy of Sciences, in recognition of his broad body of ichthyological research on the phylogenetics of reef
ishes.
Remarks. Species of Pempheris typically have 9 dorsal soft rays, but one can expect an occasional rare
individual of any species of the genus to have 10. The holotype of P. rochai, with 10 dorsal soft rays, may be such
an individual, or 10 may be the true count for the species. More specimens will provide the answer. It is distinctive
for its single row of enlarged curved teeth medially in the upper jaw (vs. two or three in other species), relatively
large eye, and the unusually broad dark brown band at the base of the anal in. The barcode mtDNA COI sequence
of the holotype of P. rochai is the same as P. connelli from South Africa and P. hollemani from Madagascar,
indicating either a recent split of closely related species or some degree of gene low across the western Indian
Ocean between the three species (Fig. 1).
Pempheris rubricauda Randall & Victor, n. sp.
Figure 32, Tables 1–3. BIN AAI3650 (in part).
Holotype. MNHN 2015-003, 72 mm SL, male, Madagascar, Nosy Be, Nosy Vorona, E side, 13°25’28” S, 48°
21’52” E, 3 m, rotenone, H. Bruggemann, N. Hubert, & S. Planes, May 8, 2008.
Paratypes. URUN 2008-4281, 3: 68.5–77 mm SL, & BPBM 41199, 72 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,36 (36–41); pectoral-in rays 17 or 18 (modally 18);
lateral-line scales 56 (55–57); gill rakers 28 (28 or 29); body depth 2.3 (2.2–2.45) in SL; body width 2.75 (2.75–
2.9) in body depth; head length 3.2 (3.05–3.3) in SL; eye relatively small, orbit diameter 8.1 (7.6–8.2) in SL;
interorbital width 11.9 (11.2–12.0) in SL; mouth oblique, forming an angle of 70° to horizontal axis of body; tip
of lower jaw slightly anterior to upper jaw when mouth fully closed; teeth in upper jaw in two rows medially,
soon narrowing to a single row; teeth in lower jaw very small and densely spaced, in a band of three or four rows
medially, progressively more retrorse posteriorly, those of inner row largest and nearly horizontal; outer layer of
scales nearly completely absent on most specimens, but enough remain on holotype to determine that those on
nape and in irst row below dorsal in are cycloid; side of chest about three-fourths cycloid, the ctenoid scales
46
Figure 32. Pempheris rubricauda, holotype, MNHN 2015-003, male, 72 mm, Nosy Be, Madagascar (H. Bruggemann).
ventroanteriorly; scales on side of caudal peduncle cycloid, those dorsally and ventrally ctenoid; predorsal length
2.65 (2.7) in SL; dorsal and caudal ins too damaged for measurement; pectoral-in length 3.45 (3.4–3.6) in SL;
prepelvic length 2.4–2.5 in SL; pelvic ins nearly or just reaching origin of anal in, 5.8 (5.8–6.0) in SL. Color
in alcohol light gray-brown below lateral line, the scale edges darker from a concentration of very small dark
brown dots; scale edges blackish above lateral line and on snout; dorsal in dusky yellowish with blackish anterior
margin and black apical spot; caudal in yellowish with blackish outer margin; anal in pale yellowish with a
narrow dark brown band at base and a broad dusky margin; paired ins yellowish, the pectoral ins without a dark
spot at base; iris dark bluish gray. Color of holotype when fresh in Fig. 32.
Etymology. This species is named Pempheris rubricauda from the Latin for red-tailed, in reference to the
bright red-orange coloration of the broad central part of the caudal in.
Remarks. The new species is presently known only from NW Madagascar. The barcode mtDNA COI
sequences of P. rubricauda (along with a sequence from P. ternay from Seychelles) form a barcode lineage very
close, only 0.66% divergent, to the lineage comprising P. tominagai from the Red Sea and P. xanthomma from
Yemen. That set of four species is sister to two other clades of schwenkii-complex lineages that occupy nonoverlapping ranges in the Indo-Paciic: 3.4% divergent from the lineage that comprises P. hadra from Maldives,
P. bineeshi from India, and probably true P. schwenkii from Indonesia (Fig. 1), as well as 3.6% divergent from a
lineage of P. aff. schwenkii from the Paciic Ocean.
Pempheris sergey Randall & Victor, n. sp.
Figure 33, Tables 1–3.
Holotype. SMF 35717 (KAU 13-671), 103 mm SL, Red Sea, Saudi Arabia, Sharm Obhur, in front of Marine
Station, 21°42’34.00” N, 39°05’45.70” E, steep slope, 12–17 m, rotenone, S.V. Bogorodsky, July 1, 2013.
Paratypes. SMF 35718 (KAU 13-672), 85 mm SL, KAUMM 370 (KAU 13-673), 104 mm SL, KAUMM 371
(KAU 13-675), 83 mm SL, & BPBM 41229 (KAU 13-674), 89 mm SL, all same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,38 (39–42); pectoral-in rays 17 (17 or 18); lateral-line
scales 51 (53–55); scales on nape, below dorsal in, and side of caudal peduncle cycloid, scales on side of chest
47
Figure 33. Pempheris sergey, holotype, SMF 35717, 103 mm SL, Sharm Obhur, Saudi Arabia, Red Sea (S.V. Bogorodsky).
cycloid except for a few ctenoid scales ventroanteriorly; gill rakers 27 (26–27); body depth 2.5 (2.5–2.55) in
SL; body width 2.65 (2.65–2.8) in body depth; head length 3.4 (3.3–3.35) in SL; eye relatively small, the orbit
diameter 7.65 (7.35–7.55) in SL; interorbital width 10.6 (11.2–11.8) in SL; mouth strongly oblique, forming an
angle of 60–65° to horizontal axis of body; recurved conical teeth in a double row in upper jaw, those of upper row
more projecting, those of lower row more strongly recurved; double row continues progressively smaller, nearly
to end of jaw where it ends as tiny teeth in a single row; teeth in lower jaw very small and slender, in a band of four
or ive, close-set, irregular rows where broadest medially, progressively more recurved inwardly; numerous black
dots in spaces between teeth; teeth smaller and in fewer rows posteriorly in jaw, ending in a few extremely small
teeth in a single row; tongue narrowly triangular or with sides slightly incurved, the tip pointed to slightly rounded;
upper lip appears smooth, but surface consists of small, close-set papillae when viewed by microscope, papillae
more evident on lower lip; predorsal length 2.7 (2.65–2.75) in SL; longest dorsal soft ray 4.05 (4.15–4.35) in SL;
caudal-in length 3.55 in SL, the caudal concavity 11.1 in SL (in damaged in all paratypes); pectoral-in length 3.1
(3.25–3.35) in SL; prepelvic length 2.6 (2.5–2.6) in SL; pelvic ins nearly or just reaching anus, 5.9 (5.75–6.3) in
SL. Color in alcohol yellowish brown, the scales narrowly darker brown; dorsal in translucent yellowish, the rays
edged in dark brown, with a broad blackish anterior margin that expands to a large distal black spot; caudal in
brownish yellow, with a very broad blackish posterior margin, the upper and lower dark margins half as wide and
not as strongly pigmented; anal in pale yellowish with a dark brown basal band that narrows posteriorly (band
is narrower when fresh; also there is a darkened border on the anal in when fresh that is no longer apparent in
alcohol); paired ins pale yellowish, pectoral ins with a dark brown band at base; eye with yellow iris when fresh,
dark bluish gray when preserved. Color when fresh as in Fig. 33.
Etymology. This species is named for the collector, Sergey V. Bogorodsky, in recognition of his extensive
research documenting the ishes of the Red Sea. The speciic epithet is a noun in apposition.
Remarks. This species is known from a single collection at Obhur Creek, 36 km north of Jeddah, Saudi
Arabia; the topography is a broad tidal channel with steep marginal reefs. S. Bogorodsky observed feeding on
zooplankton by this species and P. lavicycla near the surface from the light of a nearby boat landing. Pempheris
sergey is one of seven species of the schwenkii complex from the Red Sea; it is distinguished by color in life,
having two rows of teeth medially in the upper jaw, and a slender body, i.e. the body depth more than 2.5 in SL.
The color is similar to the closely related P. viridis: morphological differences include three rows of teeth medially
in the upper jaw and a straight dorsal proile in the latter. Both P. sergey and P. viridis have relatively narrow dark
bands at the base of the anal in, vs. P. xanthomma and some others. Pempheris tominagai differs by having three
rows of teeth in the medial upper jaw, a wider body, and at most dusky edges to the caudal in. Mitochondrial
DNA sequences from the type specimen (obtained during review) match those of P. tominagai; more specimens
are needed to resolve the genetic relationships in this group.
48
Pempheris shimoni Randall & Victor, n. sp.
Figure 34, Tables 1–3.
Holotype. SAIAB 51257, female, 133 mm SL, Kenya, Shimoni, 4.6500° S, 39.8333° E, J.L.B. & M.M.
Smith, Jan. 1, 1952.
Paratypes. SAIAB 51256, 7: 31–48 mm SL, same data as holotype but Oct. 1, 1950; SAIAB 200843, 3: 41–
51 mm SL, same data as holotype; USNM 402231, 65 mm SL, Kenya, Andromache Reef, just south of entrance to
Port Kilindini, Mombasa Harbor, off Ras Mwa Kisenge, 4° 5’5” S, 39° 40’39” E, reef lat, 0–1.3 m, dead coral &
sand, rotenone, H. Fehlmann et al., Anton Bruun Cruise 9, International Indian Ocean Expedition, Nov. 15, 1964.
Description. Dorsal-in rays VI,9; anal-in rays III,40 (37–41); pectoral-in rays 17 (17–19, modally 18);
lateral-line scales 56 (53–58); gill rakers 30 (28–31); body depth 2.2 in SL; body width 3.0 in body depth; head
length 3.25 in SL; eye diameter 7.3 in SL; interorbital width 11.3 in SL; mouth oblique, forming an angle of 63°
to horizontal axis of body; upper-jaw dentition with two irregular rows of sharply recurved teeth on each side of
symphyseal gap, reduced to a single row of gradually smaller teeth about half way back in jaw; lower jaw with
a close-set band of four or ive irregular rows of very small teeth that seem to be only nodular, but microscopic
inspection reveals them to be sharply pointed, progressively larger, and more recurved inwardly in mouth; tongue
deeply concave on each side, resulting in a narrow, rectangular, anterior part, that ends in a rounded lexible lap;
lower jaw projecting when mouth fully closed; scales on nape, below dorsal in, and on about ventroanterior twothirds of chest ctenoid; predorsal length 2.7 in SL; dorsal and caudal ins damaged; pectoral-in length 3.15 in SL;
prepelvic length 2.5 in SL; pelvic ins reaching anus, 6.05 in SL. Color in alcohol as in Fig. 34; the reddish brown
band across the base of the pectoral ins was probably dark brown in life. Color in life unknown.
Etymology. This species is named for the type locality, the small town of Shimoni, a port near the border of
Kenya and Tanzania. The speciic epithet is a noun in apposition.
Remarks. Measurements in the above description are from our only adult specimen (the paratypes are all
juveniles). We regret not having more information on the ish collection made at Shimoni by the Smiths 63 years
ago. No DNA sequences are available for P. shimoni.
Figure 34. Pempheris shimoni, holotype, SAIAB 51257, female, 133 mm SL, Shimoni, Kenya (H.A. Randall).
49
Pempheris shirleen Randall & Victor, n. sp.
Figure 35, Tables 1–3.
Holotype. USNM 402228, female, 105 mm SL, Red Sea, Egypt, Hurghada, 300 yards north of pier of Institute
of Oceanography & Fisheries, 27°16’38” N, 33°47’1” E, reef pool surrounded by profuse staghorn coral, rotenone,
H. Fehlmann et al., Jan. 4, 1965.
Paratype. USNM 436759, female, 103 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,37 (38); pectoral-in rays 17 or 18; lateral-line scales 50
(54); scales on nape, below dorsal in, and side of caudal peduncle cycloid, scales on side of chest cycloid except
for a few ctenoid scales ventroanteriorly; gill rakers 28 (27); body depth 2.35 (2.4) in SL; body width 2.75 (2.85)
in body depth; head length 3.25 (3.2) in SL; eye relatively small, the orbit diameter 7.35 (7.45) in SL; interorbital
width 11.8 (11.5) in SL; mouth oblique, forming an angle of 66° (63°) to horizontal axis of body; teeth medially
in upper jaw small, progressively more recurved inwardly, in three irregular rows (many medial teeth of holotype
and nearly all of paratype missing); teeth reduced to a single row about half way to end of jaw; teeth in lower jaw
smaller, more slender, in a band of three or four, close-set, irregular rows where broadest, and progressively more
recurved inwardly; teeth smaller and in fewer rows posteriorly in jaw, ending in a few extremely small teeth in
a single row; tongue triangular with nearly straight sides, the tip pointed to slightly rounded; upper lip appears
smooth, but surface consists of small, close-set papillae when viewed by microscope; predorsal length 2.6 (2.7)
in SL; all ins except the pelvics too damaged for measurement; prepelvic length 2.5 (2.55) in SL; pelvic ins very
short, not reaching anus, 7.45 (7.3) in SL. Color in alcohol as in Fig. 35. Color in life unknown.
Etymology. This species is named for Shirleen Smith of the Division of Fishes, United States National
Museum of Natural History, in appreciation of the many loans of Pempheris and other ishes that she has prepared
for the senior author. The speciic epithet is a noun in apposition.
Remarks. This species is known from a single collection at Hurghada in Egypt. Pempheris shirleen is one
of seven species of the schwenkii complex from the Red Sea; it is distinguished by very short pelvic ins, having
three rows of teeth medially in the upper jaw, cycloid scales on the nape and most of chest, a thinner body (body
width more than 2.7 in body depth), and a moderately deep body (body depth less than 2.4 in SL). No DNA
sequences are available for P. shirleen.
Figure 35. Pempheris shirleen, holotype,USNM 402228, female, 105 mm SL, Hurghada, Egypt, Red Sea (H.A. Randall).
50
Pempheris smithorum Randall & Victor, n. sp.
Figure 36, Tables 1–3.
Holotype. SAIAB 51259, male, 126.5 mm SL, Tanzania, Zanzibar, Chumbe Island, 6.1666° S, 39.1833° E
J.L.B. & M.M. Smith, Sept. 5, 1952.
Paratypes. BPBM 37176, female, 139 mm SL, Tanzania, Maia Island, Chole Island, rocky islet, 0–2 m,
small cave, rotenone, J.E. Randall, L.G. Westfeldt, & A. Klosser, Dec. 11, 1973.
Description. Dorsal-in rays VI,9; anal-in rays III,41 (40); pectoral-in rays 17 (17 or 18); lateral-line scales
59 (56); gill rakers 30 (31); body depth 2.35 (2.3) in SL; body width 2.6 (2.8) in body depth; head length 3.25 (3.3)
in SL; eye diameter 7.0 (7.4) in SL; interorbital width 11.7 (11.8) in SL; mouth oblique, forming an angle of 61–
66° to horizontal axis of body; upper-jaw dentition typical of the genus, with two irregular rows of recurved teeth
on each side of symphyseal gap, one protruding forward, the other strongly recurved, reduced to a single row of
gradually smaller teeth nearly half way back in jaw; tip of lower jaw expanded to a distinct knob, extending both
anteriorly and posteriorly, and strongly protruding when jaws fully closed; lips with well-developed bristle-like
papillae; lower-jaw teeth to either side large for the genus, beginning medially with seven rows, those of outer row
erect or slightly recurved, the remaining inner teeth progressively longer and more recurved, teeth smaller and in
fewer rows posteriorly in jaw; tongue triangular, straight-sided, with irm pointed tip; scales on nape, below dorsal
in, and on chest ctenoid; scales on side of caudal peduncle cycloid; predorsal length 2.7 in SL; prepelvic length
2.45–2.5 in SL; dorsal, caudal, and pectoral ins damaged; pelvic ins reaching anus, 6.35 in SL. Color of holotype
in alcohol as in Fig. 36; large spot at pectoral-in base probably dark brown in life. Color in life unknown.
Etymology. This species is named in honor of James Leonard Brierly Smith and Margaret Mary Smith, not
only for being the collectors of the holotype, but also for the major contributions they have made in pioneering
ichthyology in South Africa, describing hundreds of species, and writing numerous publications on the ishes of
southern Africa.
Remarks. Two specimens of Pempheris were collected at Chumbe Island in 1952 by the Smiths, each the
holotype of a new ish, i.e. the present species and P. leiolepis, the former with ctenoid scales on the chest and the
latter with cycloid scales and three rows of upper-jaw teeth. The paratype of P. smithorum was collected by the
irst author and associates from nearby Maia Island with no suspicion at the time that it might be an undescribed
species. No photographs and no color notes were taken. It is more copper-colored than the holotype. No DNA
sequences are available for P. smithorum.
Figure 36. Pempheris smithorum, holotype, SAIAB 51259, male, 126.5 mm SL, Zanzibar (H.A. Randall).
51
Pempheris tau Randall & Victor, n. sp.
Figure 37, Tables 1–3.
Holotype. HUJ 10979, male, 95 mm SL, Egypt, Gulf of Aqaba, east coast of Sinai Peninsula, Nuweiba, A.
Ben-Tuvia, Aug. 3, 1976.
Paratype. BPBM 41232, female, 99 mm SL, same data as holotype; SMNHTAU P.3650, male, 116.5 mm
SL, Sharm el Sheikh, Naama Bay (Marsa el At), 27°54’38” N, 34°19’44” E, Egypt, Red Sea, L. Fishelson, Oct.
16, 1968.
Description. Dorsal-in rays VI,9; anal-in rays III,39 (38 or 39); pectoral-in rays 17 (17 or 18); lateral-line
scales 50 (52–55); scales on nape, below dorsal in, and side of caudal peduncle cycloid, scales on side of chest
cycloid except for a few ctenoid scales ventroanteriorly; gill rakers 31 (30–33); body depth 2.5 (2.3–2.6) in SL;
body width 2.45 (2.55–2.7) in body depth; head length 3.3 (3.2–3.3) in SL; dorsal proile of head slightly convex;
eye very small, orbit diameter 7.9 (8.1–8.65) in SL; interorbital width 11.5 (11.3–11.8) in SL; mouth oblique,
forming an angle of 63°(62–63°) to horizontal axis of body; tip of lower jaw projecting anterior to upper jaw when
mouth fully closed; teeth conical and recurved, those medially in upper jaw in three irregular rows, soon reduced
to two rows, and inally to a single row of very small teeth near end of jaw; teeth in lower jaw similar but much
smaller, more slender, in three to four irregular rows medially, then progressively fewer and smaller toward end
of jaw; lips yellowish brown, densely stippled with small purple papillae; tongue triangular, straight-sided, with
pointed tip; predorsal length 2.75 (2.7–2.75) in SL; prepelvic length 2.5 (2.5–2.6) in SL; longest dorsal-in ray
3.8 (4.0–4.1) in SL; all caudal-in rays broken; pectoral-in length 3.05 (3.15–3.3) in SL; pelvic ins reaching or
extending slightly posterior to anus, their length 6.3 (6.8–6.85) in SL. Color in alcohol as in Fig. 37. Color in life
unknown.
Etymology. This species is named for the acronym of Tel-Aviv University, in appreciation of their large loan
of Red Sea specimens of the genus Pempheris, including the largest specimen of the present species. The speciic
epithet is a noun in apposition.
Remarks. The most distinctive morphological character of this species is the small eye, smaller than for the
other six members of the schwenkii complex in the Red Sea. Only P. micromma from Oman has a smaller eye;
note that the preserved holotype of P. micromma in Fig. 25 has a different body shape and has different tooth row
patterns. Pempheris zajonzi from Yemen shares the small eye and three tooth rows in the upper jaw, but has a
larger head and fewer gill rakers. No DNA sequences are available for P. tau.
Figure 37. Pempheris tau, holotype, HUJ 10979, male, 95 mm SL, near Nuweiba, Gulf of Aqaba, Egypt (H.A. Randall).
52
Pempheris ternay Randall & Victor, n. sp.
Figure 38, Tables 1–3. BIN AAI3650 (in part).
Pempheris tominagai [non Koeda, Yoshino, Imai & Tachihara] Koeda et al. 2014: 317, in part.
Holotype. SAIAB 77935, male, 91 mm SL, Seychelles, Mahé, Ternay Marine National Park, Cape Matoopa,
southern end, 4°39.03’ S, 55°22.71’ E, coral reef, surrounded by sand, 6 m, spear, M. Smale, A. Bentley, E.
Heemstra, & M. Mwale, April 21, 2005.
Description. Dorsal-in rays VI,9; anal-in rays III,36; pectoral-in rays 17; lateral-line scales 51; gill rakers 28;
body depth 2.45 in SL; body width 2.85 in body depth; head length 3.25 in SL; eye relatively small and precisely
circular, the orbit diameter 7.45 in SL; lips smooth; interorbital width 11.3 in SL; mouth oblique, forming an
angle of 60° to horizontal axis of body; tip of lower jaw slightly anterior to upper jaw when mouth fully closed;
teeth small, pointed, and recurved, those in upper jaw in two rows medially, the teeth of the upper row projecting
forward, those of the lower row curving downward; teeth progressively smaller and narrowing to a single row
about half way back in jaw; teeth in lower jaw very small and densely spaced, in a band of three or four rows
medially, progressively more retrorse inwardly, those of inner row largest and nearly horizontal; outer layer of
scales totally missing, and many of inner layer as well; predorsal length 2.65 in SL; longest dorsal-in ray (from
photograph) 2.7 in SL; pectoral-in length 3.0 in SL; prepelvic length 2.5 in SL; pelvic ins just reaching origin of
anal in, 6.0 in SL. Scaleless color silvery gray with a vertical brown line on each scale pocket; color when fresh
as shown in Fig. 38.
Etymology. This species is named for the type location, Ternay Marine National Park on the island of Mahé,
where one can no longer collect ishes, but where one can enjoy viewing and photographing them at close range.
The speciic epithet is a noun in apposition.
Remarks. This species shares the black-bordered red caudal in with P. rubricauda from Madagascar; it is
separated by length of the pectoral ins, the number of lateral-line scales, and the angle of the maxilla. Pempheris
ternay is the second schwenkii-complex species identiied from Seychelles; P. argyrea shares most meristics with
P. ternay, but does not have the prominent red tail. The barcode mtDNA COI sequence of P. ternay is almost the
same as for P. rubricauda, and the two form a lineage diverging by only 0.66% from the lineage including both
P. tominagai from the Red Sea and P. xanthomma from Yemen (Fig. 1). That set of four species diverges by 3.4%
from the lineage comprising P. hadra from Maldives, P. bineeshi from India, and probable true P. schwenkii from
Bali, Indonesia.
Figure 38. Pempheris ternay, holotype, SAIAB 77935, male, 91 mm SL, Mahé, Seychelles (A. Bentley).
53
Pempheris tilman Randall & Victor, n. sp.
Figure 39, Tables 1–3.
Holotype. SMF 35779 [ex SMF 23763], male, 96.5 mm SL, Red Sea, Jordan, Aqaba, F. Krupp, 1980.
Paratypes. BPBM 41233, female, 75 mm SL, & SMF 23763, female, 88 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,40 (36–41); pectoral-in rays 17 (17); lateral-line scales
50 (49–52); gill rakers 30 (29–30); body depth 2.45 (2.4–2.55) in SL; body width 2.5 (2.55–2.65) in body depth;
head length 3.0 (2.9–3.2) in SL; dorsal proile of head slightly convex; eye small, orbit diameter 7.7 (7.6–7.65)
in SL; interorbital width 11.1 (11.4–11.8) in SL; mouth oblique, forming an angle of 64° (65–67°) to horizontal
axis of body; tip of lower jaw projecting anterior to upper jaw when mouth fully closed (very slightly on smallest
paratype); teeth conical and recurved, those at front of upper jaw in two irregular rows, reduced opposite nostrils
to a single irregular row of progressively smaller teeth ending as a row of tiny nodular teeth in lower jaw very
small, in three to four irregular rows medially, progressively fewer and smaller toward end of jaw; lips brown with
numerous, very small, dark brown papillae; tongue a straight-sided triangle with pointed tip; many outer-layer
scales missing from nape, below dorsal in, and on chest of all type specimens, but enough to conclude that these
areas have cycloid scales, except for a few ctenoid scales ventrally on chest near opercle; predorsal length 2.75
(2.75–2.8) in SL; prepelvic length 2.7 (2.7–2.8) in SL; longest dorsal-in ray 3.85 (broken on paratypes) in SL;
all caudal-in rays damaged; pectoral-ins broken on all but 88-mm paratype, its length 3.6 in SL; pelvic ins just
reaching anus, their length 6.2 (6.45–6.8) in SL. Color of holotype in alcohol as in Fig. 39; color in life unknown.
Etymology. This species is named for Dr. Tilman J. Alpermann of the Senckenberg Research Institute and
Natural History Museum Frankfurt in recognition of his studies on Red Sea ishes and his assistance with loans
and collection data from the Museum. The speciic epithet is a noun in apposition.
Remarks. The three known specimens of this species came on loan with the original label only as follows:
“Pempheris vanicolensis, Roten Meer, Aqaba, 1980”. They represent one of the seven schwenkii complex species
in the Red Sea, due to their small adult size (the smallest paratype is a fully mature female), slender body, and
dark band at the base of the anal in. They are distinguished from other members of the complex in the Red Sea
by having two rows of teeth medially in the upper jaw, a short prepelvic length, and the dark line on the pectoralin base at the insertion of the rays (vs. a band lower on the base). No DNA sequences are available for P. tilman.
Figure 39. Pempheris tilman, holotype, SMF 35779, male, 96.5 mm SL, Aqaba, Jordan, Red Sea (H.A. Randall).
54
Pempheris tiran Randall & Victor, n. sp.
Figure 40, Tables 1–3.
Holotype. CAS 236818, male, 122 mm SL, Red Sea, Saudi Arabia, Strait of Tiran, 28° 11.051’ N, 34° 38.372’
E, wreck of metal ferry on reef, small caves on side of reef, 6 m, spear, L. Rocha, M. Berumen, M. Priest, & T.
Sinclair-Taylor, Oct. 1, 2013.
Paratype. BPBM 41230, female, 121 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,42 (44); pectoral-in rays 18 (one with 17 on one side);
lateral-line scales 62 (64); scales on nape, below dorsal in, and side of caudal peduncle cycloid, scales on side of
chest missing; gill rakers 31 (32); body depth 2.5 (2.45) in SL; body width 2.7 (2.7) in body depth; head length
3.45 (3.4) in SL; eye relatively small, the orbit diameter 7.95 (7.7) in SL; interorbital width 10.8 (10.9) in SL;
mouth strongly oblique, forming an angle of 64° (65°) to horizontal axis of body; teeth large for the genus, those
of medial third of upper jaw of male in three irregular rows, the upper-row teeth conical and forward-projecting,
those of inner rows strongly recurved and inwardly projecting; remaining teeth in jaw in two rows, progressively
smaller, except for a few tiny teeth in a single row near end of jaw; teeth of female about half the size of those
of male, those of medial third of upper jaw in a single irregular row, the remaining teeth in two irregular rows,
except for very small teeth in a single irregular row for last fourth of jaw; band of teeth in lower jaw of both sexes
strongly outcurved medial to narrow symphysis, the teeth in four rows, progressively larger and more recurved
inwardly in female; largest teeth of male in two outer rows, erect or slightly recurved; outer medial teeth of lower
jaw visible in both sexes when mouth fully closed; upper lip relatively smooth, the surface with close-set, low
papillae; lower lip with smaller and more widely separated papillae; tongue with sides slightly incurved, the tip
rounded and lexible; predorsal length 2.8 (2.75) in SL; longest dorsal-in soft ray 3.75 (4.05) in SL; tips of caudal
lobes broken, but slight curvature of broad middle part of in indicates a low caudal concavity; prepelvic length
2.8 (2.75) in SL; pelvic ins extending nearly to origin of anal in, in length 5.55 (5.65) in SL. Color in alcohol as
in Fig. 40, dark brown, scale edges light gray; row of scales above and below lateral line darker than remaining
scales of body; dorsal in grayish brown with a darker brown anterior margin leading to a large black apical spot
(nearly half length of anterior edge of in); caudal in dark gray-brown with a very broad black posterior margin
Figure 40. Pempheris tiran, holotype, CAS 236818, male, 122 mm SL, Strait of Tiran, Saudi Arabia, Red Sea (H.A. Randall).
55
Figure 41. Collection location on wreck of ferry for type specimens of Pempheris tiran, Strait of Tiran, Saudi Arabia, Red
Sea (J. DiBattista).
(width nearly equal to pupil diameter) and narrow black dorsal and ventral margins; anal in dark brown at base,
this color merging with dark brown of body, a middle zone of light yellowish gray, and an outer black border
more than half in width anteriorly, progressively narrower posteriorly; paired ins light yellowish gray; iris bluish
black, greenish gray lens nearly encircled by two pale yellowish arcs. Color in life brownish gray with red-tinged
in membranes, iris with a golden ring around pupil, anal in translucent and colorless below black band along in
margin.
Etymology. This species is named for the type location, the Strait of Tiran between the tip of Sinai and Saudi
Arabia. The speciic epithet is a noun in apposition.
Remarks. The two type specimens were irst identiied as P. vanicolensis (a Paciic species) because of the
prominent black border on the anal in and the in-ray and lateral-line scale counts (P. vanicolensis can be quickly
distinguished by lacking the prominent dark spot around the base of the pectoral in). However, when the dentition
was examined, the new species showed a distinctive tooth pattern shared only with P. malabarica in the region: an
external patch of teeth on the lower jaw that is apparent when the jaws are closed. Pempheris tiran can be clearly
distinguished from P. malabarica by non-overlapping gill-raker counts and very different colors and marking
patterns. Both the markings and meristics of P. tiran are similar to those of P. bexillon, from the southwest Indian
Ocean (see Mooi & Randall 2014), in particular the black margin along the anal in; the main difference is the
distinctive external patch of teeth on the lower lip of P. tiran. In addition, P. bexillon has a predominantly yellow
coloration on the ins, not apparent on the fresh photograph of the holotype of P. tiran. Pempheris lavicycla is
found in the Red Sea and also shares the dark margin of the anal in, but has a bright yellow iris, a lower range of
lateral-line scale counts, and no external patch of teeth. No DNA sequences are available for P. tiran.
56
Pempheris trinco Randall & Victor, n. sp.
Figure 42, Tables 1–3.
Pempheris schwenkii [non Bleeker] Randall & Bineesh 2014: 36.
Holotype. BPBM 18771, male, 104 mm SL, Sri Lanka, Trincomalee, Dutch Point, substratum of large
boulders, 5 m, spear, J.E. Randall, April 2, 1975.
Paratypes. USNM 343709, 7: 90–107 mm SL, Sri Lanka, Trincomalee, outside harbor, large outcrop about
50 yards offshore, 1–5 m, rotenone, W.F. Smith-Vaniz, July 1, 1969; USNM 343764, 97 mm SL, Sri Lanka,
Negombo, P.C. Heemstra, Sept. 15, 1969; USNM 343760, 4: 93–106 mm SL, Sri Lanka, Weligama, 200 yards
offshore from rest house, 1–3 m, rotenone, C. Koenig, Feb. 14, 1970; USNM 343765, 91 mm SL, Sri Lanka,
Eastern Province, Batticaloa District, Korali Pattu, Passakudah Bay, reef 500 yards NE of outlet of small stream,
6–8 m, rotenone, T. Iwamoto, June 10, 1970; BPBM 41213, 4: 85–104 mm SL, same data as holotype; BPBM
27196, 2: 81–88.5 mm SL, Sri Lanka, Hikkaduwa, about one-fourth mile offshore, rocky substrate, 10–12 m,
rotenone, J.E. Randall, Feb. 24, 1979.
Description. Dorsal-in rays VI,9; anal-in rays III,40 (37–41); pectoral-in rays 17 or 18, modally 18; lateralline scales 50 (49–58); gill rakers 26 (26–28); body depth 2.45 (2.3–2.5) in SL; body width 2.6 in body depth;
head length 3.25 (3.2–3.3) in SL; eye diameter 7.4 (7.1–7.75) in SL; interorbital width 12.6 (12.6–12.9) in SL;
mouth oblique, forming an angle of 65° (61–68°) to horizontal axis of body; tip of lower jaw slightly anterior to
upper jaw when mouth fully closed; teeth in upper jaw in two rows medially, the outer row slightly recurved, the
inner larger and more strongly recurved, narrowing to a single row posteriorly; teeth in lower jaw similar, but
smaller, in three irregular rows and more strongly recurved; tongue strongly concave on sides, creating a slender,
slightly rounded tip; outer layer of scales nearly completely absent on most specimens, but enough scales remain
on holotype to determine that those on nape and in irst row below dorsal in are cycloid; scales on side of chest
cycloid, except for a few ctenoid scales ventroanteriorly; scales on side of caudal peduncle cycloid; predorsal
length 2.7 (2.6–2.7) in SL; longest dorsal-in ray 3.85 (3.5–3.95) in SL; caudal in damaged in all specimens; the
in is intact on photograph of holotype, showing that it is forked, with a caudal concavity about 5 in head length;
pectoral-in length 3.25 (3.0–3.35) in SL; prepelvic length 2.6 (2.55–2.65) in SL; pelvic ins varying from reaching
Figure 42. Pempheris trinco, holotype, BPBM 18771, male, 104 mm SL, Trincomalee, Sri Lanka (J.E. Randall).
57
anus to nearly reaching origin of anal in, their length 6.5 (5.7–6.55) in SL. Fig. 42 of the holotype shows the fresh
coloration with the scales intact; note the scale centers are silvery gray, and the scale edges copper-colored. A dark
band along the base of the pectoral in when fresh. Holotype in alcohol orangish brown (the brown from very ine
dark brown stippling), the scale edges silvery gray.
Etymology. This species is named for the type location, Trincomalee, which is both a district and a major port
city on the east coast of Sri Lanka. It is generally called Trinco by Sri Lankans. The speciic epithet is a noun in
apposition.
Remarks. Pempheris trinco is a relatively small species of the schwenkii complex; our largest specimen
measures only 107 mm SL. The species is usually encountered on reefs or rocky substrata in small aggregations.
Presently known only from Sri Lanka, but the range may extend more widely in the appropriate habitat. The
schwenkii-complex species in neighboring south India is P. bineeshi, which differs by having a shorter predorsal
and prepelvic distance, a narrower body, and a lower range of lateral-line scales. Other Pempheris species found
in Sri Lanka comprise P. pathirana of the rhomboidea complex, the unrelated P. malabarica (see Appendix Fig.
2), and P. mangula might be expected on the island as well. No DNA sequences are available for P. trinco.
Pempheris viridis Randall & Victor, n. sp.
Figure 43, Tables 1–3.
Holotype. SMF 33629 (KAU11-018), 79.5 mm SL, Red Sea, Saudi Arabia, off Al Lith, 20°15.111’ N,
39°57.825’ E, fringing reef, 3 m, a large coral block rising to within 0.5 m of surface; rotenone set in cave in coral
block, S.V. Bogorodsky, & T. J. Alpermann, March 28, 2011.
Description. Dorsal-in rays VI,9; anal-in rays III,37; pectoral-in rays 17 (one in removed); lateral-line
scales 53; scales on nape, below dorsal in, and side of caudal peduncle cycloid; scales on side of chest cycloid
except for 6 ctenoid scales in two rows on right side, and 12 ctenoid scales in four rows on left side at angle of
Figure 43. Pempheris viridis, holotype, SMF 3363, 79.5 mm SL, Al Lith, Saudi Arabia, Red Sea (S.V. Bogorodsky).
58
opercle; gill rakers 25; body depth 2.65 in SL; body width 2.7 in body depth; head length 3.2 in SL; dorsal proile
of head nearly straight; eye diameter 7.25 in SL; interorbital width 11.4 in SL; mouth steeply oblique, forming
an angle of 73° to horizontal axis of body; teeth medially in upper jaw small, slender, conical, and recurved in
three or four irregular rows, reduced laterally in jaw to two well-separated rows of larger, strongly recurved teeth,
and posteriorly to a single row of progressively smaller teeth, ending as small nodules; lower jaw with a band
of very small teeth at symphysis, gradually reduced posteriorly to a single row of tiny teeth; tongue triangular
in shape with pointed tip; anterior lateral line very highly arched, the vertical height above level of origin of
lateral line equal to two-thirds orbit diameter, 11.5 in SL; predorsal length 2.6 in SL; prepelvic length 2.5 in SL;
longest dorsal-in rays broken on holotype, but 4.1 in SL from photograph; caudal-in length 4.35 in SL; pectoralin length 3.4 in SL; pelvic ins reaching posterior margin of anus, their length 6.9 in SL. Color in alcohol: side
of body light orangish brown, the margin of scales darker; head and back grayish brown, the head darker; dark
brown band at base of anal in. Color when fresh as in Fig. 43.
Etymology. This species is named for its principal color, i.e. Latin viridis for green.
Remarks. Pempheris viridis is one of seven species in the schwenkii complex in the Red Sea. The unique
specimen was irst identiied by the senior author as P. tominagai, but differs from the underwater photograph of
the holotype of P. tominagai (Fig. 44) by its more slender body (the holotype described in Koeda et al. 2014 has
body depth 2.35 in SL), a straight dorsal proile (vs. convex in P. tominagai), a highly arched anterior lateral line
(in P. tominagai of equal length, the vertical rise is two-ifths orbit diameter, 17.6 in SL), and in color. Pempheris
viridis is bright metallic green, has a bright yellow eye, and has a very dark band at the rear margin of the caudal
in. Pempheris sergey is similar in color to P. viridis but differs by having two rows of teeth medially in the upper
jaw and a convex dorsal proile. Mitochondrial DNA sequences from the type specimens of P. viridis and P.
sergey (obtained during review) match those of P. tominagai; more specimens are needed to resolve the genetic
relationships in this group.
Figure 44. Pempheris tominagai, underwater photograph of holotype, Sharm el-Sheikh, Egypt, Red Sea (S.V. Bogorodsky).
59
Pempheris wilsoni Randall & Victor, n. sp.
Figures 45–48, Tables 1–3. BIN AAD1777 (in part).
Holotype. CAS 236523, male, 123 mm SL, Oman, Gulf of Oman, Fahal Island, 23°40’52” N, 58°30’11.3” E,
Station LAR-85, spear, L. Rocha, J. DiBattista, M. Priest, T. Sinclair, & J. Choat, March 28, 2013.
Description. Dorsal-in rays VI,9; anal-in rays III,45; pectoral-in rays 17; lateral-line scales 59; gill rakers
29; body depth 2.35 in SL; body width 2.7 in body depth; head length 3.45 in SL; eye diameter 7.3 in SL;
interorbital width 11.7 in SL; mouth oblique, forming an angle of 60° to horizontal axis of body; dentition as
described for P. smithorum, but teeth of lower jaw smaller; tongue concave-sided with rounded tip; outer layer of
scales almost entirely missing; predorsal length 2.7 in SL; prepelvic length 2.85 in SL; longest dorsal-in ray 3.4
in SL; caudal in damaged; pectoral-in length 3.15 in SL; pelvic ins reaching anus, 5.9 in SL. Color in alcohol as
in Fig. 45; black pigment on dorsal in conined to tip; caudal in with a posterior black border, but none on dorsal
and ventral margins of in. Color in life as in Figs. 46–48.
Etymology. This species is named for Keith D.P. Wilson, an expert on the dragonlies of China, who collected
ishes in Oman for our studies on Pempheris and took the photographs of Figs. 46 and 48.
Remarks. K.D.P. Wilson and Brian W. Darvell went to Oman at the request of the irst author to collect
and photograph species of Pempheris. Because they were passengers on a commercial dive boat, they had little
opportunity to collect ish specimens, succeeding only in three of P. darvelli described in Randall & Victor (2014).
We were fortunate to ind a specimen of P. wilsoni in a loan from the California Academy of Sciences for the
holotype. It was collected at Fahal Island in the Gulf of Oman, about 4 km from the dive sites of Wilson and
Darvell. The label indicates ive collectors. All divers were spearing various species of ishes, so no attempt was
made to link each specimen to a collector.
The underwater photographs of Figs. 46–48 were identiied as P. wilsoni by eye size, the 60° angle of the
maxilla, predorsal and prepelvic lengths, lengths of the longest rays of the dorsal in as well as the paired ins,
and the dark markings on the ins. Figs. 47 and 48 were irst believed to represent different species, but counts
and measurements from the photographs fail to show any differences, and the group in Fig. 47 has individuals of
intermediate color.
Figure 45. Pempheris wilsoni, holotype, CAS 236523, male, 123 mm SL, Fahal Island, Oman (H.A. Randall).
60
Figure 46. Pempheris wilsoni, underwater photograph, Oman (K.D.P. Wilson).
Figure 47. Pempheris wilsoni, aggregation, Gulf of Oman (B.W. Darvell).
61
Figure 48. Pempheris wilsoni, underwater photograph, Oman (K.D.P. Wilson).
The barcode mtDNA COI sequence of the holotype of P. wilsoni is the same as several other species of the
rhomboidea complex, mostly species from other locations in the region, i.e. P. rhomboidea from the Red Sea
and Mediterranean, P. russellii from Pakistan, P. eatoni from southern Africa, but also P. kuriamuria which is
described from the southern coast of Oman (vs. P. wilsoni from the northern coast in the Gulf of Oman)(Fig. 1).
Pempheris xanthomma Randall & Victor, n. sp.
Figures 49 & 50, Tables 1–3. BIN AAI3650 (in part).
Pempheris tominagai [non Koeda, Yoshino, Imai & Tachihara] Randall et al. 2014: 387.
Holotype. SMF 35712, female, 96 mm SL, Gulf of Aden, Yemen, Ras Majdaha, 14°00.868’ N, 48°25.572’
E, steep rocky slope to 6 m, collected with rotenone from 8–11 m, live coral cover less than 20%, isolated corals,
mainly Porites, some Pocillopora, rotenone, F. Krupp, M. Schneider, & E. Lavergne, May 20, 2005.
Paratypes. SMF 35489, 10: 57–100 mm SL, NHCY-P-10, 9: 58–98 mm SL, BPBM 41227, 74 mm SL, &
USNM 436600, 81 mm SL, all with same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,38 (36–41); pectoral-in rays 17 (16–18, strongly modal
17); lateral-line scales 55 (49–57); scales cycloid on nape, below dorsal in, on side of caudal peduncle, and on
chest except from a few scales to as many as one-fourth ctenoid ventroanteriorly; gill rakers 28 (26–29); body
depth 2.35 (2.15–2.45) in SL; body width 3.1 (2.45–3.5) in body depth; head length 3.25 (3.05–3.35) in SL;
eye relatively small, the orbit diameter 7.5 (7.55–7.75) in SL; interorbital width 12.0 (10.9–13.2) in SL; mouth
strongly oblique, forming an angle of 69° (67–70°) to horizontal axis of body; recurved, conical teeth in a double
62
Figure 49. Pempheris xanthomma, holotype, SMF 35712, female, 96 mm SL, Gulf of Aden, Yemen, off Majdaha (H.A.
Randall).
row in upper jaw, those of upper row more projecting, those of lower row more strongly recurved; double row
continuing nearly to end of jaw where ending in a cluster of very small nodules; teeth in lower jaw much smaller,
in four to ive rows medially, progressively more recurved inwardly; tongue an equilateral triangle; upper lip
with tiny papillae, most separated by a distance two or more times the diameter of the papillae; predorsal length
2.8 (2.7–2.8) in SL; longest dorsal-in ray 4.0 (3.9–4.4) in SL; caudal in moderately forked (none fully intact);
pectoral-in length 3.2 (3.15–3.35) in SL; prepelvic length 2.55 (2.5–2.6) in SL; pelvic ins reaching or extending
slightly posterior to anus, 6.0 (5.7–6.8) in SL. Color in alcohol as in Fig. 49; dark brown to black band on anterior
margin of dorsal in expanded to a distal spot on only four of the larger type specimens, including holotype; dark
band at base of anal in broad; a narrow dark brown band across base of pectoral ins; scales of pectoral and
abdominal region of large adults with light golden centers and brown margins; iris of eye yellow, except for a
black linear dorsal rim, the yellow lighter and brighter around pupil. Color in life silvery to brownish with a broad
dark band at anal-in base; orangish caudal in without a dark posterior margin (Fig. 50).
Figure 50. Pempheris xanthomma, underwater photograph, except P. kruppi on the far right with tail cropped, Ras Majdaha,
Yemen (M.S. Aideed).
63
Etymology. This species is named Pempheris xanthomma from the Greek for yellow-eyed, in reference to the
dominant color of the iris in both live individuals and preserved specimens.
Remarks. This species was irst identiied by us as P. tominagai. Pempheris xanthomma is distinguished from
the seven schwenkii-complex species in the neighboring Red Sea by having a particularly wide dark band at the
base of the anal in and two rows of teeth medially in the upper jaw. The new species is separated from P. zajonzi,
the neighboring schwenkii-complex species from offshore Socotra Archipelago of Yemen, by having the normal
two rows of teeth medially in the upper jaw (vs. three).
The barcode mtDNA COI sequence of P. xanthomma is almost the same as the sequences for P. tominagai
(only 0.16% divergent) and close to P. ternay from the Seychelles and P. rubricauda from Madagascar. The group
of four species diverge by 3.4% from the lineage comprising P. hadra from Maldives, P. bineeshi from India, and
probably true P. schwenkii from Bali, Indonesia (Fig. 1).
Pempheris zajonzi Randall & Victor, n. sp.
Figure 51, Tables 1–3.
Holotype. SMF 35713, male, 89 mm SL, ST-024/F4, Yemen, Socotra Archipelago, Socotra Island, Ras
Qatanin, 150 m off coast on shipwreck, 12°20’03” N 53°34”26” E, 2–4 m, rotenone, U. Zajonz & M. Khalaf,
March 13, 1999.
Paratypes. SMF 35714, juveniles, 4: 41–49 mm SL, same data as holotype.
Description. Dorsal-in rays VI,9; anal-in rays III,38 (36–38); pectoral-in rays 17; lateral-line scales 50
(49–54); scales on nape, below dorsal in, and side of caudal peduncle cycloid; scales on chest cycloid and about
one-fourth ctenoid ventroanteriorly; gill rakers 28 (28–30); body depth 2.4 in SL; body width 2.7 in body depth;
dorsal proile from above eye to origin of dorsal in nearly straight; head length 3.15 in SL; eye relatively small,
the orbit diameter 8.0 in SL; interorbital width 11.8 in SL; maxilla forming an angle of 64° to horizontal axis of
body; front of upper jaw with three irregular rows of well-developed recurved teeth for two-thirds the length of
Figure 51. Pempheris zajonzi, holotype, SMF 35713, male, 89 mm SL, Socotra Archipelago, Yemen (H.A. Randall).
64
the tooth-bearing part of jaw, the uppers more protruding, and the lowers more strongly recurved; teeth anteriorly
in lower jaw very small, in three to four irregular close-set rows, progressively more recurved and larger inwardly;
lower jaw not protruding when mouth irmly shut; tongue an equilateral triangle with pointed tip; lips smooth;
caudal in damaged; predorsal length 2.55 in SL; longest dorsal-in ray 4.45 in SL; pectoral-in length 3.25 in SL;
prepelvic length 2.55 in SL; pelvic ins extending a short distance beyond anus, their length 6.6 in SL. Color in
alcohol uniform brown except for paler scale edges (Fig. 51); leading edge of dorsal in black, expanding slightly
distally; caudal in yellow with a very broad black posterior margin and narrow black upper and lower margins;
black band at base of anal in very broad, about equal in width to the outer ray part of in; base of pectoral in only
slightly darker than adjacent body; iris very dark bluish gray. Color in life unknown.
Etymology. This species is named for the collector, Uwe Zajonz, of the Senckenberg Museum, Frankfurt,
who plans to collect more specimens and obtain photographs of the new species.
Remarks. Pempheris zajonzi was irst identiied by us as P. tominagai, but that species differs in having fewer
lateral-line scales, and predorsal and prepelvic lengths of about 2.45 in SL. The new species is also separated from
P. xanthomma, the neighboring schwenkii-complex species from the mainland coast of Yemen, by having three
rows of teeth medially in the upper jaw (vs. two in P. xanthomma). Uwe Zajonz plans to make a special effort to
collect more specimens and take photographs when travel to Socotra Archipelago becomes possible. No DNA
sequences are available for P. zajonzi.
Acknowledgments
We thank the following for their efforts to collect and/or photograph species of Pempheris for our research on
the genus: Moteah Shaikh Aideed, Gerald R. Allen, Andrew Bentley, Giacomo Bernardi, K.K. Bineesh, Sergey
V. Bogorodsky, Simon Chater, Allan D. Connell, Brian W. Darvell, Joseph D. DiBattista, Alain Diringer, Patrick
Eaton, Hiromitsu Endo, Mark V. Erdmann, M. Farooq, Bob Fenner, Richard F. Field, Ofer Gon, Gavin Gouws,
Phillip and Elaine Heemstra, Nicolas Hubert, Dennis King, Jonathan Mee, Robert F. Myers, Sean O’Hara, Yohan
Pathirana, Serge Planes, Dennis Polack, P.T. Rajan, Helen A. Randall, Luiz A. Rocha, Andrey Ryanskiy, William
T. White, and Keith D.P. Wilson. We are most grateful for gifts or loans of specimens of Pempheris from James
Maclaine of the Natural History Museum, London (BMNH); David Catania of the California Academy of Sciences,
San Francisco (CAS); Alastair Graham of the Australian National Fish Collection at Hobart, Tasmania (CSIRO);
Dani Golani of the Hebrew University of Jerusalem (HUJ); Barry C. Russell of the Northern Territory Museum,
Darwin (NTM); Ukkrit Satapoomin of the Phuket Marine Biological Centre, Phuket, Thailand (PMBC); Roger
Bills and Bufo Konqobe of the South African Institute for Aquatic Biodiversity, Grahamstown (SAIAB); Tilman
J. Alpermann and Uwe Zajonz of the Senckenberg Museum, Frankfurt (SMF); Menachem Goren of Tel-Aviv
Univeristy (TAU); Henrich Bruggemann of the Laboratoire d’Ecologie Marine, Université de la Réunion (URUN),
and Shirleen Smith and Jeffrey T. Williams of the U.S. National Museum of Natural History, Washington, D.C.
(USNM). We also thank Randall D. Mooi for data on type specimens of species of Pempheris, Loreen R. O’Hara
and Arnold Y. Suzumoto of the Bishop Museum, Honolulu (BPBM) for curatorial assistance, Wouter Holleman for
taking data on type specimens at SAIAB that could not be sent on loan, Peter Psomadakis and Lloyd Paul Fanning
of FAO and Hamid Badar Osmany of the Marine Fisheries Department in Karachi for providing specimens and
photographs of Pempheris from the Sind Coast of Pakistan, Michael Berumen and Joseph D. DiBattista for their
collections in the Red Sea, and King Abdullah University of Science and Technology for logistic and inancial
support to them. Finally, our special thanks to Dawn Goebbels for her numerous underwater photographs of
species of Pempheris in the Watamu Marine National Park in Kenya. The DNA barcoding was performed at the
Biodiversity Institute of Ontario with the support of Bob Hanner and the team at BOLD. DNA barcoding was
supported by the International Barcode of Life Project (iBOL.org) with funding from the Government of Canada
via the Canadian Centre for DNA Barcoding, as well as from the Ontario Genomics Institute (2008-OGI-ICI-03),
Genome Canada, the Ontario Ministry of Economic Development and Innovation, and the Natural Sciences and
Engineering Research Council of Canada. The manuscript was reviewed by William D. Anderson, Jr., Giacomo
Bernardi, K.K. Bineesh, Sergey V. Bogorodsky, Brian W. Darvell, Richard F. Field, and Helen A. Randall.
65
References
Azzurro, E., Goren, M., Diamant, A., Galil, B. & Bernardi, G. (2015) Establishing the identity and assessing the
dynamics of invasion in the Mediterranean Sea by the dusky sweeper, Pempheris rhomboidea Kossmann &
Räuber, 1877 (Pempheridae, Perciformes). Biological Invasions, 17, 815–826.
Bleeker, P. (1855) Bijdrage tot de kennis der ichthyologische fauna van de Batoe Eilanden. Natuurkundig
Tijdschrift voor Nederlandsch Indië, 8, 305–328.
Cuvier, G. (1829) Le Règne Animal, Ed. 2. Chez Deterville, Paris, France, xv + 406 pp.
Eschmeyer, W.N. & Fricke, R. (Eds.) (2015) Catalog of Fishes (http://researcharchive.calacademy.org/research/
ichthyology/catalog/ishcatmain.asp). Electronic version accessed Nov. 21, 2015.
Ivanova, N.V., Zemlak, T.S., Hanner, R.H. & Hebert, P.D.N. (2007) Universal primer cocktails for ish DNA
barcoding. Molecular Ecology Notes, 7, 544–548.
Leis, J.M. & Carson-Ewart, B.M. (2000) The Larvae of Indo-Paciic Coastal Fishes: An Identiication Guide to
Marine Fish Larvae. Brill, Leiden, Netherlands, 850 pp.
Koeda, K., Yoshino, T., Imai, H. & Tachihara, K. (2014) A review of the genus Pempheris (Perciformes,
Pempheridae) of the Red Sea, with description of a new species. Zootaxa, 3793, 301–330.
Mooi, R.D. & Randall, J.E (2014) Pempheris bexillon, a new species of sweeper (Teleostei: Pempheridae) from
the Western Indian Ocean. Zootaxa, 3780, 388–398.
Randall, J.E. (1995) Coastal Fishes of Oman. Crawford House Publishing, Bathurst, N.S.W. & University of
Hawai‘i Press, Honolulu, HI, USA, xiii + 439 pp.
Randall, J.E. & Bineesh, K.K. (2014) Review of the ishes of the genus Pempheris (Perciformes: Pempheridae)
of India with description of a new species and a neotype for P. mangula Cuvier. Journal of the Ocean Science
Foundation, 10, 20–40.
Randall, J.E., Bogorodsky, S.V., Alpermann, T.J., Satapoomin, U., Mooi, R.D. & Mal, A.O. (2013) Pempheris
lavicycla, a new pempherid ish from the Indian Ocean, previously identiied as P. vanicolensis Cuvier.
Journal of the Ocean Science Foundation, 9, 1–23.
Randall, J.E. & Victor, B.C. (2014) Four new ishes of the genus Pempheris (Perciformes: Pempheridae) from the
western Indian Ocean. Journal of the Ocean Science Foundation, 12, 61–83.
Randall, J.E., Victor, B.C., Alpermann, T.J., Bogorodsky, S.V., Mal, A.O., Satapoomin, U. & Bineesh, K.K.
(2014) Rebuttal to Koeda et al. (2014) on the Red Sea ishes of the perciform genus Pempheris. Zootaxa,
3887, 377–392.
Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: The Barcode of Life Data System (www.barcodinglife.org).
Molecular Ecology Notes, 7(3), 355–364.
Steindachner, F. (1870) Ichthyologische Notizen (X). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Classe, 61, 623–664.
Steinke, D., Zemlak, T.S. & Hebert, P.D.N. (2009) Barcoding Nemo: DNA-Based Identiications for the
Ornamental Fish Trade. PLoS ONE 4(7) e6300. doi:10.1371/journal.pone.0006300
Victor, B.C. (2015) How many coral reef ish species are there? Cryptic diversity and the new molecular
taxonomy. In: Mora, C. (Ed.) Ecology of Fishes on Coral Reefs. Cambridge University Press, Cambridge,
United Kingdom, pp. 76–87.
Ward, R.D., Hanner, R. & Hebert, P.D.N. (2009) The campaign to DNA barcode all ishes, FISH-BOL. Journal
of Fish Biology, 74, 329–356.
Zajonz, U., Maroof, K. & Krupp, F. (2000) Coastal ish assemblages of the Socotra Archipelago. In: Conservation
and Sustainable use of Biodiversity of Socotra Archipelago, Marine Habitat, Biodiversity and Fisheries
Surveys and Management. Progress Report of Phase III. Senckenberg Research Institute and Natural History
Museum, Frankfurt, Germany, pp. 127–170.
66
Appendix A. Specimen data and GenBank accession numbers for the barcode mtDNA COI sequences used to generate the
phenogram in Fig. 1, following the order in the tree. Holotypes in bold, paratypes with P in voucher ID.
Species
Collection site GenBank #
Voucher
BOLD BIN
Collector/Source
Pempheris eatoni
South Africa
JF494126
SAIAB 80038 ADC 216.2-3
AAD1777
A. Connell
Pempheris kruppi
Yemen
KT352048
mot 151115
AAD1777
M.S. Aideed
Pempheris kruppi
Yemen
KT352041
mot 151116
AAD1777
M.S. Aideed
Pempheris kruppi
Yemen
KT352053
mot 151117
AAD1777
M.S. Aideed
Pempheris eatoni
South Africa
JF494127
SAIAB 80038 ADC 216.2-2
AAD1777
A. Connell
Pempheris eatoni
South Africa
KT207966
P BPBM 41191 ADC216.2 #20
AAD1777
A. Connell
Pempheris
Madagascar
JQ350199
NBE0182 “mangula”
AAD1777
S. Planes/N. Hubert
Pempheris eatoni
South Africa
KT207961
SAIAB 189160 ADC2013 216.2-4
AAD1777
P. Eaton/A. Connell
Pempheris eatoni
South Africa
JF494128
SAIAB 80038 ADC 216.2-1
AAD1777
A. Connell
Pempheris
South Africa
KT207952
sa142164c2
AAD1777
A. Connell
Pempheris kruppi
Yemen
KT352044
mot151118
AAD1777
M.S. Aideed
Pempheris russellii
Pakistan
KT207954
BPBM pak14px
AAD1777
P. Psomadakis
Pempheris kuriamuria
Oman
KJ937296
BPBM 41239 JR14po
AAD1777
Darvell & Wilson
Pempheris kuriamuria
Oman
KT352029
CAS236521
AAD1777
L. Rocha/CAS
Pempheris kruppi
Yemen
KT352035
mot 151114
AAD1777
M.S. Aideed
Pempheris eatoni
South Africa
KT207953
P BPBM 41191 ac1421
AAD1777
A. Connell
Pempheris wilsoni
Fahal Is., Oman
KT352058
CAS236523
AAD1777
L. Rocha/CAS
Pempheris
South Africa
KT207965
sa142164c1
AAD1777
A. Connell
Pempheris rhomboidea
Red Sea
KJ609425
PVA_EIL_091017
AAD1777
G. Bernardi
Pempheris rhomboidea
Red Sea
KJ609479
PVA_HAI_091043
AAD1777
G. Bernardi
Pempheris rhomboidea
Red Sea
KJ609414
PVA_EIL_091001
AAD1777
G. Bernardi
Pempheris rhomboidea
Red Sea
KJ609433
PVA_ALE_061005
AAD1777
G. Bernardi
Pempheris rhomboidea
Red Sea
KJ020196
NBFGR PE1
AAD1777
K.K. Bineesh
Pempheris rhomboidea
Red Sea
KJ020197
NBFGR PE2
AAD1777
K.K. Bineesh
Pempheris rhomboidea
Red Sea
KJ020198
NBFGR PM1
AAD1777
K.K. Bineesh
Pempheris rhomboidea
Red Sea
KJ020199
NBFGR PM2
AAD1777
K.K. Bineesh
Pempheris mangula
Bali, Indonesia
JN313327
BW-A11176 “adusta”
AAD1777.1
W. White/R. Ward
Pempheris mangula
Bali, Indonesia
JN313328
BW-A11177 “adusta”
AAD1777.1
W. White/R. Ward
Pempheris mangula
India
EU148571
WL-M263 “adusta”
AAD1777.1
W.S. Lakra et al.
Pempheris mangula
India
EU148572
WL-M264 “adusta”
AAD1777.1
W.S. Lakra et al.
Pempheris mangula
India
EU148574
WL-M266 “adusta”
AAD1777.1
W.S. Lakra et al.
Pempheris mangula
India
EU148573
WL-M265 “adusta”
AAD1777.1
W.S. Lakra et al.
Pempheris mangula
India
EU148575
WL-M267 “adusta”
AAD1777.1
W.S. Lakra et al.
Pempheris bineeshi
India
KJ020182
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020183
NBFGR-CH-1143
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020184
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020185
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020186
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020187
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris bineeshi
India
KJ020188
ZSI/CLT 2492
AAM9396
K.K. Bineesh
Pempheris schwenkii
Bali, Indonesia
pending
BW-A10192 CSIRO KD677
AAM9396
W. White/R. Ward
Pempheris schwenkii
Bali, Indonesia
JN312883
BW-A10194 CSIRO KD678(S)
AAM9396
W. White/R. Ward
Pempheris hadra
Maldives
pending
SAIAB 187537 MAL-022
AAM9396
O. Gon/G. Gouws
Pempheris tominagai
Red Sea
pending
PVA EIL 091005 0.82
AAM9396
G. Bernardi
Pempheris tominagai
Red Sea
pending
PVA EIL 091012 0.82
AAM9396
G. Bernardi
Pempheris tominagai
Red Sea
pending
PVA EIL 091020 0.66
AAM9396
G. Bernardi
67
68
Pempheris tominagai
Red Sea
pending
PVA EIL 091023 0.82
AAM9396
G. Bernardi
Pempheris
Chagos Arch.
KT352037
Chagos-702
AAI3650
M. Gaither/CAS
Pempheris
Chagos Arch.
KT352037
Chagos-703
AAI3650
M. Gaither/CAS
Pempheris xanthomma
Yemen
KT352036
mot151113
AAI3650
M.S. Aideed
Pempheris ternay
Seychelles
KF930243
P SAIAB 77935 KUT 6867
AAI3650
P. Heemstra et al.
Pempheris rubricauda
Madagascar
JQ350203
NBE0213 “vanicolensis”
AAI3650
S. Planes/N. Hubert
Pempheris rubricauda
Madagascar
JQ350202
NBE0214 “vanicolensis”
AAI3650
S. Planes/N. Hubert
Pempheris rubricauda
Madagascar
JQ350204
NBE0215 “vanicolensis”
AAI3650
S. Planes/N. Hubert
Pempheris bexillon
Réunion
JQ350200
REU0958 “oualensis”
AAU3528
S. Planes/N. Hubert
Pempheris bexillon
Réunion
JQ350201
REU0957 “oualensis”
AAU3528
S. Planes/N. Hubert
Pempheris ibo
South Africa
KT207950
ac14p9
AAF8820
A. Connell/SAIAB
Pempheris ibo
South Africa
HQ561454
P SAIAB 189165 ADC10_216.4 #6 AAF8820
A. Connell/SAIAB
Pempheris ibo
South Africa
KT207959
ac14p7
AAF8820
A. Connell/SAIAB
Pempheris hollemani
SE Madagascar
pending
P SAIAB 97406 AV2010-027
AAC6084
P. & E. Heemstra
Pempheris connelli
South Africa
KT207957
P SAIAB 194783 sa142164b2
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KF930242
RUSI 65242 UKFBJ655-08
AAC6084
A. Bentley/KU Tissue
Pempheris connelli
South Africa
KT207967
P SAIAB 194782 ac14p5
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207962
P SAIAB 194782 ac14p6
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207951
P SAIAB 194782 ac14p8
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207956
P SAIAB 194782 ac14p10
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207964
P SAIAB 194782 ac14p11
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207958
P SAIAB 194782 ac14p12
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
JF494131
P SAIAB 75850/1 ADC216.4-2
AAC6084
P. Heemstra/SAIAB
Pempheris connelli
South Africa
JF494130
P SAIAB 75850 ADC216.4-4
AAC6084
P. Heemstra/SAIAB
Pempheris connelli
South Africa
JF494132
P SAIAB 75776 ADC216.4-1
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KF489699
ADC12_216.4A #3
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207960
P SAIAB 194783 sa142164b1
AAC6084
A. Connell/SAIAB
Pempheris connelli
South Africa
KT207949
P SAIAB 194783 sa142164b3
AAC6084
A. Connell/SAIAB
Pempheris rochai
Oman
KT352045
CAS236522
AAC6084
L. Rocha/CAS
Pempheris sp. 1
South Africa
KT207963
L II A2 #1.1 egg DSLAF019-07
pending
A. Connell
Pempheris bruggemanni
Réunion
JQ350194
P REU0731 URUN 2007-1208
AAT9853
S. Planes/N. Hubert
Pempheris bruggemanni
Réunion
JQ350195
P REU0936 BPBM 41198
AAT9853
S. Planes/N. Hubert
Pempheris bruggemanni
Réunion
JQ350198
P REU0938 URUN 2007-1209
AAT9853
S. Planes/N. Hubert
Pempheris bruggemanni
Réunion
JQ350196
P REU0732 URON 2007-1419
AAT9853
S. Planes/N. Hubert
Pempheris bruggemanni Réunion
JQ350197
REU0937 MNHN 2015-0002
AAT9853
S. Planes/N. Hubert
Pempheris malabarica
Karachi, Pakistan
KT207955
pak14pm
ACR6962
P. Fanning/FAOPK
Pempheris lavicycla
Al Lith, Red Sea
KT352052
CAS PVA02 995
AAE7117
L. Rocha/CAS
Pempheris lavicycla
Djibouti
KT352054
CAS 235013
AAE7117
L. Rocha/CAS
Pempheris lavicycla
Red Sea
KF914197
KAU13-608
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914196
KAU11-506
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914195
KAU11-505
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914193
KAU11-016
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914194
KAU11-504
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914192
KAU11-015
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914190
KAU11-013
AAE7117
S. Bogorodsky
Pempheris lavicycla
Red Sea
KF914191
KAU11-014
AAE7117
S. Bogorodsky
Pempheris lavicycla
Seychelles
EU381029
SAIAB 77856
AAE7117
A. Bentley et al.
Pempheris lavicycla
Seychelles
EU381030
SAIAB 77805
AAE7117
A. Bentley et al.
Parapriacanthus
Tonga
KF930232
USNM 336802
AAC7602
J.T. Williams et al.
Appendix
Pempheris malabarica x Pempheris russellii
Appendix Figure 1; Appendix Table 1
A project of the Food and Agriculture Organization of the United Nations to document the ishery resources
of the Sind Coast of Pakistan provided us with the opportunity to request specimens of Pempheris from the
region. Dr. Peter Psomadakis of FAO and Hamid Badar Osmany of the Marine Fisheries Department in Karachi
sent specimens, tissue samples, and photographs of P. malabarica C. & V. and P. russellii Day from the sea off
Karachi. While making routine counts of in rays and scales of the specimens, one that looked like P. malabarica
(BPBM 41183, male, 125 mm SL) was noted to have the unusual low number of 59 lateral-line scales (60 on
the other side)(Appendix Fig. 1). Pempheris malabarica has the highest lateral-line scale count (65–75) for the
genus. Other meristic data of the specimen are within the range for P. malabarica. The eye size is precisely the
same as that of P. malabarica for the standard length, and the upper-jaw angle of 65° is the same; also it has a dark
anterior margin on the dorsal in like P. malabarica, instead of a large distal black spot found on most species of
Pempheris, including P. russellii. However, it has a deeper body like that of P. russellii and is intermediate to the
two species in its pectoral-in length (Appendix Table 1). Appendix Fig. 2 of a specimen of P. malabarica and
Appendix Fig. 3 of a specimen of P. russellii are provided for comparison.
Suspecting the specimen to be a hybrid, the dentition was carefully compared. P. malabarica has the most
distinctive dentition of the genus: the teeth in the usual two rows anteriorly in the upper jaw are nearly three
times larger than those of P. russellii and project horizontally forward instead of obliquely downward. The teeth
anteriorly in the lower jaw are in a pair of patches shaped like a comma, separated by a narrow medial gap. The
patches are half exposed when the jaws are fully closed. The teeth are progressively larger anteriorly (the reverse
is typical of the genus), the ive or six teeth at the front are stoutly conical, twice as large as remaining teeth,
Appendix Figure 1. Pempheris malabarica x Pempheris russellii, BPBM 41183, male, 125 mm SL, Karachi, Pakistan
(H.A. Randall).
69
Appendix Figure 2. Pempheris malabarica, BPBM 36185, 157 mm SL, Masirah Island, S. Oman (J.E. Randall).
Appendix Figure 3. Pempheris russellii, 137 mm SL, Sind coast of Pakistan (H. Badar Osmany).
70
APPENDIX TABLE 1
Comparison of characters of Pempheris russellii,
P. malabarica, and the hybrid P. malabarica x P. russellii*
P. russellii
hybrid
P. malabarica
Standard length (mm)
124
125
122
Sex
Lateral-line scales
Body depth
Eye diameter
Angle of maxilla
Pectoral-in length
Dorsal-in marking
male
55
2.2
7.4
70°
3.05
spot
male
59
2.2
8.35
65°
3.2
band
male
65
2.4
8.35
65°
3.3
band
*Measurements of depths and lengths are times in standard length; band refers to the
dark anterior border on the dorsal in, spot to an isolated large apical black marking on
the dorsal in.
and project directly forward. The remaining teeth in the patches are well separated, progressively smaller, more
slender, and recurved. There are two rows of small, slender, recurved teeth to the side of the jaw, narrowing to a
single row posteriorly. The dentition of P. russellii is typical of the genus Pempheris, in general; that of the hybrid
is intermediate.
The underwater photograph of Appendix Fig. 4 shows a mixed aggregation of P. malabarica (orangish and
no dorsal-in spot) and P. russellii (grayish brown with black-tipped dorsal in) on a reef off Karachi, Pakistan. In
view of the proximity of the two species, it is not dificult to predict that the ova of one might at times be fertilized
by the sperm of the other.
Appendix Figure 4. Mixed aggregation of Pempheris malabarica and P. russellii, Karachi Reef, Pakistan (M. Farooq).
71
Appendix Figure 5. Pempheris sp. USNM 343770, 98.5 mm SL, St. Brandon’s Shoals (H.A. Randall).
St. Brandon’s Shoals Fish Collections. Large ish collections were made, mainly with rotenone, in the St.
Brandon’s Shoals (Cargados Carajos) of the Indian Ocean, from a sailing vessel in April 1976. The expedition
was sponsored by Lewis H. Strauss and included the ichthyologists Victor G. Springer, Thomas H. Fraser, and
C. Lavett Smith. Among the many species of ishes collected are what seemed to be a single small species of
Pempheris of the schwenkii complex from three stations: USNM 343769, 35: 70–107 mm, SE side of Grande
Passe (16°28’ S, 59°40’ E) at 0–20 feet on April 5, 1976; USNM 343770, 44: 44–114.5 mm SL, 0.5 miles S of
Île Raphael (16°27’ S, 59°36’ E) at a depth of 0–30 feet, April 12, 1976; and USNM 343768, 8: 101.5–110.5 mm
SL, off NW shore of Albatross Island (16°15’ S, 59°15’ E), at 0–18 m, April 14, 1976. Appendix Fig.5 illustrates
a 98.5-mm specimen from the second collection.
Routine measurement and meristic data were taken of the three St. Brandon’s collections in the expectation
that all would be essentially the same. They are for body and in proportions and within one count for anal-in
rays (38–43) and lateral-line scales (49–60). However, a surprise came with gill-raker counts. The ish of the irst
collection have 27–29 gill rakers, while the next two collections have 28–30. We then searched for other characters
that might show differences. The angle of the maxilla averaged about 65° for all the specimens. Whether scales on
the side of the chest are cycloid or ctenoid is a valuable character for species of Pempheris. Unfortunately there
are usually at least some scales missing. When the chest scales are fully present or nearly so, the ish of the irst
collection have all or nearly all cycloid scales, while those of the second and third collections are mainly or all
ctenoid. Unexpectedly, we found ish of all three St. Brandon’s collections that clearly had too many of the wrong
kind of chest scales, suggesting hybridization.
To determine the status of the species of Pempheris of these three collections is beyond our present capability, in
view of the time frame to complete the manuscript. A future expedition to St. Brandon’s Shoals, with observations
on habitat, underwater photographs, and taking tissue samples for genetic analysis, should be able to resolve this
interesting complex of species.
Unidentiied species of Pempheris. Many species of Pempheris are dificult to positively identify from
underwater photographs. However, there are a few photographs that are suficiently distinct to suggest undescribed
species. Appendix Figs. 6 and 7, the former from Madagascar, the latter from Sri Lanka, are examples of new
species for which specimens are needed. We also believe Appendix Figs. 8 through 15, taken by Dawn Goebbels
in the Watamu Marine National Park in Kenya, represent undescribed species of Pempheris.
72
Appendix Figure 6. Pempheris sp., 60 km north of Tuléar, Madagascar (G.R. Allen). Koeda et al. (2014: ig. 7a) mistakenly
identiied this photograph as P. tominagai.
Appendix Figure 7. Pempheris sp., with dark edge on anal in and truncate caudal in, resembles P. lavicycla, but is missing
the characteristic dark oval at the pectoral-in base, Sri Lanka (J.E. Randall).
73
Appendix Figure 8. Pempheris sp. 1. In addition to color pattern, distinct in the deep body and slightly forked caudal in,
Watamu Marine National Park, Kenya (D. Goebbels).
Appendix Figure 9. Pempheris sp. 1: apparent subadult, Watamu Marine National Park, Kenya (D. Goebbels).
74
Appendix Figure 10. Pempheris sp. 2. Note the very long anterior part of the anal in, the black membranes of the pelvic
ins, and the copper and green striped color pattern, Watamu Marine National Park, Kenya (D. Goebbels).
Appendix Figure 11. Pempheris spp., all of the ish in this aggregation are the same species except for the striped reddish
brown ish at the lower right edge. The color difference is mainly from the angle of the light relected from the scales; all of
the ish share a reddish caudal in with dark brown dorsal and ventral margins, Watamu Marine National Park, Kenya (D.
Goebbels).
75
Appendix Figure 12. Pempheris spp., two species of Pempheris with the cleaner wrasse Labroides dimidiatus, one with
the black-tipped dorsal in probably P. nesogallica, the other probably undescribed, Watamu Marine National Park, Kenya
(D. Goebbels).
Appendix Figure 13. Pempheris of the schwenkii complex, distinct in its yellow-green color pattern. Cropped from a dense
aggregation, mainly of this species, Watamu Marine National Park, Kenya (D. Goebbels).
76
Appendix Figure 14. Pempheris sp., a reddish species, its small size apparent by the size of the blades of Thalassia and the
associated apogonid ishes, Watamu Marine National Park, Kenya (D. Goebbels).
Appendix Figure 15. Pempheris sp., the small eye and unique color pattern of the anal in indicate another probable new
species, Watamu Marine National Park, Kenya (D. Goebbels).
77