Six New Species of the Cichlid Genus Otopharynx from Lake Malaŵi (Teleostei: Cichlidae)

Abstract

The endemic Lake Malaŵi cichlid genus Otopharynx is increased from 14 species to 20 by the description of six new species collected from inshore sandy or intermediate rocky/sandy areas or trawled in depths to 59 m. Otopharynx alpha is distinguished from congeners by its unique melanin pattern with suprapectoral spot discrete but supraanal and precaudal spots connected to form a stripe, and by its distinctively shaped pharyngeal bone with numerous small, crowded teeth. Otopharynx mumboensis, thought to be endemic to Mumbo Island, is separated from most congeners by a suprapectoral spot placed entirely below the upper lateral line, and from all others by the combination of 13 gill rakers, pharyngeal bone with posterior margin nearly straight, crowns of all pharyngeal teeth in the median columns broadened and flattened, head length 30.9–32.2% SL, body depth 35.5–36.1% SL, and lower jaw 35.7–36.2% HL. Otopharynx styrax has a more elongate body (depth 24.3–28.6% SL) than any congener. Otopharynx aletes can be diagnosed by the combination of a suprapectoral spot overlapping the upper lateral line, seven or eight vertical bars below the dorsal-fin base, 10 or 11 gill rakers, 34–36 scales in the lateral line, lower pharyngeal bone with molariform posteromedian teeth, and caudal peduncle length 1.6–2.0 times its depth. Otopharynx panniculus is diagnosed by the combination of a prominent square suprapectoral spot spanning subdorsal bars 3 to 4, no dorsal midline spots, 13–15 gill rakers, 31–33 scales in the lateral line, 15 or 16 dorsal-fin spines, absence of a lacrimal notch, and slightly enlarged teeth in the median columns of the lower pharyngeal bone. Otopharynx peridodekamost closely resembles O. panniculus and shares the loss of the lacrimal notch, but has 34–35 scales in the lateral line and 11–13 gill rakers. Photographs of living or freshly collected specimens of five of the new species are provided.

Full text

Six New Species of the Cichlid Genus Otopharynx
from Lake Malaŵi (Teleostei: Cichlidae)
Michael K. Oliver
Division of Vertebrate Zoology, Peabody Museum of Natural History, Yale University,
New Haven CT 06520-8118 USA
—email: anagenys@gmail.com
ABSTRACT
The endemic Lake Malaŵi cichlid genus Otopharynx is increased from 14 species to 20 by the
description of six new species collected from inshore sandy or intermediate rocky/sandy areas or
trawled in depths to 59 m. Otopharynx alpha is distinguished from congeners by its unique
melanin pattern with suprapectoral spot discrete but supraanal and precaudal spots connected to
form a stripe, and by its distinctively shaped pharyngeal bone with numerous small, crowded
teeth. Otopharynx mumboensis, thought to be endemic to Mumbo Island, is separated from most
congeners by a suprapectoral spot placed entirely below the upper lateral line, and from all others
by the combination of 13 gill rakers, pharyngeal bone with posterior margin nearly straight,
crowns of all pharyngeal teeth in the median columns broadened and flattened, head length
30.9–32.2% SL, body depth 35.5–36.1% SL, and lower jaw 35.7–36.2% HL. Otopharynx styrax has
a more elongate body (depth 24.3–28.6% SL) than any congener. Otopharynx aletes can be diag-
nosed by the combination of a suprapectoral spot overlapping the upper lateral line, seven or eight
vertical bars below the dorsal-fin base, 10 or 11 gill rakers, 34–36 scales in the lateral line, lower
pharyngeal bone with molariform posteromedian teeth, and caudal peduncle length 1.6–2.0 times
its depth. Otopharynx panniculus is diagnosed by the combination of a prominent square
suprapectoral spot spanning subdorsal bars 3 to 4, no dorsal midline spots, 13–15 gill rakers,
31–33 scales in the lateral line, 15 or 16 dorsal-fin spines, absence of a lacrimal notch, and slightly
enlarged teeth in the median columns of the lower pharyngeal bone. Otopharynx peridodeka most
closely resembles O. panniculus and shares the loss of the lacrimal notch, but has 34–35 scales in
the lateral line and 11–13 gill rakers. Photographs of living or freshly collected specimens of five
of the new species are provided.
Zoobank.org/urn:lsid:zoobank.org:pub:66CFE44F-095D-4E30-A0D5-0A65032EFC61
KEYWORDS
Pseudocrenilabrinae, Haplochromini, endemic species, Lake Malaŵi Trawling Survey
Bulletin of the Peabody Museum of Natural History 59(2):159–197 , October 2018.
© 2018 Peabody Museum of Natural History, Yale University. All rights reserved. • http://peabody.yale.edu
Introduction
The hyperdiverse family Cichlidae likely com-
prises about 3,000 species, counting those already
described and the many that are known but await
description and extrapolating from the rate of new
discoveries. Over one-quarter of all cichlid species
occur only in Lake Malaŵi, which probably con-
tains between 800 (Snoeks 2001) and 1,000
species (Konings 2016). Currently, 406 species
of cichlid, assigned to 58 genera, have been
described from Lake Malaŵi (Oliver 2018).
The genus Otopharynx was established in a
footnote. Regan’s (1920) description of this Lake
Malaŵi cichlid genus reads, in full: “Differs from
Tilapia in that the prootic forms part of the pha-
ryngeal facet on each side; very near Chilotilapia.”
The type and, initially, only species was Tilapia
auromarginata Boulenger, 1908, but later Regan
added a second species, Otopharnyx selenurus
Regan, 1922. Trewavas (1935:69), however, was
unable to confirm Regan’s observations on the
pharyngeal facet in O. auromarginatus, noting
that “As in Haplochromis, the edge of the prootic
doi: 10.3374/014.059.0204

may at one point rise nearly or quite to the level of
the facet, but it never expands to form an impor-
tant part of it. The genus Otopharynx must, there-
fore, in the present state of our knowledge, be
merged in Haplochromis.” Subsequently, Green-
wood (1979) restricted Haplochromis to five
species in Lakes Victoria, Edward, George, and
Kivu, leaving more than 100 Lake Malaŵi species
then classified in Haplochromis without a generic
name. He proposed that Cyrtocara Boulenger,
1902, whose type species is C. moorii Boulenger,
1902, of Lake Malaŵi, be used as a provisional
generic name “until the Malaŵi species are
revised” (Greenwood 1979:317).
Eccles and Trewavas (1989) did undertake a
generic revision of the Lake Malaŵi hap-
lochromines, excluding those of the mbuna group,
describing 23 new genera and resurrecting
Otopharynx for 11 species with spots on the
flanks. In addition to O. auromarginatus and O.
selenurus, these were: O. heterodon (Trewavas,
1935), O. ovatus (Trewavas, 1935), O. tetrastigma
(Günther, 1894), O. speciosus (Trewavas, 1935), O.
decorus (Trewavas, 1935), O. argyrosoma (Regan,
1922), O. tetraspilus (Trewavas, 1935), and two
species newly described in Eccles and Trewavas
(1989): O. brooksi Oliver, 1989, and O. lithobates
Oliver, 1989. Beyond Lake Malaŵi proper, some
of these species can also be found in its outflow,
the upper Shire River. A few occur further down-
stream in Lake Malombe. One, O. tetrastigma, has
even been documented in the middle Shire at
Walkers Ferry, Malaŵi (Tweddle et al. 1979)—
some 125 km south of Lake Malaŵi, and consid-
erably farther in river distance.
In addition to the species they assigned to
Otopharynx, Eccles and Trewavas (1989) allocated
some other laterally spotted cichlids with distinc-
tive specializations to eight further genera. Their
diagnosis of Otopharynx contrasts it with five
of these other genera, but omits mention of
their newly established genera Naevochromis,
Copadichromis, and Tramitichromis, each of which
also includes one or more cichlids with suprapec-
toral and supraanal spots. Their diagnosis of
Otopharynx reads, in full: “Medium-sized hap-
lochromines endemic to Lake Malaŵi attaining
little more than 200 mm SL, characterised by the
possession of supra-pectoral and supra-anal spots
lying on or below the upper lateral line. Differ
from Hemitilapia and Trematocranus in that the
spots never extend to the dorsal surface, from
Ctenopharynx in the dentition and the usually
lower number of gill-rakers on the lower outer
arch and from Stigmatochromis and Exo-
chochromis in the jaws and dentition” (Eccles and
Trewavas 1989:154). Thus, Otopharynx is simply
an assemblage of less-apomorphic cichlids with
dark lateral spots, in effect “defined” by charac-
ters its members lack, and is almost certainly
paraphyletic.
A twelfth species of Otopharynx, O. pachy-
cheilus, was described by Arnegard and Snoeks
(2001).
Two further Otopharynx species (O. antron
and O. spelaeotes) were added by Cleaver et al.
(2009), bringing the total to 14. Their concept of
the genus was unchanged from that of Eccles and
Trewavas (1989). Cleaver et al. (2009) did provide
a new diagnosis of Otopharynx, but it consists pri-
marily of the identical characteristics they repeat
in their diagnosis of Stigmatochromis.
No comprehensive phylogenetic analysis of
Lake Malaŵi haplochromines exists. Thus, few
monophyletic genera can be delimited that com-
prise more than one species. Nevertheless, pend-
ing availability of data to define monophyletic
groups within the flock, Otopharynx sensu Eccles
and Trewavas (1989) remains pragmatically use-
ful as a formal receptacle for the 14 species
previously described and for many of the still-
undescribed, laterally spotted Lake Malaŵi cich-
lids already known or awaiting discovery. It is the
purpose of this article to describe six of these new
species from both inshore and offshore localities
in the southern third of the lake (Figure 1).
Materials and Methods
I have examined the type specimens of all species
assigned to the genus Otopharynx, with the excep-
tion of O. pachycheilus Arnegard and Snoeks,
2001, O. antron Cleaver et al., 2009, and O.
spelaeotes Cleaver et al., 2009, which are recogniz-
able from their well-illustrated original descrip-
tions. Further specimens examined are listed in
the Appendix.
I made 36 measurements (including total
length) of each specimen. All measurements—
except caudal peduncle length, a projected meas-
urement—were taken point to point, on the left
side when possible, with digital calipers (iGaging
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
160

Six New Species of the Cichlid Genus Otopharynx • Oliver 161
FIGURE 1. Southern Lake Malaŵi, showing collection localities of the type series and other specimens of newly
described Otopharynx species. Map symbols: !, O. aletes (holotype YPM ICH 031030; paratypes YPM ICH
007820). #, O. peridodeka (holotype YPM ICH 031026; paratypes YPM ICH 014120). $, O. mumboensis (holo-
type YPM ICH 031031; paratypes YPM ICH 024990). %, O. panniculus (holotype YPM ICH 031027; paratypes
YPM ICH 014162; +, nontypes AMNH I-221760 SW). ", O. alpha (holotype YPM ICH 031028; paratypes
YPM ICH 014359; (, paratype YPM ICH 028511). 9, O. styrax (holotype YPM ICH 031029; paratypes YPM
ICH 014270).

OriginCal IP54, San Clemente, CA) interfaced
directly to a spreadsheet (Microsoft Excel 2010,
Redmond, WA). Except for total length, measure-
ments were made with a Zeiss (Jena, Germany)
Stemi 2000 zoom stereo microscope (magnifica-
tion 6.5–50⫻). Angles were estimated to the near-
est degree with a digital protractor (iGaging
#35–407). Counts and measurements follow
Snoeks (2004). Some further measurements,
angles, and anatomical descriptors not available
in that source follow Barel et al. (1977) including
gape inclination, snout acuteness, and lower-jaw
length–width ratio (narrow, intermediate, or
broad). However, vertebral counts include the
compound terminal half-centrum in contrast to
Barel et al., who excluded it. Additional measure-
ments: Upper-jaw length was measured from the
most rostral point of the premaxilla to the most
ventral point of the maxilla. Lower-jaw underside
angle, premaxillary pedicel angle, interorbital
angle, and nuchal angle were measured relative to
the lateral midline (an imaginary straight line
from the premaxillary symphysis to the lower lat-
eral line at the hypural fold). The count of lateral-
line kink scales follows Oliver (2016).
In order to record measurements between
repeatable homologous landmarks (Bookstein et
al. 1985), facilitating future comparative morpho-
metric studies, I have added several further meas-
urements. Most are new or rarely used in cichlid
taxonomy, including pelvic-fin origin to anal-fin
origin, dorsal-fin origin to anal-fin origin, dorsal-
fin origin to end of anal-fin base, pelvic-fin origin
to end of dorsal-fin base, dorsal-fin origin to
pelvic-fin origin, end of dorsal-fin base to end of
anal-fin base, anal-fin origin to end of dorsal-fin
base, end of dorsal-fin base to end of hypurals at
lower lateral line, and end of anal-fin base to end
of hypurals at lower lateral line. Only experience
will decide if these novel measurements enable
enough added resolution between similar species,
and reduction of measurement error, to justify the
time and effort expended in recording them.
Most of the specimens described herein were
trawled, dumped into boxes, and sorted aboard
the ship. Therefore, many are missing quite a few
scales. I found it helpful to paint a solution of Cya-
nine Blue (Acid Blue 113, Sigma–Aldrich, St.
Louis, MO) dissolved in distilled water onto scale-
less parts of the lateral line, following the
reversible technique described by Saruwatari et al.
(1997). Often, this outlined the scale pockets and
made an accurate count possible. The dye also
facilitated examination of very small scales such
as those between pectoral- and pelvic-fin bases.
Terms for the three principal lateral spots—
“suprapectoral,” “supraanal,” and “precaudal”—
were introduced by Oliver (1977, 1984),
popularized by Eccles and Trewavas (1989), and
adopted by subsequent workers (e.g., Arnegard
and Snoeks 2001, Cleaver et al. 2009, Dierickx et
al. 2018). I also continue to refer to the series of
4–6 small, distinct dark markings spaced along
the back just below the dorsal fin of some
Otopharynx and other species as “dorsal midline
spots.”
I examined the shape of the lacrimal bone in
all six new species (Figure 2). Terminology of
lacrimal bone morphology follows Anker (1986)
with the addition of “lacrimal notch” referring to
a distinct U- or V-shaped indentation of the
orbital margin immediately ventral to the lacrimal
process.
The numbers of outer teeth in the upper and
lower jaws are bilateral totals. Teeth represented
by empty alveoli were included. If absence of
numerous teeth on one hemijaw prevented an
accurate count, the count of the other side was
doubled. In several of the species here described,
the posterior teeth in both jaws are small and
nearly or completely hidden in the mucosa, com-
plicating attempts to count them accurately. Thus,
the tooth counts herein should be considered best
estimates.
I photographed the overall habitus and heads
of the new species on a copy stand using a Nikon
(Tokyo, Japan) D90 camera with AF MicroNikkor
60mm f/2.8D macro lens. I processed the image
layers with Helicon Focus software, v. 6.3.7. The
resulting merged images were cleaned and opti-
mized for brightness, contrast, clarity, and sharp-
ness in Corel PaintShop Pro X6 software, v.
16.2.0.20 x64. Pharyngeal bones were pho-
tographed with a QImaging 01-MP5.0-RTV-
CLR-10-C CCD camera mounted on a Zeiss
Stemi 2000–CS zoom stereo microscope and con-
trolled by Q-Capture Pro7 software. Merged
image layers were processed and optimized as for
habitus photos, except that image stacks were
processed using CombineZP software (Hadley
2012) or PICOLAY v. 2017-12-06 (Cypionka
2017).
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
162

The type specimens are deposited in the Divi-
sion of Vertebrate Zoology Ichthyology collec-
tions, Peabody Museum of Natural History, Yale
University, New Haven, CT (YPM ICH). Other
institutional abbreviations: AMNH, American
Museum of Natural History, New York; BMNH,
The Natural History Museum, London; MRAC,
Royal Museum for Central Africa, Tervuren, Bel-
gium; USNM, National Museum of Natural His-
tory, Washington, D.C. Additional abbreviations
in the text: HL, head length; SL, standard length;
TL, total length.
Results
Otopharynx Regan, 1920:38
Type. Tilapia auromarginata Boulenger, 1908:241, by original
designation.
Diagnosis.
No synapomorphies have been discovered to support a hypoth-
esis that Otopharynx is a monophyletic group. Otopharynx is
distinguishable from 32 of the other 40 currently recognized
genera of non-mbuna haplochromines occurring in Lake
Malaŵi by the melanic markings, which include at least a
suprapectoral spot and usually also a supraanal spot and a pre-
caudal spot (vs. these lateral spots lacking). Otopharynx can be
separated from the remaining eight genera, with which it shares
these lateral spots, as follows: Distinguished from Ctenopharynx
by having 9–18 (vs. 16–41) gill rakers on lower limb of outer
arch, and lower pharyngeal bone varying from subtriangular to
Y-shaped but always at least moderately robust (vs. Y-shaped,
the halves slender, fragile); from Copadichromis by having pre-
maxillary pedicels relatively short, mouth not very protrusible,
species with various diets but never shoaling zooplankton feed-
ers (vs. premaxillary pedicels elongated, mouth protrusible to
form a sucking tube, shoaling species specialized for zooplank-
ton feeding in open water); from Exochochromis by a nonpreda-
tory, usually deep-bodied facies (vs. elongate, subfusiform
piscivorous predatory facies), jaws equal anteriorly or lower pro-
jecting slightly (vs. lower jaw shorter than upper), and premax-
illa not beaklike, with relatively short ascending process (pedicel)
that is not conspicuous in head profile (vs. premaxilla beaklike,
with pedicel nearly as long as its dentigerous ramus, prominent
in head profile); from Hemitilapia by outer oral teeth generally
bicuspid or unicuspid with occasional tricuspid teeth intermixed
in some species (vs. specialized teeth with elongated shafts that
are movably implanted and inclined toward symphysis, crowns
broad, convexly rounded and obliquely truncate), and suprapec-
toral and supraanal spots variously shaped and placed, but not
reaching dorsal-fin base (vs. both spots in adults extending to
Six New Species of the Cichlid Genus Otopharynx • Oliver 163
FIGURE 2. Lacrimal bones of the new Otopharynx species, showing their outline, lateral-line canals and pores.
Drawn from lacrimal of right side and reversed. A. Otopharynx alpha, YPM ICH 014359 (paratype), 117.9 mm
SL. B. O. mumboensis, YPM ICH 024990 (paratype), 107.3 mm SL. C. O. styrax, YPM ICH 014270 (paratype),
95.2 mm SL. D. O. aletes, YPM ICH 007820 (paratype), 94.5 mm SL. E. O. panniculus, YPM ICH 014162
(paratype), 69.7 mm SL. F. O. peridodeka, YPM ICH 014120 (paratype), 91.0 mm SL. Scale bars equal 2 mm.

dorsal-fin base, like two saddles); from Stigmatochromis by a
nonpredatory and usually deep-bodied facies (vs. more elon-
gate predatory facies) and by having snout shorter than postor-
bital head (vs. snout equal to or longer than postorbital head);
from Trematocranus by suprapectoral and lateral spots not
reaching dorsal-fin base (vs. both spots in adults extending to
dorsal-fin base, like saddles), and cephalic lateral-line pores and
canals not appearing inflated (vs. enlarged, inflated); from
Tramitichromis intermedius (the only spotted species assigned
to that genus) by suprapectoral and supraanal spots not reach-
ing dorsal-fin base (vs. both spots in adults extending to dorsal-
fin base, forming saddles), and blade of lower pharyngeal bone
shallow to rather deep and varying from nearly straight to
angled slightly downward (vs. blade keellike, more than twice
depth of remainder of bone, angled steeply downward ~32°rel-
ative to plane of dentigerous surface, see Trewavas 1935:109, fig.
11); from Naevochromis by outer jaw teeth with shafts and
crowns exposed, extending well beyond surface of oral mucosa
(vs. oral teeth buried to crowns in thickened mucosa), premax-
illa toothed for most of its length (vs. teeth restricted to anterior
part), and dentary at symphysis not conspicuously deeper than
premaxilla (vs. dentary about twice depth of premaxilla).
Otopharynx alpha
new species
Zoobank.org/urn:lsid:zoobank.org:act:
E162D42A-E71B-4D5A-A19A-A9CDA20D9E2D
Figures 2A, 3, 4, Tables 1, 2
Otopharynx ‘auromarginatus stripe’: Turner (1996:170, 178).
Otopharynx sp. ‘tetraspilus elongate spot’: Snoeks and Hanssens
(2004:292).
Holotype.
YPM ICH 031028, 127.8 mm SL, male; Malaŵi: Lake Malaŵi,
Nankumba Peninsula, trammel nets set overnight 75 m from
shore in 9 m depth, at about 14°01⬘45⬙S 34°49⬘43⬙E; M. K.
Oliver, K. McKaye, T. D. Kocher, 24–25 August 1980. Field num-
ber MKO80–133.
Paratypes.
YPM ICH 014359, two (male), 155.0–178.6 mm SL; collected
with the holotype. YPM ICH 028511, 1 (male), 136.8 mm SL;
Malaŵi: Lake Malaŵi, southeast arm, bottom trawl across Maz-
inzi Bay, M/L Ethelwynn Trewavas, depth 9–10 m; approx.
1-km haul at roughly 14°08⬘S 34°58⬘E; M.K. Oliver, K. McKaye,
T. D. Kocher, 28 June 1980. Field number MKO80-40. [Note
added in proof: Two further (nontype) specimens, 135.9–146.4
mm SL from Monkey Bay, YPM ICH 032353, were found in a
misidentified lot while this article was in press.]
Diagnosis,
A medium-sized (<20 cm TL) laterally spotted haplochromine
separable from all previously described species of Otopharynx
both by its unique melanin pattern and by its distinctive lower
pharyngeal bone. The suprapectoral spot is rectangular, 2–2½
scales high, longitudinally elongate, covering 7–8 consecutive
scales starting with the seventh scale of the upper lateral line,
extending vertically from just above the lateral midline to slightly
above the upper lateral line. The supraanal and precaudal spots
are conjoined, forming a continuous stripe (paler at its
midlength in some specimens) from the level of the anal-fin ori-
gin to the base of the caudal fin. The lower pharyngeal bone is
broadly triangular, somewhat heart-shaped, with numerous uni-
formly small, laterally compressed, crowded, bicuspid teeth.
Description.
Morphometric and meristic data are given in Tables 1 and 2.
Deep-bodied; body depth 34.0–38.5% SL (Figure 3A).
Dorsal profile of snout straight, profile above eye straight to
slightly concave, then evenly convex from nape to end of dorsal-
fin base. Premaxillary pedicels not prominent, their angle
42–49°, interorbital angle 39–43°, nuchal angle 24–36°. Ventral
profile nearly straight from caudal end of lower jaw to chest,
irregularly convex from pelvic-fin origin to end of anal-fin base.
Jaws equal anteriorly (Figure 1B); lips neither thickened nor
lobate; lower jaw somewhat flattened, the dentary walls appear-
ing outwardly rotated. Gape inclination 30–39°. Lower-jaw
length–width ratio intermediate, the hemijaws, as seen from
below, parallel and well separated posteriorly (Figure 3C).
Lower-jaw underside angle 37–40°. Snout acuteness 71-77°.
Eyes large, slightly oval, orbit length 32.8–34.3% HL, vertical eye
diameter 28.8–31.4% HL; pupil rounded anteriorly and poste-
riorly (Figure 3B); eye not reaching dorsal head profile. Caudal
peduncle 15.0–18.1% SL, its length 1.3–1.6 times its depth.
Soft dorsal and anal fins acutely pointed and elongated pos-
teriorly (at least in males). Caudal fin deeply emarginate; upper
and lower lobes acutely pointed, subequal. Pectoral fin long,
31.0–36.2% SL, acute, upper rays elongated, reaching level of
anus. Pelvic fins with first soft ray elongate, reaching base of first
anal-fin spine in holotype and one paratype.
Dental arcade broadly arched in each jaw. Upper jaw with
53–62 teeth (total) in outer row. Anterior and lateral teeth
unequally bicuspid, occasional tricuspid teeth laterally; poste-
rior teeth unicuspid. Anterior teeth of outer row loosely
implanted, easily lost, leaving visible holes in lips. Lower-jaw
outer tooth row of haplochromis type, with 36–46 teeth (total);
crowns as in upper jaw. Inner teeth of both jaws unequally tri-
cuspid, in 1–2 rows.
Lower pharyngeal bone dissected from holotype (Figure
3D–G) and all three paratypes; subtriangular; posterior contour
of each half convexly curved, the halves meeting at nearly a 90°
angle, giving a somewhat heart-shaped outline to the bone (Fig-
ure 3F); horns short, thin, narrowing distally. Median suture
straight (Figure 3G). Keel short, deep, scarcely descending;
dentigerous surface concave in lateral view (Figure 3E). Bone
shallow in posterior view (Figure3D). Teeth bicuspid, laterally
compressed, uniformly small except for slightly larger teeth in
posterior row; crowded, especially on posterior and lateral areas
(Figure3F). Teeth in posterior row 45–50 (holotype: 49); in each
median column 11–16 (holotype: 13–14); in each oblique row
10–14 (holotype: 13–14).
Lacrimal bone (Figure 2A) bearing four neuromasts and
five lateral-line pores; lacrimal notch distinct.
Gill rakers 12 or 13, closely spaced, rather short, of nearly
uniform length along lower limb of outer arch; unpigmented
(completely lacking melanophores).
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
164

Scales ctenoid; 35–36 in lateral line. Lateral line discontin-
uous; upper section with downward kink 1–6 scales long, or (in
one paratype, bilaterally) absent. Caudal fin scaled heavily to
near tips of upper and lower lobes, stiffening them; scaled on
basal one-half of middle rays. Soft dorsal and anal fins with a
single small scale between bases of some rays. Scales transition
gradually in size from larger on belly to smaller on chest.
Coloration in life. Unknown; however, three fresh subadult
male specimens trawled in the southeast arm are shown in
Figure 4. They appear brownish on upper head, nape, and
dorsum, silvery to white on the flanks and lower head sur-
faces. Traces of about nine narrow gray vertical bars on
flanks below dorsal-fin base. All three individuals show the
characteristic dark suprapectoral spot and posterior stripe.
Dorsal-fin lappets yellow; dorsal has dark-edged yellow
maculae throughout, most distinct in soft part of fin. Broad
upper and lower leading edges of caudal fin are pale yellow,
remainder of fin darker. Anal fin bears about four oval yel-
low eggspots in two rows along middle and near distal edge
Six New Species of the Cichlid Genus Otopharynx • Oliver 165
TABLE 1. Morphometric characters of Otopharynx alpha. Abbreviations: LLL, lower lateral line.
Holotype Paratypes
YPM ICH YPM ICH YPM ICH
031028 028511 014359
Standard length (mm) 127.8 136.8 141.0 117.9
Head length (mm) 35.7 37.4 40.4 34.3
Percentage of standard length:
Head length 28.0 27.4 28.7 29.1
Body depth 35.3 38.5 34.0 35.2
Dorsal-fin base length 59.6 61.3 59.3 58.3
Predorsal length 31.4 30.5 31.1 32.1
Prepectoral length 28.0 26.9 30.0 28.6
Prepelvic length 37.2 34.9 38.6 37.3
Preanal length 69.0 67.0 68.9 68.3
Pelvic fin origin to anal fin origin 33.3 34.2 31.4 31.9
Anal-fin base length 18.3 17.2 17.5 18.3
Caudal peduncle length 15.0 17.8 17.7 18.1
Caudal peduncle depth 11.2 11.6 10.9 11.4
Pectoral-fin length 34.2 31.0 36.2 33.2
Pelvic-fin length 28.5 32.0 34.1 30.8
Dorsal-fin origin to anal-fin origin 53.6 55.1 52.5 54.4
Dorsal-fin origin to end of anal-fin base 63.1 64.6 62.5 63.4
Pelvic-fin origin to end of dorsal-fin base 55.6 57.2 54.4 53.3
End dorsal-fin base to end anal-fin base 15.2 14.6 14.2 15.7
Anal-fin origin to end of dorsal-fin base 29.1 28.4 28.9 30.2
Dorsal-fin origin to pelvic-fin origin 29.1 37.3 34.7 34.5
End dorsal-fin base to end hypurals at LLL 18.4 19.4 17.5 18.9
End anal-fin base to end hypurals at LLL 18.1 19.2 19.7 20.1
Percentage of head length:
Head width 45.9 48.1 45.0 46.2
Interorbital width 25.6 26.7 24.7 24.7
Snout length 33.4 30.9 34.0 31.4
Snout width 32.9 33.4 33.8 34.7
Lower-jaw length 37.2 38.4 35.3 36.5
Lower-jaw width 23.7 20.3 27.2 23.5
Premaxillary pedicel length 30.1 29.5 30.6 30.2
Upper-jaw length 26.5 27.5 29.7 26.8
Cheek depth 20.9 24.5 21.0 22.2
Orbit length 33.9 34.2 32.8 34.3
Vertical eye diameter 30.6 29.7 28.8 31.4
Lacrimal (preorbital) depth 21.4 22.0 24.1 22.0
Postorbital head length 39.2 40.7 40.3 39.3

of fin. Pelvic fins yellow with white leading edges. Pectoral
fins are yellowish.
Coloration in preservative (Figure 3A–C). Head and flanks
brown above, becoming silvery below. Head with distinct oper-
cular spot and an indefinite dark oblique bar from below front
of eye to rear half of upper jaw. Underside of head and bran-
chiostegal membrane brownish (Figure 3C). Suprapectoral spot
a longitudinally elongate rectangle 2–2½ scales high, covering
7–8 consecutive scales starting with seventh scale of upper lat-
eral line, based just above lateral midline and reaching to one-
half scale above upper lateral line. Supraanal and precaudal spots
united into a continuous stripe, which may or may not be paler
at its midlength, from level of anal-fin origin to base of caudal
fin. Lower edge of stripe on scale row containing lower lateral
line, which is one scale row lower than bottom of suprapectoral
spot. About eight faint vertical bars visible on flanks of two
paratypes. Dorsal fin with series of dark-edged oval maculae
between all spines and rays, more distinct in soft dorsal; lappets
pale (but fin margin of largest paratype continuous, without lap-
pets). Caudal fin yellowish to brownish with indistinct marbling.
Anal and pelvic fins dark brown, anal with no eggspots. Pec-
torals hyaline.
Maximum size. The largest specimen known to me is a paratype
measuring 178.6 mm TL.
Parasites.
The holotype has several lernaeid copepods, Lamproglena
nyasae Fryer, 1956, attached to filaments of the outermost gill
arches bilaterally. Two individuals were removed and deposited
in the YPM invertebrate collection (YPM IZ 102202); photo-
graph available at http://collections.peabody.yale.edu/search/
Record/YPM-IZ-102202.
Distribution.
The known range of O. alpha includes the central part of the
southeast arm, where Turner (1996:178) recorded it (as O.“auro-
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
166
TABLE 2. Meristic characters of Otopharynx alpha.
Holotype Paratypes
YPM ICH YPM ICH YPM ICH
031028 028511 014359
Squamation:
Lateral-line scales 35 36 35 36
Upper lateral-line scales 29 26 24 26
Lower lateral-line scales 18 16 13 15
Lateral-line scales on caudal fin 2123
Kink length 6102
Upper transverse line scales 5556
Lower transverse line scales 10 10 10 11
Cheek scale rows 3322
Scales between pectoral- and 9 10 99
pelvic-fin bases
Fins:
Dorsal-fin spines 17 17 17 17
Dorsal-fin segmented rays 11 12 12 12
Dorsal-fin total elements 28 29 29 29
Anal-fin spines 3333
Anal-fin segmented rays 9 10 99
Anal-fin total elements 12 13 12 12
Pectoral-fin rays 15 14 14 14
Gill rakers:
Epibranchial 5455
Angle 1111
Lower limb 12 13 13 12
Total 18 18 19 18
Oral teeth:
Outer upper-jaw teeth: 28/28 = 56 27/26 = 53 31/31 = 62 29/? = ~58
left/right = total
Outer lower-jaw teeth: 22/23 = 45 22/22 = 44 23/23 = 46 18/18 = 36
left/right = total

Six New Species of the Cichlid Genus Otopharynx • Oliver 167
FIGURE 3. Otopharynx alpha. A. Holotype, YPM ICH 031028, 127.8 mm SL, male. B. Head of holotype. C. Under-
side of head of holotype. D–G. Lower pharyngeal bone of holotype, in posterior (D), left lateral (E), dorsal (F),
and ventral (G) views.

marginatus stripe”) from trawl stations off Nkope, Mazinzi,
Fowo, Chiponda, and Chekopa as well as at Chapola Shoal, all
over sand at depths of 5–30 m. Snoeks and Hanssens (2004:292)
illustrated two specimens (as Otopharynx sp. “tetraspilus elon-
gate spot”), one from Mazinzi to Kadango in the southeast arm,
the other from Chembe at the head of the Nankumba Peninsula
between the southwest and southeast arms; capture depths were
not stated. (These two specimens were not available for loan from
MRAC when I was preparing this description.) The type local-
ity (Figure 1), west of Chembe and about 1.4 km northeast of
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
168
FIGURE 4. Otopharynx alpha, freshly captured individuals trawled in 5–10 m depth on Chapola Shoal, east of
Boadzulu Island in the southeast arm of Lake Malaŵi, 31 July 1991. Apparently subadult or young nonbreeding
males. Photograph by George Turner.

Otter Point in 9 m depth, is the furthest west occurrence known
to me. It would not be surprising if it also occurs in the southwest
arm.
Etymology.
From ␣␭φ␣(alfa), first letter of the Greek alphabet. The
suprapectoral spot and posterior stripe suggest a dot and dash,
which represents “A” in International Morse Code. A noun in
apposition.
Remarks.
At the type locality, the substrate in 9 m depth consists of fine
sand with few if any macrophytes. Other species captured in the
trammel nets together with the holotype and two paratypes of
O. alpha included Buccochromis lepturus, Hemitaeniochromis
urotaenia, Hemitilapia oxyrhyncha, Protomelas spilopterus, and
Trematocranus microstoma. The remaining paratype was
trawled over sand in 9–10 m along with numerous species
including Aulonocara rostratum, Gephyrochromis moorii,
Lethrinops albus, L. cf. brevis, L. cf. christyi, L. furcifer, L. lethri-
nus, Mylochromis sphaerodon, Nyassachromis nigritaeniatus, N.
sp., Protomelas annectens, Taeniolethrinops furcicauda, T. praeor-
bitalis, Tramitichromis intermedius, T. lituris, Trematocranus
microstoma, T. placodon, and Synodontis njassae.
Gut contents were not examined. The diet, and the func-
tion of the specialized lower pharyngeal bone and its dentition,
remains to be determined.
Snoeks and Hanssens (2004:294–295) briefly described
and illustrated a cichlid resembling O. alpha that they called
Otopharynx sp. “elongate spot” (distinct from their species O.
sp. “tetraspilus elongate spot”). They stated that O. sp. “elon-
gate spot” “may be conspecific with O. ‘auromarginatus stripe’
reported by Turner (1996).” However, this is unlikely, as they
noted that their species, from Jafua Bay, Mozambique, has
enlarged median pharyngeal teeth. In contrast, Turner
(1996:178) describes the pharyngeal teeth of O. “auromar-
ginatus stripe” as “small crowded teeth, the three posterior
teeth of innermost row slightly enlarged.” The latter descrip-
tion accords well with the types of O. alpha, which also lack
clearly enlarged pharyngeal teeth in the median columns (Fig-
ure 3F).
According to Turner (1996), this species (as O. “auromar-
ginatus stripe”) was probably of minor commercial importance,
being caught by pair trawling and seining. Turner (1996)
remarked that in Malaŵi Fisheries Department records it had
been confused with Nyassachromis leuciscus, a species lacking a
suprapectoral spot. The current population status of O. alphais
unknown.
Otopharynx mumboensis
new species
Zoobank.org/urn:lsid:zoobank.org:act:
20B3BCED-DB76-487E-A084-5D956BE7BC8D
Figures 2B, 5, 6, Tables 3, 4
Protomelas sp. ‘virgatus mumbo’: Konings (2001:227, 228,
2015:208, 2016:295).
Holotype.
YPM ICH 031031, 115.7 mm SL, male; Malaŵi: Lake Malaŵi,
Mumbo Island, west shore, chased into ½⬙mesh gillnet with
SCUBA in 6–9 m depth, at about 13°59⬘30⬙S 34°45⬘10⬙E; M.K.
Oliver, K. McKaye, T. D. Kocher, 5 August 1980. Field number
MKO80-96.
Paratypes.
YPM ICH 024990, two (one male, one female), 103.1–107.3 mm
SL; collected with the holotype.
Diagnosis.
A smallish (~14 cm TL) laterally spotted haplochromine distin-
guished from most Otopharynx species by its suprapectoral spot,
which is situated entirely below the upper lateral line and sepa-
rated from it by about half the spot height. This spot is roughly
rectangular, longitudinally elongate (1–1½ scales high, covering
about seven consecutive scales), about three times as long as tall.
The placement and shape of the suprapectoral spot immediately
distinguish O. mumboensis from O. antron, O. argyrosoma, O.
auromarginatus, O. decorus, O. heterodon,O. pachycheilus,
O. selenurus, O. speciosus, O. spelaeotes, O. tetraspilus, and O.
tetrastigma, all of which have the suprapectoral spot touching
or usually extending above the upper lateral line, or (O. selenu-
rus) may have no suprapectoral spot. Only O. brooksi, O. litho-
bates, and O. ovatushave a suprapectoral spot similar to that of
O. mumboensis. Compared to O. brooksi,O. mumboensis has a
shorter head (31–32% SL vs. 36–39%), fewer outer teeth in the
upper jaw (47–50 vs. 69–87), and more gill rakers (13 on lower
limb vs. 11–12). Compared to O. lithobates, O. mumboensis is
deeper bodied (body depth 35.5–36.1% SL vs. 29.5–34.4%), has
a shorter lower jaw (35.7–36.2% HL vs.36.9–42.0), and has a
lower pharyngeal bone with all teeth of the median columns
somewhat enlarged, with subcylindrical shafts and submolari-
form crowns (vs. with slightly enlarged submolariform teeth
confined to the posteromedian area). Compared to O. ovatus,
O. mumboensis has a shorter lower jaw (35.7–36.2% HL vs.
43.2–44.3% in the lectotype and two paralectotypes), larger eye
(orbit length 34.8–35.4% HL vs. 27.9–32.9%), narrower interor-
bital width (22.6–24.0% HL vs. 26.5–29.6%), more-triangular
lower pharyngeal bone with the posterior contour nearly
straight (vs. more Y-shaped and rather deeply notched posteri-
orly), and all teeth in the median columns of the lower pharyn-
geal bone somewhat molariform, the crowns broadened and
flattened (vs. the teeth in these columns bicuspid, their crowns
not molarized).
Description.
Morphometric and meristic data are given in Tables 3 and 4.
Deep-bodied; body depth 35.5–36.1% SL (Figure 5A).
Dorsal profile straight from snout tip to above rear of orbit,
evenly rounded from there to end of dorsal-fin base, more con-
vex than ventral profile. Premaxillary pedicels not prominent,
their angle 42–46°, interorbital angle 42–44°, nuchal angle
16–20°. Ventral profile nearly straight from lower-jaw tip to
below pectoral-fin base, chest profile inclined ~20°. Jaws equal
anteriorly, lower jaw not dorsoventrally flattened; lips neither
Six New Species of the Cichlid Genus Otopharynx • Oliver 169

thickened nor lobate. Gape inclination 39–45°. Lower-jaw
length–width ratio intermediate, the hemijaws, as seen from
below, parallel or diverging and well separated posteriorly (Fig-
ure 5C). Lower-jaw underside angle 34–42°. Snout acuteness
71–77°. Eyes large, circular (Figure 5B), orbit length ~35% HL;
pupil nearly round; eye not reaching dorsal head profile. Caudal
peduncle 16.1–17.0% SL, its length 1.4–1.6 times its depth.
Dorsal and anal fins rounded posteriorly, without elon-
gated rays. Caudal fin shallowly emarginate. Pectoral fin
25.6–32.3% SL, reaching level of anus.
Dental arcade in each jaw rounded. Upper jaw with 47–51
teeth (total) in the outer row. Outer upper-jaw teeth stout, bicus-
pid (an occasional one tricuspid), crowns slightly recurved with
major cusp rounded; last three to four unicuspid or with small
shoulder of minor cusp. Anterior teeth closely spaced, inclined
slightly toward symphysis. Outer row of lower jaw of hap-
lochromis type, with 37–40 teeth (total); teeth similar to those of
upper jaw; anterior teeth erect, crowns slightly recurved. Many
outer teeth of both jaws, especially in holotype, have crowns bro-
ken or heavily worn. Inner teeth in (two) three to four rows in
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
170
TABLE 3. Morphometric characters of Otopharynx mumboensis. Abbreviations: LLL, lower lateral line.
Holotype Paratypes
YPM ICH 031031 YPM ICH 007820
Standard length (mm) 115.7 103.1 107.3
Head length (mm) 37.2 31.8 33.6
Percentage of standard length:
Head length 32.2 31.3 30.9
Body depth 36.1 35.8 35.5
Dorsal-fin base length 56.5 58.0 56.8
Predorsal length 35.9 34.8 35.7
Prepectoral length 32.4 31.6 31.2
Prepelvic length 39.6 38.7 39.3
Preanal length 66.6 64.5 67.6
Pelvic fin origin to anal fin origin 27.8 27.6 29.5
Anal-fin base length 18.7 21.6 19.9
Caudal peduncle length 17.0 17.2 16.1
Caudal peduncle depth 10.9 11.5 11.9
Pectoral-fin length 25.7 25.6 32.3
Pelvic-fin length 25.8 28.8 26.9
Dorsal-fin origin to anal-fin origin 48.9 48.1 49.5
Dorsal-fin origin to end of anal-fin base 59.7 60.9 60.7
Pelvic-fin origin to end of dorsal-fin base 53.1 54.6 55.0
End dorsal-fin base to end anal-fin base 13.6 14.5 15.4
Anal-fin origin to end of dorsal-fin base 28.3 31.3 30.8
Dorsal-fin origin to pelvic-fin origin 36.2 34.8 34.9
End dorsal-fin base to end hypurals at LLL 17.1 16.1 16.6
End anal-fin base to end hypurals at LLL 19.1 19.2 18.5
Percentage of head length:
Head width 47.6 47.5 47.0
Interorbital width 24.0 23.5 22.6
Snout length 31.7 29.6 31.3
Snout width 34.2 33.3 34.6
Lower-jaw length 36.2 35.7 36.2
Lower-jaw width 26.3 25.4 25.5
Premaxillary pedicel length 26.8 27.5 28.4
Upper-jaw length 32.7 30.8 28.9
Cheek depth 23.1 22.1 20.5
Orbit length 35.2 35.4 34.8
Vertical eye diameter 33.7 33.3 33.2
Lacrimal (preorbital) depth 19.7 19.6 20.4
Postorbital head length 37.9 38.7 38.8

each jaw; tricuspid with major cusp often semicircular; inner
teeth relatively large although clearly smaller than outer teeth;
unusual in that crowns are pigmented red-brown as in outer
teeth.
Lower pharyngeal bone dissected from holotype (Figure
5D–G) and one paratype (107.3 mm SL); subtriangular, lightly
built, dorsoventrally compressed in posterior view (Figure 5D);
posterior contour nearly straight (slightly emarginate); horns
slender, each horn terminating in an L-shaped facet whose cau-
dal flange is autapomorphically elongated into a fingerlike
process three to four times as long as its lateral flange. Median
suture straight to slightly sinuous, without interdigitations (Fig-
ure5G). Keel short, scarcely descending, its depth nearly equal
to its length, strongly convex below (Figure 5E). All teeth of
median four to six columns enlarged, stouter than more lateral
teeth, crowns somewhat molarized, more strongly so in the cau-
dal half of these columns (Figure 5F). Posterolateral teeth small,
laterally compressed, crowded, and bicuspid. Teeth in posterior
row 36 (holotype)–42; in each median column 9–10; in each
oblique row 7–8.
Lacrimal bone (Figure 2B) bearing four neuromasts and
five lateral-line pores; lacrimal notch distinct.
Gill rakers 13 on lower arch; slender, unbranched; a few
widely spaced melanophores on each raker.
Scales ctenoid; 34–35 in lateral line. Lateral line discontin-
uous, upper section with downward kink one to four scales long.
Squamation extending onto caudal fin between fin rays, to near
tips of upper and lower lobes and on basal one-half along
middle rays. Soft dorsal and anal fins with one to three small
scales between bases of some rays. Size transition gradual
between larger abdominal and smaller thoracic scales.
Coloration in life (Figure 6A, B). Live coloration of males
unknown. A young fish, possibly female, shown (as Protome-
las sp. “virgatus mumbo”) in a published underwater photo
(Konings 2015:208, 2016:295) and reproduced here as Figure
6A, shows the head and body gray to creamy white. Each flank
scale has a yellow-orange basal spot. Melanic markings com-
prise three prominent midlateral dark spots including a
suprapectoral spot about three times as long as high, situated
entirely below upper lateral line and separated from it; a
supraanal spot about twice as long as high; and a precaudal
spot at end of caudal peduncle. A row of three small dark spots
above anterior part of upper lateral line: one at the level of hind
edge of operculum and others just above lateral line over front
and rear of suprapectoral spot. Five small dorsal midline spots
just below dorsal-fin base. About nine faint, narrow vertical
bars below base of dorsal fin. A faint opercular spot. No dis-
Six New Species of the Cichlid Genus Otopharynx • Oliver 171
TABLE 4. Meristic characters of Otopharynx mumboensis.
Holotype Paratypes
YPM ICH 031031 YPM ICH 024990
Squamation:
Lateral-line scales 35 35? 34
Upper lateral-line scales 25 26+ 27
Lower lateral-line scales 16 — 16
Lateral-line scales on caudal fin 1— 2
Kink length 13 4
Upper transverse line scales 55 5
Lower transverse line scales 9 9? 10?
Cheek scale rows 33 3
Scales between pectoral- and pelvic-fin bases 86 7?
Fins:
Dorsal-fin spines 16 17 16
Dorsal-fin segmented rays 11 11 11
Dorsal-fin total elements 27 28 27
Anal-fin spines 33 3
Anal-fin segmented rays 9 10 9
Anal-fin total elements 12 13 12
Pectoral-fin rays 14 14 14
Gill rakers:
Epibranchial 55 4
Angle 11 1
Lower limb 13 13 13
Total 19 19 18
Oral teeth:
Outer upper-jaw teeth: left/right = total 23/27 = 50 25/26 = 51 23/24 = 47
Outer lower-jaw teeth: left/right = total 21/19 = 40 18/21 = 39 19/18 = 37

Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
172
FIGURE 5. Otopharynx mumboensis. A. Holotype, YPM ICH 031031, 115.7 mm SL, male. B. Head of holotype.
C. Underside of head of holotype. D–G. Lower pharyngeal bone of holotype, in posterior (D), left lateral (E),
dorsal (F), and ventral (G) views.

tinct markings on snout, nape, or fins. This melanic pattern is
essentially identical to that of O. lithobates, O. brooksi, and O.
ovatus. The yellowish scale spots and the dark markings on a
larger individual (Figure 6B) are similar; however, the dorsal
midline spots and the spots just above the upper lateral line
are darker and more prominent, and indistinct dark marks are
present between the midlateral spots. The dorsal fin has pale
yellowish lappets and there are yellow oval maculae between
the soft dorsal rays. Between the caudal-fin rays the membrane
is yellow. There appears to be a yellow eggspot in the soft anal
fin.
Coloration in preservative (Figure 5A–C). Although the type
specimens are slightly cleared, possibly because they were fixed
in inadequately buffered formalin prepared from paraformalde-
hyde, their melanic markings remain visible when the speci-
mens are rotated in the hand and closely correspond to those of
the living individuals described above. Head and body almost
uniformly brown. Suprapectoral spot 1–1½ scales high, cover-
ing about seven consecutive scales, below and separated from
upper lateral line by about half height of spot; supraanal spot
slightly longer than high; precaudal spot at end of caudal pedun-
cle, shaped like supraanal spot. Three spots above anterior part
of upper lateral line as described for living specimen. Five dor-
sal midline spots, evenly spaced, just below base of dorsal fin.
Traces of vertical bars, too faint to count accurately. Head lack-
ing bars or stripes; with an opercular spot. Dorsal-fin lappets
pale distally, brown proximally grading into poorly defined sub-
Six New Species of the Cichlid Genus Otopharynx • Oliver 173
FIGURE 6. Otopharynx mumboensis, living individuals, underwater at Mumbo Island (depths not reported).
A. Young, apparently female individual. B. Larger adult (right side reversed), apparently female, over Vallisneria
bed. Photographs by Ad Konings.

marginal dark stripe; dorsal-fin membrane brown overall, with
vague maculae between segmented rays. Caudal, anal, and
pelvic fins brown; anal fin without eggspots or other distinct
markings; pectorals hyaline.
Maximum size. The largest known specimen is the holotype,
140.6 mm TL.
Parasites.
All of the type specimens have metacercariae of a digenetic
trematode on the head, body, and fins; the holotype is especially
heavily covered (Figure 5B, C). Similar metacercarial cysts are
visible on the head, chest, and pectoral fin of the fish in Figure
6B. The cysts may be those of Astiotrema turneri, the only iden-
tified digenean from mbuna and non-mbuna cichlids of Lake
Malaŵi (Bray et al. 2006; Bray and Hendrix 2007).
Distribution.
Known only from Mumbo Island in the southwest arm of Lake
Malaŵi (Figure 1).
Etymology.
The name is an adjective referring to the type locality, Mumbo
Island.
Remarks.
The intestine, observed in situ in the holotype and one paratype,
has multiple coils. That of the holotype contains fine gravel along
much of its length, which was not noted in the examined
paratype. The diet is unknown.
Otopharynx mumboensisappears to be a rare species. The
types, apparently the only museum specimens extant, were col-
lected in 6–9 m depth in the intermediate zone (80% rocks /
20% sand). Small isolated patches of the invasive Myriophyllum
spicatum L. (Weyl and Coetzee 2014) were the only evident
macrophytes in the area. (Figure 6B shows an individual of O.
mumboensis over sand with Vallisneria spiralis L.) Other cichlid
species collected with the type series include Buccochromis het-
erotaenia, Cheilochromis euchilus, Docimodus johnstoni,
Mylochromis epichorialis, Otopharynx lithobates, O. ovatus, Stig-
matochromis woodi, Protomelas fenestratus, Tyrannochromis
macrostoma, T. nigriventer, and Labeotropheus trewavasae.
Mbuna were not specifically sought in this particular collecting
effort.
Mumbo Island occupies an isolated position in the south-
west arm of Lake Malaŵi. Its petricolous, philopatric cichlids
are isolated by the surrounding sandy substrate. The nearest
rocky or intermediate rocky/sandy shores are at Zimbawe Rock
5.3 km to the northeast, Thumbi Island West 5.7 km to the
southeast, and Domwe Island 6.7 km to the east-northeast.
Observations at Zimbawe Rock and multiple collecting stations
on various rocky and intermediate zone sites at Thumbi West
and Domwe, employing a variety of techniques, including use of
ichthyocide, yielded no individuals referable to O. mumboensis
(Oliver, unpublished).
Otopharynx mumboensis is the first formally named non-
mbuna cichlid species that is apparently restricted to Mumbo
Island. Among described mbuna species, only Labidochromis
mylodon Lewis, 1982, has a known distribution restricted to
Mumbo Island. Labeotropheus artatorostris is known from both
Mumbo and Thumbi West islands (Pauers 2017). The type
locality of Maylandia chrysomallos is Mumbo Island, but this
species also occurs on the lake’s eastern shore from Meponda to
Makanjila Point (Konings 2015). Melanochromis mellitus,
described from “off Mumbo Island,” has been synonymized with
the widespread M. melanopterus (Konings-Dudin et al. 2009).
Lundeba et al. (2011) described Petrotilapia mumboensis, but
they state that this mbuna species occurs not only at Mumbo
but also at Mbenji and Thumbi West islands. In addition, several
distinctively colored undescribed species of mbuna at Mumbo
(Konings 2015) may prove to be endemic to the island.
Although the known range of O. mumboensis is within the
boundaries of Lake Malaŵi National Park, this species should be
considered vulnerable, given its evidently restricted range, its
rarity there, and the presence of illegal net fishing operations
reported at Mumbo Island and elsewhere in Lake Malaŵi
National Park despite the legal prohibition against fishing within
100 m of the shore (e.g., Nyanyale 2005; McKenzie and Swalls
2010; Further developments 2011). The ban is “routinely flouted
by artisanal fishermen” within the park (IUCN World Heritage
Outlook 2017).
Konings (2016:295) does not explain the basis of his belief
that this species (as Protomelas sp. “virgatus mumbo”), despite
sharing the identical apomorphic, laterally spotted color pattern
of several other Otopharynx species, actually belongs in Protome-
las—a genus whose major diagnostic character is a plesiomor-
phic unbroken midlateral stripe (Eccles and Trewavas 1989).
Otopharynx styrax
new species
Zoobank.org/urn:lsid:zoobank.org:act:
8C70B6BD-8332-4C83-86C2-3117496FD7CA
Figures 2C, 7, 8, Tables 5, 6
Mylochromis sp. ‘torpedo blue’: Konings (1995:232, 260, 261).
Otopharynx ‘productus’: Turner (1996:171, 179); Duponchelle
et al. (2000:127).
Otopharynx sp. ‘torpedo blue’: Konings (2001:258, 260,
2016:341).
Otopharynx sp. ‘productus sharp snout’: Snoeks and Hanssens
(2004:292, fig. 75).
Holotype.
YPM ICH 031029, 97.3 mm SL, immature male; Malaŵi: Lake
Malaŵi, southeast arm, Lake Malaŵi Trawling Survey station
Mazinzi II, M/L Ethelwynn Trewavas, two 0.5-hr bottom
trawls off Mazinzi in 42 m depth, at about 14°07⬘17⬙S
35°00⬘14⬙E–14°06⬘15⬙S 34°59⬘53⬙E; M. K. Oliver, J. Tarbit, K.
Stride, 25 June 1971. Field number MKO71-VI-25.
Paratypes,
YPM ICH 014270, eight (five male, three female), 88.8–116.7
mm SL; collected with the holotype.
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
174

Diagnosis.
A medium-sized (~20 cm TL), laterally spotted haplochromine
distinguishable from all previously described Otopharynx
species by its more elongate body (depth 24.3–28.6% SL). The
next-most-elongate Otopharynx species are O. decorus (depth
29.4–33.3% SL) and O. lithobates(depth 29.5–34.4% SL); all oth-
ers have a body depth >31% SL (data from Oliver 1984 and
Eccles and Trewavas 1989). Otopharynx styrax is further sepa-
rated from O. decorus by its longer caudal peduncle
(length/depth 2.0–2.3, vs. 1.5–1.6 in O. decorus) and its narrower
lower pharyngeal bone, the lateral edges of the horns diverging
at ~70°(Figure7G, H) vs. 79–87°in the lectotype and a paralec-
totype of O. decorus (Oliver, unpublished). Otopharynx styrax
is further distinguished from O. lithobatesby the absence of dark
spots on the dorsum below the dorsal-fin base (vs. 4–6 distinct
dorsal midline spots in O. lithobates) and longer caudal pedun-
cle (length/depth 2.0–2.3, vs. 1.3–1.7 in O. lithobates).
Six New Species of the Cichlid Genus Otopharynx • Oliver 175
TABLE 5. Morphometric characters of Otopharynx styrax. Abbreviations: LLL, lower lateral line; S.D., standard
deviation.
Paratypes
Holotype NMean S.D. Range
Standard length (mm) 97.3 8 97.6 8.8 88.8–116.7
Head length (mm) 27.8 8 28.6 2.7 26.8–34.9
Percentage of standard length:
Head length 28.6 8 29.3 0.7 28.2–30.2
Body depth 24.3 8 26.4 1.4 24.9–28.6
Dorsal-fin base length 54.9 8 54.7 0.6 53.9–55.5
Predorsal length 31.1 8 33.2 0.7 32.1–33.8
Prepectoral length 28.5 8 29.4 0.9 27.9–31.1
Prepelvic length 32.9 8 34.5 1.3 33.3–37.0
Preanal length 63.1 8 63.7 0.5 62.5–64.1
Pelvic fin origin to anal fin origin 29.8 8 29.6 0.9 28.4–30.9
Anal-fin base length 18.6 8 18.2 0.5 17.7–18.9
Caudal peduncle length 18.6 8 19.5 0.8 18.7–20.9
Caudal peduncle depth 8.9 8 9.2 0.6 8.5–10.3
Pectoral-fin length 26.2 8 28.1 1.3 26.6–30.9
Pelvic-fin length 20.8 8 21.5 1.0 20.6–22.1
Dorsal-fin origin to anal-fin origin 42.6 8 43.3 0.8 42.1–44.3
Dorsal-fin origin to end of anal-fin base 56.4 8 56.0 1.0 54.5–57.1
Pelvic-fin origin to end of dorsal-fin base 52.6 8 52.4 0.8 51.7–53.6
End dorsal-fin base to end anal-fin base 12.7 8 12.9 0.6 12.1–14.0
Anal-fin origin to end of dorsal-fin base 26.2 8 26.5 0.7 25.9–27.6
Dorsal-fin origin to pelvic-fin origin 24.4 8 26.3 1.5 24.5–28.7
End dorsal-fin base to end hypurals at LLL 18.6 8 18.8 0.7 17.5–19.5
End anal-fin base to end hypurals at LLL 19.7 8 20.9 0.8 19.9–22.0
Percentage of head length:
Head width 42.5 8 43.5 1.0 42.6–45.5
Interorbital width 15.4 8 17.2 1.0 16.0–18.9
Snout length 32.8 8 32.6 2.0 29.0–35.8
Snout width 25.6 8 26.5 1.0 25.5–28.8
Lower-jaw length 36.2 8 36.2 1.4 33.8–37.6
Lower-jaw width 14.5 8 16.6 1.6 15.0–20.3
Premaxillary pedicel length 30.2 8 30.7 2.1 27.1–33.5
Upper-jaw length 23.8 8 24.3 1.0 23.0–26.1
Cheek depth 20.7 8 20.5 2.0 16.8–22.5
Orbit length 35.1 8 36.0 1.2 34.1–37.7
Vertical eye diameter 28.5 8 28.6 1.4 26.1–30.5
Lacrimal (preorbital) depth 22.8 8 22.4 1.8 20.2–24.5
Postorbital head length 38.1 8 37.6 1.2 35.2–39.5

Morphometric and meristic data are given in Tables 5
and 6.
Body elongate, its depth 24.3–28.6% SL (Figure 7A).
Head sharply pointed in lateral view (Figure 7B) and narrowly
rounded in dorsal view (Figure 7C). Dorsal profile evenly
convex from tip of snout to end of dorsal-fin base, except
snout profile nearly straight in some individuals. Premaxil-
lary pedicels not prominent, their angle 34–50°, interorbital
angle 27–38°, nuchal angle 15–25°. Ventral profile varying
from a virtual mirror image of the dorsal profile, to less con-
vex and with an almost straight lower head and chest profile.
Jaws equal anteriorly or lower jaw slightly projecting, lower
jaw somewhat dorsoventrally flattened; lips not thickened or
lobate. Gape inclination 10–30°. Lower-jaw length–width
ratio narrow, the hemijaws, as seen from below, converging
toward the rear and nearly touching posteriorly (Figure 7D).
Lower-jaw underside angle 10–30°. Snout acuteness 65–76°.
Eyes large, elliptical (orbit length 34–38% HL, about 25%
longer than vertical eye diameter); pupil pointed anteriorly,
rounded posteriorly; eye reaching dorsal head profile, or
nearly so. Caudal peduncle 18.6–20.9% SL, its length 2.0–2.3
times its depth.
Soft dorsal and anal fins pointed but short posteriorly. Fin
spines slender, delicate, flexible (“featherfin” condition; Snoeks
and Hanssens 2004). Caudal crescentically emarginate; upper
and lower lobes acutely pointed, subequal. Pectoral fins short,
26.2–30.9% SL, acute, reaching level of anus. Pelvic fins shorter
than pectorals (20.6–22.1% SL).
Dental arcade narrowly arched in each jaw. Upper jaw with
49–64 teeth (total) in outer row. Largest paratype (116.7 mm SL)
and most other specimens have anterior teeth subequally bicus-
pid, lateral to posterolateral teeth mostly unequally bicuspid, a
few posterior teeth unicuspid. Outer row of lower jaw of hap-
lochromis type, with 46–57 teeth (total). Anterolateral teeth
angled outward, procumbent, anterior teeth with shafts angled
toward dentary symphysis. With jaws closed, anterior and
anterolateral teeth of lower jaw lie outside upper-jaw teeth, but
lateral and posterior teeth are inside those of upper jaw. Inner
teeth of both jaws unequally tricuspid with middle cusp longest,
acute, crowns lightly or not pigmented, arranged in two to three
rows.
Lower pharyngeal bone dissected from holotype (Figure
7E–H) and four paratypes (90.1–116.7 mm SL); Y-shaped, nar-
row, rather lightly built; posterior contour deeply emarginate,
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
176
TABLE 6. Meristic characters of Otopharynx styrax. Abbreviations: Q, quartile.
Paratypes
Holotype NMedian 1st Q3
rd Q Range
Squamation:
Lateral-line scales 37 7 37 37 37 36–39
Upper lateral-line scales 28 7 27 25 28 24–30
Lower lateral-line scales ≥17 7 16 15 17 14–20
Lateral-line scales on caudal fin 282 22 2–2
Kink length 160.5 0 2.50 0–4
Upper transverse line scales 677 78 7–8
Lower transverse line scales 8810 8 10 8–11
Cheek scale rows 383 34 3–4
Scales between pectoral- and pelvic-fin bases —89 89.75 7–12
Fins:
Dorsal-fin spines 16 8 17 17 17 16–17
Dorsal-fin segmented rays 13 8 13 12.25 13 12–14
Dorsal-fin total elements 29 8 30 29.25 30 29–30
Anal-fin spines 383 33 3–3
Anal-fin segmented rays 10 8 9.5 9 10 9–10
Anal-fin total elements 13 8 12.5 12 13 12–13
Pectoral-fin rays 15 8 15 15 15 15–15
Gill rakers:
Epibranchial 585 4515–15
Angle 181 11 1–1
Lower limb 12 8 12 12 12.75 11–13
Total 18 8 17.5 17 18.75 17–19
Oral teeth:
Outer upper-jaw teeth (total both sides) 52 8 58 53 62.75 49–64
Outer lower-jaw teeth (total both sides) 46 6 54 49.5 55.50 48–57

Six New Species of the Cichlid Genus Otopharynx • Oliver 177
FIGURE 7. Otopharynx styrax. A. Holotype, YPM ICH 031029, 97.3 mm SL, immature male. B. Head of holotype.
C. Dorsal view of head of holotype. D. Underside of head of holotype. E–H. Lower pharyngeal bone of holotype,
in posterior (E), left lateral (F), dorsal (G), and ventral (H) views.

each half slightly convex, the halves meeting at an obtuse angle;
horns moderately long, slender, of uniform diameter (Figure
7G). Median suture nearly straight (Figure 7H). Keel short,
straight, moderately deep, convex below (Figure7F). Bone mod-
erately deep in posterior view (Figure7E). Teeth in lateral fields
small, closely set, laterally compressed and cuspidate, but six to
nine posteromedian teeth enlarged, their crowns molarized but
retaining a posterior cusp (Figure 7G). Teeth in posterior row
37–45 (holotype: 38); in each median column 9–10 (holotype:
9); in each oblique row 5–7 (holotype: 6–7).
Lacrimal bone (Figure 2C) bearing four neuromasts and
five lateral-line pores; lacrimal notch distinct.
Gill rakers 11–13 on lower outer arch; slender, unbranched;
lacking melanophores.
Scales ctenoid; 36–39 (median 37) in lateral line. Lateral
line discontinuous; upper section with downward kink one to
four scales long, or kink absent. Caudal fin scaled to near tips of
upper and lower lobes and on basal one-half or more along mid-
dle rays. Soft dorsal and anal fins with a few small scales between
bases of some rays. Large scales on lower flanks abruptly transi-
tion to much smaller scales on chest along a line between pec-
toral- and pelvic-fin bases (much as in Thoracochromis wingatii;
see Greenwood 1979, fig. 2).
Coloration in life. Living subadult and adult males (Figure 8A–D)
dark blue on dorsum and upper head surface, paler blue on sides
of head and flanks, whitish on belly. Longitudinally elongate,
blackish suprapectoral and supraanal spots and nearly circular
precaudal spot variably present, probably dependent on emo-
tional state. No dorsal midline spots. Seven or eight faint gray
vertical bars visible below dorsal-fin base. Dorsal-fin lappets
white, fin otherwise blue with orange maculae in both spinous
and soft sections. Caudal fin blue with yellow or orange macu-
lae. Anal fin blue with narrow orange distal margin and one to
three series of pale yellow eggspots, third row (on distal fin mar-
gin) apparently only in largest males (Figure 8D). Pelvic fins
white to pale blue with white leading edge. Pectorals hyaline.
A female shown (as Otopharynx “productus”) by Turner
(1996:171) appears grayish tan on dorsum, silver on sides of head,
silvery with bluish tinge on flanks, white on belly and underside
of head. Three lateral spots visible but indistinct. Dorsal fin shows
orangeish oval maculae between all spines and soft rays. Anal fin
hyaline with faint orange flush distally. Caudal fin plain yellow-
ish. Pelvics white flushed with orange. Pectorals hyaline.
Coloration in preservative (Figure 7A–D). Head and body light
silvery brown, slightly darker on dorsum and upper head sur-
faces. No stripes or bars on head. Lips yellowish. A vague oper-
cular spot in some specimens. Belly pale yellowish silver. Lateral
spots faint, brownish; suprapectoral spot generally more distinct
than supraanal and precaudal spots. No dorsal midline spots.
Nape with indistinct brown spot above operculum. Dorsal fin
yellowish, marbled with gray posteriorly. Anal fin yellowish with
indistinct darker margin. Caudal fin yellow with faint maculae.
Pelvics yellow, shading to brownish distally. Pectorals yellow.
Maximum size. At least 20 cm TL (see Remarks).
Distribution.
In addition to the type locality (Figure 1) in 42 m depth off Maz-
inzi Bay in the southeast arm, O. styrax has also been recorded
(as Otopharynx “productus”) in the southeast arm from trawl
stations Nkope and Chilinda in 18–20 m, and from Ulande
[Ulandi], Chekopa, near Boadzulu Island, and Chapola Shoal
in 5–50 m (Turner 1996:179). It was also trawled from unspec-
ified areas of the southwest arm in 10 m (Duponchelle et al.
2000:127) and, as Otopharynx sp. “productus sharp snout,” was
recorded from Senga Bay at an unspecified depth (Snoeks and
Hanssens 2004:292 and fig. 75). Konings (1995) reported this
species (as Mylochromis sp. “torpedo blue”) from the east shore
between Ntekete and Narungu, Malaŵi. Konings more recently
(2016:341) reports it (as Otopharynx sp. “torpedo blue”) also
from Chiloelo, Mdoka, and Nkhudzi, Malaŵi. It was exported to
the aquarium trade on one occasion from near Manda, Tanza-
nia (Figure 8D). Although most specimens mentioned in the
literature had been trawled, it is also captured near sandy shores
in depths of less than 8 m (Figure 8A–D).
Etymology.
From the Greek masculine noun ␴␶ύ␳␣␰ (styrax), a metal spike
at the lower end of a spear shaft, for the resemblance of this elon-
gate cichlid with pointed head to the shape of that classical
object. A noun in apposition.
Remarks.
Other species recovered in the same trawl hauls as the type spec-
imens included the cichlids Champsochromis spilorhynchus,
Hemitaeniochromis urotaenia, Mylochromis spilostichus, M. sub-
ocularis, Nimbochromis polystigma, Otopharynx tetraspilus,Pro-
tomelas cf. pleurotaenia, Trematocranus microstoma, and T.
placodon; noncichlids caught included Bagrus meridionalis,
unidentified Clariidae, and unspecified others.
The diet is unknown; gut contents were not examined.
Snoeks and Hanssens (2004) considered Otopharynx“pro-
ductus” (Turner 1996) to represent a different species than their
own O. sp. “productus sharp snout” solely because they found 13
lower gill rakers in their sample, whereas Turner (1996:292)
reported 11 gill rakers. None of these authors stated the number
of specimens examined for the count. In the type series of O.
styrax, from two trawl hauls made at the same site and depth in
rapid succession, the gill raker counts range from 11 to 13, with
12 the modal number. Judging by the photographs of the respec-
tive specimens, both provisional names represent the same dis-
tinctive species, now described as O. styrax. Turner (1996) stated
that the (oral) teeth are in five rows, but in the types I can find
only three to four rows (outer row plus two or three inner series).
Konings (2016) reported that the maximum total length of
this species (as O. sp. “torpedo blue”) is ~12 cm, but this is incor-
rect. The largest paratype measures 145 mm TL. The holotype
and all paratypes are either immature or sexually quiescent. The
specimen illustrated next to a ruler by Snoeks and Hanssens
(2004:293, fig. 75) is close to 20 cm TL. A living specimen cap-
tured near Manda, Tanzania (Figure 8D) was 20–22 cm TL
(Mark Smith, pers. comm.). The largest specimen reported (as
Otopharynx “productus”) by Duponchelle et al. (2000) was 134
mm SL; as some ripe females no larger than this were caught, the
latter authors provide preliminary estimates of length–weight
and fecundity–weight relationships.
This species was considered uncommon by Turner (1996)
and very rare by Konings (1995). Duponchelle et al. (2000) con-
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
178

Six New Species of the Cichlid Genus Otopharynx • Oliver 179
FIGURE 8. Otopharynx styrax, living individuals from various locations around Lake Malaŵi, all from inshore
waters over sandy substrate in depths of less than 8 m. Specimens not preserved. A. Msinje (Masinje), Malaŵi
(east shore about 3 km south of Mozambique border; right side reversed). B. Kambiri Point, Malaŵi (west shore
about 12 km north of Maleri Islands). C. Kambiri Point, Malaŵi (right side reversed). D. Vicinity of Manda, Tan-
zania (northeast shore); specimen 20–22 cm TL collected by Charles Kacirek during 1990s; photographed in
aquarium within one month of capture. All photographs by Mark Smith.

sidered it to be a rare species, captured in the southwest arm at
a rate of 0.7 kg/20-min haul in 10 m depth and not captured
below 30 m. The type series of O. styrax was, however, trawled
in 42 m in the southeast arm.
Otopharynx aletes
new species
Zoobank.org/urn:lsid:zoobank.org:act:
F48A0D30-2D2D-4A2E-8FB7-DAAD983A57EA
Figures 2D, 9, Tables 7, 8
Otopharynx sp. “molariform striped”: Snoeks and Hanssens
(2004:296).
Holotype.
YPM ICH 031030, 94.4 mm SL, male; Malaŵi: Central region,
Lake Malaŵi, bottom trawl off Bua Point, Lake Malaŵi Trawl-
ing Survey station Bua III, M/L Ethelwynn Trewavas, depth >55
m, at about 12°45⬘21⬙S 34°17⬘07⬙E–12°44⬘24⬙S 34°16⬘34⬙E; 14
November 1972.
Paratypes.
YPM ICH 007820, seven (four male, three female), 85.9–106.9
mm SL; collected with the holotype.
Diagnosis.
A smallish (<14 cm TL) laterally spotted haplochromine distin-
guished by the following characters in combination: suprapec-
toral spot mostly below upper lateral line but extending above it
by about one-half to one scale; 10 or 11 gill rakers on lower outer
arch; and lower pharyngeal bone with posteromedian teeth
molariform with stout shafts and rounded crowns. The place-
ment of the suprapectoral spot overlapping the upper lateral line
distinguishes O. aletesfrom O. brooksi, O. lithobates, and O. ova-
tus, all of which have the suprapectoral spot entirely below and
usually separated from the upper lateral line. Otopharynx aletes
has more scales in the lateral line (34–36) than O. brooksi
(31–33), O. spelaeotes (31–33), O. antron (29–31), O. tetraspilus
(30–32), and O. tetrastigma (30–33), but fewer thanO. decorus,
which has 36–38 and which also has 13–14 soft dorsal rays vs.
11–12 in O. aletes. With 10 or 11 gill rakers on the lower outer
arch, O. aletes is readily separated from O. auromarginatus
(14–18). Its longer caudal peduncle (1.6–2.0 times as long as
deep) and shallowly emarginate caudal fin distinguish O. aletes
from O. selenurus (caudal peduncle 1.0–1.5 times as long as
deep, caudal fin crescentically emarginate). The bicuspid outer
jaw teeth, molariform pharyngeal teeth, and seven to eight sub-
dorsal bars differentiate O. aletes from O. speciosus (outer jaw
teeth unicuspid; pharyngeal teeth all small, bicuspid; about four
subdorsal bars). The presence of seven to eight vertical bars
below dorsal-fin base separates O. aletes from O. argyrosoma,
which lacks vertical bars. The unthickened, nonlobate lips dif-
ferentiate O. aletes from O. pachycheilus (lips thickened, medially
lobate). The 16 dorsal-fin spines, absence of distinct equidistant
dorsal midline spots, and offshore habitat on sand or soft bottom
distinguish O. aletesfrom O. heterodon,which has 17–18 dorsal-
fin spines, about five distinct dorsal midline spots, and inhabits
rocky shores. Trematocranus brevirostris somewhat resembles
O. aletes, but in O. aletesthe suprapectoral spot is largely below
the upper lateral line but extends above it by about one-half to
one scale (vs. placed more above than below the upper lateral
line, at least in the lectotype, see Eccles and Trewavas 1989: fig.
76), the cephalic lateral-line system is not enlarged (vs. pores
and canals inflated), the dorsal fin has 11–12 (vs. 9) segmented
rays, there are 54–67 (vs. ~43) outer upper-jaw teeth, the pos-
teromedian teeth on the lower pharyngeal bone are molariform
with stout shafts and rounded crowns (vs. slightly enlarged but
cuspidate), and the caudal peduncle length/depth = 1.6–2.0 (vs.
1.2–1.4).
Otopharynx aletes and the following two new species are
superficially similar but, at least in the available samples, can be
distinguished as follows. Compared to O. panniculus, O. aletes
has more scales in the lateral line (34–36, vs. 31–33) and a heav-
ier lower pharyngeal jaw bone and dentition (compare Figs.
9C–F and 10D–G), including enlarged, molarized teeth pos-
teromedially (vs. small, laterally compressed, bicuspid teeth),
posterior horns thickened (vs. slender), and anterior blade
shorter, deeper (vs. longer, more shallow). Compared to O. peri-
dodeka, O. aletes has the hemijaws in ventral view divergent cau-
dally and distant from each other (vs. approximated, convergent;
compare Figs. 9G and 13C), and a heavier lower pharyngeal
bone, with horns thicker, posteromedian teeth enlarged, and
molarized with nearly hemispherical crowns (vs. more lightly
built, horns narrower, posteromedian teeth somewhat enlarged
but more laterally compressed, cuspidate; compare Figs. 9C–F
and 13D–G).
Description.
Morphometric and meristic data are given in Tables 7 and 8.
Deep bodied; body depth 32.1–36.9% SL (Figure9A). Dor-
sal profile straight or slightly convex from snout tip to above cen-
ter or rear of orbit (Figure 9B), about evenly rounded from there
to end of dorsal-fin base, more convex than ventral profile. (Head
profile and gape inclination are rather variable within the type
series [compare Figure 9B, H, I]; however, counts, oral dentition,
lower pharyngeal bone shape, and its distinctive dentition, and
overall habitus support the inclusion of all specimens of the type
series in a single species.) Premaxillary pedicels slightly promi-
nent, their angle 42–53°, interorbital angle 34–47°, nuchal angle
16–25°. Ventral profile slightly convex from lower-jaw tip to pos-
terior margin of branchiostegal membrane; hinge of lower jaw
often protruding. Jaws equal anteriorly or lower projecting very
slightly, lower jaw not dorsoventrally flattened; lips neither thick-
ened nor lobate. Gape inclination 22–38°. Lower-jaw length–
width ratio intermediate, the hemijaws diverging posteriorly (Fig-
ure 9G). Lower-jaw underside angle 27–40°. Snout acuteness
73–85°. Eyes large, nearly round (orbit length 34.1–37.4% HL,
slightly longer than tall); pupil rounded both anteriorly and pos-
teriorly; eye not reaching dorsal head profile. Caudal peduncle
17.6–20.8% SL, its length 1.6–2.0 times its depth.
Dorsal fin and, especially, anal fin with pungent spines,
both fins with soft portion acute posteriorly. Caudal fin emar-
ginate, lobes equal in length. Pectoral fin long, 32.3–39.3% SL,
reaching level of base of third anal-fin spine or even that of third
anal soft ray. Pelvic fins pointed.
Dental arcade of each jaw rounded. Upper jaw with 54–69
teeth (total) in outer row. Outer upper-jaw teeth small, closely
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
180

Six New Species of the Cichlid Genus Otopharynx • Oliver 181
FIGURE 9. Otopharynx aletes. A. Holotype, YPM ICH 031030, 94.4 mm SL, male. B. Head of holotype. C–F. lower
pharyngeal bone of holotype, in posterior (C), left lateral (D), dorsal (E), and ventral (F) views. G. Underside of
head of a paratype (YPM ICH 007820, 101.4 mm SL, female). H, I. Two male paratypes (YPM ICH 007820;
106.9 (H) and 105.1 (I) mm SL), showing variation in snout profile and mouth angle.

spaced, with crowns slightly recurved; anterior 11–15 on each
side unequally bicuspid, implanted with shafts angled slightly
outward; an occasional unequally tricuspid tooth among the
bicuspids; posterolateral and posterior teeth unicuspid; about
six most posterior unicuspids larger than midlateral unicuspids.
Lower jaw rather shallow but not flattened. Lower-jaw dentition
of haplochromis type, the outer row with 44–65 teeth (total).
Inner teeth of both jaws in two or (modally) three rows; unicus-
pid, crowns recurved slightly.
Lower pharyngeal bone dissected from holotype (Figure
9C–F) and four paratypes (85.9–106.9 mm SL); subtriangular,
rather heavily built; posterior contour variably concave medi-
ally a slight to moderate amount, with short robust horns that
widen distally (Figure 9E). Median suture varying from straight
or slightly sinuous to meandering posteriorly with several short
interdigitations (Figure 9F). Keel slightly descending, deep, but
with length greater than depth; strongly convex below (Figure
9D). Bone robust in posterior view (Figure 9C). Lateral edges of
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
182
TABLE 7. Morphometric characters of Otopharynx aletes. Abbreviations: LLL, lower lateral line; S.D., standard
deviation.
Holotype Paratypes
YPM ICH
031030 NMean S.D. Range
Standard length (mm) 94.4 7 98.3 7.1 85.9–106.9
Head length (mm) 30.7 7 30.9 2.5 26.9–34.1
Percentage of standard length:
Head length 32.5 7 31.3 0.7 30.2–32.4
Body depth 36.9 7 32.8 0.5 32.1–33.4
Dorsal-fin base length 55.0 7 54.1 1.3 53.0–55.6
Predorsal length 37.2 7 35.3 1.0 33.6–37.1
Prepectoral length 31.6 7 31.4 0.7 30.4–32.2
Prepelvic length 36.4 7 36.9 0.9 35.7–38.1
Preanal length 62.7 7 64.0 0.7 63.2–65.4
Pelvic fin origin to anal fin origin 26.7 7 28.1 0.9 26.4–29.1
Anal-fin base length 19.0 7 18.2 0.3 17.8–18.7
Caudal peduncle length 19.8 7 19.8 1.1 17.6–20.8
Caudal peduncle depth 11.7 7 10.7 0.3 10.1–11.2
Pectoral-fin length 39.0 7 36.0 2.8 32.3–39.3
Pelvic-fin length 28.5 7 26.3 3.1 22.6–30.0
Dorsal-fin origin to anal-fin origin 48.6 7 46.8 1.3 45.7–49.3
Dorsal-fin origin to end of anal-fin base 58.3 7 56.9 1.4 55.0–58.7
Pelvic-fin origin to end of dorsal-fin base 51.7 7 52.1 0.8 50.8–53.4
End dorsal-fin base to end anal-fin base 15.3 7 15.0 0.6 14.1–15.7
Anal-fin origin to end of dorsal-fin base 30.6 7 29.1 0.2 28.9–29.4
Dorsal-fin origin to pelvic-fin origin 36.0 7 32.5 0.4 32.0–33.0
End dorsal-fin base to end hypurals at LLL 19.9 7 19.3 0.6 18.6–20.3
End anal-fin base to end hypurals at LLL 21.8 7 21.9 0.5 20.8–22.5
Percentage of head length:
Head width 43.3 7 44.6 0.9 43.5–45.8
Interorbital width 22.1 7 21.5 0.8 20.4–22.8
Snout length 35.1 7 32.2 0.8 30.9–33.3
Snout width 32.0 7 29.9 1.0 28.5–31.1
Lower-jaw length 38.0 7 36.7 1.1 35.2–38.1
Lower-jaw width 26.1 7 22.5 3.2 18.5–26.9
Premaxillary pedicel length 30.9 7 29.3 0.9 28.1–31.0
Upper-jaw length 31.4 7 27.2 0.9 26.2–28.2
Cheek depth 22.0 7 19.3 1.2 17.1–21.2
Orbit length 34.1 7 36.0 1.2 34.3–37.4
Vertical eye diameter 30.9 7 32.7 1.2 31.4–34.0
Lacrimal (preorbital) depth 22.3 7 21.0 0.6 20.0–21.8
Postorbital head length 38.6 7 37.4 1.7 35.6–39.6

dentigerous surface with a row of regularly and closely spaced,
small, acutely cuspidate teeth. Posterior half of median dentiger-
ous area with a roughly circular patch of enlarged, molariform
teeth with domelike crowns, tooth size and degree of molariza-
tion decreasing beyond edges of circular area (Figure 9E). Teeth
in anterior half of 2–4 median columns bicuspid and smaller
than posterior molars, but clearly larger than lateral teeth. Pos-
terolateral areas with teeth small, closely spaced, laterally com-
pressed, bicuspid. Teeth in posterior row 33–41 (holotype: 33);
in each median column 7–11 (holotype: 7–8); in each oblique
row 5–7 (holotype: 6).
Lacrimal bone (Figure 2D) bearing four neuromasts and
five lateral-line pores; lacrimal notch distinct.
Gill rakers 10–11 on lower limb of outer arch; unbranched,
subequal in length; some specimens lacking any melanophores,
others having several melanophores on each lower-limb raker.
Scales ctenoid; 34–36 in lateral line. Lateral line discontin-
uous; upper section with downward kink one to two scales long,
or kink lacking. Squamation extending onto caudal fin between
fin rays, to near tips of upper and lower lobes and on basal one-
fourth to two-fifths along middle rays. Soft dorsal and anal fins
with a few small scales between bases of some rays. Size transi-
tion gradual from larger abdominal to smaller thoracic scales.
Coloration in life. Not recorded.
Coloration in preservative (Figure 9A–C). Head and flanks
medium brown, dorsal head surface and nuchal region a little
darker; no silvery or whitish areas on chest or belly (Figure 9A).
An opercular spot; an indefinite darker area between eye and
rear of upper jaw, broadest along orbital margin; no distinct head
stripes (Figure 9B). Branchiostegal membrane dark brown in
males. Seven or eight narrow dark vertical bars below dorsal-fin
base. Three lateral spots present but usually faint (most distinct
in holotype), suprapectoral spot larger and usually darker than
supraanal and precaudal spots. Suprapectoral spot trapezoidal
with anterior edge sloping up and forward, this spot overlap-
ping subdorsal bars 3 and 4, covering six to seven consecutive
upper lateral-line scales, and extending vertically from just above
lateral midline to one-half scale above upper lateral-line canals.
Supraanal spot on last subdorsal bar between upper and lower
lateral lines, when visible a little wider and darker than the bar.
Precaudal spot at end of caudal peduncle. No dorsal midline
spots. Dorsal fin with pale lappets, remainder of fin brownish
with thinly scattered melanophores; vague maculae in soft part.
Anal fin brownish, paler basally, without evident eggspots in
available specimens. Caudal fin brown, darkest on upper and
Six New Species of the Cichlid Genus Otopharynx • Oliver 183
TABLE 8. Meristic characters of Otopharynx aletes. Abbreviations: Q, quartile.
Paratypes
Holotype NMedian 1st Q3
rd Q Range
Squamation:
Lateral-line scales 34 7 35 35 36 34–36
Upper lateral-line scales 25 6 26 24.75 28 24–28
Lower lateral-line scales 14 4 14 13 16.5 13–17
Lateral-line scales on caudal fin 162 12 1–2
Kink length 140.5 0 1.75 0–2
Upper transverse line scales 675 55 5–5
Lower transverse line scales 10 5 10 10 10 10–10
Cheek scale rows 373 33 2–4
Scales between pectoral- and pelvic-fin bases 859 89.5 8–10
Fins:
Dorsal-fin spines 16 7 16 16 16 16–16
Dorsal-fin segmented rays 11 7 12 12 12 11–12
Dorsal-fin total elements 27 7 28 28 28 27–28
Anal-fin spines 373 33 3–3
Anal-fin segmented rays 979 89 8–9
Anal-fin total elements 12 7 12 11 12 11–12
Pectoral-fin rays 15 7 15 14 15 14–15
Gill rakers:
Epibranchial 474 44 4–4
Angle 171 11 1–1
Lower limb 10 7 11 11 11 10–11
Total 16 7 16 16 16 15–16
Oral teeth:
Outer upper-jaw teeth (total both sides) 69 7 60 58 65 54–67
Outer lower-jaw teeth (total both sides) 49 7 52 45 61 44–65

lower leading edges; no obvious markings. Pelvics brown on
leading half of fin, unmarked on remainder. Pectorals without
melanophores.
Maximum size. The largest paratype measures 135.0 mm TL.
Distribution.
Snoeks and Hanssens (2004:296) reported what seems to be this
species (as Otopharynx sp. “molariform striped”) from Senga
Bay (depth not provided). Otherwise, it is known only from the
type locality (Figure 1), off Bua Point in at least 55 m depth.
These two localities are about 115 km apart. Current status of the
species requires verification.
Etymology.
From the Greek masculine nominative singular noun ␭έ␶␩
(alétes), a grinder. The name alludes to the well-defined group
of conspicuously enlarged molariform teeth on the lower pha-
ryngeal bone. A noun in apposition.
Remarks.
The type series was part of the catch from a half-hour bottom
trawl. As the net was brought on deck, the fish dumped into
boxes and sorted, most specimens received damage to the scales
on the flanks, caudal peduncle, or both. As a result, accurate
counts of certain squamation characters could not be obtained
from some specimens, although application of Cyanine Blue dye
to emphasize the scale pockets permitted some scale counts not
otherwise possible.
The diet of the species is unknown; gut contents of the type
specimens were not investigated.
Three paratypes are female. Their ovaries contain imma-
ture ova approximately ¼ mm in diameter. Several males includ-
ing the holotype appear to be adult but nonbreeding.
Otopharynx panniculus
new species
Zoobank.org/urn:lsid:zoobank.org:act:
DBA01FCE-E5BC-421D-A923-C491DEAB2F8E
Figures 2E, 10, 12, Tables 9, 10
Trematocranus brevirostris [not of Trewavas, 1935]: Turner
1996:173, 188 [species more recently called Otopharynx
“brevirostris yellow,” G. Turner, pers. comm. 13 June 2018].
Holotype.
YPM ICH 031027, 72.2 mm SL, male; Malaŵi: Lake Malaŵi,
southwest arm, bottom trawl off Kasankha (or Kasanga) Bay,
Lake Malaŵi Trawling Survey station Kasankha II, M/L Ethel-
wynn Trewavas, depth ~42 m; approx. 2-km haul at about
14°06⬘S 34°50⬘E; Malaŵi Fisheries Department staff, 7 March
1972.
Paratypes.
YPM ICH 014162, 9 (most apparently male, a few undeter-
mined), 66.2–72.5 mm SL; collected with the holotype.
Additional Nontype Material.
AMNH I-221760 SW, two (females; cleared and stained, used
for fin-ray and vertebral counts only), 66.0–69.0 mm SL;
Malaŵi: Lake Malaŵi, southeast arm, bottom trawl off Nkopi
Bay, Lake Malaŵi Trawling Survey station Nkopi III, M/L Ethel-
wynn Trewavas, depth ~55 m; approx. 2-km haul at about
14°09⬘00⬙S 35°06⬘30⬙E; Malaŵi Fisheries staff, 28 February
1972.
Diagnosis.
A small (<10 cm TL), laterally spotted haplochromine which is
recognized by the following combination of characters: promi-
nent, quadrate suprapectoral spot spanning subdorsal vertical
bars 3–4, this spot below and touching the upper lateral line
and usually extending slightly above it; 13–15 lower-limb gill
rakers; and deep body (depth 34.4–38.2% SL). The shape and
placement of the suprapectoral spot distinguish O. panniculus
from O. brooksi, O. lithobates, and O. ovatus, in all of which this
spot lies entirely below and usually separated from the upper
lateral line. Its shorter lateral line of 31–33 scales discriminates
O. panniculusfrom O. decorus(36–38), O. argyrosoma (34–36),
and O. selenurus(34–35). Its higher gill-raker count (13–15 on
lower limb) distinguishes O. panniculus from O. tetrastigma
(9–11), O. speciosus (11–12), and O. antron (10–11). Its shorter
head (31.9–33.7% SL) and longer caudal peduncle (18.8–21.7%
SL) distinguish O. panniculus from O. spelaeotes (head
34.8–36.6% SL; caudal peduncle length 13.2–15.6% SL). Its
15–16 dorsal-fin spines and lack of dorsal midline spots sepa-
rate O. panniculus from O. heterodon, which has 17–18 dorsal
spines and five or six distinct, small, dark dorsal midline spots
just below the dorsal-fin base. The presence of several slightly
enlarged teeth in the median columns of the lower pharyngeal
bone separates O. panniculus from O. auromarginatus and O.
tetraspilus, which have only small, compressed, bicuspid pha-
ryngeal teeth. In O. panniculus the lips are thin and lack median
lobes, distinguishing it from O. pachycheilus (lips hypertro-
phied, with prominent median lobes). Compared to O. aletes,
O. panniculus has fewer lateral-line scales (31–33 vs. 34–36)
and a lighter lower pharyngeal jaw and dentition (compare Figs.
10D–G and 9C–F), with small, laterally compressed, bicuspid
posteromedian pharyngeal teeth (vs. these teeth enlarged,
molarized), slender posterior horns (vs. horns thickened), and
anterior blade longer, shallower (vs. blade shorter, deeper).
Compared to O. peridodeka, O. panniculus has fewer lateral-
line scales (31–33 vs. 34–35), usually has more lower-limb gill
rakers (13–15, median 14.5, vs. 11–13, median 12), and lower
pharyngeal teeth more uniform in size (compare Figs. 10F and
13F), those of the median columns scarcely larger than the
more lateral teeth (vs. posterior teeth of median columns
distinctly enlarged relative to lateral teeth). Otopharynx pan-
niculus superficially resembles Trematocranus “brevirostris
deep” (Turner 1996), but that still-undescribed species is from
substantially deeper water (90–102 m vs. ~42 m), the suprapec-
toral spot apparently spans subdorsal bars 2–3 (vs. bars 3–4 in
O. panniculus) and there are 10–13 lower-limb gill rakers (vs.
13–15) (see further comments on T. “brevirostris deep” in
Remarks under Otopharynx peridodeka, below). Trematocranus
brevirostris itself (see Figure 11) is also similar, but in O. pan-
niculus the suprapectoral spot is roughly square, covers four to
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
184

five longitudinal scales, and is located almost entirely below the
upper lateral line (vs. taller than long, covers three scales, and
at least in the lectotype is placed more above than below the
upper lateral line, see Eccles and Trewavas 1989: fig. 76), the
cephalic lateral-line system is not enlarged (vs. pores and canals
of the preorbital, nasal, dentary, and lacrimal bones inflated),
there are 52–66 outer upper-jaw teeth (vs. ~43), and 3–4 cheek
scale rows (vs. 2).
Description.
Morphometric and meristic data are given in Tables 9 and 10.
Deep-bodied, body depth 34.4–38.2% SL (Figure 10A).
Dorsal profile straight above snout, then evenly convex to end of
dorsal-fin base. Premaxillary pedicels not prominent, their angle
42–50°, interorbital angle 36–46°, nuchal angle 20–28°. Ventral
profile a less-convex mirror image of dorsal profile. Jaws equal
Six New Species of the Cichlid Genus Otopharynx • Oliver 185
TABLE 9. Morphometric characters of Otopharynx panniculus. Abbreviations: LLL, lower lateral line; S.D., stan-
dard deviation.
Paratypes
Holotype NMean S.D. Range
Standard length (mm) 72.2 9 69.6 2.1 66.2–72.5
Head length (mm) 24.3 9 22.7 0.7 21.6–23.7
Percentage of standard length:
Head length 33.6 9 32.6 0.6 31.9–33.7
Body depth 38.2 9 35.4 1.0 34.4–37.6
Dorsal-fin base length 54.9 9 54.2 0.8 52.9–55.6
Predorsal length 37.6 9 36.6 1.0 35.6–38.6
Prepectoral length 32.5 9 32.4 0.7 31.3–34.0
Prepelvic length 39.1 9 38.1 0.5 37.4–38.9
Preanal length 66.9 9 65.4 0.7 63.9–66.4
Pelvic fin origin to anal fin origin 27.9 9 28.1 0.5 27.7–29.0
Anal-fin base length 17.5 9 17.3 0.8 16.0–18.6
Caudal peduncle length 19.1 9 20.4 1.0 18.8–21.7
Caudal peduncle depth 11.4 9 11.1 0.2 10.9–11.5
Pectoral-fin length 38.7 8 40.5 1.7 38.0–42.3
Pelvic-fin length 29.3 9 27.7 2.4 23.1–31.7
Dorsal-fin origin to anal-fin origin 50.1 9 48.7 0.8 47.5–50.1
Dorsal-fin origin to end of anal-fin base 58.1 9 57.5 1.1 55.2–59.2
Pelvic-fin origin to end of dorsal-fin base 52.8 9 52.3 0.9 51.4–53.6
End dorsal-fin base to end anal-fin base 16.3 9 16.1 0.8 14.9–17.6
Anal-fin origin to end of dorsal-fin base 30.4 9 29.7 0.8 28.7–30.9
Dorsal-fin origin to pelvic-fin origin 36.5 9 34.8 0.9 33.8–37.0
End dorsal-fin base to end hypurals at LLL 19.6 9 19.0 0.6 17.9–19.9
End anal-fin base to end hypurals at LLL 21.5 9 22.3 0.8 20.7–23.1
Percentage of head length:
Head width 43.1 9 43.5 0.6 42.6–44.5
Interorbital width 19.5 9 20.6 0.8 19.6–22.2
Snout length 29.5 9 29.1 0.9 28.1–30.4
Snout width 27.5 9 27.7 1.3 26.0–29.3
Lower-jaw length 39.3 9 38.5 1.2 36.7–40.4
Lower-jaw width 16.6 9 17.4 1.4 15.2–19.3
Premaxillary pedicel length 30.7 9 28.8 0.9 27.2–30.2
Upper-jaw length 28.3 9 28.0 0.8 27.0–29.4
Cheek depth 19.9 9 19.5 1.2 17.6–21.0
Orbit length 36.8 9 37.6 0.8 36.5–38.8
Vertical eye diameter 33.5 9 34.2 1.2 32.3–36.5
Lacrimal (preorbital) depth 19.5 9 18.8 0.6 18.1–19.9
Postorbital head length 37.9 9 37.8 0.7 36.7–39.0

anteriorly or lower projecting very slightly, lower jaw dorsoven-
trally somewhat flattened; lips thin and not lobate. Gape inclina-
tion 30–42°. Lower-jaw length–width ratio narrow, the
hemijaws, as seen from below, converging toward the rear and
nearly touching posteriorly (Figure 10C). Lower-jaw underside
angle 27–40°. Snout acuteness 77–80°. Eyes large (36.5–38.8%
HL), round; pupil pointed anteriorly, rounded posteriorly; eye
nearly reaching dorsal head profile. Caudal peduncle
18.8–21.7% SL, its length 1.7–2.0 times its depth.
Soft dorsal and anal fins produced, more so in males. Cau-
dal fin emarginate. Pectoral fin 38.0–42.3% SL, reaching beyond
level of third anal-fin spine base.Pelvic fin 23.1–31.7% SL.
Dental arcade of each jaw rounded. Upper jaw with 52–66
teeth (total) in outer row; movably implanted, rather closely
spaced. Anterior and anterolateral teeth unequally bicuspid with
acute cusps, crowns slightly incurved; occasional unequally tri-
cuspid teeth present; posterior teeth unicuspid with acute
crowns. Lower-jaw outer tooth row of haplochromis type, with
40–62 teeth (total). Crowns similar to those of upper jaw. Lower
jaw somewhat flattened, dorsoventrally shallow, so that anterior
and anterolateral teeth are implanted with shafts angled out-
ward. Inner teeth small, unicuspid to weakly tricuspid, in two
rows spaced closely one behind the other.
Lower pharyngeal bone dissected from holotype (Figure
10D–G) and all nine paratypes; subtriangular (Figure 10F),
appearing lightly built in posterior view (Figure 10D); posterior
contour emarginate, each half slightly convex, the halves meeting
at an obtuse angle; horns slender, rather long (Figure 10F).
Median suture straight (Figure 10G). Keel long, shallow, straight,
its length about twice its depth; weakly convex below (Figure
10E). Dentigerous surface somewhat concave in lateral view. One
to four posterior teeth of each median column slightly enlarged
but cuspidate, not molarized (one paratype with no teeth
enlarged); lateral teeth small, bicuspid, rather crowded. Teeth in
posterior row 39–48 (holotype: 41), in each median column 9–13
(holotype: 11), in each oblique row 6–10 (holotype: 7).
Lacrimal bone (Figure 2E) bearing four neuromasts and
five lateral-line pores; lacrimal notch obsolete.
Gill rakers 13–15 (median 15) on lower arch; slender,
unbranched; melanophores variably present on outer surface of
outer-arch rakers.
Scales ctenoid; 31–33 (median 32) in lateral line. Lateral line
discontinuous, upper section with downward kink three to four
scales long, or (modally) lacking a kink. Squamation extending
onto caudal fin between fin rays, to near tips of upper and lower
lobes and on basal one-third of fin along middle rays. Soft dor-
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
186
TABLE 10. Meristic characters of Otopharynx panniculus. Fin-ray and tooth counts from two cleared and stained
specimens are included. Abbreviations: Q, quartile.
Paratypes
Holotype NMedian 1st Q3
rd Q Range
Squamation:
Lateral-line scales 33 7 32 32 32 31–33
Upper lateral-line scales 21 7 20 19 24 19–26
Lower lateral-line scales 15 6 14.5 13.5 15 12–15
Lateral-line scales on caudal fin 272 12 0–2
Kink length 070 03 0–4
Upper transverse line scales 575 45 4–5
Lower transverse line scales 989 89 8–9
Cheek scale rows 383 33 3–4
Scales between pectoral- and pelvic-fin bases 848 88.75 8–9
Fins:
Dorsal-fin spines 15 10 16 15.75 16 15–16
Dorsal-fin segmented rays 11 10 10 10 10.25 9–11
Dorsal-fin total elements 26 10 26 25.75 26 25–27
Anal-fin spines 3 10 3333–3
Anal-fin segmented rays 898 89 8–9
Anal-fin total elements 11 9 11 11 12 11–12
Pectoral-fin rays 14 9 14 14 14 14–14
Gill rakers:
Epibranchial 4 10 555.25 4–6
Angle 1 10 1111–1
Lower limb 13 10 14.5 14 15 13–15
Total 18 10 20.5 20 21.25 18–22
Oral teeth:
Outer upper-jaw teeth (total both sides) 63 10 61 55 64.25 52–66
Outer lower-jaw teeth (total both sides) —750 42 57 40–62

Six New Species of the Cichlid Genus Otopharynx • Oliver 187
FIGURE 10. Otopharynx panniculus. A. Holotype, YPM ICH 031027, 72.2 mm SL, male. B. Head of holotype.
C. Underside of head of holotype. D–G. Lower pharyngeal bone of a paratype (YPM ICH 014162, 68.9 mm SL,
not sexed), in posterior (D), left lateral (E), dorsal (F), and ventral (G) views.

sal and anal fins with a few small scales between bases of some
rays. Larger scales of lower flank transition gradually to smaller
chest and belly scales between bases of pectoral and pelvic fins.
Vertebrae (counted in two cleared and stained specimens,
see Additional material, above) 13 + 19 = 32, 14 + 18 = 32.
Coloration in life. Live coloration of males and females is
unknown. However, a freshly trawled male and female of
Otopharynx “brevirostris yellow,,” thought to be O. panniculus,
are shown in Figure 12. The male (Figure 12A) is silvery on
flanks, white on chest, with about six gray vertical bars below
dorsal-fin base. Lower half of head bright orange-yellow.
Suprapectoral spot and faint supraanal and precaudal spots steel-
blue, shaped and placed as in holotype. Dorsal fin lappets orange
distally, whitish proximally; orange maculae between soft rays.
Caudal fin brownish with orange maculae. Anal fin pale with
three or four large yellow eggspots. Pelvic fin apparently with
black leading edge. Pectorals hyaline. The presumptive female
(Figure 12B) is predominantly silvery to white on body, chest,
and head. Three distinct lateral spots; vertical bars faint. Oval
orange markings evident throughout dorsal and caudal fins.
Coloration in preservative (Figure10A–C). Upper and lateral head
surfaces light brown, nuchal area darker; upper half of lacrimal
dark brown; operculum silvery with faint brown posterodor-
sal spot (Figure 10B). Underside of head, including branchioste-
gal membrane and geniohyoid area between left and right
hemijaws, dark brown in males (Figure 10C). Flanks light
brown above midline, silvery below; no distinct dorsal midline
spots; chest peppered with dark pigment in males (Figure10C).
Eight (mode) or nine dark vertical bars below dorsal-fin base,
narrower than or equal to the interspaces; two to three fainter
bars on caudal peduncle. Suprapectoral spot square to parallel-
ogram-shaped, darker brown than bars, extending longitudi-
nally between subdorsal bars 3 and 4, covering four (mode) to
nearly five lateral-line scales, and vertically from just above lat-
eral midline of flank to one-half scale above upper lateral line.
Supraanal spot small, variably present, appearing as a darken-
ing on last subdorsal bar between lateral lines. Precaudal spot
on lower lateral line at end of caudal peduncle, slightly more
distinct than supraanal spot. Dorsal fin brownish with pale lap-
pets; no submarginal stripe; indistinct marbling in soft dorsal.
Anal fin brownish, shading in males to black on distal half; no
eggspots visible. Caudal brownish without distinct markings.
Pelvics of males blackish with narrow pale leading edge. Pec-
torals pale yellow, transparent.
Maximum size. The largest known specimen is a paratype 92.8
mm TL. A female (AMNH I-221760 SW) as small as 69 mm SL
had contained ripe ovarian ova before I cleared and stained it.
Distribution.
The type locality, in ~42 m off Kasankha Bay, is in the south-
west arm off its eastern shore (Figure 1). The species has also
been trawled in ~55 m off Nkopi (Nkope) Bay, Malaŵi, in the
southeast arm off its western shore. Current population status is
unknown.
Etymology.
From the Latin noun panniculus, a small cloth patch, with ref-
erence to the appearance of the quadrate suprapectoral spot
neatly “stitched” between two vertical bars. A noun in
apposition.
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
188
FIGURE 11. Possibly the true Trematocranus brevirostris Trewavas. This specimen (not examined) was seined in
2017 at the Palm Beach hotel located at the extreme south of the southeast arm. That locality resembles ecolog-
ically, and is near to, the type locality of T. brevirostris, “Bar House” (a previous hotel). Cuthbert Christy, who in
1925 made the collections studied by Trewavas (1935), described “Bar House and surrounding regions” as fol-
lows: “All shallow water. Coast consists of sandy bays and small inlets with almost everywhere a fringe, broken
here and there, of tall matete reeds. Near the outlet of the Shire R. bulrushes and papyrus take the place of the
tall reeds. Bottom almost everywhere sandy” (Christy 1925). Photograph by George Turner.

Remarks.
Specimen labels suggest that the Lake Malaŵi Trawling Survey
(LMTS) personnel referred to this species as Haplochromis sp. B
and Haplochromis “cf. heterodon” during the early 1970s. The
name Haplochromis sp. B has been applied to at least one other
species by LMTS personnel.
Multiple scales and outer oral teeth have been lost in the
type series, likely due to abrasion, as these small fish were brought
on deck, dumped from the trawl net into boxes, and sorted. Some
scale counts were made with the aid of Cyanine Blue dye.
Gut contents were not studied; the diet of this species
remains unknown.
Otopharynx peridodeka
new species
Zoobank.org/urn:lsid:zoobank.org:act:
CEB352C2-00C8-47E4-9863-5177BBE21900
Figures 2F, 13, 14, Tables 11, 12
? Trematocranus “brevirostris deep”: Turner (1996:173, 189).
Holotype.
YPM 031026, 92.2 mm SL, ripe male; Malaŵi: Lake Malaŵi, bot-
tom trawl off Sungu Point, Lake Malaŵi Trawling Survey sta-
Six New Species of the Cichlid Genus Otopharynx • Oliver 189
FIGURE 12. Otopharynx “brevirostris yellow,” freshly captured individuals apparently conspecific with O. pan-
niculus. A. Male trawled in 35–40 m depth northwest of Boadzulu Island, southeast arm, 29 July 1991. Exami-
nation showed 14 lower gill rakers, 31 scales in lateral line, and lower pharyngeal bone with papilliform teeth.
B. Female trawled in 19–26 m depth, center of southeast arm between Namiasi and Malindi, 19 July 1991. Pho-
tographs and data by George Turner.

tion Sungu III, M/L Ethelwynn Trewavas, depth 59 m; approx. 2-
km haul at about 13°30⬘26⬙S 34°31⬘44⬙E–13°31⬘05⬙S
34°32⬘36⬙E; Malaŵi Fisheries Department staff, 28 July 1971.
Paratypes.
YPM ICH 014120, 13 (six male, five female, two undetermined;
meristic data taken from all; two distorted specimens were
excluded from morphometrics), 62.7–92.1 mm SL; collected
with the holotype.
Diagnosis.
A small (<12 cm TL), laterally spotted haplochromine recog-
nizable by the following combination of characters: spots often
indistinct, the suprapectoral spot (when discernible) vaguely
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
190
TABLE 11. Morphometric characters of Otopharynx peridodeka. Abbreviations: LLL, lower lateral line; S.D., stan-
dard deviation.
Paratypes
Holotype NMean S.D. Range
Standard length (mm) 92.2 11 79.8 10.0 62.7–92.1
Head length (mm) 29.2 11 25.8 3.1 19.9–29.8
Percentage of standard length:
Head length 31.7 11 32.3 0.9 31.1–34.0
Body depth 33.5 11 33.0 1.4 30.7–35.0
Dorsal-fin base length 52.7 11 53.3 0.9 51.9–54.9
Predorsal length 35.4 11 36.7 1.3 34.4–38.6
Prepectoral length 31.5 11 32.4 0.6 31.6–33.9
Prepelvic length 38.6 11 38.0 0.7 36.7–39.1
Preanal length 64.5 11 64.9 0.6 63.4–65.6
Pelvic fin origin to anal fin origin 27.2 11 27.7 1.1 25.4–29.3
Anal-fin base length 17.7 11 17.9 0.8 16.3–19.0
Caudal peduncle length 20.0 11 19.7 0.8 18.2–21.0
Caudal peduncle depth 10.9 11 10.4 0.4 9.9–11.2
Pectoral-fin length 34.7 11 35.7 3.2 30.2–40.4
Pelvic-fin length 29.1 11 24.0 3.6 18.6–30.1
Dorsal-fin origin to anal-fin origin 46.8 11 46.5 1.1 44.8–48.1
Dorsal-fin origin to end of anal-fin base 56.3 11 56.2 0.9 55.2–57.8
Pelvic-fin origin to end of dorsal-fin base 50.7 11 50.8 0.8 49.7–52.6
End dorsal-fin base to end anal-fin base 16.1 11 14.7 0.6 13.4–15.7
Anal-fin origin to end of dorsal-fin base 28.8 11 28.3 0.7 27.2–29.4
Dorsal-fin origin to pelvic-fin origin 33.8 11 33.0 1.3 30.8–35.0
End dorsal-fin base to end hypurals at LLL 20.0 11 18.6 0.7 17.6–19.8
End anal-fin base to end hypurals at LLL 22.4 11 21.6 0.8 20.0–22.4
Percentage of head length:
Head width 43.4 11 43.3 1.0 42.0–44.8
Interorbital width 20.5 11 20.8 1.2 18.7–22.7
Snout length 32.5 11 31.4 1.2 29.3–33.4
Snout width 27.4 11 27.6 1.4 25.2–29.6
Lower-jaw length 36.7 11 35.9 1.2 33.9–38.0
Lower-jaw width 17.7 11 17.1 1.4 15.3–20.3
Premaxillary pedicel length 28.5 11 29.2 1.2 27.0–30.8
Upper-jaw length 28.9 11 27.1 1.1 25.9–28.8
Cheek depth 18.7 11 18.7 0.9 17.1–19.8
Orbit length 34.7 11 36.8 0.8 35.0–38.2
Vertical eye diameter 31.2 11 33.8 0.9 31.5–35.0
Lacrimal (preorbital) depth 21.8 11 21.6 1.7 19.9–24.7
Postorbital head length 38.9 11 36.9 1.2 34.7–39.1

rectangular, centered on subdorsal bar 3 or connecting bars 3
and 4, and covering four to five consecutive lateral-line scales,
extending vertically from just above lateral midline to half a scale
above upper lateral line; about seven vertical gray bars below
dorsal-fin base; 34–35 scales in lateral line; 11–13 (median 12)
gill rakers on lower limb; 48–73 teeth in outer row of upper jaw;
lower pharyngeal bone with a few posteromedian teeth enlarged
(submolariform to molariform); lacrimal bone lacking a notch
at base of lacrimal process (Figure 2F). Otopharynx peridodeka
is superficially most similar to O. panniculus, which also lacks a
lacrimal notch, but O. peridodeka has more scales in the lateral
line (34–35, vs. 31–33 in O. panniculus), usually fewer lower gill
rakers (11–13, median 12, vs. 13–15, median 14.5, in O. pan-
niculus), posteromedian pharyngeal teeth distinctly enlarged rel-
ative to more lateral teeth (vs. median and lateral teeth of nearly
uniform size), averages fewer teeth in the posterior row of the
lower pharyngeal bone (37–44, vs. 42–48 in O. panniculus), and
has a deeper lacrimal bone (19.9–24.7% HL, vs. 18.1–19.9% in
O. panniculus). Compared to O. aletes, O. peridodeka has the
hemijaws in ventral view posteriorly convergent, approximated
(vs. divergent, distant; compare Figs. 13C and 9G); and lower
pharyngeal bone more lightly built, with narrower posterior
horns, posteromedian teeth somewhat enlarged but laterally
compressed, cuspidate (vs. bone heavier, horns thickened, pos-
terior teeth of median rows enlarged, crowns molarized, nearly
hemispherical; compare Figs. 13D–G and 9C–F). Trematocranus
brevirostris (see Figure11) is somewhat similar, but in O. perido-
deka the suprapectoral spot is rectangular, covering four to five
longitudinal scales, and is located almost entirely below the
upper lateral line (vs. taller than long, covering three scales and,
at least in the lectotype, placed more above than below the upper
lateral line), the cephalic lateral-line system is not enlarged (vs.
pores and canals of the preorbital, nasal, dentary, and lacrimal
bones inflated), and there are 48–73 outer upper-jaw teeth
(vs. ~43).
Description.
Morphometric and meristic data are given in Tables 11 and 12.
Moderately deep-bodied, body depth 30.7–35.0% SL (Fig-
ure13A). Dorsal profile of snout straight or slightly convex, head
and nuchal profile nearly straight or variably convex, dorsum
convexly curved to end of dorsal-fin base. Premaxillary pedicels
slightly prominent in head profile, their angle 44–57°, interor-
bital angle 31–40°, nuchal angle 19–25°. Ventral profile almost
a mirror image of dorsal profile. Jaws equal anteriorly, lower jaw
Six New Species of the Cichlid Genus Otopharynx • Oliver 191
TABLE 12. Meristic characters of Otopharynx peridodeka. Abbreviations: Q, quartile.
Paratypes
Holotype NMedian 1st Q3
rd Q Range
Squamation:
Lateral-line scales 34 6 35 34.75 35 34–35
Upper lateral-line scales 27 7 26 24 28 24–29
Lower lateral-line scales 13+ 4 14.5 12.5 15 12–15
Lateral-line scales on caudal fin 152 0.5 2 0–2
Kink length 153 0 4.5 0–5
Upper transverse line scales 475 5 54–5
Lower transverse line scales 8710 8 10 8–10
Cheek scale rows 3 12 3332–3
Scales between pectoral- and pelvic-fin bases 879 9 98–9
Fins:
Dorsal-fin spines 17 12 16 16 16 16–17
Dorsal-fin segmented rays 10 13 11 11 12 10–13
Dorsal-fin total elements 27 12 27 27 28 26–28
Anal-fin spines 3 13 3333–3
Anal-fin segmented rays 8 13 9998–10
Anal-fin total elements 11 13 12 12 12 11–13
Pectoral-fin rays 14 13 15 14 15 14–15
Gill rakers:
Epibranchial 5 13 4454–5
Angle 1 13 1111–1
Lower limb 12 13 12 11 12 11–13
Total 18 13 17 16.5 18 16–19
Oral teeth:
Outer upper-jaw teeth (total both sides) 78 13 60 56.5 64.5 48–73
Outer lower-jaw teeth (total both sides) 60 11 58 54 59 41–63

slightly flattened dorsoventrally; lips thin and not lobate. Gape
inclination 26–36°. Lower-jaw length–width ratio narrow, the
hemijaws, as seen from below, converging toward the rear (Fig-
ure 13C). Lower-jaw underside angle 25–35°. Snout acuteness
71–84°. Eyes large (orbit length 34.7–38.2% HL), slightly oval
(vertical eye diameter 31.2–35.0% HL); pupil somewhat pointed
anteriorly, rounded posteriorly; eye nearly reaching dorsal head
profile. Caudal peduncle 18.2–21.0% SL, its length 1.7–2.1 times
its depth.
Soft dorsal and anal fins produced, more so in males. Cau-
dal fin emarginate. Pectoral fin 30.2–40.4% SL, reaching at least
to the level of first anal-fin spine base and even as far as the base
of second soft ray, the length apparently not correlated with sex,
as one of the specimens with the extreme pectoral extent is an
ovigerous female. Pelvic fin 18.6–30.1% SL, produced in males.
Dental arcade rounded in each jaw. Upper jaw with 48–73
teeth (total) in outer row; outer teeth closely spaced, larger near
symphysis. Anterior to lateral teeth unequally bicuspid with
long, acutely pointed major cusp, an occasional tricuspid tooth
intermixed; about 10–12 posterior teeth unicuspid. Outer lower-
jaw teeth similar in form to those of upper jaw; those in antero-
lateral area angled somewhat outward whereas anterior and
posterior teeth are more erectly implanted. Lower-jaw outer
tooth row of haplochromis type, with 41–63 teeth (total). Inner
teeth unequally tricuspid, in two to three rows.
Lower pharyngeal bone dissected from holotype (Figure
13D–G) and all 13 paratypes; subtriangular, somewhat lightly
built; posterior contour emarginate, the halves meeting at an
obtuse angle; horns moderately long and robust, widening dis-
tally (Figure 13F). Median suture straight to slightly sinuous (Fig-
ure 13G). Keel slightly descending, longer than deep, strongly
convex below (Figure13E). Bone of moderate depth in posterior
view (Figure 13D). Several posteromedian teeth variably enlarged
(Figure13F). In holotype, six to seven teeth on each side of mid-
line enlarged, a few with somewhat flattened crowns but only
about three teeth submolariform. In a paratype of 81.3 mm SL
about 11 posteromedian teeth enlarged to varying degree, four of
them molariform, lacking any distinct cusp and with crown
nearly flattened, other teeth less enlarged and cuspidate. In all
specimens, lateral pharyngeal teeth small, bicuspid. Teeth in pos-
terior row 37–44 (holotype: 42); in each median column 7–12
(holotype: 10–11); in each oblique row 5–8 (holotype: 6–8).
Lacrimal bone (Figure 2F) bearing four neuromasts and
five lateral-line pores; lacrimal notch obsolete.
Gill rakers 11–13 (median 12) on lower arch, rather short
(13 in only one paratype unilaterally, with lowermost two rak-
ers very short and closely spaced, 12 on other side); few
melanophores.
Scales ctenoid; 34 or (median) 35 in lateral line. Lateral line
discontinuous, upper section with downward kink three to five
scales long, or kink lacking. Squamation extending onto caudal
fin between fin rays, to near tips of upper and lower lobes and
on basal one-fourth along middle rays. Soft dorsal and anal fins
with two to three series of small scales between bases of some
rays. Larger scales of lower flanks transition gradually to smaller
chest and belly scales between pectoral- and pelvic-fin bases.
Coloration in life. Coloration of living individuals is unknown.
However, a fresh male and female identified as O. “brevirostris
deep” (Figure14) are thought to belong to this species. Both are
gray dorsally, silvery on flanks, white on belly, chest, and head
below eye; six gray bars below dorsal-fin base; indistinct gray
suprapectoral, supraanal, and precaudal spots; unpaired fins of
male yellowish, dorsal with black lappets and orange maculae
between spines and rays; pelvics yellow, those of male with dis-
tinct black leading edges; pectorals hyaline. Turner (1966:189)
provides a detailed description of the coloration of breeding
males of O. “brevirostris deep.”
Coloration in preservative(Figure 13A–C). The holotype (Fig-
ure 13A), a ripe male, is brown on dorsum and upper head
surfaces. Lower flanks silvery, grading to white ventrally on
belly and sides of chest. Operculum silvery with dark spot;
dark brown spot on upper part of lacrimal (Figure 13B).
Lower jaw, gular region, and branchiostegal membrane dark
brown (Figure 13C); ventral chest surface light brown. Seven
narrow gray vertical bars below dorsal-fin base; two bars on
caudal peduncle. Suprapectoral spot brown, darker than ver-
tical bars; supraanal and precaudal spots indistinct. No dor-
sal midline spots. Dorsal fin with whitish lappets; submarginal
fin area brown. Anal fin pale brown with no markings evi-
dent. Caudal fin brownish. Pelvics and pectorals hyaline.
Maximum size. The largest specimen known to me is a paratype
115.3 mm TL. Female paratypes as small as 65.1 mm SL have
ovaries full of mature orange ova. The smallest paratype (62.7
mm SL) is an immature female.
Distribution.
This species is known with certainty only from the type local-
ity (Figure 1) in 59 m depth off Sungu Point on the western
shore (a locality distinct from the better known Sungu Spit).
However, if, as suspected, it proves to be conspecific with
Trematocranus “brevirostris deep,” the latter is “[k]nown only
from deep water, 90–102 m depth, north of Monkey Bay”
(Turner 1996:189) which would extend the range some 60 km
to the southeast, and into considerably deeper water.
Etymology.
From the Greek adverb ␲ε␳␫ (peri: around, near) and noun
␦␷ώ␦ε␬␣ (duódeka: 12), referring to the usually 12 lower-limb
gill rakers of this species (a character separating it from the sim-
ilar-looking O. panniculus with a higher count). The compound
is treated as a noun in apposition.
Remarks.
The holotype is a ripe male with thick, sinuous testes.
All specimens in the type series are missing many scales
due to abrasion during dumping the catch and sorting it. Fur-
thermore, all had been slit for fishery research purposes from
the vent forward to a point above and in front of the right pelvic
fin, with the incision continued across the chest to in front of
the left pectoral fin. As a result, the chest with pelvic fins is
attached only by the skin and musculature of the left lower
abdominal wall. Therefore, all body depth measurements of this
species are approximate, as are measurements involving the
pelvic-fin origin.
The diet of O. peridodeka remains unknown; gut contents
were not investigated.
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
192

Six New Species of the Cichlid Genus Otopharynx • Oliver 193
FIGURE 13. Otopharynx peridodeka. A. Holotype, YPM ICH 031026, 92.2 mm SL, ripe male. B. Head of holotype.
C. Underside of head of holotype. D–G. Lower pharyngeal bone of holotype (keel broken during preparation),
in posterior (D), left lateral (E), dorsal (F), and ventral (G) views.

Acknowledgments
Gregory Watkins-Colwell (YPM) gave excellent
curatorial support throughout this study. Eric A.
Lazo-Wasem (YPM) hospitably provided access
to his photographic equipment and image-stack-
ing software. John Sparks, Melanie Stiassny, and
Thomas Vigliotta (AMNH) arranged a loan
including the specimens of Otopharynx pannicu-
lus I had cleared and stained some 35 years ago. I
am grateful to George F. Turner and an anony-
mous reviewer for providing suggestions for
improving this article. Mark Smith generously
allowed me to use his photographs of living
Otopharynx styrax specimens. Ad Konings like-
wise graciously permitted me to use his underwa-
ter photos of O. mumboensis. George Turner
newly scanned multiple photos of freshly trawled
specimens on short notice, with advice on their
identification, which I much appreciate. I thank
Lisa Boorman for pointing me toward some
online resources concerning illegal fishing at
Mumbo Island. Long ago (in 1971 and 1972), the
late P. H. Greenwood, E. Trewavas, and the staff
of the BMNH Fish Section generously hosted,
educated, and entertained me on three visits to the
museum totaling some seven months, visits
largely devoted to studying the type (and other)
specimens of the Lake Malaŵi “three-spots”
including the species now classified in Otophar-
ynx. Those visits yielded data still vital to my stud-
ies today. Research trips to Malaŵi and the
BMNH were supported by my late parents Ken-
neth A. and Madaline S. Oliver, and by Occiden-
tal College, Yale University, the former
Smithsonian Oceanographic Sorting Center under
Leslie W. Knapp, and NSF grant DEB 80-05538.
Received 5 February 2018; revised and accepted 22
June 2018.
Bulletin of the Peabody Museum of Natural History 59(2) • October 2018
194
FIGURE 14. Otopharynx “brevirostris deep,” freshly captured male (upper) and female (lower) apparently con-
specific with O. peridodeka. Trawled in 90 m depth off Monkey Bay, southeast arm, 21 May 1992. Photograph by
George Turner.

Appendix: Comparison Material
Abbreviations: KRM, K. R. McKaye; LMTS, Lake
Malaŵi Trawling Survey; MKO, M. K. Oliver; TDK,
T. D. Kocher. Note that Lake Nyassa, Lake Nyasa, and
Lake Malaŵi all refer to the same lake.
Copadichromis chrysonotus: BMNH 1908.10.27.49, 1
(lectotype); “Lake Nyassa”; E. L. Rhoades. — YPM
ICH 014471, 3; Malaŵi: Thumbi Island West; 18 Dec
1978.
Copadichromis quadrimaculatus: BMNH 1908.10.27.41,
1 (lectotype); “Lake Nyassa”; E. L. Rhoades. — YPM
ICH 009826, 6; Malaŵi: Nkhunguni Point; KRM, 19
Mar 1979.
Ctenopharynx intermedius: YPM ICH 007817, 6;
Malaŵi: station Namiasi I; LMTS.
Ctenopharynx nitidus: BMNH 1935.6.14.1763, 1 (lecto-
type); Lake Nyasa: Vua; C. Christy, 1925. — YPM
ICH 007807, 2; Malaŵi: Monkey Bay; D. H. Eccles,
12 Jul 1973.
Ctenopharynx pictus: BMNH 1935.6.14.1773, 1 (lecto-
type); Lake Nyasa: Vua; C. Christy, Aug 1925. — YPM
ICH 030223, 2; Malaŵi: Thumbi Island West; MKO,
KRM, TDK, 8 Aug 1980.
Exochochromis anagenys: USNM 304657, 1 (holotype);
Malaŵi: Thumbi Island West; MKO, KRM, TDK, 3–4
Aug 1980. — YPM ICH 023205, 1; captive.
Hemitilapia oxyrhyncha: BMNH 1906.9.7.38, 1 (lecto-
type); “Lake Nyassa”; J. E. S. Moore. — YPM ICH
007851, 5; Malaŵi: station Foo I; LMTS, 27 Sep 1972.
Naevochromis chrysogaster: BMNH 1935.6.14.1640, 1
(lectotype); Lake Nyasa: southwest arm; C. Christy,
1925. — YPM ICH 014344, 2; Malaŵi: 300 m S of
Otter Island; MKO, KRM, TDK, 10 Jul 1980. — YPM
14414, 1; Malaŵi: Thumbi Island West; MKO, KRM,
TDK, 18-19 Aug 1980. — YPM ICH 024323, 2;
Malaŵi: off W end Chembe; MKO, KRM, TDK, 6-7
Aug 1980.
Otopharynx argyrosoma: BMNH 1908.10.27.99, 1 (holo-
type); “Nyasa”; E. L. Rhoades.
Otopharynx auromarginatus: BMNH 1908.10.27.86, 1
(lectotype); “Lake Nyassa”; E. L. Rhoades. — BMNH
1908.10.27.87, 1 (paralectotype); “Lake Nyassa”; E.L.
Rhoades. — YPM ICH 030189, 1; Malaŵi: Nkhata
Bay; MKO, KRM, TDK, 25 Jul 1980.
Otopharynx brooksi: USNM 304655, 1 (holotype);
Malaŵi: Thumbi Island West; MKO, KRM, TDK,
18–19 Aug 1980. — USNM 304656, 5 (paratypes);
Malaŵi: Thumbi Island West; MKO, KRM, TDK,
18–19 Aug 1980. — USNM 314127, 1 (paratype);
Malaŵi: Thumbi Island West; MKO, KRM, TDK, 5–6
Aug 1980.
Otopharynx decorus: BMNH 1935.6.14.1651, 1 (lecto-
type); Lake Nyasa: Vua, C. Christy, 1925. — BMNH
1935.6.14.1652–1653, 2 (paralectotypes); Lake Nyasa:
Vua, C. Christy, 1925.
Otopharynx heterodon: BMNH 1935.6.14.1586, 1 (lec-
totype); Lake Nyasa: Monkey Bay. — YPM ICH
014404, 6; Malaŵi: Thumbi Island West; KRM party
(?), 23 Jun 1979.
Otopharynx lithobates: BMNH 1974.7.5:1, 1 (holotype);
Malaŵi: Thumbi Island West; 24 May 1971. — YPM
ICH 030203, 2 (paratypes); Malaŵi: Thumbi Island
West; MKO, KRM, TDK, 1 Aug 1980.
Otopharynx ovatus: BMNH 1935.6.14.1487, 1 (lecto-
type); south end of L. Nyasa; C. Christy, 1925. —
BMNH 1935.6.14.1488–1489, 2 (paralectotypes);
south end of L. Nyasa; C. Christy, 1925. — YPM ICH
014292, 2; Malaŵi: Nankumba Peninsula, Otter Point;
MKO, KRM, TDK, 29 Jul 1980.
Otopharynx selenurus: BMNH 1921.9.6.80, 1 (lecto-
type); “Lake Nyasa”; R. Wood. — BMNH 1921.9.6.81,
1 (paralectotype); “Lake Nyasa”; R. Wood.
Otopharynx speciosus: BMNH 1935.6.14.1650, 1 (lecto-
type); Lake Nyasa: Vua; C. Christy, 1925. — BMNH
1935.6.14.1649, 1 (paralectotype); Lake Nyasa: Mon-
key Bay; C. Christy, 1925. — YPM ICH 014311, 5;
Malaŵi: station Malembo III, 30-35 fathoms; LMTS,
19 Jul 1971.
Otopharynx spelaeotes: YPM ICH 026587, 3; Malaŵi:
Nkhata Bay; MKO, KRM, TDK, 23 Jul 1980. — YPM
ICH 030205, 1; Malaŵi: Nkhata Bay; MKO, KRM,
TDK, 21 Jul 1980.
Otopharynx tetraspilus: BMNH 1935.6.14.1609, 1 (lecto-
type); Lake Nyasa: southeast arm between the Bar and
Fort Maguire; C. Christy, 1925. — YPM ICH 014153,
4; Malaŵi: station Namiasi Ia, 15 fathoms; LMTS, 15
Jun 1971.
Otopharynx tetrastigma: BMNH 1893.11.15.34, 1 (lec-
totype); “Lake Nyasa and the upper Shire River”;
H. H. Johnston. — BMNH 1893.11.15.35–37, 3
(paralectotypes); “Lake Nyasa and the upper Shire
River”; H. H. Johnston. — YPM ICH 014159, 4;
Malaŵi: Monkey Bay.
Stigmatochromis modestus: BMNH 1893.1.17.5, 1 (holo-
type); “Lake Nyassa”; J. E S. Moore. — YPM ICH
014361, 1; Malaŵi: Monkey Bay; MKO, 8 Aug 1971.
Stigmatochromis pholidophorus: BMNH 1935.6.14.1544,
1 (holotype); Lake Nyasa: Vua; C. Christy, 1925. YPM
ICH 014313, 4; Malaŵi: Lake Malaŵi; LMTS.
Stigmatochromis pleurospilus: BMNH 1935.6.14.1475, 1
(holotype); Lake Nyasa: Lupembe sandbank; C.
Christy, 1925. — YPM ICH 026590, 2: Malaŵi:
Nkhata Bay; MKO, KRM, TDK, 23 Jul 1980.
Stigmatochromis woodi: YPM ICH 014150, 1; Malaŵi:
Monkey Bay; MKO, 9 Aug 1971. — YPM ICH
014593, 3; Malaŵi: Nankumba Peninsula, E of Otter
Island; MKO, KRM, TDK, 4 Jun 1980.
Tramitichromis intermedius: BMNH 1935.6.14.2081, 1
(lectotype); southwest arm or Fort Johnston; C.
Christy, 1925. — BMNH 1935.6.14.2082–2084, 3
(paralectotypes); southwest arm or Fort Johnston; C.
Christy, 1925. — YPM ICH 030234, 11; Malaŵi: off
beach at Chembe; MKO, KRM, TDK, 16 Jun 1980.
Six New Species of the Cichlid Genus Otopharynx • Oliver 195

Trematocranus brevirostris: BMNH 1935.6.14.2224, 1
(lectotype); Malaŵi: Bar House; C. Christy, 1925. —
BMNH 1935.6.14.2225, 1 (paralectotype); Malaŵi:
Bar House; C. Christy, 1925.
Trematocranus microstoma: BMNH 1935.6.14.
2232–2236, 5 (lectotype and paralectotypes); Malaŵi:
Karonga; C. Christy. — YPM ICH 014433, 21;
Malaŵi: trawled in 9–10 m across Mazinzi Bay; MKO,
KRM, TDK, 28 Jun 1980.
Trematocranus placodon: BMNH 1921.9.6.134, 1 (lecto-
type); “Lake Nyasa”; R. Wood. — YPM ICH 007800,
2; Malaŵi: 4 miles SE of Monkey Bay; LMTS, 7 Dec
1973.
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