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-FLORIDA
_MUSEUM,
OF NATURAL HISTORYTM
BULLETIN
TAXONOMIC REVISION OF LAKE TANGANYIKAN
SYNODONTIS (SILURIFORMES: MOCHOKIDAE)
Jeremy J. Wright and Lawrence M. Page
Vol. 46, No. 4, pp. 99-154
2006
UNIVERSITY OF FLORIDA
GAINESVILLE
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TAXONOMIC REVISION OF LAKE TANGANYIKAN SYNODONTIS
(SILURIFORMES: MOCHOKIDAE)
Jeremy J. Wright' and Lawrence M. Page'
ABSTRACT
The taxonomy of the Synodontis of Lake Tanganyika was investigated by examining 312 museum specimens. Morphometric, meristic, and
nontraditional internal and external characters were examined, resulting in the recognition of 11 species of Synodontis from Lake Tanganyika.
Ten of the species are endemic to the lake basin, and three are new to science. Synodontis grandiops n. sp. is most similar to S. multipunctatus
but is distinguished by measurements of the eye (64.2-81.0% of snout length vs. 44.9-62.0% in S. multipunctatus) and pectoral-fin ray counts
(7 vs. 8 in S. multipunctatus). Synodontis lucipinnis n. sp. is most similar to S. petricola but is distinguished by the lack of an axillary pore and
the presence of light-colored windows at the bases of the rayed fins. Synodontis ilebrevis n. sp. is most similar to S. polli but is distinguished
by the absence of a hindgut chamber, the shortness of the gut (0.8-1.4 times TL in S. ilebrevis vs. 4.0-5.5 times TL in S. polli), and the presence
of short, flattened papillae on the skin (vs. villous papillae in S. polli).
Key Words: Catfishes, Synodontis, Mochokidae, Lake Tanganyika, Africa.
TABLE OF CONTENTS
Introduction ..................................................... ............................................... 100
M ethods and Terminology................. ..................................................... 102
Taxonom ic Descriptions................ ......................................................... 104
Genus Synodontis........................................................................ 104
Synodontis dhonti..................... ...................................... ..................... 107
Synodontis grandiops n. sp..................... .................................................. 109
Synodontis granulosus..................... ..................................................... 114
Synodontis ilebrevis n. sp................................................ ...................... 117
Synodontis irsacae........................... ............................... ................... 122
Synodontis lucipinnis n. sp........ ....... .................................................1... 26
Synodontis melanostictus........................................... .......................... 130
Synodontis m ultipunctatus......................................... .......................... 134
Synodontis petricola.................................................. ........................... 138
Synodontis polli..................... ......................................................... 142
Synodontis tanganaicae.............................................. ............................. 147
K ey to Species.......................................................................... ...................... 15 1
D iscussion.......................... ......................................... ........................ 151
Acknowledgements.............................. ...................................................... 152
Literature C ited............................. .................................................................... 152
'Florida Museum of Natural History, P.O. Box 117800, University of Florida, Gainesville, FL 32611-7800
Wright, J.J. and L.M. Page. 2006.Taxonomic revision of Lake Tanganyikan Synodontis (Siluriformes: Mochokidae). Florida Mus. Nat. Hist. Bull.
46(4):99-154. [End of volume]
INTRODUCTION
Catfishes of the genus Synodontis are small to moder-
ately sized fishes (up to 800 mm total length) that occur
throughout sub-Saharan Africa and the Nile River Val-
ley. Poll (1971) divided the genus into eight geographi-
cally defined groups: 1) the northern supra-equatorial
group; 2) the exclusively Nilotic group; 3) the group of
species occurring exclusively in the Niger; 4) the west-
ern supra-equatorial group from the Senegal River to
the Ogowe River; 5) the Congo Basin group; 6) the south-
ernmost sub-equatorial group; 7) the Synodontis of east-
ern Africa ranging between Uebi Shebeli and the Rufigi
River; and 8) the Lake Tanganyikan group (Fig. 1).
The Synodontis of Lake Tanganyika inhabit mainly
rocky shoreline areas but also venture out over sandy
and shell bottoms (Matthes 1962; Coulter 1991 la). The
vertical distribution of these species is limited to a maxi-
mum depth of 50-100 m in the northern end of the lake
and to about 240 m in the southern end (Coulter & Spigel
1991). These depths represent the lower depth limits of
oxygen and differ with geography due to the fact that
Lake Tanganyika is composed of many sub-basins along
its length, which vary greatly in depth (from 350 m to
1470 m) (Coulter & Spigel 1991; Tiercelin & Mondeguer
1991). Locality data indicate that all species have a
lake-wide distribution, with the possible exceptions of S.
dhonti, S. ilebrevis and S. lucipinnis (see species de-
scriptions).
Although many different color patterns exist within
Synodontis, few are as striking or as instantly recogniz-
able as the pattern that is typical of most species of
Synodontis from Lake Tanganyika. This pattern con-
sists of dark triangles at the bases of all of the rayed fins
and dark spots on the upper body that may or may not
extend onto the belly. The barbels are typically white,
but may have scattered, dusky pigmentation at their
bases. Body color varies widely and may be nearly any
shade of yellow, green, brown, or gray, depending on the
species. Body size varies widely, from fairly small (maxi-
mum TL 100 mm in S. lucipinnis) to large (maximum
TL 585 mm in S. tanganaicae).
Male Synodontis have a distinct, cone-shaped
genital papilla (Matthes 1962). Minor differences in body
shape between sexes also occur in some species, with
females having slightly more robust bodies. Opercular
ornamentation has recently been discussed as a sexu-
ally dimorphic trait in some mochokid species (Friel &
Vigliotta 2006), but none of the Tanganyikan Synodontis
species have ornamented operculae. Information on
reproduction in Tanganyikan species of Synodontis is
mostly restricted to egg counts from gravid females. The
exception is S. multipunctatus, whose unique brood-
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
parasitism behavior has been well documented (Sato
1986).
Dietary information is slightly more abundant, and
a variety of prey items are utilized. Most species in-
clude insect larvae and at least a small amount of algal
matter in their diet. More specialized prey items that
may be taken include gastropods, bivalves, sponges, crus-
taceans, and eggs of other fishes (Poll 1946, 1953;
Matthes 1962; Coulter 1965-1966, 1991a).
SYSTEMATIC HISTORY
Synodontis is a genus of Mochokidae, a family of
catfishes endemic to sub-Saharan Africa and the Nile
River valley. With approximately 120 recognized spe-
cies, Synodontis is the most diverse of any African cat-
fish genus and, with the exception of Barbus, of any
African ostariophysan (Poll 1971). Poll (1971), the last
to revise Synodontis, recognized as valid seven species
from Lake Tanganyika: Synodontis multipunctatus
Boulenger, Synodontis granulosus Boulenger,
Synodontis dhonti Boulenger, Synodontis petricola
Matthes, Synodontis lacustricolus Poll (Synodontis
tanganaicae Borodin), Synodontis eurystomus
Matthes (Synodontis polli Gosse), and Synodontis
nigromaculatus Boulenger (recognized herein as
Synodontis melanostictus Boulenger). All but S.
melanostictus are endemic to the lake basin. The ten
endemic species recognized in this study can be distin-
guished from all other species of Synodontis by the pres-
ence of a black triangle at the base of the dorsal and
pectoral fins and vertical skin folds along the sides of
the body. With the exceptions of S. grandiops and S.
multipunctatus, all endemic Tanganyikan Synodontis
species also have well developed black triangles on the
base of the anal and pelvic fins. The triangles can no
longer be seen in the single known specimen of S. dhonti;
however, the original description of this species
(Boulenger 1917) and an earlier examination of this speci-
men (Matthes 1962) indicate that the color pattern of
the rayed fins in this species was consistent with that of
the remaining endemic species of Tanganyikan
Synodontis.
Poll's revision of Synodontis, while thorough for
its time, suffered from a lack of material, and appears to
have stifled taxonomic research on this genus, with only
a handful of new species described after its publication.
The Synodontis of Lake Tanganyika have largely been
ignored by taxonomists for the last 30 years. Recent
expeditions, particularly those to the Zambian coast of
the lake, which has historically been poorly sampled for
Synodontis, as well as the increased popularity of cer-
tain Tanganyikan Synodontis as aquarium species, have
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
-20 0 20 40
400 A
20"
-2 0" 6 -
-40 *
-20 0 20' 400
km
0 5001000
40
20
00
-20
-40
Figure 1. Drainage map of the African continent with Poll's geographic divisions outlined. 1 = Northern supra-
equatorial, 2 = Nile River drainage, 3 = Niger River drainage, 4 = Western supra-equatorial from the Senegal River to
the Ogowe River, 5 = Congo Basin, 6 = Southernmost sub-equatorial, 7 = Eastern Africa between Uebi Shebeli and
the Rufigi River, 8 = Lake Tanganyika drainage.
led to the availability of more specimens for study.
Schraml (2003) recently suggested that many of the speci-
mens entering the aquarium trade represent cryptic,
undescribed species. The aim of this study was to evalu-
ate the taxonomy of this group, bearing in mind that cryp-
tic species may exist. This possibility has led to the
utilization of several unconventional diagnostic charac-
ters, which have revealed undescribed species of
Synodontis from Lake Tanganyika.
Phylogenetic relationships of the endemic species
of Synodontis in Lake Tanganyika have not been inten-
sively studied. Brichard (1978) suggested that the spe-
cies do not form a monophyletic group. However, the
unique pattern of rayed fin coloration and the presence
of vertical folds in the skin constitute two synapomorphies
supporting the monophyly of the endemic species of
Synodontis in Lake Tanganyika. It is possible that S.
melanostictus does not share a most recent common
ancestor with the Tanganyikan endemics, given its wider
distribution and differences from the endemic species in
its rayed fin coloration and lack of vertical folds on the
skin.
METHODS AND TERMINOLOGY
Measurements were taken from the left side to the near-
est 0.1 mm using digital calipers and follow Poll (1971).
All counts were made using a Leica MZ75 dissecting
microscope. Vertebral and caudal-fin ray counts were
taken using lateral radiographs. Vertebral counts follow
the method of Skelton and White (1990), with the first
five fused vertebrae of the Weberian apparatus excluded
from reported values. Notation for fin ray counts is as
follows: upper case Roman numeral = fin spine, lower
case Roman numeral = unbranched rays, Arabic nu-
meral = branched fin rays. Terminology for premaxil-
lary dentition follows Skelton and White (1990). Gut
length measurements and observations of the hindgut
chamber of all species were performed only on speci-
mens that had a belly cut made prior to the beginning of
this study, with the exception of UF specimens, which
were dissected. The hindgut chamber corresponds to
the "poche intestinal" of Taverne and Aloulou-Triki
(1974). All photographs were taken using a Kodak
EasyShare CX7430 digital camera and edited using
Adobe Photoshop CS2. Maps were produced using
Adobe Illustrator. Institutional abbreviations follow
Leviton et al. (1985) except for that of the South Afri-
can Institute of Aquatic Biodiversity (SAIAB).
CHARACTERS EXAMINED
In addition to the 22 morphometric measurements
and seven meristic counts that were made for all speci-
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
mens, several new characters were found to be diag-
nostically useful. This is in contrast to most of the work
that has previously been done in this genus, which has
tended to rely mainly on morphometric ratios to diag-
nose species. These new characters are particularly
appealing because they allow for easy and accurate vi-
sual identification of specimens. Some of these charac-
ters, such as fin spine color are self-explanatory. Oth-
ers are explained below.
Foremost among these characters is the axillary
pore, an opening (or series of openings) that is located
between the base of the pectoral fin spine and ventral
margin of the humeral process (Figs. 2-5). The function
of the axillary pore is unknown in mochokids, and the
absence or presence of this structure has been noted in
only one previously described species of Synodontis
(Friel & Vigliotta 2006). The presence of a similar struc-
ture has been noted in seven other catfish genera:
Ariopsis (Ariidae) (Halstead et al., 1953), Ameiurus,
Ictalurus, and Noturus (Ictaluridae) (Reed 1907;
Birkhead 1967, 1972), Acrochordonichthys (Akysidae)
(Ng & Ng 2001), Ituglanis (Trichomycteridae) (Datovo
and Landim 2005), and is also present in Glyptothorax
(Sisoridae) (pers. obs.). In Acrochordonichthys, this
structure has been shown to produce a mucosal secre-
tion that possesses toxic properties (Ng & Ng 2001).
The utility of this secretion in the other genera has been
a matter of some debate. The presence or absence of
this character divides the 11 Tanganyikan species of
Synodontis into two groups; six species have an axil-
lary pore (Table 1) and five (including the non-endemic
S. melanostictus) do not (Table 2).
All species of Synodontis have a premaxillary
toothpad, with several rows of unicuspid, chisel-shaped
teeth. In some species this toothpad is continuous, while
in others it is clearly composed of two elements, which
are separated by a thin band of skin or a suture that
Figure 2. The axillary pore of Synodontis
multipunctatus, holotype, BMNH 1898.9.9.76,280 mm
TL, 220 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 3. Humeral processes of A) Synodontis
granulosus, UF 160945, 192 mm TL, 149 mm SL, B)
S. multipunctatus, holotype, BMNH 1898.9.9.76, 280
mm TL, 220 mm SL, C) S. grandiops, MRAC 53096-
53100, 114 mm TL, 91 mm SL, D) S. petricola, MRAC
A3-033-P-0002-0011, 117 mm TL, 98 mm SL. An axil-
lary pore is present in all of these species.
Figure 4. Humeral processes of A) Synodontis
lucipinnis, holotype SAIAB 77880, 97 mm TL, 78 mm
SL, B) S. melanostictus, holotype, BMNH 1906.9.8.72,
TL 290 mm, SL 237 mm, C) S. tanganaicae, lectotype,
MCZ 32538,415 mm TL, 320 mm SL, D) a paratype of
S. lacustricolus, MRAC 90288, 519 mm TL, 434 mm
SL, provided for comparison with C. An axillary pore is
absent in A) and B), present in C) and D).
Figure 5. Humeral processes of A) Synodontis dhonti,
holotype, MRAC 14344, 395 mm TL, 325 mm SL, B) S.
irsacae, MRAC A3-033-P-0036-0040, 139 mm TL, 116
mm SL, C) S. polli, MRAC A3-033-P-0076-0080, 160
mm TL, 132 mm SL, D) S. ilebrevis, UF 160942, holo-
type, 139 mm TL, 116 mm SL. An axillary pore is present
in A), absent in other species shown.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
transversely bisects the toothpad. The separated condi-
tion is referred to herein as interrupted, and the
unseparated condition as uninterrupted. This appears to
be the first study to examine this character in
Synodontis, although the length of the premaxillary
toothpad has often been used in previous morphological
examinations of Synodontis species.
The skin of the ten endemic species of Tanganyikan
Synodontis is unique within the genus in having numer-
ous minute, vertical skin folds. The purpose of these
folds is unknown. The folds distinguish the Tanganyikan
species from species outside the lake's basin but are not
helpful in distinguishing among species from the lake.
The papillae on the skin of certain species, however,
separates them from other Tanganyikan species in which
the skin is bare. This character is most useful in the
identification of fresh or well-preserved specimens, as
the papillae may degrade.
Some species of Synodontis have a swelling in
the posterior part of the intestine which Taverne and
Aloulou-Triki (1974) termed the "poche intestinal" and
which is referred to herein as the hindgut chamber. A
similar structure is known only in species of Kyphosidae,
a family of herbivorous, marine fishes that utilizes fer-
mentative digestion in the hindgut (Rimmer & Wiebe
1987; Clements & Choat 1997; Mountfort et al. 2002).
The presence of a similar structure in highly herbivo-
rous mochokid species such as S. polli,
Brachysynodontis batensoda, and Hemisynodontis
membranaceus suggests that the mochokid hindgut
chamber may play a similar role. If so, this is the first
time that such a behavior and structure has been found
in a freshwater fish. The presence or absence of the
hindgut chamber appears to be species-specific and can
distinguish between species for which few good exter-
nal diagnostic characters exist.
TAXONOMIC DESCRIPTIONS
Order Siluriformes Hay 1929
Family Mochokidae Jordan 1923
Genus Synodontis Cuvier 1817
Diagnosis. Lateral line complete and midlateral,
extending onto base of caudal fin. Cone-shaped genital
papilla in male; papilla absent in female. Anterior nos-
trils tubular; posterior nostrils with semicircular flaps of
skin along anterior margin. Gill opening extends from
lower edge of ventral rounded occipital-nuchal process
to point horizontal to base of pectoral fin spine. Gill
membranes broadly joined at isthmus. Well developed,
papillate, membranous flap immediately posterior to pre-
maxillary toothpad. Eye with free orbital margin.
Table 1. Diagnostic characters for Lake Tanganyikan species of Synodontis in which an axillary pore is present.
Mandibular teeth
Axillary pore
Spots on body
Fin spine color
Triangles on rayed fins
Pectoral-fin rays
Eye/Snout length
Premaxillary toothpad
Maximum TL, mm
Secondary branches
on medial mandibular
barbel
Occipito-nuchal shield
covered with skin
Papillae on body skin
Gut/Body length
Hindgut chamber
S. dhonti
22
Large
Absent
Dark
Present
8
23.1%
Uninterrupted
395
Present
No
Granular
No data
No data
S. grandiops
17-26
Large
Large
Dark
Present -
dorsal, pectoral
Absent anal,
pelvic
7
64.2-81.0%
Uninterrupted
150
Absent
Absent
0.8-0.9
Absent
S. granulosus
28-51
Large
Absent or small
Dark
Present
7-8
31.2-50.2%
Uninterrupted
270
Absent
Granular
0.5
Absent
S. multipunctatus
13-29
Large
Large
Dark
Present -
dorsal, pectoral
Absent anal,
pelvic
8
44.9-62.0%
Uninterrupted
280
Absent
Yes
Absent
0.5-0.8
Absent
S. petricola
31-50
Small
Large
White
Present
8-9
28.7-40.1%
Interrupted
135
Present
Yes
Absent
1.2-1.3
Present
S. tanganaicae
33-49
Large
Absent or small
Dark
Present- juvenile
Absent- adult
8-9
16.0-26.9%
Interrupted
585
Present
No
Granular
No data
Absent
Table 2. Diagnostic characters for Lake Tanganyikan species of Synodontis that lack an axillary pore. Traits not diagnostic within this group (premaxillary
toothpad, secondary branches on medial mandibular barbel) are included to facilitate comparison with species in Table 1.
Mandibular teeth
Spots on body
Fin spine color
Triangles on rayed fins
Pectoral-fin rays
Eye/Snout length
Body depth/SL
Premaxillary toothpad
Maximum TL, mm
Secondary branches
on medial mandibular
barbel
Occipito-nuchal shield
covered with skin
Papillae on body skin
Gut/Body length
Hindgut chamber
S. ilebrevis
50-66
Small
Dark
Present
8-9
26.0-33.4%
18.2-20.1%
Interrupted
150
Present
Yes
Flat, granular
0.8-1.4
Absent
S. irsacae
15-29
Large
Dark
Present
8-9
28.7-43.7%
17.7-28.6%
Interrupted
190
Present
Yes
Absent
0.7-0.8
Absent
S. lucipinnis
35-51
Large
White
Present
8-9
25.2-35.8%
15.7-23.9%
Interrupted
100
Present
Yes
Absent
No data
No data
S. melanostictus
23-36
Small
Dark
Absent
9-10
31.5-38.1%
20.0-23.4%
Interrupted
520
Present
No
Villous
1.3-1.6
Present
S. polli
40-70
Moderate to Large
Dark
Present
8-9
25.8-39.3%
20.2-27.0%
Interrupted
180
Present
Yes
Villous
4.0-4.5
Present
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Synodontis dhonti Boulenger, 1917
(Figs. 5A, 6, 7; Tables 1, 3)
Synodontis dhonti Boulenger, 1917:367, description,
Kilewa Bay, Lake Tanganyika; 1920:42, description,
Kilewa Bay, Lake Tanganyika. Worthington and
Ricardo, 1936:1067, 1077, 1101, note, Lake
Tanganyika. Poll, 1946:221, figs. 21, 22, descrip-
tion, Kilewa Bay, Lake Tanganyika; 1971:364, figs.
168 and 169, pls. VI.6, XI.19. Matthes, 1962:37,
fig. a, pl. 3, description, diet, Lake Tanganyika.
Material Examined.- Holotype, MRAC 14344, TL
395 mm, SL 325 mm, Lake Tanganyika, Kilewa, coll.
Stappers, 18.V.1912.
Diagnosis. -Axillary pore present, large; mandibu-
lar teeth 22; body lacking spots; fin spines brown; 8 pec-
toral-fin rays; black patches at base of rayed fins present;
eye 23.1% snout length; premaxillary toothpad uninter-
rupted; secondary branches on medial mandibular bar-
bel present; occipito-nuchal shield not covered with skin;
granular papillae present on skin of body; maximum TL
395 mm.
Synodontis dhonti can be distinguished from S.
ilebrevis, S. irsacae, S. lucipinnis, S. melanostictus,
and S. polli by the presence of an axillary pore, an unin-
terrupted premaxillary toothpad, and a smaller eye
(23.1% of snout length vs. 26.0-33.4% in S. ilebrevis,
28.7-43.7% in S. irsacae, 25.2-35.8 % in S. lucipinnis,
31.5-38.1% in S. melanostictus, and 25.8-39.3% in S.
polli). Synodontis dhonti is separated from S.
granulosus, S. ilebrevis, S. lucipinnis, S.
melanostictus, S. petricola, S. polli, and S.
tanganaicae by having a smaller number of mandibular
teeth (22 vs. 50-66 in S. ilebrevis, 28-51 in S.
granulosus, 35-51 in S. lucipinnis, 23-36 in S.
melanostictus, 31-50 in S. petricola, 40-70 in S. polli,
and 33-49 in S. tanganaicae). The presence of sec-
ondary branches on the medial mandibular barbel and a
smaller eye (23.1% of snout length vs. 64.2-81.0% in S.
grandiops, 31.2-50.2% in S. granulosus, and 44.9-
62.0% in S. multipunctatus) differentiates S. dhonti
from S. grandiops, S. granulosus, and S.
multipunctatus. Synodontis dhonti differs further from
S. grandiops, S. irsacae, S. lucipinnis, S.
multipunctatus, and S. petricola in having granular
papillae on the body (vs. papillae absent). Brown fin
spines further distinguish S. dhonti from S. lucipinnis
and S. petricola, in which they are white. A lack of
spots on the body and an elevated number (n=25) of
elongated gill rakers distinguishes S. dhonti from all other
Tanganyikan Synodontis with the exception of adult S.
tanganaicae. Synodontis dhonti can be further dis-
tinguished from S. tanganaicae by having a poorly de-
veloped adipose fin (vs. well developed) and an uninter-
rupted premaxillary toothpad (vs. interrupted in S.
tanganaicae).
Description. Morphometric and meristic data in
Table 3. Maximum TL 395 mm, SL 325 mm. Body not
compressed. Predorsal profile straight, interrupted by
convex ridge formed by orbit of eye. Preanal profile
convex. Skin on body forming numerous vertical folds
covered with granular papillae extending onto all fins.
Head slightly depressed and broad; skin covered
with granular papillae. Snout with bluntly rounded mar-
gin when viewed laterally and dorsally. Anterior nostrils
tubular; posterior nostrils with semicircular flaps of skin
along anterior margin. Occipito-nuchal shield rugose,
not covered with skin; terminating posteriorly with nar-
row, rounded process on either side of dorsal spine; ven-
trally with narrow, bluntly pointed process that extends
to upper margin of humeral process on either side of
body. Eye located dorsolaterally; ovoid; horizontal axis
longer. Interorbital area flat.
Mouth subterminal; lips wide and papillate. Man-
dibular teeth 22, unicuspid; arranged in single transverse
row. Premaxillary toothpad uninterrupted; primary, sec-
ondary and tertiary premaxillary teeth discrete; numer-
ous; arranged in 4, 5, and 1 irregular rows, respectively.
Maxillary barbel with narrow basal membrane;
lacking branches or crenelations; extending at least to
base of pectoral fin. Lateral mandibular barbel extend-
ing to point just short of anterior margin of pectoral girdle,
with 4-5 non-tuberculate branches; lacking secondary
branches. Medial mandibular barbel about 1/2 length of
lateral barbel; with 2 pairs of tuberculate branches; sec-
ondary branches present.
Dorsal fin 11,7; posterior margin straight. Dorsal-
fin spine long, striated, nearly straight, terminating in short,
dark filament; anterior margin smooth; posterior margin
with small serrations distally. Pectoral fin 1,8; posterior
margin straight. Pectoral-fin spine roughly equal in length
to dorsal-fin spine, striated, slightly curved, terminating
in short, dark filament; anterior spine margin granulous;
posterior margin with large retrorse serrations along
entire length. Adipose fin poorly developed, margin con-
vex. Pelvic fin i,6; located at vertical through posterior
base of dorsal fin; tip of appressed fin barely reaches
base of anal fin. Anal fin v,8; posterior margin rounded;
base located ventral to adipose fin. Caudal fin i,7,8,i;
forked; lobes pointed.
Humeral process narrow, elongated, granulous;
possessing distinct ridge on ventral margin; dorsal mar-
gin concave; terminating in blunted point (Fig. 5a). Large
axillary pore just ventral to humeral process.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Table 3. Morphometric and meristic counts for Synodontis dhonti. All morphometrics given in percent of base
measurement. SL = Standard length, HL = Head length, LMB = Lateral mandibular barbel, MMB = Medial mandibu-
lar barbel, HPL = Humeral process length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD =
Caudal peduncle depth.
MEASUREMENT HOLOTYPE
Body depth/SL 27.2
Head length/SL 33.3
Snout-dorsal length/SL 42.6
Adipose fin length/SL 25.1
Maxillary barbel length/SL 23.8
Dorsal spine length/SL 23.9
Pectoral spine length/SL 24.4
Head width/HL 81.7
Head depth/HL 82.5
Humeral process length/HL 49.9
Snout length/HL 48.6
Eye/HL 11.2
Interorbital width/HL 35.8
Postorbital length/HL 43.4
Mouth length/HL 34.2
Maxillary barbel length/HL 71.3
LMB/HL 33.3
MMB/HL 16.9
MMB/LMB 50.7
Dorsal spine length/HL 71.8
Pectoral spine length/HL 73.2
Eye/Snout length 23.1
Interorbital width/Snout length 73.7
Postorbital length/Snout length 89.5
Eye/Interorbital width 31.4
Eye/Postorbital length 25.8
HPL/HPD 232.7
CPL/CPD 107.8
MERISTICS
Mandibular tooth count 22
Dorsal fin count 1,7
Pectoral fin count 1,8
Pelvic fin count i,6
Anal fin count v,8
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 6. Holotype of Synodontis dhonti, MRAC 14344, 395 mm TL, 325 mm SL.
28" 30' 32" Coloration in alcohol. Body uniformly brown (Fig.
6). All barbels brown. Dorsal and pectoral-fin spines
Bur df i brown, filaments dark. All rayed fins uniformly brown-
-4 4- Distribution. Known only from Kilewa, Lake
Tanganyika (Fig. 7).
Habitat. Unknown.
Mala aI m F Diet. Unknown.
Reproduction. No information exists regarding
the reproduction of Synodontis dhonti.
\- Lukuga Tanzania -6. Taxonomic Remarks. Synodontis dhonti
Democr tic Republic Boulenger is known from a single specimen, collected in
of Congo 1912. Synodontis irsacae Matthes was placed into the
T synonymy of S. dhonti by Matthes (1962). Morpho-
logical differences between these species were said to
be due to ontogeny, but we have found no specimens
that support this claim. We consider S. dhonti and S.
-8- L- b`- 8- irsacae to be distinct species. Evidence for this conclu-
Warinpa sion is presented in the description of S. irsacae.
L Mweru
2_- Zambia Synodontis grandiops n. sp.
28 30 km 32" (Figs. 3C, 8, 9, 10; Tables 1, 4)
0 50 100
Holotype. BMNH 1982.4.13.4785, TL 129 mm,
Figure 7. Known distribution of Synodontis dhonti. T SL 96 mm, Mwakizega coastline, L. Tanganyika, coll.
denotes type locality. R. Travers.
Paratypes. (1) BMNH 1982.4.13.4786, TL 112
mm, SL 91 mm, collection information same as for holo-
type, (2) BMNH 1920.5.25.83-84, TL 98-103 mm, SL
76-79 mm, Vua Bay, L. Tanganyika, coll. L. Stappers,
(1) BMNH 1955.12.20.1833, TL 103 mm, SL 80 mm,
Moba Bay, L. Tanganyika, coll. Inst. Roy. Sci. Nat.
Belge, 7.111.1947, (1) BMNH 1955.12.20.1837, TL 101
mm, SL78 mm, Rumonge Bay, L. Tanganyika, coll. Inst.
Roy. Sci. Nat. Belge, (1) BMNH 1982.4.13.4784, TL
105 mm, SL 81 mm, Kamara Bay, L. Tanganyika, coll.
R. Travers, (3) BMNH 1982.4.13:4789-4791, TL 109-
117 mm, SL 86-92 mm, Kigonga Bay, L. Tanganyika,
coll. R. Travers, (2) BMNH 1982.4.13:4787-4788, TL
104-120 mm, SL 81-93 mm, Elephant's Foot Peninsula,
L. Tanganyika, coll. R. Travers, (3) MRAC 39186-39188,
TL 93-100 mm, SL 72-80 mm, Moba (Lake Tanganyika),
coll. Van Maldern, 1932, (2) MRAC 53096-53097, TL
107-114 mm, SL 80-91 mm, Nyanza (L. Tang.), coll. A.
Testiade, 1937, (3) MRAC 90273-90275, TL 123-147
mm, SL 96-111 mm, Stat: 63, offshore from the
Malagarasi, 10-15 km from the coast, coll. M. Poll,
3.11.1947, (2) MRAC 91606-91607, TL 114-121 mm, SL
96-98 mm, Uvira, Lake Tanganyika, coll. G. Marlier,
1 .IX. 1949, (3) SAIAB 77890, TL 98-104 mm, SL 75-81
mm, Jacobsen's Beach, Tanzania, 0454'31"S,
029036'02"E, 05.X.1992.
Diagnosis. -Axillary pore present, large; mandibu-
lar teeth 17-26; body with large spots; fin spines brown
to black; 7 pectoral-fin rays; black patches at base of
pelvic and anal fins absent or poorly developed; eye 64.2-
81.0% snout length; premaxillary toothpad uninterrupted;
secondary branches on medial mandibular barbel ab-
sent; occipito-nuchal shield covered with skin; papillae
on skin of body absent; hindgut chamber absent; maxi-
mum TL 150 mm.
This species is most similar in appearance to
Synodontis multipunctatus, but is distinguished by its
much larger eye (64.2-81.0% of snout length vs. 44.9-
62.0% in S. multipunctatus). A consistent difference
also exists in pectoral-fin ray counts, with S. grandiops
having a count of 1,7 and S. multipunctatus having a
count of 1,8, and S. grandiops reaching a much smaller
maximum TL than does S. multipunctatus (150 mm vs.
280 mm). Synodontis grandiops is distinguished from
other Tanganyikan species of Synodontis by its much
larger eye (Tables 1 and 2) and modal pectoral-fin ray
count of 1,7; other species have modal pectoral-fin ray
counts of 1,8 or 1,9.
Description. Morphometric and meristic data in
Table 4. Maximum TL 150 mm, SL 110 mm. Body
compressed laterally. Predorsal profile convex; inter-
rupted by ridge formed by dorsal rim of eye. Preanal
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
profile convex. Skin on body forming numerous vertical
folds; papillae absent.
Head somewhat depressed, broad; skin smooth.
Snout subconical when viewed laterally; rounded when
viewed dorsally. Occipito-nuchal shield completely cov-
ered with skin; terminating posteriorly with narrow,
rounded process on either side of dorsal spine; ventrally
with wide, rounded process (covered with skin in smaller
specimens) that extends to upper margin of humeral pro-
cess on either side of body, also covered with skin in
smaller specimens. Eye dorsolateral, ovoid; horizontal
axis longer. Interorbital area flat to slightly convex.
Mouth subterminal; lips wide and papillate. Man-
dibular teeth 17-26, short, unicuspid; arranged in single
transverse row. Premaxillary toothpad uninterrupted;
primary, secondary and tertiary premaxillary teeth dis-
crete, numerous, arranged in 2, 2, and 1 irregular rows,
respectively.
Maxillary barbel without basal membrane; lacking
branches or crenelations; extending at least to base of
pectoral fin. Lateral mandibular barbel extending to point
just short of anterior margin of pectoral girdle; with 4-5
non-tuberculate branches; lacking secondary branches.
Medial mandibular barbel 1/2 to 2/3 length of lateral bar-
bel; with 4-5 pairs non-tuberculate branches; lacking
secondary branches.
Dorsal fin 11,7; posterior margin straight. Dorsal-
fin spine long, striated, nearly straight, terminating in short,
white filament; anterior margin of fin spine with 0-3 small
serrations distally; posterior margin with small serrations
distally. Pectoral fin 1,7; posterior margin straight. Pec-
toral-fin spine roughly equal in length to dorsal-fin spine,
striated, slightly curved, terminating in short, white fila-
ment; anterior spine margin withmany small, antrorse
serrations; posterior margin with large, retrorse serra-
tions along entire length. Adipose fin short, poorly de-
veloped, margin convex. Pelvic fin i,6; located anterior
to vertical through origin of adipose fin; tip of appressed
fin barely reaches base of anal fin. Anal fin iii-iv,6-8;
posterior margin nearly straight; base located ventral to
adipose fin. Caudal fin i,7,8,i; forked; lobes pointed.
Humeral process narrow, elongated, granulous;
possessing distinct ridge on ventral margin; dorsal mar-
gin convex; terminating in sharp point (Fig. 3C). Large,
dark-colored axillary pore present just ventral to humeral
process. Gut length 0.8-0.9 times body length (n = 4,
MRAC 53096-53097, MRAC 90273-90275). Hindgut
chamber absent.
Coloration in alcohol. Dorsum pale yellow to
brown, covered with large black spots (Figs. 8, 9). Spots
larger, irregular, sometimes confluent in juvenile speci-
mens. Belly white, with or without black spots. Maxil-
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 4. Morphometrics and meristic counts for S. grandiops. All morphometrics given in percent of base measure-
ment. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE S. grandiops (n=25) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
27.2
30.2
39.5
33.1
26.3
27.8
78.1
75.8
56.4
41.1
30.0
34.9
35.6
28.3
95.4
53.5
27.4
51.2
91.9
72.9
85.0
86.6
85.8
84.2
286.6
107.7
20.2-27.2
27.8-32.4
35.1-43.0
20.3-33.1
26.3-37.8
21.7-31.1
22.7-32.4
73.0-92.9
69.0-83.2
52.0-62.8
39.4-46.6
27.4-31.9
26.1-37.4
31.4-41.3
28.0-37.7
90.2-127.0
45.9-68.2
20.4-44.2
44.4-65.5
73.2-103.6
76.6-108.0
64.2-81.0
58.4-85.0
69.5-100.0
77.7-120.7
73.7-100.2
241.4-431.9
107.7-189.6
23.21.8
29.61.0
39.21.8
27.93.3
30.93.4
27.02.8
27.12.8
80.64.4
75.33.6
55.92.5
42.71.8
29.71.4
31.92.9
35.32.3
31.92.5
104.310.3
55.55.9
30.34.6
54.65.7
91.210.1
92.09.9
69.74.6
74.66.4
83.07.6
94.211.8
84.57.3
310.838.4
144.418.9
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
17(1); 18(2); 19(2); 20(4); 21(6);
22*(3); 23(1); 24(4); 25(1); 26(1);
II,7*(25)
I,7*(21); I,7,i(4)
i,6*(25)
iii,6(1); iii,6,i(1); iii,7(3); iii,7,i(1);
iv,6(1); iv,6,i(1); iv,7*(13);
iv,7,i(1); iv,8(3)
i,7,8,i(3)
34(3)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 8. Synodontis multipunctatus (top), BMNH 1982.4.13.4789, 115 mm TL, 91 mm SL vs. S.
grandiops (bottom), BMNH 1982.4.13.4784, 105 mm TL, 81 mm SL. The eye of S. grandiops is
noticeably larger than that of S. multipunctatus, particularly in proportion to the length of the snout.
lary and mandibular barbels white. Iris yellowish to cop-
per colored. Dorsal and pectoral-fin spines brown to
black, filaments white. Pectoral spine with thin, light
stripe along anterior margin. Dorsal and pectoral fins
with black triangles at their base, posterior margins white
in color. Triangles may be completely solid or composed
of closely spaced spots. Anal and pelvic fins white, lack-
ing dark triangles of other Tanganyikan species, single
black spot may be present at base of these fins. Adi-
pose fin with white dorsal edge. Both lobes of caudal
fin with black bar from base to tip of fin; posterior mar-
gin of fin white.
Distribution. Lake Tanganyika (Fig. 10); appar-
ently common, though less so than Synodontis
multipunctatus.
Habitat. Likely inhabits littoral to benthic zones
over shell, sand and mud bottoms, as do most of the
other Synodontis species of the lake.
Diet. No information exists regarding the diet of
Synodontis grandiops. It is likely that S. grandiops
feeds on small gastropods, as do the other species of
Synodontis from the lake.
Reproduction. It is unknown whether Synodontis
grandiops shares the brood parasite behavior of S.
multipunctatus. In fact, without examining the speci-
mens used in the original study of this behavior (Sato
1986), it is impossible to say whether S. multipunctatus
or S. grandiops was observed in this study. Unfortu-
nately, the specimens used do not appear to have been
preserved. Differences in spawning time are likely to
exist, as one mixed lot (now split into MRAC 39186-
39188 and MRAC 39189-39192) containing specimens
of S. multipunctatus and S. grandiops of similar size
contained gravid females of the latter species, while the
females of S. multipunctatus lack mature gonads.
Etymology. -The specific name is a Latinized com-
bination of the Latin grandis, meaning large or big, and
the Greek ops, meaning eye, a reference to the in-
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 9. Holotype of Synodontis grandiops, BMNH 1982.4.13.4785, TL 129 mm, SL 96 mm.
km
0 50 100
Figure 10. Known distribution of Synodontis grandiops.
T denotes type locality.
creased size of the eye of this species, particularly in
relation to snout length. Gender: feminine.
Synodontis granulosus Boulenger, 1900
(Figs. 3A, 11, 12; Tables 1,5)
Synodontis granulosus Boulenger, 1900:480, descrip-
tion, northern end of Lake Tanganyika; 1901a:308,
description, northern end of Lake Tanganyika;
1901b: 149, pl. 16, description, northern end of Lake
Tanganyika; 1911:413, fig. 311, description, northern
end of Lake Tanganyika. Moore, 1903:166, fig.,
northern Lake Tanganyika. Worthington and
Ricardo, 1936:1067, 1077, 1101, note, Kabanga Bay,
Lake Tanganyika. -Poll, 1946:219, description, Kilewa
Bay; 1953:152, description, Lake Tanganyika;
1971:303, fig. 138, 139, pl. V, XI, description, Lake
Tanganyika. Hulot, 1950:170, Lake Tanganyika. -
Matthes, 1962:44, description, Lake Tanganyika. -
Coulter, 1965-1966:34, abundance, Lake Tanganyika;
1991a:152, 154, 181, 182, table 8.2, 8.3, 8.16, abun-
dance, ecology, Lake Tanganyika. Brichard,
1978:424, identification, Lake Tanganyika; 1989:478,
479, photo, identification. Sands, 1983:23, check-
list. Daget, Gosse, & Thys van den Audenaerde,
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
1986:129, check-list. Burgess, 1989:576, pl. 94, Lake
Tanganyika. Kobayagowa, 1989:14, photo.
Material Examined. Lectotype (Poll, 1971),
BMNH 1906.9.6.40, TL 270 mm, SL 216 mm, north end
of Lake Tanganyika, coll. J.E.S. Moore, (2)
paralectotypes, BMNH 1906.9.6.41-42, TL 220-246 mm,
SL 181-202 mm, north end of Lake Tanganyika, coll.
J.E.S. Moore, (3) BMNH 1936.6.15.1199-1201, TL 213-
249 mm, SL 165-189 mm, Lake Tanganyika, coll. Christy,
(1) MRAC 130378, TL 186 mm, SL 143 mm, Island of
Moboko, Lake Tanganyika, coll. H. Matthes, 3.IX. 1958,
(1) MRAC 94-069-P-0289, TL 153 mm, SL 124 mm,
Luhanga 15 km S. of Uvira, Lake Tanganyika (Zaire),
coll. L. DeVos, 15.II.1994, (1) MRAC 14165, TL 226
mm, SL 181 mm, Kilewa, in front of the Tombala River,
Lake Tanganyika, coll. Stappers, 25.VII.1912, (1)
MRAC 14157, TL 260 mm, SL 209 mm, Kilewa Bay,
" Lake Tanganyika, coll. Stappers, 9.IV. 1912, (1) MRAC
130465, TL 56 mm, SL 42 mm, Kalundu, Lake
Tanganyika, coll. H. Matthes (I.R.S.A.C.), 21.X.1960,
(4) MRAC 82-012-P-13-16, TL 141-194 mm, SL 112-
154 mm, Magara, Lake Tanganyika (Burundi), coll.
Schreyen, XII.1981, (1) MRAC A1-094-P-52, TL 109
mm, SL 84 mm, Crocodile Island, Lake Tanganyika (Zam-
bia), coll. Snoeks, Hanssens, Verheyen et al., 16.X.2001,
(1) UF 160945, TL 192 mm, SL 149 mm, Lake
Tanganyika at Ikola, Tanzania, via Pete Hauschner, Tropi-
cal Fish Collector, coll. V.2005.
Diagnosis.- Axillary pore present, large; mandibu-
lar teeth 28-51; body lacking large spots; fin spines brown
to black; 7-8 pectoral-fin rays; black triangles present
on bases of all rayed fins; eye 31.2-50.2% snout length;
premaxillary toothpad uninterrupted; secondary branches
on medial mandibular barbel absent; occipito-nuchal
shield not covered with skin; granular papillae present
on skin of body; humeral process length 253.8-437.2%
of humeral process width; hindgut chamber absent;
maximum TL 270 mm.
Synodontis granulosus can be distinguished from
S. ilebrevis, S. irsacae, S. lucipinnis, S. melanostictus,
and S. polli by the presence of an axillary pore.
Synodontis granulosus is further distinguished from
these species, and also from S. dhonti and S.
tanganaicae by the lack of secondary branches on the
medial mandibular barbel (vs. present). An uninterrupted
premaxillary toothpad further separates S. granulsosus
from S. ilebrevis, S. irsacae, S. lucipinnis, S.
melanostictus, S. petricola, S. polli, and S.
tanganaicae. Synodontis granulosus further differs
from S. tanganaicae by differences in humeral pro-
cess shape humerall process length 253.8-437.2% of
humeral process width in S. granulosus vs. 172.9-
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 5. Morphometric data and meristic counts for Synodontis granulosus. All morphometrics given in percent
of base measurement. *Meristic data for lectotype. Abbreviations as in Table 3.
MEASUREMENT LECTOTYPE RANGE (n=17) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
28.1
27.7
36.9
26.4
28.5
26.5
23.8
94.8
88.4
63.7
47.4
14.8
43.6
42.6
44.1
103.1
70.5
24.8
35.2
95.7
86.0
31.2
92.0
89.9
34.0
34.7
321.8
124.6
20.8-30.8
27.4-31.7
36.9-41.9
25.6-39.0
22.9-38.3
24.3-31.9
23.8-31.1
90.1-105.2
71.4-88.6
57.4-76.4
41.3-50.0
14.8-22.6
37.0-44.0
37.0-44.5
32.9-44.1
81.4-122.6
41.8-75.2
20.8-32.3
32.2-65.6
85.5-110.3
84.0-101.8
31.2-50.2
81.0-96.0
80.3-107.9
34.0-60.6
34.7-61.1
253.8-437.2
93.4-154.5
25.22.8
28.81.4
38.91.4
30.03.7
31.14.7
28.32.2
27.01.9
97.03.7
81.64.5
67.05.7
46.42.1
19.42.3
41.02.2
40.21.9
37.53.7
107.412.6
58.89.0
27.53.6
47.89.6
98.28.8
93.65.9
41.96.2
88.34.1
86.66.5
47.57.3
48.46.8
333.852.0
126.616.4
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
28(1); 32(1); 36(2); 37(1); 42(1);
43(1); 44(2); 45(1); 46*(2); 48(1);
51(1)
I1,7*(13); II,7,i(1); 11,8(3)
1,7(3); 1,8(14)
i,6*(15); i,7(2)
iii,8*(2); iii,9(1); iv,6,i(1); iv,7(3);
iv,8(2); iv,8,i(l); iv,9,i(1); v,7(1);
v,7,i(l); v,8(3);
i,7,8,i(5)
38(3), 39(1)
255.4% in S. tanganaicae) (Figs. 3A, 4C,D).
Synodontis granulosus can be further distinguished
from S. dhonti by having a greater number of mandibu-
lar teeth (28-51 vs. 22 in S. dhonti) and a larger eye
(31.2-50.2% of snout length vs. 23.1% in S. dhonti).
Synodontis granulosus differs from S.
multipunctatus and S. grandiops in lacking large spots
on the body, having a higher number of mandibular teeth
(28-51 vs. 13-29 in S. multipunctatus and 17-26 in S.
grandiops), a smaller eye (31.2-50.2% of snout length
vs. 44.9-62.0% in S. multipunctatus and 64.2-81.0% in
S. grandiops), an occipito-nuchal shield which is not
covered by skin, and body skin which is covered by
granular papillae. Synodontis granulosus is distin-
guished from S. petricola and further separated from
S. lucipinnis by lacking large spots on the body and
secondary branches on the medial mandibular barbel,
having a large axillary pore (vs. very small in S. petricola,
absent in S. lucipinnis), dark fin spines, 7-8 pectoral-fin
rays (vs. 8-9 in S. petricola and S. lucipinnis), an unin-
terrupted premaxillary toothpad, much larger body size
(maximum TL 270 mm vs. 135 mm in S. petricola, 100
mm in S. lucipinnis), an occipito-nuchal shield which is
not covered by skin, and skin which is covered by granular
papillae.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Description. Morphometric and meristic data in
Table 5. Maximum TL 270 mm, SL 216 mm. Body
compressed laterally. Predorsal profile straight, inter-
rupted by convex ridge formed by orbit of eye. Preanal
profile straight to gently convex. Skin on body forming
numerous vertical folds covered with granular papillae
extending onto all fins.
Head depressed and broad; skin covered with
granular papillae. Snout with bluntly rounded margin
when viewed laterally and dorsally. Anterior nostrils
tubular; posterior nostrils with semicircular flaps of skin
along anterior margin. Occipito-nuchal shield rugose,
not covered with skin; terminating posteriorly with nar-
row, rounded process on either side of dorsal spine; ven-
trally with narrow, bluntly pointed process that extends
to upper margin of humeral process on either side of
body. Eye dorsolateral; ovoid; horizontal axis longer.
Interorbital area flat.
Mouth subterminal; lips wide and papillate. Man-
dibular teeth 28-51, short, unicuspid; arranged in single
transverse row. Premaxillary toothpad uninterrupted;
primary, secondary and tertiary premaxillary teeth dis-
crete; numerous; arranged in 4, 4, and 2 irregular rows,
respectively.
Maxillary barbel with narrow basal membrane;
Figure 11. Synodontis granulosus, UF 160945, 192 mm TL, 149 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
lacking branches or crenelations; extending at least to
base of pectoral fin. Lateral mandibular barbel extend-
ing to point just short of anterior margin of pectoral girdle,
with 4-5 non-tuberculate branches; lacking secondary
branches. Medial mandibular barbel about 1/2 length of
lateral barbel; with 4-5 pairs of tuberculate branches;
lacking secondary branches.
Dorsal fin 11,7-8; posterior margin straight. Dor-
sal-fin spine long, striated, nearly straight, terminating in
short black filament; anterior margin smooth; posterior
margin with small serrations distally. Pectoral fin 1,7-8;
posterior margin straight. Pectoral-fin spine roughly
equal in length to dorsal-fin spine, striated, slightly curved,
terminating in short, black filament; anterior spine mar-
gin granulate in adults; many small antrorse serrations
along anterior margin in juveniles; posterior margin with
large retrorse serrations along entire length. Adipose
fin long, well developed, margin convex. Pelvic fin i,6-
7; located anterior to vertical through origin of adipose
fin; tip of appressed fin barely reaches base of anal fin.
Anal fin iii-v,7-9,i; posterior margin nearly straight; base
located ventral to adipose fin. Caudal fin i,7,8,i; forked;
lobes pointed.
Humeral process narrow, elongated, granulous; pos-
sessing distinct ridge on ventral margin; dorsal margin
concave; terminating in sharp point (Fig. 3A). Large,
dark-colored axillary pore just ventral to humeral pro-
cess. Gut 0.5 times body length (n = 1, UF 160945).
Hindgut chamber absent.
Coloration. Body slate-gray to olive brown
(Fig.11). Belly ranging from pale yellow to gray. Scat-
tered, small black spots present on juveniles. Maxillary
barbel white. Mandibular barbels white, bases some-
times with scattered dusky pigmentation. Iris copper
colored. Dorsal and pectoral-fin spines brown to black,
filaments black. All rayed fins with black triangles at
base, posterior margins white to yellowish in color. Both
lobes of caudal fin with black bar from base to tip of fin;
posterior margin of fin white to yellowish in color.
Distribution. Lake Tanganyika (Fig. 12); rare but
widely distributed.
Habitat. Littoral to benthic zones over shell, sand
and mud bottoms; to a maximum depth of 130 m (Coulter
199 l1a).
Diet. Almost nothing is known about the diet of
this species. Worthington and Ricardo (1936) reported a
single Limnothrissa sp. (Teleostei: Atherinidae) in the
stomach of one specimen. Many other Tanganyikan
species of Synodontis prey largely on mollusks and al-
gae scraped from rocky substrates. Presumably, S.
granulosus has a similar diet.
Reproduction. No information exists regarding
km
0 50 100
Figure 12. Known distribution of Synodontis
granulosus. T denotes type locality.
the reproduction of Synodontis granulosus.
Taxonomic Remarks. Synodontis granulosus
is sufficiently distinct that it is one of only two described
Tanganyikan species of Synodontis for which the tax-
onomy has remained unaltered since the original descrip-
tion (the other being S. multipunctatus). Minor differ-
ences in coloration pattern occur throughout its range
(mainly in the color of the fin margins), but they do not
appear to follow any geographic pattern.
Synodontis ilebrevis n. sp.
(Figs. 5D, 13, 14, 16; Tables 2, 6)
Holotype. UF 160942, TL 138 mm, SL 116 mm,
Lake Tanganyika at Chaitika, Zambia, via Pete
Hauschner, Tropical Fish Collector, coll. V.2005.
Paratypes. (8) UF 162562, TL 128-148 mm, SL
108-121 mm, collection data same as for holotype, (1)
SAIAB 78410, TL 141 mm, SL 118 mm, collection in-
formation same as for holotype, (5) MRAC 78-25-P-
20-24, TL 93-136 mm, SL 79-114 mm, Cape Chaitika, S.
Lake Tanganyika (Zambia); coll. P. Brichard, IV. 1978.
Diagnosis. -Axillary pore absent; mandibular teeth
50-66; body with moderately sized spots; fin spines dark;
8-9 pectoral-fin rays; black triangles on bases of all rayed
fins; eye 26.0-33.4% snout length; premaxillary toothpad
interrupted; secondary branches on medial mandibular
barbel present; occipito-nuchal shield covered with skin;
short, flat papillae on skin of body; short gut; hindgut
chamber absent; body depth 18.2-20.1% SL; maximum
TL 150 mm.
The lack of an axillary pore distinguishes
Synodontis ilebrevis from S. dhonti, S. grandiops, S.
granulosus, S. multipunctatus, S. petricola, and S.
tanganaicae. Synodontis ilebrevis is separated from
S. irsacae and S. lucipinnis and further separated from
S. dhonti and S. petricola by having a greater number
of mandibular teeth (50-66 vs. 15-29 in S. irsacae, 35-
51 in S. lucipinnis, 22 in S. dhonti, and 31-50 in S.
petricola). Synodontis ilebrevis can also be separated
from S. petricola and S. lucipinnis by having dark fin
spines (vs. white). The size of the spots on the body
distinguishes S. ilebrevis from S. granulosus, S.
melanostictus, and S. tanganaicae, and the black tri-
angle at the base of all the rayed fins further separates
S. ilebrevis from S. melanostictus and adults of S.
tanganaicae. Synodontis ilebrevis is further distin-
guished from S. dhonti, S. granulosus, S.
melanostictus, and S. tanganaicae in having the
occipito-nuchal shield covered by skin. Synodontis
ilebrevis differs from S. polli in having a short gut (0.8-
1.4 times TL vs. 4.0-5.5 times TL in S. polli) (Figs. 14,
15), no hindgut chamber (Fig. 14), slightly shallower body
(body depth 18.2-20.1% SL vs. 20.2-27.0 in S. polli),
short flat papillae on the skin (vs. villous papillae in S.
polli), and small, round, widely spaced spots on the skin
(vs. moderate to large, irregular, more closely spaced
spots in S. polli).
Description. Morphometric and meristic data in
Table 6. Maximum TL 150 mm, SL 120 mm. Body not
compressed. Predorsal profile slightly convex. Preanal
profile straight to slightly convex. Skin on body forming
numerous vertical folds; short, flat papillae present, giv-
ing skin pebbled texture.
Head slightly depressed and broad; skin covered
with papillae; papillae extend onto base of maxillary barbel
and onto paired fins. Snout with bluntly rounded margin
when viewed laterally and dorsally. Occipito-nuchal
shield covered with skin, terminating posteriorly with
wide, pointed process on either side of dorsal spine; ven-
trally with wide, rounded process that extends to upper
margin of the humeral process on either side of body.
Eyes dorsolateral; ovoid; horizontal axis longer. Interor-
bital area flat to slightly convex.
Mouth inferior; lips widened and papillate. Man-
dibular teeth 50-66, short, unicuspid; arranged in 8 short,
transverse rows. Premaxillary toothpad interrupted;
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
primary, secondary and tertiary premaxillary teeth dis-
crete; numerous; arranged in 3-4, 2, and 1-2 irregular
rows, respectively.
Maxillary barbel short; extending at least to base
of pectoral spine; small papillae at base; basal mem-
brane narrow. Lateral mandibular barbel extending to
point just past anterior margin of pectoral girdle; with 3-
6 short, weakly tuberculate branches; usually lacking
secondary branches. Medial mandibular barbel approxi-
mately 1/3 to 1/2 length of lateral barbel; with 3-5 pairs
of tuberculate branches; many secondary branches
present.
Dorsal fin 11,7; posterior margin straight to slightly
concave. Dorsal-fin spine short; striated; slightly curved,
terminating in short, dusky filament; anterior margin of
fin spine granulous, posterior margin with small serra-
tions distally. Pectoral fin 1,8-9; posterior margin straight
to broadly rounded. Pectoral-fin spine roughly equal in
length to dorsal-fin spine, striated, slightly curved, termi-
nating in short, dusky filament; anterior spine margin
granulous; posterior margin with large, retrorse serra-
tions along entire length. Adipose fin long, well devel-
oped, margin convex. Pelvic fin i,6; located at vertical
through adipose fin base; tip of appressed fin does not
reach base of anal fin. Anal fin iii-v,7-9,i; posterior mar-
gin rounded; base located at vertical through center of
adipose fin. Caudal fin i,7,8,i; forked; lobes rounded.
Humeral process triangular; granulous; poorly de-
veloped ridge on ventral margin; dorsal margin concave;
terminating in sharp point (Fig. 5D). Axillary pore ab-
sent. Gut short, 0.8-1.4 times body length (n = 8, UF
160942, 162562, SAIAB 78410). Hindgut chamber ab-
sent (Fig. 14).
Coloration. Dorsum olive brown, covered with
small, regularly shaped, widely spaced, black spots (Fig.
13). Belly lighter, with smaller, regularly shaped, black
spots. Maxillary and mandibular barbels white. Iris
copper colored. Dorsal and pectoral-fin spines dark,
terminating in short, dusky filaments. Pectoral spine with
thin, light stripe along anterior margin. All rayed fins
with black triangles at their base; posterior margins dusky.
Triangles may be completely solid or composed of closely
spaced spots. Both lobes of caudal fin with black bars
from base to tip of fin; posterior margin dusky.
Distribution. Lake Tanganyika (Fig. 16). Known
only from the Cape Chaitika area; local abundance un-
known.
Habitat. The exact habitat of this species is un-
known. Like the other small-bodied species of
Synodontis, S. ilebrevis probably inhabits fairly shal-
low, rocky coastal areas.
Diet. Most of the specimens examined were
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 6. Morphometric and meristic counts for Synodontis ilebrevis. All morphometrics given in percent of base
measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n= 15) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
19.2
29.6
36.5
43.6
13.4
17.8
20.3
83.5
67.9
37.1
55.4
18.5
35.0
36.7
54.1
45.1
32.9
19.3
58.5
60.2
68.6
33.4
63.1
66.2
52.9
50.4
163.2
101.3
18.2-20.1
27.5-30.6
34.9-38.3
31.2-44.8
12.9-26.4
16.1-22.3
17.6-23.8
78.2-86.6
63.6-73.8
36.9-46.4
53.4-59.2
14.5-18.5
33.6-40.3
34.0-36.9
49.0-57.0
44.0-92.3
32.4-53.9
16.2-26.4
34.7-63.1
55.1-74.9
59.2-79.1
26.0-33.4
61.4-73.8
57.4-68.0
38.2-53.0
42.5-52.6
149.0-228.0
94.7-117.0
19.30.6
29.21.0
36.31.1
38.44.0
18.23.8
18.61.8
21.01.9
82.92.6
68.43.0
40.92.6
56.01.6
17.11.1
36.51.7
35.51.0
52.02.2
62.514.0
43.15.8
20.82.9
49.29.1
63.65.8
71.96.6
30.62.3
65.23.2
63.43.0
47.04.3
48.32.7
169.520.4
107.07.4
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
50(1); 52(1); 54(1); 55(1); 57(1);
58(3); 59(2); 60(2); 62(1); 64(1);
66*(1)
II,7*(15)
1,8(10); I,9*(5)
i,6*(15)
iii,7(1); iv,7(1); iv,7,i(2); iv,8(5);
iv,8,i*(3); iv,9(l); iv,9,i(1); v,9(1)
i,7,8,i*(10)
35*(4); 36(6)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 13. Holotype of Synodontis ilebrevis, UF 160942, 139 mm TL, 116 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 14. Synodontis ilebrevis, UF 162562, 138 mm TL, 118 mm SL, with intestine extracted. Note the
relative shortness of the intestine and lack of a discernible hindgut chamber.
Figure 15. Synodontis polli, MRAC 131007-008, 137
TL, 112 SL, with intestine extracted. Hindgut cham-
ber indicated with white arrow.
km
0 50 100oo
Figure 16. Known distribution of Synodontis ilebrevis.
T denotes type locality.
obtained through the aquarium trade and had not been
feeding on their natural diet. Though these specimens
readily ingested algal matter, their short gut and lack of
a hindgut chamber suggest that this species is suited to a
more carnivorous diet. Examination of the stomach con-
tents of freshly caught specimens is needed to deter-
mine differences in diet between this species and
Synodontis polli, which may have led to differences in
the gut morphology of these species.
Reproduction. No information exists regarding
the reproduction of Synodontis ilebrevis.
Etymology. The specific name is a combination
of the Latin ile, meaning intestine, and the Latin brevis,
meaning short, a reference to the relatively short gut of
this species. Gender: feminine.
Synodontis irsacae Matthes, 1959
(Figs. 5B, 17, 18; Tables 2, 7)
Synodontis irsacae Matthes, 1959:78, description, Lake
Tanganyika; 1962:37, placed into synonymy of S.
dhonti.
Synodontis dhonti. Matthes, 1962:37, fig. a, pl. 3,
description, diet, Lake Tanganyika. Poll, 1971:364,
description. Brichard, 1978:423, key; 1989:479, key.
Sands, 1983:23, checklist. Daget, Gosse, & Thys
van den Audenaerde, 1986:125, check-list. Burgess,
1989:195, checklist. Kobayagawa, 1989:14, photo
(misidentified). Coulter, 1991 a: 182, 191, table 8.16,
habitat, diet, reproduction.
Material Examined. Holotype, MRAC 130315,
TL 142 mm, SL 116 mm, Kalundu, N. Lake Tanganyika,
coll. H. Matthes, 27.X.1959, (6) paratypes, MRAC
130316-321, TL 55-156 mm, SL 46-124 mm, Kalundu,
N. Lake Tanganyika, coll. H. Matthes, 14.IV.1960, (6)
MRAC 130332-130337, TL 47-102 mm, SL 38-84 mm,
Luhanga, Lake Tanganyika, coll. H. Matthes
(I.R.S.A.C.), IV.1959, (1) MRAC 77-40-P-5-7, TL 99
mm, SL 80 mm, Chipimbi, S.W. Coast of Lake Tanganyika
(Zambia), coll. P. Brichard, 6.VII. 1977, (2) MRAC 92-
081 -P-0127-0128, TL 83-105 mm, SL 69-85 mm, Ulwile
Island, northern shore, Lake Tanganyika S727.40'
E30024.20' (Tanzania), coll. Exp. Tanganyika 1992,
27.V. 1992, (2) MRAC 95-096-P-2561-2562, TL 88-107
mm, SL 71-85 mm, Kasenga Point, Zambia, 0843.31 'S,
31008.01'E, coll. Verheyen, Snoeks, Hanssens, Ruber,
Stermbauer, 4.IV. 1995, (4) MRAC A3-033-P-0036-0040,
TL 117-161mm, SL 91-133 mm, Pemba, S. of Uvira,
Lake Tanganyika (Zaire), coll. L. De Vos, 18.VIII. 1995,
(1) MRAC Al-094-P-0055-0059, TL 103 mm, SL 81
mm, Katoto, Lake Tanganyika (Zambia), coll. Snoeks,
Hanssens, Verheyen et al., 18.X.2001, (1) BMNH
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
1955.12.20.1859, TL 190 mm, SL 157 mm, Albertville,
Lulumduie village, Lake Tanganyika, coll. Inst. Roy. Sci.
Nat. Beige, 28-29.XI. 1946, (1) BMNH 2005.9.26.3, TL
90 mm, SL 72 mm, Mpulungu, Zambia, Lake Tanganyika,
coll. J. Day, 2005, (1) BMNH 2005.9.26.17, TL 116 mm,
SL 94 mm, Mpulungu, Zambia, Lake Tanganyika, coll.
J. Day, 2005, (5) SAIAB 40177, TL 86-121 mm, SL70-
97 mm, Zambia, Lake Tanganyika, Mbita Island,
08045'00"S, 031006'00"E, 08.VIII.1992, (2) SAIAB
56687, TL 94-97 mm, SL 75-79 mm, Tanzania, Lake
Tanganyika; Muzungu beach, 04054'53"S, 029035'58"E,
10.X. 1997, (2) SAIAB 76105, TL 93-98 mm, SL 76-80
mm, Zambia, Mbala, Lake Tanganyika; Mbita Island
(northwest end), 08'45.18'S, 31 '05.07'E, coll. 29.11.2004,
(1) SAIAB 76174, TL 97 mm, SL 79 mm, Zambia, Mbala,
Lake Tanganyika; Musende Rocks beach, 0845.18'S,
31005.07'E, coll. 29.II.2004, (1) SAIAB 77883, TL 133
mm, SL 108 mm, Cave, Kigoma hotel below Hill top,
04o53'03"S, 029037' 11"E, 04.X. 1997, (1) SAIAB 77892,
TL 91 mm, TL 72 mm, Tanzania; Jacobsen's Beach,
0454'31"S, 02936'02"E, 05.X.1997, (2) UF 160940,
TL 104-124 mm, SL 86-102 mm, Cape Chaitika, Lake
Tanganyika, Zambia, via Pete Hauschner, Tropical Fish
Collector, V.2005, (2) CU 88750, TL 102-121 mm, SL
85-102 mm, Tanzania; Kigoma; Jacobsen's Beach, coll.
P.B. McIntyre, 18.VII.2002.
Diagnosis. -Axillary pore absent; mandibular teeth
15-29; body with large spots; fin spines dark; 8-9 pecto-
ral-fin rays; black triangles at bases of all rayed fins;
eye 28.7-43.7% snout length; premaxillary toothpad in-
terrupted; secondary branches on medial mandibular
barbel present; occipito-nuchal shield covered with skin;
skin of body smooth; hindgut chamber absent; maximum
TL 190 mm.
The absence of an axillary pore distinguishes
Synodontis irsacae from S. dhonti, S. grandiops, S.
granulosus, S. multipunctatus, S. petricola, and S.
tanganaicae. Synodontis irsacae can be further sepa-
rated from S. dhonti, S. grandiops, S. granulosus, and
S. multipunctatus by the interrupted premaxillary
toothpad (vs. uninterrupted). The presence of second-
ary branches on the medial mandibular barbel further
distinguishes S. irsacae from S. grandiops, S.
granulosus, and S. multipunctatus. Synodontis
irsacae differs further from S. dhonti, S. granulosus,
S. melanostictus, and S. tanganaicae in having an
occipito-nuchal process which is covered by skin.
Synodontis irsacae can be distinguished from S.
ilebrevis, S. lucipinnis, S. polli, and further separated
from S. petricola, by having a lower number of man-
dibular teeth (15-29 in S. irsacae vs. 50-66 in S. ilebrevis,
35-51 in S. lucipinnis, 40-70 in S. polli, and 31-50 in S.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 7. Morphometrics and meristic counts for Synodontis irsacae. All morphometrics given in percent of base
measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n=42) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
28.6
30.4
38.1
32.3
26.8
24.4
25.3
86.8
82.3
46.3
51.1
17.3
44.3
37.3
37.4
88.1
49.5
20.9
42.2
80.3
83.2
33.8
86.7
73.1
39.0
46.2
327.8
81.5
17.7-28.6
27.9-32.8
36.7-41.8
26.9-41.5
16.1-28.4
18.8-28.0
19.0-27.5
75.9-97.1
59.1-82.3
35.7-55.8
50.4-58.2
16.0-22.6
27.3-44.3
29.8-41.0
33.7-52.8
52.4 -91.8
38.2-63.0
14.4-30.7
28.6-59.6
59.6-88.3
60.6-88.7
28.7-43.7
50.9-86.7
56.0-81.2
39.0-72.7
41.8-69.3
180.3-358.2
60.4-128.8
23.52.5
31.01.0
39.41.3
33.33.3
22.03.1
23.42.1
23.71.6
82.84.1
69.85.4
47.54.3
53.52.1
18.72.0
34.73.6
35.22.5
40.94.3
71.09.9
46.96.2
21.73.5
46.56.6
75.46.6
76.76.0
35.14.0
65.07.5
66.06.3
54.88.9
53.77.8
280.739.4
94.715.0
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
15(1); 16(2); 18(1); 19(1); 21(5);
22*(8); 23(5); 24(5); 25(4); 26(5);
27(4); 29(1)
II,7*(42)
1,8* (31), 1,9 (11),
i,6*(42)
iii,7(l1); iii,8(2); iii,8,i(2); iii,9(l1);
iv,7(10); iv,7,i(4); iv,8*(22)
i,7,8,i(5)
34(3); 35(2)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 17. Holotype of Synodontis irsacae, MRAC 130315, 142 mm TL, 116 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
28' 30'
Figure 18. Known distribution of Sy
T denotes type locality.
petricola). Synodontis irsacae is f
from S. petricola and S. lucipinnis ii
spines (vs. white fin spines). Black tria
fins of S. irsacae and the lack of pal
(vs. villous papillae) further separate
S. melanostictus, and adult S. tanga
Description. Morphometric an
Table 7. Maximum TL 190 mm, SL 1:
compressed. Predorsal profile slightly
profile straight to convex. Skin on bod
ous vertical folds; papillae absent.
Head slightly depressed; broad;
granular papillae; papillae extend onto
barbel and anterior portion of body c
with bluntly rounded margin when vie
dorsally. Occipito-nuchal shield cove
minating posteriorly with narrow, blunt
on either side of dorsal spine; veni
rounded process that extends to upp
meral process on either side of body. 1
ovoid; horizontal axis longer. Interort
Mouth inferior; lips wide and pa]
lar teeth 15-29, short, unicuspid; arrange
verse row. Premaxillary toothpad inte
secondary and tertiary premaxillary t
32' merous; arranged in 3, 1-2, and 1 irregular rows, re-
S/ spectively.
Maxillary barbel with distinct basal membrane;
extending to, or just past base of pectoral fin; granular
-1 ( papillae on base and along anterior margin. Lateral
mandibular barbel extending to point just short of ante-
rior margin of pectoral girdle, with 2-7 short, tubercu-
si1 R late branches; secondary branches sometimes present.
Medial mandibular barbel about half length of lateral
barbel; with 4-5 pairs of tuberculate branches; second-
anzania ary branches present.
Dorsal fin 11,7; posterior margin straight. Dorsal-
fin spine short; striated, slightly curved, terminating in
short, dark filament; anterior margin of fin spine smooth
in small specimens; becoming granulous with growth;
posterior margin with small serrations distally. Pectoral
i\ kam fin 1,8-9; posterior margin straight to broadly rounded.
_8' Pectoral-fin spine roughly equal in length to dorsal-fin
spine; striated, slightly curved, terminating in short, dark
ia filament; anterior spine margin granulous in adult speci-
ambia mens; smooth in juvenile specimens; posterior margin
km 32 with large retrorse serrations along entire length. Adi-
0 5o 10o pose fin long, well developed, margin convex. Pelvic fin
nodontis irsacae. i,6; located at vertical midway between posterior base
of dorsal fin and origin of adipose fins; tip of appressed
fin barely reaches base of anal fin. Anal fin iii-v,7-9;
posterior margin rounded; base located ventral to ante-
'urther separated rior third of adipose fin. Caudal fin i,7,8,i; forked; lobes
i having dark fin rounded.
ngles on the rayed Humeral process narrow, elongated, granulous;
pillae on the skin weakly developed ridge on ventral margin; dorsal mar-
this species from gin concave; terminating in rounded point (Fig. 5B).
naicae. Axillary pore absent. Gut 0.7-0.8 times body length (n =
d meristic data in 5, MRAC 130315, MRAC A3-033-P-0036-0040, UF
57 mm. Body not 160941). Hindgut chamber absent.
convex. Preanal Coloration. Dorsum grayish to cuprous brown
y forming numer- (Fig. 17). Belly coloration slightly lighter than dorsum.
Scattered, irregularly-shaped black spots present on ju-
skin covered with venile specimens. Spots becoming slightly smaller in
base of maxillary large specimens. Maxillary barbel white. Mandibular
nly. Snout long, barbels white, bases sometimes with scattered dusky
*wed laterally and pigmentation. Iris copper colored. Dorsal and pectoral-
red with skin; ter- fin spines dark, terminating in short, light to dusky col-
ly pointed process ored filaments. Anterior margin and venter of pectoral
rally with wide, fin light colored. All rayed fins with black triangles at
er margin of hu- their base in young specimens; posterior margins white
Eyes dorsolateral; in color. Both lobes of caudal fin black, posterior margin
aital area flat. white.
pillate. Mandibu-
ged in single trans-
errupted; primary,
eeth discrete; nu-
Distribution. Lake Tanganyika (Fig. 18); com-
mon and widely distributed.
Habitat. Littoral to benthic zones over shell, sand
and mud bottoms, to a maximum depth of 40 m (Coulter
1991 a).
Diet. Synodontis irsacae is omnivorous, feed-
ing on algae, sponges, ostracods, small crabs, insect lar-
vae, and fish eggs (Matthes 1959; Coulter 1991a).
Reproduction. -As with most of the other species
of Synodontis in the lake, information regarding the re-
production of S. irsacae is nonexistent.
Taxonomic Remarks. Originally described by
Matthes (1959), Synodontis irsacae was quickly placed
in the synonymy of S. dhonti Boulenger (Matthes 1962).
Specimens of S. irsacae were thought to be juvenile
individuals of S. dhonti, a species for which the holo-
type was the only known specimen. A number of onto-
genetic changes in external morphology and morpho-
metric ratios were proposed to account for the differ-
ences between specimens of S. irsacae and the holo-
type of S. dhonti. A list of these changes, given by
Matthes (1962), follows (translation by first author):
* The relative length of the snout increases.
* The relative size of the eye diminishes.
* The head and the body become more squat.
* The bone structure of the cranium, the occipito-nuchal
shield and the humeral process becomes rugose and more
strongly developed, causing a general increase in the
relative size of the head.
* The humeral process becomes more blunt.
* The number of mandibular teeth increases.
* The band of premaxillary teeth becomes wider.
* The dorsal- and pectoral-fin spines become relatively
shorter, wider and more rugose, while the anterior ser-
rations become obscured.
Examination of material identified as S. dhonti and
S. irsacae has revealed additional differences between
the S. dhonti holotype and specimens recognized herein
as S. irsacae.
* A large, conspicuous axillary pore is present in the
holotype of S. dhonti, while this structure is lacking in
S. irsacae.
* The premaxillary toothpad of S. irsacae is interrupted,
while that of the holotype of S. dhonti is uninterrupted.
* The number of primary and secondary premaxillary
tooth rows is different between S. irsacae and the ho-
lotype of S. dhonti (three and two vs. four and five,
respectively).
* No papillae are present on the body skin of S.
irsacae, while granulous papillae cover the body of the
S. dhonti holotype.
* The holotype of S. dhonti has 25 elongated gill rakers
on the first branchial arch; 10 specimens of S. irsacae
showed a range of 12-16 gill rakers, none of which was
conspicuously elongated.
In light of these new findings, Synodontis irsacae
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
is recognized herein as a distinct species from S. dhonti.
The differences between the S. dhonti holotype and
the S. irsacae specimens examined in this study do not
vary with ontogeny in other Tanganyikan species of
Synodontis, and there is no reason to believe that they
do in S. irsacae. There have been no specimens col-
lected that appear to be intermediate in size and appear-
ance between S. dhonti and S. irsacae that might lend
support to the idea that specimens of S. irsacae are
juvenile S. dhonti. A specimen cited by Matthes (1962)
as being intermediate between the two species (MRAC
90279) is actually a specimen of S. tanganaicae.
Synodontis lucipinnis n. sp.
(Figs. 4A, 19, 20; Tables 2, 8)
Holotype. SAIAB 77880, TL 97 mm, SL 78
mm, Zambia, Lake Tanganyika; Musende Rocks,
08046'00"S, 031 07'00"E, 10.X. 1992.
Paratypes. (10) SAIAB 39577, TL 58-98 mm,
SL 47-79 mm, Zambia, Mpulungu, Musende Rocks,
08046'00"S, 031051'00"E, 07.VII.1992, (3) SAIAB
77879, TL 70-85 mm, SL 56-69 mm, Zambia, Musende
Rocks, 08'46'00"S, 031 06'00"E, 05.VIII.1992, (4)
SAIAB 77882, TL 55-97 mm, SL 44-78 mm, Zambia,
Lake Tanganyika; Musende Rocks, 08046'00"S,
031007'00"E, 10.X.1992, (1) SAIAB 77894, TL 88 mm,
SL 73 mm, Zambia, Mbala, Lake Tanganyika, Mbita Is-
land (northwest end), 0845.18'S, 03105.07'E,
29.11.2004.
Diagnosis. -Axillary pore absent; mandibular teeth
35-51; body with large spots; fin spines white; 8-9 pec-
toral-fin rays; black triangles on bases of all rayed fins
with light colored window at base (except caudal fin);
eye 25.2-35.8% snout length; premaxillary toothpad in-
terrupted; secondary branches on medial mandibular
barbel present; occipito-nuchal shield covered with skin;
papillae on skin of body absent; maximum TL 100 mm.
The lack of an axillary pore distinguishes
Synodontis lucipinnis from S. dhonti, S. grandiops,
S. granulosus, S. multipunctatus, S. petricola, and S.
tanganaicae. Synodontis lucipinnis is further sepa-
rated from S. petricola by the presence of light colored
windows at the bases of the rayed fins (with the excep-
tion of the caudal fin). Morphometric measurements
vary little between these two species. Synodontis
lucipinnis differs from S. dhonti, S. granulosus, S.
ilebrevis, S. irsacae, S. melanostictus, S. polli, and S.
tanganaicae in having white fin spines (vs. brown to
black) and also differs from all of these species (with
the exception of S. irsacae) in lacking papillae on the
skin of the body. The occipito-nuchal shield being cov-
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 8. Morphometrics and meristic counts for S. lucipinnis. All morphometrics given in percent of base measure-
ment. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (N= 18) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
22.2
27.6
36.5
35.5
27.5
18.7
17.2
84.4
64.5
48.0
54.6
18.3
33.1
33.6
57.2
99.5
50.0
24.4
48.7
67.7
62.0
33.4
60.5
61.5
55.3
54.4
262.8
94.2
15.7-23.9
27.6-31.2
35.7-40.3
27.6-38.7
20.7-27.5
17.6-22.8
17.2-24.2
76.0-86.1
55.0-64.5
41.5-50.1
51.4-55.0
13.9-18.5
28.5-37.1
33.6-38.6
41.5-57.2
68.5-99.5
42.6-60.4
19.3-28.3
42.4-61.8
59.2-76.6
62.0-79.8
25.2-35.8
53.8-71.9
61.5-75.1
43.9-58.5
39.5-54.4
213.7-319.1
87.1-141.4
20.02.7
29.81.0
38.11.4
32.62.9
23.01.9
19.61.4
21.31.8
82.92.7
59.62.5
46.02.5
53.01.1
16.61.3
32.02.3
35.91.3
48.83.9
77.47.7
49.84.7
24.12.4
48.65.3
66.24.6
71.44.9
31.42.7
60.54.8
67.93.4
52.04.6
46.23.8
262.824.3
104.415.7
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
35(2); 36(3); 38(4); 39(2); 41(2);
43(1); 44(1); 47*(2); 51(1)
H,7*(17); II,7,i(1)
1,8*(17); 1,9(1)
i,6*(18)
iv,8*(12); iv,8,i(2); iv, 9(4)
i,7,8,i(10)
34(3); 35(6)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 19. Holotype of Synodontis lucipinnis, SAIAB 77880, TL 97 mm, SL 78 mm.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
ered with skin further separates S. lucipinnis from S.
dhonti, S. granulosus, S. melanostictus, and S.
tanganaicae, all of which also have a much larger maxi-
mum TL (395, 270, 520, and 585 respectively vs. 100
mm in S. lucipinnis). The black triangles at the base of
all the rayed fins further distinguish S. lucipinnis from
S. melanostictus, while the windows at the base of the
triangles further differentiate S. lucipinnis from S.
dhonti, S. grandiops, S. granulosus, S. ilebrevis, S.
irsacae, S. multipunctatus, S. polli, and S.
tanganaicae. Synodontis lucipinnis can be further
separated from S. dhonti, S. grandiops, S. granulosus,
and S. multipunctatus by having an interrupted premax-
illary toothpad (vs. uninterrupted).
Description. Morphometric and meristic data in
Table 8. Maximum TL 100 mm, SL 80 mm. Body not
compressed. Predorsal profile slightly convex. Preanal
profile slightly convex. Skin on body forming numerous
vertical folds; papillae absent.
Head slightly depressed and broad; skin covered
with villous papillae; papillae extend onto base of maxil-
lary barbel, pectoral fin and anterior portion of body only.
Snout with bluntly rounded margin when viewed later-
ally and dorsally. Occipito-nuchal shield covered with
skin, terminating posteriorly with wide, pointed process
on either side of dorsal spine, ventrally with wide,
rounded process that extends to upper margin of the
humeral process on either side of body. Eyes dorsolat-
eral; ovoid; horizontal axis longer. Interorbital area flat
to slightly convex.
Mouth inferior; lips widened and papillate. Man-
dibular teeth 35-51, short, unicuspid; arranged in 6 short,
transverse rows. Premaxillary toothpad interrupted;
primary, secondary and tertiary premaxillary teeth dis-
crete, numerous, arranged in 3, 3, and 1 irregular rows,
respectively.
Maxillary barbel short; extending at least to base
of pectoral spine; small papillae at base; basal mem-
brane narrow. Lateral mandibular barbel extending to
point just past anterior margin of pectoral girdle; with 4-
6 short, simple, weakly tuberculate branches; usually
lacking secondary branches. Medial mandibular barbel
approximately 1/3 to 1/2 length of lateral barbel; with 4-
6 pairs of tuberculate branches; many secondary
branches present.
Dorsal fin II,7-7,i; posterior margin straight to
slightly concave. Dorsal-fin spine short, striated, slightly
curved, terminating in a short, white filament; anterior
margin of fin spine smooth; posterior margin with small
serrations distally. Pectoral fin 1,8-9; posterior margin
broadly rounded. Pectoral-fin spine roughly equal in
length to dorsal-fin spine, striated, slightly curved termi-
28" 30" 32'
km
0 so 100
Figure 20. Known distribution of Synodontis lucipinnis.
T denotes type locality.
nating in short, white filament; anterior spine margin
smooth; posterior margin with large, retrorse serrations
along entire length. Adipose fin long, well developed,
margin convex. Pelvic fin i,6; located at vertical mid-
way between posterior base of dorsal fin and origin of
adipose fin; tip of appressed fin does not reach base of
anal fin. Anal fin iv,8-9; posterior margin rounded; base
located at vertical through center of adipose fin. Caudal
fin i,7,8,i; forked; lobes rounded.
Humeral process triangular; granulous; covered
with many small, villous papillae; poorly-developed ridge
on ventral margin; dorsal margin convex; terminating in
a sharp point (Fig. 4A). Axillary pore absent.
Coloration in alcohol. Dorsum yellowish to cu-
prous brown, covered with large, irregularly shaped,
black spots (Fig. 19). Belly lighter, with more regularly
shaped black spots. Maxillary and mandibular barbels
white. Iris copper colored. Dorsal and pectoral-fin spines
white, terminating in short, white filaments. All rayed
fins with black triangles at base; posterior margins white.
Triangles have large, lightly colored windows at bases,
most noticeable in dorsal and anal fins. Both lobes of
caudal fin with black bar from base to tip of fin; poste-
rior margin of fin white.
Distribution. Lake Tanganyika (Fig. 20). Known
only from the Musende Rocks area (Mpulungu), local
abundance unknown.
Habitat. Rocky, coastal areas. Maximum depth
unknown.
Diet. Unknown.
Reproduction. No information exists regarding
the reproduction of Synodontis lucipinnis.
Etymology. The specific name is a combination
of the Latin luci, meaning bright or clear, and the Latin
pinnis, meaning fin, a reference to the light patches found
at the base of the black triangles found on the rayed fins
of this species. Gender: feminine.
Synodontis melanostictus Boulenger, 1906
(Figs. 4B, 21, 22; Tables 2, 9)
Synodontis melanostictus Boulenger, 1906:553, pl. 34,
description, Lofu (Lake Tanganyika); 1911:418, fig.
314, description, Lofu (Lake Tanganyika); 1916:316,
note. Pappenheim and Boulenger, 1914:251, note.
Poll, 1946:220, description, Lake Tanganyika. -
Worthington andRicardo, 1933:1067,1077,1101, note,
Lake Tanganyika. Ricardo-Bertram, 1940:209, first
placed in synonymy of S. nigromaculatus. Matthes,
1962:41, table 2, comparison of specimens from Lake
Tanganyika and Mweru-Luapula, comparison with S.
serratus tanganaicae and S. lacustricolus. Sterba,
1963:408, fig. 610, Lake Tanganyika. Coulter, 1965-
1966:34, abundance at different depths, Lake
Tanganyika. Brichard, 1978:424, key.
Synodontis nigromaculatus. Brichard, 1989:479, key,
478, 480, 483, photos. Burgess, 1989:196, check-
list, pl. 78, 82, 83, 94.- Coulter, 1991a:152, 154, 156,
181,191, abundance, habitat, reproduction; 1991b:287,
phylogenetic relationship to Tanganyikan endemics.
Material Examined. Holotype, BMNH
1906.9.8.72, TL 290 mm, SL 237 mm, Lake Tanganyika,
coll. Cunnington, (4) MRAC 100515-100518, TL 183-
237 mm, SL 138-188 mm, Petite-Ruzizi, coll. G. Marlier
and N. Leleup, V-VII. 1955, (2) MRAC 126293-294, TL
152-172 mm, SL 122-134 mm, Uvira, Lake Tanganyika,
coll. G. Marlier, 13.IX. 1958, (1) MRAC 96-083-P-0002,
SL 199 mm, Lufubu River, lower course, beginning of
Yendwe Valley escarpment, Lake Tanganyika basin
(Zambia), coll. L. De Vos, 26.X.1995.
Comparative Material Examined. Holotype, S.
nigromaculatus, BMNH 1905.11.10.10, TL 237 mm,
SL 186 mm, Lake Bangwuelo, Zambia, coll. Esq. R.
Melland, (1) S. nigromculatus, BMNH 1908.11.6.29,
TL 244 mm, SL 177 mm, Maramba River, Upper
Zambezi, coll. G.T. Corrington, (1) S. nigromaculatus,
BMNH 1907.9.30.9, TL 319 mm, SL 238 mm, Lake
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Bangwelo, coll. R.H. Melland, (4) S. nigromaculatus,
BMNH 1920.5.26.95-102, TL58-115 mm, SL46-93 mm,
Cape Kasangeneke, Lake Mweru, coll. L. Stappers, (6)
S. nigromaculatus, BMNH 1943.7.27.414-421, TL 176-
271 mm, SL 133-208 mm, River Chambezi, Bangweulu
Swamps, and Lake Bangweulu; coll. C. Ricardo et al.,
(1) S. nigromaculatus, BMNH 1976.3.18: 2439-2440,
TL 192 mm, SL 153 mm, Thalamakane River at Maun
(Botswana), coll. K. Berne, (1) S. nigromaculatus,
MRAC 34006, TL 383 mm, SL 307 mm, Lukonzolwa,
Lake Moero, coll. DeWitte, 9-17.11.1931, (1) S.
nigromaculatus, MRAC 34007, TL 324 mm, SL 250
mm, Lukonzolwa, Lake Moero, coll. DeWitte, 9-
17.II.1931, (2) S. nigromaculatus, MRAC 37814-815,
TL 177-317 mm, SL 134-252 mm, Kasenga, Luapula
River, coll. DeWitte, 10.III.1931, (1) S. nigromaculatus,
MRAC 56319-56320, TL 132 mm, SL 104 mm, Luapula
au Bangwelo, coll. Breolo, II1.1938, (2) S.
nigromaculatus, MRAC 162207-208, TL 86-114 mm,
SL 62-87 mm, Dundo, afflt. Luachimo River called
"Musapa wa Kamakenza" by indigenous peoples
(Angola), coll. Barros Machado (Dundo Museum),
2.VI.1949, (1) S. nigromaculatus, MRAC 87132-133,
TL 154 mm, SL 129 mm, Kasaji, Katanga, coll. Rev. S.
Fischer, 5.XII.1952, (1) S. nigromaculatus, MRAC
162219, TL 183 mm, SL 141 mm, Lake Calundo, coll.
indigenous peoples, 1.1955, (2) S. nigromaculatus,
MRAC 162210-213, TL 274-306 mm, SL 209-229 mm,
Chiumbe River, near the junction with the Chilambo River
(S745', E2105') (Angola), coll. Barros Machado
(Dundo Museum), 24.XI. 1957, (1) S. nigromaculatus,
MRAC 149537, TL 181 mm, SL 142 mm, Luwohoshi
River, affl. Ruashi River, village of Shindaika, Katanga,
coll. M. Lips, 25.111.1963, (1) S. nigromaculatus,
MRAC 165806-807, TL 104 mm, SL 80 mm, Chambeshi
River (Zambia), coll. H. Matthes, 16.VI.1965, (1) S.
nigromaculatus, MRAC 172442, TL 177 mm, SL 134
mm, Lake Bangwelo, coll. Bell-Cross, (1) S.
nigromaculatus, MRAC 94-019-P-1007, TL 152 mm,
SL 116 mm, South Mofwe Lagoon, Luapula Swamp
(Zambia), coll. P. Van Zweiten, 16.1I.1993-26.111.1993,
(1) S. nigromaculatus, MRAC 94-019-P-2136, TL 208
mm, SL 158 mm, Pembe lagoon, near Katotoma, south
lagoon, Luapula swamps (Zambia), coll. P. Van Zweiten,
17.VIII.1993, (1) S. nigromaculatus, MRAC 162203-
204 (orig. I.D. S. zambesensis), TL 180 mm, SL 132
mm, Nhefo River, afflt. The river to the left of the
Luachimo, close to 50 km S. of Dundo (Angola), coll.
Barros Machado (Dundo Museum), 111.1948, (1) S.
nigromaculatus, MRAC 162206 (orig. I.D. S.
zambesensis), TL 241 mm, SL 186 mm, Dundo,
Luachimo River (Angola), coll. Barros Machado (Dundo
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Museum), XI.1948, (1) S. nigromaculatus, MRAC
162214 (orig. I.D. S. zambesensis), TL 265 mm, SL 196
mm, Dundo, Luachimo River, barrage (Angola), coll.
Casalescol (Dundo Museum), 12.V.1960, (1) S.
nigromaculatus, MRAC 162215 (orig. I.D. S.
zambesensis), TL 180 mm, SL 136 mm, Cachimo,
Tuembe River, S806', E21'29' (Angola), coll. A.L.
Terreira (Dundo Museum), 22.VIII.1961, (1) S.
nigromaculatus, MRAC 149538 (orig. I.D. S.
zambesensis), TL 160 mm, SL 122 mm, Moushoshi River
near the junction with the Kofubu River (Katanga), coll.
M. Lips (don. M. Mignolet), 6.V.1963.
Diagnosis. -Axillary pore absent; mandibular teeth
23-36; body with small spots; fin spines brown; 9-10
pectoral-fin rays; black triangles absent on all rayed fins;
eye 31.5-38.1 % of snout length; premaxillary toothpad
interrupted; secondary branches on medial mandibular
barbel absent; occipito-nuchal shield not covered by skin;
villous papillae present on skin of body; hindgut cham-
ber present; maximum TL 520 mm;
The only species of Synodontis not endemic to
the Lake Tanganyika basin, S. melanostictus lacks the
rayed fin coloration common to almost all of its
Tanganyikan congeners, having spotted fins which com-
pletely lack black triangles at their bases. The black
triangles on the fins of S. tanganaicae may become
indistinct in large specimens and the body may have small
black spots that resemble those found in S.
melanostictus. In these cases, mandibular tooth counts
(23-36 in S. melanostictus vs. 33-49 in S. tanganaicae),
differences in eye size (31.5-38.1% of snout length in S.
melanostictus vs. 16.0-26.9% in S. tanganaicae), the
villous papillae on the skin of S. melanostictus (vs. granu-
lar in S. tanganaicae), and differences in humeral pro-
cess shape (Fig. 4) distinguish these species from one
another. Synodontis melanostictus differs from S.
dhonti in having a greater number of mandibular teeth
(23-36 vs. 22), an interrupted premaxillary toothpad (vs.
uninterrupted), and small, black spots covering the body
(vs. spots absent).
Description. Morphometric and meristic data in
Table 9. Maximum TL 520 mm, maximum SL 425 mm.
Body compressed laterally. Predorsal profile straight
anterior to eye; slightly convex posterior to eye. Prea-
nal profile straight. Skin on body lacking vertical folds;
covered with villous papillae, not extending onto fins.
Head depressed and broad, skin covered with small,
granular papillae. Snout with subconical margin when
viewed laterally, bluntly pointed margin when viewed
dorsally. Occipito-nuchal shield rugose, not covered with
skin; terminating posteriorly with narrow, rounded pro-
cess on either side of dorsal spine; ventrally with wide,
rounded process that extends to upper margin of hu-
meral process on either side of body. Eyes dorsolateral,
ovoid, horizontal axis longer. Interorbital area flat to
slightly convex.
Mouth subterminal; lips wide and papillate. Man-
dibular teeth 23-36, short, unicuspid; arranged in single
transverse row. Premaxillary toothpad interrupted; pri-
mary, secondary, and tertiary premaxillary teeth discrete;
primary premaxillary teeth in 2-3 rows; secondary pre-
maxillary teeth in 2-3 rows; tertiary premaxillary teeth
in 1-2 rows.
Maxillary barbel with narrow basal membrane;
lacking branches or crenelations; extending at least to
the base of the pectoral fin. Lateral mandibular barbel
extending to point just short of anterior margin of pecto-
ral girdle; with 7-8 long, non-tuberculate branches; short
secondary branches present. Medial mandibular barbel
about half the length of the lateral barbel; with 4-5 pairs
of non-tuberculate branches; secondary branches
present.
Dorsal fin 11,7; posterior margin concave. Dorsal
fin-spine long, striated, slightly curved, terminating in short
black filament; anterior margin of fin spine smooth; pos-
terior margin with small serrations distally. Pectoral fin
1,9-10; posterior margin straight. Pectoral-fin spine
roughly equal in length to dorsal-fin spine, striated, slightly
curved, terminating in a moderate length filament; ante-
rior spine margin with many fine, long, antrorse serra-
tions along anterior margin; posterior margin with large,
retrorse serrations along entire length. Adipose fin long,
well developed, margin convex. Pelvic fin i,6; located
anterior to vertical through posterior base of dorsal fin;
tip of appressed fin barely reaches base of anal fin. Anal
fin iii-v,7-9; posterior margin nearly straight; base located
ventral to first 1/3 of adipose fin. Caudal fin forked,
i,7,8,i.
Humeral process triangular; wide at base; taper-
ing to sharp point; striated along lower half; becoming
granulous along upper margin; possessing weakly de-
veloped ridge along ventral margin; dorsal margin slightly
concave (Fig 4B). Axillary pore absent. Gut 1.3-1.6
times body length (n=2, MRAC 100515-100518). Well
developed hindgut chamber.
Coloration in alcohol. Body grayish to cuprous
brown (Fig. 21). Belly pale. Scattered, small black spots
present, sometimes confluent. Maxillary and mandibu-
lar barbels white. Iris copper colored. Dorsal- and pec-
toral-fin spines brown, terminating in dusky filaments.
All rayed fins with small, black spots, similar to those
found on body.
Distribution. Lake Tanganyika and its tributaries
(Fig. 22).
Habitat. Littoral to benthic zones over shell, sand
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Table 9. Morphometric and meristic counts for Synodontis melanostictus. All morphometrics given in percent of
base measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n=8) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
Humeral process length/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
20.0
25.8
36.3
29.4
32.4
24.9
25.2
97.3
77.0
60.0
51.3
17.0
41.0
38.8
32.6
125.5
100.8
45.3
44.9
96.6
97.8
33.1
79.8
75.6
41.5
43.8
200.1
113.0
20.0-23.4
25.8-28.4
36.2-40.3
16.1-34.5
27.5-41.6
23.3-27.5
21.5-26.6
90.6-98.6
70.9-81.8
60.0-70.8
50.6-52.1
16.4-19.3
34.1-41.0
34.5-38.8
30.9-40.0
103.6-158.2
69.5-107.3
26.6-51.2
36.3-52.6
88.6-104.0
80.7-101.3
31.5-38.1
67.2-79.8
67.4-75.6
41.5-56.7
43.8-55.8
200.1-255.9
66.9-116.4
21.80.9
26.70.9
37.51.4
25.05.0
33.44.8
25.31.7
25.01.8
95.12.8
75.34.0
66.53.6
51.40.6
17.51.0
37.82.4
36.21.4
34.33.4
125.218.8
87.814.3
41.58.2
47.25.3
94.55.5
94.07.7
34.22.1
73.64.4
70.52.7
46.75.2
48.64.0
238.217.4
100.116.8
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
23(1); 27(3); 29(1); 31(1); 35(1);
36*(1)
II,7*(8)
1,9*(7); 1,10(1)
i,6*(8)
iii,9(2); iv,8,i(3); iv,9(2); v,9*(1)
i,7,8,i(5)
38(3); 39(2)
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 21. Top Holotype of S. melanostictus, BMNH 1906.9.8.72, 290 mm TL, 237 mm SL. Bottom Holo-
type of S. nigromaculatus, BMNH 1905.11.10.10, 237 mm TL, 186 mm SL.
and mud bottoms, to a maximum depth of 150 m (Coulter
1991a).
Diet. Nothing is known about the diet of this
species.
Reproduction. Coulter (1991) cites Matthes
(1962) as reporting that gravid females S. melanostictus
were found to contain up to 3,000 eggs. The latter ref-
erence, however, includes no egg counts or other infor-
mation on the reproduction of S. melanostictus.
Taxonomic Remarks. Synodontis nigro-
maculatus Boulenger was described from Lake
Bangweulu (Zambia) and has since been reported from
the Congo, Cunene, Limpopo, Luapula, Okovango, and
Zambezi Rivers, as well as Lakes Mweru and
Tanganyika (Boulenger 1905; Poll 1971). Some of these
populations (Congo and Limpopo Rivers, Lake
Tanganyika) have been described as new species or iden-
tified as other previously described species, only to be
synonymized with S. nigromaculatus in later works,
based on similarities in morphometric ratios (Boulenger
1923, Poll 1967, 1971). While the identity of specimens
from some drainages is questionable, we can be certain
that S. nigromaculatus occurs in Lake Bangweulu, and
probably also occurs in Lake Mweru and the Luapula
River, which joins the two lakes. Tanganyikan speci-
mens were therefore compared primarily to specimens
from these areas. These comparisons have shown that
Lake Tanganyikan specimens are diagnosible from S.
nigromaculatus.
Synodontis melanostictus Boulenger was de-
scribed from the Lofu River (now the Lofubu), a tribu-
tary of Lake Tanganyika, and was distinguished from S.
nigromaculatus by having numerous, villous papillae on
the body (Boulenger 1906). Ricardo-Bertram (1940)
28' 30- 32'
km
0 50 100
Figure 22. Known distribution of Synodontis
melanostictus. T denotes type locality.
first suggested placing S. melanostictus in the synonymy
of S. nigromaculatus, stating that the presence or ab-
sence of these papillae depended upon the condition and
preservation of a particular specimen. This appears to
be the case in the holotype of S. melanostictus, which
lacks any indication of papillae on its body. While the
papillae are not as diagnostic as was once thought, other
evidence has been found to support the distinctiveness
of S. melanostictus.
The interrupted condition of the premaxillary
toothpad of Synodontis melanostictus differentiates it
from S. nigromaculatus, in which the premaxillary
toothpad is uninterrupted. Vertebral counts also differ
between these species with S. melanostictus having 38-
39 total vertebrae (n=5, BMNH 1906.9.8.72, MRAC
100515-100518) vs. 36-37 in S. nigromaculatus (n=13,
BMNH 1905.11.10.10, 1920.5.26.95-102, 1943.7.27.414-
421, 1976.3.18: 2439-2440). Finally, although small dark
spots are present in both species, these spots are dis-
crete in S. melanostictus, while S. nigromaculatus al-
ways has some confluent spots (Fig. 21). Based upon
these differences, S. melanostictus is recognized herein
as a valid species that is endemic to the Lake Tanganyika
drainage.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Synodontis multipunctatus Boulenger, 1898
(Figs. 2, 3B, 8, 23, 24; Tables 1, 10)
Synodontis multipunctatus Boulenger, 1898:24, pl. 8,
description, Sumba (Lake Tanganyika); 1899:95, note,
Lake Tanganyika; 1901:313, pl. 15, description, Sumbu,
Kalambo, Moliro, Albertville; 1901 b: 149, Kalambo
(Lake Tanganyika); 1906:553, Niamkolo (Lake
Tanganyika); 1911:420, fig. 316, description, Sumba,
Kalambo, Niamkolo.- Moore, 1903:166, fig., descrip-
tion, Sumbu (Lake Tanganyika). Worthington and
Ricardo, 1933:1067,1077, 1101, note, diet, Usumbura,
Kigoma, Kala (Lake Tanganyika). Borodin, 1936:10,
Kasanga (Lake Tanganyika). David and
Poll,1937:267, Tembwe, Moba, Rumonge, Uvira. -
Poll, 1946:220, description, Lake Tanganyika; 1953:155,
pl. 6, fig. 4, description, diet, Lake Tanganyika. Hulot,
1950:170, diet, Lake Tanganyika. Lambert, 1960:26,
figs. 26 and 27, note. Matthes, 1962:5, photo, 45,
note, sexual dimorphism, Uvira (Lake Tanganyika).
Coulter, 1965-1966:34, table, abundance at differ-
ent depths, Lake Tanganyika; 1991a:152, 154, 181,
182, table 8.2, 8.3, 8.16, abundance, ecology, Lake
Tanganyika. Brichard, 1978:360,361,423, photos,
identification, Lake Tanganyika; 1989:476, 478, 481,
483,474, 479, photos, coloration and identification. -
Sands, 1983:23, 24, 76, check-list, diet, photo.- Daget,
Gosse, & Thys van den Audenaerde, 1986:134, check-
list. Sato, 1986:58-59, brood parasitism, Lake
Tanganyika. Burgess, 1989:190, pl. 74, 77, 78, 85,
94, behavior, Lake Tanganyika. Kobayagowa,
1989:14, photo.
Material Examined. Holotype, BMNH
1898.9.9.76, TL 280 mm, SL 220 mm, Sumba, coll. J.E.S.
Moore, (2) BMNH 1906.9.8.70-71, TL 249-267 mm, SL
193-205 mm, Niamkolo, L. Tanganyika, coll.
Cunningham, (2) BMNH 1955.12.20.1840-1841, TL 213-
256 mm, SL 168-199 mm, Usumbura, L. Tanganyika,
coll. Inst. Roy. Sci. Nat. Beige, (1) BMNH
1955.12.20.1836, TL 78 mm, SL 61 mm, Nyanza Bay,
L. Tanganyika, coll. Inst. Roy. Sci. Nat. Belge, (2)
BMNH 1955.12.20.1857-1858, TL 200-214 mm, SL 152-
172 mm, Lagosa, L. Tanganyika, coll. Inst. Roy. Sci.
Nat. Belge, (1) BMNH 1955.12.20.1852, TL 99 mm, SL
75 mm, Moba Bay, Lake Tanganyika, coll. Inst. Roy.
Sci. Nat. Belge, (1) BMNH 1955.12.20.1842, TL 225
mm, SL 175 mm, Albertville, indigenous market, Lake
Tanganyika, coll. Inst. Roy. Sci. Nat. Beige, 21.XI.1946,
(1) BMNH 1955.12.20.1845, TL 213 mm, SL 168 mm,
M'Pala, L. Tanganyika, coll. Inst. Roy. Sci. Nat. Belge,
7.I.1947, (1) BMNH 1955.12.20.1833, TL 69 mm, SL
54 mm, Moba Bay, L. Tanganyika, coll. Inst. Roy. Sci.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Nat. Belge, 7.II1.1947, (2) BMNH 1955.12.20.1855-
1856, TL 150-209 mm, SL 123-166 mm, S. of Malagarasi
delta, offshore of the southern point of the Malagarasi
delta, L. Tanganyika, coll. Inst. Roy. Sci. Nat. Belge,
26.111.1947, (2) BMNH 1955.12.20.1838-1839, TL 136-
214 mm, SL 107-175 mm, opposite the Lugumba R.,
Lake Tanganyika, coll. Inst. Roy. Sci. Nat. Beige,
23.V. 1947, (2) BMNH 1982.4.13:4789-4790, TL 93-115
mm, SL 76-91 mm, Kigonga Bay, L. Tanganyika, coll.
R. Travers, (3) BMNH 2005.9.26.5-7, TL 67-79 mm,
SL 53-64 mm, Mpulungu, Lake Tanganyika (Zambia),
coll. J. Day, 2005, (3) BMNH 2005.9.26.8-10, TL 61-
69 mm, SL 48-55 mm, Edith Bay, Lake Tanganyika (Tan-
zania), coll. J. Day, 2005, (1) BMNH 2005.9.26.11-16,
TL 94 mm, SL 73 mm, Kigoma, Tanzania, coll. J. Day,
2005, (3) BMNH 2005.9.26.19-21, TL 99-106 mm, SL
77-84 mm, Democratic Republic of Congo, Lake
Tanganyika, coll. J. Day, 2005, (5) BMNH 2005.9.26.70-
74, TL 71- 99 mm, SL 54-74 mm, Kigoma, Lake
Tanganyika (Tanzania), coll. J. Day, 2005, (2) MRAC
39189-39192, TL 94-96 mm, SL 72-73 mm, Moba (Lake
Tanganyika), coll. Van Maldern, 1932, (3) MRAC 53098-
53100, TL 104-112 mm, SL 79-88 mm, Nyanza (L.
Tang.), coll. A. Testiade, 1937, (1) MRAC 90265, TL
211 mm, SL 165 mm, Albertville, coll. Poll (Expl. Hydrol.
Tang.), 21.XI.1946, (2) MRAC 90276-90277, TL 200-
216 mm, SL 152-166 mm, Stat: 63, offshore from the
Malagarasi, 10-15 km from the coast, coll. M. Poll,
3.II.1947, (2) MRAC 90283-90284, TL 94-236 mm, SL
74-187 mm, Stat. 224, Moba Bay, coll. Poll (Expl. Hydrol.
Tang.), 4.IV.1947, (1) MRAC 90287, TL 207 mm, SL
161 mm, Stat. 250, Bay of Burton, over the bottom of
the bay, coll. M. Poll (Exp. Hydr. Tang.), 18-19.IV. 1947,
(1) MRAC 91608, TL 203 mm, SL 150 mm, Uvira, Lake
Tanganyika, coll. G. Marlier, 1.IX. 1949, (1) MRAC
80538, TL 183 mm, SL 143 mm, Rumonge (L. Tang.),
coll. Buscoin, 1950, (3) MRAC A3-033-P-0050-0052,
TL 75-105 mm, SL 60-83 mm, Magara, route Bujumbura-
Nyanza Lac (Burundi), coll. L. De Vos, 19.10.1994, (1)
MRAC Al-094-P-0054, TL 80 mm, SL 61 mm,
Kasakalawe, Lake Tanganyika, S847.23', E31004.40'
(Zambia), coll. Snoeks, Hanssens, Verheyen et al., (2)
SAIAB 39578, TL 77-85 mm, SL 62-67 mm, Zambia,
Lake Tanganyika; Mpulungu, Musende Rocks,
0846'00"S, 031 51'00"E, coll. 7.VII. 1992, (3) SAIAB
40174, TL 58-96 mm, SL 45-76 mm, Zambia, Lake
Tanganyika; Musende Rocks, 0846'00"S, 031006'00"E,
coll. 05.VIII. 1992, (1) SAIAB 56212, TL 73 mm, SL 57
mm, Kigoma Bay, below Hill top Hotel, Tanzania,
0453'03"S, 029037' 11"E, coll. 08.X.1997, (1) SAIAB
56251, TL 71 mm, SL 58 mm, Jacobsen's Beach, Tan-
zania, 04o54'31"S, 029'36'02"E, coll. 04.X.1997, (2)
SAIAB 56254, TL 82-97 mm, SL 63-78 mm, Jacobsen's
Beach, Tanzania, 0454'31"S, 029036'02"E, 05.X.1997,
(1) SAIAB 76164, TL78 mm, SL 63 mm, Zambia, Mbala,
Lake Tanganyika; Mbita Island (northwest end),
845.18'S, 31005.07'E, 29.II.2004, (3) SAIAB 76175,
TL 56-76 mm, SL 44-59 mm, Zambia, Mbala, Lake
Tanganyika; Musende Rocks beach, 08045.18'S,
31005.07'E, 29.II.2004, (1) UF 160941, TL 127 mm, SL
100 mm, Cape Chaitika, Lake Tanganyika (Zambia), via
P. Hauschner, V.2005, (2) CU 90975, TL 101-103 mm,
SL 77-79 mm, Tanzania; Kigoma; Jacobsen's Beach;
coll. Catherine Wagner, 13.VIII.2005.
Diagnosis. Axillary pore present; mandibular
teeth 13-29; body with large spots; fin spines dark; 8
pectoral-fin rays; black triangles at base of pelvic and
anal fins absent or poorly developed; eye 44.9-62.0%
snout length; premaxillary toothpad uninterrupted; sec-
ondary branches on medial mandibular barbel absent;
occipito-nuchal shield usually covered with skin; papil-
lae on skin of body absent; hindgut chamber absent;
maximum TL 280 mm.
Synodontis multipunctatus can be distinguished
from S. ilebrevis, S. irsacae, S. lucipinnis, S.
melanostictus, and S. polli by the presence of an axil-
lary pore. Synodontis multipunctatus differs from S.
petricola in having a large axillary pore (vs. small in S.
petricola), brown to black fin spines (vs. white in S.
petricola), a much larger eye (44.9-62.0% of snout
length in S. multipunctatus vs. 28.7-40.1% in S.
petricola), and an uninterrupted premaxillary toothpad.
This premaxillary toothpad condition also helps to distin-
guish S. multipunctatus from S. ilebrevis, S. irsacae,
S. lucipinnis, S. melanostictus, S. polli, and S.
tanganaicae. The number of mandibular teeth further
separates S. multipunctatus from S. ilebrevis, S.
lucipinnis, S. melanostictus, S. petricola, S. polli, and
S. tanganaicae, as well as from S. granulosus (13-29
in S. multipunctatus vs. 50-66 in S. ilebrevis, 35-51 in
S. lucipinnis, 23-36 in S. melanostictus, 31-50 in S.
petricola, 40-70 in S. polli, 33-49 in S. tanganaicae,
and 28-51 in S. granulosus). Synodontis multipunctatus
can be further distinguished from other endemic
Tanganyikan species (with the exception of S.
grandiops) by its lack of well developed black triangles
at the base of the pelvic and anal fins. Synodontis
multipunctatus can be distinguished from S. dhonti and
further differentiated from S. granulosus and S.
tanganaicae by the presence of many large spots (vs.
spots absent or very small), skin covering the occipito-
nuchal shield, and lack of papillae on the body.
Synodontis multipunctatus is further distinguished from
S. dhonti and S. tanganaicae by the lack of secondary
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Table 10. Morphometrics and meristic counts for Synodontis multipunctatus. All morphometrics given in percent of
base measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n=66) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
28.8
28.6
39.2
25.2
34.6
27.0
26.2
88.2
96.8
59.8
47.1
22.2
42.7
39.1
34.3
120.7
53.6
27.0
50.4
94.2
91.4
47.1
90.7
83.0
51.9
56.8
267.7
130.8
19.6-29.4
26.2-33.2
35.8-42.7
23.9-34.9
21.5-48.6
22.7-35.3
21.1-31.3
78.5-92.7
69.3-96.7
47.8-66.0
41.8-50.8
21.6-30.2
29.7-42.7
29.7-41.7
24.2-42.3
68.3-133.8
39.9-78.1
18.2-39.0
33.0-67.2
73.2-111.3
66.9-106.0
44.9-62.0
64.3-90.7
61.7-94.5
51.9-87.6
55.6-91.0
221.2-414.9
109.8-190.2
24.32.2
30.01.3
40.01.3
29.22.2
31.15.1
26.92.4
26.81.8
84.03.1
77.75.3
57.03.9
45.81.6
25.31.7
35.02.2
37.32.1
32.83.5
102.416.4
54.48.7
27.54.5
51.06.8
89.69.2
89.37.4
55.43.9
76.64.7
81.76.0
72.56.3
68.16.9
307.636.5
142.118.6
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
13(2); 14(4); 15(3); 16(6); 17(4);
18(8); 19*(5); 20(5); 21(6);
22(10); 23(3); 24(6); 26(3); 29(1)
II,7*(66)
I,8*(66)
i,6*(66)
iii,6(9); iii,6,i(2); iii,7(28);
iii,7,i(5); iii,8*(10); iv,6 (4);
iv,6,i(2); iv,7(3); iv,7,i(1), iv,8(2)
i,7,8,i(4)
33(3); 34(1)
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
branches on the medial mandibular barbel. Synodontis
multipunctatus and S. grandiops are most reliably sepa-
rated by pectoral-fin ray counts (8 unbranched elements
in S. multipunctatus vs. 7 in S. grandiops) and eye
size (44.9-62.0% of snout length in S. multipunctatus
vs. 64.2-81.0% in S. grandiops).
Description. Morphometric and meristic data in
Table 10. Maximum TL 280 mm, SL 220 mm. Body
moderately compressed laterally. Predorsal profile
straight to slightly concave. Preanal profile convex. Skin
on body forming numerous vertical folds; papillae ab-
sent.
Head depressed and broad; skin smooth in all but
largest specimens, where ridges from underlying bone
protrude. Snout subconical when viewed laterally;
rounded when viewed dorsally. Occipito-nuchal shield
rugose; completely covered with skin in smaller speci-
mens (<150 mm), lacking skin in larger specimens; ter-
minating posteriorly with narrow, bluntly pointed process
on either side of dorsal spine; ventrally with wide,
rounded process that extends to upper margin of hu-
meral process on either side of body, also covered with
skin in smaller specimens. Eyes dorsolateral; ovoid;
horizontal axis longer. Interorbital area slightly concave
to slightly convex.
Mouth subterminal; lips slightly widened, papillate.
Mandibular teeth 13-29, short, unicuspid; arranged in a
single transverse row. Premaxillary toothpad uninter-
rupted; primary, secondary and tertiary premaxillary teeth
discrete, numerous, arranged in 3, 3, and 2 irregular rows,
respectively.
Maxillary barbel with or without thin basal mem-
brane; lacking branches or crenelations; extending any-
where from base of the pectoral fin to end of humeral
process. Lateral mandibular barbel extending to point
just beyond posterior margin of pectoral girdle; with 4-6
long, non-tuberculate branches; lacking secondary
branches. Medial mandibular barbel 1/3 to 1/2 length of
lateral barbel; with 3-4 pairs of non-tuberculate branches;
lacking secondary branches.
Dorsal fin 11,7; posterior margin slightly concave
to straight. Dorsal-fin spine long, striated, nearly straight,
terminating in short, white filament; anterior margin of
fin spine with 0-3 small serrations distally; posterior
margin with small serrations distally. Pectoral fin 1,8;
posterior margin straight. Pectoral-fin spine roughly
equal in length to dorsal-fin spine, striated, slightly curved,
terminating in short, white filament; anterior spine mar-
gin smooth in adult specimens; many small, antrorse ser-
rations along anterior margin in juvenile specimens; pos-
terior margin with large, retrorse serrations along entire
length. Adipose fin short, poorly developed, margin con-
vex. Pelvic fin i,6; located anterior to vertical through
origin of adipose fin; tip of appressed fin not reaching
base of anal fin. Anal fin iii-iv,6-8; posterior margin nearly
straight; base located ventral to center of adipose fin.
Caudal fin i,7,8,i; forked; lobes pointed.
Humeral process narrow in juveniles, becoming
wider in adults; elongated; granulous; possessing dis-
tinct ridge on its ventral margin in young specimens; ridge
becoming indistinct in adults; dorsal margin concave;
terminating in sharp point (Fig. 3B). Large, dark-col-
ored axillary pore present just ventral to humeral pro-
cess. Gut 0.5-0.8 times body length (n = 4, (2) MRAC
53098-53100, MRAC 90276-90277). Hindgut chamber
absent.
Coloration. Dorsum pale yellow to brown, cov-
ered with large black spots (Fig. 8, 23). Spots larger,
irregular, sometimes confluent in juvenile specimens.
Belly white, with or without black spots. Maxillary and
mandibular barbels white. Iris yellowish to copper col-
ored. Dorsal and pectoral-fin spines brown to black,
filaments white. Pectoral spine with thin, light stripe
along anterior margin. Dorsal and pectoral fins with
black triangles at base, posterior margins white in color.
Triangles in this species may be completely solid or com-
posed of closely spaced spots. Anal and pelvic fins white,
lacking dark triangles of other Tanganyikan species;
single black spot may be present at base of these fins.
Adipose fin with white dorsal edge. Both lobes of cau-
dal fin with black bar from base to tip of fin, posterior
margin of fin white.
Distribution. Lake Tanganyika (Fig. 24); com-
mon.
Habitat. Littoral to benthic zones over shell, sand
and mud bottoms; to a maximum depth of 170 m (Coulter
199 1a).
Diet. Synodontis multipunctatus is generally
considered to be a specialized predator of Neothauma
tanganyicense, a common Tanganyikan gastropod (Poll
1953; Brichard 1989; Coulter 1991a). The method by
which the snail is extracted from its shell is unknown.
Other authors have reported the presence of crusta-
ceans, insect larvae and lamellibranches, in addition to
Neothauma, in the stomach of S. multipunctatus
(Worthington & Ricardo 1936; Poll 1953; Coulter 1991 a),
though later accounts of this species' diet fail to mention
these prey items. Large numbers of lamellibranch shells
were present in the intestine of MRAC 90283. These
shells were packed tightly into the intestine and showed
little evidence of mechanical or chemical digestion, while
the bodies of the mollusks were mostly intact. It may be
that they are ingested incidentally, though the large num-
ber of shells found in this particular specimen suggests
otherwise.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 23. Holotype of Synodontis multipunctatus, BMNH 1898.9.9.76, 280 mm TL, 220 mm SL.
28 30' 32' Reproduction. Synodontis multipunctatus is the
only fish species known to exhibit true interspecific brood
Bur di parasitism (Sato 1986). Eggs are deposited and fertil-
ized among those of at least six species of mouth-brood-
ing cichlids (Sato 1986). These eggs are taken up by
the female cichlid and brooded with the cichlid's own
eggs. The S. multipunctatus hatch before the cichlids
SMad arasik- and after absorbing their own yolk sac, devour the cichlid
eggs and developing fry. It is unknown whether S.
multipunctatus is an obligate brood parasite, or if alter-
. _-6Tanzania nate spawning behaviors exist.
Democrtic Republic Taxonomic Remarks.- Synodontis multi-
of Congo punctatus joins S. granulosus as the only Tanganyikan
Synodontis for which no other names have been pro-
posed or mistakenly applied in the literature. Like most
species of Synodontis in the lake, slight variation in color
L, Rukwa pattern occurs among localities, but not consistently
8 L.Mweru" enough to be able to distinguish among populations.
Wantipa
LMweru \
Synodontis petricola Matthes, 1959
(Figs. 3D, 25, 26; Tables 1, 11)
28" 30* 32'
km
0 so 100 Synodontis petricola Matthes, 1959:78, description,
Lake Tanganyika; 1962:34, pl. 2, fig. a, description,
Figure 24. Known distribution of Synodontis diet, Lake Tanganyika. Poll, 1971:409, figs. 192,
multipunctatus. T denotes type locality. 193, pls. VI.18, XII.9.- Brichard, 1978:360, 424, pho-
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
tos (misidentified), key; 1989:474, 475,476, 479, 481,
483, skin texture, key, photos (some misidentifications).
Daget, Gosse, & Thys van den Audenaerde,
1986:140, check-list.- Burgess, 1989:196, 556, 559,
560, 567, check-list, photos (some misidentifications).
Kobayagowa, 1989:14, photo (misidentified). -
Coulter, 1991 a: 181, 182, table 8.16, diet, habitat, re-
production.
Synodontis multimaculatus. Worthington, E.B. and
Ricardo, C.K., 1936:1067, 1077, 1101, note. Poll,
M., 1946:223, probable absence from Lake
Tanganyika.
Material Examined. Holotype, MRAC 130357,
TL 98 mm, SL 82 mm, Kashekezi, N. Lake Tanganyika,
coll. H.Matthes, 3.IV.1959, (9) paratypes, MRAC
130368-130376, TL 40-82 mm, SL 32-67 mm, Luhanga,
South of Makobola, coll. H.Matthes, 3.IV.1959, (1)
MRAC 78-25-P-19, TL 61 mm, SL 51 mm, Cape
Chaitika, S. Lake Tanganyika (Zambia), coll. P. Brichard,
III.1978, (5) MRAC A3-033-P-0013-0017, TL 97-140
mm, SL 78-111 mm, Mahumba, km 36 route Bujumbura-
Lake Nyanza, Lake Tanganyika (Burundi), coll. L. De
Vos, 29.1.1994, (1) MRAC A3-033-P-0012, TL 102 mm,
SL 83 mm, Magara, route Bujumbara-Lake Nyanza
(Burundi), coll. L. De Vos, 8.III.1994, (1) MRAC 96-
031-P-0581, TL 134 mm, SL 113 mm, Luhanga, km 17
(Zaire), coll. M'boko, 8.III.1994, (2) MRAC 94-069-P-
0290-0291, TL 68-89 mm, SL 54-71 mm, Luhanga, Lake
Tanganyika (Zaire), coll. L. De Vos, 8.III.1994, (7)
MRAC A3-033-P-0005-0011, TL 88-117 mm, SL 69-98
mm, Magara, route Bujumbura-Lake Nyanza (Burundi),
coll. L. De Vos, 11 .V. 1994, (3) MRAC A3-033-P-0018-
0023, TL 76-92 mm, SL 61-74 mm, Karunga, close to
Ubwari, Lake Tanganyika (Zaire), coll. L. De Vos,
2.X.1995, (1) MRAC 95-096-P-2576, TL 69 mm, SL 56
mm, 08'20.24'S, 30031.21'E, Chisiki, coll. Verheyen,
Snoeks, Hanssens, Ruber, Sturmbauer, 10.IV.1995, (1)
MRAC A -094-P-53, TL 81 mm, SL 65 mm, Crocodile
Island, Lake Tanganyika (Zambia), coll. Snoeks,
Hanssens, Verheyen et al., 16.X.2001, (1) paratype,
BMNH 1936.6.14.1232, TL 88 mm, SL 71 mm, Lake
Tanganyika, coll. Christy, (1) SAIAB 77881, TL 74 mm,
SL 59 mm, Zambia, Lake Tanganyika; Musende Rocks,
0846'00"S, 03107'00"E, coll. 10.X.1992, (1) SAIAB
56255, TL 68 mm, SL 55 mm, Tanzania; Jacobsen's
Beach, 04'54'31"S, 029'36'02"E, coll. 05.X.1997, (1)
SAIAB 56225, TL 75 mm, SL61 mm, Tanzania; Kigoma
Bay, below Hill top Hotel, 04053'03"S, 02937' 11 "E, coll.
09.X.1997, (1) SAIAB 56232, TL 80 mm, SL 64 mm,
Tanzania; Cave, Kigoma Hotel below Hill top,
0453'03"S, 02937' 11"E, coll. 11.X.1997, (1) SAIAB
56244, TL 79 mm, SL 64 mm, Tanzania; Bangwe, shingle
beach, 0453'53"S, 02935'39"E, coll. 12.X. 1997.
Diagnosis. Axillary pore present; mandibular
teeth 31-50; body with large spots; fin spines white; 8-9
pectoral-fin rays; black triangles present on bases of all
rayed fins; eye 28.7-40.1% snout length; premaxillary
toothpad interrupted; secondary branches on medial
mandibular barbel present; occipito-nuchal shield cov-
ered with skin; papillae on skin of body absent; hindgut
chamber present; maximum TL 135 mm.
The presence of an axillary pore distinguishes
Synodontis petricola from S. ilebrevis, S. irsacae, S.
lucipinnis, S. melanostictus, and S. polli. The occipito-
nuchal shield being covered with skin separates S.
petricola from S. dhonti, S. granulosus, S.
melanostictus, and S. tanganaicae, all of which also
have a much larger maximum TL (395, 270, 520, and
585 mm, respectively vs. 135 mm in S. petricola).
Mandibular tooth counts serve to further distinguish S.
petricola from S. dhonti and S. irsacae (31-50 in S.
petricola vs. 22 in S. dhonti and 15-29 in S. irsacae).
Synodontis petricola differs from S. granulosus, S.
multipunctatus, and S. grandiops and further differs
from S. dhonti in having an interrupted premaxillary
toothpad and secondary branches on the medial man-
dibular barbel. It is further distinguished from all other
Tanganyikan Synodontis, with the exception of S.
lucipinnis, by having completely white fin spines.
Synodontis petricola is most easily distinguished from
S. lucipinnis by the presence of an axillary pore and
the lack of light colored patches at the bases of the dark
triangles on its fins. Synodontis petricola also some-
what resembles S. polli, but the white fin spines and
lack of papillae on the body of S. petricola (present,
villous in S. polli) serve to separate these species.
Description. Morphometric and meristic data in
Table 11. Maximum TL 135 mm, SL 115 mm. Body not
compressed. Predorsal profile slightly convex. Preanal
profile straight to slightly convex. Skin on body forming
numerous vertical folds; papillae absent.
Head slightly depressed and broad; skin covered
with villous papillae; papillae extend onto base of maxil-
lary barbel and anterior portion of body only. Snout with
nearly flattened margin when viewed laterally; bluntly
rounded when viewed dorsally. Occipito-nuchal shield
covered with skin, terminating posteriorly with wide,
pointed process on either side of dorsal spine, ventrally
with wide, rounded process that extends to upper mar-
gin of the humeral process on either side of body. Eyes
dorsolateral; ovoid; horizontal axis longer. Interorbital
area flat to slightly convex.
Mouth inferior; lips widened and papillate. Man-
dibular teeth 31-50, short, unicuspid; arranged in 6 short,
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Table 11. Morphometric measurements and meristic counts for Synodontis petricola. All morphometrics given in
percent of base measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n=38) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
21.6
27.9
36.6
33.9
18.7
22.9
21.8
84.7
66.5
39.4
52.6
16.4
41.9
37.7
42.1
67.0
59.4
28.0
47.2
82.1
77.9
31.2
79.5
71.6
39.3
43.6
143.2
84.5
14.3-25.0
26.5-30.5
32.6-40.2
22.6-44.7
12.5-29.8
17.6-27.0
18.2-26.6
75.6-95.0
53.7-76.9
39.4-57.6
49.6-60.9
15.1-22.3
27.7-47.2
31.8-41.2
37.0-52.9
45.2-97.7
37.3-72.8
16.3-36.0
34.9-68.7
61.1-98.8
64.9-97.4
28.7-40.1
53.9-83.0
52.3-75.7
38.4-62.9
41.2-60.9
192.1-333.9
66.3-129.9
20.32.6
28.51.0
36.51.6
34.34.4
22.43.6
21.92.7
21.82.0
83.64.6
63.25.4
46.64.0
53.12.3
17.81.4
35.04.6
35.31.9
43.84.1
78.512.1
54.98.1
25.34.2
46.77.1
77.69.4
77.17.0
33.62.5
65.87.0
66.64.6
51.66.2
50.75.2
245.935.4
92.617.0
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
31(1); 33(1); 34(2); 35(1); 36(2);
37(2); 38(5); 39(6); 40(4); 41*(4);
42(4); 43(1); 45(3); 47(1); 50(1)
II,6(3); II,7*(35)
I,8*(25); 1,9(13)
i,6*(38)
iii,8(2); iii,9(4); iv,7(3); iv,7,i(4);
iv,8*(18); iv,8,i(5); iv,9(2)
i,7,8,i(18)
33(2); 34(12); 35(4)
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Figure 25. Holotype of Synodontis petricola, MRAC 130357, 98 mm TL, 82 mm SL.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
transverse rows. Premaxillary toothpad interrupted;
primary, secondary and tertiary premaxillary teeth dis-
crete, numerous, arranged in 2, 2, and 1 irregular rows,
respectively.
Maxillary barbel short; extending at least to base
of pectoral spine; small papillae at base; basal mem-
brane narrow. Lateral mandibular barbel extending to
point just past anterior margin of pectoral girdle; with 4-
7 short, simple, weakly tuberculate branches; usually
lacking secondary branches. Medial mandibular barbel
approximately 1/3 to 2/3 length of lateral barbel; with 4-
6 pairs of tuberculate branches; many secondary
branches present.
Dorsal fin 11,7; posterior margin straight to slightly
concave. Dorsal-fin spine long, striated, slightly curved,
terminating in a short, white filament; anterior margin of
fin spine smooth; posterior margin with small serrations
distally. Pectoral fin 1,8-9; posterior margin broadly
rounded. Pectoral-fin spine roughly equal in length to
dorsal-fin spine, striated, slightly curved, terminating in
short, white filament; anterior spine margin smooth; pos-
terior margin with large, retrorse serrations along entire
length. Adipose fin long, well developed, margin con-
vex. Pelvic fin i,6; located at vertical midway between
posterior base of dorsal fin and origin of adipose fin; tip
of appressed fin does not reach base of anal fin. Anal
fin iii-iv,7-9; posterior margin rounded; base located at
vertical through center of adipose fin. Caudal fin i,7,8,i;
forked; lobes rounded.
Humeral process triangular; granulous; covered
with many small, villous papillae; poorly-developed ridge
on ventral margin; dorsal margin convex; terminating in
a sharp point (Fig. 3D). Axillary pore small. Gut 1.2-
1.3 times body length (n = 2, MRAC A3-033-P-0002-
0011). Well developed hindgut chamber.
Coloration in alcohol. Dorsum yellowish to cu-
prous brown, covered with large, irregularly shaped,
black spots (Fig. 25). Spots proportionately larger, some-
times confluent in juvenile specimens. Belly lighter, with
small, irregularly shaped spots. Maxillary and mandibu-
lar barbels white. Iris copper colored. Dorsal and pec-
toral-fin spines white; terminating in short, white fila-
ments. All rayed fins with black triangles at base; pos-
terior margins white. Both lobes of caudal fin with black
bars, extending from base to tip of fin; posterior margin
white.
Distribution. Lake Tanganyika (Fig. 26). The
large amount of material available in museum collec-
tions suggests that this species is more common than
previously thought (Matthes 1959; Coulter 1991 a).
Habitat. Rocky coasts within the littoral zone, to
a maximum depth of 30 m (Matthes 1959; Coulter 1991a).
28' 30'
km
0 50 100
Figure 26. Known distribution of Synodontis petricola.
T denotes type locality.
Diet. Young individuals appear to be primarily
carnivorous, subsisting mainly on hydracarians, ostra-
cods and insect larvae trichopteranss, chironomids)
(Matthes 1959). Adults feed on algae scraped from
rocky substrates, and small invertebrates (Matthes 1959,
Coulter 1991 a).
Reproduction. No information exists on the
spawning behavior of this species. One female speci-
men was found to contain approximately 100 eggs
(Matthes 1959).
Taxonomic Remarks. Synodontis petricola is
sufficiently distinct that there has been little confusion
regarding its taxonomy. Synodontis petricola shows
considerable variation in the nature of its spotted pat-
tern, with the size and spacing of the spots varying con-
siderably, even among specimens from the same local-
ity.
Synodontis polli Gosse, 1982
(Figs. 5C, 15, 27, 28; Tables 2, 12)
Synodontis eurystomus Matthes, 1959:77, description,
Lake Tanganyika; 1962:31, pl. 1, figs. B, C, descrip-
tion, diet, Lake Tanganyika. Poll, 1971:405, figs.
190, 191, pls. VII.1, XII.8, description. Brichard,
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
1978:357, 424, photos, key. Gosse, J.P., 1982:48,
junior homonym of S. eurystomus Pfeffer, replaced
with S. polli nom. nov. Sands, 1983:23, 79, check-
list, photo. Kobayagawa, 1989:14, photo
(misidentified).- Coulter, 1991a:181, 182, table 8.16,
habitat, diet, reproduction.
Synodontis polli Gosse, 1982:48, nom. nov. Daget,
Gosse, & Thys van den Audenaerde, 1986:141, check-
list. Brichard, 1989:474, 475, 478, 479, 483, skin
texture, photos (some misidentifications). Burgess,
196, 556, 559, 560, 576, checklist, photos (some
misidentifications).
Material Examined. Holotype, MRAC 130440,
TL 148 mm, SL 125 mm, Luhanga, N. Lake Tanganyika,
coll. H. Matthes, 3.IV.1959, (3) paratypes, MRAC
130444-446, TL 97-144 mm, SL 79-120 mm, Makobola,
Lake Tanganyika, coll. H. Matthes (I.R.S.A.C.),
10.XI.1958, (2) paratypes, MRAC 130442-443, TL48-
58 mm, SL 38-47 mm, Luhanga, N. Lake Tanganyika,
coll. H. Matthes (I.R.S.A.C.), 31 .VIII. 1950, (1) paratype,
MRAC 130464, TL 115 mm, SL 96 mm, Bemba, Lake
Tanganyika, coll. H. Matthes (I.R.S.A.C), 26.IV.1958,
(2) paratype, MRAC 130452-453, TL 57-60 mm, SL 46-
48 mm, Makobola, Lake Tanganyika, coll. H. Matthes
(I.R.S.A.C.), 28.VI.1960, (2) paratypes, MRAC 130462-
463, TL 69-85 mm, SL 56-70 mm, Kashkezi, Lake
Tanganyika, coll. H. Matthes (I.R.S.A.C.), 5.11.1959,
(6) paratypes, MRAC 130454-461, TL 39-116 mm, SL
31-96 mm, Makobola, coll. H. Matthes (I.R.S.A.C.),
8.XI.1960, (2) MRAC 131007-131008, TL 137-144 mm,
SL 112-116 mm, Kalungwe, Lake Tanganyika, coll. G.
Leleup, 13.VII. 1961, (2) MRAC 96-031 -P-0889-0890,
TL 117-148 mm, SL 94-123 mm, Luhanga, km 16-17, S.
of Uvira-Ubwa, Lake Tanganyika, coll. L. De Vos,
8.III.1994, (5) MRAC A3-033-P-0076-0080, TL 115-
160 mm, SL 95-132 mm, Luhanga, km 16 S. of Uvira-
Ubwari, Lake Tanganyika (Zaire), coll. M'boko,
28.VII.1995, (1) MRAC A3-033-P-0075, TL 121 mm,
SL 103 mm, Pemba, S. of Uvira, Lake Tanganyika
(Zaire), coll. L. De Vos, 4.XI.1995, (1) MRAC 75-1-P-
117, TL 101 mm, SL 84 mm, 8 km S. of Bujumbura,
Lake Tanganyika, coll. P. Brichard, VII. 1974, (1) MRAC
A3-033-P-0082, TL 146 mm, SL 120 mm, Magara, route
Bujumbura-Lake Nyanza (Burundi), coll. L. De Vos,
14.VII.1995, (1) MRAC, A4-044-P-0007, TL 120 mm,
SL 97 mm, Jakobsen's Beach, aka Mwamahunga, Lake
Tanganyika, 0457.87'S, 29'35.86'E (Tanzania), coll. G.
Kazumbe, 7.VI.2004, (2) MRAC 77-40-P-5-7, TL 88-
99 mm, SL 70-82 mm, Chipimbi, cote Sud-Ouest du Lac
Tanganyika (Zambia); coll. P. Brichard, 6.VII.1977, (1)
MRAC 78-25-P-17-18, TL 89 mm, SL 73 mm, Cap
Kachese, S. Lake Tanganyika (Zambia); coll. P.
Brichard, (6) MRAC 95-096-P-2561-2575, TL 85-108
mm, SL 71-90 mm, Kasenga Point, lake Tanganyika,
0843'31"S, 31 008'01"E [Zambia]; coll. Exp. Tanganyika
95, 4.IV. 1995, (5) MRAC 95-096-P2576-2585, TL 90-
116 mm, SL 74-98 mm, Chisiki, 08020'24"S, 3031 '2 1"E
[Zambia]; coll. Exp. Tanganyika 95, 10.IV.1995, (7)
MRAC 95-096-P-2586-2608, TL 88-125 mm, SL 70-
104 mm, Cape Kachese, lake Tanganyika, 08029'22"S,
30028'32"E [Zambia], coll. Exp. Tanganyika 95,
10.IV. 1995, (1) MRAC 95-096-P-2609, TL 135 mm, SL
112 mm, Kasaba Bay, lake Tanganyika, 0830'57"S,
30038'52"E [Zambia]; coll. Exp. Tanganyika 95,
11 .IV. 1995, (4) MRAC 95-096-P-2610-2613, TL 83-106
mm, SL 69-89 mm, Kala Bay, lake Tanganyika,
08008'47"S, 30058' 14"E [Zambia]; coll. Exp. Tanganyika
95, 19.IV.1995, (1) MRAC 95-096-P-2627, TL 108 mm,
SL 87 mm, Chikulina Bay, lake Tanganyika, 08032'41"S,
3043'36"E [Zambia]; coll. Exp. Tanganyika 95,
12.IV.1995, (1) BMNH 2005.9.26.17-18, TL 120 mm,
SL 98 mm, Mpulungu, Zambia, Lake Tanganyika, coll.
J. Day, 2005, (3) SAIAB 40171, TL 980114 mm, SL 81-
94 mm, Zambia, Lake Tanganyika; Musende Rocks,
08046'00"S, 03106'00"E, 05.VIII.1992, (1) SAIAB
42518, TL 105 mm, SL 87 mm, Zambia, Lake
Tanganyika; Musende Rocks, 08o46'00"S, 031007'00"E,
10.X. 1992, (2) SAIAB 56262, TL 122-150 mm, SL 100-
127 mm, Tanzania; Jacobsen's Beach, 0454'31"S,
029036'02"E, 10.X.1997, (1) SAIAB 76167, TL 117 mm,
SL 100 mm, Zambia, Mbala, Lake Tanganyika, Lake
Tanganyika; Mbita Island (northwest end), 08o45.18'S,
031005.07'E, 29.II.2004, (1) CU 88758, TL 180 mm, SL
145 mm, Tanzania; Kigoma; Jacobsen's Beach, coll. P.B.
McIntyre, 18.VII.2002.
Diagnosis. -Axillary pore absent; mandibular teeth
40-70; body with large spots; fin spines dark; 8-9 pecto-
ral-fin rays; black triangles on bases of all rayed fins;
eye 25.8-39.3% snout length; premaxillary toothpad in-
terrupted; secondary branches on medial mandibular
barbel present; occipito-nuchal shield covered with skin;
villous papillae present on skin of body; long gut; well
developed hindgut chamber; body depth 20.2-27.0% SL;
maximum TL 180 mm.
The lack of an axillary pore immediately distin-
guishes Synodontis polli from S. dhonti, S. grandiops,
S. granulosus, S. multipunctatus, S. petricola, and S.
tanganaicae. Additionally, S. polli generally has a
greater number of mandibular teeth than all other
Tanganyikan species (except S. ilebrevis), though some
overlap does occur (40-70 in S. polli vs. 22 in S. dhonti,
17-26 in S. grandiops, 28-51 in S. granulosus, 15-29 in
S. irsacae, 35-51 in S. lucipinnis, 13-29 in S.
multipunctatus, 23-36 in S. melanostictus, 31-50 in S.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Table 12. Morphometric measurements and meristic counts for Synodontis polli. All morphometrics given in per-
cent of base measurement. *Meristic data for holotype. Abbreviations as in Table 3.
MEASUREMENT HOLOTYPE RANGE (n=66) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
HPL/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
22.0
28.2
38.6
35.8
13.6
19.2
22.5
86.6
73.1
44.6
55.7
18.8
43.3
35.4
57.3
48.0
41.9
25.0
59.6
68.2
79.8
33.7
77.8
63.6
43.3
53.0
192.8
127.5
20.2-27.0
26.3-32.6
35.2-41.4
29.2-45.6
10.6-28.3
12.8-23.4
17.8-25.0
79.6-94.5
61.9-79.6
35.0-51.5
50.8-60.2
15.1-21.2
31.8-44.3
30.9-38.2
44.4-62.2
45.4-93.5
30.9-67.1
15.5-31.5
34.1-69.3
42.9-80.1
59.8-84.0
25.8-39.3
57.2-81.0
52.4-69.7
38.3-60.9
41.4-66.3
146.8-252.2
86.9-144.0
23.01.7
29.81.2
37.91.4
35.63.4
19.23.8
19.62.5
22.01.6
85.83.1
68.63.2
43.83.4
55.21.8
18.41.3
36.92.8
34.41.4
54.64.0
65.111.7
48.37.6
23.43.1
49.17.6
65.78.7
73.85.6
33.42.7
67.05.2
62.33.3
50.15.0
53.74.8
192.624.9
1080+12.6
MERISTICS
Mandibular tooth count
Dorsal fin count
Pectoral fin count
Pelvic fin count
Anal fin count
Caudal fin count
Total vertebrae count
40(1); 41(1); 43(3); 44(2); 45(3);
46(3);_47(3); 48(1); 49(2); 50(5);
51(1); 52(6); 53(4); 54(6); 55(3);
56(2); 57(1); 58(3); 60(2); 61(1);
62(3); 63*(3); 65(1); 66(1); 67(1);
70(1)
II,7*(66)
1,8(32); I,9*(33)
i,6*(65)
iii,8(5); iii,8,i(4); iii,9(ll); iv,7(4);
iv,8*(28); iv,8,i(4); iv,9(9); v,7(1)
i,7,8,i(8)
35(2); 36(4); 37(2)
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
petricola, and 33-49 in S. tanganaicae). Synodontis
polli further differs from S. petricola and S. lucipinnis
by having dark fin spines (vs. white in S. petricola and
S. lucipinnis). The large spots on the body distinguish
S. polli from S. dhonti, S. granulosus, S. melanostictus
and S. tanganaicae, and the black triangles at the base
of all the rayed fins further separate S. polli from S.
melanostictus and adult S. tanganaicae. Synodontis
polli is further distinguished from S. dhonti, S.
granulosus, S. melanostictus, and S. tanganaicae in
having an occipito-nuchal shield which is covered by
skin. The interrupted premaxillary toothpad further sepa-
rates S. polli from S. dhonti, S. grandiops, S.
granulosus, and S. multipunctatus, in which it is unin-
terrupted, and the presence of secondary branches on
the medial mandibular barbel further distinguishes S. polli
from S. grandiops, S. granulosus, and S.
multipunctatus. Synodontis polli differs from S.
ilebrevis in having a long gut (4.0-5.5 times TL in S.
polli vs. 0.8-1.4 times TL in S. ilebrevis) (Fig. 14, 15), a
well developed hindgut chamber (Fig. 15), deeper body
(body depth 20.2-27.0% SL in S. polli vs. 18.2-20.1%
SL in S. ilebrevis), villous papillae on the skin (vs. short,
flattened papillae in S. ilebrevis), and large, irregular,
closely spaced spots on the skin (vs. small, regular, more
widely spaced spots in S. ilebrevis).
Description. Morphometric and meristic data in
Table 12. Maximum TL 180 mm, SL 145 mm. Body not
compressed. Predorsal profile convex. Preanal profile
slightly convex. Skin on body forming numerous verti-
cal folds covered with villous papillae which do not ex-
tend onto fins, giving skin somewhat velvety texture.
Head deep and broad; skin covered with villous
papillae. Snout with broadly rounded margin when
viewed laterally and dorsally. Occipito-nuchal shield cov-
ered with skin, terminating posteriorly with wide, pointed
process on either side of dorsal spine, ventrally with wide,
rounded process that extends to upper margin of hu-
meral process on either side of body. Eyes dorsolateral,
ovoid, horizontal axis longer. Interorbital area flat to
slightly convex.
Mouth inferior; lips widened and papillate. Man-
dibular teeth 40-70, short, unicuspid; arranged in 6-8 short,
transverse rows. Premaxillary toothpad interrupted;
primary, secondary and tertiary premaxillary teeth dis-
crete; numerous; arranged in 3, 2, and 1 irregular rows,
respectively.
Maxillary barbel short; extending at least to base
of pectoral spine; lacking branches or crenelations; basal
membrane narrow. Lateral mandibular barbel extend-
ing to point just short of anterior margin of pectoral girdle;
with 4-5 short, non-tuberculate branches; lacking sec-
ondary branches. Medial mandibular barbel approxi-
mately 1/3 to 2/3 length of lateral barbel; with 3-5 pairs
of tuberculate branches; many secondary branches
present.
Dorsal fin 11,7; posterior margin slightly concave.
Dorsal-fin spine short; striated; slightly curved, termi-
nating in short, dark filament; anterior margin of fin spine
smooth; posterior margin with small serrations distally.
Pectoral fin 1,8-9; posterior margin straight. Pectoral-
fin spine roughly equal in length to dorsal-fin spine, stri-
ated, slightly curved, terminating in short, dark filament;
anterior spine margin granulate in adult specimens; many
small, antrorse serrations along anterior margin in juve-
nile specimens; posterior margin with large, retrorse ser-
rations along entire length. Adipose fin long, well devel-
oped, margin convex. Pelvic fin i,6; located at or ante-
rior to vertical through origin of adipose fin; tip of ap-
pressed fin does not reach base of anal fin. Anal fin iii-
v,7-9; posterior margin rounded; base located at vertical
through center of adipose fin. Caudal fin i,7,8,i; forked;
lobes rounded.
Humeral process wide; triangular; granulous; poorly
developed ridge on ventral margin; dorsal margin con-
vex; terminating in a sharp point (Fig. 5C). Axillary pore
absent. Gut long, 4.0-5.5 times TL (n = 3, MRAC 130440,
MRAC 131007-008). Well developed hindgut chamber
present (Fig. 15).
Coloration in alcohol. Dorsum olive brown, cov-
ered with large, irregularly shaped, black spots (Figs.
15, 27). Spots proportionately larger, irregular, some-
times confluent in juvenile specimens. Belly lighter, with
smaller, irregularly shaped spots. Maxillary and man-
dibular barbels white. Iris copper colored. Dorsal and
pectoral-fin spines dark, terminating in short, dusky fila-
ments. Pectoral spine with thin, light stripe along ante-
rior margin. All rayed fins with black triangles at their
base; posterior margins white in color, becoming dusky
as specimen grows. Triangles may be completely solid
or composed of closely spaced spots. Both lobes of
caudal fin with black bar from base to tip of fin; poste-
rior margin of fin white, becoming dusky with growth.
Distribution. Lake Tanganyika (Fig. 28); com-
mon.
Habitat. Rocky coasts within the littoral zone, to
a maximum depth of 20 m (Matthes 1959; Coulter 1991a).
Diet. Synodontis polli feeds on algae scraped
from rocky substrates and small invertebrates (e.g. os-
tracods, chironomid larvae, small shrimps) (Matthes
1959; Coulter 1991a).
Reproduction. No information exists regarding
the spawning season, behavior, or habitat of this spe-
cies, although mature specimens appear to be caught at
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 27. Holotype of Synodontis polli, MRAC 130440, 148 mm TL, 125 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
km
0 50 100
Figure 28. Known distribution of Synodontis polli. T
denotes type locality.
many different times of the year. One female specimen
has been found to contain approximately 110 eggs
(Matthes 1959).
Taxonomic Remarks. The taxonomic status of
Synodontis polli has remained fairly stable, except for
the replacement of its original name S. eurystomus
Matthes, a junior homonym of Synodontis eurystomus
Pfeffer, a synonym of Chiloglanis deckenii Peters
(Gosse 1982).
Specimens from the southern part of this species'
range show significant variation in their color pattern,
with the spots on the body becoming smaller or, in some
cases, disappearing as body size increases. In other
respects, these specimens closely resemble specimens
collected in the northern part of the lake.
Synodontis tanganaicae Borodin, 1936
(Figs. 4C, 4D, 29, 30, 31; Tables 1, 13)
Synodontis serratus tanganaicae Borodin, 1936:9, n.
sp. of S. serratus Riippel, Kasenga, Lake Tanganyika.
Synodontis lacustricolus Poll, 1953:157, fig. 18a, pl. 6,
fig. 3, description, diet, Lake Tanganyika; 1971:402,
fig. 188, 189, pl. 6, 12, description.- Matthes, 1962:41,
pl. 3, fig. a, description. -Coulter, 1965-1966:34, abun-
dance at different depths, Lake Tanganyika; 1991 a: 152,
154, 181, 182, table 8.2, 8.3, 8.16, abundance, habitat,
diet. Brichard, 1978:424, key; 1989:479, key. -
Sands, 1983, 23, check-list. Daget, Gosse, & Thys
van den Audenaerde, 1986:131, check-list. Burgess,
1989:196, check-list. De Vos and Thys van den
Audenaerde, 1998:147, synonymy with S.
tanganaicae.
Synodontis tanganyicae. De Vos, L. and Thys van
den Audenaerde, D., 1998:147.
Material Examined. Lectotype (De Vos and Thys
van den Audenaerde 1998), MCZ 32538, TL 416 mm,
SL 321 mm, Kasanga, Lake Tanganyika, coll. A.
Loveridge, 1930, (1) paralectotype, MCZ 148517, TL
403.6 mm, SL 322.41 mm, Kasanga, Lake Tanganyika,
coll. A. Loveridge, 1930, (1) holotype, S. lacustricolus,
ISRNB 197, TL 601 mm, SL 503 mm, Stat. 123: near
Karema, depth: about 30 m, coll. Expl. hydrobiol. du lac
Tanganyika, 15.II.1947, (1) paratype, S. lacustricolus,
ISRNB 198, TL 429 mm, SL 352 mm, Tembwe Bay,
Tanganyika district, Belgian Congo, coll. Explor.
hydrobiol. du lac Tanganyika, 13.11.1947, (1) paratype,
S. lacustricolus, ISRNB 199, TL 413 mm, SL 336 mm,
M'Vua Bay, Tanganyika district, Belgian Congo, coll.
Explor. hydrobiol. du Lac Tanganyika, 12.111.1947, (1)
paratype, S. lacustricolus, MRAC 130720, TL 333 mm,
SL 255 mm, Mboko Island, N. Lake Tanganyika, coll.
Marlier (I.R.S.A.C.), 7.11.1957, (1) paratype, S.
lacustricolus, MRAC 90288, paratype (S.
lacustricolus), TL 519 mm, SL 434 mm, Tembwe Bay,
Lake Tanganyika, coll. M. Poll/Expl. Hydrobio.Tang.,
13.II.1947, (1) paratype, S. lacustricolus, MRAC 90289,
TL 582 mm, SL 489 mm, Edith Bay, Lake Tanganyika,
coll. M. Poll/Expl. Hydrobio. Tang., 14-15.II.1947, (1)
MRAC 90279, SL 203.5 mm SL, Albertville, St. 103,
depth 7-10 m, trawl with panels, coll. Miss. D'Explor..
Hydr. L. Tang., 3.11.47, (2) MRAC 96-083-P-003-004,
TL 300-330 mm, SL 240-265 mm, Zambia, coll. De Vos,
1996, (1) paratype, S. lacustricolus, BMNH
1955.12.20.1849, TL 385 mm, SL 315 mm, Opposite Kala,
Lake Tanganyika, Inst. Roy. Nat. Belge, (1) paratype,
S. lacustricolus, BMNH 1955.12.20.1850, TL 361 mm,
SL 291 mm, Tembwe Bay, Lake Tanganyika, Inst. Roy.
Nat. Beige, (1) paratype, S. lacustricolus, BMNH
1955.12.20.1851, Malagarasi, Lake Tanganyika, coll. Inst.
Roy. Nat. Belge, 12.V.1947, (3) BMNH 1936.6.15.1203-
5 (orig. I.D. S. granulosus), TL 400-522 mm, SL 328-
423 mm, Lake Tanganyika, coll. Christy.
Diagnosis. Axillary pore present; mandibular
teeth 33-49; body with small or no spots; fin spines dark;
8-9 pectoral-fin rays; black triangles on bases of all rayed
fins in juveniles, absent in adults; eye 16.0-26.9% snout
length; premaxillary toothpad interrupted; secondary
branches on medial mandibular barbel present; occipito-
nuchal shield not covered with skin; granular papillae
present on skin of body; humeral-process length 172.9-
255.4% of humeral process width; maximum TL 585
mm.
The presence of an axillary pore distinguishes
Synodontis tanganaicae from S. ilebrevis, S. irsacae,
S. lucipinnis, S. melanostictus, and S. polli.
Synodontis tanganaicae differs from S. granulosus
in humeral process shape (humeral-process length 172.9-
255.4% of humeral process width in S. tanganaicae
vs. 253.8-437.2% in S. granulosus) (Figs. 3A, 4C, D).
The lack of both large spots and skin covering the
occipito-nuchal process separates S. tanganaicae from
S. grandiops, S. ilebrevis, S. lucipinnis, S.
multipunctatus, S. petricola, and S. polli. Adult speci-
mens of S. tanganaicae are further distinguished from
the other Tanganyikan endemics by the lack of black
triangles on the rayed fins.
Synodontis tanganaicae most closely resembles
S. melanostictus and S. dhonti. Synodontis
tanganaicae differs from S. melanostictus in the num-
ber of mandibular teeth (33-49 in S. tanganaicae vs.
23-36 in S. melanostictus), eye size (16.0-26.9% of snout
length in S. tanganaicae vs. 31.5-38.1% in S.
melanostictus), the granular papillae on the skin (vs.
villous in S. melanostictus), and differences in humeral
process shape (Fig. 4). The higher mandibular tooth
count (33-49 vs. 22 in S. dhonti), well-developed adi-
pose fin (vs. poorly developed in S. dhonti), and inter-
rupted premaxillary toothpad separate S. tanganaicae
from S. dhonti.
Description. Morphometric and meristic data in
Table 13. Maximum TL 585 mm, SL490 mm. Body not
compressed. Predorsal profile straight. Preanal profile
convex. Skin on body forming numerous vertical folds;
covered with papillae, which do not extend onto fins.
Head depressed and broad, skin covered with
granular papillae. Snout with blunt margin when viewed
laterally; broadly rounded margin when viewed dorsally.
Occipito-nuchal shield rugose, not covered with skin;
terminating posteriorly with narrow, bluntly pointed pro-
cess on either side of dorsal spine; ventrally with wide,
rounded process that extends to upper margin of hu-
meral process on either side of body. Eyes dorsolateral;
ovoid; horizontal axis longer. Interorbital area flat.
Mouth inferior; lips wide and papillate. Mandibu-
lar teeth 33-49, short, unicuspid; arranged in single trans-
verse row. Premaxillary toothpad interrupted; primary,
secondary and tertiary premaxillary teeth discrete; nu-
merous; arranged in irregular rows.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Maxillary barbel with well developed basal mem-
brane; lacking branches or crenelations; extending at
least to base of pectoral fin. Lateral mandibular barbel
extending to point just short of anterior margin of pecto-
ral girdle, with 4-7 non-tuberculate branches; lacking
secondary branches. Medial mandibular barbel about
1/3 length of lateral barbel; with 4-6 pairs of non-tuber-
culate branches; secondary branches present.
Dorsal fin 11,7-8; posterior margin straight. Dor-
sal-fin spine long, granulous, nearly straight, terminating
in short, dark filament; anterior margin of fin spine
granulous; posterior margin with small serrations dis-
tally. Pectoral fin 1,8-9; posterior margin nearly straight.
Pectoral-fin spine roughly equal in length to dorsal-fin
spine, striated, slightly curved, terminating in short, dark
filament; anterior spine margin granulate; posterior mar-
gin with large retrorse serrations along entire length.
Adipose fin long, well developed, margin convex. Pel-
vic fin i,6; located at vertical through posterior base of
dorsal fin; tip of appressed fin does not reach base of
anal fin. Anal fin iii-v,7-9; posterior margin broadly
rounded; base located ventral to adipose fin. Caudal fin
i,7,8,i; forked; lobes slightly rounded.
Humeral process triangular, granulous; ridge on
ventral margin absent; dorsal margin concave; termi-
nating in blunt point (Figs. 4C, 4D). Axillary pore ab-
sent.
Coloration in alcohol. Dorsum gray to reddish
brown (Figs. 29, 30). Belly lighter in color. Scattered,
small black spots present on entire body. Maxillary and
mandibular barbels white, bases sometimes with scat-
tered dusky pigmentation. Iris copper colored. Dorsal
and pectoral-fin spines dark, terminating in short, dusky
filaments. All rayed fins with small, dark spots similar to
those on body. Juvenile specimens with black triangles
at bases of rayed fins.
Distribution. Lake Tanganyika (Fig. 31); fairly
common.
Habitat. Littoral to benthic zones over shell, sand
and mud bottoms, to a maximum depth of 130 m (Coulter
1991a).
Diet. Gastropods, lamellibranchs, insect larvae,
ostracods, and shrimps (Poll 1953; Coulter 1991 a).
Reproduction. No information exists regarding
the reproduction of Synodontis tanganaicae.
Taxonomic Remarks. Synodontis serratus
tanganaicae Borodin, previously considered a subspe-
cies of S. serratus Riippell, was elevated to species
status by De Vos and Thys van den Audenaerde (1998),
who also placed S. lacustricolus Poll in the synonymy
of S. tanganaicae. Our examination of the material
available has yielded no significant morphometric or
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
Table 13. Morphometric and meristic counts for Synodontis tanganaicae. All morphometrics given in percent of
base measurement. *Meristic data for lectotype. Abbreviations as in Table 3.
MEASUREMENT LECTOTYPE RANGE (n=17) MEANSD
Body depth/SL
Head length/SL
Snout-dorsal length/SL
Adipose fin length/SL
Maxillary barbel length/SL
Dorsal spine length/SL
Pectoral spine length/SL
Head width/HL
Head depth/HL
Humeral process length/HL
Snout length/HL
Eye/HL
Interorbital width/HL
Postorbital length/HL
Mouth length/HL
Maxillary barbel length/HL
LMB/HL
MMB/HL
MMB/LMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout length
Postorbital length/Snout length
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD
23.9
36.5
48.2
32.8
40.0
23.8
24.7
81.9
72.0
42.6
57.8
13.0
38.4
36.1
45.5
109.7
42.9
16.6
38.7
65.2
67.8
22.5
66.4
62.4
33.8
36.0
198.4
149.4
20.8-26.4
31.5-36.5
41.2-48.6
21.9-32.8
29.8-46.3
21.1-27.6
19.0-27.2
79.4-94.1
66.1-84.1
38.1-55.2
51.1-60.6
9.7-14.6
35.0-45.0
35.1-40.4
30.1-47.2
90.7-138.5
36.4-66.4
12.4-23.5
29.2-43.4
63.5-87.4
57.9-83.2
16.0-26.9
60.3-83.5
59.8-78.5
23.9-38.0
23.3-39.6
172.9-255.4
98.6-149.8
24.41.5
33.11.4
43.52.2
25.82.8
38.15.1
24.12.1
23.02.4
87.54.0
73.74.0
46.45.1
55.42.2
12.81.4
39.52.6
38.01.7
43.14.0
116.315.8
52.37.0
19.63.2
37.64.2
73.37.5
69.46.8
23.23.1
71.56.4
68.65.1
32.43.6
33.24.4
211.422.5
127.815.0
MERISTICS
Mandibular tooth count 38(3); 39(1); 41(2); 42(1); 43*(3);
44(1); 46(1); 47(3); 48(1); 49(1)
Dorsal fin count II,7*(15); II,7,i(l); 11,8(1)
Pectoral fin count I,8*(4); 1,9(13)
Pelvic fin count i,6*(16); i,7(1)
Anal fin count iii,8(1); iv,7*(2); iv,8(8); iv,8,i(l);
iv,9(1); v,7(1); v,8(1); v,8,i(1)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Figure 29. Lectotype of Synodontis tanganaicae, MCZ 32538, 416 mm TL, 321 mm SL.
Figure 30. Synodontis tanganaicae, MRAC 90288, paratype of S. lacustricolus, 519 mm TL, 434 mm SL.
WRIGHT AND PAGE: Taxonomic revision of Lake Tanganyikan Synodontis
km
0 5s 100
Figure 31. Known distribution of Synodontis
tanganaicae. T denotes type locality.
meristic differences between S. tanganaicae and types
of S. lacustricolus, nor do the new characters exam-
ined in this study indicate that the specimens represent
two species. De Vos and Thys van den Audenaerde's
decision is accepted here, though we retain the spelling
of Borodin rather than using tanganyicae.
KEY TO THE SPECIES OF Synodontis
OF LAKE TANGANYIKA
l a. Large, conspicuous axillary pore.......................2...
lb. Axillary pore absent or very small.....................6...
2a. Mandibular teeth 13-29; eye large, 21.6-31.9% head
length; skin smooth; many large black spots ............3...
2b. Mandibular teeth 22-51; eye smaller, 9.7-22.6% head
length; skin covered with granula papillae; spots very
sm all or absent............................. ......................4...
3a. Pectoral fin I, 8; eye 44.9-62.0% snout length; maxi-
mum TL 280 mm........................ S. multipunctatus
3b. Pectoral fin 1,7, eye 64.2-81.0% snout length; maxi-
mum TL 145 mm................. S. grandiops
4a. Premaxillary toothpad uninterrupted; humeral pro-
cess length 253.8-437.2% humeral process width...... 5
4b. Premaxillary toothpad interrupted; humeral process
length 172.9-255.4% humeral process width...............
......................................................... S. tanganaicae
5a. 22 mandibular teeth; secondary branches on medial
mandibular barbel; eye 23.1% snout length....S. dhonti
5b. 28-51 mandibular teeth; no secondary branches on
medial mandibular barbel; eye 31.2-50.2% snout
length................... .....................S. granulosus
6a. Pectoral-fin spines white..................................7...
6b. Pectoral-fin spines dark...................................8...
7a. Very small axillary pore; black triangles on rayed
fins lack light patches at base................S.... petricola
7b. Axillary pore absent; black triangles on rayed fins
with distinct light patches at base...........S. lucipinnis
8a. Moderate to large spots; black triangles present on
rayed fins; occipito-nuchal shield covered with
sk in ............................................... ......................9...
8b. Small spots; black triangles absent on rayed fins;
occipito-nuchal shield not covered with skin...............
............ ............ ......................... S. m elanostictus
9a. M andibular teeth 40-70.................................. 10
9b. Mandibular teeth 15-29...................... S. irsacae
10a. Body depth 20.2-27.0% SL; gut length 386.0-
554.6% TL; large hindgut chamber; body and head cov-
ered with villous papillae..............................S. polli
10 Ob. Body depth 18.2-20.1% SL; gut length 78.0-136.2%
TL; hindgut chamber absent; body and head covered
with flattened, granulous papillae.............S. ilebrevis
DISCUSSION
The examination of 312 specimens of Synodontis from
Lake Tanganyika has led to the recognition of 11 spe-
cies of Synodontis from Lake Tanganyika. Ten of the
species are endemic, and three are new to science.
Synodontis irsacae and S. melanostictus are recog-
nized as valid species.
The rayed-fin color pattern and skin folds of the
endemic Tanganyikan species of Synodontis offer sup-
port for the monophyly of this group. Brooks (1950)
suggested that, in the genera of Lake Tanganyikan fishes
that are represented in the lake by a single non-endemic
species and an endemic species flock (Mochokidae and
Mastacembelidae), the endemic species probably arose
from an ancient species due to cleavage of the lake ba-
sin, while the non-endemic species in each group, e.g.
S. melanostictus, is likely to be a recent addition to the
lake fauna. Brichard's (1978) assertion that the species
of Synodontis of Lake Tanganyika are probably not
monophyletic is reasonable when S. melanostictus is
included. More confusing is his apparent belief that the
endemic species flock also does not constitute a mono-
phyletic group. Two characters examined in this study
(rayed-fin color pattern and skin folds) are apparent
synapomorphies supporting the monophyly of the
Synodontis species endemic to Lake Tanganyika.
A robust phylogeny for Synodontis will permit
comparative examination of many of the unique traits
found within this genus. The functional significance of
the skin folds, axillary pore, and hindgut chamber have
yet to be determined. A study of the axillary pore will be
particularly interesting, as Synodontis is the only Afri-
can catfish genus in which this structure has been docu-
mented. Even if this structure is found to serve an iden-
tical function to the structure found in the other families
(Akysidae, Ariidae, Ictaluridae, Sisoridae), a more de-
tailed study of the composition of its secretions will be
extremely interesting, as this has yet to be examined for
any species. The hindgut chamber may serve in a di-
gestive capacity, as its gross morphology is very similar
to that of a structure found in the Kyphosidae. Confir-
mation of the presence of fermentative gut microbes
and short chain fatty acids in the chamber is needed to
confirm that it serves a similar function to the fermenta-
tion chamber of kyphosids (D. German, pers. comm.).
Synodontis represents a group for which signifi-
cant research remains to be done. Given the high diver-
sity of species and behaviors found in this genus, its ne-
glect by past ichthyologists is confusing. It is hoped that
this study of the taxonomy of the Tanganyikan species
flock will lead to future studies of these little-known
fishes.
ACKNOWLEDGEMENTS
Funding for this study was provided by the All Catfish
Species Inventory project funded by the U.S. National
Science Foundation (DEB-0315963). The authors would
like to thank Carl Ferraris, Collete St. Mary, David Evans,
and Jonathan Armbruster for suggesting many useful
improvements to this study. We are grateful to Jacqueline
Wright for her assistance during data collection. Loans
and access to institutional specimens were provided by
Jos Snoeks, Mark Hanssens, and Miguel Parrent of the
Royal Museum for Central Africa (MRAC), James
Maclaine, Patrick Chapman, and Julia Day of the Natu-
ral History Museum (BMNH), Paul Skelton and Roger
Bills of the South African Institute of Aquatic Biodiversity
(SAIAB), John Friel of the Cornell University Museum
of Vertebrates (CU), and Karsten Hartel of the Harvard
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 46(4)
Museum of Comparative Zoology (MCZ). This study
benefited from many discussions of Synodontis with
A.W. Thomson, R.H. Robins, J.A. Lopez, and J.M.
Wright. D.P. German provided a valuable discussion of
digestive physiology and the hindgut chamber.
LITERATURE CITED
Birkhead, W.G. 1967. The comparative toxicity of stings
of the ictalurid catfish genera Ictalurus and
Schilbeodes. Comparative Biochemistry and Physi-
ology, 22:101-1ll.
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