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Taxonomic Revision of Lake Tanganyikan Synodontis (Teleostei: Mochokidae)

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Title:
Taxonomic Revision of Lake Tanganyikan Synodontis (Teleostei: Mochokidae)
Creator:
WRIGHT, JEREMY JOHN ( Author, Primary )
Copyright Date:
2008

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Subjects / Keywords:
Fish ( jstor )
Head ( jstor )
Hindgut ( jstor )
Holotypes ( jstor )
Paratypes ( jstor )
Pectorals ( jstor )
Polls ( jstor )
Skin ( jstor )
Species ( jstor )
Trucks ( jstor )

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University of Florida
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University of Florida
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Copyright Jeremy John Wright. Permission granted to University of Florida to digitize and display this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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Full Text












TAXONOMIC REVISION OF LAKE TANGANYIKAN Synodontis (TELEOSTEI:
MOCHOKIDAE)


















By

JEREMY JOHN WRIGHT


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2006




























Copyright 2006

by

Jeremy John Wright















ACKNOWLEDGMENTS

Funding for this study was provided by the All Catfish Species Inventory project

funded by the U.S. National Science Foundation (DEB-0315963). I would like to thank

Dr. Lawrence Page for directing me to the possibilities for study in the Mochokidae and

for instruction, constructive criticism, and guidance. I also thank Drs. Carl Ferraris and

Collete St. Mary for their contributions as members of my supervisory committee.

Additional thanks are due Dr. David Evans, who graciously agreed to serve as a stand-in

for Dr. Ferraris at the defense of this thesis, and Dr. Jonathan Armbruster of Auburn

University, who provided additional comments for the improvement of this thesis.

I am grateful to Jacqueline Wright for her assistance during data collection. For

help with formatting maps I thank Griffin Sheehy. Loans and access to institutional

specimens were provided by Dr. Jos Snoeks, Mark Hanssens, and Miguel Parrent of the

Royal Museum for Central Africa (MRAC); James Maclaine, Patrick Chapman, and Dr.

Julia Day of the Natural History Museum (BMNH); Drs. Paul Skelton and Roger Bills of

the South African Institute of Aquatic Biodiversity (SAIAB); and Dr. John Friel of the

Cornell University Museum of Vertebrates (CU). I additionally thank Dr. Julia Day

(BMNH) for her generosity in discussing her as-yet unpublished findings regarding the

phylogeny of Tanganyikan Synodontis with me. My study benefited from 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.









Permission to use the photographs seen in Figure 19 was arranged by K.E. Hartel of the

Museum of Comparative Zoology, Harvard University.
















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ................................................................................................. iii

LIST OF TABLES .................. .............. ................ ....... .. .............. viii

LIST OF FIGURES ......... ......................... ...... ........ ............ ix

ABSTRACT .............. ......................................... xi

CHAPTER

1 IN TR OD U CTION ............................................... .. ......................... ..
N atu ral H history ....................................................................... 1
System atic H history ...................................................... ..... .. ... ........ ..

2 M ATERIALS AND M ETHOD S ........................................... ........... ............... 6
D ata C collection ................................................ ........................... 6
C characters E xam ined ................................................................ .. ..................

3 MATERIALS EXAMINED ............ ..... .............. ............... 16
Synodontis granulosus ............................................................................. 16
Synodontis m ultipunctatus ............................ ......... ......................... ............... 17
Sy n od on tis sp 1 .................................................................19
Synodontispetricola ......................... ............ ............ ......... 20
Sy n od on tis sp 2 ................................................................2 1
Synodontis nigrom aculatus.............. ................ ................ ................. ............... 21
Synodontis tanganaicae ............... ..................... .. ........................ ............... 23
Synodontis dhonti................. ...................................24
Sy nodon tis p olli .................................................................2 5
Sy n od on tis sp 3 ................................................................2 7

4 SPECIE S D E SCRIPTION S............................................... ............................. 28
In v aria n t T raits..................................................................................................... 2 8
Synodontis granulosus Boulenger, 1900 ....................................... ............... 28
Diagnosis ............................................................................ ......... ................. 29
D description ......................................... 30
C o lo ratio n ................................................................................... 3 1
Distribution ............... .............................. 31
H a b ita t ................................................................................................................ 3 2


v









D ie t ........................................................................................................ 3 2
R ep ro du ctio n ....................................................................... 3 2
Taxonomic Remarks..................................................................................... 32
Synodontis multipunctatus Boulenger, 1898 .................................... ............... 36
D diagnosis ................................................................................................... 36
Description ......................................... 37
C o lo ratio n ..................................................... 3 9
Distribution..................... ........ 39
H a b ita t ................................................................................................................ 4 0
D ie t ................................................................... 4 0
R ep ro du ctio n ....................................................................... 4 0
T axonom ic R em arks ....................................................................... 4 1
Sy n o d o n tis sp 1 n sp ............................................................................................. 4 5
Diagnosis .................................................................................................. ........ 45
Description ......................................... 45
C o lo ratio n ..................................................... 4 7
Distribution..................... ........ 47
H a b ita t ................................................................................................................ 4 8
D ie t ................................................................... 4 8
R ep ro du ctio n ....................................................................... 4 8
T ax onom y .................................................... 4 8
Synodontispetricola Matthes, 1959 .............................. ...............52
Diagnosis .................................................................................................. ........ 52
Description ......................................... 53
C o lo ratio n ..................................................... 54
Distribution ......................................... 55
H a b ita t ................................................................................................................ 5 5
D ie t ........................................................................................................ 5 5
R ep ro du ctio n ....................................................................... 5 5
Taxonomic Remarks........................................................ 55
Synodontis sp. 2 n. sp...........................................................60
D diagnosis ................................................................................................... 60
Description ......................................... 61
C o lo ratio n ..................................................... 6 2
Distribution ......................................... 63
H a b ita t ................................................................................................................ 6 3
D ie t ........................................................................................................ 6 3
R ep ro du ctio n ....................................................................... 6 3
Taxonomic Remarks ............................................................... 63
Synodontis nigromaculatus Boulenger, 1905 ........................................ ............ 67
Diagnosis ......................................................................... ......... ......... ......... 68
Description ......................................... 69
C oloration ............................................... 7 0
Distribution..................... ........ 70
H a b ita t ................................................................................................................ 7 0
D ie t ................................................................... 7 1









R ep ro du ctio n ....................................................................... 7 1
Taxonomic Remarks......................................................... 71
Synodontis tanganaicae Borodin, 1936................................................................. 78
D diagnosis .............................................................. 78
D description ......................................... 79
C o lo ratio n ................................................... ................................ 8 0
Distribution ............... .............................. 81
H a b ita t ................................................................................................................ 8 1
D ie t ............. ............................ .. ........................................................ 8 1
Reproduction ............................................. 81
T axonom ic R em arks.................................................................. 81
Synodontis dhonti Boulenger, 1917.................................................... 87
D iag n o sis ................................................ 87
D description ......................................... 88
C o lo ratio n ..................................................... 8 9
Distribution..................... ........ 90
H a b ita t ................................................................................................................ 9 0
D ie t ................................................................... 9 0
R ep ro du ctio n ....................................................................... 9 0
Taxonomic Remarks........................................................ 91
Synodontispolli Gosse, 1982..................................................................97
D diagnosis .................................................................................................. ........ 97
D description ......................................... 98
C o lo ratio n ..................................................... 9 9
D istrib u tio n .............................................................................. 10 0
H ab itat ........................................ 10 0
D ie t ....................................................................................................... 1 0 0
R ep ro du ctio n ....................................................................... 10 0
Taxonomic Remarks......................................... 101
Synodontis sp 3 n sp ..........................................................105
D diagnosis .......................................................................................... ........ 105
D description ......................................... 106
C o lo ratio n ............................................................................... 10 7
D istrib u tio n .............................................................................. 10 8
H ab itat ........................................ 10 8
D ie t ....................................................................................................... 1 0 8
R ep ro du ctio n ....................................................................... 10 8
Taxonomic Remarks......................................... 108

5 D ISC U S SIO N ...................................................... 114

L IST O F R E F E R E N C E S ............................................................................................ 118

BIOGRAPHICAL SKETCH .............................................................................124
















LIST OF TABLES


Table pge

1 Diagnostic characters for Lake Tanganyikan Synodontis species in which an
axillary pore is present. ............................... .... .......... ................ ............. 10

2 Diagnostic characters for Lake Tanganyikan Synodontis species that lack an
ax illa ry p o re ...................................... ............. ................ ................ 1 1

3 Morphometric data and meristic counts for Synodontis granulosus........................33

4 Morphometric measurements and meristic counts for Synodontis multipunctatus.. 42

5 Morphometric and meristic counts for Synodontis sp. 1......................................49

6 Morphometric measurements and meristic counts for Synodontispetricola ..........57

7 Morphometric and meristic counts for Synodontis sp. 2......................................64

8 Morphometric and meristic counts for Synodontis nigromaculatus. .....................74

9 Morphometric and meristic counts for Synodontis tanganaicae.............................83

10 Morphometric measurements and meristic counts for Synodontis dhonti ..............93

11 Morphometric measurements and meristic counts for Synodontispolli..............102

12 Morphometric and meristic counts for Synodontis sp. 3............. ................109
















LIST OF FIGURES


Figure pge

1 Map of the African continent with the Lake Tanganyika region indicated by the red
box....... .............. ................................. ........... 5

2 Axillary pore of Synodontis multipunctatus. BMNH 1898.9.9.76, 280 mm TL, 220
m m SL ...............................................................................12

3 Humeral processes of Synodontis granulosus, S. multipunctatus, Synodontis sp. 1,
and S. petricola. An axillary pore is present in all of these species.....................13

4 Humeral processes of Synodontis sp. 2, S. nigromaculatus, S. tanganaicae, and a
paratype of S. lacustricolus. An axillary pore is absent in A and B....................14

5 Humeral processes of Synodontis dhonti, a juvenile specimen of S. dhonti, S. polli,
and Synodontis sp. 3. An axillary pore is absent in C and D. ............. ............... 15

6 Synodontis granulosus, UF 160945, 192 mm TL, 149 mm SL. Top left dorsal
view, bottom left lateral view, right ventral view of head..............................34

7 Distribution of Synodontis granulosus. T denotes type locality ...........................35

8 Synodontis multipunctatus, Holotype, BMNH 1898.9.9.76, 280 mm TL, 220 mm
SL. Top left dorsal view, bottom left lateral view, right ventral view, showing
m south and barbels. ....................... ...................... ................... .. ..... 43

9 Distribution of Synodontis multipunctatus. T denotes type locality.....................44

10 Synodontis multipunctatus (top), BMNH 1982.4.13.4789, 115 mm TL, 91 mm SL
vs. Synodontis sp. 1 (bottom), BMNH 1982.4.13.4784, 105 mm TL, 81 mm SL ...50

11 Distribution of Synodontis sp. 1. T denotes type locality. .......................................51

12 Synodontispetricola, Holotype, MRAC 130357, 98 mm TL, 82 mm SL. Top -
dorsal view, middle lateral view, bottom ventral view.................. .............58

13 Distribution of Synodontispetricola. T denotes type locality. ............................59

14 Synodontis sp. 2, SAIAB 39577, 84 mm TL, 68 mm SL. Top dorsal view, middle
lateral view, bottom ventral view. ........................................... ............... 65









15 Distribution of Synodontis sp. 2. T denotes type locality. ...................................... 66

16 Variation in body shape and coloration in Synodontis nigromaculatus ................75

17 Dorsal views of the head of left Synodontis melanostictus (Holotype) and right -
Synodontis nigromaculatus (Holotype), showing differences in occipito-nuchal
shield texture. ...................................................... ................. 76

18 Distribution of Synodontis nigromaculatus. T denotes type locality ......................77

19 Synodontis tanganaicae, Lectotype, MCZ 32538, 415 mm TL, 320 mm SL. Top
left dorsal view, bottom left lateral view, right ventral view of head ..............84

20 Synodontis tanganaicae, MRAC 90288, paratype of S. lacustricolus, 519 mm TL,
434 mm SL. Top left dorsal view, bottom left lateral view, right ventral view
of head ....................... .................. ................... .... ..... ........... ......... 85

21 Distribution of Synodontis tanganaicae. Type locality unknown...........................86

22 Synodontis dhonti, Holotype, MRAC 14344, 395 mm TL, 325 mm SL. Top left -
dorsal view, bottom left lateral view, right ventral view, showing mouth and
b arb els. ........................................................................... 9 4

23 Synodontis dhonti, MRAC 130315, Holotype of S. irsacael42 mm TL, 116 mm
SL. Top dorsal view, middle lateral view, bottom ventral view....................95

24 Distribution of Synodontis dhonti. T denotes type locality..................................96

25 Synodontispolli, Holotype, MRAC 130440, 148 mm TL, 125 mm SL. Top -
dorsal view, middle -lateral view, bottom ventral view................ ................103

26 Distribution of Synodontispolli. T denotes type locality................................... 104

27 Synodontis sp. 3, UF 160942, 139 mm TL, 116 mm SL. Top dorsal view, middle
lateral view, bottom ventral view. ............... ......................... .............. 110

28 S. polli, MRAC 131007-008, 137 TL, 112 SL, with intestine extracted. Hindgut
chamber indicated with white arrow. ..... ................... .................111

29 Synodontis sp. 3, UF 160942, 139 mm TL, 118 mm SL, with intestine extracted.
Note the relative shortness of the intestine and lack of a discernible hindgut
ch am b er. ......... .... .............. ..................................... ........................... 112

30 Distribution of Synodontis sp. 3. T denotes type locality. .................................... 113
















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

TAXONOMIC REVISION OF LAKE TANGANYIKAN Synodontis (TELEOSTEI:
MOCHOKIDAE)

By

Jeremy John Wright

May 2006

Chair: Lawrence M. Page
Major Department: Zoology

Seven species of Synodontis have historically been recognized from Lake

Tanganyika, six of which are endemic to the lake basin. The endemic species can be

distinguished from all other Synodontis by a common pattern of rayed fin coloration and

vertical folds in the skin. Though previous studies have examined these species, a

comprehensive taxonomic examination utilizing all of the material available has not been

performed.

The taxonomy of the Synodontis of Lake Tanganyika was investigated through by

examining 336 museum specimens. Morphometric, meristic, and nontraditional internal

and external characters were examined, resulting in the recognition of 10 Synodontis

species from Lake Tanganyika (nine of which are endemic and three of which are new to

science). Synodontis sp. 1 is most similar to S. multipunctatus and is distinguished from

this species 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 sp.









2 is most similar to S. petricola and is distinguished from this species by the lack of an

axillary pore and the presence of light colored windows at the bases of the rayed fins.

Synodontis sp. 3 is most similar to S. polli and is distinguished from this species by the

absence of a hindgut chamber, the shortness of the gut (0.8-1.4 times total length in S.

polli vs. 4.0-5.5 times total length in S. polli), and the presence of short, flattened papillae

on the skin (vs. villous papillae in S. polli).

Examination of the data obtained during this study offers some support for recent

molecular phylogenies obtained for this group, but additional studies are needed to

construct a clearer picture of relationships within this group and between this group and

other members of the genus. Functional studies of many characters examined in this

study must also be performed to further our understanding of the biology of these species.














CHAPTER 1
INTRODUCTION

Natural History

Catfishes of the genus Synodontis are small to moderately sized fishes (up to 800

mm) that occur throughout sub-Saharan Africa. Poll (1971) divided the genus into eight

geographically defined groups: the northern supraequatorial group; the exclusively

Nilotic group; the group of species occurring exclusively in the Niger; the western

supraequatorial group from Senegal to Ogowe; the Congo Basin group; the southernmost

subequatorial group; the Synodontis of eastern Africa ranging between Uebi Shebeli and

the Rufigi River; and the Lake Tanganyikan group (Fig. 1).

The Synodontis of Lake Tanganyika inhabit mainly rocky shoreline areas in the

lake but also venture out over sandy and shell bottoms (Matthes, 1962; Coulter, 1991).

The vertical distribution of these species is limited to a maximum depth of 50-100 m in

the northern end of the lake and to about 240 m in the southern end (Coulter and 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 and Spigel, 1991;

Tiercelin and Mondeguer, 1991). Locality data indicate that all species have a lakeside

distribution, with the exception of Synodontis sp. 2 and 3 (see Chapters 3, 4).

Although many different color patterns exist within this genus, few are as striking

or as instantly recognizable as the pattern typical of most Tanganyikan Synodontis

species. This pattern consists of dark triangles at the bases of all of the rayed fins









andlarge, dark colored 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 (max. TL 100

mm in Synodontis sp. 2) to quite large (max. TL 600 mm in S. tanganaicae).

Sexual dimorphism is known to occur in Synodontis species in the form of a

distinct, conically shaped genital papilla present only in males (Matthes, 1962). Minor

differences in body shape between sexes also occur in some species, with females having

slightly more robust bodies. Information on reproduction is largely undocumented in

Tanganyikan Synodontis species, with information being mostly restricted to egg counts

from gravid females, though some species lack even that. The glaring exception to this

statement is S. multipunctatus, whose unique brood parasitism behavior has been well

documented (Sato, 1986). Dietary information is slightly more abundant, and a variety of

prey items are utilized. Most species include 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, crustaceans, and the eggs of other fishes (Poll, 1946, 1953;

Matthes, 1962; Coulter, 1966, 1991).

Systematic History

Synodontis are members of the Mochokidae, a family of catfishes endemic to sub-

Saharan Africa. With approximately 120 currently recognized species, Synodontis is the

most diverse of any African catfish genus and, with the exception of Barbus, of any

African Ostariophysan. Poll (1971), the last to revise Synodontis, recognized as valid

seven species from Lake Tanganyika: Synodontis multipunctatus Boulenger, Synodontis

granulosus Boulenger, Synodontis dhonti Boulenger, Synodontispetricola Matthes,









Synodontis lacustricolus Poll (now Synodontis tanganaicae Borodin), Synodontis

eurystomus Matthes (now Synodontispolli Gosse), and Synodontis nigromaculatus

Boulenger. All but S. nigromaculatus are endemic to the lake. The nine endemic

Synodontis species of Lake Tanganyika recognized herein can be distinguished from all

other Synodontis species by the presence of a black triangle at the base of each rayed fin

and vertical epidermal folds along the sides of the body.

Relationships between the endemic Synodontis species of Lake Tanganyika have

yet to be intensively studied. Brichard (1978) indicates that the lake species of

Synodontis probably do not form a monophyletic group. While it is possible that

S. nigromaculatus does not share a most recent common ancestor with the

Tanganyikan endemics, given its much wider distribution and differences from the

endemic species in its rayed fin coloration and skin (lack of vertical folds), Brichard

offers no further support for his opinion. The common pattern of rayed fin coloration and

presence of vertical folds in the skin would appear to represent two synapomorphies

supporting the monophyly of the endemic Synodontis species of Lake Tanganyika;

however, a preliminary investigation of the molecular phylogeny of the endemic

Tanganyikan Synodontis species, utilizing mitochondrial (cyt b) data appears to support

Brichard's statement (Day, pers. comm.). These results are discussed later in Chapter 5.

Poll's revision of Synodontis, while thorough for its time, suffered from a lack of

material, and also appears largely 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, historically poorly






4


sampled for Synodontis, as well as the increased popularity of certain Tanganyikan

Synodontis as aquarium species, led to the availability of increased numbers of specimens

for study. Schraml (2003) suggested that many of the specimens entering the aquarium

trade may represent cryptic, undescribed species. The aim of this study was to evaluate

the taxonomy of this group, bearing in mind that cryptic species may exist. This led to

the utilization of several unconventional diagnostic characters that revealed undescribed

Synodontis species from Lake Tanganyika.



































-20 '" 20 40




2 0' -20'







-20' 0' 20' 40'
km
0 5001000



Figure 1. Map of the African continent with the Lake Tanganyika region indicated by the
red box.















CHAPTER 2
MATERIALS AND METHODS

Data Collection

Measurements were taken from the left side to the nearest 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 six, fused

vertebrae of the Weberian apparatus excluded. Notation for fin ray counts is as follows:

upper case roman numeral = fin spine, lower case roman numeral = unbranched rays,

Arabic numeral = branched fin rays. Terminology for premaxillary dentition follows

Skelton and White (1990).

Gut length measurements and observations of the hindgut chamber of all species

were performed only on specimens with a belly cut made prior to the beginning of this

study with the exception of UF specimens which were dissected over the course of this

study. 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. All maps were created using the

Online Map Creator (OMC), which can be found at http://www.aquarius.geomar.de/omc/.

Distribution points on maps were added using Adode Photoshop CS2. Points indicate

only specimens examined in this study. Institutional abbreviations follow Leviton et al.

(1985) except for the South African Institute of Aquatic Biodiversity (SAIAB).









Characters Examined

In addition to the 22 morphometric measurements and seven meristic counts which

were made for all specimens, several new characters were found to be useful in the

diagnosis of species. 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 distinguish

species. These new characters are particularly appealing because they allow for easy,

accurate visual identification of specimens. Some of these characters, such as fin spine

color are self explanatory. Those that may be unfamiliar or that are used in a taxonomic

capacity for the first time in this study 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 (which is itself a useful structure in identification) (Figs. 2-5). The

function of this structure is unknown in mochokids, and the absence or presence of this

structure has been noted in only one previously described Synodontis species (Friel and

Vigliotta, 2006). The presence of a similar structure has previously been noted in six

other catfish genera: Ariopsis (Ariidae) (Halstead et al., 1953), Ameiurus, Ictalurus, and

Noturus (Ictaluridae) (Reed, 1907; Birkhead, 1967, 1972), Ac i ,I /,l ,i c ,i iJi /11/1

(Akysidae) (Ng and Ng, 2001), and Glyptothorax (Sisoridae) (pers. obs.). In

Aci ic i,,1 iu1ui k/1i1y\, this structure has been shown to produce a mucosal secretion that

possesses toxic properties (Ng and Ng, 2001), while 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 ten Tanganyikan species of Synodontis into two groups; four species which

have an axillary pore (Table 1) and six in which it is lacking (Table 2).









All Synodontis species have a premaxillary toothpad, which has 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 transversely bisects the toothpad. The separated condition is referred to

herein as interrupted, while the unseparated condition is referred to as uninterrupted.

This appears to be the first study to examine this character in Synodontis, though the

length of the premaxillary pad has often been used in previous studies.

The skin of the nine endemic Tanganyikan Synodontis species recognized in this

study is unique among the genus in having numerous minute, vertical epidermal folds.

The purpose of these folds is unknown. These folds are useful in distinguishing members

of the Tanganyikan species flock from species outside the lake's basin, though they are

not helpful in distinguishing between species from the lake. The papillae on the skin of

certain species, however, help to separate 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 over time.

The intestinal anatomy of some Synodontis species is extremely interesting due to

the presence of a swelling in the posterior part of the intestine which Taverne and

Aloulou-Triki 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 fermentative digestion in the hindgut (Rimmer

and Wiebe, 1987; Clements and Choat, 1997; Douglas et al., 2002). The presence of this

structure in highly herbivorous mochokid species such as S. polli, Brachysynodontis

batensoda, and Hemisynodontis membranaceus suggests that a similar role may be






9


played by the mochokid hindgut chamber, marking the first time that such a behavior and

structure were 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 external diagnostic characters exist.















Table 1. Diagnostic characters for Lake Tanganyikan Synodontis species in which an axillary pore is present.


S. granulosus S. multipunctatus Synodontis sp. 1 S. petricola S. tanganaicae S. dhonti
Mandibular 28 51 13 29 17 26 31 50 33 49 15 38
teeth
Axillary pore Large Large Large Small Large Large
Large spots on Absent Present Present Present Absent Present- young
body Absent adults
Fin spine color Dark Dark Dark White Dark Dark
Pectoral-fin rays 7 8 8 7 8 9 8 9 8- 9
Eye/Snout 31.2 50.2% 44.9 62.0% 64.2 81.0% 28.7 40.1% 16.0 26.9% 23.1 43.7%
length
Premaxillary Uninterrupted Uninterrupted Uninterrupted Interrupted Interrupted Interrupted
toothpad
Maximum TL 270 mm 280 mm 150 mm 135 mm 600 mm 400 mm


Secondary
branches on
medial
mandibular
barbel
Occipito-nuchal
shield covered
with skin
Papillae on body
skin
Gut/body length
Hindgut
chamber


Absent




No


Present, granular

0.5 times

Absent


Absent




Yes


Absent

0.5-0.8 times

Absent


Absent




Yes


Absent

0.8-0.9 times

Absent


Present




Yes


Absent

1.2-1.3 times

Present


Present




No


Present, granular

No data

No data


Present




No

Present, granular

0.7 0.8 times

Absent







11


Table 2. Diagnostic characters for Lake Tanganyikan Synodontis species that lack an
axillary pore.
Synodontis sp. S. nigromaculatus S. polli Synodontis sp. 3
2


Mandibular teeth

Spots on body

Fin spine color

Black triangles on
rayed fins

Eye/Snout length

Premaxillary
toothpad
Maximum TL

Secondary branches
on medial
mandibular barbel

Occipito-nuchal
shield covered with
skin
Papillae on body
skin

Gut/body length

Hindgut chamber


38-51

Large

White


Present, with
light windows
at base
25.2- 35.8%

Interrupted

100 mm


Present



Yes


Absent

No data

No data


23 -38


Small

Dark


Absent


31.1 -48.1%

Interrupted

380 mm


Present



No


Present, villous

1.3-1.6 times


Present


40 70
Moderate to
large
Dark


Present


25.8-39.3%

Interrupted

180 mm


Present



Yes


Present,
villous


4.0-4.5 times


Present


50 66


Moderate

Dark


Present


26.0 33.4%

Interrupted

150 mm


Present



Yes


Present, flattened


0.8-1.4 times


Absent


* Certain traits not diagnostic within this group (premaxillary toothpad, secondary
branches on medial mandibular barbel) are included to facilitate comparison with species
in Table 1.



















7


Figure 2. Axillary pore of Synodontis multipunctatus. BMNH 1898.9.9.76, 280 mm TL, 220
mm SL.







































Ir I %


Figure 3. Humeral processes of A) Synodontis granulosus, UF 160945, 192 mm TL, 149
mm SL, B) S. multipunctatus, BMNH 1898.9.9.76, 280 mm TL, 220 mm SL,
C) Synodontis sp. 1, 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 axillary
pore is present in all of these species.


*i i ". r
*r.
ilr






















































Figure 4. Humeral processes of A) Synodontis sp. 2, SAIAB 42518, 97 mm TL, 78 mm
SL, B) S. nigromaculatus, BMNH 1905.11.10.10, 237 mm TL, 186 mm SL,
C) S. tanganaicae, 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 all of these species.





















































Figure 5. Humeral processes of A) Synodontis dhonti Holotype, MRAC 14344, 395 mm
TL, 325 mm SL, B) juvenile specimen of S. dhonti, 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) Synodontis sp. 3, UF 160942, 139 mm TL, 118 mm
SL. An axillary pore is absent in all of these species.















CHAPTER 3
MATERIALS EXAMINED

Number of specimens examined from each lot is indicated in parentheses.

Synodontis granulosus

(1) BMNH 1906.9.6.40, Lectotype, TL 270 mm, SL 216 mm, north end of Lake

Tanganyika, coll. J.E.S. Moore, (2) BMNH 1906.9.6.41-42, Paralectotypes, TL 220-246

mm, SL 181-202 mm, (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 (Zambia), 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, Tropical Fish Collector, coll. V.2005.









Synodontis multipunctatus

(1) BMNH 1898.9.9.76, Holotype, TL 280 mm, SL 220 mm, Sumba, coll. J.E.S.

Moore, (2) BMNH 1906.9.8.70-1, 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. Belge, (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. Belge, 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. Nat. Belge, 7.III.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.III.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. Belge, 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 (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 39186-39187,

TL 94-96 mm, SL 72-73 mm, Moba (Lake Tanganyika), coll. Van Maldern, 1932, (3)

MRAC 53096-53098, 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 90273-90274, 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 91606-91608, 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, total length 75-105 mm,

SL 60-83 mm, Magara, route Bujumbura-Nyanza Lac (Burundi), coll. L. De Vos,

19.10.1994, (1) MRAC A1-094-P-0054, TL 80 mm, SL 61 mm, Kasakalawe, Lake

Tanganyika, S 8047.23', E 3104.40' (Zambia), coll. Snoeks, Hanssens, Verheyen et al.,

(2) SAIAB 39578, TL 77-85 mm, SL 62-67 mm, Country: Zambia, System: Lake

Tanganyika; Mpulungu, Musende Rocks, 0846'00"S, 031051'00"E, coll. 7.VII.1992, (3)

SAIAB 40174, TL 58-96 mm, SL 45-76 mm, Country: Zambia, System: 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, Tanzania, 0454'31"S, 029036'02"E, coll. 04.X.1997, (4) SAIAB 56254, TL 82-

103 mm, SL 63-81 mm, Jacobsen's beach, Tanzania, 0454'31"S, 029036'02"E, coll.









05.X.1997, (1) SAIAB 76164, TL 78 mm, SL 63 mm, Zambia, Province: Northern, Area:

Mbala, System: Lake Tanganyika; Mbita Island (northwest end), 845.18'S, 31005.07'E,

coll. 29.II.2004, (3) SAIAB 76175, TL 56-76 mm, SL 44-59 mm, Country: Zambia,

Province: Northern, Area: Mbala, System: Lake Tanganyika; Musende Rocks beach,

08045.18'S, 31005.07'E, coll. 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. Wagner, Catherine,

13.VIII.2005.

Synodontis sp. 1

(2) BMNH 1920.5.25.83-84, TL 98-103 mm, SL 76-79 mm, Vua Bay, L.

Tanganyika, coll. L. Stappers, (1)BMNH 1955.12.20.1833-1835, TL 103 mm, SL 80

mm, Moba Bay, L. Tanganyika, coll. Inst. Roy. Sci. Nat. Belge, 7.III.1947, (1) BMNH

1955.12.20.1837, TL 101 mm, SL 78 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 1982.4.13:4789-4793, TL 109-117 mm, SL 86-

92 mm, Kigonga Bay, L. Tanganyika, coll. R. Travers, (2) BMNH 1982.4.13.4785-4786,

TL 112-129 mm, SL 91-96 mm, Mwakizega coastline, L. Tanganyika, coll. R. Travers,

(1) BMNH 1982.4.13:4787, TL 103 mm, SL 80 mm, Kibirzi beach, 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, (1) BMNH 2005.9.26.11-16, TL 94 mm,

SL 73 mm, Kigoma, Tanzania, coll. J. Day, 2005, (3) MRAC 39186-39192, TL 93-100

mm, SL 72-80 mm, Moba (Lake Tanganyika), coll. Van Maldem, 1932, (2) MRAC

53096-53100, TL 107-114 mm, SL 80-91 mm, Nyanza (L. Tang.), coll. A. Testiade,

1937, (3) MRAC 90273-90277, TL 123-147 mm, SL 96-111 mm, Stat: 63, offshore from









the Malagarasi, 10-15 km from the coast, coll. M. Poll, 3.II.1947, (2) MRAC 91606-

91608, TL 114-121 mm, SL 96-98 mm, Uvira, Lake Tanganyika, coll. G. Marlier,

1.IX.1949, (1) SAIAB 56254, TL 107 mm, SL 81 mm, Jacobsen's beach, Tanzania,

04054'31"S, 029036'02"E, coll. 05.X.1992.

Synodontis petricola

(1) MRAC 130357, Holotype, TL 98 mm, SL 82 mm, Kashekezi, N. Lake

Tanganyika, coll. H.Matthes, 3.IV.1959, (9) Paratypes, 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.I.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, 08020.24'S, 30031.21'E, Chisiki, coll. Verheyen, Snoeks,

Hanssens, Ruber, Sturmbauer, 10.IV.1995, (1) MRAC A1-094-P-53, TL 81 mm, SL 65

mm, Crocodile Island, Lake Tanganyika (Zaire), coll. Snoeks, Hanssens, Verheyen et al.,

16.X.2001, (1) BMNH 1936.6.14.1232, Paratype, (also I.D.'d as S.multimaculatus), TL

88 mm, SL 71 mm, Lake Tanganyika, coll. Christy, (2) SAIAB 42518, TL 70-74 mm, SL









56-59 mm, Country: Zambia, System: Lake Tanganyika; Musende Rocks, 0846'00"S,

031007'00"E, coll. 10.X.1992, (1)SAIAB 56255, TL 68 mm, SL 55 mm, Country:

Tanzania; Jacobsen's beach, 0454'31"S, 029036'02"E, coll. 05.X.1997, (1) SAIAB

56225, TL 75 mm, SL 61 mm, Country: Tanzania; Kigoma Bay, below Hill top Hotel,

04053'03"S, 029037'11"E, coll. 09.X.1997, (1) SAIAB 56232, TL 80 mm, SL 64 mm,

Country: Tanzania; Cave, Kigoma Hotel below Hill top, 04053'03"S, 029037'11"E, coll.

11.X.1997, (1) SAIAB 56244, TL 79 mm, SL 64 mm, Country: Tanzania; Bangwe,

shingle beach, 04053'53"S, 029035'39"E, coll. 12.X. 1997.

Synodontis sp. 2

(9) SAIAB 39577, TL 58-98 mm, SL 47-79 mm, Zambia; Mpulungu, Musende

Rocks, 08046'00"S, 031051'00"E, coll. 07.VII.1992, (3) SAIAB 40171, TL 70-85 mm,

SL 56-69 mm, Country: Zambia; Musende Rocks, 08046'00"S, 031006'00"E, coll.

05.VIII.1992, (4) SAIAB 42518, TL 55-97 mm, SL 44-78 mm, Country: Zambia,

System: Lake Tanganyika; Musende Rocks, 08046'00"S, 031007'00"E, coll. 10.X. 1992.

Synodontis nigromaculatus

(1) BMNH 1905.11.10.10, Holotype, TL 237 mm, SL 186 mm, Lake Bangwuelo,

Zambia, coll. Esq. R. Melland, (1) BMNH 1906.9.8.72, Holotype (S. melanostictus), TL

290 mm, SL 237 mm, Lake Tanganyika, coll. Cunnington, (1) BMNH 1908.11.6.29, TL

244 mm, SL 177 mm, Maramba River, Upper Zambezi, coll. G.T. Corrington, (1) BMNH

1907.9.30.9, TL 319 mm, SL 238 mm, Lake Bangwelo, coll. R.H. Melland, (4) BMNH

1920.5.26.95-102, TL 58-115 mm, SL 46-93 mm, Cape Kasangeneke, Lake Mweru, coll.

L. Stappers, (1) BMNH 1976.3.18: 2439-2440, TL 192 mm, SL 153 mm, Thalamakane

River at Maun (Botswana), coll. K. Berne, (1) MRAC 34006, TL 383 mm, SL 307 mm,

Lukonzolwa, Lake Moero, coll. DeWitte, 9-17.II.1931, (1) MRAC 34007, TL 324 mm,









SL 250 mm, Lukonzolwa, Lake Moero, coll. DeWitte, 9-17.II.1931, (2) MRAC 37814-

815, TL 177-317 mm, SL 134-252 mm, Kasenga, Luapula River, coll. DeWitte,

10.III.1931, (1)MRAC 56319-56320, TL 132 mm, SL 104 mm, Luapula au Bangwelo,

coll. Breolo, III.1938, (2) 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) MRAC 87132-133, TL 154 mm,

SL 129 mm, Kasaji, Katanga, coll. Rev. S. Fischer, 5.XII.1952, (1) MRAC 162219, TL

183 mm, SL 141 mm, Lake Calundo, coll. indigenous peoples, I.1955, (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 162210-213, TL 274-306 mm, SL 209-229 mm,

Chiumbe River, near the junction with the Chilambo River (S745', E21005') (Angola),

coll. Barros Machado (Dundo Museum), 24.XI.1957, (2) MRAC 126293-294, TL 152-

172 mm, SL 122-134 mm, Uvira, Lake Tanganyika, coll. G. Marlier, 13.IX.1958, (1)

MRAC 149537, TL 181 mm, SL 142 mm, Luwohoshi River, affl. Ruashi River, village

of Shindaika, Katanga, coll. M. Lips, 25.III.1963, (1) MRAC 165806-807, TL 104 mm,

SL 80 mm, Chamberi/Chambezi River (Zambia), coll. H. Matthes, 16.VI.1965, (1)

MRAC 172442, TL 177 mm, SL 134 mm, Lake Bangwelo, coll. Bell-Cross, (1) MRAC

94-019-P-1007, TL 152 mm, SL 116 mm, South Mofwe Lagoon, Luapula Swamp

(Zambia), coll. P. Van Zweiten, 16.II.1993-26.III.1993, (1) 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) 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, (1) 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), III.1948, (1) MRAC 162206 (orig. I.D. S. zambesensis), TL 241 mm, SL 186

mm, Dundo, Luachimo River (Angola), coll. Barros Machado (Dundo Museum),

XI.1948, (1) 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) MRAC 162215 (orig. I.D. S. zambesensis), TL 180 mm, SL 136 mm,

Cachimo, Tuembe River, S806', E21029' (Angola), coll. A.L. Terreira (Dundo

Museum), 22.VIII.1961, (1) 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.

Synodontis tanganaicae

(1) MRAC 130720, Paratype (S. lacustricolus), TL 333 mm, SL 255 mm, Mboko

Island, N. Lake Tanganyika, coll. Marlier (I.R.S.A.C.), 7.II.1957, (1) MRAC 90288,

Paratype (S. lacustricolus), TL 519 mm, SL 434 mm, Tembwe Bay, Lake Tanganyika,

coll. M. Poll/Expl. Hydrobiog.Tang., 13.II.1947, (1) MRAC 90289, Paratype (S.

lacustricolus), TL 582 mm, SL 489 mm, Edith Bay, Lake Tanganyika, coll. M. Poll/Expl.

Hydrobiog. Tang., 14-15.II.1947, (2) MRAC 96-083-P-003-004, TL 300-330 mm, SL

240-265 mm, Zambia, coll. De Vos, 1996, (1) BMNH 1955.12.20.1849, Paratype (S.

lacustricolus), TL 385 mm, SL 315 mm, Opposite Kala, Lake Tanganyika, Inst. Roy.

Nat. Belge, (1) BMNH 1955.12.20.1850, Paratype (S. lacustricolus), TL 361 mm, SL 291

mm, Tembwe Bay, Lake Tanganyika, Inst. Roy. Nat. Belge, (1) BMNH 1955.12.20.1851,

Paratype (S. lacustricolus), 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.

Synodontis dhonti

(1) MRAC 14344, Holotype, TL 395 mm, SL 325 mm, Lake Tanganyika, Kilewa,

coll. Stappers, 18.V.1912, (1) MRAC 130315, Holotype (S. irsacae), TL 142 mm, SL

116 mm, Kalundu, N. Lake Tanganyika, coll. H. Matthes, 27.X.1959, (6) MRAC

130316-321, Paratypes (S. irsacae), 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 (Orig. I.D. S. polli), 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 mm-85 mm, Ulwile Island, northern shore, Lake

Tanganyika S7027.40' E3024.20' (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,

08043.31'S, 31008.01'E, coll. Verheyen, Snoeks, Hanssens, Ruber, Stermbauer,

4.IV.1995, (4) MRAC A3-033-P-0036-0040, TL 117-161 mm, SL 91-133 mm, Pemba, S.

of Uvira, Lake Tanganyika (Zaire), coll. L. De Vos, 18.VIII.1995, (1) MRAC A1-094-P-

0055-0059, TL 103 mm, SL 81 mm, Katoto, Lake Tanganyika (Zambia), coll. Snoeks,

Hanssens, Verheyen et al., 18.X.2001, (1)BMNH 1955.12.20.1859, TL 192 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, SL 70-97 mm, Country: Zambia, System: Lake Tanganyika, Locality: Mbita Island,









08045'00"S, 031006'00"E, coll. 08.VIII.1992, (1) SAIAB 56231, TL 133 mm, SL 108

mm, Cave, Kigoma hotel below Hill top, 04053'03"S, 029037'11"E, coll. 04.X. 1997, (1)

SAIAB 56255, TL 91 mm, TL 72 mm, Country: Tanzania; Jacobsen's beach,

04054'31"S, 029036'02"E, coll. 05.X1997, (2) SAIAB 56687, TL 94-97 mm, SL 75-79

mm, Country: Tanzania, System: Lake Tanganyika; Muzungu beach, 04054'53"S,

029035'58"E, coll. 10.X.1997, (2) SAIAB 76105, TL 93-98 mm, SL 76-80 mm, Country:

Zambia, Province: Northern, Area: Mbala, System: Lake Tanganyika, Water body: Lake

Tanganyika; Mbita Island (northwest end), 08045.18'S, 31005.07'E, coll. 29.II.2004, (1)

SAIAB 76174, TL 97 mm, SL 79 mm, Country: Zambia, Province: Northern, Area:

Mbala, System: Lake Tanganyika, Water body: Lake Tanganyika; Musende Rocks beach,

08045.18'S, 31005.07'E, coll. 29.11.2004, (2) UF 160940, TL 104-124 mm, SL 86-102

mm, Cape Chaitika, Lake Tanganyika, Zambia, via Pete Hauschner, Tropical Fish

Collector, coll. 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.

Synodontis polli

(1) MRAC 130440, Holotype, TL 148 mm, SL 125 mm, Luhanga, N. Lake

Tanganyika, coll. H. Matthes, 3.IV.1959, (3) MRAC 130444-446, Paratypes, TL 97-144

mm, SL 79-120 mm, Makobola, Lake Tanganyika, coll. H. Matthes (I.R.S.A.C.),

10.XI.1958, (2) MRAC 130442-443, Paratypes, TL 48-58 mm, SL 38-47 mm, Luhanga,

N. Lake Tanganyika, coll H. Matthes (I.R.S.A.C.), 31.VIII.1950, (1) MRAC 130464,

Paratype, TL 115 mm, SL 96 mm, Bemba, Lake Tanganyika, coll. H. Matthes

(I.R.S.A.C), 26.IV.1958, (2) MRAC 130452-453, Paratypes, TL 57-60 mm, SL 46-48

mm, Makobola, Lake Tanganyika, coll. H. Matthes (I.R.S.A.C.), 28.VI.1960, (2) MRAC

130462-463, Paratypes, TL 69-85 mm, SL 56-70 mm, Kashkezi, Lake Tanganyika, coll.









H. Matthes (I.R.S.A.C.), 5.11.1959, (6) MRAC 130454-461, Paratypes, 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-

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, S0457.87', E29035.86'

(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, S0843'31", E3108'01" [Zambia]; coll.

Exp. Tanganyika 95, 4.IV. 1995, (5) MRAC 95-096-P2576-2585, TL 90-116 mm, SL 74-

98 mm, Chisiki, S0820'24", E30031'21" [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, S0829'22", E30028'32" [Zambia], coll. Exp. Tanganyika 95,

10.IV. 1995, (1) MRAC 95-096-P-2609, TL 135 mm, SL 112 mm, Kasaba Bay, lake









Tanganyika, S0830'57", E30038'52" [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, S0808'47", E30058'14" [Zambia]; coll. Exp. Tanganyika 95, 19.IV.1995,

(1) MRAC 95-096-P-2627, TL 108 mm, SL 87 mm, Chikulina Bay, lake Tanganyika,

S0832'41", E30043'36" [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, Country: Zambia, System:

Lake Tanganyika; Musende Rocks, 08046'00"S, 031006'00"E, coll. 05.VIII.1992, (1)

SAIAB 42518, TL 105 mm, SL 87 mm, Country: Zambia, System: Lake Tanganyika;

Musende Rocks, 08046'00"S, 031007'00"E, coll. 10.X.1992, (2) SAIAB 56262, TL 122-

150 mm, SL 100-127 mm, Country: Tanzania; Jacobsen's Beach, 04054'31"S,

029036'02"E, 10.X.1997, (1) SAIAB 76167, TL 117 mm, SL 100 mm, Country: Zambia,

Province: Northern, Area: Mbala, System: Lake Tanganyika, Water body: Lake

Tanganyika; Mbita Island (northwest end), 08045.18'S, 031005.07'E, coll. 29.II.2004, (1)

CU 88758, TL 180 mm, SL 145 mm, Tanzania; Kigoma; Jacobsen's Beach, coll. P.B.

McIntyre, 18.VII.2002.

Synodontis sp. 3

(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, (10) UF 160942, TL 128-148 mm, SL

108-121 mm, Lake Tanganyika at Chaitika, Zambia, via Pete Hauschner, Tropical Fish

Collector, coll. V.2005.















CHAPTER 4
SPECIES DESCRIPTIONS

Invariant Traits

Lateral line complete and midlateral, extending onto base of caudal fin. Distinct

genital papilla present in male; absent in female. Anterior nostrils 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. Two ovaline patches of

minute, embedded teeth posterior to mandibular tooth row(s). Well developed, papillate,

membranous flap immediately posterior to premaxillary toothpad.

Synodontis granulosus Boulenger, 1900

(Figures 3,6,7; Tables 1, 3)

Synodontis granulosus Boulenger, G.A., 1900:480, description, n. sp., northern end of
Lake Tanganyika; 1901:308, description, northern end of Lake Tanganyika;
1901:149, pl. 16, description, northern end of Lake Tanganyika; 1911:413, fig.
311, description, northern end of Lake Tanganyika. Moore, J.E.S.,
1903:166, fig., northern Lake Tanganyika. Worthington, E.B. and Ricardo,
C.K., 1936:1067, 1077, 1101, note, Kabanga Bay, Lake Tanganyika. Poll, M.,
1946:219, description, Kilewa Bay; 1953:152, description, Lake Tanganyika;
1971:303, fig. 138, 139, pl. V, XI, description, Lake Tanganyika. Hulot, A.,
1950:170, Lake Tanganyika. Matthes, H., 1962:44, description, Lake
Tanganyika. Coulter, G.W., 1965-1966:34, abundance, Lake Tanganyika;
1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance, ecology, Lake
Tanganyika. Brichard, P., 1978:424, identification, Lake Tanganyika;
1989:478, 479, photo, identification. Sands, D., 1983:23, check-list. Daget,
J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:129, check-list. -
Burgess, W.E., 1989:576, pl. 94, Lake Tanganyika. Kobayagowa, M.,
1989:14, photo. Coulter, G.W., 1991a:152, 154, 182, 191, abundance, diet,
habitat, reproduction.









Diagnosis

Axillary pore present; mandibular teeth 28-51; body lacking large spots; fin spines

brown to black; 7-8 pectoral fin rays; black triangles present on base of all rayed fins; eye

31.2-50.2% snout length; premaxillary toothpad uninterrupted; secondary branches on

internal mandibular barbels absent; occipito-nuchal shield not covered with skin; granular

papillae present on skin of body; hindgut chamber absent; maximum TL 270 mm.

Synodontis granulosus can be distinguished from Synodontis sp. 2, S.

nigromaculatus, S. tanganaicae, S. dhonti, S. polli, and Synodontis sp. 3 by the presence

of an axillary pore. Synodontis granulosus differs from S. multipunctatus and Synodontis

sp. 1 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 Synodontis sp. 1), a smaller eye (31.2-

50.2% of snout length vs. 44.9-62.0% in S. multipunctatus and 64.2-81.0% in Synodontis

sp. 1), an occipito-nuchal shield which is not covered by skin, and body skin which is

covered by granular papillae. Synodontis granulosus is distinguished from S. petricola

by lacking large spots on the body and secondary branches on the internal mandibular

barbels, having a large axillary pore (vs. very small in S. petricola), darkly colored fin

spines, 7-8 pectoral fin rays (vs. 8-9 in S. petricola), an uninterrupted premaxillary

toothpad, much larger body size (max. length 270 mm vs. 135 mm in S. petricola), an

occipito-nuchal shield which is not covered by skin, and skin which is covered by

granular papillae. Large specimens of S. granulosus may be confused with S.

tanganaicae. However, the humeral process is much narrower in S. granulosus humerall

process length 253.8-437.2% of humeral process width in S. granulosus compared to

172.9-255.4% in S. tanganaicae) (figs. 3a, 4c,d), as well as the presence of a large









axillary pore in S. granulosus (fig. 2), vs. pore absent in S. tanganaicae, are sufficient to

separate these species.

Description

Morphometric and meristic data are given in Table 3. Maximum total length 270

mm, standard length 216 mm. Body compressed. Predorsal profile straight, interrupted

by convex ridge formed by orbit of eye. Preanal profile straight to gently convex. Skin

on body forming numerous vertical epidermal folds covered with minute, 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. Occipital-nuchal shield

rugose, not covered with skin; terminating posteriorly with narrow, rounded process on

either side of dorsal spine; ventrally with narrow, bluntly pointed process which extends

to upper margin of humeral process on either side of body. Eye located laterally; ovoid;

horizontal axis longest. Interorbital area flat.

Mouth subterminal; lips wide and papillate. Mandibular teeth 28-51, short,

unicuspid; arranged in single transverse row. Premaxillary toothpad uninterrupted;

primary, secondary and tertiary premaxillary teeth discrete; numerous; arranged in 4, 4,

and 2 irregular rows, respectively.

Maxillary barbel with narrow basal membrane; lacking branches or crenelations;

extending at least to base of pectoral fin. External mandibular barbel extending to point

just short of anterior margin of pectoral girdle, with 4-5 non-tuberculate branches;

lacking secondary branches. Internal mandibular barbel about 1/2 length of external

barbel; with 4-5 pairs of tuberculate branches; lacking secondary branches.









Dorsal fin 1,7-8; posterior margin straight. Dorsal 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 margin granulate in adults; many small antrorse

serrations along anterior margin in juveniles; posterior spine margin with large retrorse

serrations along entire length. Adipose fin well developed; 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-v,7-9,i; posterior margin nearly straight; base

located ventral to adipose fin. Caudal fin 1,7,8,1; forked; lobes pointed.

Humeral process narrow, elongated, granulous; possessing distinct ridge on ventral

margin; dorsal margin concave; terminating in sharp point (Fig. 3A). Large, dark-colored

axillary pore present just ventral to humeral process. Gut 0.5 times body length (n = 1,

UF160945). Hindgut chamber absent.

Coloration

Body slate-gray to olive brown (Fig. 6). Belly ranging from pale yellow to gray.

Scattered, small black spots present on juvenile specimens. Maxillary barbels white.

Mandibular barbels white, bases sometimes 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. 7); rare.









Habitat

Littoral to benthic zones over shell, sand and mud bottoms; to a maximum depth of

130 m (Coulter, 1991).

Diet

Almost nothing is known about the diet of this species, though one publication does

reports a single Limnothrissa in the stomach of one specimen (Worthington and Ricardo,

1933). Many other Tanganyikan species of Synodontis prey largely on mollusks and

algae scraped from rocky substrates. Presumably, S. granulosus has a similar diet, but

further study is clearly needed to confirm or refute this hypothesis.

Reproduction

No information exists regarding the reproduction of S. granulosus.

Taxonomic Remarks

Synodontis granulosus is sufficiently distinct that it is one of only two described

Tanganyikan species of Synodontis that have remained unaltered with respect to their

taxonomy since their original descriptions (the other being S. multipunctatus). Minor

differences in coloration pattern occur throughout its range (mainly in the color of the fin

margins), but they do not appear to follow any discernible pattern with respect to

geography or population.











Table 3. Morphometric data and meristic counts for Synodontis granulosus. All
morphometric measurements given in percent of base measurement. Meristic
data for Lectotype is identified by a "*" superscript. SL = Standard length,
HL = Head length, XMB = External mandibular barbel, IMB = Internal
mandibular barbel, HPL = Humeral process length, HPD = humeral process
depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth.
Numbers in parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


Lectotype
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


Range (n=17)
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


Meristics
Mandibular tooth count 28(1); 32(1); 36(2); 37(1); 42(1); 43(1);
44(2); 45(1); 46*(1); 48(1); 51(1)
Dorsal fin count I,7*(13); I,7,i(1); 1,8(3)
Pectoral fin count 1,7(3); 1,8(14)
Pelvic fin count i,6*(15); i,7(2)
Anal fin count iii,8*(2); iii,9(1); iv,6,i(1); iv,7(3);
iv,8(2); iv,8,i(1); iv,9,i(1); v,7(1);
v,7,i(1); v,8(3);
Caudal fin count i,7,8,i(5)
Total vertebrae count 36(3), 37(1)


MeanSD
25.22.8
28.81.4
38.91.4
30.03.7
31.1+4.7
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



































Figure 6. Synodontis granulosus, UF 160945, 192 mm TL, 149 mm SL. Top left dorsal
view, bottom left lateral view, right ventral view of head.







kHI]


REPUBLIC


li~A


Figure 7. Distribution of Synodontis granulosus. T denotes type locality.


Le-









Synodontis multipunctatus Boulenger, 1898

(Figures 2, 3, 8, 9, 10; Tables 1, 4)

Synodontis multipunctatus Boulenger, G.A., 1898:24, pl. 8, description, n. sp., Sumba
(Lake Tanganyika); 1899:95, note, Lake Tanganyika; 1901:313, pl. 15,
description, Sumbu, Kalambo, Moliro, Albertville; 1901:149, Kalambo (Lake
Tanganyika); 1906:553, Niamkolo (Lake Tanganyika); 1911:420, fig. 316,
description, Sumba, Kalambo, Niamkolo. Moore, J.E.S., 1903:166, fig.,
description, Sumbu (Lake Tanganyika). Worthington, E.B. and Ricardo, C.K.,
1933:1067,1077, 1101, note, diet, Usumbura, Kigoma, Kala (Lake Tanganyika). -
Borodin, N.A., 1936:10, Kasanga (Lake Tanganyika). David, L. and Poll, M.,
1937:267, Tembwe, Moba, Rumonge, Uvira. Poll, M., 1946:220, description,
Lake Tanganyika; 1953:155, pl. 6, fig. 4, description, diet, Lake Tanganyika. -
Hulot, A., 1950:170, diet, Lake Tanganyika. Lambert, J., 1960:26, figs 26 and
27, note. Matthes, H., 1962:5, photo, 45, note, sexual dimorphism, Uvira (Lake
Tanganyika). Coulter, G.W., 1965-1966:34, table, abundance at different depths,
Lake Tanganyika; 1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance,
ecology, Lake Tanganyika. Brichard, P., 1978:360,361,423, photos,
identification, Lake Tanganyika; 1989:476, 478, 481, 483, 474, 479, photos,
coloration and identification. Sands, D., 1983:23, 24, 76, check-list, diet, photo.
Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:134, check-
list. Sato, T., 1986:58-59, brood parasitism, Lake Tanganyika. Burgess, W.E.,
1989:190, pl. 74, 77, 78, 85, 94, behavior, Lake Tanganyika. Kobayagowa, M.,
1989:14, photo. Coulter, G.W., 1991a:152, 154, 182, 191, abundance, diet,
habitat, reproduction.

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; secondary

branches on internal mandibular barbels absent; occipito-nuchal shield usually covered

with skin; papillae on skin of body absent; hindgut chamber absent; maximum TL 280

mm.

Synodontis multipunctatus can be distinguished from Synodontis sp. 2, S.

nigromaculatus, S. tanganaicae, S. dhonti, S. polli, and Synodontis sp. 3 by the presence

of an axillary pore and the lack of papillae on the skin. 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. The number of mandibular teeth further separates S. multipunctatus from S.

petricola, Synodontis sp. 2, S. nigromaculatus, S. tanganaicae, S. polli, and Synodontis

sp. 3, as well as from S. granulosus (13-29 in S. multipunctatus vs. 31-50 in S. petricola,

38-51 in Synodontis sp. 2, 23-38 in S. nigromaculatus, 33-49 in S. tanganaicae, 40-70 in

S. polli, 50-66 in Synodontis sp. 3, and 28-51 in S. granulosus). Synodontis

multipunctatus further differs from S. granulosus in having large spots on the body and

an occipito-nuchal shield which is covered by skin. Synodontis multipunctatus can be

further distinguished from other endemic Tanganyikan species (with the exception of

Synodontis sp. 2) by its lack of well developed black triangles at the base of the pelvic

and anal fins. Synodontis multipunctatus and Synodontis sp. 1 are most reliably separated

by pectoral fin ray counts (8 unbranched elements in S. multipunctatus vs. 7 in Synodntis

sp. 1) and eye size (44.9-62.0% of snout length in S. multipunctatus vs. 64.2-81.0% in

Synodontis sp. 1).

Description

Morphometric and meristic data are given in Table 4. Maximum total length 280

mm, standard length 220 mm. Body moderately compressed. Predorsal profile straight.

Preanal profile convex. Skin on body forming numerous vertical epidermal folds;

papillae absent.

Head somewhat depressed and broad; skin smooth in all but largest specimens,

where ridges and granulations from underlying bone protrude. Snout subconical when

viewed laterally; rounded when viewed dorsally. Occipital-nuchal shield rugose;









completely covered with skin in smaller specimens (<150 mm), lacking skin in larger

specimens; terminating posteriorly with narrow, bluntly pointed process on either side of

dorsal spine; ventrally with wide, rounded process which extends to upper margin of

humeral process on either side of body, also covered with skin in smaller specimens.

Eyes dorsolateral; ovoid; horizontal axis longest. 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 uninterrupted;

primary, secondary and tertiary premaxillary teeth discrete, numerous, arranged in 3, 3,

and 2 irregular rows, respectively.

Maxillary barbel with or without thin basal membrane; lacking branches or

crenelations; extending anywhere from base of the pectoral fin to end of humeral process.

External mandibular barbel extending to point just beyond posterior margin of pectoral

girdle; with 4-6 long, non-tuberculate branches; lacking secondary branches. Internal

mandibular barbel 1/3 to 1/2 length of external barbel; with 3-4 pairs of non-tuberculate

branches; lacking secondary branches.

Dorsal fin 1,7; 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 spine 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 margin smooth in adult specimens; many small, antrorse serrations along anterior

margin in juvenile specimens; posterior spine margin with large, retrorse serrations along









entire length. Adipose fin short; poorly developed; margin convex. 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 distinct ridge on its ventral margin in young specimens; ridge

becoming indistinct in adults; dorsal margin concave; terminating in sharp point (Fig.

3B). Large, dark-colored axillary pore present just ventral to humeral process. Gut 0.5-

0.8 times body length (n = 4, MRAC 53096-53100, MRAC 90273-90277). Hindgut

chamber absent.

Coloration

Dorsum pale yellow to brown, covered with large black spots (Fig. 8). Spots

larger, irregular, sometimes confluent in juvenile specimens. Belly white, with or

without black spots. Maxillary and mandibular barbels white. Iris yellowish to copper

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 base, posterior margins white in color. Triangles may be completely solid or

composed 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 caudal fin with black bar from base to

tip of fin, posterior margin of fin white.

Distribution

Lake Tanganyika (Fig. 9); common.









Habitat

Littoral to benthic zones over shell, sand and mud bottoms; to a maximum depth of

170 m (Coulter, 1991).

Diet

Synodontis multipunctatus is generally considered to be a specialized predator of

Neothauma tanganyicense, a common Tanganyikan gastropod (Poll, 1953; Brichard,

1989; Coulter, 1991). The method by which the snail is extracted from its shell is

unknown. Other authors have reported the presence of crustaceans, insect larvae and

lamellibranches, in addition to Neothauma, in the stomach of S. multipunctatus

(Worthington and Ricardo, 1936; Poll, 1953, Coulter, 1991), 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 number of shells found in this particular specimen would seem to

indicate otherwise. Further study of the natural feeding habits of S. multipunctatus on

both gastropods and bivalves is necessary to more completely understand the diet of this

species.

Reproduction

Synodontis multipunctatus is the only fish species known to exhibit true brood

parasitism (Sato, 1986). Eggs are deposited and fertilized among those of at least six

species of mouth-brooding cichlids (Sato, 1986). These eggs are then taken up by the

female cichlid and brooded with the cichlid's own eggs. The S. multipunctatus hatch

before the cichlids 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 alternate spawning behaviors exist.

Taxonomic Remarks

Synodontis multipunctatus joins S. granulosus as the only Tanganyikan Synodontis

for which no other names have been either proposed or mistakenly applied in the

literature. Like most Synodontis species in the lake, slight variation in color pattern

occurs among localities, but nothing which is consistent enough to be able to confidently

distinguish between populations. Another similarly patterned, but nevertheless distinct

species has been grouped with S. multipunctatus by previous authors. This species is

described in the following account.











Table 4. Morphometric measurements and meristic counts for Synodontis multipunctatus.
All morphometric measurements given in percent of base measurement.
Meristic data for Holotype is identified by a "*" superscript. SL = Standard
length, HL = Head length, XMB = External mandibular barbel, IMB =
Internal mandibular barbel, HPL = Humeral process length, HPD = humeral
process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth.
Numbers in parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


Holotype
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


Range (n=67)
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


Meristics
Mandibular tooth count 13(2); 14(4); 15(3); 16(6); 17(4); 18(9);
19*(5); 20(6); 21(6); 22(10); 23(3);
24(5); 26(3); 29(1)
Dorsal fin count 1,7*(67)
Pectoral fin count 1,8*(67)
Pelvic fin count i,6*(67)
Anal fin count iii,6(9); iii,6,i(2); iii,7(30); iii,7,i(5);
iii,8*(10); iv,6 (4); iv,6,i(2); iv,7(2);
iv,7,i(1), iv,8(2)
Caudal fin count i,7,8,i(4)
Total vertebrae count 31(3); 32(1)


MeanSD
24.32.2
30.01.3
40.01.3
29.22.2
31.1+5.1
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
81.7+6.0
72.56.3
68.16.9
307.636.5
142.118.6






















V~-,


Figure 8. Synodontis multipunctatus, Holotype, BMNH 1898.9.9.76, 280 mm TL, 220
mm SL. Top left dorsal view, bottom left lateral view, right ventral view,
showing mouth and barbels.













































krn

Figure 9. Distribution ofSynodontis multipuncttus. T denotes type locality.
Figure 9. Distribution of Synodontis multipunctatus. T denotes type locality.









Synodontis sp. 1 n. sp.

(Figures 3, 10, 11; Tables 1, 5)

Holotype: BMNH 1982.4.13.4785, TL 129 mm, SL 96 mm, Mwakizega coastline, L.
Tanganyika, coll. R. Travers.

Paratypes: (1) BMNH 1982.4.13.4786, TL 112 mm, SL 91 mm, collection information
same as for holotype.

Diagnosis

Axillary pore present; mandibular 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 internal mandibular barbels absent; occipito-nuchal shield covered

with skin; papillae on skin of body absent; hindgut chamber absent; maximum TL 150

mm.

This species is most similar in appearance to Synodontis multipunctatus, and is

distinguished from other species of Synodontis by the same traits. It is most easily

distinguished from S. multipunctatus 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 Synodontis sp. 1 having a count of 1,7 vs. 1,8 in S. multipunctatus.

Finally, Synodontis sp. 1 reaches a much smaller maximum total length than does S.

multipunctatus (150 mm vs. 280 mm).

Description

Morphometric and meristic data are given in Table 5. Maximum total length 150

mm, standard length 110 mm. Body compressed. Predorsal profile convex; interrupted

by a ridge formed by the orbit of the eye. Preanal profile convex. Skin on body forming

numerous vertical epidermal folds; papillae absent.









Head depressed and broad; skin smooth. Snout subconical when viewed laterally;

rounded when viewed dorsally. Occipital-nuchal shield completely covered with skin;

terminating posteriorly with narrow, rounded process on either side of dorsal spine;

ventrally with wide, rounded process which extends to upper margin of humeral process

on either side of body, also covered with skin in smaller specimens. Eyes dorsolateral;

ovoid; horizontal axis longest. Interorbital area flat to slightly convex.

Mouth subterminal; lips wide and papillate. Mandibular teeth 17-26, short,

unicuspid; arranged in single transverse row. Premaxillary toothpad uninterrupted;

primary, secondary and tertiary premaxillary teeth discrete, 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. External mandibular barbel extending to point

just short of anterior margin of pectoral girdle; with 4-5 non-tuberculate branches;

lacking secondary branches. Internal mandibular barbel 1/2 to 2/3 length of external

barbel; with 4-5 pairs non-tuberculate branches; lacking secondary branches.

Dorsal fin 1,7; 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 spine margin with small serrations distally. Pectoral fin 1,7;

posterior margin straight. Pectoral fin spine roughly equal in length to dorsal fin spine,

striated, slightly curved, terminating in short, white filament; anterior spine margin with

many small, antrorse serrations; posterior spine margin with large, retrorse serrations

along entire length. Adipose fin poorly developed; 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 margin convex; terminating in a 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 52096-53100, MRAC 90273-90277). Hindgut chamber

absent.

Coloration

Dorsum pale yellow to brown, covered with large black spots (Fig. 10). Spots

larger, irregular, sometimes confluent in juvenile specimens. Belly white, with or

without black spots. Maxillary and mandibular barbels white. Iris yellowish to copper

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

made up 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 caudal fin with black bar from base to

tip of fin; posterior margin of fin white.

Distribution

Lake Tanganyika (Fig. 11); apparently common, though less so than S.

multipunctatus.









Habitat

Likely inhabits littoral to benthic zones over shell, sand and mud bottoms, as do

most of the other species of the lake. The increased relative size of the eye and decreased

body size may indicate that Synodontis sp. 1 inhabits deeper water than does S.

multipunctatus, but this cannot be stated with certainty, due to the lack of depth data in

collection records.

Diet

The lack of fresh material of Synodontis sp. 1 makes a thorough examination of this

species' diet impossible. It is likely that Synodontis sp. 1 feeds on small gastropods, as

do the other species of Synodontis from the lake.

Reproduction

It is unknown whether Synodontis sp. 1 shares the brood parasite behavior of

Synodontis multipunctatus. In fact, without examining the specimens used in the original

study of this behavior (Sato, 1986), it is impossible to say which species, S.

multipunctatus or Synodontis sp. 1 was used in this study. Unfortunately, no records

exist that might indicate the location of the specimens used. Differences in spawning

time are likely to exist, as one mixed lot (MRAC 39186-39192) of S. multipunctatus and

Synodontis sp. 1 contains gravid females of the latter species, while the female specimens

of S. multipunctatus do not have mature gonads.

Taxonomy

As stated in the previous account, no additional names have previously been

suggested for S. multipunctatus. It follows that there are no names available for

Synodontis sp. 1. A proper specific epithet will be applied to this species upon

publication of this study.











Table 5. Morphometric and meristic counts for Synodontis sp. 1. All morphometric
measurements given in percent of base measurement. Ranges for S.
multipunctatus are provided for comparison. SL = Standard length, HL =
Head length, XMB = External mandibular barbel, IMB = Internal mandibular
barbel, HPL = Humeral process length, HPD = humeral process depth, CPL =
Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in
parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


S. multipunctatus
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
21.6-30.0
29.7-42.7
29.7-41.7
24.2-42.3
68.3-159.9
39.9-78.1
18.2-39.0
33.0-67.2
73.3-111.3
66.9-105.9
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


Synodontis sp. 1 (n=24)
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


Meristics
Mandibular tooth count 17(1); 18(2); 19(2); 20(4); 21(6); 22(3);
23(1); 24(3); 25(1); 26(1);
Dorsal fin count 1,7(25)
Pectoral fin count 1,7(21); I,7,i(4)
Pelvic fin count i,6(25)
Anal fin count 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)
Caudal fin count i,7,8,i(3)
Total vertebrae count 32(3)


MeanSD
23.21.8
29.61.0
39.21.8
27.93.3
30.93.4
27.02.8
27.1+2.8
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







































Figure 10. Synodontis multipunctatus (top), BMNH 1982.4.13.4789, 115 mm TL, 91 mm
SL vs. Synodontis sp. 1 (bottom), BMNH 1982.4.13.4784, 105 mm TL, 81
mm SL. Though similarly patterned, the eye of Synodontis sp. 1 is larger than
that of S. multipunctatus, particularly in proportion to the length of the snout.






51








-. BRu R 4 -1 1 -
4" ,4












TANZANIA
DEMOCR TIC REPUBLIC
OF CON O








WFtlpa




as' 20' 3;'
km
S 11so 1 00
Figure 11. Distribution of Synodontis sp. 1. T denotes type locality.









Synodontispetricola Matthes, 1959

(Figures 3, 12, 13; Tables 1, 6)

Synodontis petricola Matthes, H., 1959:78, description, n. sp., Lake Tanganyika;
1962:34, pl. 2, fig. a, description, diet, Lake Tanganyika. Poll, M., 1971:409,
figs. 192, 193, pls. VI.18, XII.9. Brichard, P., 1978:360, 424,
photos(misidentified), key; 1989:474, 475, 476, 479, 481, 483, skin texture, key,
photos(some misidentifications). Daget, J., Gosse, J.P., and Thys van den
Audenaerde, D.F.E, 1986:140, check-list. Burgess, W.E., 1989:196, 556, 559,
560, 567, check-list, photos(some misidentifications). Kobayagowa, M.,
1989:14, photo(misidentified). Coulter, G.W., 1991:181, 182, table 8.16, diet,
habitat, reproduction. Coulter, G.W., 1991a:182, table 8.16, diet, habitat,
reproduction.

Synodontis multimaculatus, non Boulenger, Worthington, E.B. and Ricardo, C.K.,
1936:1067, 1077, 1101, note. Poll, M., 1946:223, probable absence from
Lake Tanganyika.

Diagnosis

Axillary pore present; mandibular teeth 31-50; body with large spots; fin spines

white; 8-9 pectoral fin rays; black triangles present on base of all rayed fins; eye 28.7-

40.1% snout length; premaxillary toothpad interrupted; secondary branches on internal

mandibular barbels present; occipito-nuchal shield covered with skin; papillae on skin of

body absent; hindgut chamber present; maximum TL 135 mm.

The presence of an axillary pore distinguishes Synodontispetricola from

Synodontis sp. 2, S. nigromaculatus, S. tanganaicae, S. dhonti, S. polli, and Synodontis

sp. 3. The occipito-nuchal shield being covered with skin further separates S. petricola

from S. granulosus S. nigromaculatus, S. tanganaicae, and S. dhonti, all of which also

have a much larger maximum TL (270, 380, 600, and 400 mm, respectively vs. 135 mm

in S. petricola). Synodontispetricola differs from S. granulosus, S. multipunctatus, and

Synodontis sp. 1 in having an interrupted premaxillary toothpad and secondary branches

on the internal mandibular barbels. It is further distinguished from all other Tanganyikan









Synodontis, with the exception of Synodontis sp. 2, by having completely white fin

spines. Synodontispetricola is most easily distinguished from Synodontis sp. 2 by the

presence of an axillary pore and the lack of light colored windows at the bases of the dark

triangles on its fins. Synodontispetricola also somewhat 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 are given in Table 6. Maximum total length 135

mm, standard length 115 mm. Body not compressed. Predorsal profile slightly convex.

Preanal profile straight to slightly convex. Skin on body forming numerous vertical

epidermal folds; papillae absent.

Head slightly depressed and broad; skin covered with villous papillae; papillae

extend onto bases of maxillary barbels and anterior portions of body only. Snout with

nearly flattened margin when viewed laterally; bluntly rounded when viewed dorsally.

Occipital-nuchal shield covered with skin, terminating posteriorly with wide, pointed

process on either side of dorsal spine, ventrally with wide, rounded process which

extends to upper margin of the humeral process on either side of body. Eyes dorsolateral;

ovoid; horizontal axis longest. Interorbital area flat to slightly convex.

Mouth inferior; lips widened and papillate. Mandibular teeth 31-50, short,

unicuspid; arranged in 6 short, transverse rows. Premaxillary toothpad interrupted;

primary, secondary and tertiary premaxillary teeth discrete, 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 membrane narrow. External mandibular barbel extending to point just past









anterior margin of pectoral girdle; with 4-7 short, simple, weakly tuberculate branches;

usually lacking secondary branches. Internal mandibular barbel approximately 1/3 to 2/3

length of external barbel; with 4-6 pairs of tuberculate branches; many secondary

branches present.

Dorsal fin I, 7; 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 spine margin with small serrations distally. Pectoral fin I, 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; posterior spine margin with large, retrorse serrations along entire length.

Adipose fin long; well developed; margin convex. 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; extends posteriorly into body. Gut 1.2-1.3 times

body length (n = 2, MRAC A3-033-P-0002-0011). Well developed hindgut chamber.

Coloration

Dorsum yellowish to cuprous brown, covered with large, irregularly shaped, black

spots. Spots proportionately larger, sometimes confluent in juvenile specimens. Belly

lighter, with small, irregularly shaped 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 in color.









Both lobes of caudal fin with black bars, extending from base to tip of fin; posterior

margin white.

Distribution

Lake Tanganyika (Fig. 13). The large amount of material available in museum

collections would seem to indicate that this species is more common than has been stated

in previous accounts (Matthes, 1959; Coulter, 1991).

Habitat

Rocky coasts within the littoral zone, to a maximum depth of 30 m (Matthes, 1959;

Coulter, 1991).

Diet

Young individuals appear to be primarily carnivorous, subsisting mainly on

hydracarians, ostracods and insect larvae trichopteranss, chironomids) (Matthes, 1959).

Adults feed on algae scraped from rocky substrates, and ingest small invertebrates

associated with the biological cover of the rocks in which this species lives (Matthes,

1959, Coulter, 1991).

Reproduction

No information exists regarding the spawning season, behavior, or location of this

species. All that is known is that the number of eggs released is likely to be small, as a

single female specimen has been found to contain approximately 100 eggs (Matthes,

1959).

Taxonomic Remarks

Synodontispetricola is sufficiently distinct that there has historically been little

confusion regarding its taxonomy. Worthington and Ricardo (1936), in their account of

the non-cichlid fishes of Lake Tanganyika, identified a specimen collected by Christy as









Synodontis multimaculatus (BMNH 1936.6.14.1232). Poll (1946) noted that this

identification was incorrect, though he did not give a new species identification for the

specimen. No list of materials examined was given in the original description of S.

petricola (Matthes, 1959), but the type designation given to this specimen clearly refers

to this name, as a later description by the same author lists this specimen as a paratype

(Matthes, 1962).

Synodontispetricola shows considerable variation in the nature of its spotted

pattern, with the size and spacing of the spots varying considerably between specimens,

even those coming from the same locality. The coloration of the fins varies slightly as

well, though it seems to be much more consistent than the pattern of the body.

Specimens coming from one area (Musende Rocks, Mpulungu) show such a divergence

from the fin coloration of other localities, in addition to the lack of the axillary pore

found in other populations, that I consider them to be a distinct species, which is

described in the following account.











Table 6. Morphometric measurements and meristic counts for Synodontispetricola. All
morphometric measurements given in percent of base measurement. Meristic
data for Holotype is identified by a "*" superscript. SL = Standard length,
HL = Head length, XMB = External mandibular barbel, IMB = Internal
mandibular barbel, HPL = Humeral process length, HPD = humeral process
depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth.
Numbers in parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


Holotype
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


Range (n=42)
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


MERISTICS
Mandibular tooth count 31(1); 33(1); 34(3); 35(1); 36(2);
37(2); 38(5); 39(7); 40(4); 41*(4);
42(5); 43(1); 45(3); 47(1); 50(1)
Dorsal fin count 1,6(3); 1,7*(39)
Pectoral fin count 1,8*(27); 1,9(15)
Pelvic fin count i,6*(42)
Anal fin count iii,8(2); iii,9(4); iv,7(3); iv,7,i(4);
iv,8*(20); iv,8,i(6); iv,9(3)
Caudal fin count i,7,8,i(18)
Total vertebrae count 31(2); 32(12); 33(4)


MeanSD
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
46.6+4.0
53.1+2.3
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
77.1+7.0
33.62.5
65.8+7.0
66.64.6
51.66.2
50.75.2
245.935.4
92.617.0
























































Figure 12. Synodontispetricola, Holotype, MRAC 130357, 98 mm TL, 82 mm SL. Top -
dorsal view, middle lateral view, bottom ventral view.






1Yia


REPUBLIC


Figure 13. Distribution of Synodontispetricola. T denotes type locality.


tr


rz%









Synodontis sp. 2 n. sp.

(Figures 4, 14, 15; Tables 2, 7)

Holotype: SAIAB 42518, TL 90 mm, SL 74 mm, Country: Zambia, System: Lake
Tanganyika; Musende Rocks, 0846'00"S, 031 07'00"E, coll. 10.X.1992.

Paratypes: (9) SAIAB 39577, TL 58-98 mm, SL 47-79 mm, Zambia; Mpulungu,
Musende Rocks, 0846'00"S, 031051'00"E, coll. 07.VII.1992, (3) SAIAB
40171, TL 70-85 mm, SL 56-69 mm, Country: Zambia; Musende Rocks,
08046'00"S, 031006'00"E, coll. 05.VIII.1992, (3) SAIAB 42518, TL 55-97
mm, SL 44-78 mm, collection information same as for holotype.

Diagnosis

Axillary pore absent; mandibular teeth 38-51; body with large spots; fin spines

white; 8-9 pectoral fin rays; black triangles on base of all rayed fins with light colored

window at base (except caudal fin); eye 25.2-35.8% snout length; premaxillary toothpad

interrupted; secondary branches on internal mandibular barbels 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 sp. 2 from S. granulosus, S.

multipunctatus, Synodontis sp. 1, and S. petricola. Synodontis sp. 2 is further separated

from S. petricola by the presence of light colored windows at the bases of the rayed fins

(with the exception of the caudal fin). Morphometric measurements vary little between

these two species (Table 7). Synodontis sp. 2 differs from S. nigromaculatus, S.

tanganaicae, S. dhonti, S. polli, and Synodontis sp. 3 in having white fin spines (vs.

brown to black) and in lacking papillae on the skin of the body. The occipito-nuchal

shield being covered with skin further separates Synodontis sp. 2 from S. granulosus S.

nigromaculatus, S. tanganaicae, and S. dhonti, all of which also have a much larger

maximum total length (270, 380, 600, and 400 mm, respectively vs. 100 mm in

Synodontis sp. 2). The black triangles at the base of all the rayed fins further distinguish









Synodontis sp. 2 from S. nigromaculatus, while the windows at the base of the triangles

further separate Synodontis sp. 2 from S. granulosus, S. multipunctatus, Synodontis sp. 1,

S. tanganaicae, S. dhonti, S. polli, and Synodontis sp. 3.

Description

Morphometric and meristic data are given in Table 7. Maximum total length 100

mm, standard length 80 mm. Body not compressed. Predorsal profile slightly convex.

Preanal profile slightly convex. Skin on body forming numerous vertical epidermal

folds; papillae absent.

Head slightly depressed and broad; skin covered with villous papillae; papillae

extend onto bases of maxillary barbels, pectoral fins and anterior portions of body only.

Snout with bluntly rounded margin when viewed laterally and dorsally. Occipital-nuchal

shield covered with skin, terminating posteriorly with wide, pointed process on either

side of dorsal spine, ventrally with wide, rounded process which extends to upper margin

of the humeral process on either side of body. Eyes dorsolateral; ovoid; horizontal axis

longest. Interorbital area flat to slightly convex.

Mouth inferior; lips widened and papillate. Mandibular teeth 35-51, short,

unicuspid; arranged in 6 short, transverse rows. Premaxillary toothpad interrupted;

primary, secondary and tertiary premaxillary teeth discrete, 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 membrane narrow. External mandibular barbel extending to point just past

anterior margin of pectoral girdle; with 4-6 short, simple, weakly tuberculate branches;

usually lacking secondary branches. Internal mandibular barbel approximately 1/3 to 1/2









length of external barbel; with 4-6 pairs of tuberculate branches; many secondary

branches present.

Dorsal fin I, 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 spine margin with small serrations distally. Pectoral fin I, 8-

8,i; 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; posterior spine margin with large, retrorse serrations along entire

length. Adipose fin long; well developed; margin convex. 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 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

Dorsum yellowish to cuprous brown, covered with large, irregularly shaped, black

spots (Fig. 14). Belly lighter, with smaller, more regularly shaped 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; posterior margin of fin dusky.









Distribution

Lake Tanganyika (Fig. 15). 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 sp. 2.

Taxonomic Remarks

Specimens of this species have previously been identified either as Synodontis sp.

or S. petricola. While this species resembles S. petricola in most respects, it is clearly

diagnosable from S. petricola. As there are no names available to be applied to this

species, an appropriate specific epithet will be applied upon publication of this study.











Table 7. Morphometric and meristic counts for Synodontis sp. 2. All morphometric
measurements given in percent of base measurement. Ranges for S. petricola
are provided for comparison. SL = Standard length, HL = Head length, XMB
= External mandibular barbel, IMB = Internal mandibular barbel, HPL =
Humeral process length, HPD = humeral process depth, CPL = Caudal
peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses
indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


S. petricola
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
30.9-67.1
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


Synodontis sp. 2 (n=16)
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


Meristics
Mandibular tooth count 35(2); 36(2); 38(4); 39(1); 41(2); 43(1);
44(1); 47(2); 51(1)
Dorsal fin count 1,7(15); I,7,i(1)
Pectoral fin count 1,8(14); I,8,i(2)
Pelvic fin count i,6(16)
Anal fin count iv,8(11); iv,8,i(2); iv, 9(3)
Caudal fin count i,7,8,i(10)
Total vertebrae count 32(3); 33(6)


MeanSD
20.02.7
29.81.0
38.1+1.4
32.62.9
23.01.9
19.61.4
21.3+1.8
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

























-
V*4


14f
S 41i


Figure 14. Synodontis sp. 2, SAIAB 39577, 84 mm TL, 68 mm SL. Top dorsal view,
middle lateral view, bottom ventral view.


L it





66






-4"' B 1 -4,








TANZANIA
DEMOCR TIC REPUBLIC
OF CONGO




L R ulana




w' 30s'
km

Figure 15. Distribution of Synodontis sp. 2. T denotes type locality.









Synodontis nigromaculatus Boulenger, 1905

(Figures 4, 16, 17, 18; Tables 2, 8)

Synodontis nigromaculatus Boulenger, G.A., 1905:645, description, n. sp., Lake
Bangwelu; 1911:416, fig. 313, description, Lake Bangwelu. -
Worthington, E.B., 1933:50, note, Lake Bangwelu. Ricardo-Bertram, C.K.,
1940:187, 209, note, diet, Chambezi R., Bangwelu marsh, Lake Bangwelu, Congo
system, Zambezi system, comparison with S. melanostictus, considered
synonymous. Bowmaker, A.P., 1961:6, fecundity. Jackson, P.B.N., 1961:20,
87, key, note, N. Rhodesia. Jubb, R.A., 1961:43, 117, fig. 61, description,
fishing, Limpopo system, Upper Zambezi; 1963:7, 12, 15, 28, note, Upper
Zambezi (S. Afr.), = S. melanostictus; 1967,:43, 52, 56, fig. 155, description, key,
distribution table, Upper Zambezi, Okovango system, Cunene R. (S. Afr.), = S.
melanostictus. De Kimpe, P., 1964:77, Luapula-Mweru. Ladiges, W.,
1964:247, 254, 256, 257, 263, 267, diet, ford near Mulundo, Cuchi R., Cuebe R.,
Cubango R., Okovango R.- Gabie, V., 1965:386, maps 1 and 2, distribution in S.
Africa (Limpopo). Bell-Cross, G., 1955-1966a and b:13, 16, 18, 27, Upper
Zambezi, Okovango, affinities with the fauna of the Congo system (Lualaba-
Congo). Poll, M., 1967:247, fig. 115, description, Lake Calundo (Angola); Poll,
M., 1971:325, fig. 148 and 149, pl. V,11 and XI,9, description, synonymy with S.
melanostictus. Sands, D., 1983:23, check-list. Daget, J., Gosse, J.P., and Thys
van den Audenaerde, D.F.E, 1986:136, check-list. Brichard, P., 1989:479, key,
478, 480, 483, photos. -Burgess, W., 1989:196, check-list, pl. 78, 82, 83, 94. -
Kobayagawa, M., 1989:15, photo. Coulter, G.W., 1991:152, 154, 156, 181, 191,
287, abundance, habitat, reproduction. Coulter, G.W., 1991a:152, 154, 156, 182,
191, abundance, habitat, reproduction.

Synodontis melanostictus Boulenger, G.A., 1906:553, pl. 34, description, n. sp., Lofu
(Lake Tangamnyika); 1911:418, fig. 314, description, Lofu (Lake Tanganyika),
Lake Bangwelu, Lake Mweru, Maramba, Upper Zambezi; 1916:316, note.
-Pappenheim, P. and Boulenger, G.A., 1914:251, note.-Poll, M., 1933:134,
Pweto, Lukonzolwa, Lake Mweru, Katanga; 1946:220, description, Lake
Tanganyika (citation); 1948:8, Pweto, Lake Mweru (Katanga). Worthington,
E.B., 1933:50, note, Lake Bangwelu. Fowler, H.W., 1935:273, description,
Chobe R 3 miles from Kasane (Kalahari). Worthington, E.B. and Ricardo, C.K.,
1933:1067, 1077, 1101, note, Lake Tanganyika, Congo system, Zambezi system,
including Lake Malawi. Pellegrin, J., 1936:58, Humbe, Cunene, Cubango
(Angola). Ricardo-Bertram, C.K., 1940:209, first placed into synonymy of S.
nigromaculatus. Barnard, K.H., 1948:443, note, Lealui, Lake Ngami,
Victoria Falls, Seheki, Okovango R., Cunene R. Hulot, A., 1950:170, Lake
Mweru. Jubb, R.A., 1954:692, 697, S. Rhodesia; 1958:181, table 1,
Chobe R., Kobuta, Zambezi, R., Sananga. Soulsby, J.J., 1960:323, note, diet, N.
Rhodesia. Matthes, H., 1962:41, table 2, comparison of specimens from Lake
Tanganyika and Mweru-Luapula, comparison with S. serratus tanganaicae
and S. lacustricolus. Sterba, G., 1963:408, fig. 610, Zambezi, Lake
Tanganyika, Lake Bangwelu, Lake Mweru. Ladiges, W., 1964:267, Cunene,









Cubango, Okovango, Zambezi. De Kimpe, P., 1964:77, 89, 114, 116, 137, 138,
147, 185, 188, 195, 201, 222, figs 46, 72, diet, Luapula-Mweru. Coulter, G.W.,
1965-1966:34, abundance at different depths, Lake Tanganyika. Brichard,
1978:424, key.

Synodontis colyeri Boulenger, G.A., 1923:438, description, n. sp., Mansa R. (N.
Rhodesia). Jackson, P.B.N., 1961:20, 88, note, key, N. Rhodesia. De Kimpe,
P., 1964:77, 89, Mansa R. (Luapula- Moero). Bell-Cross, G., 1965-1966a:13,
18, Zambia, Congo. Poll, 1971:329, placed into synonymy with S.
nigromaculatus.

Synodontis zambesensis, non Peters, David, L. and Poll, M., 1937:234. Poll, M.,
1967:243, fig. 113, Angola, tributary of the Kasai.

Diagnosis

Axillary pore absent; mandibular teeth 23-38; body with small spots; fin spines

brown; 8-10 pectoral fin rays; black triangles absent on all rayed fins; eye 31.2-48.1% of

snout length; premaxillary toothpad interrupted; secondary branches on internal

mandibular barbels absent; occipito-nuchal shield not covered by skin; villous papillae

present on skin of body; hindgut chamber present; maximum TL 380 mm;

The only non-endemic Synodontis species found in Lake Tanganyika, S.

nigromaculatus lacks the rayed fin coloration common to all of its Tanganyikan

congeners, having spotted fins which completely lack black triangles at their bases. The

black triangles on the fins ofS. tanganaicae may become indistinct in large specimens

and the body may have small black spots that resemble those found in S. nigromaculatus.

In these cases, mandibular tooth counts (23-38 in S. nigromaculatus vs. 33-49 in S.

tanganaicae), differences in eye size (31.2-48.1% of snout length in S. nigromaculatus

vs. 16.0-26.9% in S. tanganaicae), the villous papillae on the skin of S. nigromaculatus

(vs. granular in S. tanganaicae), and differences in humeral process shape (Fig. 4)

distinguish these species from one another.









Description

Morphometric and meristic data are given in Table 8. Maximum total length 380

mm, maximum standard length 305 mm. Body compressed. Predorsal profile straight

anterior to eye; slightly convex posterior to eye. Preanal profile straight. Skin on body

lacking vertical epidermal 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.

Occipital-nuchal shield rugose, not covered with skin; terminating posteriorly with

narrow, rounded process on either side of dorsal spine; ventrally with wide, rounded

process which extends to upper margin of humeral process on either side of body. Eyes

dorsolateral, ovoid, horizontal axis longest. Interorbital area flat to slightly convex.

Mouth subterminal; lips wide and papillate. Mandibular teeth 23-38, short,

unicuspid; arranged in single transverse row. Premaxillary toothpad interrupted; primary,

secondary, and tertiary premaxillary teeth discrete; primary premaxillary teeth in 2-3

rows; secondary premaxillary 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. External mandibular barbel extending to

point just short of anterior margin of pectoral girdle; with 7-8 long, non-tuberculate

branches; short secondary branches present. Internal mandibular barbels about half the

length of the external barbels; with 4-5 pairs of non-tuberculate branches; secondary

branches present.

Dorsal fin 1,7; posterior margin concave. Dorsal fin spine long, striated, slightly

curved, terminating in short black filament; anterior margin of fin spine smooth; posterior

spine margin with small serrations distally. Pectoral fin 1,8-10; posterior margin straight.









Pectoral fin spine roughly equal in length to dorsal fin spine, striated, slightly curved,

terminating in a moderate length filament; anterior spine margin with many fine, long,

antrorse serrations along anterior margin; posterior spine margin with large, retrorse

serrations along entire length. Adipose fin 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; tapering to sharp point; striated along

lower half; becoming granulous along upper margin; possessing weakly developed ridge

along ventral margin; dorsal margin slightly concave (Fig 5B). Axillary pore absent. Gut

1.3-1.6 times body length (n=2, MRAC 100515-100518). Well developed hindgut

chamber.

Coloration

Body grayish to cuprous brown (Fig. 16). Belly pale. Scattered, small black spots

present, sometimes confluent. Maxillary and mandibular barbels white. Iris copper

colored. Dorsal and pectoral fin spines brown, terminating in dusky filaments. All rayed

fins with small, black spots, similar to those found on body.

Distribution

Lake Tanganyika, Lake Mweru, and Lake Bangwelu basins; Upper Kasai, Upper

Zambezi, Okovango, Cunene, and Limpopo Rivers; Luapula swamps (Fig. 18).

Habitat

Littoral to benthic zones over shell, sand and mud bottoms, to a maximum depth of

150 m (Coulter, 1991).









Diet

Nothing is known about the diet of this species within Lake Tanganyika.

Reproduction

No concrete information exists on the reproduction of S. nigromaculatus. Coulter

(1991) cites Matthes (1959) as reporting that gravid females of this species were found to

contain up to 3,000 eggs. However, upon reviewing this second reference, no mention of

the reproduction of S. nigromaculatus is found. Coulter (1991) states that it is likely that

S. nigromaculatus spawns in rivers within the lake basin, as this species is known to

spawn in rivers throughout its range. Coulter's assessment appears to be accurate, as

only large adult specimens of S. nigromaculatus have thus far been documented within

the lake.

Taxonomic Remarks

Synodontis melanostictus Boulenger, originally described from Lake Tanganyika,

was distinguished from S. nigromaculatus Boulenger by the presence of numerous,

villous papillae on the body (Boulenger, 1906). Ricardo-Bertram (1940) first suggested

placing S. melanostictus in the synonymy of S. nigromaculatus, stating that the presence

or absence of these papillae depended upon the condition and preservation of a particular

specimen. This is undoubtedly true to some extent, as the holotype of S. melanostictus

lacks any indication of papillae on its body. However, the holotype of S. nigromaculatus

is in excellent condition, yet it still lacks any papillae of the sort found on S.

melanostictus, as do any specimens agreeing in appearance with the holotype of S.

nigromaculatus that come from outside the Lake Tanganyika drainage.

Further distinguishing Synodontis melanostictus from S. nigromaculatus is the

shape of the snout when viewed dorsally, as well as the condition of the occipito-nuchal









shield, which is exposed and rugose in S. melanostictus, but covered with skin in S.

nigromacualtus (Fig. 17). Though the occipito-nuchal shield does become more exposed

with growth in many Synodontis species, these comparisons have been made between

specimens of similar size and appear to be consistent. While it appears that S.

melanostictus is sufficiently distinct from S. nigromaculatus to be considered a valid

species, some material at the Royal Museum for Central Africa has yet to be examined.

This material must be studied before any taxonomic changes can be proposed.

Synodontis colyeri Boulenger was described from the Mansa River in what is now

known as Zimbabwe (Boulenger, 1926). Poll (1971) considered S. colyeri to be a

synonym of S. nigromaculatus, a decision which is accepted here, due to the fact that

examination of the holotype of S. colyeri has not yet been possible, and no photographs

or drawings exist which might provide clues as to the correct placement of this specimen.

Synodontis zambesensis Peters is a widespread species known to occur in the

Pongolo, Limpopo, Pungwe, Zambezi, Lake Rukwa, and Lake Mweru drainages (Poll,

1971). Several specimens coming from the Luachimo and nearby rivers (Congo

drainage) in Angola have previously been determined to belong to this species (David

and Poll, 1937; Poll, 1967) (Fig. 16). However, adult specimens of S. zambesensis lack

spots, having a uniform, light brown coloration, while the specimens from the Luachimo

clearly have large, widely-spaced, black spots. Poll (1971) obviously realized this

mistake, as his revision of Synodontis lists these specimens as S. nigromaculatus.

Though these specimens are not S. zambesensis, they also clearly are not specimens of S.

nigromaculatus nor are they S. melanostictus. These specimens have large, widely

spaced spots, a more depressed head than either S. nigromaculatus or S. melanostictus, a






73


narrow preopercle (vs. broad), and lack any cutaneous papillae. These specimens

represent an undescribed species, which will be described in a future paper that will deal

exclusively with the taxonomy of the S. nigromaculatus species complex.











Table 8. Morphometric and meristic counts for Synodontis nigromaculatus. All
morphometric measurements given in percent of base measurement. Meristic
data for Holotype is identified by a "*" superscript. SL = Standard length,
HL = Head length, XMB = External mandibular barbel, IMB = Internal
mandibular barbel, HPL = Humeral process length, HPD = humeral process
depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth.
Numbers in parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
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


Holotype
24.9
25.9
37.8
32.9
33.8
30.3
25.0
90.6
86.8
62.7
46.5
17.2
42.1
43.5
37.9
130.5
64.5
39.0
60.4
117.0
96.3
37.1
90.5
93.6
41.0
39.6
219.1
91.7


Range (n=34)
16.7-26.2
24.9-31.4
34.3-41.1
16.1-41.8
22.2-48.0
20.5-36.0
21.5-30.9
81.7-99.2
63.4-91.7
51.5-76.1
44.5-53.7
15.4-22.0
32.5-54.2
33.9-44.0
24.9-44.0
77.4-172.7
51.2-107.6
26.6-61.3
36.3-66.8
84.1-127.2
80.0-110.8
31.1-48.1
66.8-96.8
64.8-93.6
32.9-65.2
37.7-63.2
170.9-318.2
66.9-121.8


Meristics
Mandibular tooth count 23(1); 25(1); 26(3); 27(5); 28(3);
29*(5); 30(1); 31(3); 32(6); 33(1);
34(1); 35(2); 36(3); 37(1); 38(2)
Dorsal fin count 1,7*(39)
Pectoral fin count I,8*(15); I,8,i(1); 1,9(20), ,9,i(1);
1,10(1)
Pelvic fin count i,6*(39)
Anal fin count iii,8(2); iii,8,i(2); iii,9(4); iv,7(1);
iv,7,i(4); iv,8 (8); iv,8,i(8); iv,9*(5);
v,7(1), v,8(2), v,8,i(1), v,9(1)
Caudal fin count i,7,8,i(1)
Total vertebrae count 36(1)


MeanSD
22.02.1
27.51.4
38.1+1.6
31.2+3.6
34.25.9
26.92.8
26.1+2.0
92.43.6
73.76.6
64.25.3
49.02.2
18.21.7
38.2+4.1
37.82.5
34.64.4
124.923.0
81.5+16.4
41.98.8
51.1+5.6
98.89.3
95.67.2
37.34.4
77.36.6
77.47.2
48.48.2
48.46.4
245.935.4
94.812.2






















































Figure 16. Variation in body shape and coloration in Synodontis nigromaculatus, Top
S. melanostictus, Holotype, BMNH 1906.9.8.72, 290 mm TL, 237 mm SL.
Middle S. nigromaculatus, Holotype, BMNH 1905.11.10.10, 237 mm TL,
186 mm SL. Bottom Synodontis sp. (Orig. I.D. = S. zambesensis), MRAC
162203-204, 180 mm TL, 132 mm SL.






76





























occipito-nuchal shield texture.
occipito-nuchal shield texture.










































15" 20"


30"
km
n qnn Ann


Figure 18. Distribution of Synodontis nigromaculatus. T denotes type locality.









Synodontis tanganaicae Borodin, 1936

(Figures 4, 19, 20, 21; Tables 2, 9)

Synodontis serratus tanganaicae Borodin, N.A., 1936:9, description, ssp. of S. serratus
Riippel, Kasenga, Lake Tanganyika.

Synodontis lacustricolus Poll, M., 1953:157, fig. 18a, pl. 6, fig. 3, description, n. sp., diet,
Lake Tanganyika; 1971:402, fig. 188, 189, pl. 6, 12. Matthes, H., 1962:41, pl. 3,
fig. a, description. Coulter, G.W., 1965-1966:34, abundance at different depths,
Lake Tanganyika; 1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance,
habitat, diet. Brichard, P., 1978:424, key; 1989:479, key. Sands, D., 1983, 23,
check-list. Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E,
1986:131, check-list. Burgess, W.E., 1989:196, check-list. De Vos, L. and
Thys van den Audenaerde, D., 1998:147, synonymy with S. tanganaicae. -
Coulter, G.W., 1991a:152, 154, 182, 191, abundance, habitat, reproduction.

Synodontis tanganyicae De Vos, L. and Thys van den Audenaerde, D., 1998:147.

Diagnosis

Axillary pore absent; mandibular teeth 33-49; body with small or no spots; fin

spines dark; 8-9 pectoral fin rays; black triangles on base of all rayed fins in juveniles,

absent in adults; eye 16.0-26.9% snout length; premaxillary toothpad interrupted;

secondary branches on internal mandibular barbels present; occipito-nuchal shield not

covered with skin; granular papillae present on skin of body; maximum TL 585 mm.

The lack of an axillary pore distinguishes Synodontis tanganaicae from S.

granulosus, S. multipunctatus, Synodontis sp. 1, and S. petricola. Synodontis

tanganaicae differs further from S. granulosus in humeral process shape humerall

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. multipunctatus, Synodontis

sp. 1, S. petricola, Synodontis sp. 2, S. polli, and Synodontis sp. 3. Adult specimens of S.









tanganaicae are further distinguished from the other Tanganyikan endemics (with the

exception of adult S. dhonti) by the lack of black triangles on the rayed fins.

Synodontis tanganaicae most closely resembles S. nigromaculatus and adult

specimens of S. dhonti. Synodontis tanganaicae differs from S. nigromaculatus in the

number of mandibular teeth (33-49 in S. tanganaicae vs. 23-38 in S. nigromaculatus),

eye size (16.0-26.9% of snout length in S. tanganaicae vs. 31.2-48.1% in S.

nigromaculatus), the granular papillae on the skin (vs. villous in S. nigromaculatus), and

differences in humeral process shape (Fig. 4). The only potentially reliable character that

differentiates S. tanganaicae and adult specimens of S. dhonti is the gill raker count on

the first branchial arch (27-28 in S. tanganaicae vs. 11-20 in S. dhonti) (De Vos and Thys

van den Audenaerde, 1998).

Description

Morphometric and meristic data are given in Table 9. Maximum total length 585

mm, standard length 490 mm. Body not compressed. Predorsal profile straight. Preanal

profile convex. Skin on body forming numerous vertical epidermal 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. Occipital-

nuchal shield rugose, not covered with skin; terminating posteriorly with narrow, bluntly

pointed process on either side of dorsal spine; ventrally with wide, rounded process which

extends to upper margin of humeral process on either side of body. Eyes dorsolateral;

ovoid; horizontal axis longest. Interorbital area flat.









Mouth inferior; lips wide and papillate. Mandibular teeth 33-49, short, unicuspid;

arranged in single transverse row. Premaxillary toothpad interrupted; primary, secondary

and tertiary premaxillary teeth discrete; numerous; arranged in irregular rows.

Maxillary barbel with well developed basal membrane; lacking branches or

crenelations; extending at least to base of pectoral fin. External mandibular barbel

extending to point just short of anterior margin of pectoral girdle, with 4-7 non-

tuberculate branches; lacking secondary branches. Internal mandibular barbel about 1/3

length of external barbel; with 4-6 pairs of non-tuberculate branches; secondary branches

present.

Dorsal fin 1,7-8; posterior margin straight. Dorsal fin spine long, granulous, nearly

straight, terminating in short, dark filament; anterior margin of fin spine granulous;

posterior spine margin with small serrations distally. 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 spine margin with large retrorse serrations along entire length. Adipose fin well

developed; margin convex. Pelvic 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 1,7,8,1;

forked; lobes slightly rounded.

Humeral process triangular, granulous; ridge on ventral margin absent; dorsal

margin concave; terminating in blunt point (Figs. 4C, 4D). Axillary pore absent.

Coloration

Dorsum gray to reddish brown (Figs. 19, 20). Belly lighter in color. Scattered,

small black spots present on entire body. Maxillary and mandibular barbels white, bases









sometimes with scattered 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 base of rayed

fins.

Distribution

Lake Tanganyika (Fig. 21); fairly common.

Habitat

Littoral to benthic zones over shell, sand and mud bottoms, to a maximum depth of

130 m (Coulter, 1991).

Diet

Gastropods, lamellibranchs, insect larvae, ostracods, and shrimps (Poll, 1953;

Coulter, 1991).

Reproduction

No information exists regarding the reproduction of S. tanganaicae.

Taxonomic Remarks

Synodontis serratus tanganaicae Borodin has recently been elevated to species

status, having previously been considered a subspecies of Synodontis serratus Riippell

(De Vos and Thys van den Audenaerde, 1998). De Vos and Thys van den Audenaerde

also placed Synodontis lacustricolus Poll in the synonymy of S. tanganaicae. This

decision is accepted here, however, neither of the type specimens of S. tanganaicae have

been examined.

De Vos and Thys van den Audenaerde note a strong similarity between S.

tanganaicae and S. dhonti Boulenger. This similarity is apparent when examining

specimens of S. dhonti and photographs of the Lectotype of S. tanganaicae (MCZ









32538). Differences in mandibular tooth number, adipose fin shape, and gill raker count

are cited as the characters in which S. dhonti differs from S. tanganaicae (De Vos and

Thys van den Audenaerde, 1998). However, there is significant overlap in the

mandibular tooth count of both species (20-38 in S. dhonti and 33-40 in S. tanganaicae,

33-49 when nominal specimens of S. lacustricolus are included). While the degree of

overlap between the first and last value is not large, S. dhonti is clearly distinct from

nominal specimens of S. lacustricolus. The significance of overlap is much greater

between S. dhonti and specimens identified as S. tanganaicae. Additionally, minor

variation in adipose fin morphology is common in the Synodontis species of Lake

Tanganyika. Gill rakers have not been counted for the specimens of either species that

have been examined in this study due to the necessarily destructive nature of this

procedure, therefore, the utility of this character can be neither confirmed nor refuted by

this study. Further study is clearly needed to determine whether S. tanganaicae and S.

dhonti truly represent different species, as well as to evaluate the validity of De Vos and

Thys van den Audenaerde's conclusion regarding the synonymy of S. tanganaicae and S.

lacustricolus.











Table 9. Morphometric and meristic counts for Synodontis tanganaicae. All
morphometric measurements given in percent of base measurement. Meristic
data for Lectotype is identified by a "*" superscript. All morphometric and
meristic values for the Lectotype and Paralectotype are reproduced from De
Vos & Thys van den Audenaerde, 1998. SL = Standard length, HL = Head
length, XMB = External mandibular barbel, IMB = Internal mandibular
barbel, HPL = Humeral process length, HPD = humeral process depth, CPL =
Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in
parentheses indicate number of specimens.


Measurement
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
XMB/HL
IMB/HL
IMB/XMB
Dorsal spine length/HL
Pectoral spine length/HL
Eye/Snout length
Interorbital width/Snout leng
Postorbital length/Snout leng
Eye/Interorbital width
Eye/Postorbital length
HPL/HPD
CPL/CPD


Lecto- & Paralectotype
28.4, 32.2
35.7, 34.7
49.5, 48.8
28.1, 32.2

22.6, 22.0
23.8, 25.7
85.9, 85.6
67.4, 69.2

58.2, 51.6
13.1, 13.4
39.4, 39.2
39.2, 38.5
52.0, 50.4
112.9, 127.7
47.3, 60.4
17.5, 22.7




5th
5th


Range (n=14)
20.8-32.2
31.5-35.7
41.2-49.5
23.0-32.2
29.8-43.7
21.1-27.6
19.0-26.7
79.4-94.1
66.1-84.1
38.1-55.2
51.1-60.6
9.7-14.6
35.0-42.1
35.1-40.4
30.1-52.0
90.7-138.5
47.3-66.4
14.7-23.5
29.2-43.4
63.6-87.4
57.9-83.2
16.0-26.9
60.3-81.0
59.8-78.5
26.6-38.0
26.8-39.6
172.9-255.4
98.6-149.8


Meristics
Mandibular tooth count 33(1); 38(2); 39(1); 40*(1); 42(1);
43(2); 44(1); 46(1); 47(2); 48(1); 49(1)
Dorsal fin count I,7*(12); I,7,i(1); 1,8(1)
Pectoral fin count 1,8*(5); 1,9(9)
Pelvic fin count i,6*(12); 1,7(1)
Anal fin count iii,8*(2); iv,7(1); iv,8(6); iv,8,i(1);
iv,9(1); v,7(1); v,8,i(1)


MeanSD
25.42.9
33.01.3
43.72.7
26.22.8
37.2+5.0
23.82.4
23.02.3
87.83.8
72.94.8
47.1+5.4
55.1+2.6
13.01.4
39.1+2.2
38.1+1.7
43.85.4
116.1+15.7
53.75.4
20.02.8
37.44.7
74.58.3
69.57.1
23.5+3.2
71.06.4
69.05.4
33.1+3.3
34.03.8
212.224.4
129.614.4
































Figure 19. Synodontis tanganaicae, Lectotype, MCZ 32538, 415 mm TL, 320 mm SL.
Top left dorsal view, bottom left lateral view, right ventral view of head.
Photographs courtesy of Museum of Comparative Zoology, Harvard
University.


































Figure 20. Synodontis tanganaicae, MRAC 90288, paratype of S. lacustricolus, 519 mm
TL, 434 mm SL. Top left dorsal view, bottom left lateral view, right -
ventral view of head.































































km


Figure 21. Distribution of Synodontis tanganaicae. Type locality unknown.


U_ U_ U_









Synodontis dhonti Boulenger, 1917

(Figures 5, 22, 23, 24; Tables 2, 10)

Synodontis dhonti Boulenger, G.A., 1917:367, description, n. sp., Kilewa Bay, Lake
Tanganyika; 1920:42, description, Kilewa Bay, Lake Tanganyika. -Worthington,
E.B. and Ricardo, C.K., 1936:1067, 1077, 1101, note, Lake Tanganyika. Poll,
M., 1946:221, figs. 21, 22, description, Kilewa Bay, Lake Tanganyika; 1971:364,
figs. 168 and 169, pls. VI.6 and XI.19. Matthes, H., 1962:37, fig. a, pl. 3,
description, diet, Lake Tanganyika. Brichard, P., 1978:423, key; 1989:479, key.
Sands, D., 1983:23, checklist. Daget, J., Gosse, J.P., and Thys van den
Audenaerde, D.F.E, 1986:125, check-list. Burgess, 1989:195, checklist. -
Kobayagawa, M., 1989:14, photo(misidentified). Coulter, G.W., 1991:182, 191,
table 8.16, habitat, diet, reproduction.

Synodontis irsacae Matthes, H., 1959:78, description, n. sp., Lake Tanganyika; 1962:37,
placed into synonymy of S. dhonti.

Diagnosis

Axillary pore absent; mandibular teeth 15-38; body with large spots; fin spines

dark; 8-9 pectoral fin rays; black triangles at base of all rayed fins; eye 23.1-43.7% snout

length; premaxillary toothpad interrupted; secondary branches on internal mandibular

barbels present; occipito-nuchal shield covered or not covered with skin; granular

papillae present on skin of body; hindgut chamber absent; maximum TL 400 mm.

The lack of an axillary pore distinguishes Synodontis dhonti from S. granulosus, S.

multipunctatus, Synodontis sp. 1, and S. petricola. A smaller eye (23.1-43.7% of snout

length vs. 44.9-62.0% and 64.2-81.0%, respectively) further separates juvenile S. dhonti

from S. multipunctatus and Synodontis sp. 1. Synodontis dhonti differs from S.

multipunctatus, Synodontis sp. 1, S. petricola, Synodontis sp. 2, S. polli, and Synodontis

sp. 3 in having an occipito-nuchal process which is not covered by skin. Synodontis

dhonti can be further distinguished from S. petricola, Synodontis sp. 2, S. polli, and

Synodontis sp. 3 by having a lower number of mandibular teeth (15-38 in S. dhonti vs.

31-50 in S. petricola, 38-51 in Synodontis sp. 2, 40-70 in S. polli, and 50-64 in Synodontis









sp. 3). Synodontis dhonti is further separated from S. petricola and Synodontis sp. 2 in

having dark fin spines and papillae on the body skin, (vs. white fin spines and bare skin

in S. petricola and Synodontis sp. 2). Black triangles on the rayed fins of S. dhonti

separate this species from S. nigromaculatus, where they are absent, while the presence

of granular papillae on the skin of S. dhonti (vs. villous papillae in S. nigromaculatus)

further distinguish these species. Synodontis dhonti strongly resembles specimens of S.

tanganaicae. Previous descriptions state that these species can be separated by the gill

raker count on the first branchial arch (11-20 in S. dhonti vs. 27-28 in S. tanganaicae)

(De Vos and Thys van den Audenaerde, 1998).

Description

Morphometric and meristic data are given in Table 9. Maximum total length 400

mm, standard length 325 mm. Body not compressed. Predorsal profile slightly convex.

Preanal profile straight to convex. Skin on body forming numerous vertical epidermal

folds; covered with minute, granular papillae extending onto all fins.

Head slightly depressed; broad; skin covered with granular papillae. Snout long,

with bluntly rounded margin when viewed laterally and dorsally. Occipital-nuchal shield

rugose in holotype; covered with skin in young specimens; terminating posteriorly with

narrow, bluntly pointed process on either side of dorsal spine; ventrally with wide,

rounded process which extends to upper margin of humeral process on either side of

body. Eyes dorsolateral; ovoid; horizontal axis longest. Interorbital area flat.

Mouth inferior; lips wide and papillate. Mandibular teeth 15-38, short, unicuspid;

arranged in single transverse row. Premaxillary toothpad interrupted; primary, secondary

and tertiary premaxillary teeth discrete; numerous; arranged in 3, 1-2, and 1 irregular

rows, respectively.




Full Text

PAGE 1

TAXONOMIC REVISION OF LAKE TANGANYIKAN Synodontis (TELEOSTEI: MOCHOKIDAE) By JEREMY JOHN WRIGHT A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

PAGE 2

Copyright 2006 by Jeremy John Wright

PAGE 3

iii ACKNOWLEDGMENTS Funding for this study was provided by the All Catfish Species Inventory project funded by the U.S. National Science Founda tion (DEB-0315963). I w ould like to thank Dr. Lawrence Page for directing me to the possibilities for study in the Mochokidae and for instruction, constructive criticism, and guida nce. I also thank Drs. Carl Ferraris and Collete St. Mary for their contributions as members of my supervisory committee. Additional thanks are due Dr. David Evans, w ho graciously agreed to serve as a stand-in for Dr. Ferraris at the defense of this thes is, and Dr. Jonathan Armbruster of Auburn University, who provided addi tional comments for the improvement of this thesis. I am grateful to Jacqueline Wright for her assistance during data collection. For help with formatting maps I thank Griffin Sheehy. Loans and access to institutional specimens were provided by Dr. Jos Snoeks, Ma rk Hanssens, and Miguel Parrent of the Royal Museum for Central Africa (MRAC); James Maclaine, Patrick Chapman, and Dr. Julia Day of the Natural History Museum (BMN H); Drs. Paul Skelton and Roger Bills of the South African Institute of Aquatic Biodiversity (SAIAB); and Dr. John Friel of the Cornell University Museum of Vertebrates (CU). I additionally thank Dr. Julia Day (BMNH) for her generosity in discussing her as-yet unpublished findings regarding the phylogeny of Tanganyikan Synodontis with me. My study benefited from 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.

PAGE 4

iv Permission to use the photographs seen in Fi gure 19 was arranged by K.E. Hartel of the Museum of Comparative Z oology, Harvard University.

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v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iii LIST OF TABLES...........................................................................................................viii LIST OF FIGURES...........................................................................................................ix ABSTRACT....................................................................................................................... xi CHAPTER 1 INTRODUCTION........................................................................................................1 Natural History.............................................................................................................1 Systematic History........................................................................................................2 2 MATERIALS AND METHODS.................................................................................6 Data Collection.............................................................................................................6 Characters Examined....................................................................................................7 3 MATERIALS EXAMINED.......................................................................................16 Synodontis granulosus ................................................................................................16 Synodontis multipunctatus ..........................................................................................17 Synodontis sp. 1..........................................................................................................19 Synodontis petricola ...................................................................................................20 Synodontis sp. 2..........................................................................................................21 Synodontis nigromaculatus .........................................................................................21 Synodontis tanganaicae ..............................................................................................23 Synodontis dhonti ........................................................................................................24 Synodontis polli ..........................................................................................................25 Synodontis sp. 3..........................................................................................................27 4 SPECIES DESCRIPTIONS........................................................................................28 Invariant Traits............................................................................................................28 Synodontis granulosus Boulenger, 1900...................................................................28 Diagnosis............................................................................................................29 Description.........................................................................................................30 Coloration...........................................................................................................31 Distribution.........................................................................................................31 Habitat................................................................................................................32

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vi Diet.....................................................................................................................32 Reproduction......................................................................................................32 Taxonomic Remarks...........................................................................................32 Synodontis multipunctatus Boulenger, 1898.............................................................36 Diagnosis............................................................................................................36 Description.........................................................................................................37 Coloration...........................................................................................................39 Distribution.........................................................................................................39 Habitat................................................................................................................40 Diet.....................................................................................................................40 Reproduction......................................................................................................40 Taxonomic Remarks...........................................................................................41 Synodontis sp. 1 n. sp..................................................................................................45 Diagnosis............................................................................................................45 Description.........................................................................................................45 Coloration...........................................................................................................47 Distribution.........................................................................................................47 Habitat................................................................................................................48 Diet.....................................................................................................................48 Reproduction......................................................................................................48 Taxonomy...........................................................................................................48 Synodontis petricola Matthes, 1959..........................................................................52 Diagnosis............................................................................................................52 Description.........................................................................................................53 Coloration...........................................................................................................54 Distribution.........................................................................................................55 Habitat................................................................................................................55 Diet.....................................................................................................................55 Reproduction......................................................................................................55 Taxonomic Remarks...........................................................................................55 Synodontis sp. 2 n. sp.................................................................................................60 Diagnosis............................................................................................................60 Description.........................................................................................................61 Coloration...........................................................................................................62 Distribution.........................................................................................................63 Habitat................................................................................................................63 Diet.....................................................................................................................63 Reproduction......................................................................................................63 Taxonomic Remarks...........................................................................................63 Synodontis nigromaculatus Boulenger, 1905............................................................67 Diagnosis............................................................................................................68 Description.........................................................................................................69 Coloration...........................................................................................................70 Distribution.........................................................................................................70 Habitat................................................................................................................70 Diet.....................................................................................................................71

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vii Reproduction......................................................................................................71 Taxonomic Remarks...........................................................................................71 Synodontis tanganaicae Borodin, 1936.....................................................................78 Diagnosis............................................................................................................78 Description.........................................................................................................79 Coloration...........................................................................................................80 Distribution.........................................................................................................81 Habitat................................................................................................................81 Diet.....................................................................................................................81 Reproduction......................................................................................................81 Taxonomic Remarks...........................................................................................81 Synodontis dhonti Boulenger, 1917...........................................................................87 Diagnosis............................................................................................................87 Description.........................................................................................................88 Coloration...........................................................................................................89 Distribution.........................................................................................................90 Habitat................................................................................................................90 Diet.....................................................................................................................90 Reproduction......................................................................................................90 Taxonomic Remarks...........................................................................................91 Synodontis polli Gosse, 1982.....................................................................................97 Diagnosis............................................................................................................97 Description.........................................................................................................98 Coloration...........................................................................................................99 Distribution.......................................................................................................100 Habitat..............................................................................................................100 Diet...................................................................................................................100 Reproduction....................................................................................................100 Taxonomic Remarks.........................................................................................101 Synodontis sp. 3 n. sp................................................................................................105 Diagnosis..........................................................................................................105 Description.......................................................................................................106 Coloration.........................................................................................................107 Distribution.......................................................................................................108 Habitat..............................................................................................................108 Diet...................................................................................................................108 Reproduction....................................................................................................108 Taxonomic Remarks.........................................................................................108 5 DISCUSSION...........................................................................................................114 LIST OF REFERENCES.................................................................................................118 BIOGRAPHICAL SKETCH...........................................................................................124

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viii LIST OF TABLES Table page 1 Diagnostic characters for Lake Tanganyikan Synodontis species in which an axillary pore is present.............................................................................................10 2 Diagnostic characters for Lake Tanganyikan Synodontis species that lack an axillary pore..............................................................................................................11 3 Morphometric data and meristic counts for Synodontis granulosus ........................33 4 Morphometric measurements and meristic counts for Synodontis multipunctatus ..42 5 Morphometric and meristic counts for Synodontis sp. 1..........................................49 6 Morphometric measurements and meristic counts for Synodontis petricola ...........57 7 Morphometric and meristic counts for Synodontis sp. 2..........................................64 8 Morphometric and meristic counts for Synodontis nigromaculatus ........................74 9 Morphometric and meristic counts for Synodontis tanganaicae ..............................83 10 Morphometric measurements and meristic counts for Synodontis dhonti ...............93 11 Morphometric measurements and meristic counts for Synodontis polli ................102 12 Morphometric and meristic counts for Synodontis sp. 3........................................109

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ix LIST OF FIGURES Figure page 1 Map of the African continent with the La ke Tanganyika region indicated by the red box............................................................................................................................ ..5 2 Axillary pore of Synodontis multipunctatus BMNH 1898.9.9.76, 280 mm TL, 220 mm SL......................................................................................................................12 3 Humeral processes of Synodontis granulosus S. multipunctatus Synodontis sp 1, and S. petricola An axillary pore is presen t in all of these species.......................13 4 Humeral processes of Synodontis sp 2, S. nigromaculatus S. tanganaicae and a paratype of S. lacustricolus An axillary pore is absent in A and B........................14 5 Humeral processes of Synodontis dhonti a juvenile specimen of S. dhonti S. polli and Synodontis sp 3. An axillary pore is absent in C and D..................................15 6 Synodontis granulosus UF 160945, 192 mm TL, 149 mm SL. Top left dorsal view, bottom left lateral view, right ventral view of head...................................34 7 Distribution of Synodontis granulosus T denotes type locality.............................35 8 Synodontis multipunctatus Holotype, BMNH 1898.9.9.76, 280 mm TL, 220 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view, showing mouth and barbels....................................................................................................43 9 Distribution of Synodontis multipunctatus T denotes type locality.......................44 10 Synodontis multipunctatus (top), BMNH 1982.4.13.4789, 115 mm TL, 91 mm SL vs. Synodontis sp 1 (bottom), BMNH 1982.4.13.4784, 105 mm TL, 81 mm SL...50 11 Distribution of Synodontis sp. 1. T denotes type locality........................................51 12 Synodontis petricola Holotype, MRAC 130357, 98 mm TL, 82 mm SL. Top dorsal view, middle – lateral view, bottom – ventral view.......................................58 13 Distribution of Synodontis petricola T denotes type locality................................59 14 Synodontis sp 2, SAIAB 39577, 84 mm TL, 68 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view.......................................................................65

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x 15 Distribution of Synodontis sp 2. T denotes type locality........................................66 16 Variation in body shape and coloration in Synodontis nigromaculatus ...................75 17 Dorsal views of the head of left – Synodontis melanostictus (Holotype) and right – Synodontis nigromaculatus (Holotype), showing differe nces in occipito-nuchal shield texture............................................................................................................76 18 Distribution of Synodontis nigromaculatus T denotes type locality......................77 19 Synodontis tanganaicae Lectotype, MCZ 32538, 415 mm TL, 320 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view of head..............84 20 Synodontis tanganaicae MRAC 90288, paratype of S. lacustricolus 519 mm TL, 434 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view of head......................................................................................................................85 21 Distribution of Synodontis tanganaicae Type locality unknown...........................86 22 Synodontis dhonti Holotype, MRAC 14344, 395 mm TL, 325 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view, showing mouth and barbels......................................................................................................................94 23 Synodontis dhonti MRAC 130315, Holotype of S. irsacae 142 mm TL, 116 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view......................95 24 Distribution of Synodontis dhonti T denotes type locality.....................................96 25 Synodontis polli Holotype, MRAC 130440, 148 mm TL, 125 mm SL. Top dorsal view, middle – lateral view, bottom – ventral view.....................................103 26 Distribution of Synodontis polli T denotes type locality......................................104 27 Synodontis sp 3, UF 160942, 139 mm TL, 116 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view.....................................................................110 28 S. polli MRAC 131007-008, 137 TL, 112 SL, with intestine extracted. Hindgut chamber indicated with white arrow......................................................................111 29 Synodontis sp. 3, UF 160942, 139 mm TL, 118 mm SL, with intestine extracted. Note the relative shortness of the intes tine and lack of a discernible hindgut chamber..................................................................................................................112 30 Distribution of Synodontis sp 3. T denotes type locality.....................................113

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xi Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science TAXONOMIC REVISION OF LAKE TANGANYIKAN Synodontis (TELEOSTEI: MOCHOKIDAE) By Jeremy John Wright May 2006 Chair: Lawrence M. Page Major Department: Zoology Seven species of Synodontis have historically been recognized from Lake Tanganyika, six of which are endemic to the lake basin. The endemic species can be distinguished from all other Synodontis by a common pattern of rayed fin coloration and vertical folds in the skin. Though previous studies have examined these species, a comprehensive taxonomic examination utilizing a ll of the material av ailable has not been performed. The taxonomy of the Synodontis of Lake Tanganyika wa s investigated through by examining 336 museum specimens. Morphometri c, meristic, and nontraditional internal and external characters were examine d, resulting in the recognition of 10 Synodontis species from Lake Tanganyika (nine of which are endemic and three of which are new to science). Synodontis sp. 1 is most similar to S. multipunctatus and is distinguished from this species 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 sp.

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xii 2 is most similar to S. petricola and is distinguished from th is species by the lack of an axillary pore and the presence of light colored windows at th e bases of the rayed fins. Synodontis sp. 3 is most similar to S. polli and is distinguished fr om this species by the absence of a hindgut chamber, the shortness of the gut (0.8-1.4 times total length in S. polli vs. 4.0-5.5 times total length in S. polli ), and the presence of short, flattened papillae on the skin (vs. villous papillae in S. polli ). Examination of the data obtained during th is study offers some support for recent molecular phylogenies obtained for this group, but additional studies are needed to construct a clearer picture of relationships within this group and between this group and other members of the genus. Functional stud ies of many characters examined in this study must also be performed to further our understanding of the biology of these species.

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1 CHAPTER 1 INTRODUCTION Natural History Catfishes of the genus Synodontis are small to moderately sized fishes (up to 800 mm) that occur throughout sub-Saharan Africa. Poll (1971) divided the genus into eight geographically defined groups: the northern supraequatorial gr oup; the exclusively Nilotic group; the group of sp ecies occurring exclusively in the Niger; the western supraequatorial group from Senegal to Ogowe; the Congo Basin group; the southernmost subequatorial group; the Synodontis of eastern Africa ranging between Uebi Shebeli and the Rufigi River; and the La ke Tanganyikan group (Fig. 1). The Synodontis of Lake Tanganyika inhabit mainly rocky shoreline areas in the lake but also venture out over sandy and shell bottoms (M atthes, 1962; Coulter, 1991). The vertical distribution of these species is limited to a maximum depth of 50-100 m in the northern end of the lake and to about 240 m in the southern end (Coulter and 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 (fro m 350 m to 1470 m) (Coulter and Spigel, 1991; Tiercelin and Mondeguer, 1991). Locality data indicate that all species have a lakewide distribution, with the exception of Synodontis sp 2 and 3 (see Chapters 3, 4). Although many different color patterns exist within this genus, few are as striking or as instantly recognizable as the pattern typical of most Tanganyikan Synodontis species. This pattern consists of dark tria ngles at the bases of all of the rayed fins

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2 2 andlarge, dark colored spots on the upper body th at may or may not extend onto the belly. The barbels are typically wh ite, 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 vari es widely, from fairly small (max. TL 100 mm in Synodontis sp. 2) to quite large (max. TL 600 mm in S. tanganaicae ). Sexual dimorphism is known to occur in Synodontis species in the form of a distinct, conically shaped ge nital papilla present only in males (Matthes, 1962). Minor differences in body shape between sexes also occur in some species, with females having slightly more robust bodies. Informati on on reproduction is largely undocumented in Tanganyikan Synodontis species, with information being mostly restricted to egg counts from gravid females, though some species lack even that. The glaring exception to this statement is S. multipunctatus whose unique brood parasiti sm behavior has been well documented (Sato, 1986). Dietary information is slightly more abundant, and a variety of prey items are utilized. Most species include insect larvae and at leas t a small amount of algal matter in their diet. More specializ ed prey items that may be taken include gastropods, bivalves, sponges, crustaceans, and the eggs of other fishes (Poll, 1946, 1953; Matthes, 1962; Coulter, 1966, 1991). Systematic History Synodontis are members of the Mochokidae, a family of catfishes endemic to subSaharan Africa. With approximate ly 120 currently recognized species, Synodontis is the most diverse of any African catfish genus and, with the exception of Barbus of any African Ostariophysan. 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,

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3 3 Synodontis lacustricolus Poll (now Synodontis tanganaicae Borodin), Synodontis eurystomus Matthes (now Synodontis polli Gosse), and Synodontis nigromaculatus Boulenger. All but S. nigromaculatus are endemic to the lake. The nine endemic Synodontis species of Lake Tanganyika recognized herein can be distinguished from all other Synodontis species by the presence of a black tria ngle at the base of each rayed fin and vertical epidermal fold s along the sides of the body. Relationships between the endemic Synodontis species of Lake Tanganyika have yet to be intensively studied. Brichard (1978) indicates that the lake species of Synodontis probably do not form a monophyletic group. While it is possible that S. nigromaculatus does not share a most recen t common ancestor with the Tanganyikan endemics, given its much wide r distribution and differences from the endemic species in its rayed fin coloration a nd skin (lack of vertical folds), Brichard offers no further support for his opinion. Th e common pattern of rayed fin coloration and presence of vertical folds in the skin would appear to represent two synapomorphies supporting the monophyly of the endemic Synodontis species of Lake Tanganyika; however, a preliminary inve stigation of the molecula r phylogeny of the endemic Tanganyikan Synodontis species, utilizing mitochondrial (c yt b) data appears to support BrichardÂ’s statement (Day, pers. comm.). Th ese results are discussed later in Chapter 5. PollÂ’s revision of Synodontis while thorough for its time, suffered from a lack of material, and also appears largely to have stifled taxonomic research on this genus with only a handful of new species desc ribed after its publication. The Synodontis of Lake Tanganyika have largely been ignored by ta xonomists for the last 30 years. Recent expeditions, particularly those to the Zamb ian coast of the lake, historically poorly

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4 4 sampled for Synodontis as well as the increased popul arity of certain Tanganyikan Synodontis as aquarium species, led to the availab ility of increased numbers of specimens for study. Schraml (2003) suggested that ma ny of the specimens entering the aquarium trade may represent cryptic, unde scribed species. The aim of this study was to evaluate the taxonomy of this group, bearing in mind that cryptic species may ex ist. This led to the utilization of seve ral unconventional diagnostic char acters that revealed undescribed Synodontis species from Lake Tanganyika.

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5 5 Figure 1. Map of the African c ontinent with the Lake Tanganyi ka region indicated by the red box.

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6 CHAPTER 2 MATERIALS AND METHODS Data Collection Measurements were taken from the left si de to the nearest 0.1 mm using digital calipers and follow Poll (1971). All counts were made us ing a Leica MZ75 dissecting microscope. Vertebral and caudal fin ray c ounts were taken using lateral radiographs. Vertebral counts follow the method of Skelton and White (1990), with the first six, fused vertebrae of the Weberian appa ratus excluded. Notation for fin ray counts is as follows: upper case roman numeral = fin spine, lowe r case roman numeral = unbranched rays, Arabic numeral = branched fin rays. Te rminology for premaxillary dentition follows Skelton and White (1990). Gut length measurements and observations of the hindgut chamber of all species were performed only on specimens with a bell y cut made prior to the beginning of this study with the exception of UF specimens whic h were dissected over the course of this study. The hindgut chamber corresponds to th e “poche intestinal” of Taverne and Aloulou-Triki (1974). All photographs were taken using a Kodak EasyShare CX7430 digital camera and edited using Adobe Photosho p CS2. All maps were created using the Online Map Creator (OMC), which can be found at http://www.aquarius.geomar.de/omc/ Distribution points on maps were added usi ng Adode Photoshop CS2. Points indicate only specimens examined in this study. Instit utional abbreviations follow Leviton et al. (1985) except for the South African Instit ute of Aquatic Biodiversity (SAIAB).

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7 Characters Examined In addition to the 22 morphometric measur ements and seven meristic counts which were made for all specimens, several new ch aracters were found to be useful in the diagnosis of species. 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 distinguish species. These new characters are particular ly appealing because they allow for easy, accurate visual identification of specimens. Some of these characters, such as fin spine color are self explanatory. Those that may be unfamiliar or that are used in a taxonomic capacity for the first time in th is study are explained below. Foremost among these characters is the ax illary pore, an ope ning, or series of openings that is located between the base of the pectoral fin spine and ventral margin of the humeral process (which is it self a useful structure in iden tification) (Figs. 2-5). The function of this structure is unknown in mochokids, and the absence or presence of this structure has been noted in only one previously described Synodontis species (Friel and Vigliotta, 2006). The presence of a similar structure has previously been noted in six other catfish genera: Ariopsis (Ariidae) (Halstead et al., 1953), Ameiurus Ictalurus and Noturus (Ictaluridae) (Reed, 1907; Birkhead, 1967, 1972), Acrochordonichthys (Akysidae) (Ng and Ng, 2001), and Glyptothorax (Sisoridae) (pers. obs.). In Acrochordonichthys this structure has been shown to produce a mucosal secretion that possesses toxic propertie s (Ng and Ng, 2001), while the util ity of this secretion in the other genera has been a matter of some debate. The presence or absence of this character divides the ten Tanganyikan species of Synodontis into two groups; four species which have an axillary pore (Table 1) and six in which it is lacking (Table 2).

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8 All Synodontis species have a premaxillary toot hpad, which has several rows of unicuspid, chisel-shaped teeth. In some speci es 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 transversely bisects the toot hpad. The separated c ondition is referred to herein as interrupted, while the unseparated condition is re ferred to as uninterrupted. This appears to be the first study to examine this character in Synodontis though the length of the premaxillary pad has often been used in previous studies. The skin of the nine endemic Tanganyikan Synodontis species recognized in this study is unique among the genus in having numer ous minute, vertical epidermal folds. The purpose of these folds is unknown. These folds are useful in distinguishing members of the Tanganyikan species flock from species outside the lake’s basin, though they are not helpful in distinguishing between species fr om the lake. The papillae on the skin of certain species, however, help to separate them from other Tanganyikan species in which the skin is bare. This character is most useful in the identific ation of fresh or well preserved specimens, as the papillae may degrade over time. The intestinal anatomy of some Synodontis species is extremely interesting due to the presence of a swelling in the posterior part of the intestine which Taverne and Aloulou-Triki termed the “poche intestinal” and which is referred to herein as the hindgut chamber. A similar structur e is known only in species of Kyphosidae, a family of herbivorous, marine fishes th at utilizes fermentative dige stion in the hindgut (Rimmer and Wiebe, 1987; Clements and Choat, 1997; Dougl as et al., 2002). The presence of this structure in highly herbivorous mochokid species such as S. polli Brachysynodontis batensoda and Hemisynodontis membranaceus suggests that a similar role may be

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9 played by the mochokid hindgut chamber, marking the first time that such a behavior and structure were found in a freshwater fis h. The presence or absence of the hindgut chamber appears to be species-specific and can distinguish between species for which few good external diagnos tic characters exist.

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Table 1. Diagnostic characters for Lake Tanganyikan Synodontis species in which an axillary pore is present. S. granulosus S. multipunctatus Synodontis sp. 1 S. petricola S. tanganaicae S. dhonti Mandibular teeth 28 – 51 13 – 29 17 – 26 31 – 50 33 – 49 15 – 38 Axillary pore Large Large Large Small Large Large Large spots on body Absent Present Present Present Absent Present – young Absent – adults Fin spine color Dark Dark Dark White Dark Dark Pectoral-fin rays 7 – 8 8 7 8 – 9 8 – 9 8 – 9 Eye/Snout length 31.2 50.2% 44.9 62.0% 64.2 81.0% 28.7 40.1% 16.0 26.9% 23.1 – 43.7% Premaxillary toothpad Uninterrupted Uninterrupted Uninterrupted Interrupted Interrupted Interrupted Maximum TL 270 mm 280 mm 150 mm 135 mm 600 mm 400 mm Secondary branches on medial mandibular barbel Absent Absent Absent Present Present Present Occipito-nuchal shield covered with skin No Yes Yes Yes No No Papillae on body skin Present, granular Absent Absent Absent Present, granular Present, granular Gut/body length 0.5 times 0.5-0.8 times 0.8-0.9 times 1.2-1.3 times No data 0.7 – 0.8 times Hindgut chamber Absent Absent Absent Present No data Absent 10

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11 Table 2. Diagnostic characters for Lake Tanganyikan Synodontis species that lack an axillary pore. Synodontis sp. 2 S. nigromaculatus S. polli Synodontis sp. 3 Mandibular teeth 38 – 51 23 – 38 40 70 50 – 66 Spots on body Large Small Moderate to large Moderate Fin spine color White Dark Dark Dark Black triangles on rayed fins Present, with light windows at base Absent Present Present Eye/Snout length 25.2 35.8% 31.1 48.1% 25.8 39.3% 26.0 33.4% Premaxillary toothpad Interrupted Interrupted Interrupted Interrupted Maximum TL 100 mm 380 mm 180 mm 150 mm Secondary branches on medial mandibular barbel Present Present Present Present Occipito-nuchal shield covered with skin Yes No Yes Yes Papillae on body skin Absent Present, villous Present, villous Present, flattened Gut/body length No data 1.3-1.6 tim es 4.0-4.5 times 0.8-1.4 times Hindgut chamber No data Present Present Absent Certain traits not diagnostic within this group (premaxillary toothpad, secondary branches on medial mandibular barbel) are incl uded to facilitate comparison with species in Table 1.

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12 Figure 2. Axillary pore of Synodontis multipunctatus BMNH 1898.9.9.76, 280 mm TL, 220 mm SL.

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13 Figure 3. Humeral processes of A) Synodontis granulosus UF 160945, 192 mm TL, 149 mm SL, B) S. multipunctatus BMNH 1898.9.9.76, 280 mm TL, 220 mm SL, C) Synodontis sp 1, 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 axillary pore is present in all of these species.

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14 Figure 4. Humeral processes of A) Synodontis sp 2, SAIAB 42518, 97 mm TL, 78 mm SL, B) S. nigromaculatus BMNH 1905.11.10.10, 237 mm TL, 186 mm SL, C) S. tanganaicae 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 all of these species.

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15 Figure 5. Humeral processes of A) Synodontis dhonti Holotype, MRAC 14344, 395 mm TL, 325 mm SL, B) juvenile specimen of S. dhonti MRAC A3-033-P-00360040, 139 mm TL, 116 mm SL, C) S. polli MRAC A3-033-P-0076-0080, 160 mm TL, 132 mm SL, D) Synodontis sp 3, UF 160942, 139 mm TL, 118 mm SL. An axillary pore is abse nt in all of these species.

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16 CHAPTER 3 MATERIALS EXAMINED Number of specimens examined from each lot is indicated in parentheses. Synodontis granulosus (1) BMNH 1906.9.6.40, Lectotype, TL 270 mm, SL 216 mm, north end of Lake Tanganyika, coll. J.E.S. Moore, (2 ) BMNH 1906.9.6.41-42, Parale ctotypes, TL 220-246 mm, SL 181-202 mm, (3) BMNH 1936.6.15.1199-1201, TL 213-249 mm, SL 165-189 mm, Lake Tanganyika, coll. Christ y, (1) MRAC 130378, TL 186 mm, SL 143 mm, Island of Moboko, Lake Tanganyika, coll. H. Matthes, 3.IX.1958, (1) MRAC 94-069-P0289, 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 82012-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 (Zambia), 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 Ha uschner, Tropical Fish Collector, coll. V.2005.

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17 Synodontis multipunctatus (1) BMNH 1898.9.9.76, Holotype, TL 280 mm, SL 220 mm, Sumba, coll. J.E.S. Moore, (2) BMNH 1906.9.8.70-1, 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. Belge, (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. Ro y. Sci. Nat. Belge, (1) BMNH 1955.12.20.1852, TL 99 mm, SL 75 mm, Moba Bay, La ke 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. Belge, 21.XI.1946, (1) BMNH 1955.12.20.1845, TL 213 mm, SL 168 mm, MÂ’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. Na t. Belge, 7.III.1947, (2) BMNH 1955.12.20.18551856, TL 150-209 mm, SL 123-166 mm, S. of Malagara si delta, offshore of the southern point of the Malagarasi delta, L. Tanganyika coll. Inst. Roy. Sci. Nat. Belge, 26.III.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. Belge, 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 6779 mm, SL 53-64 mm, Mpulungu, Lake Tanganyika (Zambia), coll. J. Day, 2005, (3) BMNH 2005.9.26.8-10, TL 6169 mm, SL 48-55 mm, Edith Bay, Lake Tanganyika (T anzania), 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 7199 mm, SL 54-74

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18 mm, Kigoma, Lake Tanganyika (Tanzan ia), coll. J. Day, 2005, (2) MRAC 39186-39187, TL 94-96 mm, SL 72-73 mm, Moba (Lake Tanganyika), coll. Van Maldern, 1932, (3) MRAC 53096-53098, 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 90273-90274, 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.), 1819.IV.1947, (1) MRAC 91606-91608, 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, total length 75-105 mm, SL 60-83 mm, Magara, route Bujumbura-Nyan za Lac (Burundi), coll. L. De Vos, 19.10.1994, (1) MRAC A1-094-P-0054, TL 80 mm, SL 61 mm, Kasakalawe, Lake Tanganyika, S 847.23’, E 3104.40’ (Zambia), coll. Snoeks, Hanssens, Verheyen et al., (2) SAIAB 39578, TL 77-85 mm, SL 62-67 mm, Country: Zambia, System: Lake Tanganyika; Mpulungu, Musende Rocks, 08 46’00”S, 03151’00”E, coll. 7.VII.1992, (3) SAIAB 40174, TL 58-96 mm, SL 45-76 mm, Country: Zambia, System: Lake Tanganyika; Musende Rocks, 0846’00”S, 03106’00”E, coll. 05.VIII.1992, (1) SAIAB 56212, TL 73 mm, SL 57 mm, Kigoma Bay, below Hill top Hotel, Tanzania, 0453’03”S, 02937’11”E, coll. 08.X.1997, (1) SAIAB 56251, TL 71 mm, SL 58 mm, Jacobsen’s beach, Tanzania, 0454’31”S, 02936’02”E, coll. 04.X.1997, (4) SAIAB 56254, TL 82103 mm, SL 63-81 mm, Jacobsen’s beach, Tanzania, 0454’31”S, 02936’02”E, coll.

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19 05.X.1997, (1) SAIAB 76164, TL 78 mm, SL 63 mm, Zambia, Province: Northern, Area: Mbala, System: Lake Tanganyika; Mbita Is land (northwest end), 845.18Â’S, 3105.07Â’E, coll. 29.II.2004, (3) SAIAB 76175, TL 56-76 mm, SL 44-59 mm, Country: Zambia, Province: Northern, Area: Mbala, System : Lake Tanganyika; Musende Rocks beach, 0845.18Â’S, 3105.07Â’E, coll. 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 101103 mm, SL 77-79 mm, Tanzania; Kigoma; Jacobs enÂ’s Beach; coll. Wagner, Catherine, 13.VIII.2005. Synodontis sp. 1 (2) BMNH 1920.5.25.83-84, TL 98-103 mm, SL 76-79 mm, Vua Bay, L. Tanganyika, coll. L. Stappers, (1) BMNH 1955.12.20.1833-1835, TL 103 mm, SL 80 mm, Moba Bay, L. Tanganyika, coll. Inst. Roy. Sci. Nat. Belge, 7.III.1947, (1) BMNH 1955.12.20.1837, TL 101 mm, SL 78 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 1982.4.13:4789-4793, TL 109-117 mm, SL 8692 mm, Kigonga Bay, L. Tanganyika, co ll. R. Travers, (2) BMNH 1982.4.13.4785-4786, TL 112-129 mm, SL 91-96 mm, Mwakizega coastlin e, L. Tanganyika, coll. R. Travers, (1) BMNH 1982.4.13:4787, TL 103 mm, SL 80 mm, Ki birzi beach, L. Tanganyika, coll. R. Travers, (2) BMNH 1982.4.13:4787-4788, TL 104120 mm, SL 81-93 mm, ElephantÂ’s Foot Peninsula, L. Tanganyika, coll. R. Travers, (1) BMNH 2005.9.26.11-16, TL 94 mm, SL 73 mm, Kigoma, Tanzania, coll. J. Day, 2005, (3) MRAC 39186-39192, TL 93-100 mm, SL 72-80 mm, Moba (Lake Tanganyika ), coll. Van Maldern, 1932, (2) MRAC 53096-53100, TL 107-114 mm, SL 80-91 mm, Nyanza (L. Tang.), coll. A. Testiade, 1937, (3) MRAC 90273-90277, TL 123-147 mm, SL 96-111 mm, Stat: 63, offshore from

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20 the Malagarasi, 10-15 km from the coast, coll. M. Poll, 3.II.1947, (2) MRAC 9160691608, TL 114-121 mm, SL 96-98 mm, Uvira, La ke Tanganyika, coll. G. Marlier, 1.IX.1949, (1) SAIAB 56254, TL 107 mm, SL 81 mm, Jacobsen’s beach, Tanzania, 0454’31”S, 02936’02”E, coll. 05.X.1992. Synodontis petricola (1) MRAC 130357, Holotype, TL 98 mm, SL 82 mm, Kashekezi, N. Lake Tanganyika, coll. H.Matthes, 3.IV.1959, (9 ) Paratypes, TL 40-82 mm, SL 32-67 mm, Luhanga, South of Makobola, coll. H.Ma tthes, 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 97140 mm, SL 78-111 mm, Mahumba, km 36 route Bujumbura-Lake Nyanza, Lake Tanganyika (Burundi), coll. L. De Vos, 29.I.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-P0581, 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 Tanga nyika (Zaire), coll. L. De Vos, 2.X.1995, (1) MRAC 95-096P-2576, TL 69 mm, SL 56 mm, 0820.24’S, 3031.21’E, Chisiki, coll. Verheyen, Snoeks, Hanssens, Ruber, Sturmbauer, 10.IV.1995, (1) MRAC A1-094-P-53, TL 81 mm, SL 65 mm, Crocodile Island, Lake Tanganyika (Zaire), coll. Snoeks, Hanssens, Verheyen et al., 16.X.2001, (1) BMNH 1936.6.14.1232, Paratype, (also I. D.’d as S.multimaculatus), TL 88 mm, SL 71 mm, Lake Tanganyika, coll. Christy, (2) SAIAB 42518, TL 70-74 mm, SL

PAGE 33

21 56-59 mm, Country: Zambia, System: Lake Tanganyika; Musende Rocks, 0846’00”S, 03107’00”E, coll. 10.X.1992, (1)SAIAB 56255, TL 68 mm, SL 55 mm, Country: Tanzania; Jacobsen’s beach, 0454’31” S, 02936’02”E, coll. 05.X.1997, (1) SAIAB 56225, TL 75 mm, SL 61 mm, Country: Tanzania ; Kigoma Bay, below Hill top Hotel, 0453’03”S, 02937’11”E, coll. 09.X.1997, (1) SAIAB 56232, TL 80 mm, SL 64 mm, Country: Tanzania; Cave, Kigoma Hotel belo w Hill top, 0453’03”S, 02937’11”E, coll. 11.X.1997, (1) SAIAB 56244, TL 79 mm, SL 64 mm, Country: Tanzania; Bangwe, shingle beach, 0453’53”S, 02935’39”E, coll. 12.X.1997. Synodontis sp. 2 (9) SAIAB 39577, TL 58-98 mm, SL 47-79 mm, Zambia; Mpulungu, Musende Rocks, 0846’00”S, 03151’00”E, coll. 07.VII.1992, (3) SAIAB 40171, TL 70-85 mm, SL 56-69 mm, Country: Zambia; Musende Rocks, 0846’00”S, 03106’00”E, coll. 05.VIII.1992, (4) SAIAB 42518, TL 55-97 mm, SL 44-78 mm, Country: Zambia, System: Lake Tanganyika; Musende Rock s, 0846’00”S, 03107’00”E, coll. 10.X.1992. Synodontis nigromaculatus (1) BMNH 1905.11.10.10, Holotype, TL 237 mm, SL 186 mm, Lake Bangwuelo, Zambia, coll. Esq. R. Melland, (1) BMNH 1906.9.8.72, Holotype ( S. melanostictus ), TL 290 mm, SL 237 mm, Lake Tanganyika, coll. Cunnington, (1) BMNH 1908.11.6.29, TL 244 mm, SL 177 mm, Maramba River, Upper Zambezi, coll. G.T. Corrington, (1) BMNH 1907.9.30.9, TL 319 mm, SL 238 mm, Lake Bangwelo, coll. R.H. Melland, (4) BMNH 1920.5.26.95-102, TL 58-115 mm, SL 46-93 mm, Cape Kasangeneke, Lake Mweru, coll. L. Stappers, (1) BMNH 1976.3.18: 2439-2440, TL 192 mm, SL 153 mm, Thalamakane River at Maun (Botswana), coll. K. Be rne, (1) MRAC 34006, TL 383 mm, SL 307 mm, Lukonzolwa, Lake Moero, coll. DeWitt e, 9-17.II.1931, (1) MRAC 34007, TL 324 mm,

PAGE 34

22 SL 250 mm, Lukonzolwa, Lake Moero, co ll. DeWitte, 9-17.II.1931, (2) MRAC 37814815, TL 177-317 mm, SL 134-252 mm, Kasenga, Luapula River, coll. DeWitte, 10.III.1931, (1) MRAC 56319-56320, TL 132 mm, SL 104 mm, Luapula au Bangwelo, coll. Breolo, III.1938, (2) MRAC 162207208, 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) MRAC 87132-133, TL 154 mm, SL 129 mm, Kasaji, Katanga, coll. Rev. S. Fischer, 5.XII.1952, (1) MRAC 162219, TL 183 mm, SL 141 mm, Lake Calundo, coll. indigenous peoples, I.1955, (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 162210-213, TL 274-306 mm, SL 209-229 mm, Chiumbe River, near the junction with th e Chilambo River (S745’, E2105’) (Angola), coll. Barros Machado (Dundo Museum), 24.XI.1957, (2) MRAC 126293-294, TL 152172 mm, SL 122-134 mm, Uvira, Lake Tanga nyika, coll. G. Marlier, 13.IX.1958, (1) MRAC 149537, TL 181 mm, SL 142 mm Luwohoshi River, affl. Ruashi River, village of Shindaika, Katanga, coll. M. Li ps, 25.III.1963, (1) MRAC 165806-807, TL 104 mm, SL 80 mm, Chamberi/Chambezi River (Zam bia), coll. H. Matthes, 16.VI.1965, (1) MRAC 172442, TL 177 mm, SL 134 mm, Lake Ba ngwelo, coll. Bell-Cross, (1) MRAC 94-019-P-1007, TL 152 mm, SL 116 mm, S outh Mofwe Lagoon, Luapula Swamp (Zambia), coll. P. Van Zweiten, 16. II.1993-26.III.1993, (1) MR AC 94-019-P-2136, TL 208 mm, SL 158 mm, Pembe lagoon, near Ka totoma, south lagoon, Luapula swamps (Zambia), coll. P. Van Zweiten, 17.V III.1993, (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, (1) MRAC 162203-204 (orig. I.D. S.

PAGE 35

23 zambesensis ), TL 180 mm, SL 132 mm, Nhefo River, a fflt. The river to the left of the Luachimo, close to 50 km S. of Dundo (Angola), coll. Barros Machado (Dundo Museum), III.1948, (1) MRAC 162206 (orig. I.D. S. zambesensis ), TL 241 mm, SL 186 mm, Dundo, Luachimo River (Angola), co ll. Barros Machado (Dundo Museum), XI.1948, (1) 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) MRAC 162215 (orig. I.D. S. zambesensis ), TL 180 mm, SL 136 mm, Cachimo, Tuembe River, S806Â’, E2129Â’ (Angola), coll. A.L. Terreira (Dundo Museum), 22.VIII.1961, (1) MRAC 149538 (orig. I.D. S. zambesensis ), TL 160 mm, SL 122 mm, Moushoshi River near the junction wi th the Kofubu River (Katanga), coll. M. Lips (don. M. Mignolet), 6.V.1963. Synodontis tanganaicae (1) MRAC 130720, Paratype ( S. lacustricolus ), TL 333 mm, SL 255 mm, Mboko Island, N. Lake Tanganyika, coll. Marlier (I.R.S.A.C.), 7.II.1957, (1) MRAC 90288, Paratype ( S. lacustricolus ), TL 519 mm, SL 434 mm, Temb we Bay, Lake Tanganyika, coll. M. Poll/Expl. Hydrobiog.Tang., 13.II.1947, (1) MRAC 90289, Paratype ( S. lacustricolus ), TL 582 mm, SL 489 mm, Edith Bay, Lake Tanganyika, coll. M. Poll/Expl. Hydrobiog. Tang., 14-15.II.1947, (2) MRAC 96-083-P-003-004, TL 300-330 mm, SL 240-265 mm, Zambia, coll. De Vos, 1996, (1) BMNH 1955.12.20.1849, Paratype ( S. lacustricolus ), TL 385 mm, SL 315 mm, Opposite Ka la, Lake Tanganyika, Inst. Roy. Nat. Belge, (1) BMNH 1955.12.20.1850, Paratype ( S. lacustricolus ), TL 361 mm, SL 291 mm, Tembwe Bay, Lake Tanganyika, In st. Roy. Nat. Belge, (1) BMNH 1955.12.20.1851, Paratype ( S. lacustricolus ), Malagarasi, Lake Tanganyika coll. Inst. Roy. Nat. Belge,

PAGE 36

24 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. Synodontis dhonti (1) MRAC 14344, Holotype, TL 395 mm, SL 325 mm, Lake Tanganyika, Kilewa, coll. Stappers, 18.V.1912, (1) MRAC 130315, Holotype ( S. irsacae ), TL 142 mm, SL 116 mm, Kalundu, N. Lake Tanganyika, co ll. H. Matthes, 27.X.1959, (6) MRAC 130316-321, Paratypes (S. irsacae), TL 55-156 mm, SL 46-124 mm, Kalundu, N. Lake Tanganyika, coll. H. Matthes, 14.I V.1960, (6) MRAC 130332-130337, TL 47-102 mm, SL 38-84 mm, Luhanga, Lake Tanganyika, co ll. H. Matthes (I.R.S.A.C.), IV.1959, (1) MRAC 77-40-P-5-7 (Orig. I.D. S. polli), 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-01270128, TL 83-105 mm, SL 69 mm-85 mm, Ulwile Island, northern shore, Lake Tanganyika S727.40Â’ E3024.20Â’ (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, 0843.31Â’S, 3108.01Â’E, coll. Verheyen, Snoeks, Hanssens, Ruber, Stermbauer, 4.IV.1995, (4) MRAC A3-033-P-0036-0040, TL 117161 mm, SL 91-133 mm, Pemba, S. of Uvira, Lake Tanganyika (Zaire), coll. L. De Vos, 18.VIII.1995, (1) MRAC A1-094-P0055-0059, TL 103 mm, SL 81 mm, Katoto, Lake Tanganyika (Zambia), coll. Snoeks, Hanssens, Verheyen et al., 18.X.2001, (1) BMNH 1955.12.20.1859, TL 192 mm, SL 157 mm, Albertville, Lulumduie village, Lake Ta nganyika, coll. Inst. Roy. Sci. Nat. Belge, 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, SL 70-97 mm, Country: Zambia, System: Lake Tanganyika, Locality: Mbita Island,

PAGE 37

25 0845’00”S, 03106’00”E, coll. 08.VIII.1992, (1) SAIAB 56231, TL 133 mm, SL 108 mm, Cave, Kigoma hotel be low Hill top, 0453’03”S, 02937’11”E, coll. 04.X.1997, (1) SAIAB 56255, TL 91 mm, TL 72 mm, Country : Tanzania; Jacobsen's beach, 0454’31”S, 02936’02”E, coll. 05.X1997, (2) SAIAB 56687, TL 94-97 mm, SL 75-79 mm, Country: Tanzania, System: Lake Tanganyika; Muzungu beach, 0454’53”S, 02935’58”E, coll. 10.X.1997, (2) SAIAB 7610 5, TL 93-98 mm, SL 76-80 mm, Country: Zambia, Province: Northern, Area: Mbala, System: Lake Tanganyika, Water body: Lake Tanganyika; Mbita Island (northwest e nd), 0845.18’S, 3105.07’E, coll. 29.II.2004, (1) SAIAB 76174, TL 97 mm, SL 79 mm, Country: Zambia, Province: Northern, Area: Mbala, System: Lake Tanganyika, Water body: Lake Tanganyika; Musende Rocks beach, 0845.18’S, 3105.07’E, coll. 29.II.2004, (2) UF 160940, TL 104-124 mm, SL 86-102 mm, Cape Chaitika, Lake Ta nganyika, Zambia, via Pete Hauschner, Tropical Fish Collector, coll. 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. Synodontis polli (1) MRAC 130440, Holotype, TL 148 mm, SL 125 mm, Luhanga, N. Lake Tanganyika, coll. H. Matthes, 3.IV.1959, (3) MRAC 130444-446, Paratypes, TL 97-144 mm, SL 79-120 mm, Makobola, Lake Tanganyika, coll. H. Matthes (I.R.S.A.C.), 10.XI.1958, (2) MRAC 130442-443, Paratypes, TL 48-58 mm, SL 38-47 mm, Luhanga, N. Lake Tanganyika, coll H. Matthes (I.R.S.A.C.), 31.VIII.1950, (1) MRAC 130464, Paratype, TL 115 mm, SL 96 mm, Bemba, Lake Tanganyika, coll. H. Matthes (I.R.S.A.C), 26.IV.1958, (2) MRAC 130452-453, Paratypes, TL 57-60 mm, SL 46-48 mm, Makobola, Lake Tanganyika, coll. H. Matthes (I.R.S.A.C.), 28.VI.1960, (2) MRAC 130462-463, Paratypes, TL 69-85 mm, SL 56-70 mm, Kashkezi, Lake Tanganyika, coll.

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26 H. Matthes (I.R.S.A.C.), 5.II.1959, (6) MRAC 130454-461, Paratypes, TL 39-116 mm, SL 31-96 mm, Makobola, coll. H. Matthes (I.R.S.A.C.), 8.XI.1960, (2) MRAC 131007131008, TL 137-144 mm, SL 112-116 mm Kalungwe, Lake Tanganyika, coll. G. Leleup, 13.VII.1961, (2) MRAC 96-031-P-08890890, 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 A3-033P-0075, TL 121 mm, SL 103 mm, Pemba, S. of Uvir a, 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-00820082, TL 146 mm, SL 120 mm, Magara, route Bu jumbura-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, La ke Tanganyika, S0457.87’, E2935.86’ (Tanzania), coll. G. Kazumbe, 7.VI.2004, (2) MRAC 77-40-P-5-7, TL 88-99 mm, SL 7082 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. Bricha rd (6) MRAC 95-096-P-2561-2575, TL 85-108 mm, SL 71-90 mm, Kasenga Point, lake Tanganyi ka, S0843’31”, E3108’01” [Zambia]; coll. Exp. Tanganyika 95, 4.IV.1995, (5) MRAC 95-096-P2576-2585, TL 90-116 mm, SL 7498 mm, Chisiki, S0820’24”, E3031’ 21” [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, S0829’22”, E30 28’32” [Zambia], coll. Exp. Tanganyika 95, 10.IV.1995, (1) MRAC 95-096-P-2609, TL 135 mm, SL 112 mm, Kasaba Bay, lake

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27 Tanganyika, S0830’57”, E3038’52” [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, S0808’47”, E3058’14” [Zambia]; coll. Exp. Tanganyika 95, 19.IV.1995, (1) MRAC 95-096-P-2627, TL 108 mm, SL 87 mm, Chikulina Bay, lake Tanganyika, S0832’41”, E3043’36” [Zambia]; coll. Exp. Tanganyika 95, 12.IV.1995, (1) BMNH 2005.9.26.17-18, TL 120 mm, SL 98 mm, Mpulungu, Zamb ia, Lake Tanganyika, coll. J. Day, 2005, (3) SAIAB 40171, TL 980114 mm, SL 81-94 mm, Country: Zambia, System: Lake Tanganyika; Musende Rocks, 0846’00”S, 03106’00”E, coll. 05.VIII.1992, (1) SAIAB 42518, TL 105 mm, SL 87 mm, Country : Zambia, System: Lake Tanganyika; Musende Rocks, 0846’00”S, 03107’00”E, coll. 10.X.1992, (2) SAIAB 56262, TL 122150 mm, SL 100-127 mm, Country: Tanzan ia; Jacobsen’s Beach, 0454’31”S, 02936’02”E, 10.X.1997, (1) SAIAB 76167, TL 117 mm, SL 100 mm, Country: Zambia, Province: Northern, Area: Mbala, Syst em: Lake Tanganyika, Water body: Lake Tanganyika; Mbita Island (northwest e nd), 0845.18’S, 03105.07’E, coll. 29.II.2004, (1) CU 88758, TL 180 mm, SL 145 mm, Tanzania; Ki goma; Jacobsen’s Beach, coll. P.B. McIntyre, 18.VII.2002. Synodontis sp. 3 (5) MRAC 78-25-P-20-24, TL 93-136 mm, SL 79-114 mm, Cape Chaitika, S. Lake Tanganyika (Zambia); coll. P. Bricha rd, IV.1978, (10) UF 160942, TL 128-148 mm, SL 108-121 mm, Lake Tanganyika at Chaitika, Zamb ia, via Pete Hauschner, Tropical Fish Collector, coll. V.2005.

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28 CHAPTER 4 SPECIES DESCRIPTIONS Invariant Traits Lateral line complete and midlateral, exte nding onto base of caudal fin. Distinct genital papilla present in male; absent in fe male. Anterior nostrils tubular; posterior nostrils with semicircular flaps of skin al ong anterior margin. Gill opening extends from lower edge of ventral rounded occipital-nuchal process to poi nt horizontal to base of pectoral fin spine. Gill me mbranes broadly joined at isth mus. Two ovaline patches of minute, embedded teeth posterior to mandibular tooth row(s) Well developed, papillate, membranous flap immediately posteri or to premaxillary toothpad. Synodontis granulosus Boulenger, 1900 (Figures 3,6,7; Tables 1, 3) Synodontis granulosus Boulenger, G.A., 1900:480, descrip tion, n. sp., northern end of Lake Tanganyika; 1901:308, description, northern end of Lake Tanganyika; 1901:149, pl. 16, description, northern e nd of Lake Tanganyika; 1911:413, fig. 311, description, northern end of La ke Tanganyika. Moore, J.E.S., 1903:166, fig., northern Lake Tanganyika – Worthington, E.B. and Ricardo, C.K., 1936:1067, 1077, 1101, note, Kabanga Ba y, Lake Tanganyika. – Poll, M., 1946:219, description, Kilewa Bay; 1953: 152, description, Lake Tanganyika; 1971:303, fig. 138, 139, pl. V, XI, descrip tion, Lake Tanganyika. – Hulot, A., 1950:170, Lake Tanganyika. Matthes H., 1962:44, description, Lake Tanganyika. – Coulter, G.W., 19651966:34, abundance, Lake Tanganyika; 1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance, ecology, Lake Tanganyika. – Brichard, P., 1978:424, identification, Lake Tanganyika; 1989:478, 479, photo, identification. – Sands D., 1983:23, check-list. – Daget, J., Gosse, J.P., and Thys van den A udenaerde, D.F.E, 1986:129, check-list. – Burgess, W.E., 1989:576, pl. 94, La ke Tanganyika. – Kobayagowa, M., 1989:14, photo. – Coulter, G.W., 1991a:152, 154, 182, 191, abundance, diet, habitat, reproduction.

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29 Diagnosis Axillary pore present; mandibular teeth 28-51 ; body lacking large spots; fin spines brown to black; 7-8 pectoral fi n rays; black triangles present on base of all rayed fins; eye 31.2-50.2% snout length; premaxillary toothpad uninterrupted; secondary branches on internal mandibular barbels abse nt; occipito-nuchal shield not covered with skin; granular papillae present on skin of body; hindgut chamber absent; maximum TL 270 mm. Synodontis granulosus can be distinguished from Synodontis sp. 2, S. nigromaculatus S. tanganaicae S. dhonti S. polli and Synodontis sp. 3 by the presence of an axillary pore. Synodontis granulosus differs from S. multipunctatus and Synodontis sp. 1 in lacking large spots on the body, ha ving a higher number of mandibular teeth (2851 vs. 13-29 in S. multipunctatus and 17-26 in Synodontis sp. 1), a smaller eye (31.250.2% of snout length vs. 44.9-62.0% in S. multipunctatus and 64.2-81.0% in Synodontis sp. 1), an occipito-nuchal shield which is not covered by skin, and body skin which is covered by granular papillae. Synodontis granulosus is distinguished from S. petricola by lacking large spots on the body and seconda ry branches on the internal mandibular barbels, having a large axillar y pore (vs. very small in S. petricola ), darkly colored fin spines, 7-8 pectoral fin rays (vs. 8-9 in S. petricola ), an uninterrupted premaxillary toothpad, much larger body size (ma x. length 270 mm vs. 135 mm in S. petricola ), an occipito-nuchal shield which is not cove red by skin, and skin which is covered by granular papillae. Large specimens of S. granulosus may be confused with S. tanganaicae However, the humeral process is much narrower in S. granulosus (humeral process length 253.8-437.2% of humeral process width in S. granulosus compared to 172.9-255.4% in S. tanganaicae ) (figs. 3a, 4c,d), as well as the presence of a large

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30 axillary pore in S. granulosus (fig. 2), vs. pore absent in S. tanganaicae are sufficient to separate these species. Description Morphometric and meristic data are given in Table 3. Maximum total length 270 mm, standard length 216 mm. Body compressed. Predorsal profile straight, interrupted by convex ridge formed by orbit of eye. Preanal profile straight to gently convex. Skin on body forming numerous vertical epiderma l folds covered with minute, granular papillae extending onto all fins. Head depressed and broad; skin covered with granular papillae. Snout with bluntly rounded margin when viewed laterally and dorsa lly. Anterior nostrils tubular; posterior nostrils with semicircular flaps of skin along anterior margin. Occipital-nuchal shield rugose, not covered with skin; terminating posteriorly with narro w, rounded process on either side of dorsal spine; ventrally with narrow, bluntly pointed process which extends to upper margin of humeral process on either side of body. Eye lo cated latera lly; ovoid; horizontal axis longest. Interorbital area flat. Mouth subterminal; lips wide and papill ate. Mandibular teeth 28-51, short, unicuspid; arranged in single transverse row. Premaxillary toothpad uninterrupted; primary, secondary and tertiary premaxillary teeth discrete; nume rous; arranged in 4, 4, and 2 irregular rows, respectively. Maxillary barbel with narrow basal membra ne; lacking branches or crenelations; extending at least to base of pectoral fin. External mandibular barb el extending to point just short of anterior margin of pectoral girdle, with 4-5 non-tuberculate branches; lacking secondary branches. Internal mandi bular barbel about 1/2 length of external barbel; with 4-5 pairs of tuberculate branches; lacking secondary branches.

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31 Dorsal fin I,7-8; posterior margin straight Dorsal fin spine long, striated, nearly straight, terminating in short black filamen t; anterior margin smooth; posterior margin with small serrations distally. Pectoral fin I ,7-8; posterior margin st raight. Pectoral fin spine roughly equal in length to dorsal fin spin e, striated, slightly curved, terminating in short, black filament; anterior spine margin granulate in adults; many small antrorse serrations along anterior margin in juveniles; posterior spine margin with large retrorse serrations along entire length. Adipose fin we ll developed; margin convex. Pelvic fin i,6; located anterior to verti cal through origin of adipose fi n; tip of appressed fin barely reaches base of anal fin. Anal fin iii-v,79,i; posterior margin nearly straight; base located ventral to adipose fin. Cauda l fin I,7,8,I; forked; lobes pointed. Humeral process narrow, elongated, granul ous; possessing distinct ridge on ventral margin; dorsal margin concave; terminating in sharp point (Fig. 3A). Large, dark-colored axillary pore present just ventral to humeral process. Gut 0.5 times body length (n = 1, UF160945). Hindgut chamber absent. Coloration Body slate-gray to olive brow n (Fig. 6). Belly ranging fr om pale yellow to gray. Scattered, small black spots present on juveni le specimens. Maxillary barbels white. Mandibular barbels white, bases sometimes with scattered dusky pigmentation. Iris copper colored. Dorsal and pectoral fin sp ines brown to black, filaments black. All rayed fins with black triangles at base, poste rior margins white to yellowish in color. Both lobes of caudal fin with black bar from ba se to tip of fin; posterior margin of fin white to yellowish in color. Distribution Lake Tanganyika (Fig. 7); rare.

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32 Habitat Littoral to benthic zones over shell, sand and mud bottoms; to a maximum depth of 130 m (Coulter, 1991). Diet Almost nothing is known about the diet of this species, though one publication does reports a single Limnothrissa in the stomach of one specimen (Worthington and Ricardo, 1933). Many other Tanganyikan species of Synodontis prey largely on mollusks and algae scraped from rocky substrates. Presumably, S. granulosus has a similar diet, but further study is clearly needed to co nfirm or refute this hypothesis. Reproduction No information exists regarding the reproduction of S. granulosus Taxonomic Remarks Synodontis granulosus is sufficiently distinct that it is one of only two described Tanganyikan species of Synodontis that have remained unalte red with respect to their taxonomy since their original descriptions (the other being S. multipunctatus ). Minor differences in coloration pattern occur throughou t its range (mainly in the color of the fin margins), but they do not appear to follow any discernible pattern with respect to geography or population.

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33 Table 3. Morphometric data and meristic counts for Synodontis granulosus All morphometric measurements given in per cent of base measurement. Meristic data for Lectotype is identified by a “*” superscript. SL = Standard length, HL = Head length, XMB = External mandibular barbel, IMB = Internal mandibular barbel, HPL = Humeral pro cess length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 28.1 20.8-30.8 25.22.8 Head length/SL 27.7 27.4-31.7 28.81.4 Snout-dorsal length/SL 36.9 36.9-41.9 38.91.4 Adipose fin length/SL 26.4 25.6-39.0 30.03.7 Maxillary barbel length/SL 28.5 22.9-38.3 31.14.7 Dorsal spine length/SL 26.5 24.3-31.9 28.32.2 Pectoral spine length/SL 23.8 23.8-31.1 27.01.9 Head width/HL 94.8 90.1-105.2 97.03.7 Head depth/HL 88.4 71.4-88.6 81.64.5 HPL/HL 63.7 57.4-76.4 67.05.7 Snout length/HL 47.4 41.3-50.0 46.42.1 Eye/HL 14.8 14 .8-22.6 19.42.3 Interorbital width/HL 43.6 37.0-44.0 41.02.2 Postorbital length/HL 42.6 37.0-44.5 40.21.9 Mouth length/HL 44.1 32.9-44.1 37.53.7 Maxillary barbel length/HL 103.1 81.4-122.6 107.412.6 XMB/HL 70.5 41.8-75.2 58.89.0 IMB/HL 24.8 20.8-32.3 27.53.6 IMB/XMB 35.2 32 .2-65.6 47.89.6 Dorsal spine length/HL 95.7 85.5-110.3 98.28.8 Pectoral spine length/HL 86.0 84.0-101.8 93.65.9 Eye/Snout length 31.2 31.2-50.2 41.96.2 Interorbital width/Snout length 92.0 81.0-96.0 88.34.1 Postorbital length/Snout length 89.9 80.3 -107.9 86.66.5 Eye/Interorbital width 34.0 34.0-60.6 47.57.3 Eye/Postorbital length 34.7 34.7-61.1 48.46.8 HPL/HPD 321.8 253.8-437.2 333.852.0 CPL/CPD 124.6 93.4-154.5 126.616.4 Meristics Mandibular tooth count 28(1); 32(1); 36(2); 37(1); 42(1); 43(1); 44(2); 45(1); 46*(1); 48(1); 51(1) Dorsal fin count I,7*(13); I,7,i(1); I,8(3) Pectoral fin count I,7(3); I,8(14) Pelvic fin count i,6*(15); i,7(2) Anal fin count iii,8*(2); iii,9(1); iv,6,i(1); iv,7(3); iv,8(2); iv,8,i(1); iv,9,i(1); v,7(1); v,7,i(1); v,8(3); Caudal fin count i,7,8,i(5) Total vertebrae count 36(3), 37(1) Measurement Lectotype Range (n=17) MeanSD

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34 Figure 6. Synodontis granulosus UF 160945, 192 mm TL, 149 mm SL. Top left dorsal view, bottom left lateral view, right ventral view of head.

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35 Figure 7. Distribution of Synodontis granulosus T denotes type locality.

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36 Synodontis multipunctatus Boulenger, 1898 (Figures 2, 3, 8, 9, 10; Tables 1, 4) Synodontis multipunctatus Boulenger, G.A., 1898:24, pl. 8, description, n. sp., Sumba (Lake Tanganyika); 1899:95, note, Lake Tanganyika; 1901:313, pl. 15, description, Sumbu, Kalambo, Moliro, Albertville; 1901:149, Kalambo (Lake Tanganyika); 1906:553, Niamkolo (Lak e Tanganyika);1911:420, fig. 316, description, Sumba, Kalambo, Ni amkolo. – Moore, J.E.S., 1903:166, fig., description, Sumbu (Lake Tanganyika). Worthington, E.B. and Ricardo, C.K., 1933:1067,1077, 1101, note, diet, Usumbura, Kigo ma, Kala (Lake Tanganyika). – Borodin, N.A., 1936:10, Kasanga (Lake Tanga nyika). – David, L. and Poll, M., 1937:267, Tembwe, Moba, Rumonge, Uvira. – Poll, M., 1946:220, description, Lake Tanganyika; 1953:155, pl. 6, fig. 4, de scription, diet, Lake Tanganyika. – Hulot, A., 1950:170, diet, Lake Tanganyi ka. – Lambert, J., 1960:26, figs 26 and 27, note. – Matthes, H., 1962:5, photo, 45, no te, sexual dimorphism, Uvira (Lake Tanganyika). – Coulter, G.W., 1965-1966:34, ta ble, abundance at different depths, Lake Tanganyika; 1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance, ecology, Lake Tanganyika. Brichard, P., 1978:360,361,423, photos, identification, Lake Tanganyika; 1989:476, 478, 481, 483, 474, 479, photos, coloration and identification. – Sands, D., 1983:23, 24, 76, check-list, diet, photo. – Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:134, checklist. – Sato, T., 1986:58-59, brood parasitism Lake Tanganyika. Burgess, W.E., 1989:190, pl. 74, 77, 78, 85, 94, behavior, La ke Tanganyika. – Kobayagowa, M., 1989:14, photo. – Coulter, G.W., 1991a:152, 154, 182, 191, abundance, diet, habitat, reproduction. Diagnosis Axillary pore present; mandibular teeth 13-29 ; body with large spots; fin spines dark; 8 pectoral fin rays; black triangles at ba se of pelvic and anal fins absent or poorly developed; eye 44.9-62.0% snout length; premaxillary toothpad uninterrupted; secondary branches on internal mandibular barbels abse nt; occipito-nuchal shield usually covered with skin; papillae on skin of body absen t; hindgut chamber absent; maximum TL 280 mm. Synodontis multipunctatus can be distinguished from Synodontis sp. 2, S. nigromaculatus S. tanganaicae S. dhonti S. polli and Synodontis sp. 3 by the presence of an axillary pore and the lack of papillae on the skin. Synodontis multipunctatus differs

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37 from S. petricola in having a large axill ary pore (vs. small in S. petricola ), brown to black fin spines (vs. white in S. petricola ), a much larger eye (44.962.0% of snout length in S. multipunctatus vs. 28.7-40.1% in S. petricola ), and an uninterrupted premaxillary toothpad. The number of mandibul ar teeth further separates S. multipunctatus from S. petricola Synodontis sp. 2, S. nigromaculatus S. tanganaicae S. polli and Synodontis sp. 3, as well as from S. granulosus (13-29 in S. multipunctatus vs. 31-50 in S. petricola 38-51 in Synodontis sp 2, 23-38 in S. nigromaculatus 33-49 in S. tanganaicae 40-70 in S. polli 50-66 in Synodontis sp. 3, and 28-51 in S. granulosus ). Synodontis multipunctatus further differs from S. granulosus in having large spots on the body and an occipito-nuchal shield which is covered by skin. Synodontis multipunctatus can be further distinguished from other endemic Tanganyikan species (with the exception of Synodontis sp. 2) by its lack of well developed black triangles at the base of the pelvic and anal fins. Synodontis multipunctatus and Synodontis sp 1 are most reliably separated by pectoral fin ray counts (8 unbranched elements in S. multipunctatus vs. 7 in Synodntis sp 1) and eye size (44.9-62.0% of snout length in S. multipunctatus vs. 64.2-81.0% in Synodontis sp. 1) Description Morphometric and meristic data are give n in Table 4. Maximum total length 280 mm, standard length 220 mm. Body moderately compressed. Predorsa l profile straight. Preanal profile convex. Skin on body formi ng numerous vertical epidermal folds; papillae absent. Head somewhat depressed and broad; skin smooth in all but largest specimens, where ridges and granulations from underlyi ng bone protrude. Snout subconical when viewed laterally; rounded when viewed dorsa lly. Occipital-nuch al shield rugose;

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38 completely covered with skin in smaller specimens (<150 mm), lacking skin in larger specimens; terminating posteriorly with narrow, bluntly pointed process on either side of dorsal spine; ventrally with wide, rounded process which extends to upper margin of humeral process on either side of body, also covered with skin in smaller specimens. Eyes dorsolateral; ovoid; horizontal axis longes t. Interorbital area slightly concave to slightly convex. Mouth subterminal; lips slightly widened, pa pillate. Mandibular teeth 13-29, short, unicuspid; arranged in a single transverse row. Premaxillary toothpad uninterrupted; primary, secondary and tertiary premaxillary teeth discrete, nume rous, arranged in 3, 3, and 2 irregular rows, respectively. Maxillary barbel with or without thin basal membrane; lacking branches or crenelations; extending anywhere fr om base of the pectoral fin to end of humeral process. External mandibular barbel extending to point just beyond posterior margin of pectoral girdle; with 4-6 long, non-tube rculate branches; lacking seco ndary branches. Internal mandibular barbel 1/3 to 1/2 length of extern al barbel; with 3-4 pa irs of non-tuberculate branches; lacking secondary branches. Dorsal fin I,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 spine margin with small serrations distally. Pectoral fin I,8; poste rior margin straight. Pectoral fin spine roughly equal in length to dorsal fin spine, striated, slightly curved, terminating in short, white filament; anterior spine margin smooth in adult specimens; many small, antrorse serrations along anterior margin in juvenile specimens; posterior spine margin with large, retrorse serrations along

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39 entire length. Adipose fin short; poorly de veloped; margin convex. Pelvic fin i,6; located anterior to ver tical 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 distinct ridge on its ventral margin in young specimens; ridge becoming indistinct in adults; dorsal margin concave; terminating in sharp point (Fig. 3B). Large, dark-colored axillary pore pres ent just ventral to hume ral process. Gut 0.50.8 times body length (n = 4, MRAC 53096-53100, MRAC 90273-90277). Hindgut chamber absent. Coloration Dorsum pale yellow to brown, covered w ith large black spots (Fig. 8). Spots larger, irregular, sometimes confluent in ju venile specimens. Belly white, with or without black spots. Maxill ary and mandibular barbels white Iris yellowish to copper colored. Dorsal and pectoral fin spines brown to black, fila ments white. Pectoral spine with thin, light stripe along ante rior margin. Dorsal and pector al fins with black triangles at base, posterior margins wh ite 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. Adipose fin with white dorsal edge. Both lobe s of caudal fin with black bar from base to tip of fin, posterior margin of fin white. Distribution Lake Tanganyika (Fig. 9); common.

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40 Habitat Littoral to benthic zones over shell, sand and mud bottoms; to a maximum depth of 170 m (Coulter, 1991). Diet Synodontis multipunctatus is generally considered to be a specialized predator of Neothauma tanganyicense a common Tanganyikan gastropod (Poll, 1953; Brichard, 1989; Coulter, 1991). The method by which th e snail is extracted from its shell is unknown. Other authors have reported the pr esence of crustaceans, insect larvae and lamellibranches, in addition to Neothauma in the stomach of S. multipunctatus (Worthington and Ricardo, 1936; Poll, 1953, C oulter, 1991), though later accounts of this speciesÂ’ diet fail to mention these prey ite ms. Large numbers of lamellibranch shells were present in the intestine of MRAC 90283. These shells we re packed tightly into the intestine and showed little evidence of m echanical or chemical digestion, while the bodies of the mollusks were mostly intact. It may be that they are ingested incidentally, though the large number of shells found in this particular specimen would seem to indicate otherwise. Further study of the natural feeding habits of S. multipunctatus on both gastropods and bivalves is necessary to more completely understand the diet of this species. Reproduction Synodontis multipunctatus is the only fish species known to exhibit true brood parasitism (Sato, 1986). Eggs are deposited a nd fertilized among those of at least six species of mouth-brooding cichlids (Sato, 1986). These eggs are then taken up by the female cichlid and brooded with the cichlidÂ’s own eggs. The S. multipunctatus hatch before the cichlids and after absorbing their own yolk sac, devour the cichlid eggs and

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41 developing fry. It is unknown whether S. multipunctatus is an obligate br ood parasite, or if alternate spawning behaviors exist. Taxonomic Remarks Synodontis multipunctatus joins S. granulosus as the only Tanganyikan Synodontis for which no other names have been either proposed or mistakenly applied in the literature. Like most Synodontis species in the lake, slight variation in color pattern occurs among localities, but nothing which is co nsistent enough to be able to confidently distinguish between populations Another similarly patterne d, but nevertheless distinct species has been grouped with S. multipunctatus by previous authors. This species is described in the following account.

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42 Table 4. Morphometric measuremen ts and meristic counts for Synodontis multipunctatus All morphometric measurements given in percent of base measurement. Meristic data for Holotype is identifie d by a “*” superscript. SL = Standard length, HL = Head length, XMB = Ex ternal mandibular barbel, IMB = Internal mandibular barbel, HPL = Hu meral process length, HPD = humeral process depth, CPL = Caudal peduncle le ngth, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 28.8 19.6-29.4 24.32.2 Head length/SL 28.6 26.2-33.2 30.01.3 Snout-dorsal length/SL 39.2 35.8-42.7 40.01.3 Adipose fin length/SL 25.2 23.9-34.9 29.22.2 Maxillary barbel length/SL 34.6 21.5-48.6 31.15.1 Dorsal spine length/SL 27.0 22.7-35.3 26.92.4 Pectoral spine length/SL 26.2 21.1-31.3 26.81.8 Head width/HL 88.2 78.5-92.7 84.03.1 Head depth/HL 96.8 69.3-96.7 77.75.3 HPL/HL 59.8 47.8-66.0 57.03.9 Snout length/HL 47.1 41.8-50.8 45.81.6 Eye/HL 22.2 21.6-30.2 25.31.7 Interorbital width/HL 42.7 29.7-42.7 35.02.2 Postorbital length/HL 39.1 29.7-41.7 37.32.1 Mouth length/HL 34.3 24.2-42.3 32.83.5 Maxillary barbel length/HL 120.7 68.3-133.8 102.416.4 XMB/HL 53.6 39.9-78.1 54.48.7 IMB/HL 27.0 18.2-39.0 27.54.5 IMB/XMB 50.4 33.0-67.2 51.06.8 Dorsal spine length/HL 94.2 73.2-111.3 89.69.2 Pectoral spine length/HL 91.4 66.9-106.0 89.37.4 Eye/Snout length 47.1 44.9-62.0 55.43.9 Interorbital width/Snout length 90.7 64.3-90.7 76.64.7 Postorbital length/Snout length 83.0 61.7-94.5 81.76.0 Eye/Interorbital width 51.9 51.9-87.6 72.56.3 Eye/Postorbital length 56.8 55.6-91.0 68.16.9 HPL/HPD 267.7 221.2-414.9 307.636.5 CPL/CPD 130.8 109.8-190.2 142.118.6 Meristics Mandibular tooth count 13(2); 14(4); 15(3); 16(6); 17(4); 18(9); 19*(5); 20(6); 21(6); 22(10); 23(3); 24(5); 26(3); 29(1) Dorsal fin count I,7*(67) Pectoral fin count I,8*(67) Pelvic fin count i,6*(67) Anal fin count iii,6(9); iii, 6,i(2); iii,7(30); iii,7,i(5); iii,8*(10); iv,6 (4); iv,6,i(2); iv,7(2); iv,7,i(1), iv,8(2) Caudal fin count i,7,8,i(4) Total vertebrae count 31(3); 32(1) Measurement Holotype Range (n=67) MeanSD

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43 Figure 8. Synodontis multipunctatus Holotype, BMNH 189 8.9.9.76, 280 mm TL, 220 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view, showing mouth and barbels.

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44 Figure 9. Distribution of Synodontis multipunctatus T denotes type locality.

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45 Synodontis sp. 1 n. sp. (Figures 3, 10, 11; Tables 1, 5) Holotype : BMNH 1982.4.13.4785, TL 129 mm, SL 96 mm, Mwakizega coastline, L. Tanganyika, coll. R. Travers. Paratypes : (1) BMNH 1982.4.13.4786, TL 112 mm, SL 91 mm, collection information same as for holotype. Diagnosis Axillary pore present; mandibular teeth 17-26 ; body with large spots; fin spines brown to black; 7 pectoral fin rays; black patche s at base of pelvic a nd anal fins absent or poorly developed; eye 64.2-81.0% snout length; premaxillary toothpad uninterrupted; secondary branches on internal mandibular ba rbels absent; occipitonuchal shield covered with skin; papillae on skin of body absen t; hindgut chamber absent; maximum TL 150 mm. This species is most similar in appearance to Synodontis multipunctatus and is distinguished from other species of Synodontis by the same traits. It is most easily distinguished from S. multipunctatus by its much larger eye (64.2-81.0% of snout length vs. 44.9-62.0% in S. multipunctatus ). A consistent difference al so exists in pectoral fin ray counts, with Synodontis sp 1 having a count of I,7 vs. I,8 in S. multipunctatus Finally, Synodontis sp 1 reaches a much smaller maximum total length than does S. multipunctatus (150 mm vs. 280 mm). Description Morphometric and meristic data are give n in Table 5. Maximum total length 150 mm, standard length 110 mm. Body compresse d. Predorsal profile convex; interrupted by a ridge formed by the orbit of the eye. Preanal profile convex. Skin on body forming numerous vertical epiderma l folds; papillae absent.

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46 Head depressed and broad; skin smooth. Snout subconical when viewed laterally; rounded when viewed dorsally. Occipital-nuch al shield completely covered with skin; terminating posteriorly with narrow, rounded process on either side of dorsal spine; ventrally with wide, rounded process which extends to upper margin of humeral process on either side of body, also covered with skin in smaller specimens. Eyes dorsolateral; ovoid; horizontal axis longe st. Interorbital area fl at to slightly convex. Mouth subterminal; lips wide and papill ate. Mandibular teeth 17-26, short, unicuspid; arranged in single transverse row. Premaxillary toothpad uninterrupted; primary, secondary and tertiary premaxillary teeth discrete, nume rous, 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. External mandibular barb el extending to point just short of anterior margin of pectoral girdle; with 4-5 non-tuberculate branches; lacking secondary branches. Internal mandibul ar barbel 1/2 to 2/3 length of external barbel; with 4-5 pairs no n-tuberculate branches; lacking secondary branches. Dorsal fin I,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 spine margin with small serrations distall y. Pectoral fin I,7; posterior margin straight. Pectoral fin spin e roughly equal in lengt h to dorsal fin spine, striated, slightly curved, terminating in short, white filament; anterior spine margin with many small, antrorse serrations ; posterior spine margin with large, retrorse serrations along entire length. Adipose fin poorly develope d; margin convex. Pelvic fin i,6; located anterior to vertical through or igin of adipose fin; tip of a ppressed fin barely reaches base

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47 of anal fin. Anal fin iii-iv,6-8; posterior marg in nearly straight; base located ventral to adipose fin. Caudal fin i,7,8,i; forked; lobes pointed. Humeral process narrow, elongated, granul ous; possessing distinct ridge on ventral margin; dorsal margin convex; terminating in a sharp point (Fig. 3C). Large, darkcolored axillary pore present just ventral to humeral process. Gut length 0.8-0.9 times body length (n = 4, MRAC 52096-53100, MRAC 90273-90277). Hindgut chamber absent. Coloration Dorsum pale yellow to brown, covered w ith large black spots (Fig. 10). Spots larger, irregular, sometimes confluent in ju venile specimens. Belly white, with or without black spots. Maxill ary and mandibular barbels white Iris yellowish to copper colored. Dorsal and pectoral fin spines brown to black, fila ments white. Pectoral spine with thin, light stripe along ante rior margin. Dorsal and pector al fins with black triangles at their base, posterior margins white in colo r. Triangles may be completely solid or made up 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 lobe s of caudal fin with black bar from base to tip of fin; posterior margin of fin white. Distribution Lake Tanganyika (Fig. 11); appare ntly common, though less so than S. multipunctatus

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48 Habitat Likely inhabits littoral to benthic zones over shell, sand and mud bottoms, as do most of the other species of the lake. The in creased relative size of the eye and decreased body size may indicate that Synodontis sp. 1 inhabits deeper water than does S. multipunctatus but this cannot be stated with certai nty, due to the lack of depth data in collection records. Diet The lack of fresh material of Synodontis sp 1 makes a thorough examination of this speciesÂ’ diet impossible. It is likely that Synodontis sp. 1 feeds on small gastropods, as do the other species of Synodontis from the lake. Reproduction It is unknown whether Synodontis sp 1 shares the brood parasite behavior of Synodontis multipunctatus In fact, without examining the specimens used in the original study of this behavior (Sato, 1986), it is impossible to say which species, S. multipunctatus or Synodontis sp 1 was used in this study. Unfortunately, no records exist that might indicate the location of th e specimens used. Differences in spawning time are likely to exist, as one mixed lot (MRAC 39186-39192) of S. multipunctatus and Synodontis sp. 1 contains gravid females of the la tter species, while the female specimens of S. multipunctatus do not have mature gonads. Taxonomy As stated in the previous account, no a dditional names have previously been suggested for S. multipunctatus It follows that there are no names available for Synodontis sp 1. A proper specific epithet will be applied to this species upon publication of this study.

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49 Table 5. Morphometric and meristic counts for Synodontis sp. 1. All morphometric measurements given in percent of base measurement. Ranges for S. multipunctatus are provided for comparison. SL = Standard length, HL = Head length, XMB = External mandibular barbel, IMB = Internal mandibular barbel, HPL = Humeral process lengt h, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caud al peduncle depth. Numbers in parentheses indicate number of specimens. Body depth/SL 19.6-29.4 20.2-27.2 23.21.8 Head length/SL 26.2-33.2 27.8-32.4 29.61.0 Snout-dorsal length/SL 35.8-42.7 35.1-43.0 39.21.8 Adipose fin length/SL 23.9-34.9 20.3-33.1 27.93.3 Maxillary barbel length/SL 21.5-48.6 26.3-37.8 30.93.4 Dorsal spine length/SL 22.7-35.3 21.7-31.1 27.02.8 Pectoral spine length/SL 21.1-31.3 22.7-32.4 27.12.8 Head width/HL 78.5-92.7 73.0-92.9 80.64.4 Head depth/HL 69.3-96.7 69.0-83.2 75.33.6 HPL/HL 47.8-66.0 52.0-62.8 55.92.5 Snout length/HL 41.8-50.8 39.4-46.6 42.71.8 Eye/HL 21.6-30.0 27.4-31.9 29.71.4 Interorbital width/HL 29.7-42.7 26.1-37.4 31.92.9 Postorbital length/HL 29.7-41.7 31.4-41.3 35.32.3 Mouth length/HL 24.2-42.3 28.0-37.7 31.92.5 Maxillary barbel length/HL 68.3-159.9 90.2-127.0 104.310.3 XMB/HL 39.9-78.1 45.9-68. 2 55.55.9 IMB/HL 18.2-39.0 20.4-44.2 30.34.6 IMB/XMB 33.0-67.2 44.4-65.5 54.65.7 Dorsal spine length/HL 73.3-111.3 73.2-103.6 91.210.1 Pectoral spine length/HL 66.9-105.9 76.6-108.0 92.09.9 Eye/Snout length 44.9-62.0 64.2-81.0 69.74.6 Interorbital width/Snout length 64.3-90.7 58.4-85.0 74.66.4 Postorbital length/Snout length 61.7-94.5 69.5-100.0 83.07.6 Eye/Interorbital width 51.9-87.6 77.7-120.7 94.211.8 Eye/Postorbital length 55.6-91.0 73.7-100.2 84.57.3 HPL/HPD 221.2-414.9 241.4-431.9 310.838.4 CPL/CPD 109.8-190.2 107.7-189.6 144.418.9 Meristics Mandibular tooth count 17(1); 18(2); 19(2); 20(4); 21(6); 22(3); 23(1); 24(3); 25(1); 26(1); Dorsal fin count I,7(25) Pectoral fin count I,7(21); I,7,i(4) Pelvic fin count i,6(25) Anal fin count 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) Caudal fin count i,7,8,i(3) Total vertebrae count 32(3) Measurement S. multipunctatus Synodontis sp 1 (n=24) MeanSD

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50 Figure 10. Synodontis multipunctatus (top), BMNH 1982.4.13.4789, 115 mm TL, 91 mm SL vs. Synodontis sp 1 (bottom), BMNH 1982.4.13.4784, 105 mm TL, 81 mm SL. Though similarly patterned, the eye of Synodontis sp 1 is larger than that of S. multipunctatus particularly in proportion to the length of the snout.

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51 Figure 11. Distribution of Synodontis sp. 1. T denotes type locality.

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52 Synodontis petricola Matthes, 1959 (Figures 3, 12, 13; Tables 1, 6) Synodontis petricola Matthes, H., 1959:78, description, n. sp., Lake Tanganyika; 1962:34, pl. 2, fig. a, description, diet, Lake Tanganyika. – Poll, M., 1971:409, figs. 192, 193, pls. VI.18, XII .9. – Brichard, P., 1978:360, 424, photos(misidentified), key; 1989:474, 475, 476, 479, 481, 483, skin texture, key, photos(some misidentifications). – Dage t, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:140, check-list. – Burgess, W.E., 1989:196, 556, 559, 560, 567, check-list, photos(some misiden tifications). – Kobayagowa, M., 1989:14, photo(misidentified). – Coulte r, G.W., 1991:181, 182, table 8.16, diet, habitat, reproduction. – Coulter, G. W., 1991a:182, table 8.16, diet, habitat, reproduction. Synodontis multimaculatus non Boulenger, Worthington E.B. and Ricardo, C.K., 1936:1067, 1077, 1101, note. – Poll, M., 1946:223, probable absence from Lake Tanganyika. Diagnosis Axillary pore present; mandibular teeth 31-50 ; body with large spots; fin spines white; 8-9 pectoral fin rays; black triangles present on base of all rayed fins; eye 28.740.1% snout length; premaxillary toothpad interrupted; secondary branches on internal mandibular barbels present; occipi to-nuchal shield c overed 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 Synodontis sp. 2, S. nigromaculatus S. tanganaicae S. dhonti S. polli and Synodontis sp. 3. The occipito-nuchal shield bei ng covered with skin further separates S. petricola from S. granulosus S. nigromaculatus, S. tanganaicae, and S. dhonti all of which also have a much larger maximum TL (27 0, 380, 600, and 400 mm, respectively vs. 135 mm in S. petricola ). Synodontis petricola differs from S. granulosus S. multipunctatus and Synodontis sp. 1 in having an interrupted premax illary toothpad and secondary branches on the internal mandibular barbel s. It is further distinguis hed from all other Tanganyikan

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53 Synodontis with the exception of Synodontis sp. 2, by having completely white fin spines. Synodontis petricola is most easily distinguished from Synodontis sp 2 by the presence of an axillary pore and the lack of light colored windows at the bases of the dark triangles on its fins. Synodontis petricola also somewhat 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 are give n in Table 6. Maximum total length 135 mm, standard length 115 mm. Body not compre ssed. Predorsal profile slightly convex. Preanal profile straight to slightly convex. Skin on body forming numerous vertical epidermal folds; papillae absent. Head slightly depressed and broad; skin covered with villous papillae; papillae extend onto bases of maxillary barbels and anterior portions of body only. Snout with nearly flattened margin when viewed laterall y; bluntly rounded when viewed dorsally. Occipital-nuchal shield cove red with skin, terminating pos teriorly with wide, pointed process on either side of dor sal spine, ventrally with wide, rounded process which extends to upper margin of the humeral proce ss on either side of body. Eyes dorsolateral; ovoid; horizontal axis longe st. Interorbital area fl at to slightly convex. Mouth inferior; lips widened and papilla te. Mandibular teeth 31-50, short, unicuspid; arranged in 6 short, transverse rows. Premaxillary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete, nume rous, arranged in 2, 2, and 1 irregular rows, respectively. Maxillary barbel short; extendi ng at least to base of pector al spine; small papillae at base; basal membrane narrow. External mandi bular barbel extending to point just past

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54 anterior margin of pectoral girdle; with 47 short, simple, weakly tuberculate branches; usually lacking secondary branches. Internal mandibular barbel appr oximately 1/3 to 2/3 length of external barbel; with 4-6 pair s of tuberculate branches; many secondary branches present. Dorsal fin I, 7; posterior marg in straight to slightly concave. Dorsal fin spine short, striated, slightly curved, terminating in a s hort, white filament; anterior margin of fin spine smooth; posterior spine margin with sma ll serrations distally. Pectoral fin I, 8-9; posterior margin broadly rounded. Pectoral fin spine roughly e qual in length to dorsal fin spine, striated, slightly curve d, terminating in short, white filament; anterior spine margin smooth; posterior spine margin with large, retrorse serrations along entire length. Adipose fin long; well develope d; margin convex. Pelvic fi n i,6; located at vertical midway between posterior base of dorsal fin an d origin of adipose fi n; tip of appressed fin does not reach base of anal fin. Anal fi n iii-iv, 7-9; posterior margin rounded; base located at vertical through cen ter of adipose fin. Caudal fin i,7,8,i; forked; l obes rounded. Humeral process triangular; granulous; covered with many small, villous papillae; poorly-developed ridge on ventra l margin; dorsal margin conve x; terminating in a sharp point (Fig. 3D). Axillary pore small; extends posteriorly into body. Gut 1.2-1.3 times body length (n = 2, MRAC A3-033-P-0002-0011) Well developed hindgut chamber. Coloration Dorsum yellowish to cuprous brown, covere d with large, irregul arly shaped, black spots. Spots proportionately larger, sometimes confluent in juvenile specimens. Belly lighter, with small, irregularly shaped spot s. Maxillary and mandibular barbels white. Iris copper colored. Dorsal a nd pectoral fin spines white; terminating in short, white filaments. All rayed fins with black triangles at base; posterior marg ins white in color.

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55 Both lobes of caudal fin with black bars, exte nding from base to tip of fin; posterior margin white. Distribution Lake Tanganyika (Fig. 13). The large am ount of material available in museum collections would seem to indi cate that this species is more common than has been stated in previous accounts (Matth es, 1959; Coulter, 1991). Habitat Rocky coasts within the littoral zone, to a maximum depth of 30 m (Matthes, 1959; Coulter, 1991). Diet Young individuals appear to be primar ily carnivorous, subsisting mainly on hydracarians, ostracods and insect larvae (tri chopterans, chironomids) (Matthes, 1959). Adults feed on algae scraped from rocky s ubstrates, and ingest small invertebrates associated with the biological cover of the rocks in which this species lives (Matthes, 1959, Coulter, 1991). Reproduction No information exists regarding the spaw ning season, behavior, or location of this species. All that is known is that the number of eggs released is likely to be small, as a single female specimen has been found to contain approximately 100 eggs (Matthes, 1959). Taxonomic Remarks Synodontis petricola is sufficiently distinct that th ere has historically been little confusion regarding its taxonomy. Worthington and Ricardo (1936), in their account of the non-cichlid fishes of Lake Tanganyika, id entified a specimen collected by Christy as

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56 Synodontis multimaculatus (BMNH 1936.6.14.1232). Poll (1946) noted that this identification was incorrect, though he did not give a new species identification for the specimen. No list of materials examined wa s given in the original description of S. petricola (Matthes, 1959), but the ty pe designation given to this specimen clearly refers to this name, as a later description by the same author lists this specimen as a paratype (Matthes, 1962). Synodontis petricola shows considerable variation in the nature of its spotted pattern, with the size and spacing of the spot s varying considerably between specimens, even those coming from the same locality. The coloration of the fins varies slightly as well, though it seems to be much more c onsistent than the pattern of the body. Specimens coming from one area (Musende Ro cks, Mpulungu) show such a divergence from the fin coloration of other localities, in addition to the lack of the axillary pore found in other populations, that I consider them to be a di stinct species, which is described in the following account.

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57 Table 6. Morphometric measuremen ts and meristic counts for Synodontis petricola All morphometric measurements given in per cent of base measurement. Meristic data for Holotype is identified by a “*” s uperscript. SL = Standard length, HL = Head length, XMB = External mandibular barbel, IMB = Internal mandibular barbel, HPL = Humeral pro cess length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 21.6 14.3-25.0 20.32.6 Head length/SL 27.9 26.5-30.5 28.51.0 Snout-dorsal length/SL 36.6 32.6-40.2 36.51.6 Adipose fin length/SL 33.9 22.6-44.7 34.34.4 Maxillary barbel length/SL 18.7 12.5-29.8 22.43.6 Dorsal spine length/SL 22.9 17.6-27.0 21.92.7 Pectoral spine length/SL 21.8 18.2-26.6 21.82.0 Head width/HL 84.7 75.6-95.0 83.64.6 Head depth/HL 66.5 53.7-76.9 63.25.4 HPL/HL 39.4 39.4-57.6 46.64.0 Snout length/HL 52.6 49.6-60.9 53.12.3 Eye/HL 16.4 15.1-22.3 17.81.4 Interorbital width/HL 41.9 27.7-47.2 35.04.6 Postorbital length/HL 37.7 31.8-41.2 35.31.9 Mouth length/HL 42.1 37.0-52.9 43.84.1 Maxillary barbel length/HL 67.0 45.2-97.7 78.512.1 XMB/HL 59.4 37.3-72.8 54.98.1 IMB/HL 28.0 16.3-36.0 25.34.2 IMB/XMB 47.2 34.9-68.7 46.77.1 Dorsal spine length/HL 82.1 61.1-98.8 77.69.4 Pectoral spine length/HL 77.9 64.9-97.4 77.17.0 Eye/Snout length 31.2 28.7-40.1 33.62.5 Interorbital width/Snout length 79.5 53.9-83.0 65.87.0 Postorbital length/Snout length 71.6 52.3-75.7 66.64.6 Eye/Interorbital width 39.3 38.4-62.9 51.66.2 Eye/Postorbital length 43.6 41.2-60.9 50.75.2 HPL/HPD 143.2 192.1-333.9 245.935.4 CPL/CPD 84.5 66.3-129.9 92.617.0 MERISTICS Mandibular tooth count 31(1); 33(1); 34(3); 35(1); 36(2); 37(2); 38(5); 39(7); 40(4); 41*(4); 42(5); 43(1); 45(3); 47(1); 50(1) Dorsal fin count I,6(3); I,7*(39) Pectoral fin count I,8*(27); I,9(15) Pelvic fin count i,6*(42) Anal fin count iii,8(2); iii,9(4); iv,7(3); iv,7,i(4); iv,8*(20); iv,8,i(6); iv,9(3) Caudal fin count i,7,8,i(18) Total vertebrae count 31(2); 32(12); 33(4) Measurement Holotype Range (n=42) MeanSD

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58 Figure 12. Synodontis petricola Holotype, MRAC 130357, 98 mm TL, 82 mm SL. Top dorsal view, middle – lateral view, bottom – ventral view.

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59 Figure 13. Distribution of Synodontis petricola T denotes type locality.

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60 Synodontis sp. 2 n. sp. (Figures 4, 14, 15; Tables 2, 7) Holotype : SAIAB 42518, TL 90 mm, SL 74 mm, Country: Zambia, System: Lake Tanganyika; Musende Rocks, 0846’00”S, 03107’00”E, coll. 10.X.1992. Paratypes : (9) SAIAB 39577, TL 58-98 mm, SL 47-79 mm, Zambia; Mpulungu, Musende Rocks, 0846’00”S, 03151’00”E, coll. 07.VII.1992, (3) SAIAB 40171, TL 70-85 mm, SL 56-69 mm, Country: Zambia; Musende Rocks, 0846’00”S, 03106’00”E, coll. 05.VIII.1992, (3) SAIAB 42518, TL 55-97 mm, SL 44-78 mm, collection information same as for holotype. Diagnosis Axillary pore absent; mandibular teeth 38-51; body with large spots; fin spines white; 8-9 pectoral fin rays; black triangles on base of all rayed fins with light colored window at base (except caudal fin); eye 25.235.8% snout length; premaxillary toothpad interrupted; secondary branches on internal ma ndibular barbels pres ent; occipito-nuchal shield covered with skin ; papillae on skin of body absent; maximum TL 100 mm. The lack of an axillary pore distinguishes Synodontis sp 2 from S. granulosus S. multipunctatus Synodontis sp 1, and S. petricola Synodontis sp 2 is further separated from S. petricola by the presence of light colored wi ndows at the bases of the rayed fins (with the exception of the caudal fin). Morphometric m easurements vary little between these two species (Table 7). Synodontis sp. 2 differs from S. nigromaculatus S. tanganaicae S. dhonti S. polli and Synodontis sp. 3 in having white fin spines (vs. brown to black) and in lacking papillae on the skin of the body. The occipito-nuchal shield being covered with skin further separates Synodontis sp. 2 from S. granulosus S. nigromaculatus, S. tanganaicae, and S. dhonti all of which also have a much larger maximum total length (270, 380, 600, and 400 mm, respectively vs. 100 mm in Synodontis sp. 2). The black triangles at the base of all the rayed fins further distinguish

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61 Synodontis sp 2 from S. nigromaculatus while the windows at the base of the triangles further separate Synodontis sp. 2 from S. granulosus S. multipunctatus Synodontis sp. 1, S. tanganaicae, S. dhonti S. polli and Synodontis sp. 3. Description Morphometric and meristic data are give n in Table 7. Maximum total length 100 mm, standard length 80 mm. Body not compre ssed. Predorsal profile slightly convex. Preanal profile slightly convex. Skin on body forming numerous vertical epidermal folds; papillae absent. Head slightly depressed and broad; skin covered with villous papillae; papillae extend onto bases of maxillary barbels, pector al fins and anterior portions of body only. Snout with bluntly rounded margin when view ed laterally and dorsa lly. Occipital-nuchal shield covered with skin, terminating posteri orly with wide, pointed process on either side of dorsal spine, ventrally with wide, rounded process which extends to upper margin of the humeral process on either side of body. Eyes dorsolateral; ovoid; horizontal axis longest. Interorbital area flat to slightly convex. Mouth inferior; lips widened and papilla te. Mandibular teeth 35-51, short, unicuspid; arranged in 6 short, transverse rows. Premaxillary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete, nume rous, arranged in 3, 3, and 1 irregular rows, respectively. Maxillary barbel short; extendi ng at least to base of pector al spine; small papillae at base; basal membrane narrow. External mandi bular barbel extending to point just past anterior margin of pectoral girdle; with 46 short, simple, weakly tuberculate branches; usually lacking secondary branches. Internal mandibular barbel appr oximately 1/3 to 1/2

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62 length of external barbel; with 4-6 pair s of tuberculate branches; many secondary branches present. Dorsal fin I, 7-7,i; posterior margin straight to slightly concave. Dorsal fin spine short, striated, slightly curve d, terminating in a short, white filament; anterior margin of fin spine smooth; posterior spine margin with sm all serrations distally. Pectoral fin I, 88,i; posterior margin broadly rounded. Pect oral fin spine roughl y equal in length to dorsal fin spine, striated, slightly curved te rminating in short, white filament; anterior spine margin smooth; posterior spine margin with large, retrorse serrations al ong entire length. Adipose fin long; well developed; ma rgin convex. Pelvic fin i,6; located at vertical midway between posterior base of dor sal 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 ventra l margin; dorsal margin conve x; terminating in a sharp point (Fig. 4A). Axillary pore absent. Coloration Dorsum yellowish to cuprous brown, covere d with large, irregul arly shaped, black spots (Fig. 14). Belly lighter, with smaller, more regularly shaped spots. Maxillary and mandibular barbels white. Iris copper colored. Dorsal and pectoral fin spines white, terminating in short, white filaments. A ll rayed fins with black triangles at base; posterior margins white. Triangles have la rge, lightly colored windows at bases, most noticeable in dorsal and anal fins. Both lobe s of caudal fin with black bar from base to tip of fin; posterior margin of fin dusky.

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63 Distribution Lake Tanganyika (Fig. 15). 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 sp. 2. Taxonomic Remarks Specimens of this species have prev iously been identified either as Synodontis sp. or S. petricola While this species resembles S. petricola in most respects, it is clearly diagnosable from S. petricola As there are no names available to be applied to this species, an appropriate specifi c epithet will be applied upon publication of this study.

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64 Table 7. Morphometric and meristic counts for Synodontis sp. 2. All morphometric measurements given in percent of base measurement. Ranges for S. petricola are provided for comparison. SL = St andard length, HL = Head length, XMB = External mandibular barbel, IMB = Internal mandibular barbel, HPL = Humeral process length, HPD = hu meral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indicate number of specimens. Body depth/SL 14.3-25.0 15.7-23.9 20.02.7 Head length/SL 26.5-30.5 27.6-31.2 29.81.0 Snout-dorsal length/SL 32.6-40.2 35.7-40.3 38.11.4 Adipose fin length/SL 22.6-44.7 27.6-38.7 32.62.9 Maxillary barbel length/SL 12.5-29.8 20.7-27.5 23.01.9 Dorsal spine length/SL 17.6-27.0 17.6-22.8 19.61.4 Pectoral spine length/SL 18.2-26.6 17.2-24.2 21.31.8 Head width/HL 75.6-95.0 76.0-86.1 82.92.7 Head depth/HL 53.7-76.9 55.0-64.5 59.62.5 HPL/HL 39.4-57.6 41.5-50.1 46.02.5 Snout length/HL 49.6-60.9 51.4-55.0 53.01.1 Eye/HL 15.1-22.3 13.9-18.5 16.61.3 Interorbital width/HL 27.7-47.2 28.5-37.1 32.02.3 Postorbital length/HL 31.8-41.2 33.6-38.6 35.91.3 Mouth length/HL 37.0-52.9 41.5-57.2 48.83.9 Maxillary barbel length/HL 45.2-97.7 68.5-99.5 77.47.7 XMB/HL 30.9-67.1 42.6-60.4 49.84.7 IMB/HL 16.3-36.0 19.3-28.3 24.12.4 IMB/XMB 34.9-68.7 42.4-61.8 48.65.3 Dorsal spine length/HL 61.1-98.8 59.2-76.6 66.24.6 Pectoral spine length/HL 64.9-97.4 62.0-79.8 71.44.9 Eye/Snout length 28.7-40.1 25.2-35.8 31.42.7 Interorbital width/Snout length 53.9-83.0 53.8-71.9 60.54.8 Postorbital length/Snout length 52.3-75.7 61.5-75.1 67.93.4 Eye/Interorbital width 38.4-62.9 43.9-58.5 52.04.6 Eye/Postorbital length 41.2-60.9 39.5-54.4 46.23.8 HPL/HPD 192.1-333.9 213.7-319.1 262.824.3 CPL/CPD 66.3-129.9 87.1-141.4 104.415.7 Meristics Mandibular tooth count 35(2); 36(2); 38(4); 39(1); 41(2); 43(1); 44(1); 47(2); 51(1) Dorsal fin count I,7(15); I,7,i(1) Pectoral fin count I,8(14); I,8,i(2) Pelvic fin count i,6(16) Anal fin count iv,8(11); iv,8,i(2); iv, 9(3) Caudal fin count i,7,8,i(10) Total vertebrae count 32(3); 33(6) Measurement S. petricola Synodontis sp 2 (n=16) MeanSD

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65 Figure 14. Synodontis sp 2, SAIAB 39577, 84 mm TL, 68 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view.

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66 Figure 15. Distribution of Synodontis sp 2. T denotes type locality.

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67 Synodontis nigromaculatus Boulenger, 1905 (Figures 4, 16, 17, 18; Tables 2, 8) Synodontis nigromaculatus Boulenger, G.A., 1905:645, description, n. sp., Lake Bangwelu; 1911:416, fig. 313, de scription, Lake Bangwelu. – Worthington, E.B., 1933:50, note, Lake Ba ngwelu. – Ricardo-Bertram, C.K., 1940:187, 209, note, diet, Chambezi R., Bangwelu marsh, Lake Bangwelu, Congo system, Zambezi system, comparison with S. melanostictus considered synonymous. – Bowmaker, A.P., 1961:6, f ecundity. – Jackson, P.B.N., 1961:20, 87, key, note, N. Rhodesia. – Jubb, R.A., 1961:43, 117, fig. 61, description, fishing, Limpopo system, Upper Zambezi; 1963:7, 12, 15, 28, note, Upper Zambezi (S. Afr.), = S. melanostictus ; 1967,:43, 52, 56, fig. 155, description, key, distribution table, Upper Zambezi, Okovango system, Cunene R. (S. Afr.), = S. melanostictus – De Kimpe, P., 1964:77, Luapula-Mweru. – Ladiges, W., 1964:247, 254, 256, 257, 263, 267, diet, ford near Mulundo, Cuchi R., Cuebe R., Cubango R., Okovango R.Gabie, V., 1965: 386, maps 1 and 2, distribution in S. Africa (Limpopo). – Bell-Cross, G ., 1955-1966a and b:13, 16, 18, 27, Upper Zambezi, Okovango, affinities with the fa una of the Congo system (LualabaCongo). – Poll, M., 1967:247, fig. 115, desc ription, Lake Calundo (Angola); Poll, M., 1971:325, fig. 148 and 149, pl. V,11 and XI,9, description, synonymy with S. melanostictus – Sands, D., 1983:23, check-list. – Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:136, check-list. – Brichard, P., 1989:479, key, 478, 480, 483, photos. – Burgess, W., 1989: 196, check-list, pl. 78, 82, 83, 94. – Kobayagawa, M., 1989:15, photo. – Coulter, G.W., 1991:152, 154, 156, 181, 191, 287, abundance, habitat, reproduction. – Coulter, G.W., 1991a:152, 154, 156, 182, 191, abundance, habitat, reproduction. Synodontis melanostictus Boulenger, G.A., 1906:553, pl 34, description, n. sp., Lofu (Lake Tangamnyika); 1911:418, fig. 314, de scription, Lofu (Lake Tanganyika), Lake Bangwelu, Lake Mweru, Maramba, Upper Zambezi; 1916:316, note. – Pappenheim, P. and Boulenger, G. A., 1914:251, note. – Poll, M., 1933:134, Pweto, Lukonzolwa, Lake Mweru, Ka tanga; 1946:220, description, Lake Tanganyika (citation); 1948:8, Pweto, La ke Mweru (Katanga). – Worthington, E.B., 1933:50, note, Lake Bangwelu. – Fowler, H.W., 1935:273, description, Chobe R 3 miles from Kasane (Kalahar i). – Worthington, E.B. and Ricardo, C.K., 1933:1067, 1077, 1101, note, Lake Tanganyika, Congo system, Zambezi system, including Lake Malawi. – Pellegr in, J., 1936:58, Humbe, Cunene, Cubango (Angola). – Ricardo-Bertram, C.K., 1940: 209, first placed into synonymy of S. nigromaculatus – Barnard, K.H., 1948:443, note, Lealui, Lake Ngami, Victoria Falls, Seheki, Okovango R., Cunene R. – Hulot, A., 1950:170, Lake Mweru. – Jubb, R.A., 1954:692, 697, S. Rhodesia; 1958:181, table 1, Chobe R., Kobuta, Zambezi, R., Sananga. – Soulsby, J.J., 1960:323, note, diet, N. Rhodesia. – Matthes, H., 1962:41, table 2, comparison of specimens from Lake Tanganyika and Mweru-Luapula, comparison with S. serratus tanganaicae and S. lacustricolus – Sterba, G., 1963:408, fi g. 610, Zambezi, Lake Tanganyika, Lake Bangwelu, Lake Mweru. – Ladiges, W., 1964:267, Cunene,

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68 Cubango, Okovango, Zambezi. – De Kimpe, P., 1964:77, 89, 114, 116, 137, 138, 147, 185, 188, 195, 201, 222, figs 46, 72, diet, Luapula-Mweru. – Coulter, G.W., 1965-1966:34, abundance at different dept hs, Lake Tanganyika. – Brichard, 1978:424, key. Synodontis colyeri Boulenger, G.A., 1923:438, desc ription, n. sp., Mansa R. (N. Rhodesia). – Jackson, P.B.N., 1961:20, 88, note, key, N. Rhodesia. – De Kimpe, P., 1964:77, 89, Mansa R. (LuapulaMo ero). – Bell-Cross, G., 1965-1966a:13, 18, Zambia, Congo. – Poll, 1971:329, placed into synonymy with S. nigromaculatus Synodontis zambesensis non Peters, David, L. and Poll, M., 1937:234. – Poll, M., 1967:243, fig. 113, Angola, tributary of the Kasai. Diagnosis Axillary pore absent; mandibular teeth 23-38; body with small spots; fin spines brown; 8-10 pectoral fin ra ys; black triangles absent on all rayed fins; eye 31.2-48.1% of snout length; premaxillary toothpad interrupted; secondary branches on internal mandibular barbels absent; occipito-nuchal sh ield not covered by skin; villous papillae present on skin of body; hindgut ch amber present; maximum TL 380 mm; The only non-endemic Synodontis species found in Lake Tanganyika, S. nigromaculatus lacks the rayed fin coloration common to all of its Tanganyikan congeners, having spotted fins which completely 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. nigromaculatus In these cases, mandibular tooth counts (23-38 in S. nigromaculatus vs. 33-49 in S. tanganaicae ), differences in eye size (31.2-48.1% of snout length in S. nigromaculatus vs. 16.0-26.9% in S. tanganaicae ), the villous papillae on the skin of S. nigromaculatus (vs. granular in S. tanganaicae ), and differences in hume ral process shape (Fig. 4) distinguish these species from one another.

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69 Description Morphometric and meristic data are give n in Table 8. Maximum total length 380 mm, maximum standard length 305 mm. Body compressed. Predorsal profile straight anterior to eye; slightly c onvex posterior to eye. Preanal profile straight. Skin on body lacking vertical epidermal 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, blun tly pointed margin when viewed dorsally. Occipital-nuchal shield rugos e, not covered with skin; te rminating posteriorly with narrow, rounded process on either side of dorsal spine; ventrally with wide, rounded process which extends to upper margin of hume ral process on either side of body. Eyes dorsolateral, ovoid, horizontal ax is longest. Interorbital ar ea flat to slightly convex. Mouth subterminal; lips wide and papill ate. Mandibular teeth 23-38, short, unicuspid; arranged in single transverse row. Premaxillary toothpad interrupted; primary, secondary, and tertiary premax illary teeth discrete; primar y premaxillary teeth in 2-3 rows; secondary premaxillary teeth in 2-3 rows ; tertiary premaxillary teeth in 1-2 rows. Maxillary barbel with narrow basal membra ne; lacking branches or crenelations; extending at least to the base of the pectoral fin. Extern al mandibular barbel extending to point just short of anterior margin of p ectoral girdle; with 7-8 long, non-tuberculate branches; short secondary branch es present. Internal mandi bular barbels about half the length of the external barbels; with 4-5 pa irs of non-tuberculate branches; secondary branches present. Dorsal fin I,7; posterior margin concave. Dorsal fin spine lon g, striated, slightly curved, terminating in short bl ack filament; anterior margin of fin spine smooth; posterior spine margin with small serrations distally. Pe ctoral fin I,8-10; posteri or margin straight.

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70 Pectoral fin spine roughly equa l in length to dorsal fin spine, striated, slightly curved, terminating in a moderate length filament; an terior spine margin with many fine, long, antrorse serrations along anterior margin; pos terior spine margin with large, retrorse serrations along entire length. Adipose fin we ll developed; margin convex. Pelvic fin i,6; located anterior to verti cal through posterior base of dor sal fin; tip of appressed fin barely reaches base of anal fin. Anal fin ii i-v,7-9; posterior margin nearly straight; base located ventral to first 1/3 of ad ipose fin. Caudal fin forked, i,7,8,i. Humeral process triangular; wide at base; tapering to sharp point; striated along lower half; becoming granulous along upper margin; possessing weakly developed ridge along ventral margin; dorsal margin slightly co ncave (Fig 5B). Axillary pore absent. Gut 1.3-1.6 times body length (n=2, MRAC 100515-100518). Well developed hindgut chamber. Coloration Body grayish to cuprous brown (Fig. 16). Belly pale. Scattered, small black spots present, sometimes confluent. Maxillary and mandibular barbels white. Iris copper colored. Dorsal and pectoral fin spines brown, terminating in dusky filaments. All rayed fins with small, black spot s, similar to those found on body. Distribution Lake Tanganyika, Lake Mweru, and Lake Bangwelu basins; Upper Kasai, Upper Zambezi, Okovango, Cunene, and Limpopo Ri vers; Luapula swamps (Fig. 18). Habitat Littoral to benthic zones over shell, sand and mud bottoms, to a maximum depth of 150 m (Coulter, 1991).

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71 Diet Nothing is known about the diet of this species within Lake Tanganyika. Reproduction No concrete information ex ists on the reproduction of S. nigromaculatus Coulter (1991) cites Matthes (1959) as reporting that gravid females of this species were found to contain up to 3,000 eggs. However, upon review ing this second reference, no mention of the reproduction of S. nigromaculatus is found Coulter (1991) states that it is likely that S. nigromaculatus spawns in rivers within the lake basin, as this species is known to spawn in rivers throughout its range. CoulterÂ’s assessment appears to be accurate, as only large adult specimens of S. nigromaculatus have thus far been documented within the lake. Taxonomic Remarks Synodontis melanostictus Boulenger, originally desc ribed from Lake Tanganyika, was distinguished from S. nigromaculatus Boulenger by the presence of numerous, villous papillae on the body (Boulenger, 1906) Ricardo-Bertram ( 1940) first suggested placing S. melanostictus in the synonymy of S. nigromaculatus stating that the presence or absence of these papillae depended upon the condition and preservation of a particular specimen. This is undoubtedly true to some extent, as the holotype of S. melanostictus lacks any indication of papillae on its body. However, the holotype of S. nigromaculatus is in excellent condition, yet it still lacks any papillae of the sort found on S. melanostictus as do any specimens agreeing in appearance with the holotype of S. nigromaculatus that come from outside th e Lake Tanganyika drainage. Further distinguishing Synodontis melanostictus from S. nigromaculatus is the shape of the snout when viewed dorsally, as well as the condition of the occipito-nuchal

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72 shield, which is exposed and rugose in S. melanostictus but covered with skin in S. nigromacualtus (Fig. 17). Though the occipito-nucha l shield does become more exposed with growth in many Synodontis species, these comparisons have been made between specimens of similar size and appear to be consistent. Wh ile it appears that S. melanostictus is sufficiently distinct from S. nigromaculatus to be considered a valid species, some material at the Royal Museum for Central Africa has yet to be examined. This material must be studied before any taxonomic changes can be proposed. Synodontis colyeri Boulenger was described from th e Mansa River in what is now known as Zimbabwe (Boulenger, 1926). Poll (1971) considered S. colyeri to be a synonym of S. nigromaculatus a decision which is accepted here, due to the fact that examination of the holotype of S. colyeri has not yet been possi ble, and no photographs or drawings exist which might provide clues as to the correc t placement of this specimen. Synodontis zambesensis Peters is a widespread sp ecies known to occur in the Pongolo, Limpopo, Pungwe, Zambezi, Lake Rukw a, and Lake Mweru drainages (Poll, 1971). Several specimens coming from the Luachimo and nearby rivers (Congo drainage) in Angola have previously been de termined to belong to this species (David and Poll, 1937; Poll, 1967) (Fig. 16) However, adult specimens of S. zambesensis lack spots, having a uniform, light brown colora tion, while the specimens from the Luachimo clearly have large, widely-spaced, black s pots. Poll (1971) obvi ously realized this mistake, as his revision of Synodontis lists these specimens as S. nigromaculatus Though these specimens are not S. zambesensis they also clearly are not specimens of S. nigromaculatus nor are they S. melanostictus These specimens have large, widely spaced spots, a more depressed head than either S. nigromaculatus or S. melanostictus a

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73 narrow preopercle (vs. broad), and lack any cutaneous papillae. These specimens represent an undescribed species, which will be described in a future paper that will deal exclusively with th e taxonomy of the S. nigromaculatus species complex.

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74 Table 8. Morphometric and meristic counts for Synodontis nigromaculatus All morphometric measurements given in per cent of base measurement. Meristic data for Holotype is identified by a “*” supe rscript. SL = Standard length, HL = Head length, XMB = External mandibular barbel, IMB = Internal mandibular barbel, HPL = Humeral pro cess length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 24.9 16.7-26.2 22.02.1 Head length/SL 25.9 24.9-31.4 27.51.4 Snout-dorsal length/SL 37.8 34.3-41.1 38.11.6 Adipose fin length/SL 32.9 16.1-41.8 31.23.6 Maxillary barbel length/SL 33.8 22.2-48.0 34.25.9 Dorsal spine length/SL 30.3 20.5-36.0 26.92.8 Pectoral spine length/SL 25.0 21.5-30.9 26.12.0 Head width/HL 90.6 81.7-99.2 92.43.6 Head depth/HL 86.8 63.4-91.7 73.76.6 Humeral process length/HL 62.7 51.5-76.1 64.25.3 Snout length/HL 46.5 44.5-53.7 49.02.2 Eye/HL 17.2 15.4-22.0 18.21.7 Interorbital width/HL 42.1 32.5-54.2 38.24.1 Postorbital length/HL 43.5 33.9-44.0 37.82.5 Mouth length/HL 37.9 24.9-44.0 34.64.4 Maxillary barbel length/HL 130.5 77.4-172.7 124.923.0 XMB/HL 64.5 51.2-107.6 81.516.4 IMB/HL 39.0 26.6-61.3 41.98.8 IMB/XMB 60.4 36.3-66.8 51.15.6 Dorsal spine length/HL 117.0 84.1-127.2 98.89.3 Pectoral spine length/HL 96.3 80.0-110.8 95.67.2 Eye/Snout length 37.1 31.1-48.1 37.34.4 Interorbital width/Snout length 90.5 66.8-96.8 77.36.6 Postorbital length/Snout length 93.6 64.8-93.6 77.47.2 Eye/Interorbital width 41.0 32.9-65.2 48.48.2 Eye/Postorbital length 39.6 37.7-63.2 48.46.4 HPL/HPD 219.1 170.9-318.2 245.935.4 CPL/CPD 91.7 66.9-121.8 94.812.2 Meristics Mandibular tooth count 23(1); 25(1); 26(3); 27(5); 28(3); 29*(5); 30(1); 31(3); 32(6); 33(1); 34(1); 35(2); 36(3); 37(1); 38(2) Dorsal fin count I,7*(39) Pectoral fin count I,8*(15); I,8,i(1); I,9(20), I,9,i(1); I,10(1) Pelvic fin count i,6*(39) Anal fin count iii,8(2); iii, 8,i(2); iii,9(4); iv,7(1); iv,7,i(4); iv,8 (8); iv,8,i(8); iv,9*(5); v,7(1), v,8(2), v,8,i(1), v,9(1) Caudal fin count i,7,8,i(1) Total vertebrae count 36(1) Measurement Holotype Range (n=34) MeanSD

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75 Figure 16. Variation in body shape and coloration in Synodontis nigromaculatus Top – S. melanostictus Holotype, BMNH 1906.9.8.72, 290 mm TL, 237 mm SL. Middle – S. nigromaculatus Holotype, BMNH 1905.11.10.10, 237 mm TL, 186 mm SL. Bottom – Synodontis sp. (Orig. I.D. = S. zambesensis ), MRAC 162203-204, 180 mm TL, 132 mm SL.

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76 Figure 17. Dorsal views of the head of left – Synodontis melanostictus (Holotype) and right – Synodontis nigromaculatus (Holotype), showing differences in occipito-nuchal shield texture.

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77 Figure 18. Distribution of Synodontis nigromaculatus T denotes type locality.

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78 Synodontis tanganaicae Borodin, 1936 (Figures 4, 19, 20, 21; Tables 2, 9) Synodontis serratus tanganaicae Borodin, N.A., 1936:9, description, ssp. of S. serratus Rppel, Kasenga, Lake Tanganyika. Synodontis lacustricolus Poll, M., 1953:157, fig. 18a, pl. 6, fig. 3, description, n. sp., diet, Lake Tanganyika; 1971:402, fig. 188, 189, pl 6, 12. – Matthes, H., 1962:41, pl. 3, fig. a, description. – Coulter, G.W., 1 965-1966:34, abundance at different depths, Lake Tanganyika; 1991:152, 154, 181, 182, table 8.2, 8.3, 8.16, abundance, habitat, diet. – Brichard, P., 1978: 424, key; 1989:479, key. – Sands, D., 1983, 23, check-list. – Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:131, check-list. – Burgess, W.E., 1989:196, check-list. – De Vos, L. and Thys van den Audenaerde, D., 1998:147, synonymy with S. tanganaicae – Coulter, G.W., 1991a:152, 154, 182, 191, a bundance, habitat, reproduction. Synodontis tanganyicae De Vos, L. and Thys van den Audenaerde, D., 1998:147. Diagnosis Axillary pore absent; mandibular teeth 33-49; body with small or no spots; fin spines dark; 8-9 pectoral fin ra ys; black triangles on base of all rayed fins in juveniles, absent in adults; eye 16.0-26.9% snout le ngth; premaxillary toothpad interrupted; secondary branches on internal mandibular ba rbels present; occipi to-nuchal shield not covered with skin; granular papillae present on skin of body; maximum TL 585 mm. The lack of an axillary pore distinguishes Synodontis tanganaicae from S. granulosus S. multipunctatus Synodontis sp. 1, and S. petricola Synodontis tanganaicae differs further from S. granulosus in humeral process shape (humeral process length 172.9-255.4% of humeral process width in S. tanganaicae vs. 253.8437.2% 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. multipunctatus Synodontis sp. 1 S. petricola Synodontis sp. 2 S. polli and Synodontis sp. 3. Adult specimens of S.

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79 tanganaicae are further distinguished from the other Tanganyikan endemics (with the exception of adult S. dhonti ) by the lack of black triangles on the rayed fins. Synodontis tanganaicae most closely resembles S. nigromaculatus and adult specimens of S. dhonti Synodontis tanganaicae differs from S. nigromaculatus in the number of mandibular teeth (33-49 in S. tanganaicae vs. 23-38 in S. nigromaculatus ), eye size (16.0-26.9% of snout length in S. tanganaicae vs. 31.2-48.1% in S. nigromaculatus ), the granular papillae on the skin (vs. villous in S. nigromaculatus ), and differences in humeral process shape (Fig. 4). The only potentially reli able character that differentiates S. tanganaicae and adult specimens of S. dhonti is the gill raker count on the first branchial arch (27-28 in S. tanganaicae vs. 11-20 in S. dhonti ) (De Vos and Thys van den Audenaerde, 1998). Description Morphometric and meristic data are give n in Table 9. Maximum total length 585 mm, standard length 490 mm. Body not compre ssed. Predorsal profile straight. Preanal profile convex. Skin on body forming numerous vertical epidermal 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 rounde d margin when viewed dorsally. Occipitalnuchal shield rugose, not covered with skin; terminating posteriorly with narrow, bluntly ponted process on either side of dorsal spine; ventrally with wide, rounded process which extends to upper margin of humeral process on either side of body. Eyes dorsolateral; ovoid; horizontal axis longest Interorbital area flat.

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80 Mouth inferior; lips wide and papillate. Mandibular teeth 33-49, short, unicuspid; arranged in single transverse row. Premax illary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete; numerous; arranged in irregular rows. Maxillary barbel with well developed basal membrane; lacking branches or crenelations; extending at leas t to base of pectoral fin. External mandibular barbel extending to point just short of anterior margin of pectoral girdle, with 4-7 nontuberculate branches; lacking secondary branches. Internal mandibular barbel about 1/3 length of external barbel; with 4-6 pairs of non-tuberculate branches ; secondary branches present. Dorsal fin I,7-8; posterior margin straight Dorsal fin spine long, granulous, nearly straight, terminating in short, dark filamen t; anterior margin of fin spine granulous; posterior spine margin with small serrations di stally. Pectoral fin I,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; anteri or spine margin granulate; posterior spine margin with large retrorse se rrations along entire leng th. Adipose fin well developed; margin convex. Pelvic fin i,6; lo cated at vertical thr ough posterior base of dorsal fin; tip of appressed fin does not reac h base of anal fin. Anal fin iii-v,7-9; posterior margin broadly rounded; base locate d 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; terminating in blunt point (Figs. 4C, 4D). Axillary pore absent. Coloration Dorsum gray to reddish brown (Figs. 19, 20) Belly lighter in color. Scattered, small black spots present on entire body. Ma xillary and mandibular barbels white, bases

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81 sometimes with scattered dusky pigmentation. Iris copper colored. Dorsal and pectoral fin spines dark, terminating in short, dusky f ilaments. All rayed fins with small, dark spots similar to those on body. Juvenile specime ns with black triangles at base of rayed fins. Distribution Lake Tanganyika (Fi g. 21); fairly common. Habitat Littoral to benthic zones over shell, sand and mud bottoms, to a maximum depth of 130 m (Coulter, 1991). Diet Gastropods, lamellibranchs, insect larvae, ostracods, and shrimps (Poll, 1953; Coulter, 1991). Reproduction No information exists regarding the reproduction of S. tanganaicae Taxonomic Remarks Synodontis serratus tanganaicae Borodin has recently be en elevated to species status, having previously been considered a subspecies of Synodontis serratus Rppell (De Vos and Thys van den Audenaerde, 1998). De Vos and Thys van den Audenaerde also placed Synodontis lacustricolus Poll in the synonymy of S. tanganaicae This decision is accepted here, however, ne ither of the type specimens of S. tanganaicae have been examined. De Vos and Thys van den Audenaerde note a strong similarity between S. tanganaicae and S. dhonti Boulenger. This similarity is apparent when examining specimens of S. dhonti and photographs of the Lectotype of S. tanganaicae (MCZ

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82 32538). Differences in mandibular tooth number, adipose fin shape, and gill raker count are cited as the characters in which S. dhonti differs from S. tanganaicae (De Vos and Thys van den Audenaerde, 1998). However, there is signifi cant overlap in the mandibular tooth count of both species (20-38 in S. dhonti and 33-40 in S. tanganaicae 33-49 when nominal specimens of S. lacustricolus are included). While the degree of overlap between the first and last value is not large, S. dhonti is clearly distinct from nominal specimens of S. lacustricolus The significance of overlap is much greater between S. dhonti and specimens identified as S. tanganaicae Additionally, minor variation in adipose fin morphology is common in the Synodontis species of Lake Tanganyika. Gill rakers have not been counted for the specimens of either species that have been examined in this study due to th e necessarily destructive nature of this procedure, therefore, the utility of this ch aracter can be neither confirmed nor refuted by this study. Further study is clear ly needed to determine whether S. tanganaicae and S. dhonti truly represent different species, as well as to evaluate the validity of De Vos and Thys van den AudenaerdeÂ’s conc lusion regarding the synonymy of S. tanganaicae and S. lacustricolus

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83 Table 9. Morphometric and meristic counts for Synodontis tanganaicae All morphometric measurements given in per cent of base measurement. Meristic data for Lectotype is identified by a “* ” superscript. All morphometric and meristic values for the Lectotype and Paralectotype are reproduced from De Vos & Thys van den Audenaerde, 1998. SL = Standard length, HL = Head length, XMB = External mandibular ba rbel, IMB = Internal mandibular barbel, HPL = Humeral process lengt h, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caud al peduncle depth. Numbers in parentheses indicate number of specimens. Body depth/SL 28.4, 32.2 20.8-32.2 25.42.9 Head length/SL 35.7, 34.7 31.5-35.7 33.01.3 Snout-dorsal length/SL 49.5, 48.8 41.2-49.5 43.72.7 Adipose fin length/SL 28.1, 32.2 23.0-32.2 26.22.8 Maxillary barbel length/SL 29.8-43.7 37.25.0 Dorsal spine length/SL 22.6, 22.0 21.1-27.6 23.82.4 Pectoral spine length/SL 23.8, 25 .7 19.0-26.7 23.02.3 Head width/HL 85.9, 85.6 79.4-94.1 87.83.8 Head depth/HL 67.4, 69.2 66.1-84.1 72.94.8 Humeral process length/HL 38.1-55.2 47.15.4 Snout length/HL 58.2, 51.6 51.1-60.6 55.12.6 Eye/HL 13.1, 13.4 9.7-14.6 13.01.4 Interorbital width/HL 39.4, 39.2 35.0-42.1 39.12.2 Postorbital length/HL 39.2, 38.5 35.1-40.4 38.11.7 Mouth length/HL 52.0, 50.4 30.1-52.0 43.85.4 Maxillary barbel length/HL 112.9, 127.7 90.7-138.5 116.115.7 XMB/HL 47.3, 60.4 47.3-66.4 53.75.4 IMB/HL 17.5, 22.7 14.7-23.5 20.02.8 IMB/XMB 29.2-43.4 37.44.7 Dorsal spine length/HL 63.6-87.4 74.58.3 Pectoral spine length/HL 57.9-83.2 69.57.1 Eye/Snout length 16.0-26.9 23.53.2 Interorbital width/Snout length 60.3-81.0 71.06.4 Postorbital length/Snout length 59.8-78.5 69.05.4 Eye/Interorbital width 26.6-38.0 33.13.3 Eye/Postorbital length 26.8-39.6 34.03.8 HPL/HPD 172.9-255.4 212.224.4 CPL/CPD 98.6-149.8 129.614.4 Meristics Mandibular tooth count 33(1); 38(2); 39(1); 40*(1); 42(1); 43(2); 44(1); 46(1); 47(2); 48(1); 49(1) Dorsal fin count I,7*(12); I,7,i(1); I,8(1) Pectoral fin count I,8*(5); I,9(9) Pelvic fin count i,6*(12); I,7(1) Anal fin count iii,8*(2); iv,7(1); iv,8(6); iv,8,i(1); iv,9(1); v,7(1); v,8,i(1) Measurement Lecto& Paralectotype Range (n=14) MeanSD

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84 Figure 19. Synodontis tanganaicae Lectotype, MCZ 32538, 415 mm TL, 320 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view of head. Photographs courtesy of Museum of Comparative Zoology, Harvard University.

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85 Figure 20. Synodontis tanganaicae MRAC 90288, paratype of S. lacustricolus 519 mm TL, 434 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view of head.

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86 Figure 21. Distribution of Synodontis tanganaicae Type locality unknown.

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87 Synodontis dhonti Boulenger, 1917 (Figures 5, 22, 23, 24; Tables 2, 10) Synodontis dhonti Boulenger, G.A., 1917:367, descrip tion, n. sp., Kilewa Bay, Lake Tanganyika; 1920:42, description, Kilewa Bay, Lake Tanganyika. – Worthington, E.B. and Ricardo, C.K., 1936:1067, 1077, 1101, note, Lake Tanganyika. – Poll, M., 1946:221, figs. 21, 22, description, K ilewa Bay, Lake Tanganyika; 1971:364, figs. 168 and 169, pls. VI.6 and XI.19. – Matthes, H., 1962:37, fig. a, pl. 3, description, diet, Lake Tanganyika. – Brichard, P., 1978:423, key; 1989:479, key. – Sands, D., 1983:23, checklist. – Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:125, check-li st. – Burgess, 1989:195, checklist. – Kobayagawa, M., 1989:14, photo(misiden tified). – Coulter, G.W., 1991:182, 191, table 8.16, habitat, diet, reproduction. Synodontis irsacae Matthes, H., 1959:78, description, n. sp., Lake Tanganyika; 1962:37, placed into synonymy of S. dhonti Diagnosis Axillary pore absent; mandibular teeth 15-38; body with large spots; fin spines dark; 8-9 pectoral fin rays; bl ack triangles at base of all rayed fins; eye 23.1-43.7% snout length; premaxillary toothpa d interrupted; secondary bran ches on internal mandibular barbels present; occipito-nucha l shield covered or not c overed with skin; granular papillae present on skin of body; hindgut chamber absent; maximum TL 400 mm. The lack of an axillary pore distinguishes Synodontis dhonti from S. granulosus S. multipunctatus Synodontis sp 1 and S. petricola A smaller eye (23.1-43.7% of snout length vs. 44.9-62.0% and 64.2-81.0%, respecti vely) further separates juvenile S. dhonti from S. multipunctatus and Synodontis sp 1 Synodontis dhonti differs from S. multipunctatus Synodontis sp. 1, S. petricola Synodontis sp. 2, S. polli and Synodontis sp. 3 in having an occipito-nuchal pro cess which is not covered by skin. S ynodontis dhonti can be further distinguished from S. petricola, Synodontis sp 2 S. polli, and Synodontis sp 3 by having a lower number of mandibular teeth (15-38 in S. dhonti vs. 31-50 in S. petricola 38-51 in Synodontis sp. 2, 40-70 in S. polli and 50-64 in Synodontis

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88 sp. 3). Synodontis dhonti is further separated from S. petricola and Synodontis sp 2 in having dark fin spines and papillae on the body skin, (vs. white fin spines and bare skin in S. petricola and Synodontis sp. 2). Black triangles on the rayed fins of S. dhonti separate this species from S. nigromaculatus where they are absent, while the presence of granular papillae on the skin of S. dhonti (vs. villous papillae in S. nigromaculatus ) further distinguish these species. Synodontis dhonti strongly resembles specimens of S. tanganaicae Previous descriptions state that these species can be separated by the gill raker count on the first br anchial arch (11-20 in S. dhonti vs. 27-28 in S. tanganaicae ) (De Vos and Thys van den Audenaerde, 1998). Description Morphometric and meristic data are give n in Table 9. Maximum total length 400 mm, standard length 325 mm. Body not compre ssed. Predorsal profile slightly convex. Preanal profile straight to convex. Skin on body forming numerous vertical epidermal folds; covered with minute, granul ar papillae extending onto all fins. Head slightly depressed; broad; skin c overed with granular papillae. Snout long, with bluntly rounded margin when viewed late rally and dorsally. Occipital-nuchal shield rugose in holotype; covered w ith skin in young specimens; terminating posteriorly with narrow, bluntly pointed process on either side of dorsal spine; ventrally with wide, rounded process which extends to upper margin of humeral process on either side of body. Eyes dorsolateral; ovoid; horizontal ax is longest. Interorbital area flat. Mouth inferior; lips wide and papillate. Mandibular teeth 15-38, short, unicuspid; arranged in single transverse row. Premax illary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete; numerous; arranged in 3, 1-2, and 1 irregular rows, respectively.

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89 Maxillary barbel with distinct basal memb rane; extending to, or just past base of pectoral fin; granular papillae on base and along anterior margin. External mandibular barbel extending to point just short of anterior margin of pectoral girdle, with 2-7 short, tuberculate branches; secondary branches some times present. Internal mandibular barbel about half length of external barbel; with 4-5 pairs of t uberculate branches; secondary branches present. Dorsal fin I,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 spine margin with small serrations distally. Pectoral fin I,8-9; pos terior margin straight to broadly rounded. Pectoral fin spine roughly equal in length to dorsal fin; striated, slightly curved, terminating in short, dark filament; anterior spine margin granulous in adult specimens; smooth in juvenile specimens; posterior spine margin with large retrorse serrations along entire length. Adipose fin we ll developed; margin convex. Pelvic fin i,6; located at vertical through posterior base of dorsal fin; tip of appresse d fin barely reaches base of anal fin. Anal fin iii-v,7-9; posterior margin rounded; base located ventral to anterior third of adipose fin. Caudal fin I,7,8,I; forked; lobes rounded. Humeral process narrow, elongated, granul ous; weakly develope d ridge on ventral margin; dorsal margin concave; terminating in blunt point (Figs. 5A, 5B). Axillary pore absent. Gut 0.7-0.8 times body length (n = 5, MRAC 130315, MRAC A3-033-P-00360040, UF 160941). Hindgut chamber absent. Coloration Dorsum grayish to cuprous brown (Figs. 22, 23). Belly coloration slightly lighter than dorsum. Scattered, irre gularly-shaped black spots pr esent on juvenile specimens.

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90 Spots becoming smaller and eventually disa ppearing in large specimens. Maxillary barbels white. Mandibular barbels white bases sometimes with scattered dusky pigmentation. Iris copper colored. Dorsal and pectoral fin spines dark, terminating in short, light to dusky colored filaments. Anteri or margin and venter of pectoral fin light colored. All rayed fins with black triangles at their base in young specimens; posterior margins white in color. Black triangles indist inct in holotype. Bo th lobes of caudal fin black, posterior margin white. Distribution Lake Tanganyika (Fig. 24); common. Habitat Littoral to benthic zones over shell, sand and mud bottoms, to a maximum depth of 40 m (Coulter, 1991). Diet Synodontis dhonti is omnivorous, feeding on algae, sponges, ostracods, small crabs, insect larvae, and fish eggs in the stomach of this species (Matthes 1959; Coulter, 1991). Reproduction As with the other Synodontis species of the lake, detail ed information regarding the reproduction of S. dhonti is nonexistent. Matthes (1962) found mature female specimens to contain between 1,000 and 3,000 eggs. Coul ter (1991), noting that this large number closely resembles the number found in S. nigromaculatus suggests that this may indicate that S. dhonti spawns in lake tributaries. This hyp othesis suffers from a lack of published support, as no specimens of S. dhonti have been reported outside of the lake and the number of eggs produced by S. nigromaculatus is not actually reported in the publication cited by Coulter. It is reas onable however, to assume that S. dhonti differs in its

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91 reproductive behavior from the other, smaller Synodontis species of the lake given the large differences in egg number between these species and S. dhonti A detailed study of the reproduction of Tanganyikan Synodontis is clearly needed before detailed statements can be made about the reproduction of any of these species. Taxonomic Remarks Synodontis dhonti Boulenger has been a source of some taxonomic confusion due to the major morphological differences ex isting between the un ique, large-bodied holotype and smaller specimens of this spec ies. Juvenile specimens were described under the name Synodontis irsacae by Matthes (1959), who qui ckly decided that they were, in fact, juvenile specimens of S. dhonti (Matthes, 1962). A number of ontogenetic changes in external morphology and morpho metric ratios serve to complicate the identification of this species. A list of these changes, given by Matthes (1962), follows (translated): 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 shie ld and the humeral process becomes rugose and more strongly de veloped, causing a general increase in the relative size of the head. The humeral process becomes more blunt. The number of mandibul ar teeth increases. The band of premaxillary teeth becomes wider. The dorsal and pectoral fin spines beco me relatively shorter, wider and more rugose, while the anterior serrations become obscured.

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92 The lack of adult specimens (excepting the unique holotype) of this species is confusing, given the relatively large numb er of young specimens in collections. However, the material examined in this study has indicated that S. tanganaicae Borodin may be a synonym of S. dhonti and not of S. lacustricolus Poll, as previously believed (De Vos and Thys van den Audenaerde, 1998). S. tanganaicae attains the same size as the holoype of S. dhonti and the lectotype of S. tanganaicae whose photograph appears on Harvard UniversityÂ’s Museum of Compar ative Zoology online database looks more like S. dhonti in most respects than it does nominal specimens of S. lacustricolus Furthermore, there is significant overlap in the number of mandibular teeth possessed by these species (15-38 in S. dhonti 33-49 in S. tanganaicae when nominal specimens of S. lacustricolus are included, 33-40 when they are excluded), one of the main differences cited as separating these tw o species (De Vos and Thys van den Audenaerde, 1998). If this hypothesis were confirmed, it would e xplain the lack of adult specimens of S. dhonti in collections, as they would have been identified as S. tanganaicae This remains speculation however, as the types of S. tanganaicae have yet to be examined.

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93 Table 10. Morphometric measurements and meristic counts for Synodontis dhonti All morphometric measurements given in per cent of base measurement. Meristic data for Holotype is identified by a “*” s uperscript. SL = Standard length, HL = Head length, XMB = External ma ndibular barbel, IMB = Internal mandibular barbel, HPL = Humeral pro cess length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 27.2 17.7-28.6 23.52.6 Head length/SL 33.3 27.9-33.3 31.01.0 Snout-dorsal length/SL 42.6 36.7-42.6 39.41.3 Adipose fin length/SL 25.1 25.1-41.5 33.33.3 Maxillary barbel length/SL 23.8 16.1-28.4 22.03.1 Dorsal spine length/SL 23.9 18.8-28.0 23.42.1 Pectoral spine length/SL 24.4 19.0-27.5 23.71.6 Head width/HL 81.7 75.9-97.1 82.84.1 Head depth/HL 82.5 59.1-82.5 69.85.4 HPL/HL 49.9 35.7-55.8 47.54.3 Snout length/HL 48.6 48.6-58.2 53.52.1 Eye/HL 11.2 11.2-22.6 18.62.0 Interorbital width/HL 35.8 27.3-44.3 34.73.6 Postorbital length/HL 43.4 29.8-43.4 35.22.5 Mouth length/HL 34.2 33.7-52.8 40.94.3 Maxillary barbel length/HL 71.3 52.4 -91.8 71.09.9 XMB/HL 33.3 33.3-63.0 46.96.2 IMB/HL 16.9 14.4-30.7 21.73.5 IMB/XMB 50.7 28.6-59.6 46.56.6 Dorsal spine length/HL 71.8 59.6-88.3 75.46.6 Pectoral spine length/HL 73.2 60.6-88.7 76.76.0 Eye/Snout length 23.1 23.1-43.7 35.14.0 Interorbital width/Snout length 73.7 50.9-86.7 65.07.5 Postorbital length/Snout length 89.5 56.0-89.5 66.06.3 Eye/Interorbital width 31.4 31.4-72.7 54.88.9 Eye/Postorbital length 25.8 25.8-69.3 53.77.8 HPL/HPD 232.7 180.3-358.2 280.739.4 CPL/CPD 107.8 60.4-301.0 94.715.0 Meristics Mandibular tooth count 15(1); 16(2); 18(1); 19(1); 21(5); 22*(8); 23(4); 24(6); 25(4); 26(5); 27(4); 29(1); 38(1) Dorsal fin count I,7*(43) Pectoral fin count I,8* (30), I,9 (14), Pelvic fin count i,6*(43) Anal fin count iii,7(1); iii, 8(2); iii,8,i( 2); iii,9(1); iv,7(10) iv,7,i(4); iv,8(23); v,8*(1) Caudal fin count i,7,8,i(5) Total vertebrae count 32(3); 33(2) Measurement Holotype Range (n=43) MeanSD

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94 Figure 22. Synodontis dhonti Holotype, MRAC 14344, 395 mm TL, 325 mm SL. Top left – dorsal view, bottom left – lateral view, right – ventral view, showing mouth and barbels.

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95 Figure 23. Synodontis dhonti MRAC 130315, Holotype of S. irsacae 142 mm TL, 116 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view.

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96 Figure 24. Distribution of Synodontis dhonti T denotes type locality.

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97 Synodontis polli Gosse, 1982 (Figures 5, 25, 26, 28; Tables 2, 11) Synodontis polli Gosse, J.P., 1982:48, nom. nov. – Daget, J., Gosse, J.P., and Thys van den Audenaerde, D.F.E, 1986:141, check-l ist. – Brichard, P., 1989:474, 475, 478, 479, 483, skin texture, photos(some misiden tifications). – Burgess, W.E., 196, 556, 559, 560, 576, checklist, photos(so me misidentifications). Synodontis eurystomus Matthes, H., 1959:77, descripti on, n. sp., Lake Tanganyika; 1962:31, pl. 1, figs. B, C, description, diet, Lake Tanganyika. – Poll, M., 1971:405, figs. 190, 191, pls. VII.1, XII. 8, description. – Brichard, P., 1978:357, 424, photos, key. – Sands, D., 1983:23, 79, checklist, photo.Kobayagawa, M., 1989:14, photo(misiden tified). – Coulter, G.W., 1991:181, 182, table 8.16, habitat, diet, reproduction. Diagnosis Axillary pore absent; mandibular teeth 40-70; body with large spots; fin spines dark; 8-9 pectoral fin rays; black triangles on base of all rayed fins; eye 25.8-39.2% snout length; premaxillary toothpa d interrupted; secondary bran ches on internal mandibular barbels present; occipito-nuchal shield covered with skin; vill ous papillae present on skin of body; long gut; well-developed hi ndgut chamber; maximum TL 160 mm. The lack of an axillary pore immediately distinguishes Synodontis polli from S. granulosus S. multipunctatus Synodontis sp 1, and S. petricola Additionally, S. polli differs from these species and S. dhonti by generally having a greater number of mandibular teeth (40-70 in S. polli vs. 28-51 in S. granulosus 13-29 in S. multipunctatus 17-26 in Synodontis sp. 1, 31-50 in S. petricola 38-51 in Synodntis sp 2, and 15-38 in S. dhonti ). Synodontis polli also differs from S. petricola and Synodontis sp. 2 by having dark fin spines. The larg e spots on the body distinguish S. polli from S. granulosus S. nigromaculatus and S. tanganaicae and the black triangles at the base of all the rayed fins further separate S. polli from S. nigromaculatus and S. tanganaicae Synodontis polli is further distinguished from S. granulosus S. nigromaculatus S. tanganaicae and S.

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98 dhonti in having an occipito-nuchal sh ield which is covered by skin. Synodontis polli differs from Synodontis sp 3 in having a long gut (4.0-5.5 times TL in S. polli vs. 0.751.35 times TL in Synodontis sp 3) (fig. 28), a well-develope d hindgut chamber (fig. 28), villous papillae on the skin (vs. short, flattened papillae in Synodontis sp 3 ), and large, irregular, closely spaced spots on the skin (vs. small, regul ar, more widely spaced spots in Synodontis sp. 3). Description Morphometric and meristic data are give n in Table 11. Maximum total length 180 mm, standard length 145 mm. Body not compre ssed. Predorsal profile slightly convex. Preanal profile convex. Skin on body forming numerous vertical epidermal folds covered with minute, villous papillae which do not extend 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. Occipital-nuchal shield covered with skin, terminating posteriorly with wide pointed process on either side of dorsal spine, ventrally with wide, rounded process which extends to upper margin of humeral process on either side of body. Eyes dorso lateral, ovoid, horizont al axis longest. Interorbital area flat to slightly convex. Mouth inferior; lips widened and papilla te. Mandibular teeth 40-70, short, unicuspid; arranged in 8 short, transverse rows. Premaxillary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete; nume rous; arranged in 3, 2, and 1 irregular rows, respectively. Maxillary barbel short; ex tending at least to base of pectoral spine; lacking branches or crenelations; basal membrane na rrow. External mandibular barbel extending

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99 to point just short of anterior margin of pectoral girdle; with 4-5 short, non-tuberculate branches; lacking secondary branches. Intern al mandibular barbel approximately 1/3 to 2/3 length of external barbel; with 3-5 pa irs of tuberculate branches; many secondary branches present. Dorsal fin I, 7; posterior margin slightly concave. Dorsal fin spine short; striated; slightly curved, terminating in short, dark filament; anterior margin of fin spine smooth; posterior spine margin with small serrations dist ally. Pectoral fin I, 8-9; posterior margin straight. Pectoral fin spine roughly equal in length to dorsal fin, stri ated, slightly curved, terminating in short, dark filament; anterior spine margin granulate in adult specimens; many small, antrorse serrations along anteri or margin in juvenile specimens; posterior spine margin with large, retrorse serrations along entire length. Adipose fin long; well developed; margin convex. Pelvic fin i,6; locat ed at or anterior to vertical through origin of adipose fin; tip of appressed fin does not reach base of anal fi n. Anal fin iii-v,7-9; posterior margin rounded; base located at ve rtical through center of adipose fin. Caudal fin i,7,8,i; forked; lobes rounded. Humeral process wide; triangular; granul ous; poorly developed ridge on ventral margin; dorsal margin convex; 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. 28). Coloration Dorsum olive brown, covered with large, i rregularly shaped, black spots (Fig. 25). Spots proportionately larger, irregular, sometim es confluent in juvenile specimens. Belly lighter, with smaller, irregular ly shaped spots. Maxillary and mandibular barbels white. Iris copper colored. Dorsal a nd pectoral fin spines dark, terminating in short, dusky

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100 filaments. Pectoral spine with thin, light stripe along anterior 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 made up of closely spaced spots. Both lobes of caudal fin with black bar from ba se to tip of fin; posterior margin of fin white, becoming dusky with growth. Distribution Lake Tanganyika (Fig. 26); common. Habitat Rocky coasts within the littoral zone, to a maximum depth of 20 m (Matthes, 1959; Coulter, 1991). Diet S. polli feeds on algae scraped from rocky substr ates, as well as small invertebrates (e.g. ostracods, chironomid larvae, small shri mps) (Matthes, 1959; Coulter, 1991). The digestive morphology of this sp ecies (long intestine, well-d eveloped hindgut chamber) is similar to highly herbivorous mochokid genera ( Hemisynodontis and Brachysynodontis ), suggesting that invertebrates are only ingested incidenta lly along with the algae upon which they live. Reproduction No information exists regarding the spaw ning season, behavior, or location of this species, though mature specimens appear to be caught at many different times of the year. It is known is that the number of eggs releas ed is likely to be small, as a single female specimen has been found to contain approximately 110 eggs (Matthes, 1959).

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101 Taxonomic Remarks The taxonomy of S. polli has remained fairly stable, except for the replacement of its original name S. eurystomus due to the fact that Synodontis eurystomus Matthes was a junior homonym of Synodontis eurystomus Pfeffer, a synonym of Chiloglanis deckenii Peters (Gosse, 1982). Specimens from the southern part of this speciesÂ’ range show significant variation in their coloration pattern, w ith the spots on the body becoming smaller or in some cases, disappearing entirely. In a ll other respects however, these specimens closely resemble specimens collected in the northern part of the lake. The exceptions to this statement are specimens coming from Cape Chaitika on the so uthwestern coast, which are described as a distinct species in the following account.

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102 Table 11. Morphometric measurements and meristic counts for Synodontis polli All morphometric measurements given in per cent of base measurement. Meristic data for Holotype is identified by a “*” s uperscript. SL = Standard length, HL = Head length, XMB = External ma ndibular barbel, IMB = Internal mandibular barbel, HPL = Humeral pro cess length, HPD = humeral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indi cate number of specimens. Body depth/SL 22.0 20.2-27.0 23.01.7 Head length/SL 28.2 26.3-32.6 29.81.2 Snout-dorsal length/SL 38.6 35.2-41.4 37.91.4 Adipose fin length/SL 35.8 29.2-45.6 35.63.4 Maxillary barbel length/SL 13.6 10.6-28.3 19.23.8 Dorsal spine length/SL 19.2 12.8-23.4 19.62.5 Pectoral spine length/SL 22.5 17.8-25.0 22.01.6 Head width/HL 86.6 79.6-94.5 85.83.1 Head depth/HL 73.1 61.9-79.6 68.63.2 HPL/HL 44.6 35.0-51.5 43.83.4 Snout length/HL 55.7 50.8-60.2 55.21.8 Eye/HL 18.8 15.1-21.2 18.41.3 Interorbital width/HL 43.3 31.8-44.3 36.92.8 Postorbital length/HL 35.4 30.9-38.2 34.41.4 Mouth length/HL 57.3 44.4-62.2 54.64.0 Maxillary barbel length/HL 48.0 45.4-93.5 65.111.7 XMB/HL 41.9 30.9-67.1 48.37.6 IMB/HL 25.0 15.5-31.5 23.43.1 IMB/XMB 59.6 34.1-69.3 49.17.6 Dorsal spine length/HL 68.2 42.9-80.1 65.78.7 Pectoral spine length/HL 79.8 59.8-84.0 73.85.6 Eye/Snout length 33.7 25.8-39.3 33.42.7 Interorbital width/Snout length 77.8 57.2-81.0 67.05.2 Postorbital length/Snout length 63.6 52.4-69.7 62.33.3 Eye/Interorbital width 43.3 38.3-60.9 50.15.0 Eye/Postorbital length 53.0 41.4-66.3 53.74.8 HPL/HPD 192.8 146.8-252.2 192.624.9 CPL/CPD 127.5 86.9-144.0 108.012.6 Meristics Mandibular tooth 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) Dorsal fin count I,7*(66) Pectoral fin count I,8(32); I,9*(33) Pelvic fin count i,6*(65) Anal fin count iii,8(5); iii, 8,i(4); iii,9(11); iv,7(4); iv,7,i(2); iv,8*(26); iv,8,i(4); iv,9(9); v,7(1) Caudal fin count i,7,8,i(8) Total vertebrae count 33(2); 34(4); 35(2) Measurement Holotype Range (n=66) MeanSD

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103 Figure 25. Synodontis polli Holotype, MRAC 130440, 148 mm TL, 125 mm SL. Top dorsal view, middle – lateral view, bottom – ventral view.

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104 Figure 26. Distribution of Synodontis polli T denotes type locality.

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105 Synodontis sp. 3 n. sp. (Figures 5, 27, 29, 30; Tables 2, 12) Holotype : UF 160942, TL 138 mm, SL 117 mm, Lake Tanganyika at Chaitika, Zambia, via Pete Hauschner, Tropical Fish Collector, coll. V.2005. Paratypes : (9) UF 160942, TL 128-148 mm, SL 108-121 mm, collection data same as for holotype, (5) MRAC 78-25-P20-24, 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 fi n rays; black triangles on base of all rayed fins; eye 26.033.4% snout length; premaxillary toothpad interrupted; secondary branches on internal mandibular barbels present; occipi to-nuchal shield c overed with skin; short, flat papillae on skin of body; short gut; hindgut chamber absent; maximum TL 150 mm. The lack of an axillary pore distinguishes Synodontis sp 3 from S. granulosus S. multipunctatus Synodontis sp 1, and S. petricola Synodontis sp 3 is further separated from S. petricola as well as Synodontis sp 2 and S. dhonti by having dark fin spines (vs. white in S. petricola and Synodontis sp. 2), and having a greater number of mandibular teeth (50-66 in Synodontis sp. 3 vs. 31-50 in S. petricola 38-51 in Synodntis sp. 2, and 15-38 in S. dhonti ). The spots on the body distinguish Synodontis sp. 3 from S. granulosus S. nigromaculatus and S. tanganaicae and the black triangles at the base of all the rayed fins further separate Synodontis sp. 3 from S. nigromaculatus and S. tanganaicae Synodontis sp. 3 is further distinguished from S. granulosus S nigromaculatus S. tanganaicae and S. dhonti in having an occipito -nuchal shield which is covered by skin. Synodontis sp 3 differs from S. polli in having a short gut (0.8-1.4 times TL in S. polli vs. 4.0-5.5 times TL in S. polli ) (Fig. 28, 29), no hindgut chamber,

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106 short flat papillae on the skin (vs. villous papillae in S. polli ), and moderately sized, round, widely spaced spots on the skin (vs. mode rate to large, irregular, more closely spaced spots in S. polli ). Description Morphometric and meristic data are give n in Table 12. Maximum total length 150 mm, standard length 120 mm. Body not compre ssed. Predorsal profile slightly convex. Preanal profile straight to slightly convex. Skin on body forming numerous vertical epidermal folds; short, flat papillae pres ent, giving skin pebbled texture. Head slightly depressed and broad; skin covered with papillae; papillae extend onto bases of maxillary barbels as well as all pa ired fins. Snout with bluntly rounded margin when viewed laterally and dorsally. Occi pital-nuchal shield covered with skin, terminating posteriorly with wide, pointed proc ess on either side of dorsal spine, ventrally with wide, rounded process which extends to upper margin of the humeral process on either side of body. Eyes dorsolateral; ovoid; horizontal axis longest. Interorbital area flat to slightly convex. Mouth inferior; lips widened and papilla te. Mandibular teeth 50-66, short, unicuspid; arranged in 8 short, transverse rows. Premaxillary toothpad interrupted; primary, secondary and tertiary premaxillary teeth discrete; numerous; arranged in 3-4, 2, and 1-2 irregular rows, respectively. Maxillary barbel short; extendi ng at least to base of pector al spine; small papillae at base; basal membrane narrow. External mandi bular barbel extending to point just past anterior margin of pectoral girdle; with 36 short, weakly tuberc ulate branches; usually lacking secondary branches. Internal mandibul ar barbel approximately 1/3 to 1/2 length

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107 of external barbel; with 3-5 pairs of t uberculate branches; many secondary branches present. Dorsal fin I, 7; posterior marg in straight to slightly concave. Dorsal fin spine short; striated; slightly curved, terminating in short, dusky filament; anterior margin of fin spine granulous; posterior spine margin with small serrations distally. Pectoral fin I, 8-9; posterior margin straight to broadly rounded. Pectoral fi n spine roughly equal in length to dorsal fin, striated, slightly curved, terminating in short, dusky filament; anterior spine margin granulous; posterior spine margin wi th large, retrorse serrations along entire length. Adipose fin long; well developed; ma rgin convex. Pelvic fin i,6; located at vertical through adipose fin base; tip of appr essed fin does not reach base of anal fin. Anal fin iii-v, 7-9,i; posterior margin rounded; base located at vert ical through center of adipose fin. Caudal fin i,7,8,i; forked; lobes rounded. Humeral process triangular; granulous; poorly developed ridge on ventral margin; dorsal margin concave; terminating in sharp point (Fig. 5D). Axillary pore absent. Gut short, 0.8-1.4 times body length (n = 8, 160942). Hindgut chamber absent (Fig. 29). Coloration Dorsum olive brown, covered with modera tely sized, regularly shaped, widely spaced, black spots (Fig. 27). Belly lighter, with smaller, regularly shaped spots. Maxillary and mandibular barbel s white. Iris copper colored. Dorsal and pectoral fin spines dark, terminating in shor t, dusky filaments. Pectoral spine with thin, light stripe along anterior margin. All rayed fins with bl ack triangles at their base; posterior margins dusky. Triangles may be completely solid or made up of closely spaced spots. Both lobes of caudal fin with black bars from ba se to tip of fin; posterior margin dusky.

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108 Distribution Lake Tanganyika (Fig. 30). Known only from the Cape Chaitika area, local abundance unknown. Habitat The exact habitat of this sp ecies is unknown. It is likel y that, like the other smallbodied Synodontis species, Synodontis sp. 3 inhabits coastal, fairly shallow, rocky areas. Diet Most of the specimens of this species examined during this study were obtained through the aquarium trade, and had not b een feeding upon their natural diet. Though these specimens, when alive, readily ingested algal matter, their short gut and lack of a hindgut chamber suggests that this species is suited to a more carnivorous diet. Examination of the stomach contents of fres hly caught specimens is needed to determine differences in diet between this species and S. polli which may have led to the differences in gut morphology seen between these species. Reproduction No information exists regarding the reproduction of Synodontis sp. 3. Taxonomic Remarks Superficial examination of specimens belongi ng to this species might lead to their identification as specimens of S. polli However, a closer examination, taking into account such factors as the nature of the pa pillae on the body, differen ces in color pattern, and especially differences in gut mor phology, shows that these specimens clearly represent a new species. As there are no name s available which might be applied to this species, a new specific epithet will be a pplied to this species upon this studyÂ’s publication.

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109 Table 12. Morphometric and meristic counts for Synodontis sp. 3. All morphometric measurements given in percent of base measurement. Ranges for S. polli are provided for comparison. SL = Standa rd length, HL = Head length, XMB = External mandibular barbel, IMB = In ternal mandibular barbel, HPL = Humeral process length, HPD = hu meral process depth, CPL = Caudal peduncle length, CPD = Caudal peduncle depth. Numbers in parentheses indicate number of specimens. Body depth/SL 20.2-27.0 18.2-20.1 19.30.6 Head length/SL 26.3-32.6 27.5-30.6 29.21.0 Snout-dorsal length/SL 35.5-41.4 34.9-38.3 36.31.1 Adipose fin length/SL 29.2-45.6 31.2-44.8 38.44.0 Maxillary barbel length/SL 10.6-28.3 12.9-26.4 18.23.8 Dorsal spine length/SL 12.8-23.4 16.1-22.3 18.61.8 Pectoral spine length/SL 17.8-25.0 17.6-23.8 21.01.9 Head width/HL 79.6-101.6 78.2-86.6 82.92.6 Head depth/HL 61.9-79.6 63.6-73.8 68.43.0 HPL/HL 35.0-51.5 36.9-46.4 40.92.6 Snout length/HL 50.8-60.2 53.4-59.2 56.01.6 Eye/HL 15.1-21.2 14.5-18.5 17.11.1 Interorbital width/HL 31.8-44.3 33.6-40.3 36.51.7 Postorbital length/HL 30.9-36.7 34.0-36.9 35.51.0 Mouth length/HL 44.4-62.2 49.0-57.0 52.02.2 Maxillary barbel length/HL 45.4-93.5 44.0-92.3 62.514.0 XMB/HL 30.9-67.1 32.4-53.9 43.15.8 IMB/HL 15.5-31.5 16.2-26.4 20.82.9 IMB/XMB 34.1-69.3 34.7-63.1 49.29.1 Dorsal spine length/HL 42.9-80.1 55.1-74.9 63.65.8 Pectoral spine length/HL 59.8-84.0 59.2-79.1 71.96.6 Eye/Snout length 25.8-39.3 26.0-33.4 30.62.3 Interorbital width/Snout length 57.2-81.0 61.4-73.8 65.23.2 Postorbital length/Snout length 52.4-68.0 57.4-68.0 63.43.0 Eye/Interorbital width 38.3-60.9 38.2-53.0 47.04.3 Eye/Postorbital length 41.4-66.3 42.5-52.6 48.32.7 HPL/HPD 146.8-252.2 149.0-228.0 169.520.4 CPL/CPD 86.9-144.0 94.7-117.0 107.07.4 Meristics Mandibular tooth count 50(1); 52(1); 54(1); 55(1); 57(1); 58(3); 59(2); 60(2); 62(1); 64(1); 66(1) Dorsal fin count I,7(15) Pectoral fin count I,8(10); I,9(5) Pelvic fin count i,6(15) Anal fin count iii,7(1); iv,7(1); iv,7,i(2); iv,8(5); iv,8,i(3); iv,9(1); iv,9,i(1); v,9(1) Caudal fin count i,7,8,i(3) Total vertebrae count 33(2); 34(1) Measurement S. polli Synodontis sp 3 (n=15) MeanSD

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110 Figure 27. Synodontis sp 3, UF 160942, 139 mm TL, 116 mm SL. Top – dorsal view, middle – lateral view, bottom – ventral view.

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111 Figure 28. S. polli MRAC 131007-008, 137 TL, 112 SL, with intestine extracted. Hindgut chamber indicated with white arrow.

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112 Figure 29. Synodontis sp. 3, UF 160942, 139 mm TL, 118 mm SL, with intestine extracted. Note the relative shortness of the intestine and lack of a discernible hindgut chamber.

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113 Figure 30. Distribution of Synodontis sp 3. T denotes type locality.

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114 CHAPTER 5 DISCUSSION The examination of 336 specimens of Synodontis from Lake Tanganyika has led to the recognition of ten Synodontis species within the Lake Tanga nyika basin, nine of which are endemic and three of which are new to science. The addition of these new species represents a significan t contribution to our knowledge of Tanganyikan Synodontis species. Furthermore, this study represents the first comprehensive examination of the taxonomy of this species group in over 30 years and has revealed that significant work remains to be done. Examination of S. nigromaculatus has revealed that it most likely represents a complex of seve ral undescribed species, one of which was identified during this study. This species, from the Luachim o River in the Congo River drainage, will be described in a later publication. Th e examination of the taxonomy of S. tanganaicae and S. dhonti has revealed several problems in previous taxonomic studies. However, solutions to these problems will require fu rther analysis and the collection of new comparative material. Finally, the charac ters examined in this study offer some preliminary insights into the possible relationships among the endemic species of Lake Tanganyika. These insights and their impli cations for future research are briefly discussed below. The rayed fin color pattern and skin folds of the endemic Tanganyikan Synodontis species offer support for the monophyly of th is group. Brooks (1950) suggested that, in the genera of Lake Tanganyikan fishes that are represented in the lake by a single

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115 nonendemic species and an endemic species flock (Mochokidae and Mastacembelidae), the endemic species probably arose from an an cient species due to cleavage of the lake basin, while the nonendemic species, e.g. S. nigromaculatus is likely to be a recent addition to the lake fauna. Bric hardÂ’s (1978) assertion that the Synodontis species of Lake Tanganyika are probably not monophyletic is reasonable when S. nigromaculatus is included. More confusing is his apparent be lief that the endemic sp ecies flock also does not constitute a monophyletic group. Howeve r, this hypothesis has gained support from a molecular phylogeny generated using data fr om the cytochrome b gene (Day, pers. comm.). In this phylogeny, a Lake Victoria species, S. victoriae Boulenger, splits the Tanganyikan endemics into two clades, one consisting of S. granulosus S. multipunctatus and apparent specimens of Synodontis sp. 1, with the other clade consisting of the remaining lake endemics. The appearance of S. victoriae within the Tanganyikan species flock is surprising, as this species shows few morphological affinities to the endemic species of Lake Tanganyika. Though based on data from only one mitochondrial gene, certain aspects of the tree topology are supported by the current study. The grouping of S. granulosus S. multipunctatus and Synodontis sp. 1 appears to be morphologically supported by the unint errupted condition of the premaxillary toothpad, and the lack of secondary bran ches on the medial mandibular barbel. Morphological examination found no charac ters to support the second clade of Tanganyikan Synodontis or species relationships within this clade. Because the placement of S. victoriae within the Tanganyikan species flock may be due to its true

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116 sister species being excluded from the phylogene tic analysis, or limitations of the single gene studied, further studies, preferably uti lizing more gene regions and taxa, are needed. Synodontis species from lakes Victoria and Malawi, as well as the Congo River drainage were included in the abovementioned analysis. Synodontis nigromaculatus was placed in a clade with S. njassae Keilhack, and this clade was placed sister to the clade composed of the endemic Tanganyikan species and S. victoriae This is an interesting result, as the origins of the endemic Synodontis of Lake Tanganyika, and the fishes of Lake Tanganyika in general are unknown. Three possible connections between other African waterways and Lake Tanganyika have been proposed, from which the species of Lake Tanganyika may have originated: 1) a co nnection with the ancient Nile system, 2) a connection with Lake Malawi via Lake Rukwa, and 3) a connection with the Congo River system (Coulter, 1991). While molluscan and general (family level) ichthyological affinities, along with geographic proximity suggest that the third hypothesis is the most likely one (Coulter, 1991), cyt b data clearly indicate that in the case of the endemic Tanganyikan Synodontis species, the second hypothesis is supported; however, the number of species from the Congo included in the molecular phylogeny is limited (seven of approximately 40 species). A more comprehensive comparison of the Synodontis species of Lake Tanganyika and the neighboring drainages is clearly necessary before the phylogenetic position of the Tanganyikan species flock within the genus Synodontis can be proposed with any confidence. A robust phylogeny for Synodontis will permit comparative examination of many of the unique traits found within this genus. The functional significances of the epidermal skin folds, axillary pore, and hindgut chamber have yet to be determined. A

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117 study of the axillary pore will be particularly interesting, as Synodontis appears to be the only African catfish genus in which this stru cture has been documented. Even if this structure is found to serve an identical function to the st ructure found in the aforementioned families (Akysidae, Ariidae, Ictaluridae, Sisoridae), a more detailed study of the composition of its s ecretions will be extremely inte resting, as this has yet to be examined for any species. The hindgut cham ber may serve in a digestive capacity, as its gross morphology is very similar to that of a structure found in the Kyphosidae. An analysis of the contents of the hindgut chamber of Synodontis for the presence of fermentative gut microbes and short chain fatty acids must be performed to confirm that it serves a similar function to the fermentation chamber of kyphosids. Synodontis clearly represents a group for whic h significant research remains to be done. Given the high diversity of species and behaviors found in this genus, its neglect by past ichthyologists is conf using. It is hoped that this study of the taxonomy of the Tanganyikan species flock will serve as a springboard for many future studies.

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118 LIST OF REFERENCES Barnard, K.H. 1948. Report on a collection of fi shes from the Okovango river, with notes on Zambesi fishes. Annals of th e South African Museum, 36: 407-458. Bell-Cross, G. 1965-1966a. The distribution of fishes in Central Africa. Fisheries Research Bulletin of Zambia, 4: 3-20. Bell-Cross, G. 1965-1966b. Preliminary observations on Hydrocynus vittatus in the Upper Zambezi River system. Fisheries Research Bulletin of Zambia, 4: 21-27. Birkhead, W.G. 1967. The comparative toxicity of stings of the ictalurid catfish genera Ictalurus and Schilbeodes Comparative Biochemistry and Physiology, 22: 101111. Birkhead, W.G. 1972. Toxicity of stings of ariid and ictalurid catfishes. (4): 790-807. Borodin, N.A. 1936. On a collection of freshw ater fishes from Lake Nyasa, Tanganyika and Victoria in Central Africa. Zoologisches Jahrbuc h Jena (Syst.), 68: 1-34. Boulenger, G.A. 1898. Report on the collection of fishes made by Mr. J.E.S. Moore in Lake Tanganyika during his expedition, 1895-1896. With an appendix by J.E.S. Moore. Transactions of the Zoological Society of London, XV: 1-30. Boulenger, G.A., 1899. Second contribution to the ichthyology of lake Tanganyika. On the fishes obtained by the Congo Free Stat e Expedition under Lieut. Lemaire in 1898. Transactions of the Zoologi cal Society of London, XV: 87-96. Boulenger, G.A. 1900. Diagnoses of new fishes discovered by Mr. J.E.S. Moore in Lake Tanganyika. 1. Cyprinidae, Siluridae. Anna ls and Magazine of Natural History, 7, (VI): 478-481. Boulenger, G.A. 1901a. Les poissons du bassin du Congo. Publication de lÂ’Etat indpendant du Congo. Brussels, 8: I-LXII, 1-532. Boulenger, G.A. 1901b. Third contribution to the ichthyology of lake Tanganyika. Report on the collection of fishes made by Mr. J.E.S. Moore in Lakes Tanganyika and Kivu during his second expedition, 18991900. Transactions of the Zoological Society of London, XVI: 137-178.

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119 Boulenger, G.A. 1905. On a collection of fishes from Lake Bangwelo. Annals and Magazine of Natural History, 7, (XVI): 642-647. Boulenger, G.A. 1906. Fourth contribution to the ichthyology of Lake Tanganyika. Report on the collection of fishes made by Dr. W.A. Cunnington during the third Tanganyika Expedition. Transactions of the Zoological Society of London, XVII: 537-576. Boulenger, G.A. 1911. Catalogue of the fres h-water fishes of Africa in the British Museum (Natural History) London, vol. 2: I-XIII, 1-529. Boulenger, G.A. 1916. Catalogue of the fres h-water fishes of Africa in the British Museum (Natural History) London, vol. 4: I-XXVII, 1-392. Boulenger, G.A. 1917. Descripti ons of new fishes from La ke Tanganyika, forming part of the collection made by the late Dr. L. Stappers for the Belgian Government. Annals and Magazine of Na tural History, 8, (XX): 363-368. Boulenger, G.A. 1923. Descriptions of thr ee new freshwater fishes from Northern Rhodesia. Annals of the South African Museum, 13, (7): 437-438. Bowmaker, A.P. Research results, lac Ba ngweulu. Report, Joint Fisheries Research Organization of Northern Rhode sia and Nyasaland, n 11: 6-20. Brichard, P. 1978. Fishes of Lake Tanganyi ka. T.F.H. Publications, Inc. Hong Kong. Brichard, P. 1989. Cichlids and all the ot her Fishes of Lake Tanganyika. T.F.H. Publications, Inc. Hong Kong. Brooks, J.L. 1950. Speciation in ancient lakes. Quarterly Review of Biology, Baltimore, 25(2): 30-36, 131-176. Burgess, W.E. 1989. An Atlas of Freshwater a nd Marine Catfishes. T.F.H. Publications, Inc. Hong Kong. Clements, K.D. and Choat, J.H. 1997. Compar ison of herbivory in the closely-related marine fish genera Girella and Kyphosus Marine Biology, 127: 579-586. Coulter, G.W. 1965-1966. The deep benthic fishes at the South of Lake Tanganyika with special reference to distribution and feeding in Bathybates species, Hemibates stenosoma and Chrysichthys species. Fisheries Research Bulletin of Zambia, 4: 33-38. Coulter, G.W. 1991a. The benthic fish communities. In Lake Tanganyika and its Life, G.W. Coulter, editor. Oxford University Press, New York: 151-199.

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124 BIOGRAPHICAL SKETCH Jeremy J. Wright was born on December 30, 1981, in Postville Iowa. One of two children, he developed a keen interest in natu re at an early age, and could usually be found searching local streams and marshes fo r fishes and amphibians when he was not otherwise occupied with reading about them. He received his Bachelor of Science degree in marine biology and his Bachelor of Arts de gree in chemistry from the University of Rhode Island, where he graduated summa cum laude in May, 2004. He moved to Gainesville, Florida in June, 2004 and enroll ed in UFÂ’s Department of Zoology as a graduate student, with a research assistan tship from the All Catfish Species Inventory Project funded by the U.S. National Science Fo undation. He is curr ently investigating options for future work on catfish systematics.