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Taxonomic Revision of the Amphilius uranoscopus Group of East-Central Africa (Teleostei

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Title:
Taxonomic Revision of the Amphilius uranoscopus Group of East-Central Africa (Teleostei Amphiliidae)
Creator:
Thomson, Alfred William
Place of Publication:
[Gainesville, Fla.]
Florida
Publisher:
University of Florida
Publication Date:
Language:
english
Physical Description:
1 online resource (73 p.)

Thesis/Dissertation Information

Degree:
Master's ( M.S.)
Degree Grantor:
University of Florida
Degree Disciplines:
Zoology
Committee Chair:
Page, Lawrence M.
Committee Members:
Kimball, Rebecca T.
Armbruster, Jonathan
Graduation Date:
8/11/2007

Subjects

Subjects / Keywords:
Anus ( jstor )
Fish ( jstor )
Head ( jstor )
Natural history ( jstor )
Pectorals ( jstor )
Peduncle ( jstor )
Pigments ( jstor )
Rivers ( jstor )
Species ( jstor )
Vertebrae ( jstor )
Zoology -- Dissertations, Academic -- UF
africa, amphiliidae, amphilius, taxonomy
Genre:
bibliography ( marcgt )
theses ( marcgt )
government publication (state, provincial, terriorial, dependent) ( marcgt )
born-digital ( sobekcm )
Electronic Thesis or Dissertation
Zoology thesis, M.S.

Notes

Abstract:
The taxonomy of species of the Amphilius uranoscopus group in east-central Africa is reviewed, and the Amphilius uranoscopus group is recognized by the following unique combination of characters: absence of a crenellated epidermal fold at the base of the caudal fin, usually 8+9 (i,7,8,i) principal caudal-fin rays, small bilateral bony swimbladder capsules, 36-39 total and 20-23 caudal vertebrae, and leading pterygiophore of the dorsal fin intercepting the vertebral column at the first, second or third post-Weberian vertebrae. The examination of all available museum specimens of the genus Amphilius from east-central Africa reveals the existence of at least five species of the Amphilius uranoscopus group in the region, A. grandis, A. oxyrhinus, A. krefftii, A. uranoscopus and Amphilius sp. 1. Three of the species were previously believed to be synonyms of Amphilius uranoscopus, and Amphilius sp. 1 is new to science. Amphilius sp. 1 is distinguished from A. grandis, A. oxyrhinus, A. krefftii and A. uranoscopus by its distinctive adult pigment pattern consisting of many small darks spots on the head, body and fins (vs. body, head and fins lacking dark spots in A. grandis, A. krefftii and A. uranoscopus or large dark spots present on body in A. oxyrhinus). It is additionally distinguished from these species by its absence of a visible epidermal core extending posteriorly from the mandibular barbels (vs. presence in A. oxyrhinus and A. krefftii), rounded caudal-fin lobes (vs. pointed in A. grandis and A. oxyrhinus), forked caudal fin and absence of light dorsal saddles (vs. caudal fin shallowly emarginate and light dorsal saddles present at the base of the dorsal and adipose fin in A. krefftii and A. uranoscopus), and fewer branchiostegal rays (6-7 vs. 8-9 in A. uranoscopus). The caudal fin shape and pigment patterns observed suggest that a northern and a southern group of species exist in east-central Africa. The northern species (A. grandis, A. oxyrhinus and Amphilius sp. 1) have a more deeply forked caudal fin and dark markings either as adults or juveniles, and lack light dorsal saddles. The southern species (A. krefftii and A. uranoscopus) have no dark markings, shallowly emarginate caudal fins, and light dorsal saddles present at the bases of the dorsal and adipose fin. Despite differences observed in pigment pattern and intensity between the Amphilius in the Wami, Ruvu and Rufiji drainages, all of these populations are tentatively recognized as a single species because most of the specimens examined were juveniles and only juvenile pigmentation could be compared. The three other nominal species synonymized with A. uranoscopus, A. hargeri, A. brevidorsalis and A. platychir cubangoensis, are discussed, but their status is beyond the geographic scope of this study and only tentative conclusions are made about the identity of these species. ( en )
General Note:
In the series University of Florida Digital Collections.
General Note:
Includes vita.
Bibliography:
Includes bibliographical references.
Source of Description:
Description based on online resource; title from PDF title page.
Source of Description:
This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis:
Thesis (M.S.)--University of Florida, 2007.
Local:
Adviser: Page, Lawrence M.
Statement of Responsibility:
by Alfred William Thomson.

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UFRGP
Rights Management:
Copyright Thomson, Alfred William. Permission granted to the University of Florida to digitize, archive and distribute 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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TABLE OF CONTENTS

page
A C K N O W L E D G M E N T S ............................................... ..................................... .....................3

LIST OF TA BLES .............. ................................................................................... ..... 6

L IST O F FIG U R E S ............................................................................... 7

ABSTRAC T ...........................................................................................

CHAPTER

1 IN TR O D U C T IO N ................................................................................ 10

N atu ra l H isto ry ................................................................................................................. 1 0
S y stem atic H isto ry ............................................................................................................ 12

2 M A TERIALS AN D M ETH OD S ........................................................................ ..18

3 M ATERIALS EXAM INED ....................................................... 21

Amphilius grandis ..................................... 21
A mp hilius krefftii ................................................................21
Amphilius oxyrhinus ........................................ .. ....... ........22
Amphilius uranoscopus...... ...... .................................. .. ...... ............ ....22
Amphilius sp. 1 ................................................................. ........ 23

4 SY STEM A TIC A CCOU N TS... ......................................................................... ...............24

A mphilius uranoscop us group ......................................................................... ....................24
Amphilius grandis Boulenger 1905 ................................. .......................... ............. 25
D diagnosis ...... .... .............. .... ........ .. ....... ....... ... ....... .............. 25
D e scrip tio n ..........................................................................................................2 6
C oloration .................................................... ..... ....................... ..........................2 7
D distribution ...........................................................................................................28
Amphilius krefftii Boulenger 1911 .............................................. .............................. 28
D iag n o sis ......... .... .............. ..................................... ...........................2 8
D e scrip tio n ................................................................2 9
C o lo ra tio n ...................... .. ............. .. ....................................................... 3 1
D distribution ............... ..... ............... ............... ..................................... 3 1
Amphilius oxyrhinus Boulenger 1912.............................................................................. 31
D ia g n o sis ................................................................................................................... 3 2
D e scrip tio n ................................................................3 2
Coloration ........................... ..................... ............ .........34
D istrib u tio n .................... ...... ......... .. .................................................3 5
Amphilius uranoscopus (Pfeffer 1889).............................................................................. ...35
D iag n o sis ................................. .................. .. ............................ .......................... 3 6


4









LIST OF TABLES


Table page

4-1 External traits diagnostic for species of the Amphilius uranoscopus group in east-central
A frica .......................................................... ...................................4 3

4-2 Morphometric data for Amphilius grandis. Range and mean include the syntypes ..............44

4-3 Morphometric data for Amphilius krefftii. Range and mean include the syntypes.................45

4-4 Morphometric data for Amphilius oxyrhinus. Range and mean include the holotype. ...........46

4-5 Morphometric data for Amphilius uranoscopus. Range and mean include the specimens
from the W am i drainage. ......................................................................... ....................47

5-1 Branchiostegal ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus ........... ..... ......... ................. 64

5-2 Total gill raker counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three
populations of A. uranoscopus............... ............................... ............... 64

5-3 Branched pectoral-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii
and three populations of A. uranoscopus............. .............. ............. ............... 64

5-4 Branched anal-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus ........... ..... ......... ................. 64

5-5 Total vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three
populations of A. uranoscopus............... ............................... ............... 65

5-6 Preanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus ........... ..... ......... ................. 65

5-7 Postanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus ........... ..... ......... ................. 65

5-8 First dorsal pterygiophore intercept counts in A. oxyrhinus, A. grandis, Amphilius sp. 1,
A. krefftii and three populations of A. uranoscopus...................................... ....................65










Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Eusso Nyiro drainage]; Harry,
1953: 189, synonymy; Copley, 1958: 98 (in part), [Tana drainage].

Amphiliusplatychir (non Ginther) Whitehead, 1958: 198 (in part), [Eusso Nyiro drainage].

Amphilius uranoscopus (non Pfeffer) Skelton, 1994: 126, [Tana drainage]; Seegers et al., 2003:
37 (in part), [Tana drainage].

Diagnosis

Visible epidermal core extending posteriorly from the mandibular barbels present; caudal

fin forked with tips of lobes pointed; adults with large dark spots or blotches on body; spots on

head and fins absent or indistinct; dorsal saddles absent.

The presence of a visible epidermal core extending posteriorly from the bases of the

inner and outer mandibular barbels distinguishes Amphilius oxyrhinus from A. grandis,

Amphilius sp. 1 and A. uranoscopus. Amphilius oxyrhinus is distinguished from A. krefftii and

further distinguished from A. uranoscopus by its forked caudal fin with the tips of the caudal-fin

lobes pointed (vs. caudal fin shallowly emarginate with rounded lobes) and the absence of light

dorsal saddles (vs. light dorsal saddles present at the base of the dorsal and adipose fin). It is

further distinguished from Amphilius sp. 1 by the larger spots or blotches on the body of adults

(vs. small spots present on head and body), its pointed caudal-fin lobes (vs. caudal fin with

rounded lobes), and the absence of spots on the fins. Adults ofAmphilius oxyrhinus are further

distinguished from adults of A. grandis, Amphilius sp. 1 and A. uranoscopus by the presence of

dark spots or blotches on the body of adults (vs. adults without dark spots or blotches on body).

Description

Morphometric data as in Table 4-4. Body elongate, ventral profile flattened ventrally to

anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from

tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest









body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated

epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin,

closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line

complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae

37 (4) or 38 (4). Preanal vertebrae 21 (7) or 22 (1). Postanal vertebra 16 (5) or 17 (3). First

dorsal pterygiophore intercept count 1 (4) or 2 (4).

Head and anterior part of body depressed and broad. Head wedge shaped in lateral view.

Snout broad, pointed when viewed from above. Head becoming wider from tip of snout to

pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped

or deeply concave connection. Membrane connected by a small frenum at midline, but with

shallow groove at posterior margin.

Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe

large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed.

Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary

teeth short and conical, tooth patches forming U-shaped band, separated medially.

Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and

flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle

pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of

lowerjaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates

anterolaterally of inner mandibular barbel, extending just short of edge of branchiostegal

membrane. Visible epidermal core extending posteriorly from bases of mandibular barbels.

Branchiostegal membrane with 7 (4), 8 (6) or 9 (2) rays. Gill rakers on first epibranchial 3 (13);










301E


-5E




-10Et


30E 35E 40E
km
0 200 400
Figure 4-10. Known distribution ofAmphilius sp. 1. Some symbols represent more than one
collection site. T denotes type locality.


40o









Diagnosis

Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal

fin emarginate with tips of lobes rounded; no dark spots present on head, body or fins of adults;

light dorsal saddles present at the base of the dorsal and adipose fin; branchiostegal rays 8-9.

The absence of a visible epidermal core extending posteriorly from the bases of the inner

and outer mandibular barbels distinguishes Amphilius uranoscopus from A. oxyrhinus and A.

krefftii. Amphilius uranoscopus is distinguished from A. grandis, Amphilius sp. 1 and further

distinguished from A. grandis by the shape of the caudal fin emarginatee in A. uranoscopus vs.

shallowly emarginate or forked in A. grandis and forked in Amphilius sp. 1 and A. oxyrhinus)

and the presence of light dorsal saddles at the base of the dorsal and adipose fin (vs. light dorsal

saddles absent). Additionally the caudal-fin lobes are rounded in A. uranoscopus (vs. pointed in

A. grandis and A. oxyrhinus). Amphilius uranoscopus is further distinguished from A. oxyrhinus

and Amphilius sp. 1 by the absence of dark spots or blotches on the head and body in adults. Its

greater number of branchiostegal rays (8-9 vs. 6-7) and lack of spots on the fins further

distinguishes it from Amphilius sp. 1.

Description

Morphometric data as in Table 4-5. Body elongate, ventral profile flattened ventrally to

anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from

tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest

body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated

epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin,

closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line

complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae









caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown with light distal

edge; ventral surfaces light yellow. Maxillary and mandibular barbels brown. Caudal fin with

dark crescent-shaped band at base. Juvenile coloration: medium-sized dark spots on head and

body in specimens larger than 40.0 mm SL, but spots usually absent on specimens under 40.0

mm SL. Spots on body arranged in line along lateral line. All fins light yellow, with medial

bands of brown pigment all rayed fins. Pectoral and pelvic fins with first unbranched ray dark

brown dorsally.

Distribution

Tana and Eusso Nyiro (Ewaso Ng'iro) River drainages, Kenya.

Amphilius krefftii Boulenger 1911

(Figures 4-3 and 4-4; Table 4-3)

Amphilius krefftii Boulenger, 1911: 356, fig. 276, Original description, Type locality: Usambara,
in rivulet running from Anani Hills to Sigi River. [Sigi River drainage, Tanzania]; Harry,
1953: 189, synonymy; Copley, 1958: 100 [Pangani drainage]; Whitehead, 1958: 198
[Kenya and Tanzania]; Bailey, 1969: 192 [Sigi and Pangani drainages]; Bernacek, 1980:
36 [Tanzania]; Skelton, 1984: 45, in synonymy of A. uranoscopus.

Pimelodus (Amphilius) uranoscopus (non Pfeffer) Hilgendorf, 1905: 411 (in part), [Pangani
drainage].

Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Galina drainage] [BMNH
1969.3.24.63-69, BMNH 1969.3.24.70]; Copley, 1958: 100 [Galina drainage] [BMNH
1969.3.24.63-69, BMNH 1969.3.24.70]; Bailey, 1969: 192 (in part), [Pangani drainage].

Amphilius uranoscopus (non Pfeffer) Harry, 1953: 189 (in part), synonymy; Skelton, 1994: 126
(in part), [Galina and Pangani drainages]; Seegers et al., 2003: 37 (in part), [Pangani
drainage].

Amphiliusplatychir (non Gunther) Copley, 1958: 100 (in part), [Pangani drainage].

Diagnosis

Visible epidermal core extending posteriorly from the mandibular barbels present; caudal

fin emarginate with tips of lobes rounded; adults without dark spots on head, body or fins; light









CHAPTER 3
MATERIALS EXAMINED

Amphilius grandis

Eusso Nyiro drainage: BMNH 1908.9.17.13-18, Kenya: Nyiro-Narok; Niro-Narok

system, elev. 4000-5000 ft., 016'00"N, 360 31'33"E (6: 93.4-181.4); BMNH 1912.3.22.119,

Kenya, Eusso Nyiro, below falls, 012'00"N, 38 00'00"E (1: 113.6). Tana drainage: BMNH

1904.12.23.50-52, Kenya, Chania R. of Tetse, Tana system, Kenya, elev. 7000 ft., 102'58"N,

370 05'34"E (2: 136.1-160.0, Syntypes); BMNH 1937.6.4.36-42, Kenya, Thika River, Ndula

Falls, 102'58"N, 370 05'34"E (12: 32.1-65.9).


Amphilius krefftii

Galina drainage: BMNH 1969.3.24.63-69, Kenya, Tsabo [Tasvo] River tributary of

Athi [Galina] River, mountain Mbololo, 3 0' N, 38 20'E, (7: 43.6-99.5); BMNH 1969.3.24.70,

Kenya, Voi River, Teita hills, Voi District, 322'44"N, 380 34'07"E (1: 106.6). Pangani

drainage: BMNH 1909.10.19.26-27, Tanzania, Usambara, in rivulet running from Anani Hills

to Sigi R., 50 O'N, 380 48'E (2: 47.5-86.0, Syntypes); BMNH 1905.7.25.41-42, Kenya, Kibosho,

Kilimandjaro, 3 12', 37 19' (2: 77.9-94.9); BMNH 1968.10.25.3, Tanzania, River Lume (upper

reaches of Pangani River) at source, 5 miles east of Lake Chala, 323'22"N, 370 43'45"E (1:

106.6); BMNH 1968.10.25.5-6, Tanzania, Kisiwani, near Amani, East Usambura mountains,

elev. 1,475 ft., 511'34"N, 380 37'39"E (2: 36.2-43.1); BMNH 1968.10.25.7, Tanzania, River

Sigi, east Usumbura Mountains, 5 O'N, 38 48'E (1: 72.7); BMNH 1968.10.25.8, Tanzania,

River Lume (upper reaches of Pangani River) east of Taveta, 3023'22"N, 370 43'45"E (1: 67.2);

FMNH 111678, Tanzania, East Usambara Mountains, 4.5 km ESE Amani, Monga Tea Estate, 5

6'N, 38 36'E (1: 112.0); FMNH 111684, Tanzania, West Usambara Mts., in river (stream) near

Ambangulu Tea Estate factory, 5004'54"N, 380 25'55"E (1: 29.0).









Amphilius sp. 1 by having the lobes of the caudal fin pointed (vs. rounded) and the absence of

spots on the fins.

Description

Morphometric data as in Table 4-2. Body elongate, ventral profile flattened ventrally to

anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from

tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest

body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated

epidermal fold. Anus and urogenital openings located at posteriormost extent of pelvic fin, closer

to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete,

extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 38 (3), 39

(6) or 40 (1). Preanal vertebrae 20 (1), 21 (3) or 22 (6). Postanal vertebra 16 (1), 17 (4) or 18

(5). First dorsal pterygiophore intercept count 1 (2) or 2 (8).

Head and anterior part of body depressed and broad. Head wedge shaped in lateral view.

Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to

pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped

connection. Membranes connected by a small frenum at midline, but with shallow groove at

posterior margin.

Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe

large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed.

Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary

teeth short and conical, tooth patches forming U-shaped band, separated medially.

Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and

flattened with pointed tip; barbel extending posterolaterally from corner of mouth to just short of

pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of










30fE


-_-\j- -It














30E 35E 40E
km
0 200 400

Figure 4-2. Known distribution ofAmphilius grandis. Some symbols represent more than one
collection site. T denotes type locality.


40F









posterior nares with prominent tubular rims; nares separate but relatively close to each other.

Posterior nare located about midway between eye and tip of snout.

Dorsal-fin origin at point over middle of pectoral fin. Dorsal fin with i,5 (1) or i,6 (53)

rays, and fin margin slightly convex. Pectoral fin with i,8 (3), i,9 (23) or i,10 (28) rays with first

ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight,

with 4-5 innermost rays progressively shorter making inner part of posterior fin margin rounded.

Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (54) rays with first ray

unbranched and greatly thickened. Pelvic fin with straight posterior margin.

Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base,

extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded

edge, not deeply incised posteriorly. Caudal fin emarginate with tips of lobes rounded, with

i,7,6,i (1), i,7,7,i (1) or i,7,8,i (52) principal rays. Anal fin with short base, origin posterior to

origin of adipose-fin base, with ii,6 (6), iii,5 (2), iii,6 (15), iii7 (18) or iii,8 (11) rays. Anal fin

margin almost straight.

Coloration

Dorsal and lateral surfaces of head and body brown. Ventral region light yellow. Light

dorsal saddles at anterior and posterior of dorsal-fin and adipose-fin bases. Dorsal, adipose,

caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown, ventral surfaces

light yellow. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped

band at base. Juvenile coloration: Similar to that of adults, except all fins light yellow, with

small blotches of brown pigment on the branched rays. Pectoral and pelvic fins with first

unbranched ray dark brown dorsally.

Distribution

Wami, Ruvu and River Rufiji drainages, Tanzania.










Amphilius oxyrhinus

Eusso Nyiro drainage: BMNH 1912.22.120, Kenya, Eusso Mara, a swift mountain

stream, tributary of Eusso Nyiro, 0 12'N, 38 00'E (1: 166.0, Holotype). Tana drainage:

BMNH 1965.12.7.125, Kenya, Rogati River, Sagana, 040'42"N, 370 16'18"E (1: 42.9); BMNH

1965.12.7.126-128, Kenya, Rogati River, Sagana, 040'42"N, 370 16'18"E (3: 60.9-89.7);

BMNH 1965.12.7.129, Kenya, Rogati River, Sagana, 040'42"N, 370 16'18"E (1: 156.8);

BMNH 1965.12.7.130-131, Kenya, Rogati River, Sagana, 040'42"N, 37 16'18"E (2: 74.9-

98.7); BMNH 1965.12.7.132, Kenya, Rogati River, Sagana, 040'42"N, 370 16'18"E (1: 51.8);

BMNH 1966.6.28.2-3, Kenya, Rogati River, Sagana, 040'42"N, 37 16'18"E (2: 65.9-102.6);

BMNH 1966.8.25.18, Kenya, Rogati River, Sagana, 040'42"N, 370 16'18"E (1: 34.8); 34.8;

MRAC 74-48-P-12-13, Kenya, upper Tana, side creek, 00042'S, 37014'E (2: 52.6-60.3).


Amphilius uranoscopus

Rufiji drainage: FMNH 111679, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest

Reserve, 4 km W, 5 km N Chita, along the Chita-Ihimbo trail, elev. 1460 m, 8028'30"S,

35054'25"E (8: 24.4-40.4); FMNH 111680, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest

Reserve, 4.5 km W Chita, along Chita-Ihimbo trail, elev. 600 m, 8031'10"S, 35054'15"E (18:

25.0-68.4.0); FMNH 111681, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest Reserve, 4.5

km W Chita, along Chita-Ihimbo trail, elev. 600 m, 8031'10"S, 35054'15"E (14: 25.5-77.4).

Ruvu drainage: FMNH 111682, Tanzania, Uluguru Mountains, Uluguru North Forest Reserve,

3 km W, 1.3 km N Tegetero, elev. 1345 m, 60 55' 45" S, 370 42' 20" E (1:55.7); FMNH 111683,

Tanzania, Uluguru Mts., Uluguru North Forest Reserve, 5.1 km W, 2.3 km N Tegetero, elev.

1535 m, 6055'12"S, 37041'0"E (3: 26.4-52.4); MNHN 1897-0003, Tanzania, Torrent of Mrogoro

at Zanguebar, Oukami, 6046'00"S, 37043'45"E (1: 117.3, Holotype of Chimarrhoglanus leroyi),









small dark spots (although a few larger blotches may be present), and spots are also always

present on the head. In A. oxyrhinus, the dark markings are always in the form of large blotches

or spots, with only a few smaller spots or blotches present. Spots on the head are usually absent

or indistinct. The pectoral, pelvic, dorsal, anal, adipose and caudal fins ofAmphilius sp. 1 are

also heavily spotted. Amphilius oxyrhinus lacks spots on the fins, although some dark pigment

may be present. Light dorsal saddles at the base of the dorsal and adipose fins are present in A.

krefftii and A. uranoscopus (except specimens from the Ruvu drainage, see discussion below).

These light dorsal saddles are not present in A. grandis, A. oxyrhinus or Amphilius sp. 1.

Two forms of the caudal fin were observed. In Amphilius grandis, A. oxyrhinus and

Amphilius sp. 1, the caudal fin is deeply emarginate or moderately forked, but in A. krefftii and A.

uranoscopus the caudal fin is feebly emarginate. In A. grandis and A. oxyrhinus, the tips of the

caudal-fin lobes are pointed, while in Amphilius sp. 1, A. krefftii andA. uranoscopus the tips of

the caudal-fin lobes are noticeably more rounded.

A visible epidermal core extending posteriorly from the mandibular barbels is always

present in Amphilius oxyrhinus and A. krefftii and always absent in A. grandis, Amphilius sp. 1

and A. uranoscopus. The visible epidermal core is a vein-like structure extending posteriorly

from the bases of the inner and outer mandibular barbels. The structure is pigmented, but even

in old specimens in which the pigment has faded, the structure is still noticeable.

Frequency tables were constructed for counts of eight different meristic characters.

Branchiostegal ray count was the only meristic character found to be useful for distinguishing

species. Amphilius sp. 1 had the fewest branchiostegal rays, with all specimens having 6 or 7

branchiostegal rays. Amphilius uranoscopus had the most, with all specimens having 8 or 9

branchiostegal rays. Branchiostegal ray count usually distinguishes A. krefftii from A.









rounded caudal-fin lobes (vs. pointed in A. grandis and A. oxyrhinus), forked caudal fin and

absence of light dorsal saddles (vs. caudal fin shallowly emarginate and light dorsal saddles

present at the base of the dorsal and adipose fin in A. krefftii and A. uranoscopus), and fewer

branchiostegal rays (6-7 vs. 8-9 in A. uranoscopus).

The caudal fin shape and pigment patterns observed suggest that a northern and a

southern group of species exist in east-central Africa. The northern species (A. grandis, A.

oxyrhinus and Amphilius sp. 1) have a more deeply forked caudal fin and dark markings either as

adults or juveniles, and lack light dorsal saddles. The southern species (A. krefftii and A.

uranoscopus) have no dark markings, shallowly emarginate caudal fins, and light dorsal saddles

present at the bases of the dorsal and adipose fin.

Despite differences observed in pigment pattern and intensity between the Amphilius in

the Wami, Ruvu and Rufiji drainages, all of these populations are tentatively recognized as a

single species because most of the specimens examined were juveniles and only juvenile

pigmentation could be compared. The three other nominal species synonymized with A.

uranoscopus, A. hargeri, A. brevidorsalis and A. platychir cubangoensis, are discussed, but their

status is beyond the geographic scope of this study and only tentative conclusions are made

about the identity of these species.









pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped

or deeply concave connection. Membrane connected by a small frenum at midline, but with

shallow groove at posterior margin.

Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe

large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed.

Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary

teeth short and conical, tooth patches forming U-shaped band, separated medially.

Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and

flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle

pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of

lowerjaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates

anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane.

Branchiostegal membrane with 7 (16) or 8 (1) rays. Gill rakers on first epibranchial 2 (11), 3 (3),

or 4 (1); rakers on first ceratobranchial 5 (5), 6 (8) or 7 (2); total gill rakers on first arch 7 (5), 8

(4), 9 (5) or 10 (1).

Eyes small, positioned dorsolaterally approximately midway between tip of snout and

posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter.

Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and

posterior nares with prominent tubular rims; nares separate but relatively close to each other.

Posterior nare located about midway between eye and tip of snout.

Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (17) rays, and fin

margin slightly convex. Pectoral fin with i,9 (11), i,10 (5) or i, 1 (1) rays with first ray

unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with









emarginate) and the absence of light dorsal saddles (vs. light dorsal saddles present at the base of

the dorsal and adipose fin). It is distinguished from A. uranoscopus by its fewer branchiostegal

rays (6-7 vs. 8-9).

Description

Morphometric data as in Table 4-6. Body elongate, ventral profile flattened ventrally to

anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from

tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest

body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated

epidermial fold. Anus and urogenital openings located just posterior to base of pelvic fin, closer

to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete,

extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 37 (2), 38

(7) or 39 (3). Preanal vertebrae 20 (2), 21 (3) or 22 (7). Postanal vertebra 15 (1), 16 (5), 17 (4)

or 18 (2). First dorsal pterygiophore intercept count 1 (12) or 2 (1).

Head and anterior part of body depressed and broad. Head wedge shaped in lateral view.

Snout broad, moderatly pointed when viewed from above. Head becoming wider from tip of

snout to pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-

shaped or deeply concave connection. Membrane connected by a small frenum at midline, but

with shallow groove at posterior margin.

Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe

large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed.

Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary

teeth short and conical, tooth patches forming U-shaped band, separated medially.

Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and

flattened with pointed tip; barbel extending posterolaterally from corner of mouth to just short of









Table 4-1. External traits diagnostic for species of the Amphilius uranoscopus group in east-
central Africa.


Visible epidermal Absent
core extending
posteriorly from
the mandibular
barbels
Caudal-fin shape Deeply


Caudal lobes

Dark spots on
body (adult)

Dark spots on
head


Spots on fins


Dorsal saddles

Branchiostegal
rays


Present





Emarginate


Rounded


Absent


Absent



Absent


Present

Usually 7


A. oxyrhinus


Present





Forked


Pointed

Large


Usually
absent,
indistinct if
present
Absent


Absent

7-9


A. uranoscopus

Absent





Emarginate


Rounded


Absent


Absent



Absent


Present

8-9


Amphilius sp. 1


Absent





Forked


Rounded

Small


Present



Present


Absent

6-7


Character


A. grandis A. krefftii


emarginate
or forked
Pointed


Absent


Absent



Absent


Absent

7-8










Amphilius grandis Boulenger 1905

(Figure 4-1 and 4-2; Table 4-2)

Amphilius grandis Boulenger 1905a: 63, P1. 7 (fig. 3), Original description, Type locality:
Chania R. of Tetse, Tana system, Kenya, elev. 7000 ft.; Boulenger 1905b: 48,
Headwaters of the Tana system; Boulenger 1911: 353, 355, fig. 275 (in part), in key,
description, East Africa (Tana to Athi River systems) [Tana drainage only]; Boulenger,
1912: 675, diagnosis from Amphilius oxyrhinus, [Eusso Nyiro drainage], [BMNH
1912.3.22.119]; Boulenger, 1916: 306, Nairobi, Eusso Nyiro, below falls; Pellegrin 1936:
57, similarity to Amphiliusplatychir var. cubangoensis; Copley, 1941: 15 [Eusso Nyiro
and Tana drainages], Harry, 1953: 189 (in part), synonymy; Copley, 1958: 98, [Tana
drainage], [BMNH 1937.6.4.36-42], Whitehead, 1958: 198, [Kenya and Tanzania],
Bailey, 1969: 192 (in part), eastern rivers of Kenya; Skelton, 1984: 45, in synonymy of A.
uranoscopus.

Amphiliusplatychir (non Ginther) Whitehead, 1958: 198 (in part), [Eusso Nyiro drainage].

Amphilius uranoscopus (non Pfeffer) Skelton, 1994: 126, [Tana drainage]; Seegers et al., 2003:
37 (in part), [Tana drainage].

Diagnosis

Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal

fin deeply emarginate or forked with tips of lobes pointed; adults plain-colored, without dark

spots or blotches on body, head or fins; dorsal saddles absent.

The absence of a visible epidermal core extending posteriorly from the bases of the inner

and outer mandibular barbels distinguishes Amphilius grandis from A. oxyrhinus and A. krefftii.

Amphilius grandis is distinguished from A. uranoscopus and A. krefftii by its more deeply

emarginate caudal fin with the tips of the caudal-fin lobes pointed (vs. caudal fin shallowly

emarginate with rounded lobes) and the absence of light dorsal saddles (vs. light dorsal saddles

present at the base of the dorsal and adipose fin). Amphilius grandis is distinguished from

Amphilius sp. 1 and A. oxyrhinus by the absence of dark spots on the head or body of adults (vs.

adults of Amphilius sp. 1 with many small dark spots on head and body, and adults of A.

oxyrhinus with dark spots or blotches on body). Amphilius grandis is distinguished from










Table 5-5. Total vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus
Total Vertebrae 36 37 38 39 40
A. oxyrhinus 4 4
A. grandis 3 6 1
Amphilius sp. 1 2 7 3
A. krefftii 7 3 1
A. uranoscopus (Wami) 3
A. uranoscopus (Ruvu) 2 2
A. uranoscopus (Rufiji) 11 12 2

Table 5-6. Preanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus
Preanal Vertebrae 19 20 21 22
A. oxyrhinus 7 1
A. grandis 1 3 6
Amphilius sp. 1 2 3 7
A. krefftii 7 5
A. uranoscopus (Wami) 1 1 1
A. uranoscopus (Ruvu) 2 2
A. uranoscopus (Rufiji) 1 6 11 7

Table 5-7. Postanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus
Postanal Vertebrae 15 16 17 18
A. oxyrhinus 5 3
A. grandis 1 4 5
Amphilius sp. 1 1 5 4 2
A. krefftii 1 9 1
A. uranoscopus (Wami) 1 1 1
A. uranoscopus (Ruvu) 4
A. uranoscopus (Rufiji) 9 15 1

Table 5-8. First dorsal pterygiophore intercept counts in A. oxyrhinus, A. grandis, Amphilius sp.
1, A. krefftii and three populations of A. uranoscopus
1st DPI 1 2
A. oxyrhinus 4 4
A. grandis 2 8
Amphilius sp. 1 12 1
A. krefftii 11 1
A. uranoscopus (Wami) 3
A. uranoscopus (Ruvu) 4
A. uranoscopus (Rufiji) 23 1









The status of three other species synonymized with Amphilius uranoscopus, A. hargeri,

A. brevidorsalis and A. platychir cubangoensis, are beyond the geographic scope of this study,

but tentative conclusions can be made about the identity of these species. The type of A. hargeri

is similar to A. uranoscopus, but the first-dorsal pterygiophore intercepts the third post-Weberian

vertebra. Skelton (1984) hypothesized that it could be a hybrid specimen, but concluded that it

was best considered a junior synonym of A. uranoscopus. This character separates A. hargeri

from the species in east-central Africa. All the specimens examined in east-central Africa have

the dorsal pterygiophore intercepting the first or second post-Weberian vertebra, with the

majority of specimens examined having the dorsal pterygiophore intercepting the first post-

Weberian vertebra.

The type ofAmphilius brevidorsalis from Mozambique is most similar to A. uranoscopus

and may be a synonym of A. uranoscopus. The type of A. brevidorsalis lacks dark markings and

has distinct light dorsal saddles. Additionally it a visible epidermal core extending posteriorly

from the mandibular barbels and has a caudal fin shape characteristic ofA. uranoscopus. The

type specimens examined ofA. platychir cubangoensis have a caudal fin shape similar to that of

A. krefftii and A. uranoscopus, but differ from these species by having distinct dark markings on

the body. Additionally, a visible epidermal core extending posteriorly from the mandibular

barbelsis present in the type specimens examined, further distinguishing A. platychir

cubangoensis from A. uranoscopus.









uranoscopus; 18 specimens of A. krefftii had a count of 7 and one specimen had 8 branchiostegal

rays (Table 5-1). Total gill-raker counts did not distinguish between species, but the counts for

A. uranoscopus in the Wami, Ruvu and Rufiji did indicate differences in those populations

(Table 5-2). No clear patterns were observed in the frequency-distribution tables of branched

pectoral-fin rays, branched anal-fin rays, total vertebrae, preanal vertebrae, postanal vertebrae

and first dorsal pterygiophore intercept count (Tables 5-3 to 5-8).

The characters examined in this study offer some preliminary insights into the possible

relationships among species of the Amphilius uranoscopus group in east-central Africa and the

biogeography of the region. Amphilius grandis, A. oxyrhinus and Amphilius sp. 1, the species

with deeply emarginate or forked caudal fins, are distributed in the northern part of the east-

central region, while all A. krefftii and A. uranoscopus populations are distributed to the south of

these species. Additionally, A. grandis, A. oxyrhinus and Amphilius sp. 1 all have dark markings

either as adults or juveniles, while A. krefftii and A. uranoscopus never have dark markings.

Finally, light dorsal saddles are absent in the northern species, but present in the southern

species.

The caudal fin shape and pigment patterns observed suggest that a northern and a

southern group of species exist in east-central Africa. These northern and southern species

groups correspond to the physical geography of the region. The Eusso Nyiro, Tana and Galina

drainages all have high-gradient tributaries in the Mount Kenya-Aberdare highlands (Skelton,

1994). Amphilius grandis and A. oxyrhinus overlap in distribution in the Eusso Nyiro and Tana

drainages. Amphilius sp. 1 is restricted to the upper reaches of the Galina drainage in close

proximity to the Eusso Nyiro and Tana drainages.










CHAPTER 5
RESULTS

The present study examined all available museum specimens of the genus Amphilius

from east-central Africa and revealed the existence of at least five species of the Amphilius

uranoscopus group in the region. Previously, only one species of the Amphilius uranoscopus

group was recognized from the region. Three species previously believed to be synonyms of

Amphilius uranoscopus are determined to be valid species, and one species new to science is

described. Additionally, pigment differences were noticed in the A. uranoscopus from the

Wami, Ruvu and Rufiji drainages, but because only a few adult specimens are available from

these drainages only a single species is recognized from these drainages.

The recognition of Amphilius uranoscopus as a single widespread species has been based

largely on the lack of clear meristic and morphometric differences between populations.

Additionally, previous studies never looked at large number of specimens from the east-central

region. Species that had been described from the region were based on just a few specimens, and

only the type specimens of those species had been compared. Although differences in pigment

pattern and caudal-fin shaped have been noticed previously (Copley, 1958; Seegers, 1996b), this

is the first study to examine variation in these characters thoroughly.

Although there is some degree of intraspecific variation in pigment pattern, certain

aspects are consistent and are useful in distinguishing the species. Amphilius oxyrhinus and

Amphilius sp. 1 both have dark markings on the body in the form of blotches or spots as adults

and juveniles. Adults of A. grandis lack dark markings on the body, but juveniles have dark

spots. Dark markings are generally absent in adults and juveniles of A. krefftii and A.

uranoscopus. Amphilius krefftii may have a few dark areas on its body but never has distinct

dark blotches or spots. In Amphilius sp. 1, the dark markings are always present in the form of
















































Figure 4-1. Amphilius grandis, BMNH 1912.3.22.119, 113.6 mm SL; lateral, dorsal and ventral
view.









pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of

lowerjaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates

anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane.

Branchiostegal membrane with 6 (1) or 7 (21) rays. Gill rakers on first epibranchial 3 (19) or 4

(3); rakers on first ceratobranchial 6 (8) or 7 (14); total gill rakers on first arch 6 (1), 9 (7), 10

(12) or 11 (2).

Eyes small, positioned dorsolaterally approximately midway between tip of snout and

posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter.

Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and

posterior nares with prominent tubular rims; nares separate but relatively close to each other.

Posterior nare located about midway between eye and tip of snout.

Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (22) rays, and fin

margin straight. Pectoral fin with i,9 (12) or i, 10 (9) rays with first ray unbranched and greatly

thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays

progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted

posteriorly to dorsal-fin base. Pelvic fin with i,5 (22) rays with first ray unbranched and greatly

thickened. Pelvic fin with straight posterior margin.

Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base,

extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded

edge, not deeply incised posteriorly. Caudal fin deeply emarginate or forked with tips of lobes

rounded, with i,5,6,i (1), i,6,7,i (1), i,7,7,i (2), i,7,8,i (16) or i,8,8,i (1) principal rays. Anal fin

with short base, origin posterior to origin of adipose-fin base, with iii, 5 (7), iii,6 (13) or iii, 7 (2)

rays. Anal fin margin almost straight.











Table 4-2. Morphometric data for Amphilius grandis. Range and mean include the syntypes.
Syntypes (n=2) Range (n=21) MeanSD
%SL
Head length 25.8-27.3 23.1-27.8 25.81.1
Head width 21.1-22.1 19.7-23.5 21.31.1
Head height 12.9-13.0 11.0-14.9 13.51.0
Body depth 14.6-15.4 12.2-17.6 13.91.2
Body depth at anus 11.9-14.4 11.6-14.4 12.90.7
Predorsal length 36.1-37.8 33.7-40.4 37.21.4
Prepectoral length 22.3-23.8 19.6-24.5 21.91.4
Preanal length 75.1-77.1 65.7-78.4 73.42.6
Dorsal-fin base length 8.4-9.2 7.9-13.9 10.21.5
Adipose-fin base length 18.6-19.0 15.0-24.9 20.42.6
Anal-fin base length 8.2-8.9 8.2-17.8 11.1+2.3
Pelvic-fin length 14.6-16.7 14.6-20.8 18.61.8
Pectoral-fin length 17.1-18.4 17.1-26.5 21.52.5
Anal-fin length 14.3-15.1 14.3-21.1 17.91.9
Caudal peduncle length 18.6-19.0 14.6-19.0 16.81.3
Caudal peduncle depth 9.9-10.1 9.3-12.6 10.60.8
Anus-to-anal fin length 13.8-14.0 8.6-14.0 11.31.7
Prepelvic length 55.9-56.3 48.5-56.4 53.31.8
Postpelvic length 47.0-48.2 47.0-51.0 48.91.1
End-of-dorsal to adipose fin 42.2-44.9 38.9-48.9 43.22.8
Front-of-dorsal to caudal fin 62.1-64.9 57.1-70.8 64.23.6
Preanus length 61.0-62.5 58.9-64.1 61.71.4

%HL
Snout length 48.5-51.3 43.6-57.4 48.64.1
Interorbital distance 26.1-26.7 24.0-31.2 27.21.9
Maxillary barbel length 57.0-58.3 55.6-81.3 66.76.5
Inner mandibular barbel length 24.5-30.6 21.4-39.6 30.94.3
Outer mandibular barbel length 39.1-43.5 39.1-60.8 48.65.6
Eye diameter 10.8-11.9 8.0-17.3 13.42.7
Postorbital length 35.5-39.6 31.2-43.3 37.53.0









The Pangani, Ruvu and Wami drainages drain the south-east slopes of Kilimanjaro and

the eastern side of the Gregory Rift (Skelton, 1994). The present-day Rufiji drainage has grown

from a series of captures (Willis, 1936). Its largest tributary may once have drained into the

Indian Ocean via the Wami (Bailey, 1969) or the Pangani (Haldemann, 1962). Although

Amphilius krefftii also occurs in the Galina drainage, it is found only in tributaries of the lower

reaches of the river. These tributaries are in close proximity to the Pangani drainage, where the

species also occurs.

The Amphilius in the Wami, Ruvu and Rufiji drainages are tentatively recognized here as

a single species despite differences observed in pigment pattern and intensity. Because most of

the specimens examined were juveniles, only juvenile pigmentation could be compared. All

specimens examined from the Ruvu drainage, including the type of Chimarrhoglanis leroyi were

lighter in color than specimens from the Wami and Rufiji (Fig. 5-1). Additionally, these

specimens did not have the light dorsal saddles that were always observed in specimens from the

Wami and Rufiji. Given that amount of variation in darkness observed in other species of

Amphilius, the lighter color of the Ruvu population is not considered a significant taxonomic

character. Additionally, the lighter color of the type of C. leroyi may be due to the age and

preservation of the specimen, and the absence of light dorsal saddles in all specimens examined

may only be because of the lighter color of these specimens. The Wami and Rufiji populations

were similar in body coloration, but specimens from the Wami always had darker fins. Although

this difference was consistent, no additional differences were observed. Although specimens

from the different populations appeared to differ in body shape, no morphometric differences

were found.









Systematic History

In 1864, Gunther described Amphilius as a section of the genus Pimelodus for a new

species he described as Pimelodusplatychir from Sierra Leone, western Africa. Soon thereafter

he recognized Amphilius as a genus (Gunther, 1865). In 1889, Pfeffer, apparently unaware of

Gunther's description, created the genus Anoplopterus for a new species he described as An.

uranoscopus from the Wami River drainage in Tanzania. Nine years later, Vaillant created the

genus Chimarrhoglanis for a supposedly new species, which he described as C. leroyi from the

nearby Ruvu River drainage. Boulenger (1898a) synonymized C. leroyi with Pimelodus

platychir, but overlooked Ginther's description of the genus Amphilius and synonymized

Chimarrhoglanis with Anoplopterus. Boulenger (1898a) also recognized Pfeffer's A.

uranoscopus and suggested that the types of A. platychir did not come from Sierra Leone.

Poche (1902a) and Gunther (1902) refuted Boulenger's (1898a) suggestion that the types

of Amphiliusplatychir were not from Sierra Leone, and Poche (1902b), recognizing Amphilius as

the correct name for the genus, synonymized Anoplopterus and Chimarrhoglanis with

Amphilius. Despite Poche and Gunther's assertions, Boulenger continued to question the type

locality of Sierra Leone for A. platychir and recognized A. platychir as a species in eastern Africa

(Boulenger, 1902, 1905a, 1907a,1907b).

In 1905, Boulenger described Amphilius grandis from the Tana River drainage in Kenya.

He distinguished this species from A. uranoscopus by its longer head (length four times in total

length v. five times in total length) and shorter barbels maxillaryy barbel not reaching posterior

border of head vs. extending beyond posterior border of head). He distinguished this species

from A. platychir by its longer head (length 4 times in total length v. 4 to 4 1/2 times in total

length), longer snout (snout length 1 2/3 interocular width vs. 1 1/3 to 1 1/2 interocular width),

and a shorter caudal peduncle (not longer than deep vs. longer than deep).










30fE


-_-\j- -It














30E 35E 40E
km
0 200 400

Figure 4-6. Known distribution ofAmphilius oxyrhinus. Some symbols represent more than one
collection site. T denotes type locality.


40F









All available museum specimens from the Ewaso Nyiro, Tana, Galana, Pangani, Wami,

Ruvu and Rifiji drainages were examined. In addition, the type specimens ofAmphilius hargeri,

A. brevidorsalis and A. platychir cubangoensis were examined; however, because large series of

specimens from outside the east-central region could not be examined, only tentative conclusions

can be made about the identity of these species.

Measurements were made point-to-point with digital calipers, and data were recorded to

tenths of a millimeter. Subunits of the head are presented as proportions of the head length (HL).

Head length and measurements of the body parts are given as proportions of the standard length

(SL).

For the fin-ray counts, the numbers of unbranched soft rays are indicated by Roman

numerals, branched soft rays are indicated by Arabic numerals. The number of anterior

unbranched rays in the anal fin is difficult to determine, and the counts were checked with

radiographs whenever possible. Amphiliids typically have a small spinelet in front of the first

unbranched dorsal-fin ray. The spinelet is not included in the counts. Vertebrae were counted by

means of radiographs following the methods of Skelton (1976). Counts exclude 5 Weberian

vertebrae and include the ural centrum as one.

Material examined is given under each species account and is listed by the drainage

followed by the catalog number, country, locality, geographic coordinates, and in parentheses,

the number of specimens and the size range in mm SL. Geographic coordinates were determined

from maps. An effort was made to provide geographic coordinates for all specimens; however,

these coordinates are only estimates and some lots with different specific localities have the same

geographic coordinates because some localities could not be found on a map.









Abstract of Dissertation 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 THE Amphilius uranoscopus GROUP OF EAST-CENTRAL
AFRICA (TELEOSTEI: AMPHILIIDAE)

By

Alfred W. Thomson

August 2007

Chair: Lawrence M. Page
Major: Zoology

The taxonomy of species of the Amphilius uranoscopus group in east-central Africa is

reviewed, and the Amphilius uranoscopus group is recognized by the following unique

combination of characters: absence of a crenellated epidermal fold at the base of the caudal fin,

usually 8+9 (i,7,8,i) principal caudal-fin rays, small bilateral bony swimbladder capsules, 36-39

total and 20-23 caudal vertebrae, and leading pterygiophore of the dorsal fin intercepting the

vertebral column at the first, second or third post-Weberian vertebrae. The examination of all

available museum specimens of the genus Amphilius from east-central Africa reveals the

existence of at least five species of the Amphilius uranoscopus group in the region, A. grandis, A.

oxyrhinus, A. krefftii, A. uranoscopus and Amphilius sp. 1.

Three of the species were previously believed to be synonyms of Amphilius uranoscopus,

and Amphilius sp. 1 is new to science. Amphilius sp. 1 is distinguished from A. grandis, A.

oxyrhinus, A. krefftii and A. uranoscopus by its distinctive adult pigment pattern consisting of

many small darks spots on the head, body and fins (vs. body, head and fins lacking dark spots in

A. grandis, A. krefftii and A. uranoscopus or large dark spots present on body in A. oxyrhinus). It

is additionally distinguished from these species by its absence of a visible epidermal core

extending posteriorly from the mandibular barbels (vs. presence in A. oxyrhinus and A. krefftii),









TAXONOMIC REVISION OF THE Amphilius uranoscopus GROUP OF EAST-CENTRAL
AFRICA (TELEOSTEI: AMPHILIIDAE)




















By

ALFRED WILLIAM THOMSON


A THESIS 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

2007









Table 4-5. Morphometric data for Amphilius uranoscopus. Range and mean include the


specimens from the Wami drainage.
Wami dr. (n=3)


%SL
Head length
Head width
Head height
Body depth
Body depth at anus
Predorsal length
Prepectoral length
Preanal length
Dorsal-fin base length
Adipose-fin base length
Anal-fin base length
Pelvic-fin length
Pectoral-fin length
Anal-fin length
Caudal peduncle length
Caudal peduncle depth
Anus to anal fin length
Prepelvic length
Postpelvic length
End of dorsal to adipose
Front of dorsal to caudal
Preanus length


%HL
Snout length
Interorbital distance
Maxillary barbel length
Inner mandibular barbel length
Outer mandibular barbel length
Eye diameter
Postorbit length


24.9-26.1
22.7-23.3
12.3-13.6
13.8-14.8
12.9-15.0
36.0-36.2
20.3-21.4
73.2-74.9
9.8-10.2
20.8-25.0
9.2-10.6
18.2-20.6
19.9-23.5
18.0-19.4
16.3-18.4
10.4-13.2
10.2-11.2
50.9-53.2
47.6-50.0
42.4-47.0
63.9-67.3
60.6-64.1


46.6-51.7
27.8-30.3
54.5-82.8
29.4-34.3
53.5-64.9
12.1-14.9
37.7-39.6


Range(n=55)


23.4-30.1
19.3-25.5
10.0-16.4
9.5-16.7
9.6-16.0
33.8-40.8
17.0-25.4
60.2-78.6
7.9-12.7
19.1-29.3
7.0-19.4
15.0-21.0
17.7-23.8
15.4-21.6
14.5-19.7
7.8-13.2
7.5-13.5
48.6-57.6
44.4-54.1
39.2-49.8
63.2-72.5
58.0-69.4


40.1-52.4
23.7-32.3
54.5-112.8
22.6-45.4
36.5-77.4
9.5-20.0
31.0-45.7


MeanSD

25.81.4
21.31.4
13.31.3
12.71.6
11.81.2
36.61.5
20.61.9
72.1+3.9
10.3+1.1
24.72.4
11.8+2.4
18.41.5
21.31.3
18.3+1.7
17.01.1
9.8+1.1
10.41.2
52.71.6
49.31.9
44.22.5
66.3+2.0
62.3+2.0


46.1+3.2
28.33.3
78.7+11.1
33.5+5.5
55.29.4
14.72.2
38.33.1









All maps were created using the Online Map Creator (OMC), which can be found at

http://www.aquarius.ifm-geomar.de/ (Weinelt, 1996-2006). Distribution points on maps were

added using Abode Photoshop CS2. Points indicate only specimens examined in this study.

Synonymies include all accounts of the species in east-central Africa. The first page of

the account of the species and all figures are listed. If the species is also listed in a key on a

separate page from the account, that page is also listed. The type of information in the reference

is given followed by the locality for the species in the account. Additionally, any specimens

known that the account may be based on are listed. If the account is only based in part on the

species, only the information that is applicable to that species is listed. Accounts of species

outside east-central Africa that give only a general distribution in east-central Africa are

excluded.









epidermal fold and usually has 8 + 9 principal caudal-fin rays. Amphiliusplatychir was

identified as a member of the first group, and all records of the species from eastern and southern

Africa were determined to be based on a single widespread species. Skelton (1984) recognized

the name A. uranoscopus for this widespread eastern and southern Africa species and considered

C. leroyi, A. grandis, A. krefftii and A. oxyrhinus to be synonyms of this species. Three other

taxa were also synonymized with A. uranoscopus: Amphilius hargeri from Mlangi, British

Central Africa, Amphilius brevidorsalis from the Riviere Revue (Mozambique) and Amphilius

platychir cubangoensis from Angola. Figure 1-2 shows the type localities ofA. uranoscopus and

the seven nominal species that Skelton synonymized with it.

Amphilius uranoscopus is the only Amphilius species of Skelton's eastern and southern

African group currently recognized in east-central Africa. A single species of Skelton's western

African Amphilius group, A. jacksonii, has been reported to occur in the Rufiji and Wami

drainages in Tanzania (Seegers, 1996a) and is easily distinguished from A. uranoscopus by its

deeply forked caudal fin, crenellated epidermal fold, and 6 + 7 principal caudal-fin rays.

Although Skelton examined the type specimens of the five nominal species from east-central

Africa, his recognition ofA. uranoscopus as a single widespread species was based largely on

the examination of specimens from South Africa. Additionally, his conclusions were based on

the lack of clear meristic and morphometric differences between populations. He did not

examine variation in other characters like pigment pattern and caudal fin shape. Variation in

these characters in specimens identified as A. uranoscopus, suggests that the validity of the

nominal species synonomized with A. uranoscopus and the acceptance of A. uranoscopus as a

single species needs to be investigated.









UF 84882, Tanzania, Morogoro Region, southern side of Uluguro Mountains, about 1 km N of

Ruvu River, S of Kibungo Village, Kimboza Forest, 7000'00"S, 37048'00"E (7: 21.9-40.4).

Wami drainage: CAS 80494, Tanzania, Nguru Mountains, Nguru South Forest Reserve, 607'S,

37032'E (2: 49.1-101.4); BMNH 1969.1.15.1, Tanzania, Stream at Arusha in the foothills of

Mount Meru, 3021'57"S, 36040'28"E (1: 84.7).


Amphilius sp. 1

Galina drainage: BMNH 1905.12.11.2, Kenya, Nairobi River, Kikaya, 116'28"S,

36048'47"E (1: 160.3), BMNH 1909.11.15.21-23, Nairobi River, elev. 2500 ft., 116'28"S,

36048'47"E (3: 89.1-119.5), BMNH 1910. 10.31.31, Nairobi, Kenya, 116'28"S, 36048'47"E (1:

150.0), BMNH 1928.11.10.11-12, Nairobi River, elev. 6500 ft., 116'28"S, 36048'47"E (3: 59.6-

140.5), BMNH 1928.11.10.13-18, Kenya, Riara River, elev. 5700 ft., 110'01"S, 36049'19"E (6:

37.6-92.6), BMNH 1937.12.11.16-18, Kenya, Mbakasi River (3: 58.1-110.3), BMNH

1969.3.24.61-62, Kenya, Ngong River, tributary of Nairobi River, 116'28"S, 36048'47"E (2:

123.6-146.9), MCZ 32518, Kenya, Riara River, elev. 5700 ft., 110'01"S, 36049'19"E (1: 96.5),

USNM 72922, Kenya, Nairobi R., near Nairobi, 116'28"S, 36048'47"E (2: 113.4-159.4).
















































Figure 4-3. Amphilius krefftii, BMNH, 1909.10.19.26-27, Syntype 86.0 mm SL; lateral, dorsal
and ventral view. Photograph by Mark Allen.













51










-20E 0 E20E 40 60E
40t W S 40t







20E [20






OE O






-201 '"20L

-___ -____-1-




-20E OE 20E 40E 60E
km
0 5001000

Figure 1-1. Map of Africa with the east-central region indicated by the red box.









CHAPTER 1
INTRODUCTION

Natural History

The catfish family Amphiliidae is widely distributed throughout sub-Saharan Africa and

it includes 12 genera and about 66 species (Ferraris, 2007). Most amphiliid species are found in

fast-flowing water of clear, rocky streams at high elevations or rapids of large lowland rivers

(Berra, 2001; Roberts, 2003). Recent workers have recognized three subfamilies within

Amphiliidae, with Amphilius and Paramphilius placed in the subfamily Amphiliinae. Twenty

species are currently recognized in the genus Amphilius, with most of the species occurring in the

drainages of western Africa and the Congo basin.

Two species of Amphilius are currently recognized in east-central Africa, an area herein

defined as the area from and including the Ewaso Nyiro drainage in Kenya south to and

including the Rufiji drainage in Tanzania (Fig. 1-1). This area generally corresponds to the east

coast ichthyofaunal province of Roberts (1975). Roberts' ichthyofaunal province also includes

the Lake Victoria basin and all eastern-flowing rivers south to but not including the Zambezi

River. Amphilius has been reported from the Yala River, a tributary of Lake Victoria

(Whitehead, 1958; 1959) and the Ruvuma drainage, the next large drainage south of the Rufiji

(Tweddle, 1983), but these records are not supported by vouchers.

Species of the genus Amphilius are habitat specialists and are adapted for fast flow and

rocky habitats (Skelton, 1986). Morphological adaptations for these habitats include expanded

pectoral and pelvic fins with an expanded first ray, depressed body, upwardly directed eyes and

reduced gas bladder (Skelton, 1986; Walsh et al., 2000). The large pelvic fins have been

observed to form a feeble sucking disc, which in conjunction with the body, enable the fish to

cling to rocks (Jackson, 1961b). The epidermal microstructure of the anterior ray of the pectoral


















































Figure 4-5. Amphilius oxyrhinus, BMNH 1912.3.22.120, Holotype, 166.0 mm SL; lateral, dorsal
and ventral view. Photograph by Mark Allen.











53









The goals of this study are to examine morphological variation in Amphilus in east-

central Africa and to recognize valid taxa. In the process, the identities of the five nominal

species ofAmphilius described from east-central Africa will be determined.









In 1911, Boulenger published volume two of his "Catalogue of Freshwater Fishes of

Africa" and recognized four species in the drainages of east-central Africa: A. uranoscopus, A.

platychir, A. grandis, and A. krefftii, a new species that he described from the Sigi River,

Tanzania. Amphilius krefftii, like A. grandis, was distinguished from the other species of

Amphilius by relative proportions of it head, barbel and caudal peduncle lengths. Soon

thereafter, he described a fifth species, A. oxyrhinus, from the Eusso Mara, a tributary of the

Eusso Nyiro in Kenya. He distinguished this species from only A. grandis, the only other

species of Amphilius he recorded from the Eusso Nyiro.

Since Boulenger's description of Amphilius grandis, no new species of Amphilius have

been described from east-central Africa and there has been limited study of the genus in east-

central Africa. Most workers continued to recognize all of the species that Boulenger recognized

(Copley, 1941; Harry, 1953; Whitehead, 1958), but Bailey (1969: 192) questioned the validity of

A. krefftii and stated that it is "very probably a synonym of A. grandis". Meanwhile, A. grandis

was reported from throughout eastern and southern Africa (Van der Horst, 1931; Poll, 1952;

Crass, 1960; Jubb, 1961; Crass, 1964).

Amphiliusplatychir was also being reported from these areas (Ricardo, 1939a, 1939b;

Jackson, 1959; Maar, 1960; Jackson, 1961a; Jubb, 1963; Bell-Cross, 1972). Bell-Cross & Jubb

(1973) referred records of A. grandis from eastern and southern Africa to A. platychir, and A.

platychir became accepted as being widespread throughout eastern and southern Africa.

Skelton (1984) revised the genus Amphilius from eastern and southern Africa and

identified two groups within Amphilius. One group is primarily western African, has a

crenellated epidermal fold at the base of the caudal fin and 6 + 7 principal caudal-fin rays. The

second group is primarily distributed in eastern and southern Africa, lacks the crenellated









CHAPTER 2
MATERIALS AND METHODS

Use of the terms origin and insertion to designate, respectively, anterior and posterior

base of all fins follows Cailliet et al. (1986). Counts and measurements were made on the left

side of specimens when possible and follow Skelton (1981, 1984, 1986), except head length was

measured as the distance from the top of snout to the dorsal edge of the opercular cavity. In

addition, the following measurements were made:

* Body depth at anus: depth of the body at the anus.

* Postorbital length: distance from the posterior margin of the orbit to the dorsal edge of the
opercular cavity.

* Prepectoral length: distance from the origin of the pectoral fin to the tip of the snout.

* Preanal length: distance from the origin of the anal fin to the tip of the snout.

* Dorsal-fin base length: distance from the structural base of the first ray to the point where the
membrane behind the last ray contacts the body.

* Adipose-fin base length: distance from the origin of the adipose fin to the posteriormost
connection of the adipose fin to the body.

* Pelvic- and pectoral-fin lengths: distance from the origin of the fin to the tip of the fin.

* Anus to anal-fin length: distance from the origin of the anal fin to the center of the anus.

* Prepelvic length: distance from the origin of the pelvic fin to the tip of the snout.

* Postpelvic length: distance from the origin of the pelvic fin to the base of the caudal fin.

* Distance from insertion of dorsal fin to insertion of adipose fin (end dorsal end adipose):
distance from the point where the membrane behind the last dorsal-fin ray contacts the body
to the posteriormost connection of the adipose fin to the body.

* Distance from origin of dorsal fin to base of caudal fin (front dorsal caudal): distance from
the origin of the dorsal fin to the base of the caudal fin.

* Preanus length: distance from tip of the snout to opening of the anus.









Coloration

Dorsal and lateral surfaces of head and body brown with many small black spots. Ventral

region of head dark yellow or brown. Ventral region of body brown to dark yellow. Light dorsal

saddles absent. Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and

pelvic fins brown with light distal edge, ventral surfaces light yellow. Small black spots present

on all fins. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band

at base. Large juvenile coloration similar to that of adults.

Distribution

Known only from the Galana River drainage, Kenya.









D e scrip tio n ................................................................3 6
C o lo ratio n ......... ...... ...................................................... ................................3 8
D istrib u tion ......... ...... ... ................. ..........................................3 8
A mphilius sp. 1 n. sp. ...................................................................39
D ia g n o sis .................................................................................................................... 3 9
D e scrip tio n ................................................................4 0
C oloration ................................ ......... .. ..................... ............................. 4 2
Distribution ............... .. ... ........................ ...............42

5 R E S U L T S ..........................................................................5 9

L IST O F R E F E R E N C E S ...................... ................ .....................................................67

B IO G R A PH IC A L SK E T C H .................................................................................................... 73










301E


-5E




-10Et


30E 35E 40E
km
0 200 400
Figure 4-4. Known distribution ofAmphilius krefftii. Some symbols represent more than one
collection site. T denotes type locality.


40F

































2007 Alfred William Thomson









lower jaw, extending to origin of pectoral-fin. Inner mandibular barbel originates anterolaterally

of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal

membrane with 7 (10) or 8 (9) rays. Gill rakers on first epibranchial 2 (7) or 3 (12); rakers on

first ceratobranchial 4 (1), 5(3), 6 (6), 7 (8) or 8 (1); total gill rakers on first arch 6 (1), 7 (2), 8

(2), 9 (8), 10 (5) or 11 (1).

Eye small, positioned dorsolaterally approximately midway between tip of snout and

posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter.

Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and

posterior nares with prominent tubular rims; nares separate but relatively close to each other.

Posterior nare located about midway between eye and tip of snout.

Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (18) or i,7 (1) rays,

and fin margin straight. Pectoral fin with i,9 (3) or i,10 (16) rays with first ray unbranched and

greatly thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays

progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted

posteriorly to dorsal-fin base. Pelvic fin with i,5 (19) rays with first ray unbranched and greatly

thickened. Pelvic fin with straight posterior margin.

Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base,

extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded

edge, not deeply incised posteriorly. Caudal fin deeply emarginate or forked with tips of lobes

pointed, with i,7,8,i (18) or i,8,9,i (1) principal rays. Anal fin with short base, origin posterior to

origin of adipose-fin base, with iii,6 (19) rays. Anal fin margin almost straight.

Coloration

Dorsal and lateral surfaces of head and body brown. Ventral region of head dark yellow.

Ventral region of body brown to dark yellow. Light dorsal saddles absent. Dorsal, adipose,

















































Figure 5-1. Amphilius uranoscopus (juveniles) from the Wami (top: CAS 80494, 49.1 mm SL),
Ruvu (middle: UF 84882, 40.4 mm SL) and Rufiji (bottom: FMNH 111680, 46.0 mm
SL) drainage.









66









fin as observed by a scanning electron microscope (Bell-Cross & Jubb, 1973) is similar to the

epidermal microstructure of the adhesive organs in Asian sisorid catfishes (Singh & Agarwal,

1991; Das & Nag, 2004, 2005).

In Amphilius species, males and females are generally similar, but the adductor

mandibulae muscles attached to the head are sometimes greatly enlarged in mature males (J.

Friel, Pers. Comm. 2007). This dimorphism has also been observed in species of Paramphilius,

and was believed to be a derived character of Paramphilius by Skelton (1989).

Little information is available on the biology ofAmphilius species. Skelton (2001) stated

that South African species breed during the summer. Marriott et al. (1997) studied the

reproductive biology of A. natalensis in South Africa and concluded that the species had a long

breeding period extending from August to February. They observed that breeding coincided

with the rainy season and postulated that spawning was initiated in response to increased water

flow rate or changes in water quality following periods of rainfall (Marriott et al., 1997). Walsh

et al. (2000) were not able to determine the breeding season ofA. jacksonii, but suggested that it

may have two breeding periods per year.

The diet of Amphilius species has been reported to predominantly consist of benthic

aquatic insects (Marriott et al., 1997; Skelton, 2001; Walsh et al., 2000). Marriott et al. (1997)

conducted a detailed stomach analysis of specimens ofA. natalensis and found the diet to consist

mostly of larval chironomids and ephemeropterans. Additionally, they found that small fishes

fed predominantly on chironomid larvae, while in larger fishes, chironomid and ephemeropteran

larvae contributed almost equally in volume. Walsh et al. (2000) analyzed the stomach contents

of 118 specimens ofA. jacksonii, and similarly found the diet of the species to consist mainly of

larval Diptera, Ephemeroptera and Trichoptera.































Figure 4-7. Amphilius uranoscopus, CAS 80494, 101.4 mm SL; lateral, dorsal and ventral view.


B~ '"









LIST OF REFERENCES


BAILEY, R. G. 1969. The non-cichlid fishes of the eastward flowing rivers of Tanzania, East
Africa. Revue de Zoologie et Botanique Africaines 80:170-199.

BELL-CROSS, G. 1972. The fish fauna of the Zambezi River system. Arnoldia (Rhodesia) 5:1-19.

BELL-CROSS, G., AND R. A. JUBB, 1973. The Amphiliidae of southern Africa and record of
Amphilius lampei Pietschmann, 1913, from the Inyanga Mountains, Rhodesia. Aronoldia
(Rhodesia) 6:1-9.

BERNACEK, G. M. 1980. Introduction to the freshwater fishes of Tanzania. University of Dar es
Salaam, Deptment of Zoology, Dar es Salaam.

BERRA, T. M. 2001. Freshwater Fish Distribution. Academic Press, San Diego, California.

BERTIN, L., AND R. ESTEVE. 1950. Catalogue des Types de Poissons du Museum National
d'Histoire Naturelle. 5e partie.- Ostariophysaires (Siluriformes). Imprimerie Nationale,
Paris.

BOULENGER, G. A. 1898a. On the habit of the siluroid fish Anoplopterusplatychir, Gthr. Annals
and Magazine of Natural History (Ser. 7) 8:447-448.

BOULENGER, G. A. 1898b. Report on the collection of fishes made by Mr. J. E. S. Moore in Lake
Tanganyika during his expedition, 1895-96. Transactions of the Zoological Society of
London 15:1-30, pls. 1-8.

BOULENGER, G. A. 1901. Description of a new silurid fish of the genus Anoplopterus, from
Cameroon. Annals and Magazine of Natural History (Ser. 7) 8: 447-448.

BOULENGER, G. A. 1902. Additions a la faune ichthyologique de bassin du Congo. Materiaux
pour la faune du Congo. Annales du Musee du Congo, Zoologie 2:19-57, pls. 7-16.

BOULENGER, G. A. 1905a. On a second collection of fishes made by Mr. S. L. Hinde in the
Kenya District, East Africa. Proceedings of the Zoological Society of London 1:62-64,
pl. 7.

BOULENGER, G. A. 1905b. A list of freshwaters fishes of Africa. Annals and Magazine of
Natural History (Ser. 7) 16: 36-60.

BOULENGER, G. A. 1907a. Descriptions of three new fishes from Central Africa. Annals and
Magazine of Natural History (Ser. 7) 20: 487-489.

BOULENGER, G. A. 1907b. On a collection of fishes, batrachians and reptiles, made by Mr. S. A.
Neave in Rhodesia, north of the Zambesi with field notes by the collector. Memoirs and
Proceedings of the Manchester Literary & Philosophical Society 51: 1-12.










301E


-5E




-10Et


30E 35E 40E
km
0 200 400
Figure 4-8. Known distribution ofAmphilius uranoscopus. Some symbols represent more than
one collection site.


40F









Table 5-1. Branchiostegal ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus
Branchiostegal Rays 6 7 8 9
A. oxyrhinus 4 9 2
A. grandis 9 8
Amphilius sp. 1 1 21
A. krefftii 18 1
A. uranoscopus (Wami) 3
A. uranoscopus (Ruvu) 6 5
A. uranoscopus (Rufiji) 17 9

Table 5-2. Total gill raker counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and
three populations of A. uranoscopus
Total Gill Rakers 6 7 8 9 10 11
A. oxyrhinus 1 3 10
A. grandis 1 2 2 8 7 1
Amphilius sp. 1 1 7 11 2
A. krefftii 5 5 6 1
A. uranoscopus (Wami) 1 1 1
A. uranoscopus (Ruvu) 1 4 3 3
A. uranoscopus (Rufiji) 6 23 8 1

Table 5-3. Branched pectoral-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A.
krefftii and three populations of A. uranoscopus
Branched Pectoral-Fin Rays 8 9 10 11
A. oxyrhinus 12 2
A. grandis 3 17 1
Amphilius sp. 1 12 9
A. krefftii 11 7 1
A. uranoscopus (Wami) 2 1
A. uranoscopus (Ruvu) 8 4
A. uranoscopus (Rufiji) 3 14 23

Table 5-4. Branched anal-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii
and three populations of A. uranoscopus
Branched Anal-Fin Rays 5 6 7 8
A. oxyrhinus 14
A. grandis 21
Amphilius sp. 1 7 13 2
A. krefftii 17 2
A. uranoscopus (Wami) 1 2
A. uranoscopus (Ruvu) 6 6
A. uranoscopus (Rufiji) 2 17 10 11









HARRY, R. R. 1953. A contribution to the classification of the African catfishes of the family
Amphiliidae, with description of collections from Cameroon. Revue de Zoologie et de
Botanique Africaines 47: 177-200; 201-232.

HALDEMANN, E. G. 1962. The geology of the Rufiji Basin, with reference to proposed dam sites
in eastern and southern Tanganyika. Bulletin Geological Survey of Tanganyika 33:1-
250.

HILGENDORF, F. M. 1905. Fische von Deutsch und Englisch Ost-Africa. Zoologische Jahrbucher.
Abteilung fur Systematik, Geographie und Biologie der Tiere (Jena) 22:405-420.

JACKSON, P. B. N. 1959. New records and little-known species offish from Rhodesia and
Nyasaland. Occasional papers of the National Museums of Southern Rhodesia, Natural
Sciences 3:295-305.

JACKSON, P. B. N. 1961a. Check-list of the fishes of Nyasaland. Occasional Papers of the
National Museums of Southern Rhodesia, Natural sciences 3:535-621.

JACKSON, P. B. N. 1961b. The fishes of Northern Rhodesia: a checklist if indigenous species.
Lusaka: The Government Printer.

JUBB, R. A. 1961. An illustrated guide to the freshwater fishes of the Zambezi River, Lake
Kariba, Pungwe, Subi, Lundi and Limpopo Rivers. Bulawayo: Stuart Manning.

JUBB, R. A. 1963. A revised list of the freshwater fishes of southern Africa. Annals of the Cape
Provincial Museums (Natural History) 3:40-43.

MAAR, A. 1960. Introductory check list of fish of the Ethiopian region. Proceedings of the
Federal Science Congress 1:1-8.

MARRIOTT, M.S., A. J. BOOTH, AND P. H. SKELTON. 1997. Reproductive and feeding biology of
the Natal mountain catfish Amphilius natalensis (Siluroidei: Amphiliidae).
Environmental Biology of Fishes 49:461-470.

MATTHES, H. 1967. The fishes and fisheries of the Ruaha River basin, Tanzania. East African
Freshwater Fisheries Research Organisation (EAFFO) Occasional Papers No. 9:1-19.

PELLEGRIN, J. 1905. Mission scientifique de Ch. Alluaud en Afrique orientale (Juin 1903 -- Mai
1904). Poissons. II. Systematique. Memoires de la Societe Zoologie de France 17: 167-
185, pl. 16.

PELLEGRIN, J. 1936. Contribution a l'Ichtyologie de l'Angola. Arquivos do Museu Bocage,
Lisboa 7:45-62.

PFEFFER, G. J. 1889. Ubersicht der von Herrn Dr. Franz Stuhlmann in Agypten, auf Sanzibar und
dem gegeniberliegenden Festlande gesammelten Reptilien, Amphibien, Fische,









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 thank Dr. Lawrence Page for

directing me on this topic of study and for instruction, and for his constructive criticism and

guidance. I also thank Drs. Jon Armbruster and Rebecca Kimball for their contributions and

guidance. Mark Allen kindly provided photographs for figures 4-3 and 4-5. For specimen loans

and access to institutional specimens I thank Ralf Britz and James Maclain (BMNH), William N.

Eschmeyer, Tomio Iwamoto and David Catania (CAS), Mary A. Rogers (FMNH), Karsten

Hartel (MCZ), Guy Duhamel and Patrice Pruvost (MNHN), Jos Snoeks and Mark Hanssens

(MRAC), Robert H. Robins (UF) and Jeffery T. Williams (USNM).


















































Figure 4-9. Amphilius sp. 1, BMNH 1910.10.31.31, Holotype, 150.0 mm SL; lateral, dorsal and
ventral view.














57









Table 4-6. Morphometric data for Amphilius sp. 1. Range and mean include the holotype.

Holotype Range (n=21) MeanSD
%SL
Head length 26.9 24.0-29.3 25.81.2
Head width 22.7 19.7-26.2 21.91.4
Head height 11.3 11.3-14.9 12.71.0
Body depth 13.0 12.4-17.3 14.81.4
Body depth at anus 11.8 11.1-15.3 13.31.1
Predorsal length 37.9 33.3-39.6 37.21.5
Prepectoral length 24.1 19.5-25.6 22.01.7
Preanal length 79.6 60.8-79.6 73.25.2
Dorsal-fin base length 9.2 8.0-11.7 9.50.9
Adipose-fin base length 18.7 18.5-24.3 20.51.8
Anal-fin base length 8.7 8.6-16.6 10.41.8
Pelvic-fin length 16.1 16.0-19.7 17.71.2
Pectoral-fin length 16.1 17.4-23.4 19.71.5
Anal-fin length 18.7 14.5-19.2 17.01.4
Caudal peduncle length 15.7 14.6-18.3 16.51.1
Caudal peduncle depth 15.8 10.1-12.6 11.00.7
Anus to anal fin length 13.0 10.4-14.1 12.1+0.9
Prepelvic length 56.1 50.9-58.3 54.21.9
Postpelvic length 49.9 42.8-51.2 47.22.3
End of dorsal to adipose 45.3 42.3-49.9 45.21.7
Front of dorsal to caudal 64.9 62.4-67.6 65.41.1
Preanus length 66.3 58.1-66.3 62.02.0
%HL
Snout length 53.8 46.7-56.6 52.52.3
Interorbital distance 26.8 23.9-37.0 28.73.1
Maxillary barbel length 68.1 60.4-90.6 75.07.7
Inner mandibular barbel length 37.7 26.9-43.9 36.84.2
Outer mandibular barbel length 61.7 49.3-71.9 59.07.0
Eye diameter 9.0 9.0-15.3 11.4+2.0
Postorbit length 36.6 35.6-43.1 39.82.2









SKELTON, P. H. 1976. Preliminary observations on the relationships of Barbus species from Cape
coastal rivers, South Africa (Cypriniformes: Cyprinidae). Zoologie Africaine 11: 399-
411.

SKELTON, P. H. 1981. The description and osteology of a new species of Gephyroglanis
(Siluriformes, Bagridae) from the Olifants River, South West Cape, South Africa. Annals
of the Cape Provincial Museums, Natural History 13:217-249.

SKELTON, P. H. 1984. A systematic revision of species of the catfish genus Amphilius
(Siluroidei, Amphiliidae) from east and southern Africa. Annals of the Cape Provincial
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SKELTON, P. H. 1986. Two new Amphilius (Pisces, Siluroidei, Amphiliidae) from the Zaire River
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48:1-13.

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Bulletin du Museum d'Histoire Naturelle, Paris 3:81-84.

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14:246-250.

WALSH, S. J., L. J. CHAPMAN, A. E. ROSENBERGER, AND C. A. CHAPMAN. 2000. Redescription of
Amphiliusjacksonii (Siluriformes: Amphiliidae) with habitat and life-history notes.
Ichthyological Exploration of Freshwaters 11:163-174.

WEINELT, M. 1996-2006. Online Map Creation. http://www.aquarius.ifm-geomar.de/. (July,
2007).









rakers on first ceratobranchial 4 (1), 5(3) or 6 (9); total gill rakers on first arch 7 (1), 8 (3) or 9

(9).

Eyes small, positioned dorsolaterally approximately midway between tip of snout and

posterior margins of operculum. Horizontal diameter of eye slightly wider than vertical

diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head.

Anterior and posterior nares with prominent tubular rims; nares separate but relatively close to

each other. Posterior nare located about midway between eye and tip of snout.

Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (13) rays, and fin

margin straight. Pectoral fin with i, 10 (11) or i, 11 (2) rays with first ray unbranched and greatly

thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays

progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted

posteriorly to dorsal-fin base. Pelvic fin with i,5 (13) rays with first ray unbranched and greatly

thickened. Pelvic fin with straight posterior margin.

Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base,

extending over posterior end of anal-fin base. Margin strongly convex with sharply rounded

edge, not deeply incised posteriorly. Caudal fin forked with tips of lobes pointed, with i,7,7,i (1)

or i,7,8,i (11) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin

base, with iii,6 (12) rays. Anal fin margin almost straight.

Coloration

Dorsal and lateral surfaces of head and body brown. Ventral and lateral regions of body

covered with large black spots or blotches. Ventral region of head and body dark yellow. Light

dorsal saddles absent, but pale regions sometimes present in front of dorsal and anal-fin bases.

Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown

with light distal edge, ventral surfaces light yellow. Medial band of dark pigment on all rayed









LIST OF FIGURES


Figure page

1-1 Map of Africa with the east-central region indicated by the red box................................ 16

1-2 Map of Africa indicating type localities of species of the Amphilius uranoscopus group
as defined by Skelton (1984). ................................................ ................................ 17

4-1 Amphilius grandis, BMNH 1912.3.22.119, 113.6 mm SL; lateral, dorsal and ventral
v iew .................................................................. ...........................4 9

4-2 Known distribution of Amphilius grandis. Some symbols represent more than one
collection site. T denotes type locality. ........................................ ......................... 50

4-3 Amphilius krefftii, BMNH, 1909.10.19.26-27, Syntype 86.0 mm SL; lateral, dorsal and
ventral view. Photograph by M ark Allen ......... ... .......... .. .................. .............. 51

4-4 Known distribution of Amphilius krefftii. Some symbols represent more than one
collection site. T denotes type locality. ........................................ ......................... 52

4-5 Amphilius oxyrhinus, BMNH 1912.3.22.120, Holotype, 166.0 mm SL; lateral, dorsal and
ventral view. Photograph by Mark Allen..................................................53

4-6 Known distribution of Amphilius oxyrhinus. Some symbols represent more than one
collection site. T denotes type locality. ........................................ ......................... 54

4-7 Amphilius uranoscopus, CAS 80494, 101.4 mm SL; lateral, dorsal and ventral view...........55

4-8 Known distribution of Amphilius uranoscopus. Some symbols represent more than one
collection site. .............................................................................56

4-9 Amphilius sp. 1, BMNH 1910.10.31.31, Holotype, 150.0 mm SL; lateral, dorsal and
ventral view ............... ..... ......... ............................... ... .. ....... .. 57

4-10 Known distribution of Amphilius sp. 1. Some symbols represent more than one
collection site. T denotes type locality. ........................................ ......................... 58

5-1 Amphilius uranoscopus (juveniles) from the Wami (top: CAS 80494, 49.1 mm SL),
Ruvu (middle: UF 84882, 40.4 mm SL) and Rufiji (bottom: FMNH 111680, 46.0
m m SL ) drain ag e .................................................... ................. 66









CHAPTER 4
SYSTEMATIC ACCOUNTS

Amphilius uranoscopus group

All species of Amphilius in east-central Africa belong to the Amphilius uranoscopus

group and are recognized by the following unique combination of characters: absence of a

crenellated epidermal fold at the base of the caudal fin, usually 8+9 (i,7,8,i) principal caudal fin

rays, small bilateral bony swimbladder capsules, 36-39 total and 20-23 caudal vertebrae, and

leading pterygiophore of the dorsal fin intercepting the vertebral column at the first, second or

third post-Weberian vertebrae. Skelton (1984) recognized this species group as a single species,

A. uranoscopus, but did not formally diagnose it.

Externally, species of the A. uranoscopus group are similar to A. natalensis, A. lampei

and A. kivuensis but these species differ by having the leading pterygiophore of the dorsal fin

intercepting the vertebral column after the third post-Weberian vertebrae. Amphilius

cryptobullatus is also similar externally, but differs from A. uranoscopus in the development and

extremely large size of its bilateral bony swimbladder capsules. Following are diagnoses and

descriptions of valid species of the Amphilius uranoscopus group in east-central Africa. Species

are arranged alpabetically. Diagnostic traits are summarized in Table 4-1.









fins. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band at

base Juvenile coloration: Similar to that of adults, except all fins lighter brown, with medial

bands of dark pigment all rayed fins. Pectoral and pelvic fins with first unbranched ray dark

brown dorsally.

Distribution

Tana and Eusso Nyiro (Ewaso Ng'iro) River drainages, Kenya.

Amphilius uranoscopus (Pfeffer 1889)

(Figures 4-7 and 4-8; Table 4-5)

Anoplopterus uranoscopus Pfeffer 1889: 16. Type locality: Ushonda and Monda, upper ranges of
Wami R., Tanzania [Wami River drainage]; Pfeffer, 1893: 161, [Wami drainage]; Pfeffer,
1896: 33, fig. 14, [Wami drainage]; Boulenger, 1898a: 255, reference to original
description; Boulenger, 1901: 447, diagnosis; Poche, 1902a: 21; Poche, 1902b: 211.

Chimarrhoglanis leroyi Vaillant, 1897: 82, Original description, Type locality: Zanquebar,
torrent de Mroboro, 600 meters altitude [Ruvu drainage]; Boulenger, 1898a: 254, as a
synonym ofAmphiliusplatychir; Boulenger, 1898b: 4 [Ruvu drainage]; Poche, 1902a:
121; Poche, 1902b: 211; Pellegrin, 1905a: 177, type information; Pellegrin 1936: 57,
similarity to Amphilius platychir var. cubangoensis; Harry, 1953: 186, in synonymy of
Amphiliusplatychir; Bertin & Esteve, 1950: 35, information on type; Skelton, 1984: 45,
in synonymy of Amphilius uranoscopus.

Amphilius uranoscopus Boulenger, 1902: 41, in key; Boulenger, 1905b: 48, [Wami drainage];
Boulenger 1911: 253, 354, fig. 277, Description, [Wami drainage]; Harry, 1953: 187,
synonymy; Copley, 1958: 99 [Wami drainage]; Bailey, 1969: 192 [Wami drainage];
Bernacek, 1980: 36, [Wami, Ruvu and Rufiji drainages]; Skelton, 1984: 41, information
on types; Skelton, 1994: 126 [Wami, Ruvu and Rufiji drainages]; Seegers, 1996a: 192,
figs. 136-137, information on types.

Pimelodus (Amphilius) uranoscopus (non Pfeffer) Hilgendorf, 1905: 411 (in part), [Rufiji
drainage].

Amphiliusplatychir (non Giinther) Matthes, 1967: 5, 15 [Rufiji drainage]; Bell-Cross, 1972: 18,
table 7, [Rufiji drainage].

Amphilius grandis (non Boulenger) Bailey, 1969: 192 (in part) [Ruvu drainage].










WHITEHEAD, P. J. P. 1958. A new species of Chiloglanis (Pisces, Mochocidae) in Kenya. Annals
and Magazine of Natural History (Ser. 13) 1:197-208.

WHITEHEAD, P. J. P. 1959. The anadromous fishes of Lake Victoria. Revue de Zoologie et
Botanique Africaines 59:329-363.

WILLIS, B. 1936. East African plateaus and rift valleys. Studies in Comparative Seismology,
Carnegie Institute of Washington Publication 470:1-358.









36 (11), 37 (14) or 38 (2). Preanal vertebrae 19 (2), 20 (7), 21 (14) or 22 (9). Postanal vertebra

15 (9), 16 (19) or 17 (1). First dorsal pterygiophore intercept count 1 (30) or 2 (1).

Head and anterior part of body depressed and broad. Head wedge shaped in lateral view.

Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to

pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped

connection. Membrane connected by a small frenum at midline, but with shallow groove at

posterior margin.

Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe

large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed.

Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary

teeth short and conical, tooth patches forming U-shaped band, separated medially.

Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and

flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle of

pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of

lower jaw, extending middle pectoral-fin base. Inner mandibular barbel originates anterolaterally

of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal

membrane with 8 (25) or 9 (29) rays. Gill rakers on first epibranchial 2 (35), 3 (18) or 4(1);

rakers on first ceratobranchial 4 (9), 5 (26), 6 (14), 7 (4) or 8 (1); total gill rakers on first arch 6

(5), 7 (24), 8 (15), 9 (5), 10 (4) or 11 (1).

Eyes small, positioned dorsolaterally approximately midway between tip of snout and

posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter.

Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and









BIOGRAPHICAL SKETCH

Alfred W. Thomson was born December 30, 1978, in Englewood, New Jersey. One of

three children, he was interested in the natural world from an early age and has always been

especially interested in fishes. He received his bachelor of science degree from Mansfield

University in August 2001, and moved to Kenosha, Wisconsin to work as a research technician

for the Illinois Natural History Survey at the Lake Michigan Biological Station in Zion, Illinois.

In January 2004 he moved to Gainesville, Florida and worked as a research technician on the All

Catfish Species Inventory Project at the Florida Museum of Natural History. In August 2005 he

enrolled in UF's Department of Zoology as a full-time graduate student with a research

assistantship from the All Catfish Species Inventory Project. He is currently continuing his

research on catfish systematics as a Ph.D. student in UF's Department of Zoology.










Table 4-4. Morphometric data for Amphilius oxyrhinus. Range and mean include the holotype.
Holotype Range (n=14) MeanSD
%SL
Head length 25.5 23.1-27.0 25.51.1
Head width 22.0 19.5-23.9 21.51.1
Head height 11.8 11.3-14.6 12.70.9
Body depth 13.2 9.6-15.2 12.91.7
Body depth at anus 11.6 11.0-14.2 12.91.0
Predorsal length 38.5 35.2-40.3 36.71.5
Prepectoral length 23.4 17.3-23.4 20.61.8
Preanal length 75.4 70.6-76.1 74.1+1.8
Dorsal-fin base length 8.4 8.4-12.7 10.91.2
Adipose-fin base length 15.6 15.6-26.0 21.52.9
Anal-fin base length 9.3 9.3-15.0 11.41.5
Pelvic-fin length 14.9 14.9-22.1 19.91.7
Pectoral-fin length 17.8 17.8-26.2 22.92.0
Anal-fin length 15.8 15.8-19.5 18.01.1
Caudal peduncle length 17.0 14.8-18.0 16.90.9
Caudal peduncle depth 9.0 9.0-12.1 10.90.9
Anus to anal fin length 13.0 9.5-14.1 11.71.3
Prepelvic length 55.6 49.9-57.4 53.02.0
Postpelvic length 46.0 46.0-52.1 49.1+2.0
End of dorsal to adipose 42.7 41.1-48.6 44.22.1
Front of dorsal to caudal 63.3 61.5-68.3 64.72.4
Preanus length 63.0 59.8-66.0 61.81.7

%HL
Snout length 53.2 47.4-54.5 51.1+2.5
Interorbital distance 23.0 23.0-33.3 28.03.1
Maxillary barbel length 51.9 51.9-85.5 73.88.5
Inner mandibular barbel length 31.3 21.1-59.1 34.08.7
Outer mandibular barbel length 42.7 30.4-64.2 51.810.7
Eye diameter 8.1 8.1-16.5 13.52.6
Postorbit length 38.9 33.4-42.3 38.52.6










-20E OE 20E 40E 60E








20E 20E
4 0 1E --.-,-.---- .'-'., -, .- i ". a 4 01E





















.. pIltychir
-20E -ignensi-20L
.m .'orvrhmu.




A.". h ref/i t




A' breviiorsatis


-20E 0E 20E 40E 60E
km
0 5001000

Figure 1-2. Map of Africa indicating type localities of species of the Amphilius uranoscopus
group as defined by Skelton (1984).









dorsal saddles present at the base of the dorsal and adipose fin; branchiostegal rays usually

seven.

The presence of a visible epidermal core extending posteriorly from the bases of the

inner and outer mandibular barbels distinguishes Amphilius krefftii from A. grandis, Amphilius

sp. 1 and A. uranoscopus. Amphilius krefftii is distinguished from A. oxyrhinus, A. grandis and

Amphilius sp. 1 by the shape of the caudal fin emarginatee in A. krefftii vs. deeply emarginate or

forked in A. grandis and forked in A. oxyrhinus and Amphilius sp. 1) and the presence of light

dorsal saddles at the base of the dorsal and adipose fin (vs. light dorsal saddles absent).

Additionally the caudal-fin lobes are rounded in A. krefftii (vs. pointed in A. grandis and A.

oxyrhinus). Amphilius krefftii is further distinguished from Amphilius sp. 1 by the absence of

dark spots or blotches on the head, body or fins in adults and further distinguished from A.

uranoscopus by having usually seven branchiostegal rays (vs. eight or nine).

Description

Morphometric data as in Table 4-3. Body elongate, ventral profile flattened ventrally to

anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from

tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest

body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated

epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin,

closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line

complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae

36 (7), 37 (3) or 38 (1). Preanal vertebrae 20 (7) or 21 (5). Postanal vertebra 15 (1), 16 (9) or 17

(1). First dorsal pterygiophore intercept count 1 (11) or 2 (1).

Head and anterior part of body depressed and broad. Head wedge shaped in lateral view.

Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to









BOULENGER, G. A. 1911. Catalogue of the fresh-water fishes of Africa in the British Museum
(Natural History), Vol. 2. British Museum (Natural History) Trustees, London.

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East Africa to the East of Lake Baringo. Proceedings of the Zoological Society of
London 1912:672-676, pls. 78-80.

BOULENGER, G. A. 1916. Catalogue of the fresh-water fishes of Africa in the British Museum
(Natural History), Addenda. British Museum (Natural History) Trustees, London.

CAILLIET, G. M., M. S. LOVE AND A. W. EBELING. 1986. Fishes: a field and laboratory manual
on their structure, identification, and natural history. Wadsworth Publishing Company,
Belmont, CA.

COPLEY, H. 1941. A short account of the freshwater fishes of Kenya. Journal of the East Africa
and Uganda Natural History Society 16:24 pp., 8 pls.

COPLEY, H. 1958. Common freshwater fishes of east Africa. H. F. & G. Witherby Ltd., London.

CRASS, R. S. 1960. Notes on the freshwater fishes of Natal with descriptions of four new species.
Annals of the Natal Museum 14: 405-458.

CRASS, R. S. 1964. Freshwater fishes of Natal. Pietermaritzburg: Shuter & Shooter.

DAS, D., AND T. C. NAG. 2004. Adhesion by paired pectoral and pelvic fins in a mountain-
stream catfish, Pseudocheneis sulcatus (Sisoridae). Environmental Biology of Fishes
71:1-5.

DAS, D., AND T. C. NAG. 2005. Structure of adhesive organ of the mountain-stream catfish,
Pseudocheneis [sic.] sulcatus (Teleostei: Sisoridae). Acta Zoologica 86:231-237.

FERRARIS, C. J., Jr. 2007. Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes),
and catalogue of siluriform primary types. Zootaxa 1418:1-628.

GONTHER, A. 1864. Catalogue of the Fishes in the British Museum, vol. 5.- Catalogue of the
Physostomi, Containing the Families Siluridae, Characinidae, Haplochitonidae,
Sternoptychidae, Scopelidae, Stomiatidae in the Collection of the British Museum.
Trustees, London.

GUNTHER, A. 1865. Pisces. The record of zoological literature. Zoological Record 1864:133-188.

GUNTHER, A. 1902. Last account of fishes collected by Mr. R. B. N. Walker, C. M. Z. S., on the
Gold Coast. Proceeding of the Zoological Society of London 2:330-339.










Table 4-3. Morphometric data for Amphilius krefftii. Range and mean include the syntypes.
Syntypes (n=2) Range (n=19) MeanSD
%SL
Head length 22.7-22.8 21.9-25.9 23.81.1
Head width 20.2-20.6 18.7-23.6 21.21.1
Head height 12.6-12.8 9.1-14.7 11.71.7
Body depth 11.6-11.6 9.1-17.2 12.22.3
Body depth at anus 13.5-13.5 10.8-15.5 12.91.3
Predorsal length 34.3-34.5 31.8-36.9 34.71.5
Prepectoral length 19.8-20.8 17.4-23.9 20.1+1.8
Preanal length 72.8-73.7 62.5-76.3 72.1+3.9
Dorsal-fin base length 8.8-9.5 8.2-13.5 10.41.5
Adipose-fin base length 21.0-22.7 19.1-29.8 23.2+3.0
Anal-fin base length 10.6-10.7 8.8-15.0 11.21.5
Pelvic-fin length 19.3-19.8 17.6-20.6 19.50.9
Pectoral-fin length 19.0-21.3 19.0-24.8 21.51.4
Anal-fin length 16.2-17.9 15.4-21.7 18.62.1
Caudal peduncle length 17.3-18.6 14.9-18.6 16.51.0
Caudal peduncle depth 10.5-11.5 8.1-12.6 11.1+0.9
Anus to anal fin length 9.3-11.9 7.2-13.9 11.1+1.8
Prepelvic length 52.1-52.6 49.4-54.5 52.61.5
Postpelvic length 47.4-50.0 46.8-50.8 48.91.1
End of dorsal to adipose 46.9-50.2 42.5-50.8 47.1+2.5
Front of dorsal to caudal 65.5-67.0 63.4-71.3 66.92.0
Preanus length 61.3-63.2 59.3-64.8 62.31.7

%HL
Snout length 45.4-45.4 45.4-53.1 49.62.4
Interorbital distance 24.5-26.9 23.2-31.3 26.92.4
Maxillary barbel length 72.2-86.7 52.6-95.9 73.712.2
Inner mandibular barbel length 28.1-30.6 20.7-47.8 30.86.2
Outer mandibular barbel length 48.1-55.6 36.8-68.6 53.98.7
Eye diameter 12.8-16.7 11.8-19.5 15.41.8
Postorbit length 34.2-40.7 34.2-43.1 38.72.1









four to five innermost rays progressively shorter making inner part of posterior fin margin

rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (17) rays with first

ray unbranched and greatly thickened. Pelvic fin with straight posterior margin.

Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base,

extending over posterior end of anal-fin base. Margin strongly convex with sharply rounded

edge, not deeply incised posteriorly. Caudal fin emarginate with tips of lobes rounded, with

i,5,6,i (1) or i,7,8,i (15) principal rays. Anal fin with short base, origin posterior to origin of

adipose-fin base, with iii,6 (15) or iii7 (2) rays. Anal fin margin almost straight.

Coloration

Dorsal and lateral surfaces of head and body brown. Ventral region light brown. Light

dorsal saddles at anterior and posterior of dorsal-fin and adipose-fin bases. Dorsal, adipose,

caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown, ventral surfaces

light yellow. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped

band at base. Juvenile coloration: similar to that of adults, except all fins light yellow, with small

blotches of brown pigment on the branched rays. Pectoral and pelvic fins with first unbranched

ray dark brown dorsally.

Distribution

Lower Galina and Pangani River drainages, Kenya and Tanzania.

Amphilius oxyrhinus Boulenger 1912

(Figures 4-5 and 4-6; Table 4-4)

Amphilius oxyrhinus Boulenger 1912: 675, P1. 80, Original description, Type locality: Eusso
Mara, a swift mountain stream, Kenya [Eusso Nyiro (Ewaso Ng'iro) drainage];
Boulenger, 1916: 306, [Eusso Nyiro drainage]; Skelton, 1984: 45, in synonymy of A.
uranoscopus.

Amphilius leroyi (non Vaillant) Pellegrin, 1905: 177, Nairobi; Harry, 1953: 189 (in part),
synonymy.









Mollusken und Krebse. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten,
Hamburg 6:1-36.

PFEFFER, G. J. 1893. Ostafrikanische Fische gesammelt von Herrn Dr. F. Stuhlmann im Jahre
1888 und 1889. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Hamburg
10:131-177, 3 pls.

PFEFFER, G. J. 1896. Die Fische Ost-Afrikas, p. xviii + 72 p. In: Die Thierwelt Ost-Afrikas und
der Nachbargebiete. Vol. 1, Lief. V. K. Mobius (ed.), D. Reimer, Berlin.

POCHE, F. 1902a. Uber das Vorkommen von Anoplopterusplatychir (Gthr) in Westafrika.
Zoologischer Anzeiger 25:121-122.

POCHE, F. 1902b. Richtigstellung eines Gattungsnamens unter den Siluriden. Zoologischer
Anzeiger 25:211.

POLL, M. 1952. Poissons de rivieres de la region des lacs Tanganika et Kivu recuellis par G.
Marlier. Revue de Zoologie et de Botanique Africaines 46:221-236.

RICARDO, C. K. 1939a. The fishes of Lake Rukwa. Journal of the Linnean Society of London,
Zoology 40:625-657.

RICARDO, C. K. 1939b. Report on the fish and fisheries of Lake Rukwa in Tanganika Territory
and the Bangweulu regions in Northern Rhodesia. Crown Agent, London.

ROBERTS, T. R. 1975. Geographical distribution of African freshwater fishes. Zoological Journal
of the Linnean Society 57:249-319.

ROBERTS, T. R. 2003. Systematics and osteology of Leptoglaninae, a new subfamily of the
African catfish family Amphiliidae, with descriptions of three new genera and six new
species. Proceedings of the California Academy of Sciences 54:81-132.

SEEGERS, L. 1996a. The fishes of the Lake Rukwa drainage. Annales de la Societe Zoologique
de Belgique 278:1-407.

SEEGERS, L., 1996b. Die Amphilius-Arten Ostafrikas. Die Aquarien- und Terrarienzeitschrift
(DATZ) 49:249-255.

SEEGERS, L., L. DE VOS AND D. O. OKEYO. 2003. Annotated checklist of the freshwater fishes of
Kenya (excluding the lacustrine haplochromines from Lake Victoria). Journal of East
African Natural History 92:11-47.

SINGH, A., AND N. K. AGARWAL. 1991. SEM surface structure of the adhesive organ of the
hillstream fish Glyptothoraxpectinopterus (Teleostei: Sisoridae) from the Garhwal Hills.
Functional and Developmental Morphology 1:11-13.









Amphilius sp. 1 n. sp.

(Figures 4-9 and 4-10; Table 4-6)

Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Galina drainage] [BMNH
1937.12.11.16-18, BMNH 1969.3.24.61-62]; Copley, 1958: 98 (in part), [Galina
drainage] [BMNH 1937.12.11.16-18, BMNH 1969.3.24.61-62].

Amphilius uranoscopus Skelton, 1994: 126 (in part) [Galina drainage]; Seegers, 1996b: 255, fig.
16 (in part), Athi River; Seegers et al., 2003: 37 (in part), Athi River system.

Holotype: BMNH 1910.10.31.31, Nairobi, Kenya, 116'28"S, 36048'47"E (1: 150.0),

Paratypes: BMNH 1905.12.11.2, Kenya, Nairobi River, Kikaya, 116'28"S, 36048'47"E (1:
160.3), BMNH 1909.11.15.21-23, Nairobi River, elev. 2500 ft., 116'28"S, 36048'47"E
(3: 89.1-119.5), BMNH 1928.11.10.11-12, Nairobi River, elev. 6500 ft., 116'28"S,
36048'47"E (3: 59.6-140.5), BMNH 1928.11.10.13-18, Kenya, Riara River, elev. 5700 ft.,
P10'01"S, 3649'19"E (6: 37.6-92.6), BMNH 1937.12.11.16-18, Kenya, Mbakasi River
(3: 58.1-110.3), BMNH 1969.3.24.61-62, Kenya, Ngong River, tributary of Nairobi
River, 116'28"S, 36048'47"E (2: 123.6-146.9), MCZ 32518, Kenya, Riara River, elev.
5700 ft., 110'01"S, 36049'19"E (1: 96.5), USNM 72922 Kenya, Nairobi R., near
Nairobi, 116'28"S, 36048'47"E (2: 113.4-159.4).


Diagnosis

Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal

fin forked with tips of lobes rounded; many small dark spots present on head, body and fins of

adults; dorsal saddles absent; branchiostegal rays 6-7.

Amphilius sp. 1 is distinguished from A. grandis, A. oxyrhinus, A. krefftii and A.

uranoscopus by its distinctive adult pigment pattern consisting of many small darks spots on the

head, body and fins (vs. body, head and fins lacking dark spots in A. grandis, A. krefftii and A.

uranoscopus or large dark spots present on body in A. oxyrhinus). The absence a visible

epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels

distinguishes Amphilius sp. 1 from A. oxyrhinus and A. krefftii. Amphilius sp. 1 is distinguished

from A. grandis and A. oxyrhinus by its rounded (vs. pointed) caudal-fin lobes. It is

distinguished from A. krefftii and A. uranoscopus by its forked caudal (vs. caudal fin shallowly




Full Text

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TAXONOMIC REVISION OF THE Amphilius uranoscopus GROUP OF EAST-CENTRAL AFRICA (TELEOSTEI: AMPHILIIDAE) By ALFRED WILLIAM THOMSON A THESIS 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 2007 1

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2007 Alfred William Thomson 2

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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 thank Dr. Lawrence Page for directing me on this topic of study and for instruction, and for his constructive criticism and guidance. I also thank Drs. Jon Armbruster and Rebecca Kimball for their contributions and guidance. Mark Allen kindly provided photographs for figures 4-3 and 4-5. For specimen loans and access to institutional specimens I thank Ralf Britz and James Maclain (BMNH), William N. Eschmeyer, Tomio Iwamoto and David Catania (CAS), Mary A. Rogers (FMNH), Karsten Hartel (MCZ), Guy Duhamel and Patrice Pruvost (MNHN), Jos Snoeks and Mark Hanssens (MRAC), Robert H. Robins (UF) and Jeffery T. Williams (USNM). 3

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TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................3 LIST OF TABLES ...........................................................................................................................6 LIST OF FIGURES .........................................................................................................................7 ABSTRACT .....................................................................................................................................8 CHAPTER 1 INTRODUCTION..................................................................................................................10 Natural History.......................................................................................................................10 Systematic History..................................................................................................................12 2 MATERIALS AND METHODS...........................................................................................18 3 MATERIALS EXAMINED...................................................................................................21 Amphilius grandis...................................................................................................................21 Amphilius krefftii....................................................................................................................21 Amphilius oxyrhinus...............................................................................................................22 Amphilius uranoscopus...........................................................................................................22 Amphilius sp. 1........................................................................................................................23 4 SYSTEMATIC ACCOUNTS.................................................................................................24 Amphilius uranoscopus group................................................................................................24 Amphilius grandis Boulenger 1905........................................................................................25 Diagnosis.........................................................................................................................25 Description......................................................................................................................26 Coloration........................................................................................................................27 Distribution......................................................................................................................28 Amphilius krefftii Boulenger 1911..........................................................................................28 Diagnosis.........................................................................................................................28 Description......................................................................................................................29 Coloration........................................................................................................................31 Distribution......................................................................................................................31 Amphilius oxyrhinus Boulenger 1912.....................................................................................31 Diagnosis.........................................................................................................................32 Description......................................................................................................................32 Coloration........................................................................................................................34 Distribution......................................................................................................................35 Amphilius uranoscopus (Pfeffer 1889)...................................................................................35 Diagnosis.........................................................................................................................36 4

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Description......................................................................................................................36 Coloration........................................................................................................................38 Distribution......................................................................................................................38 Amphilius sp. 1 n. sp...............................................................................................................39 Diagnosis.........................................................................................................................39 Description......................................................................................................................40 Coloration........................................................................................................................42 Distribution......................................................................................................................42 5 RESULTS...............................................................................................................................59 LIST OF REFERENCES...............................................................................................................67 BIOGRAPHICAL SKETCH.........................................................................................................73 5

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LIST OF TABLES Table page 4-1 External traits diagnostic for species of the Amphilius uranoscopus group in east-central Africa.................................................................................................................................43 4-2 Morphometric data for Amphilius grandis. Range and mean include the syntypes................44 4-3 Morphometric data for Amphilius krefftii. Range and mean include the syntypes..................45 4-4 Morphometric data for Amphilius oxyrhinus. Range and mean include the holotype............46 4-5 Morphometric data for Amphilius uranoscopus. Range and mean include the specimens from the Wami drainage....................................................................................................47 5-1 Branchiostegal ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus..................................................................................64 5-2 Total gill raker counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus...........................................................................................64 5-3 Branched pectoral-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus...........................................................................64 5-4 Branched anal-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus..................................................................................64 5-5 Total vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus...........................................................................................65 5-6 Preanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus..................................................................................65 5-7 Postanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus..................................................................................65 5-8 First dorsal pterygiophore intercept counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus...........................................................65 6

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LIST OF FIGURES Figure page 1-1 Map of Africa with the east-central region indicated by the red box......................................16 1-2 Map of Africa indicating type localities of species of the Amphilius uranoscopus group as defined by Skelton (1984).............................................................................................17 4-1 Amphilius grandis, BMNH 1912.3.22.119, 113.6 mm SL; lateral, dorsal and ventral view....................................................................................................................................49 4-2 Known distribution of Amphilius grandis. Some symbols represent more than one collection site. T denotes type locality..............................................................................50 4-3 Amphilius krefftii, BMNH, 1909.10.19.26-27, Syntype 86.0 mm SL; lateral, dorsal and ventral view. Photograph by Mark Allen..........................................................................51 4-4 Known distribution of Amphilius krefftii. Some symbols represent more than one collection site. T denotes type locality..............................................................................52 4-5 Amphilius oxyrhinus, BMNH 1912.3.22.120, Holotype, 166.0 mm SL; lateral, dorsal and ventral view. Photograph by Mark Allen..........................................................................53 4-6 Known distribution of Amphilius oxyrhinus. Some symbols represent more than one collection site. T denotes type locality..............................................................................54 4-7 Amphilius uranoscopus, CAS 80494, 101.4 mm SL; lateral, dorsal and ventral view...........55 4-8 Known distribution of Amphilius uranoscopus. Some symbols represent more than one collection site.....................................................................................................................56 4-9 Amphilius sp. 1, BMNH 1910.10.31.31, Holotype, 150.0 mm SL; lateral, dorsal and ventral view........................................................................................................................57 4-10 Known distribution of Amphilius sp. 1. Some symbols represent more than one collection site. T denotes type locality..............................................................................58 5-1 Amphilius uranoscopus (juveniles) from the Wami (top: CAS 80494, 49.1 mm SL), Ruvu (middle: UF 84882, 40.4 mm SL) and Rufiji (bottom: FMNH 111680, 46.0 mm SL) drainage................................................................................................................66 7

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Abstract of Dissertation 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 THE Amphilius uranoscopus GROUP OF EAST-CENTRAL AFRICA (TELEOSTEI: AMPHILIIDAE) By Alfred W. Thomson August 2007 Chair: Lawrence M. Page Major: Zoology The taxonomy of species of the Amphilius uranoscopus group in east-central Africa is reviewed, and the Amphilius uranoscopus group is recognized by the following unique combination of characters: absence of a crenellated epidermal fold at the base of the caudal fin, usually 8+9 (i,7,8,i) principal caudal-fin rays, small bilateral bony swimbladder capsules, 36-39 total and 20-23 caudal vertebrae, and leading pterygiophore of the dorsal fin intercepting the vertebral column at the first, second or third post-Weberian vertebrae. The examination of all available museum specimens of the genus Amphilius from east-central Africa reveals the existence of at least five species of the Amphilius uranoscopus group in the region, A. grandis, A. oxyrhinus, A. krefftii, A. uranoscopus and Amphilius sp. 1. Three of the species were previously believed to be synonyms of Amphilius uranoscopus, and Amphilius sp. 1 is new to science. Amphilius sp. 1 is distinguished from A. grandis, A. oxyrhinus, A. krefftii and A. uranoscopus by its distinctive adult pigment pattern consisting of many small darks spots on the head, body and fins (vs. body, head and fins lacking dark spots in A. grandis, A. krefftii and A. uranoscopus or large dark spots present on body in A. oxyrhinus). It is additionally distinguished from these species by its absence of a visible epidermal core extending posteriorly from the mandibular barbels (vs. presence in A. oxyrhinus and A. krefftii), 8

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rounded caudal-fin lobes (vs. pointed in A. grandis and A. oxyrhinus), forked caudal fin and absence of light dorsal saddles (vs. caudal fin shallowly emarginate and light dorsal saddles present at the base of the dorsal and adipose fin in A. krefftii and A. uranoscopus), and fewer branchiostegal rays (6-7 vs. 8-9 in A. uranoscopus). The caudal fin shape and pigment patterns observed suggest that a northern and a southern group of species exist in east-central Africa. The northern species (A. grandis, A. oxyrhinus and Amphilius sp. 1) have a more deeply forked caudal fin and dark markings either as adults or juveniles, and lack light dorsal saddles. The southern species (A. krefftii and A. uranoscopus) have no dark markings, shallowly emarginate caudal fins, and light dorsal saddles present at the bases of the dorsal and adipose fin. Despite differences observed in pigment pattern and intensity between the Amphilius in the Wami, Ruvu and Rufiji drainages, all of these populations are tentatively recognized as a single species because most of the specimens examined were juveniles and only juvenile pigmentation could be compared. The three other nominal species synonymized with A. uranoscopus, A. hargeri, A. brevidorsalis and A. platychir cubangoensis, are discussed, but their status is beyond the geographic scope of this study and only tentative conclusions are made about the identity of these species. 9

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CHAPTER 1 INTRODUCTION Natural History The catfish family Amphiliidae is widely distributed throughout sub-Saharan Africa and it includes 12 genera and about 66 species (Ferraris, 2007). Most amphiliid species are found in fast-flowing water of clear, rocky streams at high elevations or rapids of large lowland rivers (Berra, 2001; Roberts, 2003). Recent workers have recognized three subfamilies within Amphiliidae, with Amphilius and Paramphilius placed in the subfamily Amphiliinae. Twenty species are currently recognized in the genus Amphilius, with most of the species occurring in the drainages of western Africa and the Congo basin. Two species of Amphilius are currently recognized in east-central Africa, an area herein defined as the area from and including the Ewaso Nyiro drainage in Kenya south to and including the Rufiji drainage in Tanzania (Fig. 1-1). This area generally corresponds to the east coast ichthyofaunal province of Roberts (1975). Roberts ichthyofaunal province also includes the Lake Victoria basin and all eastern-flowing rivers south to but not including the Zambezi River. Amphilius has been reported from the Yala River, a tributary of Lake Victoria (Whitehead, 1958; 1959) and the Ruvuma drainage, the next large drainage south of the Rufiji (Tweddle, 1983), but these records are not supported by vouchers. Species of the genus Amphilius are habitat specialists and are adapted for fast flow and rocky habitats (Skelton, 1986). Morphological adaptations for these habitats include expanded pectoral and pelvic fins with an expanded first ray, depressed body, upwardly directed eyes and reduced gas bladder (Skelton, 1986; Walsh et al., 2000). The large pelvic fins have been observed to form a feeble sucking disc, which in conjunction with the body, enable the fish to cling to rocks (Jackson, 1961b). The epidermal microstructure of the anterior ray of the pectoral 10

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fin as observed by a scanning electron microscope (Bell-Cross & Jubb, 1973) is similar to the epidermal microstructure of the adhesive organs in Asian sisorid catfishes (Singh & Agarwal, 1991; Das & Nag, 2004, 2005). In Amphilius species, males and females are generally similar, but the adductor mandibulae muscles attached to the head are sometimes greatly enlarged in mature males (J. Friel, Pers. Comm. 2007). This dimorphism has also been observed in species of Paramphilius, and was believed to be a derived character of Paramphilius by Skelton (1989). Little information is available on the biology of Amphilius species. Skelton (2001) stated that South African species breed during the summer. Marriott et al. (1997) studied the reproductive biology of A. natalensis in South Africa and concluded that the species had a long breeding period extending from August to February. They observed that breeding coincided with the rainy season and postulated that spawning was initiated in response to increased water flow rate or changes in water quality following periods of rainfall (Marriott et al., 1997). Walsh et al. (2000) were not able to determine the breeding season of A. jacksonii, but suggested that it may have two breeding periods per year. The diet of Amphilius species has been reported to predominantly consist of benthic aquatic insects (Marriott et al., 1997; Skelton, 2001; Walsh et al., 2000). Marriott et al. (1997) conducted a detailed stomach analysis of specimens of A. natalensis and found the diet to consist mostly of larval chironomids and ephemeropterans. Additionally, they found that small fishes fed predominantly on chironomid larvae, while in larger fishes, chironomid and ephemeropteran larvae contributed almost equally in volume. Walsh et al. (2000) analyzed the stomach contents of 118 specimens of A. jacksonii, and similarly found the diet of the species to consist mainly of larval Diptera, Ephemeroptera and Trichoptera. 11

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Systematic History In 1864, Gnther described Amphilius as a section of the genus Pimelodus for a new species he described as Pimelodus platychir from Sierra Leone, western Africa. Soon thereafter he recognized Amphilius as a genus (Gnther, 1865). In 1889, Pfeffer, apparently unaware of Gnthers description, created the genus Anoplopterus for a new species he described as An. uranoscopus from the Wami River drainage in Tanzania. Nine years later, Vaillant created the genus Chimarrhoglanis for a supposedly new species, which he described as C. leroyi from the nearby Ruvu River drainage. Boulenger (1898a) synonymized C. leroyi with Pimelodus platychir, but overlooked Gnthers description of the genus Amphilius and synonymized Chimarrhoglanis with Anoplopterus. Boulenger (1898a) also recognized Pfeffers A. uranoscopus and suggested that the types of A. platychir did not come from Sierra Leone. Poche (1902a) and Gnther (1902) refuted Boulengers (1898a) suggestion that the types of Amphilius platychir were not from Sierra Leone, and Poche (1902b), recognizing Amphilius as the correct name for the genus, synonymized Anoplopterus and Chimarrhoglanis with Amphilius. Despite Poche and Gnthers assertions, Boulenger continued to question the type locality of Sierra Leone for A. platychir and recognized A. platychir as a species in eastern Africa (Boulenger, 1902, 1905a, 1907a,1907b). In 1905, Boulenger described Amphilius grandis from the Tana River drainage in Kenya. He distinguished this species from A. uranoscopus by its longer head (length four times in total length v. five times in total length) and shorter barbels (maxillary barbel not reaching posterior border of head vs. extending beyond posterior border of head). He distinguished this species from A. platychir by its longer head (length 4 times in total length v. 4 to 4 1/2 times in total length), longer snout (snout length 1 2/3 interocular width vs. 1 1/3 to 1 1/2 interocular width), and a shorter caudal peduncle (not longer than deep vs. longer than deep). 12

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In 1911, Boulenger published volume two of his Catalogue of Freshwater Fishes of Africa and recognized four species in the drainages of east-central Africa: A. uranoscopus, A. platychir, A. grandis, and A. krefftii, a new species that he described from the Sigi River, Tanzania. Amphilius krefftii, like A. grandis, was distinguished from the other species of Amphilius by relative proportions of it head, barbel and caudal peduncle lengths. Soon thereafter, he described a fifth species, A. oxyrhinus, from the Eusso Mara, a tributary of the Eusso Nyiro in Kenya. He distinguished this species from only A. grandis, the only other species of Amphilius he recorded from the Eusso Nyiro. Since Boulengers description of Amphilius grandis, no new species of Amphilius have been described from east-central Africa and there has been limited study of the genus in east-central Africa. Most workers continued to recognize all of the species that Boulenger recognized (Copley, 1941; Harry, 1953; Whitehead, 1958), but Bailey (1969: 192) questioned the validity of A. krefftii and stated that it is very probably a synonym of A. grandis. Meanwhile, A. grandis was reported from throughout eastern and southern Africa (Van der Horst, 1931; Poll, 1952; Crass, 1960; Jubb, 1961; Crass, 1964). Amphilius platychir was also being reported from these areas (Ricardo, 1939a, 1939b; Jackson, 1959; Maar, 1960; Jackson, 1961a; Jubb, 1963; Bell-Cross, 1972). Bell-Cross & Jubb (1973) referred records of A. grandis from eastern and southern Africa to A. platychir, and A. platychir became accepted as being widespread throughout eastern and southern Africa. Skelton (1984) revised the genus Amphilius from eastern and southern Africa and identified two groups within Amphilius. One group is primarily western African, has a crenellated epidermal fold at the base of the caudal fin and 6 + 7 principal caudal-fin rays. The second group is primarily distributed in eastern and southern Africa, lacks the crenellated 13

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epidermal fold and usually has 8 + 9 principal caudal-fin rays. Amphilius platychir was identified as a member of the first group, and all records of the species from eastern and southern Africa were determined to be based on a single widespread species. Skelton (1984) recognized the name A. uranoscopus for this widespread eastern and southern Africa species and considered C. leroyi, A. grandis, A. krefftii and A. oxyrhinus to be synonyms of this species. Three other taxa were also synonymized with A. uranoscopus: Amphilius hargeri from Mlangi, British Central Africa, Amphilius brevidorsalis from the Rivire Revue (Mozambique) and Amphilius platychir cubangoensis from Angola. Figure 1-2 shows the type localities of A. uranoscopus and the seven nominal species that Skelton synonymized with it. Amphilius uranoscopus is the only Amphilius species of Skeltons eastern and southern African group currently recognized in east-central Africa. A single species of Skeltons western African Amphilius group, A. jacksonii, has been reported to occur in the Rufiji and Wami drainages in Tanzania (Seegers, 1996a) and is easily distinguished from A. uranoscopus by its deeply forked caudal fin, crenellated epidermal fold, and 6 + 7 principal caudal-fin rays. Although Skelton examined the type specimens of the five nominal species from east-central Africa, his recognition of A. uranoscopus as a single widespread species was based largely on the examination of specimens from South Africa. Additionally, his conclusions were based on the lack of clear meristic and morphometric differences between populations. He did not examine variation in other characters like pigment pattern and caudal fin shape. Variation in these characters in specimens identified as A. uranoscopus, suggests that the validity of the nominal species synonomized with A. uranoscopus and the acceptance of A. uranoscopus as a single species needs to be investigated. 14

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The goals of this study are to examine morphological variation in Amphilus in east-central Africa and to recognize valid taxa. In the process, the identities of the five nominal species of Amphilius described from east-central Africa will be determined. 15

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Figure 1-1. Map of Africa with the east-central region indicated by the red box. 16

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Figure 1-2. Map of Africa indicating type localities of species of the Amphilius uranoscopus group as defined by Skelton (1984). 17

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CHAPTER 2 MATERIALS AND METHODS Use of the terms origin and insertion to designate, respectively, anterior and posterior base of all fins follows Cailliet et al. (1986). Counts and measurements were made on the left side of specimens when possible and follow Skelton (1981, 1984, 1986), except head length was measured as the distance from the top of snout to the dorsal edge of the opercular cavity. In addition, the following measurements were made: Body depth at anus: depth of the body at the anus. Postorbital length: distance from the posterior margin of the orbit to the dorsal edge of the opercular cavity. Prepectoral length: distance from the origin of the pectoral fin to the tip of the snout. Preanal length: distance from the origin of the anal fin to the tip of the snout. Dorsal-fin base length: distance from the structural base of the first ray to the point where the membrane behind the last ray contacts the body. Adipose-fin base length: distance from the origin of the adipose fin to the posteriormost connection of the adipose fin to the body. Pelvicand pectoral-fin lengths: distance from the origin of the fin to the tip of the fin. Anus to anal-fin length: distance from the origin of the anal fin to the center of the anus. Prepelvic length: distance from the origin of the pelvic fin to the tip of the snout. Postpelvic length: distance from the origin of the pelvic fin to the base of the caudal fin. Distance from insertion of dorsal fin to insertion of adipose fin (end dorsal end adipose): distance from the point where the membrane behind the last dorsal-fin ray contacts the body to the posteriormost connection of the adipose fin to the body. Distance from origin of dorsal fin to base of caudal fin (front dorsal caudal): distance from the origin of the dorsal fin to the base of the caudal fin. Preanus length: distance from tip of the snout to opening of the anus. 18

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All available museum specimens from the Ewaso Nyiro, Tana, Galana, Pangani, Wami, Ruvu and Rifiji drainages were examined. In addition, the type specimens of Amphilius hargeri, A. brevidorsalis and A. platychir cubangoensis were examined; however, because large series of specimens from outside the east-central region could not be examined, only tentative conclusions can be made about the identity of these species. Measurements were made point-to-point with digital calipers, and data were recorded to tenths of a millimeter. Subunits of the head are presented as proportions of the head length (HL). Head length and measurements of the body parts are given as proportions of the standard length (SL). For the fin-ray counts, the numbers of unbranched soft rays are indicated by Roman numerals, branched soft rays are indicated by Arabic numerals. The number of anterior unbranched rays in the anal fin is difficult to determine, and the counts were checked with radiographs whenever possible. Amphiliids typically have a small spinelet in front of the first unbranched dorsal-fin ray. The spinelet is not included in the counts. Vertebrae were counted by means of radiographs following the methods of Skelton (1976). Counts exclude 5 Weberian vertebrae and include the ural centrum as one. Material examined is given under each species account and is listed by the drainage followed by the catalog number, country, locality, geographic coordinates, and in parentheses, the number of specimens and the size range in mm SL. Geographic coordinates were determined from maps. An effort was made to provide geographic coordinates for all specimens; however, these coordinates are only estimates and some lots with different specific localities have the same geographic coordinates because some localities could not be found on a map. 19

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All maps were created using the Online Map Creator (OMC), which can be found at http://www.aquarius.ifm-geomar.de/ ( Weinelt 1996-2006). Distribution points on maps were added using Abode Photoshop CS2. Points indicate only specimens examined in this study. Synonymies include all accounts of the species in east-central Africa. The first page of the account of the species and all figures are listed. If the species is also listed in a key on a separate page from the account, that page is also listed. The type of information in the reference is given followed by the locality for the species in the account. Additionally, any specimens known that the account may be based on are listed. If the account is only based in part on the species, only the information that is applicable to that species is listed. Accounts of species outside east-central Africa that give only a general distribution in east-central Africa are excluded. 20

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CHAPTER 3 MATERIALS EXAMINED Amphilius grandis Eusso Nyiro drainage: BMNH 1908.9.17.13-18, Kenya: Nyiro-Narok; Niro-Narok system, elev. 4000-5000 ft., 0N, 36 31E (6: 93.4-181.4); BMNH 1912.3.22.119, Kenya, Eusso Nyiro, below falls, 012N, 38 00E (1: 113.6). Tana drainage: BMNH 1904.12.23.50-52, Kenya, Chania R. of Tetse, Tana system, Kenya, elev. 7000 ft., 1N, 37 05E (2: 136.1-160.0, Syntypes); BMNH 1937.6.4.36-42, Kenya, Thika River, Ndula Falls, 1N, 37 05E (12: 32.1-65.9). Amphilius krefftii Galina drainage: BMNH 1969.3.24.63-69, Kenya, Tsabo [Tasvo] River tributary of Athi [Galina] River, mountain Mbololo, 3 0' N, 38 20'E, (7: 43.6-99.5); BMNH 1969.3.24.70, Kenya, Voi River, Teita hills, Voi District, 322N, 38 34E (1: 106.6). Pangani drainage: BMNH 1909.10.19.26-27, Tanzania, Usambara, in rivulet running from Anani Hills to Sigi R., 5 0'N, 38 48'E (2: 47.5-86.0, Syntypes); BMNH 1905.7.25.41-42, Kenya, Kibosho, Kilimandjaro, 3 12', 37 19' (2: 77.9-94.9); BMNH 1968.10.25.3, Tanzania, River Lume (upper reaches of Pangani River) at source, 5 miles east of Lake Chala, 323N, 37 43E (1: 106.6); BMNH 1968.10.25.5-6, Tanzania, Kisiwani, near Amani, East Usambura mountains, elev. 1,475 ft., 5N, 38 37E (2: 36.2-43.1); BMNH 1968.10.25.7, Tanzania, River Sigi, east Usumbura Mountains, 5 0'N, 38 48'E (1: 72.7); BMNH 1968.10.25.8, Tanzania, River Lume (upper reaches of Pangani River) east of Taveta, 3N, 37 43E (1: 67.2); FMNH 111678, Tanzania, East Usambara Mountains, 4.5 km ESE Amani, Monga Tea Estate, 5 6'N, 38 36'E (1: 112.0); FMNH 111684, Tanzania, West Usambara Mts., in river (stream) near Ambangulu Tea Estate factory, 5N, 38 25E (1: 29.0). 21

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Amphilius oxyrhinus Eusso Nyiro drainage: BMNH 1912.22.120, Kenya, Eusso Mara, a swift mountain stream, tributary of Eusso Nyiro, 0 12'N, 38 00'E (1: 166.0, Holotype). Tana drainage: BMNH 1965.12.7.125, Kenya, Rogati River, Sagana, 0N, 37 16E (1: 42.9); BMNH 1965.12.7.126-128, Kenya, Rogati River, Sagana, 040N, 37 16E (3: 60.9-89.7); BMNH 1965.12.7.129, Kenya, Rogati River, Sagana, 0N, 37 16E (1: 156.8); BMNH 1965.12.7.130-131, Kenya, Rogati River, Sagana, 0N, 37 16E (2: 74.9-98.7); BMNH 1965.12.7.132, Kenya, Rogati River, Sagana, 0N, 37 16E (1: 51.8); BMNH 1966.6.28.2-3, Kenya, Rogati River, Sagana, 0N, 37 16E (2: 65.9-102.6); BMNH 1966.8.25.18, Kenya, Rogati River, Sagana, 0N, 37 16E (1: 34.8); 34.8; MRAC 74-48-P-12-13, Kenya, upper Tana, side creek, 00'S, 37'E (2: 52.6-60.3). Amphilius uranoscopus Rufiji drainage: FMNH 111679, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest Reserve, 4 km W, 5 km N Chita, along the Chita-Ihimbo trail, elev. 1460 m, 8'30"S, 35'25"E (8: 24.4-40.4); FMNH 111680, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest Reserve, 4.5 km W Chita, along Chita-Ihimbo trail, elev. 600 m, 8'10"S, 35'15"E (18: 25.0-68.4.0); FMNH 111681, Tanzania, Udzungwa Mts., Udzungwa Scarp Forest Reserve, 4.5 km W Chita, along Chita-Ihimbo trail, elev. 600 m, 8'10"S, 35'15"E (14: 25.5-77.4). Ruvu drainage: FMNH 111682, Tanzania, Uluguru Mountains, Uluguru North Forest Reserve, 3 km W, 1.3 km N Tegetero, elev. 1345 m, 6 55' 45" S, 37 42' 20" E (1:55.7); FMNH 111683, Tanzania, Uluguru Mts., Uluguru North Forest Reserve, 5.1 km W, 2.3 km N Tegetero, elev. 1535 m, 6'12"S, 37'0"E (3: 26.4-52.4); MNHN 1897-0003, Tanzania, Torrent of Mrogoro at Zanguebar, Oukami, 6'00"S, 37'45"E (1: 117.3, Holotype of Chimarrhoglanus leroyi), 22

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UF 84882, Tanzania, Morogoro Region, southern side of Uluguro Mountains, about 1 km N of Ruvu River, S of Kibungo Village, Kimboza Forest, 7'00"S, 37'00"E (7: 21.9-40.4). Wami drainage: CAS 80494, Tanzania, Nguru Mountains, Nguru South Forest Reserve, 6'S, 37'E (2: 49.1-101.4); BMNH 1969.1.15.1, Tanzania, Stream at Arusha in the foothills of Mount Meru, 3'57"S, 36'28"E (1: 84.7). Amphilius sp. 1 Galina drainage: BMNH 1905.12.11.2, Kenya, Nairobi River, Kikaya, 1'28"S, 36'47"E (1: 160.3), BMNH 1909.11.15.21-23, Nairobi River, elev. 2500 ft., 1'28"S, 36'47"E (3: 89.1-119.5), BMNH 1910. 10.31.31, Nairobi, Kenya, 1'28"S, 36'47"E (1: 150.0), BMNH 1928.11.10.11-12, Nairobi River, elev. 6500 ft., 1'28"S, 36'47"E (3: 59.6-140.5), BMNH 1928.11.10.13-18, Kenya, Riara River, elev. 5700 ft., 1'01"S, 36'19"E (6: 37.6-92.6), BMNH 1937.12.11.16-18, Kenya, Mbakasi River (3: 58.1-110.3), BMNH 1969.3.24.61-62, Kenya, Ngong River, tributary of Nairobi River, 1'28"S, 36'47"E (2: 123.6-146.9), MCZ 32518, Kenya, Riara River, elev. 5700 ft., 1'01"S, 36'19"E (1: 96.5), USNM 72922, Kenya, Nairobi R., near Nairobi, 1'28"S, 36'47"E (2: 113.4-159.4). 23

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CHAPTER 4 SYSTEMATIC ACCOUNTS Amphilius uranoscopus group All species of Amphilius in east-central Africa belong to the Amphilius uranoscopus group and are recognized by the following unique combination of characters: absence of a crenellated epidermal fold at the base of the caudal fin, usually 8+9 (i,7,8,i) principal caudal fin rays, small bilateral bony swimbladder capsules, 36-39 total and 20-23 caudal vertebrae, and leading pterygiophore of the dorsal fin intercepting the vertebral column at the first, second or third post-Weberian vertebrae. Skelton (1984) recognized this species group as a single species, A. uranoscopus, but did not formally diagnose it. Externally, species of the A. uranoscopus group are similar to A. natalensis, A. lampei and A. kivuensis but these species differ by having the leading pterygiophore of the dorsal fin intercepting the vertebral column after the third post-Weberian vertebrae. Amphilius cryptobullatus is also similar externally, but differs from A. uranoscopus in the development and extremely large size of its bilateral bony swimbladder capsules. Following are diagnoses and descriptions of valid species of the Amphilius uranoscopus group in east-central Africa. Species are aranged alpabetically. Diagnostic traits are summarized in Table 4-1. 24

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Amphilius grandis Boulenger 1905 (Figure 4-1 and 4-2; Table 4-2) Amphilius grandis Boulenger 1905a: 63, Pl. 7 (fig. 3), Original description, Type locality: Chania R. of Tetse, Tana system, Kenya, elev. 7000 ft.; Boulenger 1905b: 48, Headwaters of the Tana system; Boulenger 1911: 353, 355, fig. 275 (in part), in key, description, East Africa (Tana to Athi River systems) [Tana drainage only]; Boulenger, 1912: 675, diagnosis from Amphilius oxyrhinus, [Eusso Nyiro drainage], [BMNH 1912.3.22.119]; Boulenger, 1916: 306, Nairobi, Eusso Nyiro, below falls; Pellegrin 1936: 57, similarity to Amphilius platychir var. cubangoensis; Copley, 1941: 15 [Eusso Nyiro and Tana drainages], Harry, 1953: 189 (in part), synonymy; Copley, 1958: 98, [Tana drainage], [BMNH 1937.6.4.36-42], Whitehead, 1958: 198, [Kenya and Tanzania], Bailey, 1969: 192 (in part), eastern rivers of Kenya; Skelton, 1984: 45, in synonymy of A. uranoscopus. Amphilius platychir (non Gnther) Whitehead, 1958: 198 (in part), [Eusso Nyiro drainage]. Amphilius uranoscopus (non Pfeffer) Skelton, 1994: 126, [Tana drainage]; Seegers et al., 2003: 37 (in part), [Tana drainage]. Diagnosis Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal fin deeply emarginate or forked with tips of lobes pointed; adults plain-colored, without dark spots or blotches on body, head or fins; dorsal saddles absent. The absence of a visible epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels distinguishes Amphilius grandis from A. oxyrhinus and A. krefftii. Amphilius grandis is distinguished from A. uranoscopus and A. krefftii by its more deeply emarginate caudal fin with the tips of the caudal-fin lobes pointed (vs. caudal fin shallowly emarginate with rounded lobes) and the absence of light dorsal saddles (vs. light dorsal saddles present at the base of the dorsal and adipose fin). Amphilius grandis is distinguished from Amphilius sp. 1 and A. oxyrhinus by the absence of dark spots on the head or body of adults (vs. adults of Amphilius sp. 1 with many small dark spots on head and body, and adults of A. oxyrhinus with dark spots or blotches on body). Amphilius grandis is distinguished from 25

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Amphilius sp. 1 by having the lobes of the caudal fin pointed (vs. rounded) and the absence of spots on the fins. Description Morphometric data as in Table 4-2. Body elongate, ventral profile flattened ventrally to anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated epidermal fold. Anus and urogenital openings located at posteriormost extent of pelvic fin, closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 38 (3), 39 (6) or 40 (1). Preanal vertebrae 20 (1), 21 (3) or 22 (6). Postanal vertebra 16 (1), 17 (4) or 18 (5). First dorsal pterygiophore intercept count 1 (2) or 2 (8). Head and anterior part of body depressed and broad. Head wedge shaped in lateral view. Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped connection. Membranes connected by a small frenum at midline, but with shallow groove at posterior margin. Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed. Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary teeth short and conical, tooth patches forming U-shaped band, separated medially. Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and flattened with pointed tip; barbel extending posterolaterally from corner of mouth to just short of pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of 26

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lower jaw, extending to origin of pectoral-fin. Inner mandibular barbel originates anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal membrane with 7 (10) or 8 (9) rays. Gill rakers on first epibranchial 2 (7) or 3 (12); rakers on first ceratobranchial 4 (1), 5(3), 6 (6), 7 (8) or 8 (1); total gill rakers on first arch 6 (1), 7 (2), 8 (2), 9 (8), 10 (5) or 11 (1). Eye small, positioned dorsolaterally approximately midway between tip of snout and posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and posterior nares with prominent tubular rims; nares separate but relatively close to each other. Posterior nare located about midway between eye and tip of snout. Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (18) or i,7 (1) rays, and fin margin straight. Pectoral fin with i,9 (3) or i,10 (16) rays with first ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (19) rays with first ray unbranched and greatly thickened. Pelvic fin with straight posterior margin. Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base, extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded edge, not deeply incised posteriorly. Caudal fin deeply emarginate or forked with tips of lobes pointed, with i,7,8,i (18) or i,8,9,i (1) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin base, with iii,6 (19) rays. Anal fin margin almost straight. Coloration Dorsal and lateral surfaces of head and body brown. Ventral region of head dark yellow. Ventral region of body brown to dark yellow. Light dorsal saddles absent. Dorsal, adipose, 27

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caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown with light distal edge; ventral surfaces light yellow. Maxillary and mandibular barbels brown. Caudal fin with dark crescent-shaped band at base. Juvenile coloration: medium-sized dark spots on head and body in specimens larger than 40.0 mm SL, but spots usually absent on specimens under 40.0 mm SL. Spots on body arranged in line along lateral line. All fins light yellow, with medial bands of brown pigment all rayed fins. Pectoral and pelvic fins with first unbranched ray dark brown dorsally. Distribution Tana and Eusso Nyiro (Ewaso Ng'iro) River drainages, Kenya. Amphilius krefftii Boulenger 1911 (Figures 4-3 and 4-4; Table 4-3) Amphilius krefftii Boulenger, 1911: 356, fig. 276, Original description, Type locality: Usambara, in rivulet running from Anani Hills to Sigi River. [Sigi River drainage, Tanzania]; Harry, 1953: 189, synonymy; Copley, 1958: 100 [Pangani drainage]; Whitehead, 1958: 198 [Kenya and Tanzania]; Bailey, 1969: 192 [Sigi and Pangani drainages]; Bernacek, 1980: 36 [Tanzania]; Skelton, 1984: 45, in synonymy of A. uranoscopus. Pimelodus (Amphilius) uranoscopus (non Pfeffer) Hilgendorf, 1905: 411 (in part), [Pangani drainage]. Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Galina drainage] [BMNH 1969.3.24.63-69, BMNH 1969.3.24.70]; Copley, 1958: 100 [Galina drainage] [BMNH 1969.3.24.63-69, BMNH 1969.3.24.70]; Bailey, 1969: 192 (in part), [Pangani drainage]. Amphilius uranoscopus (non Pfeffer) Harry, 1953: 189 (in part), synonymy; Skelton, 1994: 126 (in part), [Galina and Pangani drainages]; Seegers et al., 2003: 37 (in part), [Pangani drainage]. Amphilius platychir (non Gnther) Copley, 1958: 100 (in part), [Pangani drainage]. Diagnosis Visible epidermal core extending posteriorly from the mandibular barbels present; caudal fin emarginate with tips of lobes rounded; adults without dark spots on head, body or fins; light 28

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dorsal saddles present at the base of the dorsal and adipose fin; branchiostegal rays usually seven. The presence of a visible epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels distinguishes Amphilius krefftii from A. grandis, Amphilius sp. 1 and A. uranoscopus. Amphilius krefftii is distinguished from A. oxyrhinus, A. grandis and Amphilius sp. 1 by the shape of the caudal fin (emarginate in A. krefftii vs. deeply emarginate or forked in A. grandis and forked in A. oxyrhinus and Amphilius sp. 1) and the presence of light dorsal saddles at the base of the dorsal and adipose fin (vs. light dorsal saddles absent). Additionally the caudal-fin lobes are rounded in A. krefftii (vs. pointed in A. grandis and A. oxyrhinus). Amphilius krefftii is further distinguished from Amphilius sp. 1 by the absence of dark spots or blotches on the head, body or fins in adults and further distinguished from A. uranoscopus by having usually seven branchiostegal rays (vs. eight or nine). Description Morphometric data as in Table 4-3. Body elongate, ventral profile flattened ventrally to anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin, closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 36 (7), 37 (3) or 38 (1). Preanal vertebrae 20 (7) or 21 (5). Postanal vertebra 15 (1), 16 (9) or 17 (1). First dorsal pterygiophore intercept count 1 (11) or 2 (1). Head and anterior part of body depressed and broad. Head wedge shaped in lateral view. Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to 29

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pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped or deeply concave connection. Membrane connected by a small frenum at midline, but with shallow groove at posterior margin. Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed. Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary teeth short and conical, tooth patches forming U-shaped band, separated medially. Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of lower jaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal membrane with 7 (16) or 8 (1) rays. Gill rakers on first epibranchial 2 (11), 3 (3), or 4 (1); rakers on first ceratobranchial 5 (5), 6 (8) or 7 (2); total gill rakers on first arch 7 (5), 8 (4), 9 (5) or 10 (1). Eyes small, positioned dorsolaterally approximately midway between tip of snout and posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and posterior nares with prominent tubular rims; nares separate but relatively close to each other. Posterior nare located about midway between eye and tip of snout. Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (17) rays, and fin margin slightly convex. Pectoral fin with i,9 (11), i,10 (5) or i,11 (1) rays with first ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with 30

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four to five innermost rays progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (17) rays with first ray unbranched and greatly thickened. Pelvic fin with straight posterior margin. Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base, extending over posterior end of anal-fin base. Margin strongly convex with sharply rounded edge, not deeply incised posteriorly. Caudal fin emarginate with tips of lobes rounded, with i,5,6,i (1) or i,7,8,i (15) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin base, with iii,6 (15) or iii7 (2) rays. Anal fin margin almost straight. Coloration Dorsal and lateral surfaces of head and body brown. Ventral region light brown. Light dorsal saddles at anterior and posterior of dorsal-fin and adipose-fin bases. Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown, ventral surfaces light yellow. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band at base. Juvenile coloration: similar to that of adults, except all fins light yellow, with small blotches of brown pigment on the branched rays. Pectoral and pelvic fins with first unbranched ray dark brown dorsally. Distribution Lower Galina and Pangani River drainages, Kenya and Tanzania. Amphilius oxyrhinus Boulenger 1912 (Figures 4-5 and 4-6; Table 4-4) Amphilius oxyrhinus Boulenger 1912: 675, Pl. 80, Original description, Type locality: Eusso Mara, a swift mountain stream, Kenya [Eusso Nyiro (Ewaso Ng'iro) drainage]; Boulenger, 1916: 306, [Eusso Nyiro drainage]; Skelton, 1984: 45, in synonymy of A. uranoscopus. Amphilius leroyi (non Vaillant) Pellegrin, 1905: 177, Nairobi; Harry, 1953: 189 (in part), synonymy. 31

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Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Eusso Nyiro drainage]; Harry, 1953: 189, synonymy; Copley, 1958: 98 (in part), [Tana drainage]. Amphilius platychir (non Gnther) Whitehead, 1958: 198 (in part), [Eusso Nyiro drainage]. Amphilius uranoscopus (non Pfeffer) Skelton, 1994: 126, [Tana drainage]; Seegers et al., 2003: 37 (in part), [Tana drainage]. Diagnosis Visible epidermal core extending posteriorly from the mandibular barbels present; caudal fin forked with tips of lobes pointed; adults with large dark spots or blotches on body; spots on head and fins absent or indistinct; dorsal saddles absent. The presence of a visible epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels distinguishes Amphilius oxyrhinus from A. grandis, Amphilius sp. 1 and A. uranoscopus. Amphilius oxyrhinus is distinguished from A. krefftii and further distinguished from A. uranoscopus by its forked caudal fin with the tips of the caudal-fin lobes pointed (vs. caudal fin shallowly emarginate with rounded lobes) and the absence of light dorsal saddles (vs. light dorsal saddles present at the base of the dorsal and adipose fin). It is further distinguished from Amphilius sp. 1 by the larger spots or blotches on the body of adults (vs. small spots present on head and body), its pointed caudal-fin lobes (vs. caudal fin with rounded lobes), and the absence of spots on the fins. Adults of Amphilius oxyrhinus are further distinguished from adults of A. grandis, Amphilius sp. 1 and A. uranoscopus by the presence of dark spots or blotches on the body of adults (vs. adults without dark spots or blotches on body). Description Morphometric data as in Table 4-4. Body elongate, ventral profile flattened ventrally to anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest 32

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body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin, closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 37 (4) or 38 (4). Preanal vertebrae 21 (7) or 22 (1). Postanal vertebra 16 (5) or 17 (3). First dorsal pterygiophore intercept count 1 (4) or 2 (4). Head and anterior part of body depressed and broad. Head wedge shaped in lateral view. Snout broad, pointed when viewed from above. Head becoming wider from tip of snout to pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped or deeply concave connection. Membrane connected by a small frenum at midline, but with shallow groove at posterior margin. Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed. Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary teeth short and conical, tooth patches forming U-shaped band, separated medially. Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of lower jaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates anterolaterally of inner mandibular barbel, extending just short of edge of branchiostegal membrane. Visible epidermal core extending posteriorly from bases of mandibular barbels. Branchiostegal membrane with 7 (4), 8 (6) or 9 (2) rays. Gill rakers on first epibranchial 3 (13); 33

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rakers on first ceratobranchial 4 (1), 5(3) or 6 (9); total gill rakers on first arch 7 (1), 8 (3) or 9 (9). Eyes small, positioned dorsolaterally approximately midway between tip of snout and posterior margins of operculum. Horizontal diameter of eye slightly wider than vertical diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and posterior nares with prominent tubular rims; nares separate but relatively close to each other. Posterior nare located about midway between eye and tip of snout. Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (13) rays, and fin margin straight. Pectoral fin with i,10 (11) or i,11 (2) rays with first ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (13) rays with first ray unbranched and greatly thickened. Pelvic fin with straight posterior margin. Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base, extending over posterior end of anal-fin base. Margin strongly convex with sharply rounded edge, not deeply incised posteriorly. Caudal fin forked with tips of lobes pointed, with i,7,7,i (1) or i,7,8,i (11) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin base, with iii,6 (12) rays. Anal fin margin almost straight. Coloration Dorsal and lateral surfaces of head and body brown. Ventral and lateral regions of body covered with large black spots or blotches. Ventral region of head and body dark yellow. Light dorsal saddles absent, but pale regions sometimes present in front of dorsal and anal-fin bases. Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown with light distal edge, ventral surfaces light yellow. Medial band of dark pigment on all rayed 34

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fins. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band at base Juvenile coloration: Similar to that of adults, except all fins lighter brown, with medial bands of dark pigment all rayed fins. Pectoral and pelvic fins with first unbranched ray dark brown dorsally. Distribution Tana and Eusso Nyiro (Ewaso Ng'iro) River drainages, Kenya. Amphilius uranoscopus (Pfeffer 1889) (Figures 4-7 and 4-8; Table 4-5) Anoplopterus uranoscopus Pfeffer 1889: 16. Type locality: Ushonda and Monda, upper ranges of Wami R., Tanzania [Wami River drainage]; Pfeffer, 1893: 161, [Wami drainage]; Pfeffer, 1896: 33, fig. 14, [Wami drainage]; Boulenger, 1898a: 255, reference to original description; Boulenger, 1901: 447, diagnosis; Poche, 1902a: 21; Poche, 1902b: 211. Chimarrhoglanis leroyi Vaillant, 1897: 82, Original description, Type locality: Zanquebar, torrent de Mroboro, 600 meters altitude [Ruvu drainage]; Boulenger, 1898a: 254, as a synonym of Amphilius platychir; Boulenger, 1898b: 4 [Ruvu drainage]; Poche, 1902a: 121; Poche, 1902b: 211; Pellegrin, 1905a: 177, type information; Pellegrin 1936: 57, similarity to Amphilius platychir var. cubangoensis; Harry, 1953: 186, in synonymy of Amphilius platychir; Bertin & Estve, 1950: 35, information on type; Skelton, 1984: 45, in synonymy of Amphilius uranoscopus. Amphilius uranoscopus Boulenger, 1902: 41, in key; Boulenger, 1905b: 48, [Wami drainage]; Boulenger 1911: 253, 354, fig. 277, Description, [Wami drainage]; Harry, 1953: 187, synonymy; Copley, 1958: 99 [Wami drainage]; Bailey, 1969: 192 [Wami drainage]; Bernacek, 1980: 36, [Wami, Ruvu and Rufiji drainages]; Skelton, 1984: 41, information on types; Skelton, 1994: 126 [Wami, Ruvu and Rufiji drainages]; Seegers, 1996a: 192, figs. 136-137, information on types. Pimelodus (Amphilius) uranoscopus (non Pfeffer) Hilgendorf, 1905: 411 (in part), [Rufiji drainage]. Amphilius platychir (non Gnther) Matthes, 1967: 5, 15 [Rufiji drainage]; Bell-Cross, 1972: 18, table 7, [Rufiji drainage]. Amphilius grandis (non Boulenger) Bailey, 1969: 192 (in part) [Ruvu drainage]. 35

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Diagnosis Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal fin emarginate with tips of lobes rounded; no dark spots present on head, body or fins of adults; light dorsal saddles present at the base of the dorsal and adipose fin; branchiostegal rays 8-9. The absence of a visible epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels distinguishes Amphilius uranoscopus from A. oxyrhinus and A. krefftii. Amphilius uranoscopus is distinguished from A. grandis, Amphilius sp. 1 and further distinguished from A. grandis by the shape of the caudal fin (emarginate in A. uranoscopus vs. shallowly emarginate or forked in A. grandis and forked in Amphilius sp. 1 and A. oxyrhinus) and the presence of light dorsal saddles at the base of the dorsal and adipose fin (vs. light dorsal saddles absent). Additionally the caudal-fin lobes are rounded in A. uranoscopus (vs. pointed in A. grandis and A. oxyrhinus). Amphilius uranoscopus is further distinguished from A. oxyrhinus and Amphilius sp. 1 by the absence of dark spots or blotches on the head and body in adults. Its greater number of branchiostegal rays (8-9 vs. 6-7) and lack of spots on the fins further distinguishes it from Amphilius sp. 1. Description Morphometric data as in Table 4-5. Body elongate, ventral profile flattened ventrally to anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated epidermial fold. Anus and urogenital openings located at posteriormost extent of pelvic fin, closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 36

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36 (11), 37 (14) or 38 (2). Preanal vertebrae 19 (2), 20 (7), 21 (14) or 22 (9). Postanal vertebra 15 (9), 16 (19) or 17 (1). First dorsal pterygiophore intercept count 1 (30) or 2 (1). Head and anterior part of body depressed and broad. Head wedge shaped in lateral view. Snout broad, blunt when viewed from above. Head becoming wider from tip of snout to pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped connection. Membrane connected by a small frenum at midline, but with shallow groove at posterior margin. Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed. Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary teeth short and conical, tooth patches forming U-shaped band, separated medially. Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and flattened with pointed tip; barbel extending posterolaterally from corner of mouth to middle of pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of lower jaw, extending middle pectoral-fin base. Inner mandibular barbel originates anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal membrane with 8 (25) or 9 (29) rays. Gill rakers on first epibranchial 2 (35), 3 (18) or 4(1); rakers on first ceratobranchial 4 (9), 5 (26), 6 (14), 7 (4) or 8 (1); total gill rakers on first arch 6 (5), 7 (24), 8 (15), 9 (5), 10 (4) or 11 (1). Eyes small, positioned dorsolaterally approximately midway between tip of snout and posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and 37

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posterior nares with prominent tubular rims; nares separate but relatively close to each other. Posterior nare located about midway between eye and tip of snout. Dorsal-fin origin at point over middle of pectoral fin. Dorsal fin with i,5 (1) or i,6 (53) rays, and fin margin slightly convex. Pectoral fin with i,8 (3), i,9 (23) or i,10 (28) rays with first ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (54) rays with first ray unbranched and greatly thickened. Pelvic fin with straight posterior margin. Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base, extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded edge, not deeply incised posteriorly. Caudal fin emarginate with tips of lobes rounded, with i,7,6,i (1), i,7,7,i (1) or i,7,8,i (52) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin base, with ii,6 (6), iii,5 (2), iii,6 (15), iii7 (18) or iii,8 (11) rays. Anal fin margin almost straight. Coloration Dorsal and lateral surfaces of head and body brown. Ventral region light yellow. Light dorsal saddles at anterior and posterior of dorsal-fin and adipose-fin bases. Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown, ventral surfaces light yellow. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band at base. Juvenile coloration: Similar to that of adults, except all fins light yellow, with small blotches of brown pigment on the branched rays. Pectoral and pelvic fins with first unbranched ray dark brown dorsally. Distribution Wami, Ruvu and River Rufiji drainages, Tanzania. 38

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Amphilius sp. 1 n. sp. (Figures 4-9 and 4-10; Table 4-6) Amphilius grandis (non Boulenger) Copley, 1941: 15 (in part), [Galina drainage] [BMNH 1937.12.11.16-18, BMNH 1969.3.24.61-62]; Copley, 1958: 98 (in part), [Galina drainage] [BMNH 1937.12.11.16-18, BMNH 1969.3.24.61-62]. Amphilius uranoscopus Skelton, 1994: 126 (in part) [Galina drainage]; Seegers, 1996b: 255, fig. 16 (in part), Athi River; Seegers et al., 2003: 37 (in part), Athi River syatem. Holotype: BMNH 1910.10.31.31, Nairobi, Kenya, 1'28"S, 36'47"E (1: 150.0), Paratypes: BMNH 1905.12.11.2, Kenya, Nairobi River, Kikaya, 1'28"S, 36'47"E (1: 160.3), BMNH 1909.11.15.21-23, Nairobi River, elev. 2500 ft., 1'28"S, 36'47"E (3: 89.1-119.5), BMNH 1928.11.10.11-12, Nairobi River, elev. 6500 ft., 1'28"S, 36'47"E (3: 59.6-140.5), BMNH 1928.11.10.13-18, Kenya, Riara River, elev. 5700 ft., 1'01"S, 36'19"E (6: 37.6-92.6), BMNH 1937.12.11.16-18, Kenya, Mbakasi River (3: 58.1-110.3), BMNH 1969.3.24.61-62, Kenya, Ngong River, tributary of Nairobi River, 1'28"S, 36'47"E (2: 123.6-146.9), MCZ 32518, Kenya, Riara River, elev. 5700 ft., 1'01"S, 36'19"E (1: 96.5), USNM 72922 Kenya, Nairobi R., near Nairobi, 1'28"S, 36'47"E (2: 113.4-159.4). Diagnosis Visible epidermal core extending posteriorly from the mandibular barbels absent; caudal fin forked with tips of lobes rounded; many small dark spots present on head, body and fins of adults; dorsal saddles absent; branchiostegal rays 6-7. Amphilius sp. 1 is distinguished from A. grandis, A. oxyrhinus, A. krefftii and A. uranoscopus by its distinctive adult pigment pattern consisting of many small darks spots on the head, body and fins (vs. body, head and fins lacking dark spots in A. grandis, A. krefftii and A. uranoscopus or large dark spots present on body in A. oxyrhinus). The absence a visible epidermal core extending posteriorly from the bases of the inner and outer mandibular barbels distinguishes Amphilius sp. 1 from A. oxyrhinus and A. krefftii. Amphilius sp. 1 is distinguished from A. grandis and A. oxyrhinus by its rounded (vs. pointed) caudal-fin lobes. It is distinguished from A. krefftii and A. uranoscopus by its forked caudal (vs. caudal fin shallowly 39

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emarginate) and the absence of light dorsal saddles (vs. light dorsal saddles present at the base of the dorsal and adipose fin). It is distinguished from A. uranoscopus by its fewer branchiostegal rays (6-7 vs. 8-9). Description Morphometric data as in Table 4-6. Body elongate, ventral profile flattened ventrally to anal-fin base, then tapered dorsally to end of caudal peduncle. Dorsal profile rising gently from tip of snout to origin of dorsal fin, then nearly horizontal to end of caudal peduncle. Greatest body depth at dorsal-fin origin. Caudal peduncle laterally compressed, without crenellated epidermial fold. Anus and urogenital openings located just posterior to base of pelvic fin, closer to posterior base of pelvic fin than to origin of anal fin. Skin smooth. Lateral line complete, extending from dorsal edge of opercular cavity to base of caudal fin. Total vertebrae 37 (2), 38 (7) or 39 (3). Preanal vertebrae 20 (2), 21 (3) or 22 (7). Postanal vertebra 15 (1), 16 (5), 17 (4) or 18 (2). First dorsal pterygiophore intercept count 1 (12) or 2 (1). Head and anterior part of body depressed and broad. Head wedge shaped in lateral view. Snout broad, moderatly pointed when viewed from above. Head becoming wider from tip of snout to pectoral-fin base. Branchiostegal membranes moderately joined at isthmus forming a V-shaped or deeply concave connection. Membrane connected by a small frenum at midline, but with shallow groove at posterior margin. Mouth broad, gently curved, subterminal. Lips moderately fleshy, papillate. Rictal lobe large and papillate. Anterior portion of premaxillary tooth band exposed with mouth closed. Premaxillary tooth patches joined, forming U-shaped band, with short conical teeth. Dentary teeth short and conical, tooth patches forming U-shaped band, separated medially. Three pairs of simple tapered circumoral barbels. Maxillary barbel large, fleshy and flattened with pointed tip; barbel extending posterolaterally from corner of mouth to just short of 40

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pectoral-fin base. Outer mandibular barbel thin with pointed tip, origin at posterior corner of lower jaw, extending to edge of branchiostegal membrane. Inner mandibular barbel originates anterolaterally of inner mandibular barbel, extending to edge of branchiostegal membrane. Branchiostegal membrane with 6 (1) or 7 (21) rays. Gill rakers on first epibranchial 3 (19) or 4 (3); rakers on first ceratobranchial 6 (8) or 7 (14); total gill rakers on first arch 6 (1), 9 (7), 10 (12) or 11 (2). Eyes small, positioned dorsolaterally approximately midway between tip of snout and posterior margin of operculum. Horizontal diameter of eye slightly wider than vertical diameter. Eye without free orbit; covered with skin confluent with dorsal surface of head. Anterior and posterior nares with prominent tubular rims; nares separate but relatively close to each other. Posterior nare located about midway between eye and tip of snout. Dorsal-fin origin at point over tip of pectoral fin. Dorsal fin with i,6 (22) rays, and fin margin straight. Pectoral fin with i,9 (12) or i,10 (9) rays with first ray unbranched and greatly thickened. Outer part of posterior margin of pectoral fin straight, with 4-5 innermost rays progressively shorter making inner part of posterior fin margin rounded. Pelvic fin inserted posteriorly to dorsal-fin base. Pelvic fin with i,5 (22) rays with first ray unbranched and greatly thickened. Pelvic fin with straight posterior margin. Adipose-fin base longer than anal-fin base, origin anterior to origin of anal-fin base, extending beyond posterior end of anal-fin base. Margin strongly convex with sharply rounded edge, not deeply incised posteriorly. Caudal fin deeply emarginate or forked with tips of lobes rounded, with i,5,6,i (1), i,6,7,i (1), i,7,7,i (2), i,7,8,i (16) or i,8,8,i (1) principal rays. Anal fin with short base, origin posterior to origin of adipose-fin base, with iii, 5 (7), iii,6 (13) or iii, 7 (2) rays. Anal fin margin almost straight. 41

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Coloration Dorsal and lateral surfaces of head and body brown with many small black spots. Ventral region of head dark yellow or brown. Ventral region of body brown to dark yellow. Light dorsal saddles absent. Dorsal, adipose, caudal and anal fins brown. Dorsal surfaces of pectoral and pelvic fins brown with light distal edge, ventral surfaces light yellow. Small black spots present on all fins. Maxillary and mandibular barbels brown. Caudal fin with dark crescent shaped band at base. Large juvenile coloration similar to that of adults. Distribution Known only from the Galana River drainage, Kenya. 42

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Table 4-1. External traits diagnostic for species of the Amphilius uranoscopus group in east-central Africa. Character A. grandis A. krefftii A. oxyrhinus A. uranoscopus Amphilius sp. 1 Visible epidermal core extending posteriorly from the mandibular barbels Absent Present Present Absent Absent Caudal-fin shape Deeply emarginate or forked Emarginate Forked Emarginate Forked Caudal lobes Pointed Rounded Pointed Rounded Rounded Dark spots on body (adult) Absent Absent Large Absent Small Dark spots on head Absent Absent Usually absent, indistinct if present Absent Present Spots on fins Absent Absent Absent Absent Present Dorsal saddles Absent Present Absent Present Absent Branchiostegal rays 7-8 Usually 7 7-9 8-9 6-7 43

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Table 4-2. Morphometric data for Amphilius grandis. Range and mean include the syntypes. Syntypes (n=2) Range (n=21) MeanSD %SL Head length 25.8-27.3 23.1-27.8 25.8.1 Head width 21.1-22.1 19.7-23.5 21.31.1 Head height 12.9-13.0 11.0-14.9 13.51.0 Body depth 14.6-15.4 12.2-17.6 13.91.2 Body depth at anus 11.9-14.4 11.6-14.4 12.9.7 Predorsal length 36.1-37.8 33.7-40.4 37.21.4 Prepectoral length 22.3-23.8 19.6-24.5 21.91.4 Preanal length 75.1-77.1 65.7-78.4 73.42.6 Dorsal-fin base length 8.4-9.2 7.9-13.9 10.2.5 Adipose-fin base length 18.6-19.0 15.0-24.9 20.42.6 Anal-fin base length 8.2-8.9 8.2-17.8 11.1.3 Pelvic-fin length 14.6-16.7 14.6-20.8 18.61.8 Pectoral-fin length 17.1-18.4 17.1-26.5 21.52.5 Anal-fin length 14.3-15.1 14.3-21.1 17.91.9 Caudal peduncle length 18.6-19.0 14.6-19.0 16.8.3 Caudal peduncle depth 9.9-10.1 9.3-12.6 10.6.8 Anus-to-anal fin length 13.8-14.0 8.6-14.0 11.3.7 Prepelvic length 55.9-56.3 48.5-56.4 53.31.8 Postpelvic length 47.0-48.2 47.0-51.0 48.91.1 End-of-dorsal to adipose fin 42.2-44.9 38.9-48.9 43.2.8 Front-of-dorsal to caudal fin 62.1-64.9 57.1-70.8 64.2.6 Preanus length 61.0-62.5 58.9-64.1 61.71.4 %HL Snout length 48.5-51.3 43.6-57.4 48.64.1 Interorbital distance 26.1-26.7 24.0-31.2 27.21.9 Maxillary barbel length 57.0-58.3 55.6-81.3 66.76.5 Inner mandibular barbel length 24.5-30.6 21.4-39.6 30.9.3 Outer mandibular barbel length 39.1-43.5 39.1-60.8 48.6.6 Eye diameter 10.8-11.9 8.0-17.3 13.4.7 Postorbital length 35.5-39.6 31.2-43.3 37.53.0 44

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Table 4-3. Morphometric data for Amphilius krefftii. Range and mean include the syntypes. Syntypes (n=2) Range (n=19) MeanSD %SL Head length 22.7-22.8 21.9-25.9 23.81.1 Head width 20.2-20.6 18.7-23.6 21.21.1 Head height 12.6-12.8 9.1-14.7 11.7.7 Body depth 11.6-11.6 9.1-17.2 12.2.3 Body depth at anus 13.5-13.5 10.8-15.5 12.9.3 Predorsal length 34.3-34.5 31.8-36.9 34.71.5 Prepectoral length 19.8-20.8 17.4-23.9 20.11.8 Preanal length 72.8-73.7 62.5-76.3 72.13.9 Dorsal-fin base length 8.8-9.5 8.2-13.5 10.4.5 Adipose-fin base length 21.0-22.7 19.1-29.8 23.23.0 Anal-fin base length 10.6-10.7 8.8-15.0 11.2.5 Pelvic-fin length 19.3-19.8 17.6-20.6 19.50.9 Pectoral-fin length 19.0-21.3 19.0-24.8 21.51.4 Anal-fin length 16.2-17.9 15.4-21.7 18.62.1 Caudal peduncle length 17.3-18.6 14.9-18.6 16.5.0 Caudal peduncle depth 10.5-11.5 8.1-12.6 11.1.9 Anus to anal fin length 9.3-11.9 7.2-13.9 11.1.8 Prepelvic length 52.1-52.6 49.4-54.5 52.61.5 Postpelvic length 47.4-50.0 46.8-50.8 48.91.1 End of dorsal to adipose 46.9-50.2 42.5-50.8 47.1.5 Front of dorsal to caudal 65.5-67.0 63.4-71.3 66.9.0 Preanus length 61.3-63.2 59.3-64.8 62.31.7 %HL Snout length 45.4-45.4 45.4-53.1 49.62.4 Interorbital distance 24.5-26.9 23.2-31.3 26.92.4 Maxillary barbel length 72.2-86.7 52.6-95.9 73.712.2 Inner mandibular barbel length 28.1-30.6 20.7-47.8 30.8.2 Outer mandibular barbel length 48.1-55.6 36.8-68.6 53.9.7 Eye diameter 12.8-16.7 11.8-19.5 15.41.8 Postorbit length 34.2-40.7 34.2-43.1 38.72.1 45

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Table 4-4. Morphometric data for Amphilius oxyrhinus. Range and mean include the holotype. Holotype Range (n=14) MeanSD %SL Head length 25.5 23.1-27.0 25.5.1 Head width 22.0 19.5-23.9 21.5.1 Head height 11.8 11.3-14.6 12.7.9 Body depth 13.2 9.6-15.2 12.9.7 Body depth at anus 11.6 11.0-14.2 12.9.0 Predorsal length 38.5 35.2-40.3 36.7.5 Prepectoral length 23.4 17.3-23.4 20.6.8 Preanal length 75.4 70.6-76.1 74.1.8 Dorsal-fin base length 8.4 8.4-12.7 10.9.2 Adipose-fin base length 15.6 15.6-26.0 21.5.9 Anal-fin base length 9.3 9.3-15.0 11.4.5 Pelvic-fin length 14.9 14.9-22.1 19.9.7 Pectoral-fin length 17.8 17.8-26.2 22.9.0 Anal-fin length 15.8 15.8-19.5 18.0.1 Caudal peduncle length 17.0 14.8-18.0 16.9.9 Caudal peduncle depth 9.0 9.0-12.1 10.9.9 Anus to anal fin length 13.0 9.5-14.1 11.7.3 Prepelvic length 55.6 49.9-57.4 53.0.0 Postpelvic length 46.0 46.0-52.1 49.1.0 End of dorsal to adipose 42.7 41.1-48.6 44.2.1 Front of dorsal to caudal 63.3 61.5-68.3 64.7.4 Preanus length 63.0 59.8-66.0 61.8.7 %HL Snout length 53.2 47.4-54.5 51.1.5 Interorbital distance 23.0 23.0-33.3 28.0.1 Maxillary barbel length 51.9 51.9-85.5 73.88.5 Inner mandibular barbel length 31.3 21.1-59.1 34.0.7 Outer mandibular barbel length 42.7 30.4-64.2 51.8.7 Eye diameter 8.1 8.1-16.5 13.5.6 Postorbit length 38.9 33.4-42.3 38.5.6 46

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Table 4-5. Morphometric data for Amphilius uranoscopus. Range and mean include the specimens from the Wami drainage. Wami dr. (n=3) Range (n=55) MeanSD %SL Head length 24.9-26.1 23.4-30.1 25.81.4 Head width 22.7-23.3 19.3-25.5 21.31.4 Head height 12.3-13.6 10.0-16.4 13.31.3 Body depth 13.8-14.8 9.5-16.7 12.7.6 Body depth at anus 12.9-15.0 9.6-16.0 11.8.2 Predorsal length 36.0-36.2 33.8-40.8 36.61.5 Prepectoral length 20.3-21.4 17.0-25.4 20.61.9 Preanal length 73.2-74.9 60.2-78.6 72.13.9 Dorsal-fin base length 9.8-10.2 7.9-12.7 10.3.1 Adipose-fin base length 20.8-25.0 19.1-29.3 24.72.4 Anal-fin base length 9.2-10.6 7.0-19.4 11.8.4 Pelvic-fin length 18.2-20.6 15.0-21.0 18.41.5 Pectoral-fin length 19.9-23.5 17.7-23.8 21.31.3 Anal-fin length 18.0-19.4 15.4-21.6 18.31.7 Caudal peduncle length 16.3-18.4 14.5-19.7 17.0.1 Caudal peduncle depth 10.4-13.2 7.8-13.2 9.8.1 Anus to anal fin length 10.2-11.2 7.5-13.5 10.4.2 Prepelvic length 50.9-53.2 48.6-57.6 52.71.6 Postpelvic length 47.6-50.0 44.4-54.1 49.31.9 End of dorsal to adipose 42.4-47.0 39.2-49.8 44.2.5 Front of dorsal to caudal 63.9-67.3 63.2-72.5 66.3.0 Preanus length 60.6-64.1 58.0-69.4 62.32.0 %HL Snout length 46.6-51.7 40.1-52.4 46.13.2 Interorbital distance 27.8-30.3 23.7-32.3 28.33.3 Maxillary barbel length 54.5-82.8 54.5-112.8 78.711.1 Inner mandibular barbel length 29.4-34.3 22.6-45.4 33.5.5 Outer mandibular barbel length 53.5-64.9 36.5-77.4 55.2.4 Eye diameter 12.1-14.9 9.5-20.0 14.7.2 Postorbit length 37.7-39.6 31.0-45.7 38.33.1 47

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Table 4-6. Morphometric data for Amphilius sp. 1. Range and mean include the holotype. Holotype Range (n=21) MeanSD %SL Head length 26.9 24.0-29.3 25.8.2 Head width 22.7 19.7-26.2 21.9.4 Head height 11.3 11.3-14.9 12.7.0 Body depth 13.0 12.4-17.3 14.8.4 Body depth at anus 11.8 11.1-15.3 13.3.1 Predorsal length 37.9 33.3-39.6 37.2.5 Prepectoral length 24.1 19.5-25.6 22.0.7 Preanal length 79.6 60.8-79.6 73.2.2 Dorsal-fin base length 9.2 8.0-11.7 9.5.9 Adipose-fin base length 18.7 18.5-24.3 20.5.8 Anal-fin base length 8.7 8.6-16.6 10.4.8 Pelvic-fin length 16.1 16.0-19.7 17.7.2 Pectoral-fin length 16.1 17.4-23.4 19.7.5 Anal-fin length 18.7 14.5-19.2 17.0.4 Caudal peduncle length 15.7 14.6-18.3 16.5.1 Caudal peduncle depth 15.8 10.1-12.6 11.0.7 Anus to anal fin length 13.0 10.4-14.1 12.1.9 Prepelvic length 56.1 50.9-58.3 54.2.9 Postpelvic length 49.9 42.8-51.2 47.2.3 End of dorsal to adipose 45.3 42.3-49.9 45.2.7 Front of dorsal to caudal 64.9 62.4-67.6 65.4.1 Preanus length 66.3 58.1-66.3 62.0.0 %HL Snout length 53.8 46.7-56.6 52.5.3 Interorbital distance 26.8 23.9-37.0 28.7.1 Maxillary barbel length 68.1 60.4-90.6 75.07.7 Inner mandibular barbel length 37.7 26.9-43.9 36.8.2 Outer mandibular barbel length 61.7 49.3-71.9 59.0.0 Eye diameter 9.0 9.0-15.3 11.4.0 Postorbit length 36.6 35.6-43.1 39.8.2 48

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Figure 4-1. Amphilius grandis, BMNH 1912.3.22.119, 113.6 mm SL; lateral, dorsal and ventral view. 49

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Figure 4-2. Known distribution of Amphilius grandis. Some symbols represent more than one collection site. T denotes type locality. 50

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Figure 4-3. Amphilius krefftii, BMNH, 1909.10.19.26-27, Syntype 86.0 mm SL; lateral, dorsal and ventral view. Photograph by Mark Allen. 51

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Figure 4-4. Known distribution of Amphilius krefftii. Some symbols represent more than one collection site. T denotes type locality. 52

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Figure 4-5. Amphilius oxyrhinus, BMNH 1912.3.22.120, Holotype, 166.0 mm SL; lateral, dorsal and ventral view. Photograph by Mark Allen. 53

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Figure 4-6. Known distribution of Amphilius oxyrhinus. Some symbols represent more than one collection site. T denotes type locality. 54

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Figure 4-7. Amphilius uranoscopus, CAS 80494, 101.4 mm SL; lateral, dorsal and ventral view. 55

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Figure 4-8. Known distribution of Amphilius uranoscopus. Some symbols represent more than one collection site. 56

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Figure 4-9. Amphilius sp. 1, BMNH 1910.10.31.31, Holotype, 150.0 mm SL; lateral, dorsal and ventral view. 57

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Figure 4-10. Known distribution of Amphilius sp. 1. Some symbols represent more than one collection site. T denotes type locality. 58

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CHAPTER 5 RESULTS The present study examined all available museum specimens of the genus Amphilius from east-central Africa and revealed the existence of at least five species of the Amphilius uranoscopus group in the region. Previously, only one species of the Amphilius uranoscopus group was recognized from the region. Three species previously believed to be synonyms of Amphilius uranoscopus are determined to be valid species, and one species new to science is described. Additionally, pigment differences were noticed in the A. uranoscopus from the Wami, Ruvu and Rufiji drainages, but because only a few adult specimens are available from these drainages only a single species is recognized from these drainages. The recognition of Amphilius uranoscopus as a single widespread species has been based largely on the lack of clear meristic and morphometric differences between populations. Additionally, previous studies never looked at large number of specimens from the east-central region. Species that had been described from the region were based on just a few specimens, and only the type specimens of those species had been compared. Although differences in pigment pattern and caudal-fin shaped have been noticed previously (Copley, 1958; Seegers, 1996b), this is the first study to examine variation in these characters thoroughly. Although there is some degree of intraspecific variation in pigment pattern, certain aspects are consistent and are useful in distinguishing the species. Amphilius oxyrhinus and Amphilius sp. 1 both have dark markings on the body in the form of blotches or spots as adults and juveniles. Adults of A. grandis lack dark markings on the body, but juveniles have dark spots. Dark markings are generally absent in adults and juveniles of A. krefftii and A. uranoscopus. Amphilius krefftii may have a few dark areas on its body but never has distinct dark blotches or spots. In Amphilius sp. 1, the dark markings are always present in the form of 59

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small dark spots (although a few larger blotches may be present), and spots are also always present on the head. In A. oxyrhinus, the dark markings are always in the form of large blotches or spots, with only a few smaller spots or blotches present. Spots on the head are usually absent or indistinct. The pectoral, pelvic, dorsal, anal, adipose and caudal fins of Amphilius sp. 1 are also heavily spotted. Amphilius oxyrhinus lacks spots on the fins, although some dark pigment may be present. Light dorsal saddles at the base of the dorsal and adipose fins are present in A. krefftii and A. uranoscopus (except specimens from the Ruvu drainage, see discussion below). These light dorsal saddles are not present in A. grandis, A. oxyrhinus or Amphilius sp. 1. Two forms of the caudal fin were observed. In Amphilius grandis, A. oxyrhinus and Amphilius sp. 1, the caudal fin is deeply emarginate or moderately forked, but in A. krefftii and A. uranoscopus the caudal fin is feebly emarginate. In A. grandis and A. oxyrhinus, the tips of the caudal-fin lobes are pointed, while in Amphilius sp. 1, A. krefftii and A. uranoscopus the tips of the caudal-fin lobes are noticeably more rounded. A visible epidermal core extending posteriorly from the mandibular barbels is always present in Amphilius oxyrhinus and A. krefftii and always absent in A. grandis, Amphilius sp. 1 and A. uranoscopus. The visible epidermal core is a vein-like structure extending posteriorly from the bases of the inner and outer mandibular barbels. The structure is pigmented, but even in old specimens in which the pigment has faded, the structure is still noticeable. Frequency tables were constructed for counts of eight different meristic characters. Branchiostegal ray count was the only meristic character found to be useful for distinguishing species. Amphilius sp. 1 had the fewest branchiostegal rays, with all specimens having 6 or 7 branchiostegal rays. Amphilius uranoscopus had the most, with all specimens having 8 or 9 branchiostegal rays. Branchiostegal ray count usually distinguishes A. krefftii from A. 60

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uranoscopus; 18 specimens of A. krefftii had a count of 7 and one specimen had 8 branchiostegal rays (Table 5-1). Total gill-raker counts did not distinguish between species, but the counts for A. uranoscopus in the Wami, Ruvu and Rufiji did indicate differences in those populations (Table 5-2). No clear patterns were observed in the frequency-distribution tables of branched pectoral-fin rays, branched anal-fin rays, total vertebrae, preanal vertebrae, postanal vertebrae and first dorsal pterygiophore intercept count (Tables 5-3 to 5-8). The characters examined in this study offer some preliminary insights into the possible relationships among species of the Amphilius uranoscopus group in east-central Africa and the biogeography of the region. Amphilius grandis, A. oxyrhinus and Amphilius sp. 1, the species with deeply emarginate or forked caudal fins, are distributed in the northern part of the east-central region, while all A. krefftii and A. uranoscopus populations are distributed to the south of these species. Additionally, A. grandis, A. oxyrhinus and Amphilius sp. 1 all have dark markings either as adults or juveniles, while A. krefftii and A. uranoscopus never have dark markings. Finally, light dorsal saddles are absent in the northern species, but present in the southern species. The caudal fin shape and pigment patterns observed suggest that a northern and a southern group of species exist in east-central Africa. These northern and southern species groups correspond to the physical geography of the region. The Eusso Nyiro, Tana and Galina drainages all have high-gradient tributaries in the Mount Kenya-Aberdare highlands (Skelton, 1994). Amphilius grandis and A. oxyrhinus overlap in distribution in the Eusso Nyiro and Tana drainages. Amphilius sp. 1 is restricted to the upper reaches of the Galina drainage in close proximity to the Eusso Nyiro and Tana drainages. 61

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The Pangani, Ruvu and Wami drainages drain the south-east slopes of Kilimanjaro and the eastern side of the Gregory Rift (Skelton, 1994). The present-day Rufiji drainage has grown from a series of captures (Willis, 1936). Its largest tributary may once have drained into the Indian Ocean via the Wami (Bailey, 1969) or the Pangani (Haldemann, 1962). Although Amphilius krefftii also occurs in the Galina drainage, it is found only in tributaries of the lower reaches of the river. These tributaries are in close proximity to the Pangani drainage, where the species also occurs. The Amphilius in the Wami, Ruvu and Rufiji drainages are tentatively recognized here as a single species despite differences observed in pigment pattern and intensity. Because most of the specimens examined were juveniles, only juvenile pigmentation could be compared. All specimens examined from the Ruvu drainage, including the type of Chimarrhoglanis leroyi were lighter in color than specimens from the Wami and Rufiji (Fig. 5-1). Additionally, these specimens did not have the light dorsal saddles that were always observed in specimens from the Wami and Rufiji. Given that amount of variation in darkness observed in other species of Amphilius, the lighter color of the Ruvu population is not considered a significant taxonomic character. Additionally, the lighter color of the type of C. leroyi may be due to the age and preservation of the specimen, and the absence of light dorsal saddles in all specimens examined may only be because of the lighter color of these specimens. The Wami and Rufiji populations were similar in body coloration, but specimens from the Wami always had darker fins. Although this difference was consistent, no additional differences were observed. Although specimens from the different populations appeared to differ in body shape, no morphometric differences were found. 62

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The status of three other species synonymized with Amphilius uranoscopus, A. hargeri, A. brevidorsalis and A. platychir cubangoensis, are beyond the geographic scope of this study, but tentative conclusions can be made about the identity of these species. The type of A. hargeri is similar to A. uranoscopus, but the first-dorsal pterygiophore intercepts the third post-Weberian vertebra. Skelton (1984) hypothesized that it could be a hybrid specimen, but concluded that it was best considered a junior synonym of A. uranoscopus. This character separates A. hargeri from the species in east-central Africa. All the specimens examined in east-central Africa have the dorsal pterygiophore intercepting the first or second post-Weberian vertebra, with the majority of specimens examined having the dorsal pterygiophore intercepting the first post-Weberian vertebra. The type of Amphilius brevidorsalis from Mozambique is most similar to A. uranoscopus and may be a synonym of A. uranoscopus. The type of A. brevidorsalis lacks dark markings and has distinct light dorsal saddles. Additionally it a visible epidermal core extending posteriorly from the mandibular barbels and has a caudal fin shape characteristic of A. uranoscopus. The type specimens examined of A. platychir cubangoensis have a caudal fin shape similar to that of A. krefftii and A. uranoscopus, but differ from these species by having distinct dark markings on the body. Additionally, a visible epidermal core extending posteriorly from the mandibular barbelsis present in the type specimens examined, further distinguishing A. platychir cubangoensis from A. uranoscopus. 63

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Table 5-1. Branchiostegal ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Branchiostegal Rays 6 7 8 9 A. oxyrhinus 4 9 2 A. grandis 9 8 Amphilius sp. 1 1 21 A. krefftii 18 1 A. uranoscopus (Wami) 3 A. uranoscopus (Ruvu) 6 5 A. uranoscopus (Rufiji) 17 9 Table 5-2. Total gill raker counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Total Gill Rakers 6 7 8 9 10 11 A. oxyrhinus 1 3 10 A. grandis 1 2 2 8 7 1 Amphilius sp. 1 1 7 11 2 A. krefftii 5 5 6 1 A. uranoscopus (Wami) 1 1 1 A. uranoscopus (Ruvu) 1 4 3 3 A. uranoscopus (Rufiji) 6 23 8 1 Table 5-3. Branched pectoral-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Branched Pectoral-Fin Rays 8 9 10 11 A. oxyrhinus 12 2 A. grandis 3 17 1 Amphilius sp. 1 12 9 A. krefftii 11 7 1 A. uranoscopus (Wami) 2 1 A. uranoscopus (Ruvu) 8 4 A. uranoscopus (Rufiji) 3 14 23 Table 5-4. Branched anal-fin ray counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Branched Anal-Fin Rays 5 6 7 8 A. oxyrhinus 14 A. grandis 21 Amphilius sp. 1 7 13 2 A. krefftii 17 2 A. uranoscopus (Wami) 1 2 A. uranoscopus (Ruvu) 6 6 A. uranoscopus (Rufiji) 2 17 10 11 64

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Table 5-5. Total vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Total Vertebrae 36 37 38 39 40 A. oxyrhinus 4 4 A. grandis 3 6 1 Amphilius sp. 1 2 7 3 A. krefftii 7 3 1 A. uranoscopus (Wami) 3 A. uranoscopus (Ruvu) 2 2 A. uranoscopus (Rufiji) 11 12 2 Table 5-6. Preanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Preanal Vertebrae 19 20 21 22 A. oxyrhinus 7 1 A. grandis 1 3 6 Amphilius sp. 1 2 3 7 A. krefftii 7 5 A. uranoscopus (Wami) 1 1 1 A. uranoscopus (Ruvu) 2 2 A. uranoscopus (Rufiji) 1 6 11 7 Table 5-7. Postanal vertebrae counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus Postanal Vertebrae 15 16 17 18 A. oxyrhinus 5 3 A. grandis 1 4 5 Amphilius sp. 1 1 5 4 2 A. krefftii 1 9 1 A. uranoscopus (Wami) 1 1 1 A. uranoscopus (Ruvu) 4 A. uranoscopus (Rufiji) 9 15 1 Table 5-8. First dorsal pterygiophore intercept counts in A. oxyrhinus, A. grandis, Amphilius sp. 1, A. krefftii and three populations of A. uranoscopus 1st DPI 1 2 A. oxyrhinus 4 4 A. grandis 2 8 Amphilius sp. 1 12 1 A. krefftii 11 1 A. uranoscopus (Wami) 3 A. uranoscopus (Ruvu) 4 A. uranoscopus (Rufiji) 23 1 65

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Figure 5-1. Amphilius uranoscopus (juveniles) from the Wami (top: CAS 80494, 49.1 mm SL), Ruvu (middle: UF 84882, 40.4 mm SL) and Rufiji (bottom: FMNH 111680, 46.0 mm SL) drainage. 66

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LIST OF REFERENCES BAILEY, R. G. 1969. The non-cichlid fishes of the eastward flowing rivers of Tanzania, East Africa. Revue de Zoologie et Botanique Africaines 80:170-199. BELL-CROSS, G. 1972. The fish fauna of the Zambezi River system. Arnoldia (Rhodesia) 5:1-19. BELL-CROSS, G., AND R. A. JUBB, 1973. The Amphiliidae of southern Africa and record of Amphilius lampei Pietschmann, 1913, from the Inyanga Mountains, Rhodesia. Aronoldia (Rhodesia) 6:1-9. BERNACEK, G. M. 1980. Introduction to the freshwater fishes of Tanzania. University of Dar es Salaam, Deptment of Zoology, Dar es Salaam. BERRA, T. M. 2001. Freshwater Fish Distribution. Academic Press, San Diego, California. BERTIN, L., AND R. ESTVE. 1950. Catalogue des Types de Poissons du Musum National d'Histoire Naturelle. 5 e partie. Ostariophysaires (Siluriformes). Imprimerie Nationale, Paris. BOULENGER, G. A. 1898a. On the habit of the siluroid fish Anoplopterus platychir, Gthr. Annals and Magazine of Natural History (Ser. 7) 8:447-448. BOULENGER, G. A. 1898b. Report on the collection of fishes made by Mr. J. E. S. Moore in Lake Tanganyika during his expedition, 1895-96. Transactions of the Zoological Society of London 15:1-30, pls. 1-8. BOULENGER, G. A. 1901. Description of a new silurid fish of the genus Anoplopterus, from Cameroon. Annals and Magazine of Natural History (Ser. 7) 8: 447-448. BOULENGER, G. A. 1902. Additions la faune ichthyologique de bassin du Congo. Matriaux pour la faune du Congo. Annales du Muse du Congo, Zoologie 2:19-57, pls. 7-16. BOULENGER, G. A. 1905a. On a second collection of fishes made by Mr. S. L. Hinde in the Kenya District, East Africa. Proceedings of the Zoological Society of London 1:62-64, pl. 7. BOULENGER, G. A. 1905b. A list of freshwaters fishes of Africa. Annals and Magazine of Natural History (Ser. 7) 16: 36-60. BOULENGER, G. A. 1907a. Descriptions of three new fishes from Central Africa. Annals and Magazine of Natural History (Ser. 7) 20: 487-489. BOULENGER, G. A. 1907b. On a collection of fishes, batrachians and reptiles, made by Mr. S. A. Neave in Rhodesia, north of the Zambesi with field notes by the collector. Memoirs and Proceedings of the Manchester Literary & Philosophical Society 51: 1-12. 67

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BOULENGER, G. A. 1911. Catalogue of the fresh-water fishes of Africa in the British Museum (Natural History), Vol. 2. British Museum (Natural History) Trustees, London. BOULENGER, G. A. 1912. On a collection of fishes made by Mr. A. Blayney Percival in British East Africa to the East of Lake Baringo. Proceedings of the Zoological Society of London 1912:672-676, pls. 78-80. BOULENGER, G. A. 1916. Catalogue of the fresh-water fishes of Africa in the British Museum (Natural History), Addenda. British Museum (Natural History) Trustees, London. CAILLIET, G. M., M. S. LOVE AND A. W. EBELING. 1986. Fishes: a field and laboratory manual on their structure, identification, and natural history. Wadsworth Publishing Company, Belmont, CA. COPLEY, H. 1941. A short account of the freshwater fishes of Kenya. Journal of the East Africa and Uganda Natural History Society 16:24 pp., 8 pls. COPLEY, H. 1958. Common freshwater fishes of east Africa. H. F. & G. Witherby Ltd., London. CRASS, R. S. 1960. Notes on the freshwater fishes of Natal with descriptions of four new species. Annals of the Natal Museum 14: 405-458. CRASS, R. S. 1964. Freshwater fishes of Natal. Pietermaritzburg: Shuter & Shooter. DAS, D., AND T. C. NAG. 2004. Adhesion by paired pectoral and pelvic fins in a mountain-stream catfish, Pseudocheneis sulcatus (Sisoridae). Environmental Biology of Fishes 71:1. DAS, D., AND T. C. NAG. 2005. Structure of adhesive organ of the mountain-stream catfish, Pseudocheneis [sic.] sulcatus (Teleostei: Sisoridae). Acta Zoologica 86:231-237. FERRARIS, C. J., Jr. 2007. Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types. Zootaxa 1418:1-628. GNTHER, A. 1864. Catalogue of the Fishes in the British Museum, vol. 5. Catalogue of the Physostomi, Containing the Families Siluridae, Characinidae, Haplochitonidae, Sternoptychidae, Scopelidae, Stomiatidae in the Collection of the British Museum. Trustees, London. GNTHER, A. 1865. Pisces. The record of zoological literature. Zoological Record 1864:133-188. GNTHER, A. 1902. Last account of fishes collected by Mr. R. B. N. Walker, C. M. Z. S., on the Gold Coast. Proceeding of the Zoological Society of London 2:330-339. 68

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HARRY, R. R. 1953. A contribution to the classification of the African catfishes of the family Amphiliidae, with description of collections from Cameroon. Revue de Zoologie et de Botanique Africaines 47: 177-200; 201-232. HALDEMANN, E. G. 1962. The geology of the Rufiji Basin, with reference to proposed dam sites in eastern and southern Tanganyika. Bulletin Geological Survey of Tanganyika 33:1-250. HILGENDORF, F. M. 1905. Fische von Deutsch und Englisch Ost-Africa. Zoologische Jahrbcher. Abteilung fr Systematik, Geographie und Biologie der Tiere (Jena) 22:405-420. JACKSON, P. B. N. 1959. New records and little-known species of fish from Rhodesia and Nyasaland. Occasional papers of the National Museums of Southern Rhodesia, Natural Sciences 3:295-305. JACKSON, P. B. N. 1961a. Check-list of the fishes of Nyasaland. Occasional Papers of the National Museums of Southern Rhodesia, Natural sciences 3:535-621. JACKSON, P. B. N. 1961b. The fishes of Northern Rhodesia: a checklist if indigenous species. Lusaka: The Government Printer. JUBB, R. A. 1961. An illustrated guide to the freshwater fishes of the Zambezi River, Lake Kariba, Pungwe, Subi, Lundi and Limpopo Rivers. Bulawayo: Stuart Manning. JUBB, R. A. 1963. A revised list of the freshwater fishes of southern Africa. Annals of the Cape Provincial Museums (Natural History) 3:40-43. MAAR, A. 1960. Introductory check list of fish of the Ethiopian region. Proceedings of the Federal Science Congress 1:1-8. MARRIOTT, M.S., A. J. BOOTH, AND P. H. SKELTON. 1997. Reproductive and feeding biology of the Natal mountain catfish Amphilius natalensis (Siluroidei: Amphiliidae). Environmental Biology of Fishes 49:461-470. MATTHES, H. 1967. The fishes and fisheries of the Ruaha River basin, Tanzania. East African Freshwater Fisheries Research Organisation (EAFFO) Occasional Papers No. 9:1-19. PELLEGRIN, J. 1905. Mission scientifique de Ch. Alluaud en Afrique orientale (Juin 1903 -Mai 1904). Poissons. II. Systmatique. Mmoires de la Socit Zoologie de France 17: 167-185, pl. 16. PELLEGRIN, J. 1936. Contribution l'Ichtyologie de l'Angola. Arquivos do Museu Bocage, Lisboa 7:45-62. PFEFFER, G. J. 1889. bersicht der von Herrn Dr. Franz Stuhlmann in gypten, auf Sanzibar und dem gegenberliegenden Festlande gesammelten Reptilien, Amphibien, Fische, 69

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Mollusken und Krebse. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Hamburg 6:1-36. PFEFFER, G. J. 1893. Ostafrikanische Fische gesammelt von Herrn Dr. F. Stuhlmann im Jahre 1888 und 1889. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Hamburg 10:131-177, 3 pls. PFEFFER, G. J. 1896. Die Fische Ost-Afrikas, p. xviii + 72 p. In: Die Thierwelt Ost-Afrikas und der Nachbargebiete. Vol. 1, Lief. V. K. Mbius (ed.), D. Reimer, Berlin. POCHE, F. 1902a. ber das Vorkommen von Anoplopterus platychir (Gthr) in Westafrika. Zoologischer Anzeiger 25:121-122. POCHE, F. 1902b. Richtigstellung eines Gattungsnamens unter den Siluriden. Zoologischer Anzeiger 25:211. POLL, M. 1952. Poissons de rivires de la rgion des lacs Tanganika et Kivu recuellis par G. Marlier. Revue de Zoologie et de Botanique Africaines 46:221-236. RICARDO, C. K. 1939a. The fishes of Lake Rukwa. Journal of the Linnean Society of London, Zoology 40:625-657. RICARDO, C. K. 1939b. Report on the fish and fisheries of Lake Rukwa in Tanganika Territory and the Bangweulu regions in Northern Rhodesia. Crown Agent, London. ROBERTS, T. R. 1975. Geographical distribution of African freshwater fishes. Zoological Journal of the Linnean Society 57:249-319. ROBERTS, T. R. 2003. Systematics and osteology of Leptoglaninae, a new subfamily of the African catfish family Amphiliidae, with descriptions of three new genera and six new species. Proceedings of the California Academy of Sciences 54:81-132. SEEGERS, L. 1996a. The fishes of the Lake Rukwa drainage. Annales de la Socit Zoologique de Belgique 278:1-407. SEEGERS, L., 1996b. Die Amphilius-Arten Ostafrikas. Die Aquarienund Terrarienzeitschrift (DATZ) 49:249-255. SEEGERS, L., L. DE VOS AND D. O. OKEYO. 2003. Annotated checklist of the freshwater fishes of Kenya (excluding the lacustrine haplochromines from Lake Victoria). Journal of East African Natural History 92:11-47. SINGH, A., AND N. K. AGARWAL. 1991. SEM surface structure of the adhesive organ of the hillstream fish Glyptothorax pectinopterus (Teleostei: Sisoridae) from the Garhwal Hills. Functional and Developmental Morphology 1:11-13. 70

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BIOGRAPHICAL SKETCH Alfred W. Thomson was born December 30, 1978, in Englewood, New Jersey. One of three children, he was interested in the natural world from an early age and has always been especially interested in fishes. He received his bachelor of science degree from Mansfield University in August 2001, and moved to Kenosha, Wisconsin to work as a research technician for the Illinois Natural History Survey at the Lake Michigan Biological Station in Zion, Illinois. In January 2004 he moved to Gainesville, Florida and worked as a research technician on the All Catfish Species Inventory Project at the Florida Museum of Natural History. In August 2005 he enrolled in UFs Department of Zoology as a full-time graduate student with a research assistantship from the All Catfish Species Inventory Project. He is currently continuing his research on catfish systematics as a Ph.D. student in UFs Department of Zoology. 73