<%BANNER%>

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

Permanent Link: http://ufdc.ufl.edu/UFE0021228/00001

Material Information

Title: Taxonomic Revision of the Amphilius uranoscopus Group of East-Central Africa (Teleostei Amphiliidae)
Physical Description: 1 online resource (73 p.)
Language: english
Creator: Thomson, Alfred William
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: africa, amphiliidae, amphilius, taxonomy
Zoology -- Dissertations, Academic -- UF
Genre: Zoology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

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.
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.
Statement of Responsibility: by Alfred William Thomson.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Page, Lawrence M.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021228:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021228/00001

Material Information

Title: Taxonomic Revision of the Amphilius uranoscopus Group of East-Central Africa (Teleostei Amphiliidae)
Physical Description: 1 online resource (73 p.)
Language: english
Creator: Thomson, Alfred William
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: africa, amphiliidae, amphilius, taxonomy
Zoology -- Dissertations, Academic -- UF
Genre: Zoology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

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.
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.
Statement of Responsibility: by Alfred William Thomson.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Page, Lawrence M.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021228:00001


This item has the following downloads:


Full Text
xml version 1.0 encoding UTF-8
REPORT xmlns http:www.fcla.edudlsmddaitss xmlns:xsi http:www.w3.org2001XMLSchema-instance xsi:schemaLocation http:www.fcla.edudlsmddaitssdaitssReport.xsd
INGEST IEID E20101113_AAAAQW INGEST_TIME 2010-11-14T00:43:49Z PACKAGE UFE0021228_00001
AGREEMENT_INFO ACCOUNT UF PROJECT UFDC
FILES
FILE SIZE 4532 DFID F20101113_AADYPG ORIGIN DEPOSITOR PATH thomson_a_Page_16thm.jpg GLOBAL false PRESERVATION BIT MESSAGE_DIGEST ALGORITHM MD5
b7f752188505defe9913030ab8a79362
SHA-1
61203d744f5889fb6ce96331156bb5c9098cbd83
109893 F20101113_AADYFM thomson_a_Page_62.jp2
d0b1742dd8dfbfcf2e52b1c2221db966
34f3fa28ffd2f796a7871b3b9b981b283272a99f
29592 F20101113_AADYAP thomson_a_Page_05.jpg
ff8a1396ff7f6c558ff0332c82e380d8
18eaa653dccc23b1e786cf48a5fdd4609f1e4d27
42928 F20101113_AADYKJ thomson_a_Page_47.pro
9b45f450eed99ff016d1ec9890efaedb
47fade5a2971929945360e364903e52ea8e8294a
16109 F20101113_AADYPH thomson_a_Page_17.QC.jpg
959241bdc8b842c67374ed6895996003
ef2725811001f4e92076a5e5e777c4a710dbc787
97575 F20101113_AADYFN thomson_a_Page_63.jp2
e04928f3bfb9fddcf2f66d7e86469d3f
22d784d5cb17deb5b9a683a69f3a82117d4d506a
79274 F20101113_AADYAQ thomson_a_Page_06.jpg
e47afced252828db39d5c385ebdc804e
eb250a92abe32438b11f68ec2319e2d7da8757fd
45918 F20101113_AADYKK thomson_a_Page_48.pro
a97ce8b52a2e7e3c07d6d3ce6f58cf8c
1889378677b0fcb03c79b17b401a1a2c727e098a
4863 F20101113_AADYPI thomson_a_Page_17thm.jpg
b53d7708c2151bd1c099a22a1745c196
8dca48100b0ab18e7fe5e2051e5e863e71ad9aea
79735 F20101113_AADYFO thomson_a_Page_64.jp2
d06e63dfb69cfb0981c5dcece267918f
91d2183b9366e5b26abe31fb2f11c063025d8461
6169 F20101113_AADYKL thomson_a_Page_50.pro
2c98ec7cb512a07676bc0be0d239eb75
3b3ab29bf1024b47105559590e7c7bab9d66e92e
23422 F20101113_AADYPJ thomson_a_Page_18.QC.jpg
c6ba3f6ef74142a5e0f8ebbd005bae66
5bb84152346cd31a1d6fec775752a94895f3c73b
78942 F20101113_AADYFP thomson_a_Page_65.jp2
ef4db9dc4387653208a0b594cfcdc018
557392fb2dfddc60577ac3f746e86867cfc6dc65
81858 F20101113_AADYAR thomson_a_Page_07.jpg
c7545f5fe3ee6010f11b0ffb0096bd38
b8eafe93d670683f7244b35730d5719e98a390c5
3912 F20101113_AADYKM thomson_a_Page_51.pro
e34e95b7bb650188391929cbdfb7afd1
0213073cf6f84ec16884626b47140e47073ec97a
6465 F20101113_AADYPK thomson_a_Page_18thm.jpg
4a202173680e1509c39e6c1d8ab6ddeb
eb96e26af16381d94f4aa273200ec219f5dd53a3
876330 F20101113_AADYFQ thomson_a_Page_66.jp2
6513e66b327e15ca9f1e51146e814ef3
289dfbc2e5dc2cabaec129815f76e3436b540780
70886 F20101113_AADYAS thomson_a_Page_08.jpg
f5f58d9d2a18843ee5b7c0c0d2f38431
9b4e78b88b1b1f233af198db7bd4881be6d8581f
5716 F20101113_AADYKN thomson_a_Page_52.pro
2bcd145321a0384bd2c1b9a3a7e3048e
2f3cf8bd08b22090a5fb50291fa166532ee5b637
23302 F20101113_AADYPL thomson_a_Page_19.QC.jpg
b281ab83e458046bfa42667b9399e07f
ad4ff44329c10ffccf564e9b4d3013edaa2552bc
124437 F20101113_AADYFR thomson_a_Page_68.jp2
c6c1b726f3758a6e5b2033d938f60c9a
31e45fc5149c73116460f19ac203240db0041b6c
55224 F20101113_AADYAT thomson_a_Page_09.jpg
0d361b66740f23c1387eb4c6efa20699
9d41750f5f52d5497a86ac82ba95b802c178ecb7
3957 F20101113_AADYKO thomson_a_Page_53.pro
301dc8f1738222032a57fcae07e1aaf3
67d0a35222b039d4357a8c44573364c448852d23
6770 F20101113_AADYPM thomson_a_Page_19thm.jpg
4222eee0317d7cf6daedf0970f6a34df
8735af1628f5df83df1bce1f3c3aed49330d14e4
130686 F20101113_AADYFS thomson_a_Page_69.jp2
c5febb96daa58b348d281fb6d0a49554
3b8b477c5d9070cc12c3316e55b4af632611c0da
74458 F20101113_AADYAU thomson_a_Page_11.jpg
4694a56c399eb68a8a5eecd8b46a5731
6438c7be52ffaef641640972fd0c1c46af548dce
6192 F20101113_AADYKP thomson_a_Page_54.pro
106d33ad07785e78f87930baf2ea6e5a
dc7a05c5f2ff02c3084c29ed93ce2e838257065f
13966 F20101113_AADYPN thomson_a_Page_20.QC.jpg
731b50753fa3714df819b31746e641ee
7e075f67db96bf242690333ee2571f934f6c8782
122434 F20101113_AADYFT thomson_a_Page_70.jp2
15124795121a048b213474752a5f9716
3d94255005891b2d4440c7f495a99a812e1b98f8
78355 F20101113_AADYAV thomson_a_Page_12.jpg
fc81a738cc9b96ca2d76fc8eaaccc6e6
48fd62ba73754c28f3c043a11b4d6929f5ddae6b
2898 F20101113_AADYKQ thomson_a_Page_55.pro
31b3c43a75af749bb4e2e4dc8da3e134
40c1af3a75ebbf7c520950345c51a1b3da7d3c5d
4075 F20101113_AADYPO thomson_a_Page_20thm.jpg
86841271c33b5733082b5a8f7449d7a8
769fb7be9b12df285077fef844695b6e0ffc69b7
76439 F20101113_AADYAW thomson_a_Page_13.jpg
8b26cd8ad5dffd9a0fe86523944d7188
239ddeddd829435c18f43ac2f97084e602e6928e
5144 F20101113_AADYKR thomson_a_Page_56.pro
641d0b3dbb39559c30d8c222878f2824
4d3eb06ffc4221263f06b787ad7c3cb2ae037bca
1051967 F20101113_AADYFU thomson_a_Page_71.jp2
ef9bc5cb8c500cede0945d1b2eb81816
e45ad807e8d15e94ddaac170f98ca4084a9ee86c
23670 F20101113_AADYPP thomson_a_Page_21.QC.jpg
9f6a49c19c7accb27aa605c0bf2fc961
ceed3f1d8432714d15b2706deb035d7b8431d0c7
74630 F20101113_AADYAX thomson_a_Page_14.jpg
280ef24da83ddd7c7888da056535b302
3d6795d65a1925cecb33a55e227e4096bfc35ee0
3132 F20101113_AADYKS thomson_a_Page_57.pro
8df29ddd06904dec49069c0032d60954
c75158d96c13bc5790097c4685270fc96a8cf1c3
27742 F20101113_AADYFV thomson_a_Page_72.jp2
a57ca4ed5763d553c8e2550f87337967
f30b8e401511c0e0952364eb91f05ebe823578e3
6484 F20101113_AADYPQ thomson_a_Page_21thm.jpg
e7010b6da5a1f9b4e9dcf1e15ec97dbf
92f2284d48a5fbdb4318ecc4880eae66ff240e3f
16865 F20101113_AADYAY thomson_a_Page_15.jpg
9927a2205706788132e165c5b7405dec
a240b86f237f2e1fd0c0410d507053e6e8a3fb77
5648 F20101113_AADYKT thomson_a_Page_58.pro
706faac9dcca317261d7d6748fbaa463
5524a52fe8981e061875657c7c2d0a9cd8ff06d5
25043 F20101113_AADYPR thomson_a_Page_22.QC.jpg
8652a1a21a79b4c8527779b7be54149d
280551cbe992981a4e5a5ffc9314518518bebb47
50204 F20101113_AADYAZ thomson_a_Page_16.jpg
e1410423dd2a8fe853d0a439b671e8d5
640a17b5640a2e43a312e6cb027174d6e8b21a0a
53136 F20101113_AADYKU thomson_a_Page_59.pro
2c10f0ab9a2fac53c374a1bdc610ffd7
17dc72e5a14d917bd1c0a146c9c8a3da7e8b22d4
56099 F20101113_AADYFW thomson_a_Page_73.jp2
da67db0b41088c707806025b257b49ac
d428449f4032676dbc5e108daae1cffdc55fd7c2
17188 F20101113_AADYPS thomson_a_Page_23.QC.jpg
2201c4ba424ad95d9762b19d793a612b
762fd5bb41640dbbfd01ec5858d89c666ef5724f
55747 F20101113_AADYKV thomson_a_Page_60.pro
78119f730214724bff8635cec9ba2e4d
07498824530fd712067421ba3252534adba0f184
1053954 F20101113_AADYFX thomson_a_Page_01.tif
f848dab8d7246e7eb31baf16492bcb36
ae9e64afea22e3aac437e1f3086a8e1abf38086b
4854 F20101113_AADYPT thomson_a_Page_23thm.jpg
bdad6bc6207e4de761404200462787c4
06bc61fff5786ebcbeae4501312d2db4cc35a98c
50550 F20101113_AADYKW thomson_a_Page_61.pro
0054b985dd7d284cc589d40fdd2c3b0f
9083db0e56db34c8b0be8fb4c819798fa59a81d1
87125 F20101113_AADYDA thomson_a_Page_69.jpg
81a8d95657b94b74c2839fadcbe39f62
3b73c5270ca4efe0aff1de86e69db47829781ec0
F20101113_AADYFY thomson_a_Page_02.tif
ebaa4e02b71f68b1382ae9fee3c302a5
a71666d3810bfeff784c68b1bb5e32c78097bfcf
17134 F20101113_AADYPU thomson_a_Page_24.QC.jpg
3dc0da93bfc5bc6d661ec5c7a3b64c24
f37b1b079e65e03e266cd7f991972319f4b54024
49983 F20101113_AADYKX thomson_a_Page_62.pro
03a5db30e0971b5588dc39f3f2b1874f
a0e082ed266085b200fb38e1e935400039c3daf2
78441 F20101113_AADYDB thomson_a_Page_70.jpg
2d80921d8b11da867c5abddca5c7c553
c3bea38ab41444df190faf48ab9baf8f306978aa
F20101113_AADYFZ thomson_a_Page_03.tif
134c003d204cc7735502d1009cdc6730
95baebefd41510457b97490463a6d9ce4bbe3e77
4895 F20101113_AADYPV thomson_a_Page_24thm.jpg
32f57f3d44294ceb4b85e0e7d7aec48d
eb1e25f54e7c215822c43c8305975d1a67320466
43094 F20101113_AADYKY thomson_a_Page_63.pro
4723bcbf4c99b7afafb5bfa73e43e08f
6c78fcad8f1ed9cb7cb6f14dd534661fc691a69a
89081 F20101113_AADYDC thomson_a_Page_71.jpg
faa04f573fa48cbd5270029e90429719
efa67e897837a11efbc155d4e883a8d79f55ae7c
25511 F20101113_AADYPW thomson_a_Page_25.QC.jpg
10e67ae981240db40e7641a70f208726
eb276373bee6aa42f8156d8b3ce9b646b4731226
25271604 F20101113_AADYIA thomson_a_Page_57.tif
2676dacf77aa195c45ffa4617d3bd21f
b9072560fec54f8b1370304477cc551def03c89a
45385 F20101113_AADYKZ thomson_a_Page_64.pro
95fcde6722d9d9d93fdfb7242cbfb873
de1c5230ad237ea14a3e6295f5d99aa81f5f0441
22937 F20101113_AADYDD thomson_a_Page_72.jpg
d7ed3aa03f7758f22f7c84c38d922139
7c6ef18b5448604182ad68a4c173dca194eabd79
6768 F20101113_AADYPX thomson_a_Page_25thm.jpg
9a4de69688bc8af6aa011ab61a9fac92
fdc921343cae4494be352afe6d8afd734425eafb
F20101113_AADYIB thomson_a_Page_58.tif
caf52384cf9845b3ee05322c2e45f4c5
abc2ee6081d04c56341b1d439a920ec2343838dd
40409 F20101113_AADYDE thomson_a_Page_73.jpg
c0c250a36d9844a01dec1b1a3680a9a5
045ff9fa36eb749e7a4d9e13b6756ca0a16ab3e4
25044 F20101113_AADYPY thomson_a_Page_26.QC.jpg
b305fefdd6497f72bc894ce627de8b1a
f0e7b6d900244f6234eacec17d46e7aadcaa1b06
F20101113_AADYIC thomson_a_Page_59.tif
b300722193168fdd5b5f061c7cbb7f9e
2669859be33fd3d91f431b562068f2cbc9c2b628
25729 F20101113_AADYDF thomson_a_Page_01.jp2
e99d08295421ee51a3a4b8b3141fdf5c
e398eda91dcf39051c62b4ed87bb13bbbb739bde
6805 F20101113_AADYPZ thomson_a_Page_26thm.jpg
86177aaa6f24fad59f683dd76faee2e2
eac1e649bbb44bc8c767390e4136450b01343fd3
F20101113_AADYID thomson_a_Page_60.tif
f0cc1831eef2fdccc898e68cc3d3a63f
1029b24e9021cdf9f2b4b773d211d0241774038c
5834 F20101113_AADYDG thomson_a_Page_02.jp2
f3789957113021688e1f34d77b87b4d8
ef3c75c19002640e33961084dda91a39fee66330
2366 F20101113_AADYNA thomson_a_Page_44.txt
9392265b258f8a73bbae22369bfeb9f2
cdc0cf40cb4d31b0c160f9c5d0ceeb6ff306d7d7
F20101113_AADYIE thomson_a_Page_61.tif
9b9cf03cf816ba2214396ef24702402a
46bc917c0ac65f60e623bf593d88e32d89f177b3
50369 F20101113_AADYDH thomson_a_Page_03.jp2
43f316a9e7de95ff68de77126ec31b74
d836f886241fdf38ff9ba67024dbc9024b81690c
2369 F20101113_AADYNB thomson_a_Page_45.txt
5976b10aab02a4cd0faed1de8cd434dd
983be21451cc52e24713e15176105cd4c0101687
4314 F20101113_AADYSA thomson_a_Page_53thm.jpg
5abc3c7344bc43d84d63da5240b72d52
5ec4bc9390cab76d1aa58856a7be9bec4de39f77
F20101113_AADYIF thomson_a_Page_62.tif
376ba2819c14e1a54c205404293739b7
10da5fb82541d1760522e7358a8c420cc1b982f7
1051978 F20101113_AADYDI thomson_a_Page_04.jp2
9c7948a56b97cbab7bdfc6ae3ddb1d3a
d0a6faffef2fcfbb79aec591296967b0a5edea16
2387 F20101113_AADYNC thomson_a_Page_46.txt
9d383b204fe7c08583a37a477207f10a
c0cc168bf237ef8df8af04327f7e0fdcc17faa07
14881 F20101113_AADYSB thomson_a_Page_54.QC.jpg
014f19a09632cb3f6c30613d1ce1f2aa
c5685b2d429eabcbb96612d00983a46b9b46c13b
F20101113_AADYIG thomson_a_Page_63.tif
0edee1135cdeab343094971fb9743ae5
f0e29b939d085665f054e8873603c38e7debad84
652249 F20101113_AADYDJ thomson_a_Page_05.jp2
28a1981ee1a01d7f723648fe41b2d665
11057ed26198f71541a4ad923796fb4bc615e9db
1793 F20101113_AADYND thomson_a_Page_47.txt
7548c900c24a4e109bc9c39525101d2d
fe9e6d80b75554f1e39ca170a1702fdd5453d1b6
4661 F20101113_AADYSC thomson_a_Page_54thm.jpg
6cdde0c5c4a8e4ff7c0422f4a7d4062f
d17738c4220ba6cbd9947b1c095aad3a82cbb726
F20101113_AADYIH thomson_a_Page_64.tif
00bac3509ce3bfac5d82805f8a4d5d28
82dc99257141702daf03412dce9e0b9c6673259b
1051983 F20101113_AADYDK thomson_a_Page_06.jp2
a188d2455f8493048a38c2d285fe9f69
016e05c71149f449c39224bdf2b5d2df8ff831b3
2390 F20101113_AADYNE thomson_a_Page_48.txt
d88f739bfd9cfa61cc086b068a613aa2
795330c0203ce284a65e268d5769d8e4c6d5248a
F20101113_AADYII thomson_a_Page_65.tif
ee5fd9e7edff6d9b612b0e0b46ad2900
ccad5f52b766266d385e01601fb3182f9afeeb0a
1051985 F20101113_AADYDL thomson_a_Page_07.jp2
82b63c145b063606b490b317d669ca7d
d4f5609aa971e79895b6d51c9aad4bd2980936fc
203 F20101113_AADYNF thomson_a_Page_49.txt
8e7fe2e6224cd33f4103f31f774e9bfa
32ae3cb9199788df62f0a808404985535aad4133
13458 F20101113_AADYSD thomson_a_Page_55.QC.jpg
9b03e3c07839cb0c2d82f8a25f369542
2891cf4804bb890f3e6c3229a848a75ca13c6656
F20101113_AADYIJ thomson_a_Page_66.tif
27f441cc9599d58676836e0d3f3ee0ff
112ab2fd01163f0605dde0b02b17af1d615e9fa8
107401 F20101113_AADYDM thomson_a_Page_08.jp2
2088c15615a0203cfe427561747cbd3e
8bb5528a8451f5ab0b5a1aad05ecc861ff30c434
496 F20101113_AADYNG thomson_a_Page_50.txt
d73e8422bcb2a2519e69d88fd2d49647
04cb034b591ab916f934e961e8009214c42065df
4899 F20101113_AADYSE thomson_a_Page_55thm.jpg
7b136fd8b906b7ac7583bfe2b05780b8
7ab74ed3ae6fb802df7dc66a74cd6cbd06072a7f
F20101113_AADYIK thomson_a_Page_68.tif
b693f536da4b9cefe0b930adca1b84c3
a7258bf118e2de51f0944c1af60d5b1effbb9c6d
85150 F20101113_AADYDN thomson_a_Page_09.jp2
85f2debe097ffb915e0f73dc49a10899
26a61e0f1b35e90424a82d9155001483c1394d32
313 F20101113_AADYNH thomson_a_Page_51.txt
b685b9ce45c3fba7d46b47e6a21d290f
11d3d25b583a8d59ae86e9269ef76451bf44eb3f
14700 F20101113_AADYSF thomson_a_Page_56.QC.jpg
426b97fb8da05b5d23a446341fff7ffa
af99c03fbf602350b3f8abbe4d357c6481e082f0
F20101113_AADYIL thomson_a_Page_69.tif
c72bbeb14a362cefedec569364842720
7fe80a8f67c467a2160f82dc346fce39a48202e5
111656 F20101113_AADYDO thomson_a_Page_10.jp2
c130fdf364ffd2ce08d00273d9e69d9f
a7b4677fed3bf1dc5bc23da493bf45e72a4c3a93
409 F20101113_AADYNI thomson_a_Page_52.txt
17ce876a22bd6139b6ff22348ae20e32
6605ea0a339045ea2ac37fbfe70995fa2b51b43e
4559 F20101113_AADYSG thomson_a_Page_56thm.jpg
c3d26ab68cde331890c499a44c285f57
72297d9f6c79b00cd515ddd3766e9216318cec15
F20101113_AADYIM thomson_a_Page_70.tif
1e71d9f2e82d6c10b80ccc0ccc4df9eb
14afab9fedf2dd8a9d6c2653d8a64bf4c65ba966
114492 F20101113_AADYDP thomson_a_Page_11.jp2
c2e7dfd1c4dfbad951822bf18bd6efab
c103e35373e2a915556be193670bcd4172f90c3a
498 F20101113_AADYNJ thomson_a_Page_54.txt
d64cf4f51448e9ddee47b7f21c4ab0ab
fece4eb463c7b51552194ec44b51360d8c968b60
12587 F20101113_AADYSH thomson_a_Page_57.QC.jpg
3c1d69d95199340090c0cd087a6dda3d
2aa83d48bc9d6ece54683077795e45680d15e3e8
F20101113_AADYIN thomson_a_Page_71.tif
dd2a46de8f903f8c9ac20fb733808dd5
5a5e4a0ca6be854892820a1e80a8ef8f2dd7a50c
122943 F20101113_AADYDQ thomson_a_Page_12.jp2
f4717990a39ebfb20f390464ead0e6d9
b2f1c0d497760ddb31dbfdcf376fc85bfc1f8c9d
164 F20101113_AADYNK thomson_a_Page_55.txt
030b35df64dcddf1ad6dfad6858d9023
2ea855c6bb5fefd055bd573e54ecae2942f0a49a
4543 F20101113_AADYSI thomson_a_Page_57thm.jpg
13eb45540c53d3b22f0a6f2020240034
981089cad9c837c710e14be5b7048e6d3eacea11
F20101113_AADYIO thomson_a_Page_72.tif
e9625766c731e35e82b992a3c2fa455e
e47413a66dfb930b874b5da06e8c79c57820a4ad
18410 F20101113_AADYDR thomson_a_Page_15.jp2
e4462dfc736be7c51ee1ecbc8c3167ee
af53f4e5dffec09b7bf38c901efbe505b58b6ba1
380 F20101113_AADYNL thomson_a_Page_56.txt
42b1d9798789dfbb59b8c0bf801036be
94a1e31c7146304517aa2ae70856bb40bdc3d634
14750 F20101113_AADYSJ thomson_a_Page_58.QC.jpg
395143ac758107df42c7ffaf13497a97
c2205c26996913db0cd35624a7c0fb951553a468
F20101113_AADYIP thomson_a_Page_73.tif
9816944d802e06314d178540bf6259dc
65bffd52ad1cb8a217ae4cc6e5c8067311e3e858
823282 F20101113_AADYDS thomson_a_Page_16.jp2
755f43ada1a215600e73f81e39d05205
9fad1bd43040ab8506708b9998dfa889715dfc9d
288 F20101113_AADYNM thomson_a_Page_57.txt
e30edf54f9b4282f7da52f6c7b770545
e893401886359d83a17f07ce9c04d188c8f05689
4549 F20101113_AADYSK thomson_a_Page_58thm.jpg
26989527ac89f7af117454c2a0d5b990
0fba0da6037574ad443cc957a3f7c892595b32a2
920355 F20101113_AADYDT thomson_a_Page_17.jp2
0b7c98a95b8c435c6d9add731737e697
254ab7b3344461a00f1201d7a27d12772fc00c6f
404 F20101113_AADYNN thomson_a_Page_58.txt
77281e0b07f7d1255f4f97f285cfdded
879093119dc64af201936922e816c0ff12e3e4bb
8471 F20101113_AADYIQ thomson_a_Page_01.pro
e1795cc44f6cbcf365e1bfaf898ed17f
175874d4ead78ee0d2c346bb17ef6bce154c64ac
25033 F20101113_AADYSL thomson_a_Page_59.QC.jpg
4f2f09f62bf38dbea01f5e4ccd5d2234
0d1e81da0fa7dcd72328689360ef8659d0e175b2
2172 F20101113_AADYNO thomson_a_Page_59.txt
36f0f5e5f37ca3695fbd174dbe647ca8
e22410eaa69df091ca80a5fd2584f56f4a320582
1043 F20101113_AADYIR thomson_a_Page_02.pro
4f44b0ef7fc4d0b0e691cb66228a26d1
20a307d48d3e2961f5ebe20a478547046e7e67e3
6819 F20101113_AADYSM thomson_a_Page_59thm.jpg
ef2adb55e07156bea48c6b8ff8883264
f3740614f5cfd37785e0efd9a7097259c14255be
106028 F20101113_AADYDU thomson_a_Page_18.jp2
b4d777948c73a3cbf5c4b83197592977
fb45480cb88e1e0630cddc528e61cdf2b1708f8a
2197 F20101113_AADYNP thomson_a_Page_60.txt
933214741c6a09d4ac96502c13b7eca9
e8b059e913938b08ddecf4076dc2d7b6c249875d
20911 F20101113_AADYIS thomson_a_Page_03.pro
d9e15734624c254fdd1893c823c2e504
1328a97ab692c64926f02d2bcbba2d8bc3b94823
25830 F20101113_AADYSN thomson_a_Page_60.QC.jpg
2ba51b892afd85eefe5fff059153e858
39c3a31765a9f672a4e1203988e909374b2ee815
107614 F20101113_AADYDV thomson_a_Page_19.jp2
09c44229678ee31e2659bc9df00f3308
e45d83287190da29dd4d4d1edd68a57931a607e1
1979 F20101113_AADYNQ thomson_a_Page_62.txt
4bbb2d09eec646c0de20a88a8cc20efd
c5c6ca99ad533b1f8f8ec798594785aab82f22ad
101071 F20101113_AADYIT thomson_a_Page_04.pro
1c1ad569f74e79994f442b49bbc582a8
2d08346f759363b63df26d4f99cae0590d0f1e40
6854 F20101113_AADYSO thomson_a_Page_60thm.jpg
38e05903bd8ce33a6dc5786555ac0c45
2ccd6df812b9f1b545fe80cccbdffab24974bd4c
585397 F20101113_AADYDW thomson_a_Page_20.jp2
db22f9b21062bc6844a79e821eacdb22
de664aff23c091711800ac27cd4d354830e66ab9
1721 F20101113_AADYNR thomson_a_Page_63.txt
c58d0c0941da77dffcce1b8116615a26
668f05e2f0c4d573f09ecb8009796877899a50a3
34598 F20101113_AADYIU thomson_a_Page_05.pro
8491387b68e133810325c02dab05fb7f
cd373d1ec09875d27f0c4c770dc2326e8fee7211
23080 F20101113_AADYSP thomson_a_Page_61.QC.jpg
7787caee920022873b63ff0b6e2d88d8
ce81a263696bd704153dd325e61fa01a705c1720
106265 F20101113_AADYDX thomson_a_Page_21.jp2
16d2bdd7308f1dbcac600d4f614b27e0
c7505bad0403e56686f2b9f6e8d3f514226cc33a
2220 F20101113_AADYNS thomson_a_Page_64.txt
83bf89ad74d4e3e8d907f28cc2268e60
7b1fb23cfe87c3120b34e3eddc32c75b746f02c0
59773 F20101113_AADYIV thomson_a_Page_06.pro
69b6424363602e46b690746775063cdf
bac25b7e8df5ad8e8edb43b73dfbaa0a30809d07
24131 F20101113_AADYSQ thomson_a_Page_62.QC.jpg
536f9c58ea7600cecc50c34acff11060
694e839b5344e995854d9fe520c1adb5d3608bca
55363 F20101113_AADYBA thomson_a_Page_17.jpg
7ddb4ff7c6a9f56a80d9668696ba60ca
6d0757bb8be790e3dfcf740cb67f7244c13cf7e9
112672 F20101113_AADYDY thomson_a_Page_22.jp2
f3c3fdfa832cbc7d76ab18ac9a811d9d
3b41ef13ab9b886b897cb4d04f2f1d350f43332c
2135 F20101113_AADYNT thomson_a_Page_65.txt
82669a411aabd43342c0a18b602c2d86
5dc64f1287fb65a8d9b83e00a7f15e911a139f81
64782 F20101113_AADYIW thomson_a_Page_07.pro
07b76c5642ef9b201fcff12126479823
110a3cac2725552f4ee5de61fe7ba2b453302b92
6540 F20101113_AADYSR thomson_a_Page_62thm.jpg
95d86a5324a2536385b0085b1c431fb7
29b1e0b46ef22ecaf7cca1a6b3d8625c5ff3bb37
72437 F20101113_AADYBB thomson_a_Page_18.jpg
17b533505e17022a780aafd77db68809
a1ee7f834cf52bcc8d1dc09706783463d0f145b4
72518 F20101113_AADYDZ thomson_a_Page_23.jp2
3b18cbd40f4b48053d942a32b01da9f6
ca76dc3c33610cf00b2d585abd7d5df9044ac793
358 F20101113_AADYNU thomson_a_Page_66.txt
5ad0adca123cee32aff94fee93189967
47efa2882c3cbd43c3ddb44a75b1dc7dac96fab9
48355 F20101113_AADYIX thomson_a_Page_08.pro
71360c7d16e9ee86636515fa2762c28f
878fa9f8f8434c22917567a9da5d0713710333ff
20737 F20101113_AADYSS thomson_a_Page_63.QC.jpg
28cc4b9b2066514db8419f6d41348369
801f0755d606ccc280658de51b3fc47e5bce84ab
70855 F20101113_AADYBC thomson_a_Page_19.jpg
22b43b52134bf53af3be7d5dbced2e7a
820967c6e7aaa5cd69de40e1330316c19c08be93
2429 F20101113_AADYNV thomson_a_Page_67.txt
d98482b73b4021c45a30ed6c8382b97e
9b33fe00c9814d08170afd9725978989cc8434ad
50972 F20101113_AADYIY thomson_a_Page_10.pro
66f4e85a79e5a10d51e854bbcf5dd0b3
27aa8e3ad36134462c4b9fd00d38e4ba241c6db5
5861 F20101113_AADYST thomson_a_Page_63thm.jpg
0d0f1d88d12a2e85bc9d963825d40102
b0f58f68540d7b26fcd46c57b70d26424186a37a
2424 F20101113_AADYNW thomson_a_Page_68.txt
c24029270254efab67e45521288b441b
8d569b5fdad9e1406f7db1b5bac2d1333b36da50
F20101113_AADYGA thomson_a_Page_04.tif
0002ca1a083dfb5fe93c888797a17e93
f05c149c83521a7e4cee9cd72a38a8e67c7d3879
44425 F20101113_AADYBD thomson_a_Page_20.jpg
7356568c1223dc870b33907706232b2c
52eab013d4de996994fc473ff24cf56347fd9cc6
17569 F20101113_AADYSU thomson_a_Page_64.QC.jpg
5d4e2baacc044c33e25c8ca866f84c82
8cc84d7f62f9ce39954ec5b9f791df1b8e619f5a
2533 F20101113_AADYNX thomson_a_Page_69.txt
e643f072da4cd33df6525e0999d76971
c3e5daa7d2cfeb1c9b10608f04edc4669d88e3f9
F20101113_AADYGB thomson_a_Page_05.tif
a4a426e1ddddc05ccbcf5bed9bd2ebbd
8b41285faa211381e4e37601a48508daca2d189a
52245 F20101113_AADYIZ thomson_a_Page_11.pro
15aa8b54922bb80079f85ea7356e75d7
952a86023972ca243923f0be8bd4976bcacee5c8
73566 F20101113_AADYBE thomson_a_Page_21.jpg
79010361f9bb3287e1236e895aec0e56
680c7a34fab1cebd0551590b51dc24d40ec664d2
4875 F20101113_AADYSV thomson_a_Page_64thm.jpg
1f4ba33772d262a2a0951450621ee9f9
5d7b9bee82256e541f909da2800d642fed61e14a
2348 F20101113_AADYNY thomson_a_Page_70.txt
6c7df0d5fe6b779cec6541a6b56b7954
ce4181de2ce51c58d3e7ba82559f22f328ab6b69
F20101113_AADYGC thomson_a_Page_07.tif
c8129cddbe3e7f348b98d566380c1bd6
ba98234981f6b54bb1c6c8811768985d79458847
76529 F20101113_AADYBF thomson_a_Page_22.jpg
e9d94c8589f47d1c8f6ff491a82ce8c2
3a154e04cce2151cb24cad86964f24f0392eba85
17031 F20101113_AADYSW thomson_a_Page_65.QC.jpg
416b2766799698e3ff05d26bfe19648e
e94299025e662de447245ee974a4e0f5a119f56c
44233 F20101113_AADYLA thomson_a_Page_65.pro
be97e18be631267a8a05fa8776294e96
94a71e0077fcd459f426988a19bad47c6c65b623
2361 F20101113_AADYNZ thomson_a_Page_71.txt
2ba62cfb0c0e413e509699ebae80732f
6e48b27c31eedfbb5d2e15ddc287095e387114af
F20101113_AADYGD thomson_a_Page_08.tif
855c0f4bf7411ef2c79283aeb1a32fb9
b70e940ccc600122cfb5171702cd605e0e577c3f
51800 F20101113_AADYBG thomson_a_Page_23.jpg
832bf390c6a763fa04dc6d1d59e6d5fc
c7d5e11a45c4c05afce19b89224a19ccb6f88ac8
4836 F20101113_AADYSX thomson_a_Page_65thm.jpg
5c567cc06a53ac08ae320934ca6c84a1
920a761c9020af50de4aa98f4cd7ea5a5024f2d2
5209 F20101113_AADYLB thomson_a_Page_66.pro
cd58fa26e985a7171398495500eaee51
4432c169f9d72e5f5f6c186091a2fa81429d5d26
F20101113_AADYGE thomson_a_Page_09.tif
1ccf20937399d89789d385da014e9daf
82969e88b465502e0671d6fabb9685ec1758c353
53075 F20101113_AADYBH thomson_a_Page_24.jpg
6b6c3eb54a8635a16903e2281ed7c408
bb876a5a2df38a58e2ac236d3f0c3b1cc6166c9c
14271 F20101113_AADYSY thomson_a_Page_66.QC.jpg
724207adfae838db7a59b6d4ea6f0181
40e0139e06722ea65387bb86d39d315af1a758db
25140 F20101113_AADYQA thomson_a_Page_27.QC.jpg
d6a087e4c4e01d9a4742e0bf563b78d5
8536cb8d0bbdb104498d18c008bbf44ffd45fb87
59811 F20101113_AADYLC thomson_a_Page_67.pro
2db4aba952cc2c899d5812ccb394ac87
bad0321c1a345c2948d5a0d6bbe05684b1d67a3e
F20101113_AADYGF thomson_a_Page_10.tif
9a9d01e532a5d530230d1b7f5afa178c
b0ffe77ca7a95459cb1da342b078b58fffa293bf
83630 F20101113_AADYBI thomson_a_Page_25.jpg
ac2d59c0b40ee95a4d52a6e3279c97b5
a62909ca1e6c27cd2da8ecb5f9900e3a4ceb4275
5237 F20101113_AADYSZ thomson_a_Page_66thm.jpg
e4afc1a285d2855c608a8c9c6388a5b6
bd7751e2d25da23da309a52ecb0db00e2bf6a830
59456 F20101113_AADYLD thomson_a_Page_68.pro
413fa4664787a9bf59cb96fe91847b1f
c8711e8fba2db1754316cdf96db8f11814331cab
F20101113_AADYGG thomson_a_Page_11.tif
4d17a3b9d945bf17aeb2a438ef33269a
8976449aa026882c585cf340a500dcdc9212f4e4
74371 F20101113_AADYBJ thomson_a_Page_26.jpg
d5994e3b357ed1a48eb5f69826a96963
858ad7294edfd65d182c50d59f0882e6f58eff14
7160 F20101113_AADYQB thomson_a_Page_27thm.jpg
adb100798a0196266d1082fb88726fd2
ca5cc16b028bfeed6d5654df98da4e2a98710e1c
61936 F20101113_AADYLE thomson_a_Page_69.pro
af7dd65fb4ddd49b7105e0284759c14a
cc789d003fab19ffc5aa10b6ee24f2afaf7b461a
F20101113_AADYGH thomson_a_Page_12.tif
b5ad50cb03da844148622bbb0743989f
c85fb10e953ecfcc29c1765ee2a3f9bfd794b9e0
76757 F20101113_AADYBK thomson_a_Page_27.jpg
618c48a75ee0b1411d53f4143f82186c
dff965f7a95f5db68b015865dd86f83bf38ae8e6
23196 F20101113_AADYQC thomson_a_Page_28.QC.jpg
808a575233f03f7e9fbaf27751fc87c7
fd55b648fd59166ad907682d2ce12793ef105734
57244 F20101113_AADYLF thomson_a_Page_70.pro
d37acc4683c41b0c685bd6e357239fad
eb05f75da2f4645e54a84c1f0934bc1c99779d24
F20101113_AADYGI thomson_a_Page_13.tif
ff117a2c0178f35f55cee57d8694dac0
6a1bd04725663498214d8b6f5fb765ea7490629f
74145 F20101113_AADYBL thomson_a_Page_28.jpg
880e32ce21bb2aff002a13b6172a6b0e
e8032bade20d8b4fe64bcfe37c0bf2d67924d688
6438 F20101113_AADYQD thomson_a_Page_28thm.jpg
319226e23d3d6942e85f836ec4221e19
e8bc869226bf8533bd908975560dfb74e71aeee1
57106 F20101113_AADYLG thomson_a_Page_71.pro
777bc0493ea0a63b297030ccfe10bd83
eec49dfa3a62da215f25a684db732754fd8a14f0
F20101113_AADYGJ thomson_a_Page_14.tif
d372d4e91a5b2a207bd1d3b11925265c
e1fcc4ddead5a76b4c42390c7d8d6cea2f592e6f
75357 F20101113_AADYBM thomson_a_Page_29.jpg
24b03fda9733b579a129224ea3b835ce
851824f5d1a7b1f5abafbba25ea477ffdd02d2ab
25072 F20101113_AADYQE thomson_a_Page_29.QC.jpg
e705b6f4ac7576a6eca69e2610ce6796
94626dfa351593158dd02ef1c8a454654f1105ee
11089 F20101113_AADYLH thomson_a_Page_72.pro
484b370ffbdea53e634a33f361206087
5bba99bd765e3f5fabaf8f4d08a565d02a8c4c05
F20101113_AADYGK thomson_a_Page_15.tif
296154e81d173d777e1c4c038800f092
be9428f0a0fbfd728147bdd73f35564d816fd6b4
73800 F20101113_AADYBN thomson_a_Page_30.jpg
9a6447ae5880470ae9cddac9e2646920
939f7f410bb335ee9670857b8456690958aae778
6741 F20101113_AADYQF thomson_a_Page_29thm.jpg
af8620e74aa930cb27182d99400744f4
b5efc1a3824c96d32c12e5f46789788fbcf74c6b
24488 F20101113_AADYLI thomson_a_Page_73.pro
34a27c963a2e3dae51f0360a51bb1470
a49cc6222b7acfba17d1890fb07ca7e589b1eb1e
F20101113_AADYGL thomson_a_Page_16.tif
fa8970357f801eb8f344ab56ecdf9c9e
01a46a35c13ae1fa683bf2e3008e1e1aa8ded911
70330 F20101113_AADYBO thomson_a_Page_31.jpg
f95e77c52d9615541c5e9733e9b5e269
d22664fccbe6426ab7016d3daa2236ed8a23918c
24751 F20101113_AADYQG thomson_a_Page_30.QC.jpg
819d4c57d20ecce7e3426f36eb5b7d1d
9169730c5f101ba635726454941951483f9a7515
508 F20101113_AADYLJ thomson_a_Page_01.txt
14311af02d0588c24a9f428f54701f34
9f17788c6367fc218b33d5fefe303aa74c4e3fe9
F20101113_AADYGM thomson_a_Page_17.tif
d73cce595429a5ab2e6b6260dc41327b
ccbbfcaf149b204c664ecc0c192e32a7f2287ef5
74729 F20101113_AADYBP thomson_a_Page_32.jpg
e6bb713fe0c76f48939d9605a080b07d
afcb392af15e565b8f8b9fffdee5c5f8605bcab1
6825 F20101113_AADYQH thomson_a_Page_30thm.jpg
cc130b9878f7af1dbb792f5b857f9ff2
30fddb5d3bd0e88fa8667476dc2457bf899312a4
96 F20101113_AADYLK thomson_a_Page_02.txt
fffc440e0250f28e5d55cbf2a56379ea
5153f8a5780c443b6caa1b2552d78a0eb7377173
F20101113_AADYGN thomson_a_Page_18.tif
f9af31015d73d50a7f900603f0e5f917
4c3ba620c91bd5b9f6324adbe2922c56b25f994d
73215 F20101113_AADYBQ thomson_a_Page_33.jpg
cdf61f1ef021281a36a563901da19d7c
5250ad9e75929d31021ba2ee9538dee439c2dcb1
6551 F20101113_AADYQI thomson_a_Page_31thm.jpg
21261ec3e0804eeafd5eac50babdad49
e25bf6bffeff1536187eb54e0597d061d6283824
868 F20101113_AADYLL thomson_a_Page_03.txt
1ec6bbddd06cb39e90d3bf09deac8155
2a582c3e4bc9835c482e0a8a38a1d14eaf2874b2
F20101113_AADYGO thomson_a_Page_19.tif
d56f05d5759a6a0114d86f4a444453d2
c8a38802ec94c279e76ca73b8bfcd500c8591511
73246 F20101113_AADYBR thomson_a_Page_34.jpg
c70e6c709dc5c63ae9ea8ad5d2d3bd94
bee059026dd4320953bcc4e68400b663e65d44d7
24339 F20101113_AADYQJ thomson_a_Page_32.QC.jpg
7f55ba6a8819d40709ea6f9ec15d9db4
f1c1a9193895fd3263efc9f759873c0a436dfa98
F20101113_AADYGP thomson_a_Page_20.tif
4a4a2078899a916408f48956b8ed8207
25565e60bcae72cd4f79f83a5391ed37a5066da1
4154 F20101113_AADYLM thomson_a_Page_04.txt
0e89db03495ed8491b0b75fc23409298
a661d0cde93485dd13692b0c93aa97d8d6a113cf
6683 F20101113_AADYQK thomson_a_Page_32thm.jpg
a619c3b70691b30ef5f2689793641b6d
d4e46e725e994d02024098da9e945fefd2b3777e
F20101113_AADYGQ thomson_a_Page_21.tif
589dcf24e15a9dc694b83d36a0970b41
6a28f9d5ca9e87f602294cd9d5eea839ce279fa3
79604 F20101113_AADYBS thomson_a_Page_35.jpg
44e0cdad23aa7346558f2f274ed4d886
e26af140b5b571553b5dc2c439bafffbb8000975
1390 F20101113_AADYLN thomson_a_Page_05.txt
a3eaa91cba081e9ce02fa6b7c5c205eb
c2a3961f699720ecc3fef283f246d46d0297b9a0
23681 F20101113_AADYQL thomson_a_Page_33.QC.jpg
f9df8b4850f535bcd48f0474b5a67b7e
f3f491784787be425fef42fc16f3fa02a256520a
F20101113_AADYGR thomson_a_Page_22.tif
4bae75bf19b155ce805c27d92494774d
ef5178246cee77b6a2ef0a3d48233bf879d870c8
2823 F20101113_AADXZT thomson_a_Page_49.pro
524b84cd24d24c30aec29f717e040bd8
b21ebed3cac30e956c518cfdff052ca33cf7347c
74520 F20101113_AADYBT thomson_a_Page_36.jpg
4aa64eb67ac7f9a2d4c523a8a994a527
7a29268753ead0f5ca278fc664b2da585d7f7bb9
2450 F20101113_AADYLO thomson_a_Page_06.txt
9ffd8eb8d858a4a0ff9d86827dd73464
719a4e52c8b60000485964f7b54ac7ab18faae78
F20101113_AADYQM thomson_a_Page_33thm.jpg
8533e29431915117270be26cc758507c
f3abf6d4db270b99ad039875e67d329a3850e824
F20101113_AADYGS thomson_a_Page_23.tif
91ce5404336c97a1e53c4dec5221de54
3182d985ff50323dc7679d28118f24df8e230f10
73999 F20101113_AADXZU thomson_a_Page_10.jpg
ea76af80a42c5927d0c8030c18fde35e
1a97fa39be726696e046437c887fabdccc451296
72459 F20101113_AADYBU thomson_a_Page_37.jpg
fecd3db57b88f721553e50402b4ed2f9
1c4c4197c01fb8949d829831c42e3c0ccdb93771
2658 F20101113_AADYLP thomson_a_Page_07.txt
b64c675dbe60c8ff95a59a3191d35eed
aa3152de48dc998571ce5be564badb11b22bed20
24293 F20101113_AADYQN thomson_a_Page_34.QC.jpg
486bb8aa5b80b080e3052194b22d522f
cb3377f9a89fe32ff2f357039040b65517eda56a
F20101113_AADYGT thomson_a_Page_24.tif
c6c8f6775c82f373b0e0f9a674136001
d07684903903502cca2b95cc2f62b4096caa3459
312 F20101113_AADXZV thomson_a_Page_53.txt
4e1967090e1e23ab144ffebb0441d0d7
eb9f0b42ad37f252314d5723240a90b63e5cb772
70583 F20101113_AADYBV thomson_a_Page_38.jpg
4e7d1afdbed23fd359ee0edaa751aa15
cd9356a60e79f8a44222a5cf0d5a85dcb9be8664
2101 F20101113_AADYLQ thomson_a_Page_08.txt
038840429dde93b81febee8a74c558d0
578ca52fd4d1876469e637ef3804244bbc0b6292
6573 F20101113_AADYQO thomson_a_Page_34thm.jpg
5fd4ca5be90c6701c268bc4b9af9a222
46b6fc8f617a4285bf36c4642060fa9b7b8f03de
F20101113_AADYGU thomson_a_Page_25.tif
ad6c5dbb1d46c53cd00c75a1e1abcd00
2c45fe80032a99822486b40357a36da30e1ed501
1998 F20101113_AADXZW thomson_a_Page_61.txt
3e1ec85b649397ec186b184be9eff9bc
61d1753347202add03ccadb3142ac5a8a8e20c46
84905 F20101113_AADYBW thomson_a_Page_39.jpg
eea0f991240fa1a8baeff3d7201432f0
806063b8f39fbd5ea84fe264338a1da81749615f
1503 F20101113_AADYLR thomson_a_Page_09.txt
c5f7de12c88bdce0ca216d4a445a08c5
fea2a4395ddf46be8649662373ba0eb78ebc42b9
23410 F20101113_AADYQP thomson_a_Page_35.QC.jpg
15d908f9abb938e9d3469c929b41950c
b96bb59ddd3e9a76792c9d4834b08d9142c0a1e6
F20101113_AADYGV thomson_a_Page_26.tif
d6cf8190ac4a1c7fe6c74cf308d00217
dbe9db51d06fea7faeea835cef3e879760cc4da1
F20101113_AADXZX thomson_a_Page_67.tif
216281a6c88245bb2c6b0b6c197db583
3ecd45c67a437d13ad8ce624eb54c1473974c7bb
75736 F20101113_AADYBX thomson_a_Page_40.jpg
3e905cd19cea65748996f8e7f4bf19e1
75731b24b652a611459af9f98c2b23b68b492ae9
2123 F20101113_AADYLS thomson_a_Page_10.txt
55d76e0804576cd7cc3d7f780b2d6a4e
25a31ec1388680a91f3061969a9218ad66d1e370
6452 F20101113_AADYQQ thomson_a_Page_35thm.jpg
4bdaa94a758a6084f6f4a733e8d5be81
a361650178ced885ac81fc37c94c0d7156f69282
F20101113_AADYGW thomson_a_Page_27.tif
51ccb47d8f63cbc4921c91a252269e81
bc09c84e8531e73dbc88d31c749b0b95feb264df
7374 F20101113_AADXZY thomson_a_Page_01.QC.jpg
7d930f7488b2708b4c3934183554ab3f
17327400bc34b0bab7476112cd87288920a81286
73223 F20101113_AADYBY thomson_a_Page_41.jpg
926433fbda8a7eeb07b3ca0e0331240b
4bbe3c467479ce5c98313da627587d8c5cc3e4ff
2065 F20101113_AADYLT thomson_a_Page_11.txt
8260024f7383b55fcf67c9000c89b00d
5f32eef221c4c5c2ea488c855dc12c9dd765cdc6
24663 F20101113_AADYQR thomson_a_Page_36.QC.jpg
b5508a71ab0035f365db6bd137f2b4db
11d05fc9fd29ec9c14b3ad26852954f667266b0b
22685 F20101113_AADXZZ thomson_a_Page_31.QC.jpg
9036a810ba32bfdf40756af9f46cc087
15dc80b6ce7dbdad2b0cc47ce7b4bc90bb5ba7e0
29893 F20101113_AADYBZ thomson_a_Page_42.jpg
3216437f364c8a4275487efe10ec566f
5bda15a7f0d29c38cbc69f49f6eed87617b990c7
2191 F20101113_AADYLU thomson_a_Page_12.txt
1d5119cef2570af536886328fd281715
37699fa1764808c1478b62152034db7dd5393d82
6579 F20101113_AADYQS thomson_a_Page_36thm.jpg
dd86d77824185e66b6f280940ce2447b
19ee723d19782218087e94f8b4ccee4b8c6f6908
F20101113_AADYGX thomson_a_Page_28.tif
32f29d53b15b68807a4e196b67d324c5
48b71a5a795e00447905413b904fe67b034e5c1c
2112 F20101113_AADYLV thomson_a_Page_13.txt
572ec4da84b971e0a5c77fffe1ae34b2
b42c465f7b15d9575f594e7824886760908492de
23719 F20101113_AADYQT thomson_a_Page_37.QC.jpg
7e3ac32495682c66b62e0a5d355788f7
1005e11586a11c07dedcf13cf6adc3cfc6374756
F20101113_AADYGY thomson_a_Page_29.tif
8166d0597320aa674a3dad27b336f090
4092137ada6652620b4c87995c142b19ee0c108c
2028 F20101113_AADYLW thomson_a_Page_14.txt
de67de498a35820498022ce367b7030e
c7aeb0252e5a012b7c2ce32399c7c8819a680067
78924 F20101113_AADYEA thomson_a_Page_24.jp2
2a355ea3a694fbf536b0003d4faed3b2
fdf3b4497fd6b4c816f5b1dc6c09c6349a65c22e
6552 F20101113_AADYQU thomson_a_Page_37thm.jpg
f2293269cac0e2d9822486e68af2be96
54752b87a9c7c69939b63fa6df05f316a7943d14
F20101113_AADYGZ thomson_a_Page_30.tif
3cf98d367052adb8cdca0a0cab065a2e
fe8ee7c251b2d1fd96279a157baf6c168acc49a5
283 F20101113_AADYLX thomson_a_Page_15.txt
594ba95891377e91f0ee40f3231fe02c
8d147ec4d796d272fad4ee11374d918ba624acfe
128118 F20101113_AADYEB thomson_a_Page_25.jp2
70ccfc12303ab1bad74e399683305054
c0a08df3fce4cc0752da507c32bef98b4a571d8b
23021 F20101113_AADYQV thomson_a_Page_38.QC.jpg
a06661761bc450d5b0a33510c8bf6891
b0edf30f3f64f1418a7f709602b91a03152d4fbd
760 F20101113_AADYLY thomson_a_Page_16.txt
8077a13029df5024fc85fa5b1bbe860a
92e293245a913dc940a87fac42832e58bfc07741
112409 F20101113_AADYEC thomson_a_Page_26.jp2
24d544b75897b60ed63b2e9497939425
ba33fee58f527cb12299ba2f98b03a7b88e9c34e
6603 F20101113_AADYQW thomson_a_Page_38thm.jpg
523dad9f2c5b544341ad06b7920e1307
994e9ea6a43198b6afb1fe2bebd906d7c339eaa0
54719 F20101113_AADYJA thomson_a_Page_12.pro
91c69ba34729e4652beebd1abcb4e110
6153e87fa3114ea912009e6339c28b9bd732adb3
1080 F20101113_AADYLZ thomson_a_Page_17.txt
ed072e4e3cf297da83861d84dc6a0e0c
b41639a576a1a1560c3138f15a74cab946cc2150
115072 F20101113_AADYED thomson_a_Page_27.jp2
faaedb41b0f1a29021ff049217c5b08c
9970004960213135bee99d31060e6f1bb19f7931
24737 F20101113_AADYQX thomson_a_Page_39.QC.jpg
9cca0012462422acdeca49e1bc4510d9
4d14bb87d48d699fe8e35bb59ca8feaa7e0a7df1
53330 F20101113_AADYJB thomson_a_Page_13.pro
f7fe2b0f69cdd9ccfa01582e9b0b61eb
c7496059caaa50be949791010dd094a6136a9bda
114013 F20101113_AADYEE thomson_a_Page_28.jp2
69f624c45f36d6c1e0f6dfd3487b81bd
13b408c1f76881ecab664c49d82fdb5f2d99d20b
6696 F20101113_AADYQY thomson_a_Page_39thm.jpg
837f9151325a4f4ec79c2ad540d312c8
97de4cd01cb731c2444dbefb325701b7b32f5751
51499 F20101113_AADYJC thomson_a_Page_14.pro
d7d653723f04c57fc0986820f710541f
7367a583bfa554390b74f0dab07fe6c26393aed6
115906 F20101113_AADYEF thomson_a_Page_29.jp2
aea520e3de07bf83c42891c87006519f
743b7736e23020540911b589ffbbf812383f9e0b
25292 F20101113_AADYQZ thomson_a_Page_40.QC.jpg
c4bde8e2260d72475d11245a5612fd6e
e4bb676990c2356c6e6601e1ba78afc709545cec
462 F20101113_AADYOA thomson_a_Page_72.txt
97aca2f03772921bbacd062fe1071fc5
5a740fbf66386fe2460abd46c405c262b2b43956
6895 F20101113_AADYJD thomson_a_Page_15.pro
1d5c8b6133bff30f7f687b5d292b2327
900200f77a15547cfa24b2324d784026b6179717
111855 F20101113_AADYEG thomson_a_Page_30.jp2
99d72ceb37cb13be674f5c741a21c839
f1d886dd548be5fa9a7c4270ca423a7bbbac45fb
1009 F20101113_AADYOB thomson_a_Page_73.txt
9c2e69f12bfe104c54d64d3ed77ae1f9
d48c7557da8b6f4923bbca55ae7910cd403e4492
10763 F20101113_AADYJE thomson_a_Page_16.pro
046a6ec1b0ce21b4351d22e500c7ffd1
abe35a6209f7f621f4a375d264ba3dfe52058658
105637 F20101113_AADYEH thomson_a_Page_31.jp2
31c00d8bfc569682dbe2464dacb8503f
8b32e9378971028f4d71c73a44b41c5c46ff12a5
24724 F20101113_AADYTA thomson_a_Page_67.QC.jpg
e5839e9ffdd6430939feefdfc290b340
32cf5890293ed5142ce168b00c66f99156b11941
2474091 F20101113_AADYOC thomson_a.pdf
382ee530e3e870679eeee932db20b268
17123e5dd4bbe834385ab0c9a66b2843a7ce31a0
13168 F20101113_AADYJF thomson_a_Page_17.pro
cba25ae9a646c726c0fe7cf74d32a4c7
ee10c9064a3933a35d504f63c391c4f60d88092f
113695 F20101113_AADYEI thomson_a_Page_32.jp2
2300267bb623934113c6d6f654df9364
de71f1e0d9ca480c3b6efc20e013db6227826a22
7033 F20101113_AADYTB thomson_a_Page_67thm.jpg
70be0c0f9b03b519023c31a2d0bf106c
c467741d1c8ec6ee059ab533284990efa75e4fb1
2342 F20101113_AADYOD thomson_a_Page_01thm.jpg
2ad468612c977ce337de2610542cdc8a
3f35a59561166f483245ae36f14c97ab2815d88b
51772 F20101113_AADYJG thomson_a_Page_18.pro
df2e6b92ebb141967a58183ef45edfe8
ac7123843f33fd0bd948bdf77f75b29efa14e618
110101 F20101113_AADYEJ thomson_a_Page_33.jp2
bf625e8092cbe8c5a207f47068bc3674
06fa7a596156f67231ca276e4c680986a7c69e00
24813 F20101113_AADYTC thomson_a_Page_68.QC.jpg
bd7eab1f386891fca480d3495c8b7b61
67d8a7346149019c0a5f738cc51b3546b8a9799f
3147 F20101113_AADYOE thomson_a_Page_02.QC.jpg
5d5ffd4388a6ec98aba2c1f4c0f390d3
7cd8509048776d0c1ac8019044367e7d7c10ee61
48729 F20101113_AADYJH thomson_a_Page_19.pro
b3873b5e728af542c85a02da4422a73e
c70f35b818fc168a753f4dd7ed865e6d1adc0f7b
112078 F20101113_AADYEK thomson_a_Page_34.jp2
a0d1c5fe845dbc6d142f1e30b99019e5
02057c031b27b0f6d15e6f17ac6abb93df4e4064
6823 F20101113_AADYTD thomson_a_Page_68thm.jpg
450504ab0f58423aa6f8f160506a7144
25f48f59165742a14eb0f84a7dd6a41099caf7cc
1350 F20101113_AADYOF thomson_a_Page_02thm.jpg
c0b0d96efb9fcbda750b355b675c0527
78c69926da8bdd4e929988d672f6d16a0eb1ae4d
24670 F20101113_AADYJI thomson_a_Page_20.pro
7adccc90ddc7fae251f04701a47f9268
4781d0e3745955134aa12f914612fab1df3e5c13
122010 F20101113_AADYEL thomson_a_Page_35.jp2
94f28ff0bcd6b346f583fbf56280f062
101d815e67d7d3a55e610fb3c306f9a7e475f58d
12372 F20101113_AADYOG thomson_a_Page_03.QC.jpg
a86437b4e82f0a988b3c6cb7c94421c3
089a6220c9cc3869ea029ef21ce94f4074ece389
47498 F20101113_AADYJJ thomson_a_Page_21.pro
1ea96aab10e85cf5df0981fb82b264e9
2a34fc64d8a3da38e30da8d83a8488a2a0b51ae3
112328 F20101113_AADYEM thomson_a_Page_36.jp2
2dfabab086c0080fd9c2b1035267fedf
1b405d0be4b268f93449ac54b261527148c853b1
26318 F20101113_AADYTE thomson_a_Page_69.QC.jpg
fe459cd79896679fc9e7a6d56e893d3e
4404a68a940e4020d53645ff20db5ea90e47d183
3608 F20101113_AADYOH thomson_a_Page_03thm.jpg
49eb19ecfa4b2f1c6127dcb5e4e44605
708ba61150afdc29a869072fef85651ba0471834
50080 F20101113_AADYJK thomson_a_Page_22.pro
22496a424187cbe74303eeabf03dad03
3a1c52024438dc05dea8c0a4cdf83b14e975744f
108148 F20101113_AADYEN thomson_a_Page_37.jp2
86997b8d74a5925cb20da38d11edbc41
0745f381b3d9665f1c7cd717a86539166d6554c6
7369 F20101113_AADYTF thomson_a_Page_69thm.jpg
e769175ffa047d269d6f33f3690e932e
35791c430c3685d77164bffe178a4852b90e0770
31828 F20101113_AADYJL thomson_a_Page_23.pro
218c458e50ef028831ba24411b2ee63e
a2fc1578d775ac7db495c4c31f423caa90e60cb1
106672 F20101113_AADYEO thomson_a_Page_38.jp2
c34b7bdae6788413910c26053da83f63
b8f0757f28f9acb9b526b5e216e39ccb4768201e
19787 F20101113_AADYOI thomson_a_Page_04.QC.jpg
c1ea9596c21d1af93eba132e3d53840d
7a8d3780ac45ec54ab7b0b791bcd8a7079de27ed
24416 F20101113_AADYTG thomson_a_Page_70.QC.jpg
65de6b18bbe0f3d7e331ab25192ff74d
0d4c55b31c37e708e9295cfe8bb0f7751b196c9a
34354 F20101113_AADYJM thomson_a_Page_24.pro
3698ee517beca71e106ed5b908416eba
8c782390c0e267eb0a8edfeab282200399ed83fa
126378 F20101113_AADYEP thomson_a_Page_39.jp2
2e3cb11dfbdc0e6d92be4638d0e2a52b
d6a423651843cbef69a73d32f79bfa31044b1437
4985 F20101113_AADYOJ thomson_a_Page_04thm.jpg
efc20f83e7f7d354cfac17b29c333fae
b8c2902ec7958da7410cddd30f7d661333c068ea
6676 F20101113_AADYTH thomson_a_Page_70thm.jpg
fe7c0d86f5d1ac82490f645fb33dc8be
020f3a3b09e3ff99677147eb8d72786be08d1fc6
59415 F20101113_AADYJN thomson_a_Page_25.pro
494aa161b3cf7feda27ee1dee4922087
899c9121d540a79eafd4c1e2948a3019e8ad7709
114994 F20101113_AADYEQ thomson_a_Page_40.jp2
b52bafc0cd8a5bc5dbc010856aee36a4
2b0b80a0572741ecc4aecaeecf709470faf30104
7800 F20101113_AADYOK thomson_a_Page_05.QC.jpg
575c17e90dd0ded4964ab16de2637c77
07a7ad176551e828544ebb543f4ab60290a005be
25040 F20101113_AADYTI thomson_a_Page_71.QC.jpg
47c5da2a42301c27284ff91e4f5101f8
ad9915de70b9f04b59f7b68a96a60bc8f31b4aa0
52401 F20101113_AADYJO thomson_a_Page_26.pro
64f1b0ae3db67efdd681d8e72ad17efa
ec847748f9f420e0186b2d7183b937c723445152
110957 F20101113_AADYER thomson_a_Page_41.jp2
ad9475b81fc536e93cad450d8b421c0c
8d3bef2bc19955e12cb4616fd825bd9af0478ff3
22787 F20101113_AADYOL thomson_a_Page_06.QC.jpg
1fff11c3decc77d9f52f3f0867ac15cb
f0a474aa8678e26fdc9521255d0d4b4c76c68c78
7000 F20101113_AADYTJ thomson_a_Page_71thm.jpg
69527354de7b9d58b76d0f2a45cc73f2
76cda22184aba0812f629936b871072d41ad8fa2
54671 F20101113_AADYJP thomson_a_Page_27.pro
0703523199a13c955bf48407d1d7c4b5
af283e91889648c9db445647286f686731e40170
38527 F20101113_AADYES thomson_a_Page_42.jp2
07eb43cf71f4f22b4917d7ff859e1f0c
8a9f44a059f52020ec45452745edd4497baea505
6106 F20101113_AADYOM thomson_a_Page_06thm.jpg
810f0ffeb35782a968ea355a966913ed
8624929fc72b921ce09f0d96409719b8ed9f21ec
7050 F20101113_AADYTK thomson_a_Page_72.QC.jpg
79c15e85edc5ee9a94d073c1cf6547b0
2bd1add54d38702e23051b3b0f9b9f4c5b908894
52160 F20101113_AADYJQ thomson_a_Page_28.pro
f5372d95ed16cad802756b5da1bdeeb7
88ac4ca04cb5e83152ef4865eb1fa31f4db15903
49718 F20101113_AADYET thomson_a_Page_43.jp2
2d6c03d481b669f0d4748f87fbecbf09
2ad95d781bdb9297ecbb81ae376644db086793da
23769 F20101113_AADYON thomson_a_Page_07.QC.jpg
c497b8e6e05d7d21456a90b29ad38861
d94bfa6736c2271347ae9404cec76ccb9eeae571
2384 F20101113_AADYTL thomson_a_Page_72thm.jpg
c89e736ae6a357e22eeb8fe7c429a7b0
b7c0613a44d546b983cdab7eaa7a4392f9de23eb
54143 F20101113_AADYJR thomson_a_Page_29.pro
0468c68b70fa7355b06fb22c27083386
bcb49c3ffaf73da5680dec4f8b342e2052ae8345
87274 F20101113_AADYEU thomson_a_Page_44.jp2
e0b1a72182cb5ebe4abe92ca540c7950
9493c819065eb0b474d3d831536fb0ae289fc47d
6206 F20101113_AADYOO thomson_a_Page_07thm.jpg
c32d4232065fa7472ef07db4e4291e13
b513f7e7b41ce0efa72f4133501cb4f4a1cdf881
13357 F20101113_AADYTM thomson_a_Page_73.QC.jpg
482a3cf6ab0b93fdcf413ed486c07b88
8fbc01381c76fcf1a978bb11782494b9750af031
52518 F20101113_AADYJS thomson_a_Page_30.pro
00c3e7e305024324d826c6e632915a01
6ef55f5b31a4787f72b5b6b5e8285c471975fc24
22099 F20101113_AADYOP thomson_a_Page_08.QC.jpg
be27088d553b8fdcdbcc1230ea83d377
2b9ed49b4844f3c4dcba24aa2052105c3c3a3f12
3775 F20101113_AADYTN thomson_a_Page_73thm.jpg
5e34260fcf31d4678eb2eb3584fa884e
1ca13bfb240550781f9192d95235c3f3e3da29ff
49291 F20101113_AADYJT thomson_a_Page_31.pro
68970bad38eb07e8b9e5eed4d05631e7
ab4c11810ba9855888f58d3b7a267b4da81d0150
87128 F20101113_AADYEV thomson_a_Page_45.jp2
0b30f01414f407c0f636cd509d7b6d20
e47eeb770f3b8f3e98c6a943fbf7917db28b1a7d
6341 F20101113_AADYOQ thomson_a_Page_08thm.jpg
41d7a71a6ba456e0e17db88403fe27f9
12d20c8427994f76658aedd8ea7209e25a9921bf
86222 F20101113_AADYTO UFE0021228_00001.mets FULL
65dc3ad06824af106b0334f9efe0c3ec
cbae07eb10a1e0ebbf95f29d8ad8948bdd3d4ef2
52264 F20101113_AADYJU thomson_a_Page_32.pro
f31815782c5b23b1628665a1f52ecc0e
28f60f2f23a9ff162921f44c9309a7b72200a55d
79391 F20101113_AADYEW thomson_a_Page_46.jp2
3253425cc570812860f84ba22d259394
924e0787330ac9f98e3338678acf9d67009371e8
18558 F20101113_AADYOR thomson_a_Page_09.QC.jpg
8d3f86ef056d4aa4597dd50c05b123d3
7680d05447689d535f5119b332079f6153bfdd93
50744 F20101113_AADYJV thomson_a_Page_33.pro
215a5caf0814260a62ac6c4f63b38861
5838ee8e13c407c615bb56361facffa1e171eeda
88680 F20101113_AADYEX thomson_a_Page_47.jp2
c80abda1fc86cb05ca034797ce51030b
cf46729ba336addb8079ebc1b86638c1dc8ade40
5425 F20101113_AADYOS thomson_a_Page_09thm.jpg
7408087012084dba9bf94989c21dd9df
e010dedc62c3d23fafefb89aebaf4b284cb94416
53556 F20101113_AADYJW thomson_a_Page_34.pro
424f1364e7dd2cbe9b4b2327d3374f1b
5da5e4a01a62d5d1c15393b8c44437c628190f8f
35699 F20101113_AADYCA thomson_a_Page_43.jpg
adb2e6829243058521da01cd7eaea8bb
25510b39158f989109c0b4bbf3521cf10f9d6d09
79116 F20101113_AADYEY thomson_a_Page_48.jp2
87d1fb0abfd735a063f6bbbf1ef7535f
a43ac5c738c71df4d5ff231f999bf78a0258f398
23998 F20101113_AADYOT thomson_a_Page_10.QC.jpg
c731f02daa4c31a9f31bcac311fab5d4
a1d0fed4a51e65d8e8713d200e143917a47e6dbe
55730 F20101113_AADYJX thomson_a_Page_35.pro
d0f3da8f5eb5b642e76c421340a06e2e
3e1cefbe5b4288c1d6c86f14064b271d717e30c4
61683 F20101113_AADYCB thomson_a_Page_44.jpg
83ca07ad6dbb7d3a24d7ceac54107129
c6d2f05356478fed17974512229f92360f8807cc
826796 F20101113_AADYEZ thomson_a_Page_49.jp2
a5c147f6be1d6df64e7c3422b88ae37b
bde2168f1fefc58fc9966112733301ca6bd0735d
6697 F20101113_AADYOU thomson_a_Page_10thm.jpg
00f3392e6a00a303ed51c8087d4407bc
804ec5f915aea254c0972dc73cb1ecc30aa6d9f1
F20101113_AADYJY thomson_a_Page_36.pro
cc317a1f9670ac10c750c83adc5ff7a0
1491001d31c252f83c24c7032c04add88c9eba9f
61746 F20101113_AADYCC thomson_a_Page_45.jpg
f18c5e90aa594313709b8b0f5b69460f
83bcb1c0797c95692d9b37b8cb829fee86e8a059
24693 F20101113_AADYOV thomson_a_Page_11.QC.jpg
60aef116987ab54392c952b55bcb8155
4a078be152231673be184082820f853dc2f8ade6
50352 F20101113_AADYJZ thomson_a_Page_37.pro
c03f5d1481aa583f0f6d53f17e79758c
427268424012b221d004a8f7eab1dc4d49495a4f
56903 F20101113_AADYCD thomson_a_Page_46.jpg
b72928b12f3c8be941cce532a74c61c4
d7647566aa42b0f44d373ba98942a6788ae7c4aa
6863 F20101113_AADYOW thomson_a_Page_11thm.jpg
52299ec84906e1c8eefc640ccf07da93
c16614cf5c4aa70e6bc5d6d7b00f3ef1c1d75df8
F20101113_AADYHA thomson_a_Page_31.tif
83ec6307ef14a2bbf17524c149e67260
4f82a3958bee6195dae462cff5f95802b125fcc6
62626 F20101113_AADYCE thomson_a_Page_47.jpg
7d3470fa008e225b6f5bfd487771e301
fa507276e3e7b699f7b51d9117bca77f1930d3ee
25963 F20101113_AADYOX thomson_a_Page_12.QC.jpg
43ba60404b2074d07a4fefd56994046b
95b3d32eba9e1a2941ac3ad55dbd8573a633e949
F20101113_AADYHB thomson_a_Page_32.tif
4eda0c126131235c372cf824cd528e42
4a7045098759037ca94057fcc33cb807014ab1cc
56986 F20101113_AADYCF thomson_a_Page_48.jpg
c28a38a706d9333a180b64e2a3ef0a52
882db84b6bd62402e7507bd02347f8ef557d695b
6944 F20101113_AADYOY thomson_a_Page_12thm.jpg
e2891e27c0a6cc830b33ca8f7f632204
03b7ef015790172fb33f310026b3ffc1c07c1cbf
F20101113_AADYHC thomson_a_Page_33.tif
a204bbda73fed8083505466c569991e2
d7e9cd71ef7c5bcc6ab0fabf6364e5219e513edc
35118 F20101113_AADYCG thomson_a_Page_49.jpg
66aeb4940e64de8614feb86a1cbd86e1
f2e7697486eaaa9a875eab25539b25593760c41f
2136 F20101113_AADYMA thomson_a_Page_18.txt
0ddf49bafa4629d99d4ae8f1afb756e8
1e8193ca219a527e61cbabcf386510b8aead88ee
25468 F20101113_AADYOZ thomson_a_Page_13.QC.jpg
53718e4c98f58f0301f414be89718e51
5ead19ad2f1862039288959bd3757bf56bd93820
F20101113_AADYHD thomson_a_Page_34.tif
12dedf8abe7aa775be36e8eb9f66cc3f
0b21a32002571ccfd2caa56db52cedb0b221c394
48451 F20101113_AADYCH thomson_a_Page_50.jpg
947d975b49e57d45cded7a32fd9ec4a5
ac4346a7f5f3de7945634387d9e78efb57640276
1939 F20101113_AADYMB thomson_a_Page_19.txt
1c0c2daf984e19c9941d755e6e4b0de3
5d1f0538158b8f4f5a0dfbfb6bff393b33d2f732
F20101113_AADYHE thomson_a_Page_35.tif
0fb8196839ac569a1aeb6261af151b5e
da0740d08d2379b144871967f067ccafc41bf3ca
6934 F20101113_AADYRA thomson_a_Page_40thm.jpg
a42b498df68cf8bb36348721269f31f3
e825856bb4c95c180e5e10170e224451f65dbc7c
34919 F20101113_AADYCI thomson_a_Page_51.jpg
fc7ddba0b095291bd1c7261a414ccd7a
d9a77a25bfd99e29536b4e1ea79ff65d23de24bb
993 F20101113_AADYMC thomson_a_Page_20.txt
04d5dd247349f30b081c00e0604e1bf9
33bc2b87fda0450d3d6498571d6bee284d344126
F20101113_AADYHF thomson_a_Page_36.tif
4020a222b6335a514cd3d7f81e4faeb1
80f6e5776ae954294d76cad27aa333712b374fca
24010 F20101113_AADYRB thomson_a_Page_41.QC.jpg
02491beb0d3cce8e754c5e293c5497a5
a98091c5a1bc083e3eb223ff5bf7ed624697308d
48605 F20101113_AADYCJ thomson_a_Page_52.jpg
65d4d10a52cd8814ee2f528d7fb7c807
c73a0ec8eca05152bd8fd2213ca12871b89a9920
F20101113_AADYMD thomson_a_Page_21.txt
81622d9f90aa5cf11c640ebe0b87dfac
c9ef76666f6265b88c60d6deadd2b5d22ded22c4
F20101113_AADYHG thomson_a_Page_37.tif
3a00cd86beb5aba53c5688412a498c80
23cdb0333f2fdc60d7416d819ff8d26d7a9d41f6
36995 F20101113_AADYCK thomson_a_Page_53.jpg
81c1300754b88ee5a6c9d3a5ce38beaf
13f550a79d261c670677ba885c3e13a909dea265
2052 F20101113_AADYME thomson_a_Page_22.txt
d368b4b5da3a1cdef7b8c1d8e2957570
e87a380b87baeb5a82b6798dadaf4b12132ca311
F20101113_AADYHH thomson_a_Page_38.tif
e3f5f77e760f0102abe667900292e80b
79cf520b3039b746719c896b84f1f507f8b7f002
6756 F20101113_AADYRC thomson_a_Page_41thm.jpg
625d2e7ce6e734c20ccfb0f3e9ef1392
d22c62c80fbe155c6731a765a5198020d86f601b
48564 F20101113_AADYCL thomson_a_Page_54.jpg
9384a129023a36a78f3d8992ccb91739
1a41eee6bfdefcceb705b00b97080d488403567b
1319 F20101113_AADYMF thomson_a_Page_23.txt
a6c5a2e5f9428fcfb3bb948e74567e7c
2273b6edde3414c3cd85e9cf172f7457b1510850
F20101113_AADYHI thomson_a_Page_39.tif
33e7819d893dfab3b847eaf5c1bf9a64
b9fd49187ebb347a51037c17f766f7c0b0040f12
9796 F20101113_AADYRD thomson_a_Page_42.QC.jpg
a79f1e6040d327abb64a96bb9d26a468
c6387bdea42466d0e12f1c70a75ff628f66a308e
37513 F20101113_AADYCM thomson_a_Page_55.jpg
0f80d2e2c0081954de8905b462fcfd51
f9a4ae3bdb8e9c81ee477b87c69f0ecdaf48193a
1479 F20101113_AADYMG thomson_a_Page_24.txt
74d01447d8f36d37ab7d20f571d9fddf
7998b442cb23839ecf87872de8d1fe290e11ab76
F20101113_AADYHJ thomson_a_Page_40.tif
29e323f974a07ead46b7d632a389131e
cfb50a4182b05e5a94157c92f6fe09abcfe89f5f
3043 F20101113_AADYRE thomson_a_Page_42thm.jpg
a0c0125c4affa74d960d520c83627797
43f11464a77019b4150356738950f8e56cd8c6da
47825 F20101113_AADYCN thomson_a_Page_56.jpg
f6d0daaa9745bf9933a7032713542a59
3a8a94ebcfd2c5aa85f53aab7de8cf0b2fb1fec1
2456 F20101113_AADYMH thomson_a_Page_25.txt
6301a874f06309044217b7f3c8fda2b5
895414912b885677d800e733f5aa2da4185f5644
F20101113_AADYHK thomson_a_Page_41.tif
5456e44e95796e63d58eeceab6e6c30d
7489f4c60ded495d44d28dc07c6972f35895962e
12378 F20101113_AADYRF thomson_a_Page_43.QC.jpg
0359bc6be5943c3d30e09e3ccd7990e2
0f7745cfe0abd3d187b2b8195c85d67192efaa75
36658 F20101113_AADYCO thomson_a_Page_57.jpg
a2a116091a68c1ea48b856ae1b158734
22d2094ed103f46f0678a923f2ea002a31074bf9
2073 F20101113_AADYMI thomson_a_Page_26.txt
c3b87ab30b266446422507489af1ffeb
f69fab343b5d571822ada46161c75d33ffc576b5
F20101113_AADYHL thomson_a_Page_42.tif
e9733ef6698bf406f7b2181ccb0024a7
6b637a4b536e2cdc8a84609ab69e0978c76f273e
4073 F20101113_AADYRG thomson_a_Page_43thm.jpg
d7bebff2d1d3be4d91a50e0ca8706ba3
4e6202d30072431557e8c174ffa5e2af0a79965b
48252 F20101113_AADYCP thomson_a_Page_58.jpg
827a4cb5f35df50e4ed7e87d627a220c
b815b21a52c97e2204cdcfb407e7b823dcf99862
2166 F20101113_AADYMJ thomson_a_Page_27.txt
521a9807fabc0b5670cc1f172eb8deff
6da17b6a08f36ab45fc5174acf3f66cb1af862ee
F20101113_AADYHM thomson_a_Page_43.tif
fbecd57c3c6eb7fd108a8ed347c2df5f
fba3a0c9d3a62e5ccde76fe41e4e4805c6f4c5a7
18197 F20101113_AADYRH thomson_a_Page_44.QC.jpg
28e5e48fef2aa7a0a568b774aa575ba9
0dcd5448736ca03c8d42c4a57f28704ff9762995
76226 F20101113_AADYCQ thomson_a_Page_59.jpg
6c5f664a3243b0971d36d43eb4431595
b423cde8d927302fab5ed3ca02d64bfd81e24b32
2171 F20101113_AADYMK thomson_a_Page_28.txt
03b5a3195cef24171481831385472d0d
e7cbf100c2bb3bbd17a216983958a91153ed2e81
F20101113_AADYHN thomson_a_Page_44.tif
ec77fdfa4ef8d7f981fb924b7eb2a65a
44189edc87b1e7ffe3827397223db641040de78b
4757 F20101113_AADYRI thomson_a_Page_44thm.jpg
1ea439fc8d38510208daf772d93940b9
54f79fc7ff30b3d58cceb4c0180d5c75b02ad765
77042 F20101113_AADYCR thomson_a_Page_60.jpg
dd623e921a13bc64be29388069b39178
ef32cd987b4d283a719c1621a8f91c9364d3c3d5
F20101113_AADYML thomson_a_Page_29.txt
3d5cb6fc0c58877d311b9d50fd5d8dcb
a129dc5334fb4f575fd4c1964fbd79d71081667c
F20101113_AADYHO thomson_a_Page_45.tif
c1d7d5a979d243ee7be977497522b242
d818e300d2d35c6515c00cc43f1711fa32142862
18096 F20101113_AADYRJ thomson_a_Page_45.QC.jpg
40dab2c7739f337f1c390ef8e09b86c9
de7e96a33cdb49d6fb7ab67d2eb391563c3d9a3e
71245 F20101113_AADYCS thomson_a_Page_61.jpg
d87c197071b0008118bb81bbd9b1d755
3baeb9c151df4b115e2cabfdc2977e66249cac13
2081 F20101113_AADYMM thomson_a_Page_30.txt
87184417426bf9c80bb23772c42ea582
589c81a21f30aba736bfe9394b66e8e961dde2d9
F20101113_AADYHP thomson_a_Page_46.tif
c9a722169f64cb4b4c7e8356b0759a4c
8a94835ebb6b7c91c29ac27c4f779d5743b4c6d0
4633 F20101113_AADYRK thomson_a_Page_45thm.jpg
9328412664c7ddd93d63a25b6934cfe0
c3eaa27176b7ed8fcce9dd06bbed37230478953c
2031 F20101113_AADYMN thomson_a_Page_31.txt
c9eaf2858573716dd442035421d7dc3c
0ba549ec6f92ee566cdfbd6adaa8ab64b7c621da
F20101113_AADYHQ thomson_a_Page_47.tif
844a48c1e5f85a07dc9c4faad639ed29
d53405c56926436da86d6e7e71601e81b2439793
16681 F20101113_AADYRL thomson_a_Page_46.QC.jpg
ecec2138dae154ca4c5f995c36f1d25d
4743ea2ac356ea3293369fce786e0eeb66653131
73077 F20101113_AADYCT thomson_a_Page_62.jpg
20b1152734e0654775b03c3a4c60c2ca
741397015b09a83ac02bba95fe1ccd24fd72da7f
2082 F20101113_AADYMO thomson_a_Page_32.txt
3329e17c9221f6aaff2b0cd61396350e
db6c1084a650871c457590963078db0d74eed32a
F20101113_AADYHR thomson_a_Page_48.tif
7a845cbd8b2491ddcea020ef724b7e8e
289e2d56d11fafa563718f456dd476d0d745be0e
F20101113_AADYRM thomson_a_Page_46thm.jpg
ca7bf6abbc63b4327fd748a07c4b0b5c
735c5ca8eea51c56173f319a1646904dc6c68e9b
62745 F20101113_AADYCU thomson_a_Page_63.jpg
581ca79ab494420d07d7e1d5d3b019ec
6ee453050df0b843c14e81bea0425d5877145c09
2009 F20101113_AADYMP thomson_a_Page_33.txt
5d5baea94f9ee05dc39b3ce208b801fe
99ee30075cbd4bd14118183844076994865c3ea6
F20101113_AADYHS thomson_a_Page_49.tif
53de7fcbd175c909ff6b21fd5fef2b75
d7753d7417ee5f35adb4b318bb2714257f13c8b9
18382 F20101113_AADYRN thomson_a_Page_47.QC.jpg
7c7d3652b26e414158ee5adcd11bc0d1
933b040b5ebc40a9f491d384ca365cd10769d449
55752 F20101113_AADYCV thomson_a_Page_64.jpg
65707fa384b3767f72ac88e6a53ea95b
88d34763c39c5023d03d184cb1993455aa973899
2116 F20101113_AADYMQ thomson_a_Page_34.txt
6f3e94557d20e14efa3eafe3085af8b4
e2b24cdfe8d6e55baf1df2b2033e7997be79cb47
F20101113_AADYHT thomson_a_Page_50.tif
d9880dd2cf7caf5de73c56f5f1e28a33
39239998efc9e0a6965a68a3ea65068ec713b60e
4642 F20101113_AADYRO thomson_a_Page_47thm.jpg
a8675efb59cc2943c2c6b378e17365b3
01262435cfcfcf0a44871da3daee38552518544b
54735 F20101113_AADYCW thomson_a_Page_65.jpg
9ddc6b2367ea6aefb3023a342e31a558
105d24b3ce1a1175300a0404041e465a2f8d393f
2346 F20101113_AADYMR thomson_a_Page_35.txt
cb8e38a4c3077c693e6aea8af471c013
0d7dc46a95b73c59bfa37ec10558d845a58d6e34
F20101113_AADYHU thomson_a_Page_51.tif
f5fafc5fb1716f4ccf43ad3935adbeee
99b7e14297d13c4a98131e7e24a4fc9305b119f5
16867 F20101113_AADYRP thomson_a_Page_48.QC.jpg
0192a79b4632023f9895fa001d0b470b
17557093294379d3048df8882971f4c5c09d8de0
2046 F20101113_AADYMS thomson_a_Page_36.txt
b26a52682d49a4f9d2ed0e5d7ecee0c7
01a14ec6797bf022ceece750b349b5e418fabcec
F20101113_AADYHV thomson_a_Page_52.tif
b9f0d93216c0ee92d37dd08273b42468
1eca3a0ad195708d1aba5e500c51b3ad75ea9557
40398 F20101113_AADYCX thomson_a_Page_66.jpg
5a59ef4f043c6b5f33879560c19ccd05
3b44795741e0fc8db71c14db7e81d5e6421fa851
4336 F20101113_AADYRQ thomson_a_Page_48thm.jpg
bd8e62ebfab36ae5b96873fcd7b57bad
8acdd31205fe28af6af3df2e65b283ee48083bfe
2001 F20101113_AADYMT thomson_a_Page_37.txt
6a2da02bc68a08da8f64dd7d2241d9d4
33da15d707aca8a04e18599f0663e20d44b604ed
F20101113_AADYHW thomson_a_Page_53.tif
8296a7dd3c416eec14da5f397df7370a
c4b28362e8720f180360abdb4e58bc1dde5b5a14
37552 F20101113_AADYAA thomson_a_Page_09.pro
70d8fc529e89749d3c1796d3191ccaea
7b85c1b98910baa7b355952041aec055d3ecf139
83825 F20101113_AADYCY thomson_a_Page_67.jpg
e87895b1d7346e0b5ea19365a6189528
215ef26bd63205bc6360ce9724ea0c9cc6118410
11795 F20101113_AADYRR thomson_a_Page_49.QC.jpg
99a3b31f9a5f62663f34a4343c75a9dc
9d5a3961e74f6594cbe491f6b87532c54bdc9590
2007 F20101113_AADYMU thomson_a_Page_38.txt
315eb26378605a9e8b2336274714744b
af69787fe7b6322ab415a14d34e5ca4ca4b09a68
F20101113_AADYHX thomson_a_Page_54.tif
d935221300c16eb7400f8de058f33c2c
faddf75349a3042aee7b21e631a06c5365442af0
6694 F20101113_AADYAB thomson_a_Page_61thm.jpg
ed0b1f3d5e5d861e861a893dc75bbb27
6e794a56f78273a99fd7cbe0d411a23c7f4615fe
83863 F20101113_AADYCZ thomson_a_Page_68.jpg
5da2c57021a4eb62e060338e579c0b9a
35fc60eb2b11c00de44b42c9ca7790a553815b66
4599 F20101113_AADYRS thomson_a_Page_49thm.jpg
aa4b547154475e45e0a360860f824bd9
660b6ecf363445cb40024936cb96024041a42198
2483 F20101113_AADYMV thomson_a_Page_39.txt
6f209d23bbcea68909ea07547205b5e9
52885f4d84fb096edbd01aad1888d606f4145bee
6920 F20101113_AADYAC thomson_a_Page_22thm.jpg
dac852b917d2d2222c46521114e9d822
38f3e5d85bd3a5657b9ffa9615fad088e339e667
14867 F20101113_AADYRT thomson_a_Page_50.QC.jpg
d47d7e5b608548c1d133672d5e213bbc
96a6fdbd76b96a7f20905924f382778730007d58
2120 F20101113_AADYMW thomson_a_Page_40.txt
f87fedf84ff86b1601d62148fe14ccf2
3b10ce762dca070bd9c2c78db6fcc4eae57e2d26
578471 F20101113_AADYFA thomson_a_Page_50.jp2
e208e5c299de439a6881d72b7cd895a2
fb99f9bea0930d955b3f917aa5a1149ba429ca4c
F20101113_AADYHY thomson_a_Page_55.tif
29c45d6722029da34144cb6a027ead59
a383c22328a3a648e264ff145bbfd60d3a6b5d55
126504 F20101113_AADYAD thomson_a_Page_67.jp2
8e4e08cb41c2c1d4dc1cdcdddb8a5d5e
dac66cd4d3ebddad683cb56daeced7ce7d132a4f
4634 F20101113_AADYRU thomson_a_Page_50thm.jpg
f9c3f4c183ca529997d7d667caa5caca
a9e2c54cb29c07fcb66580460641cf3402861842
2093 F20101113_AADYMX thomson_a_Page_41.txt
2b77f1c6315b3676749fb3e315832595
a51ba678d20ed6141df47877dce92d7f746913b0
883289 F20101113_AADYFB thomson_a_Page_51.jp2
f5d9f6c6ce3dc547ddf11619fb80020f
ae213fa41575c1dbd927bd2e0b3c36b12bffe8bd
F20101113_AADYHZ thomson_a_Page_56.tif
bbe181f29cfd986308636f92eb4ec9b1
7649e20c9b368a9432835d382b0fef590c17cced
117386 F20101113_AADYAE thomson_a_Page_13.jp2
d710e2b44cf9fa2351dba2ee51811f56
50889c0cc8ade9a1094935147e22e9b66fa68dd3
11709 F20101113_AADYRV thomson_a_Page_51.QC.jpg
d77e7fef2ab1f9407f0af182c1e2f9db
be556aa498fb5705f5b96dcc45f214ad6458cadb
660 F20101113_AADYMY thomson_a_Page_42.txt
207d79f1a3b905dd34ac6cf0718a8e1d
9ed8afa413613e0fc8ce57b897f7feb208d318c6
572407 F20101113_AADYFC thomson_a_Page_52.jp2
541f7f8bb2f35ae373904923eb8fbb34
f7ab369ca8d177379bd0045c7a98f56b12212d13
115395 F20101113_AADYAF thomson_a_Page_14.jp2
9046656a5392ff26dec32c309088b04d
00d357e8948ac34e8ed16d561d4ff51ca4b2e0cf
4331 F20101113_AADYRW thomson_a_Page_51thm.jpg
3154ab723fe79d77fd20e44123277b18
faa1c68b158da76b223090de4fe2cf7865ab234b
1036 F20101113_AADYMZ thomson_a_Page_43.txt
69350a89e5bf76394a1a6efdfa99d04c
55125878c77723efc60c8666ee54cc3b3f21ca2e
977417 F20101113_AADYFD thomson_a_Page_53.jp2
162586a26df0c32afc17f09e5f66a718
9007993088da459a68674ca44dc33ade03315bdf
F20101113_AADYAG thomson_a_Page_06.tif
4e96f6c848eeff28927f7345804107ca
14ed869e2dae1c4a083b7b83ad8d538b44bab029
50339 F20101113_AADYKA thomson_a_Page_38.pro
a1f38e1f12605fadf8724d3eb2abb1a1
e1bc2901eff540919cd24a5976edfaab4d7e5f02
14857 F20101113_AADYRX thomson_a_Page_52.QC.jpg
616dc4fb279143ae898e7f1dbb0ce778
340540df6f8170d1c55fc9a7ba658a79d0fd14f0
578928 F20101113_AADYFE thomson_a_Page_54.jp2
94ed2cb344a55e1366ac3010ee60aeff
0698e1b3b41ac5a9a73e35c7efde982d959238eb
2338 F20101113_AADYAH thomson_a_Page_05thm.jpg
6da15e4e7c0d81471e79e1d42024e643
46b6e7134f1d98e0f426bc2d5b4ddebcbb1c266c
59645 F20101113_AADYKB thomson_a_Page_39.pro
2968e6e71abcece9d9ee9ee136ceb43a
8b41df7dea20e087f1e9104a302815bb3d77867b
4550 F20101113_AADYRY thomson_a_Page_52thm.jpg
ac25255722be2b486176086c525def1c
bac0e64ed7331b99a7aaa1773c2ff553d3003421
897175 F20101113_AADYFF thomson_a_Page_55.jp2
dc130784c7855980694657a580e2ce0e
dbd4ce8a131319ab5e09425f924888e0df537237
111517 F20101113_AADYAI UFE0021228_00001.xml
4701ec2a764851092d03c0bee87a665a
4c5e9d3ce1f045c1c197c9c57d52dc1894704fec
53610 F20101113_AADYKC thomson_a_Page_40.pro
a36b5063b072ee9fb6030d6c8995e7d6
263571179a411180f626a01ea845303f930ce2df
11841 F20101113_AADYRZ thomson_a_Page_53.QC.jpg
d3779b666d14e350adb99d470606b23d
23d01d887bc4159fb11cb30efe52ad2a356af106
567275 F20101113_AADYFG thomson_a_Page_56.jp2
2efd8e378db187d32218376e03176eaf
f09737fc9159e155c394664a1f846c2eb86f08dd
6866 F20101113_AADYPA thomson_a_Page_13thm.jpg
c3e5203e0fca062a01cbc50948bae886
07ef0d6f3cf6472f0ec14a99fdc9a66482eeabfb
52999 F20101113_AADYKD thomson_a_Page_41.pro
62cfccbb439c4e87d6774a450497b68e
6885e867bf5a2e8e5fed3600c6f5b98ad5516a65
804519 F20101113_AADYFH thomson_a_Page_57.jp2
e226c607682f2638c483b22a6a37b743
a526bdd3e3638c93b480445a4d28929d7758c74a
24669 F20101113_AADYPB thomson_a_Page_14.QC.jpg
adebc0930129f4c08e34164052b87845
23e2ebc58415c1165574f02af71c5cb2eed5f317
16137 F20101113_AADYKE thomson_a_Page_42.pro
6674e7e74aba894a422bc98245442125
ea90e73dbceeb94a62467b6417f6f964e16a9fcf
572271 F20101113_AADYFI thomson_a_Page_58.jp2
6eabfebd1db9471961045a7ed7b53a63
591ca28c394c4668d21ddeca12d39883708c6dd9
6705 F20101113_AADYPC thomson_a_Page_14thm.jpg
b9a57e4344e1906e19b179777ac57fdb
4d88f7561e6957aa898fd3e6fc0f875848f016af
24485 F20101113_AADYAL thomson_a_Page_01.jpg
9b2b2e3bda5a2549714103b7b6cc0bf3
babc84f1f94d0d2cd19a4b3bef2f392446031d83
21235 F20101113_AADYKF thomson_a_Page_43.pro
1f1d7cc8dae6204e0a7394c9e3dd29bf
d65a23ff66031bec3c8b0b8e4c84e2032fde0043
117003 F20101113_AADYFJ thomson_a_Page_59.jp2
6cc98d78de0cc8814944e47a6b6df609
9d142ff9b45f424a8f83722930a7ccb4eec83307
5772 F20101113_AADYPD thomson_a_Page_15.QC.jpg
b85f98bd36b690bda6dd378be55852b0
9de8765c9ebc58c13885842dd5ffce4b48f416c4
10184 F20101113_AADYAM thomson_a_Page_02.jpg
67e6c5445cd03eebb7aa6b72214be68f
9ea7c1a78ec95e51b1e981e2b86fc63e74a0abe4
47994 F20101113_AADYKG thomson_a_Page_44.pro
7804c74fa971ee049420bb47a6ab1c09
40f12c615205e4e1490f94a2a08391120113bfa1
120634 F20101113_AADYFK thomson_a_Page_60.jp2
63a860231dcc7b0b69c6ceebb31de34a
c17b6eb8ab4f34bd0bb85edec62da39fbec72513
2029 F20101113_AADYPE thomson_a_Page_15thm.jpg
774d9dee303d71c76b59b79461245351
6103bedb417e6219a701d6b291da5f2e3b882983
38010 F20101113_AADYAN thomson_a_Page_03.jpg
ed5ca2efb4e1b03e9cb701b2a74a0e5c
50d7fd4dc63765fb53964e1a6332e846db5eae41
47942 F20101113_AADYKH thomson_a_Page_45.pro
9d90c078cea59b1900a48a9de85e686e
204b553bcf945cda7f498603e2f547891d3a0d7d
14753 F20101113_AADYPF thomson_a_Page_16.QC.jpg
877c8f56be939082dd1a5c2f4491bc02
67cac07431705bf79df4b8177622d05475793d55
110456 F20101113_AADYFL thomson_a_Page_61.jp2
4379fba34a259e8d0567e538899b425e
e9664f818a7dbd263689567dfc8ad8c08175c0ef
73431 F20101113_AADYAO thomson_a_Page_04.jpg
fb06e99a302a02c6b07491856f324553
e37afff5f44f2889774a3514256f95e2fed88f15
46010 F20101113_AADYKI thomson_a_Page_46.pro
a7750de6e4977f9e4cfd4aea77ebbe1e
6abe96e24e0ac55bee198b6eef8000d9bb3f23ce







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

































2007 Alfred William Thomson









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).










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









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









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









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









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),









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.









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









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.









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).









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









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.









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.










-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.










-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).









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.









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.









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.









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 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),









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).









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.










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









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









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,









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









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









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









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.










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);









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









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].









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









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









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 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









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









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.









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.









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











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










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










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









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









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
















































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










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
















































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










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


















































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










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































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


B~ '"










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


















































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














57










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










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









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.









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.









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.









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.









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










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

















































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









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.









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
71:1-5.

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.









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,









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.









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
Museums, Natural History 16:41-71.

SKELTON, P. H. 1986. Two new Amphilius (Pisces, Siluroidei, Amphiliidae) from the Zaire River
system, Africa. Revue de Zoologie Africaine 99:263-291.

SKELTON, P. H. 1989. Descriptions of two new species of West African amphiliid catfishes
(Siluroidei: Amphiliidae). J. L. B. Smith Institute of Ichthyology, Special Publication no.
48:1-13.

SKELTON, P. H., 1994. Diversity and distribution of freshwater fishes in East and Southern
Africa. p.95-131. In G.G.Teugels, J.F. Guegan and J.J. Albaret (eds.) Biological diversity
of African fresh- and brackish water fishes. Geographical overviews presented at the
PARADI Symposium, Senegal, 15-20 November 1993. Annales Musee Royal de
1'Afrique Centrale Zoologiques.

SKELTON, P. H. 2001. A Complete Guide to the Freshwater Fishes of Southern Africa, Second
Edition. Struik Publishers, Cape Town.

TWEDDLE, D. 1983. The fish and fisheries of Lake Chiuta. Luso: Journal of Science and
Technology (Malawi) 4:55-81.

VAILLANT, L. L. 1897. Siluroide nouveau de l'Afrique orientale (Chimarrhoglanis leroyi).
Bulletin du Museum d'Histoire Naturelle, Paris 3:81-84.

VANDER HORST, C. J. 1931. Some South African siluroid fishes. Annals of the Transvaal Museum
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).










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.









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.





PAGE 1

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

PAGE 2

2007 Alfred William Thomson 2

PAGE 3

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

PAGE 4

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

PAGE 5

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

PAGE 6

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

PAGE 7

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

PAGE 8

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

PAGE 9

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

PAGE 10

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

PAGE 11

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

PAGE 12

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

PAGE 13

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

PAGE 14

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

PAGE 15

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

PAGE 16

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

PAGE 17

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

PAGE 18

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

PAGE 19

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

PAGE 20

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

PAGE 21

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

PAGE 22

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

PAGE 23

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

PAGE 24

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

PAGE 25

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

PAGE 26

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

PAGE 27

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

PAGE 28

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

PAGE 29

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

PAGE 30

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

PAGE 31

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

PAGE 32

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

PAGE 33

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

PAGE 34

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

PAGE 35

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

PAGE 36

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

PAGE 37

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

PAGE 38

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

PAGE 39

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

PAGE 40

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

PAGE 41

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

PAGE 42

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

PAGE 43

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

PAGE 44

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

PAGE 45

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

PAGE 46

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

PAGE 47

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

PAGE 48

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

PAGE 49

Figure 4-1. Amphilius grandis, BMNH 1912.3.22.119, 113.6 mm SL; lateral, dorsal and ventral view. 49

PAGE 50

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

PAGE 51

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

PAGE 52

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

PAGE 53

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

PAGE 54

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

PAGE 55

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

PAGE 56

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

PAGE 57

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

PAGE 58

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

PAGE 59

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

PAGE 60

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

PAGE 61

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

PAGE 62

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

PAGE 63

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

PAGE 64

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

PAGE 65

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

PAGE 66

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

PAGE 67

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

PAGE 68

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

PAGE 69

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

PAGE 70

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

PAGE 71

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 Museums, Natural History 16:41-71. SKELTON, P. H. 1986. Two new Amphilius (Pisces, Siluroidei, Amphiliidae) from the Zare River system, Africa. Revue de Zoologie Africaine 99:263-291. SKELTON, P. H. 1989. Descriptions of two new species of West African amphiliid catfishes (Siluroidei: Amphiliidae). J. L. B. Smith Institute of Ichthyology, Special Publication no. 48:1-13. SKELTON, P. H., 1994. Diversity and distribution of freshwater fishes in East and Southern Africa. p.95-131. In G.G.Teugels, J.F. Gugan and J.J. Albaret (eds.) Biological diversity of African freshand brackish water fishes. Geographical overviews presented at the PARADI Symposium, Senegal, 15-20 November 1993. Annales Muse Royal de lAfrique Centrale Zoologiques. SKELTON, P. H. 2001. A Complete Guide to the Freshwater Fishes of Southern Africa, Second Edition. Struik Publishers, Cape Town. TWEDDLE, D. 1983. The fish and fisheries of Lake Chiuta. Luso: Journal of Science and Technology (Malawi) 4:55-81. VAILLANT, L. L. 1897. Silurode nouveau de l'Afrique orientale (Chimarrhoglanis leroyi). Bulletin du Musum dHistoire Naturelle, Paris 3:81-84. VAN DER HORST, C. J. 1931. Some South African siluroid fishes. Annals of the Transvaal Museum 14:246. WALSH, S. J., L. J. CHAPMAN, A. E. ROSENBERGER, AND C. A. CHAPMAN. 2000. Redescription of Amphilius jacksonii (Siluriformes: Amphiliidae) with habitat and life-history notes. Ichthyological Exploration of Freshwaters 11:163. WEINELT M. 1996-2006. Online Map Creation. http://www.aquarius.ifm-geomar.de/ (July, 2007). 71

PAGE 72

WHITEHEAD, P. J. P. 1958. A new species of Chiloglanis (Pisces, Mochocidae) in Kenya. Annals and Magazine of Natural History (Ser. 13) 1:197. 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. 72

PAGE 73

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