Systematic revision of piscivorous cichlids of Lake Victoria

A systematic revision of the piscivorous haplochromine Cichlidae
(Pisces: Teleostei) of Lake Victoria (East Africa). Part I
M.J.P. v a n O i j e n
Oijen, M.J.P. van. A systematic revision of the piscivorous haplochromine Cichlidae (Pisces:
Teleostei) of Lake Victoria (East Africa). Part I.
Zool. Verh. Leiden 272,24.xii.1991:1-95, figs 1-114, tables 1-18.— ISSN 0024-1652.
Key words: Cichlidae; piscivorous haplochromine cichlids; Lake Victoria; Haplochromis; Harpago-
chromis; Prognathochromis; new species; geographical variation.
The distinction between Harpagochromis and Prognathochromis, and the distinction between these
and other cichlid genera of Lake Victoria as proposed by Greenwood (1980), is demonstrated to be ar-
tificial. The BM(NH) collections of Haplochromis dichrourus and H. pellegrini were found to be polyspe-
cific and are redescribed. On the basis of specimens formerly included in H. pellegrini a new species is
described. Five new species are described based on material from the Mwanza Gulf and Maisome
Island. Specimens of H. dichrourus from different areas of Lake Victoria are described separately to
demonstrate geographic variation.
M.J.P. van Oijen, Nationaal Natuurhistorisch Museum (Rijksmuseum van Natuurlijke Historie)
Postbus 9517, 2300 R A Leiden, The Netherlands.
Contents
Introduction 3
Material, methods and techniques 7
Taxonomy 9
The generic classification of the Lake Victoria haplochromine Cichlidae 9
Species descriptions 21
Haplochromis bareli spec. nov. 21
Haplochromis dichrourus Regan 30
H. dichrourus "Uganda" 31
H. dichrourus "Majita" 37
H. dichrourus "Mwanza" 40
Haplochromis maisomei spec. nov. 50
Haplochromis kunjunjui spec. nov. 54
Haplochromis pyrrhopteryx spec. nov. 57
Haplochromis chrysogynaion spec. nov. 67
Haplochromis pellegrini Regan 73
Haplochromis harpakteridion spec. nov. 76
Concluding remarks 81
Acknowledgements 82
References 82
Figures 104-114 86
Introduction
U n t i l recently, L a k e V i c t o r i a , l i k e L a k e s Tanganjika a n d M a l a w i w a s a p p r o p r i a t e -
l y c a l l e d a " c i c h l i d l a k e " ( G r e e n w o o d , 1984; W i t t e , 1984a). C i c h l i d s , w i t h a n estimat-
e d n u m b e r o f m o r e t h a n 300 species (Witte & v a n O i j e n , 1990), d o m i n a t e d the n o n -
Ν
g
un
η
χ
Figs. 1-2, Haplochromis barelispec.nov.; 1 (left), R M N H 30635, SL 97 mm, sexual active d"; 2 (right), R M N H 30388, SL 100 mm, spent 9. m
<
71
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Figs. 3-4, Haplochromis dichrourus "Mwanza" Regan, 1922; 3 (left), R M N H 30064, SL 89 mm, sexually active <f; 4 (right), R M N H 30060, SL 132 mm,ripe<?.
g
m
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Ο
Ο
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Figs 56 Haplochromis pyrrhoptetyx spec, nov.; 5 (left), R M N H 30038, SL 137.1 mm, quiescent c? ; 6 (right), R M N H 30038, SL 178 mm, spent 5. -a %
r
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m
η
η
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1
r
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Fig. 7 (left), Haplochromis chrysogynaion spec, nov.; R M N H 30466, SL 89 mm, maturing 9. Fig. 8 (right), H. maisomei spec, nov., R M N H 30122, SL 98.1 mm, quiescent cf.
Ol
6 ZOOLOGISCHE V E R H A N D E L I N G E N 272 (1991)
c i c h l i d fishes (11 f a m i l i e s a n d c. 50 species), not o n l y i n n u m b e r of species b u t also i n

b i o m a s s . K u d h o n g a n i a & C o r d o n e (1974) estimated 83% of the i c h t h y o m a s s of L a k e
V i c t o r i a to consist of h a p l o c h r o m i n e c i c h l i d s . Since the e x p l o s i v e increase of the N i l e
p e r c h p o p u l a t i o n , the t e r m " c i c h l i d l a k e " n o l o n g e r s e e m s a p p l i c a b l e to L a k e
V i c t o r i a . F o r m e r l y , h a p l o c h r o m i n e c i c h l i d s w e r e f o u n d i n a l l habitats of the l a k e ,
whereas they are n o w restricted to v e r y s h a l l o w areas (Witte et a l . , i n press). Espe-
c i a l l y i n the deeper w a t e r (> 4m) the N i l e p e r c h has h a d a d e v a s t a t i n g effect o n the
o r i g i n a l f a u n a . W i t h the e x c e p t i o n of a f e w p e l a g i c z o o p l a n k t i v o r o u s species a l l
h a p l o c h r o m i n e s h a v e d i s a p p e a r e d . L a r g e n o n - c i c h l i d s l i k e Protopterus aethiopicus
H e c k e l , 1851, Bagrus docmak (Forskál, 1775) a n d Clarias gariepinus ( B u r c h e l l , 1822)
occur o n l y i n v e r y s m a l l n u m b e r s ( G o u d s w a a r d , L i g t v o e t , W i t t e , pers. c o m m . ) . O n l y
the s m a l l p e l a g i c z o o p l a n k t i v o r o u s c y p r i n i d Rastrineobola argentea ( P e l l e g r i n , 1904)
has t h r i v e d , a p p a r e n t l y p r o f i t i n g f r o m the absence of c o m p e t i t i o n for f o o d w i t h h a p -
l o c h r o m i n e s ( W a n i n k , 1991; L i g t v o e t , G o u d s w a a r d , pers. c o m m . ) .
A s changes i n the species c o m p o s i t i o n i n the M w a n z a G u l f h a d a l r e a d y been
n o t e d before the p o p u l a t i o n e x p l o s i o n of the N i l e p e r c h i n 1984 ( G o u d s w a a r d , 1988;
W i t t e & G o u d s w a a r d , 1985) the d i s a p p e a r a n c e of the c i c h l i d fishes i n this area n o
d o u b t w a s q u i c k e n e d b y the h i g h f i s h i n g p r e s s u r e of the c o m m e r c i a l t r a w l e r s .
H o w e v e r , the s i t u a t i o n i n the M w a n z a G u l f is at present not different f r o m areas out-
s i d e the G u l f w h e r e h a p l o c h r o m i n e fisheries never existed (Witte et a l . , i n press). T h e
deeper waters are n o w c o m p l e t e l y d o m i n a t e d b y N i l e p e r c h f o r m i n g 95-100% of the
d e m e r s a l i c h t h y o m a s s ( G o u d s w a a r d , 1988; L i g t v o e t & M k u m b o , 1990). N e a r rocks
a n d i n s h a l l o w bays w i t h reed a n d p a p y r u s m a r g i n s , areas not frequented b y large
p i s c i v o r o u s N i l e p e r c h , the o r i g i n a l f a u n a seems m o r e or less intact. H o w e v e r , as
these areas f o r m e r l y w e r e u s e d as nurseries b y m a n y h a p l o c h r o m i n e species f r o m
deeper waters, the species c o m p o s i t i o n there has also c h a n g e d . It has been suggested
that the s h a l l o w areas w o u l d serve as refugia for species f r o m deeper waters. T h i s
s e e m s u n l i k e l y f o r t w o reasons. F i r s t l y , the s p e c i f i c r e q u i r e m e n t s of d e e p w a t e r
species cannot be f o u n d i n s h a l l o w water habitats, a n d secondly, it is i m p r o b a b l e that
there w o u l d be space for ' a l i e n ' species i n the s h a l l o w water c o m m u n i t y w h i c h a l -
w a y s has been the m o s t speciose of the L a k e ( G r e e n w o o d , 1974; W i t t e , 1981). There-
fore w e m u s t a s s u m e that the species w h i c h h a v e d i s a p p e a r e d f r o m the deeper w a -
ters, are extinct.
Since 1977 the H a p l o c h r o m i s E c o l o g y S u r v e y T e a m ( H E S T ) , u s i n g a v a r i e t y of
f i s h i n g techniques (van O i j e n et a l . , 1981), has m a d e a n extensive collection of h a p l o -
c h r o m i n e c i c h l i d s f r o m the M w a n z a G u l f area. T h e o r i g i n a l p u r p o s e of this collec-
t i o n , i.e. m a k i n g a n i n v e n t o r y of the species i n h a b i t i n g the M w a n z a G u l f as a basis
f o r e c o l o g i c a l research to p r o v i d e d a t a for l a s t i n g fisheries o n h a p l o c h r o m i n e c i -
c h l i d s , u n f o r t u n a t e l y has been m a d e s u p e r f l u o u s b y the N i l e p e r c h . H o w e v e r , the
H E S T c o l l e c t i o n n o w f o r m s a u n i q u e r e c o r d of the f i s h f a u n a before, d u r i n g , a n d
after the N i l e p e r c h i n v a s i o n . T h e collection p r o b a b l y is b o t h the last a n d the m o s t
c o m p r e h e n s i v e of the o r i g i n a l f a u n a at a n y p a r t i c u l a r area of L a k e V i c t o r i a . T h e m a -
jority of the species i n this collection is still u n d e s c r i b e d . T h i s p a p e r deals w i t h the
d e s c r i p t i o n of six n e w species of p i s c i v o r o u s h a p l o c h r o m i n e c i c h l i d s a n d the redes-
c r i p t i o n of t w o p i s c i v o r o u s species. Six of these species h a v e m o d e r a t e l y to s t r o n g l y
c u r v e d teeth. A l l species, except one, are exceptional a m o n g s t the L a k e V i c t o r i a h a p -
l o c h r o m i n e p i s c i v o r e s i n h a v i n g p o l y c h r o m a t i c females. U s u a l l y , the females of h a -
V A N OIJEN: PISCIVOROUS H A P L O C H R O M I N E CICHLIDAE O F L A K E VICTORIA 7
p l o c h r o m i n e c i c h l i d s h a v e a n i n c o n s p i c u o u s c o l o r a t i o n a n d o n l y t h e m a l e s are
brightly coloured.
To f i n d o u t w h i c h genus c o u l d accomodate the n e w species, it w a s necessary to
analyse the d e s c r i p t i o n s of Harpagochromis a n d Prognathochromis b o t h i n t r o d u c e d b y
G r e e n w o o d (1980).
Material, methods a n d techniques
T h e major part of the m a t e r i a l d e s c r i b e d b e l o w w a s collected f r o m t r a w l catches

m a d e o v e r m u d b o t t o m s i n the M w a n z a G u l f of L a k e V i c t o r i a . F o r p r o c e d u r e s of
p r e s e r v a t i o n , l a b e l l i n g a n d c o l o u r p h o t o g r a p h y see v a n O i j e n et a l . (1981). Besides
c o l o u r slides of l i v e fishes, p h o t o g r a p h s w e r e also m a d e of s p e c i m e n s w h i c h w e r e
p r e s e r v e d o n m e l t i n g ice f o r some time. T h e c o l o r a t i o n of these specimens i n t e n s i f i e d
rather t h a n decreased d u r i n g p r e s e r v a t i o n . I n a d d i t i o n to c o l o u r p h o t o g r a p h y , notes
o n the c o l o r a t i o n of specimens w e r e m a d e r i g h t after their capture.
T e r m i n o l o g y a n d m e a s u r e m e n t s f o l l o w the d e f i n i t i o n s of B a r e l et a l . , (1976),
G r e e n w o o d (1980), H o o g e r h o u d (1984), H o o g e r h o u d & W i t t e (1981), W i t t e (1984b)
Fig. 1. Measuring points for Premaxillary Pedicel Length (PPL) and Upper Jaw Length (UJL). Fig. 2.
Examples of different lower jaw side obliqueness. A, almost flat, B, moderately oblique, C , strongly
oblique.
a n d W i t t e & W i t t e - M a a s (1981). T w o n e w measurements a n d a q u a l i t a t i v e character

p r o v e d to be u s e f u l i n d i s t i n g u i s h i n g p i s c i v o r o u s species a n d are d e s c r i b e d below.
T h e m e a s u r e m e n t s s h o u l d be taken f r o m intact specimens w i t h the m o u t h closed or
slightly opened.
— U p p e r J a w L e n g t h ( U J L , f i g . 1) is d e f i n e d as the distance b e t w e e n the rostral-
m o s t p o i n t of the p r e m a x i l l a r y s y m p h y s i s a n d the c a u d o - v e n t r a l t i p of the m a x i l l a
(the s h a n k process of Barel et a l . , 1976).
— P r e m a x i l l a r y P e d i c e l L e n g t h ( P P L , f i g . 1) is d e f i n e d as the distance b e t w e e n
the r o s t r a l m o s t p o i n t of the p r e m a x i l l a r y s y m p h y s i s a n d the d o r s a l t i p of the pre-
m a x i l l a r y p e d i c e l . (This is i n fact the external m e a s u r e m e n t of the P r e m a x i l l a r y A s -
c e n d i n g A r m L e n g t h of H o o g e r h o u d (1984) a n d W i t t e (1984b)).
— T h e obliqueness of the l o w e r j a w sides is d e f i n e d i n a s p e c i m e n i n a rostral
v i e w , as the angle b e t w e e n a p l a n e t h r o u g h the lateral side of the l o w e r j a w a n d the
m e d i a l p l a n e (fig. 2).
S p e c i m e n s w e r e m e a s u r e d u s i n g a p a i r of calipers. M e a s u r e m e n t s i n the h e a d
r e g i o n w e r e o f t e n m a d e u n d e r a W I L D M 5 b i n o c u l a r m i c r o s c o p e , w h i c h w a s also
u s e d for the e x a m i n a t i o n of the d e n t i t i o n , the s q u a m a t i o n , a n d the skeletal elements.
D r a w i n g s of skeletal elements w e r e m a d e u s i n g the s a m e m i c r o s c o p e m o u n t e d
w i t h a camera lucida.
A l l t y p e s p e c i m e n s of p i s c i v o r o u s h a p l o c h r o m i n e c i c h l i d s f r o m L a k e V i c t o r i a
f r o m the collections of the A c a d e m y of N a t u r a l Sciences of P h i l a d e l p h i a ( A N S P ) ; the
N a t u r a l H i s t o r y M u s e u m ( B M N H ) , L o n d o n ; the M u s e u m of C o m p a r a t i v e Z o o l o g y
( M C Z ) , C a m b r i d g e ; the M u s e u m N a t i o n a l d ' H i s t o i r e N a t u r e l l e ( M N H N ) , P a r i s ; the
M u s e o C i v i c o d i Storia N a t u r a l e ( M S N G ) , G e n o v a ; the N a t u r h i s t o r i s c h e s M u s e u m
( N H W ) , V i e n n a , a n d the M u s e u m für N a t u r k u n d e der H u m b o l d t Universität ( Z M B ) ,
B e r l i n w e r e e x a m i n e d . O n l y the h o l o t y p e of Haplochromis serranus (Pfeffer, 1896)
c o u l d not be located.
D e f i n i t i o n of terms. W h e n e v e r a character is c a l l e d g e n e r a l i z e d it is s i m i l a r to
that of H. elegans T r e w a v a s , 1933, a m o r p h o l o g i c a l l y m o d a l h a p l o c h r o m i n e c i c h l i d
( A n k e r , 1978; Barel et a l . , 1976). W h e n an element is c a l l e d r e l a t i v e l y s m a l l or rela-
t i v e l y large it is w i t h reference to a m o d a l p i s c i v o r o u s species. " P i s c i v o r e s " here is
u s e d as a collective n a m e for a l l h a p l o c h r o m i n e c i c h l i d s for w h i c h fishes g e n e r a l l y
are the d o m i n a n t f o o d , e x c l u d i n g the peadophages. P i s c i v o r e s h a v e a d u l t sizes be-
t w e e n 70 a n d 280 m m S L . F o r the sake of convenience a n u m b e r of size g r o u p s are
d i s t i n g u i s h e d ; s m a l l (70-110 m m ) , m e d i u m (110-150 m m ) m o d e r a t e l y large (150-190
m m ) , a n d large (> 190 m m ) .
T h e r e l a t i v e l y l a r g e s i z e of p i s c i v o r e s m a k e s it d i f f i c u l t to c o m p a r e b e t w e e n
species characters w h i c h are a l l o m e t r i c . P r o p o r t i o n a l m e a s u r e m e n t s w h i c h s h o w
a l l o m e t r y can o n l y effectively be c o m p a r e d if the a l l o m e t r i c ratio of the characters
are k n o w n , o r b e t w e e n specimens d i f f e r i n g little i n s t a n d a r d l e n g t h . To facilitate the
c o m p a r i s o n , the specimens u s e d for the descriptions w e r e d i v i d e d i n t o classes of 10
m m . If t w o successive classes h a d o n l y f e w representatives they w e r e u n i t e d to f o r m
o n e class of 20 m m . I n o n e case, three classes w e r e u n i t e d to f o r m o n e g r o u p of 66.2-
80.5 m m .
In the d r a w i n g s p r e m a x i l l a r y teeth are n u m b e r e d f r o m m e d i o - r o s t r a l to c a u d a l ,
l o w e r p h a r h y n g e a l teeth are n u m b e r e d f r o m m e d i a l to lateral.
Taxonomy
S u p r a g e n e r i c classification of the L a k e V i c t o r i a h a p l o c h r o m i n e C i c h l i d a e
F o l l o w i n g G r e e n w o o d (1974: 17) the H E S T m e m b e r s h a v e u s e d the t e r m ' h a p -
l o c h r o m i n e c i c h l i d s ' a n d ' h a p l o c h r o m i n e s ' , to refer to a l l species o f Haplochromis H i l -
gendorf, 1888 a n d the closely r e s e m b l i n g m o n o t y p i c genera (i.e. Macropleurodus B o u -
lenger, 1906, Hoplotilapia H i l g e n d o r f , 1888, Platytaeniodus Boulenger, 1906, Paralabi-
dochromis G r e e n w o o d , 1956 a n d Astatoreochromis P e l l e g r i n , 1904).
Recently, Eccles & Trewaves (1989: 21) d e f i n e d the tribe H a p l o c h r o m i n i as 'mater-
n a l m o u t h - b r o o d i n g c i c h l i d s of A f r i c a a n d the J o r d a n V a l l e y i n w h i c h the b a s i o c c i p i -
tal b o n e participates w i t h the p a r a s p h e n o i d to f o r m the a p o p h y s i s f o r the u p p e r p h a -
r y n g e a l bones'. F o l l o w i n g this d e f i n i t i o n a l l h a p l o c h r o m i n e c i c h l i d s fishes of L a k e
V i c t o r i a are H a p l o c h r o m i n i .
T h e g e n e r i c c l a s s i f i c a t i o n of the L a k e V i c t o r i a h a p l o c h r o m i n e C i c h l i d a e
G r e e n w o o d ' s r e v i s i o n of the genus Haplochromis: i n t r o d u c t i o n

G r e e n w o o d (1979), u s i n g a cladistic a p p r o a c h , attempted to break u p the genus
Haplochromis i n t o a n u m b e r of m o n o p h y l e t i c l i n e a g e s . A c c o r d i n g to G r e e n w o o d
(1979), the h i g h n u m b e r o f species (c. 390), the w i d e g e o g r a p h i c a l d i s t r i b u t i o n a n d
the w i d e range of a n a t o m i c a l a n d m o r p h o l o g i c a l v a r i a t i o n of Haplochromis i n d i c a t e d
that the genus w a s ' v e r y i l l - d e f i n e d ' . Strictly a d h e r i n g to H i l g e n d o r f 's ' d e f i n i t i o n ' of
the t o o t h s h a p e , G r e e n w o o d restricted the g e n u s Haplochromis to f i v e (lacustrine)
species. F o r reasons o u t l i n e d b e l o w , this d e f i n i t i o n of the genus Haplochromis w i l l n o t
be u s e d i n this paper. I n the second part of h i s r e v i s i o n G r e e n w o o d (1980) p l a c e d the
h a p l o c h r o m i n e species f r o m lakes V i c t o r i a , N a b u g a b o , E d w a r d , G e o r g e a n d K i v u i n
20 genera. In the i n t r o d u c t i o n to this p a p e r G r e e n w o o d (1980: 3) stated ' A t present,
the f o r m e r genus Haplochromis c a n be r e s o l v e d i n t o a n u m b e r of a p p a r e n t l y m o n o -
p h y l e t i c lineages; the search f o r characters u n i t i n g these lineages t h r o u g h v a r i o u s
levels o f c o m m o n ancestry m u s t c o n t i n u e ' . F u r t h e r h e a d m i t t e d that h e d i d n o t suc-
ceed i n c o n s t r u c t i n g d i c h o t o m o u s l y b r a n c h i n g p h y l o g e n i e s at either inter- o r i n t r a -
lineage levels, a n d suggested that m o r e d i s c r i m i n a t i n g a n a t o m i c a l studies a n d b i o -
c h e m i c a l t e c h n i q u e s m i g h t r e v e a l l i n k a g e s so far u n d i s c o v e r e d . I n the c o n c l u s i o n
G r e e n w o o d (1980: 95), q u o t i n g R e g a n (1920: 160), stated that he d i d n o t r e g a r d h i s
n e w classification as entirely satisfactoriy. G r e e n w o o d ' s r e v i s i o n has been c h a l l e n g e d
b y m a n y researchers w o r k i n g at different levels o f b i o l o g i c a l o r g a n i s a t i o n . A s u m -
m e r y of this critique is presented b e l o w .
A n a l y s e s o n the m o l e c u l a r l e v e l
Sage et a l . (1984), m e a s u r i n g the extent of s t r u c t u r a l gene d i v e r g e n c e b y means of
starch g e l electrophoresis of m u s c l e a n d eye proteins, f o u n d extremely s m a l l genetic
differences a m o n g t e n h a p l o c h r o m i n e species f r o m l a k e V i c t o r i a ( b e l o n g i n g to s i x
genera a c c o r d i n g to G r e e n w o o d , 1980). T h e y f o u n d it barely possible to d i s t i n g u i s h
Hoplotilapia f r o m Haplochromis. H o w e v e r , there w a s a clear difference b e t w e e n these
t w o genera a n d Astatoreochromis. A c c o r d i n g to Sage et al. (1984) the s m a l l genetic dif-
ferences are consistent w i t h the p o s s i b i l i t y that the h a p l o c h r o m i n e species flock arose

f r o m a s i n g l e ancestral species after L a k e V i c t o r i a w a s f o r m e d a n d i n d i c a t e recent
speciation. Verheijen et a l . (1985,1986,1988), s t u d y i n g a l l o z y m e v a r i a t i o n b y a n elec-
t r o p h o r e t i c a n a l y s i s of m u s c l e e n z y m e s , e l e c t r o p h o r e t i c m o b i l i t y of h a e m o g l o b i n
b a n d s a n d iso-electric f o c u s e d eye-lens p r o t e i n patterns, d i d n o t f i n d a n y differences
a m o n g e i g h t Haplochromis species ( b e l o n g i n g to at least f o u r g e n e r a after G r e e n -
w o o d , 1980), a n d the m o n o t y p i c genera Hoplotilapia a n d Macropleurodus. T h e y s u g -
gested that speciation does n o t h a v e to i n v o l v e electrophoretically detectable differ-
e n t i a t i o n , a n d a d d e d that the genetic s i m i l a r i t y of these species p o i n t s to a v e r y
recent speciation. Verheijen & V a n R o m p a e y (1986) a n d V a n R o m p a e y et a l . (1988)
d e m o n s t r a t e d f o r s o m e c i c h l i d s f r o m L a k e V i c t o r i a a correlation b e t w e e n the general
electrophoretic pattern o f m u s c l e proteins a n d ecological characteristics. T h e y con-
c l u d e d that i n h a p l o c h r o m i n e c i c h l i d s f r o m L a k e V i c t o r i a t r o p h i c differentiation m a y
h a v e o c c u r r e d after habitat (or substrate) selection. Verheijen (1989) stated that the
t a x o n o m i e , ecological a n d m o r p h o l o g i c a l d i v e r s i t y i n L a k e V i c t o r i a h a p l o c h r o m i n e s
is n o t m a t c h e d b y a s i g n i f i c a n t l e v e l of e l e c t r o p h o r e t i c a l l y detectable change. H e
stressed that his results are i n accordance w i t h literature data s u g g e s t i n g that G r e e n -
w o o d ' s generic s u b d i v i s i o n of the 'genus' Haplochromis m a y be i n v a l i d .
D o r i t (1986), i n a s t u d y of m i t o c h o n d r i a l D N A of ten h a p l o c h r o m i n e species f r o m
G r e e n w o o d ' s Psammochromis-Macropleurodus s u p e r l i n e a g e ( b e l o n g i n g to 6 genera
after G r e e n w o o d , 1980) f o u n d that the trees generated b y u s i n g the presence or ab-
sence of p a r t i c u l a r restriction sites as indicators of h o m o l o g y , bear 'both s t r i k i n g s i m -
ilarities a n d interesting differences w i t h the c l a d o g r a m suggested b y G r e e n w o o d o n
m o r p h o l o g i c a l g r o u n d s '(Dorit, 1986: 69). T h e results of D o r i c s research i m p l y some
interesting p r o b l e m s ; it w a s possible for e x a m p l e to d i s t i n g u i s h Haplochromis " v e l v e t
b l a c k " specimens f r o m six localities i n the M w a n z a G u l f b u t it p r o v e d i m p o s s i b l e to
separate specimens o f Haplochromis sauvagei (Pfeffer, 1894) f r o m those of H. prodomus
T r e w a v a s , 1935.
Recently, M e y e r et a l . (1990) suggested a m o n o p h y l e t i c o r i g i n for the L a k e V i c -
t o r i a h a p l o c h r o m i n e c i c h l i d s o n the basis of research o n m i t o c h o n d r i a l D N A se-
quences. F o u r t e e n e n d e m i c h a p l o c h r o m i n e species s h o w e d a l m o s t n o d i f f e r e n t i a t i o n
at a b o u t 800 n u c l e o t i d e p o s i t i o n s i n the c y t o c h r o m e b gene a n d i n m i t o c h o n d r i a l
D N A that bears part of t w o transfer R N A genes a n d the n o r m a l l y h i g h l y v a r i a b l e
p o r t i o n o f the c o n t r o l r e g i o n . I n the e x a m i n e d s e q u e n c e , h a p l o c h r o m i n e c i c h l i d
species f r o m L a k e M a l a w i differed f r o m those i n L a k e V i c t o r i a b y 54-55 base substi-
tutions, whereas the L a k e V i c t o r i a h a p l o c h r o m i n e s d i f f e r e d a m o n g s t each other b y
a n average of o n l y three m i t o c h o n d r i a l m u t a t i o n s . M e y e r et a l . (1990) date the o r i g i n
of the extant L a k e V i c t o r i a h a p l o c h r o m i n e s at less t h a n 200.000 years ago a n d f i n d it
l i k e l y that the V i c t o r i a flock arose w i t h i n the lake. H o w e v e r , the last statement is pre-
m a t u r e as the authors d i d not yet e x a m i n e h a p l o c h r o m i n e c i c h l i d s f r o m L a k e s K i v u
and George.
I m p l i c a t i o n s of the revisions for t a x o n o m y

A t the a l p h a - t a x o n o m i c l e v e l , researchers w e r e most d i r e c t l y c o n f r o n t e d w i t h the
consequences of G r e e n w o o d ' s generic r e v i s i o n . V a r i o u s n e w h a p l o c h r o m i n e species
f r o m L a k e s K i v u a n d V i c t o r i a cannot be p l a c e d u n e q u i v o c a l l y i n o n e of G r e e n w o o d ' s
genera. A m o n g species f r o m L a k e K i v u : (1) Haplochromis murakaze C o e n e n et a l , 1984
c o m b i n e s characters of three genera; it is intermediate b e t w e e n Astatotilopia, Labro-

chromis a n d Gaurochromis; (2) Haplochromis scheffersi Snoeks et a l , 1987, is p a r t l y refer-
able to Astatotilapia b u t the ranges of certain of its characters a m p l y e x c e e d e d the
ranges of this genus as d e f i n e d b y G r e e n w o o d (1979); (3) Haplochromis occultidens
S n o e k s , 1988, agrees i n s o m e characters w i t h Lipochromis (Cleptochromis) b u t this
(sub)genus cannot really accommodate the s p e c i e s u n l e s s t h e g e n e r i c l i m i t s are
e x t e n d e d ; (4) Haplochromis olivaceus a n d H. crebidens Snoeks et a l . , 1990, b o t h e p i l i t h i c
algae scrapers, cannot be p l a c e d i n a n y of the genera of G r e e n w o o d (1980). S i m i l a r l y
Haplochromis erythromaculatus d e Vos, Snoeks & T h y s v a n d e n A u d e n a e r d e , 1990, a
species o c c u r r i n g i n L a k e s B u l e r o a n d R u h o n d o cannot be p l a c e d i n a n y of these
genera (de Vos et a l . , 1990).
H o o g e r h o u d (1984), w o r k i n g o n m o l l u s c i v o r o u s c i c h l i d s , e n c o u n t e r e d s i m i l a r
p r o b l e m s w i t h species f r o m L a k e V i c t o r i a : Haplochromis iris H o o g e r h o u d & W i t t e ,
1981 a n d H. hiatus H o o g e r h o u d & Witte, 1981, b r i d g e the g a p b e t w e e n Gaurochromis
(Mylacochromis) G r e e n w o o d , 1980, a n d Gaurochromis (Gaurochromis) G r e e n w o o d ,
1980. R e e x a m i n a t i o n o f G r e e n w o o d ' s o r i g i n a l m a t e r i a l s h o w e d that a c c e p t i n g the
d e f i n i t i o n o f Gaurochromis w o u l d m e a n t h a t s p e c i m e n s o f Haplochromis ptistes
G r e e n w o o d & Barel, 1978 a n d H. iris h a v e to be d i v i d e d o v e r Labrochromis R e g a n ,
1920 a n d Gaurochromis, a n d that Haplochromis humilior (Boulenger, 1911) is i n fact
i n t e r m e d i a t e b e t w e e n the genera Gaurochromis a n d Enterochromis G r e e n w o o d , 1980.
W i t t e & W i t t e - M a a s (1987), d e s c r i b i n g n e w s p e c i e s o f z o o p l a n k t i v o r o u s h a p -
l o c h r o m i n e s f r o m L a k e V i c t o r i a , p r o v e d that s o m e of these species b r i d g e the g a p
b e t w e e n Yssichromis G r e e n w o o d , 1980 a n d Astatotilapia P e l l e g r i n , 1903. T h e y s h o w e d
that several characters u s e d b y G r e e n w o o d (1980) to separate the genera are i n v a l i -
d a t e d b y G r e e n w o o d & G e e (1969). W i t t e & W i t t e - M a a s (op. cit.) also suggested that
the g a p b e t w e e n Yssichromis a n d Psammochromis is b r i d g e d b y certain u n d e s c r i b e d
species.
Because they realised that generic b o u n d a r i e s m i g h t h a v e to be a d a p t e d after the
d e s c r i p t i o n of almost e v e r y n e w species the above m e n t i o n e d authors d e c i d e d n o t to
accept G r e e n w o o d ' s genera a n d to use the genus Haplochromis i n the sense i t w a s
u s e d before the generic r e v i s i o n of G r e e n w o o d .
G r e e n w o o d ' s generic classification of the p i s c i v o r o u s species: i n t r o d u c t i o n

G r e e n w o o d (1980) p l a c e d the majority of the p i s c i v o r o u s h a p l o c h r o m i n e c i c h l i d s
i n t w o g e n e r a , v i z . Harpagochromis G r e e n w o o d , 1 9 8 0 a n d Prognathochromis G r e e n -
w o o d , 1980, w h i c h , a c c o r d i n g to G r e e n w o o d , c a n b e d i s t i n g u i s h e d f r o m the other
c i c h l i d genera i n L a k e V i c t o r i a b y their relatively l o n g l o w e r jaw. G r e e n w o o d (1980)
considers the r e l a t i v e l y l o n g l o w e r j a w a n a p o m o r h p i c character. G r e e n w o o d gave as
ranges f o r the L o w e r J a w L e n g t h / H e a d L e n g t h r a t i o of Harpagochromis a n d Prog-
nathochromis 4 3 - 6 1 % a n d 41-62%, r e s p e c t i v e l y . H o w e v e r , s o m e species p l a c e d b y
G r e e n w o o d (1980) i n Psammochromis G r e e n w o o d , 1980, v i z . Haplochromis cassius
G r e e n w o o d & Barel, 1978, a n d H. acidens G r e e n w o o d , 1967 h a v e L J L / H L ratio's of
43-48% a n d 44-50 %, respectively. T h e L J L / H L ratio therefore cannot be u s e d to dis-
t i n g u i s h Harpagochromis a n d Prognathochromis f r o m Psammochromis. F o r s i m i l a r rea-
sons this m e a s u r e m e n t cannot be u s e d to d i s t i n g u i s h the p i s c i v o r o u s genera f r o m
Pixichromis G r e e n w o o d , 1980 a n d Lipochromis R e g a n , 1920. O t h e r characters w h i c h
a c c o r d i n g to G r e e n w o o d (1980) sepatate these genera f r o m the p i s c i v o r o u s genera
h a v e n o t yet been c r i t i c a l l y e x a m i n e d .
T h e p r i n c i p a l d i a g n o s t i c features G r e e n w o o d u s e d to differentiate Harpagochro-
mis f r o m Prognathochromis are f o u n d i n the n e u r o c r a n i u m . T h e n e u r o c r a n i u m of
Harpagochromis is of a g e n e r a l i z e d type, b u t w o u l d h a v e a s h a l l o w e r otic r e g i o n a n d a
h i g h e r s u p r a o c c i p i t a l crest, generally near p y r a m i d i c a l i n o u t l i n e . I n contrast, Prog-
nathochromis w o u l d possess a n elongate, slender a n d s h a l l o w n e u r o c r a n i u m w i t h a
l o w s u p r a o c c i p i t a l crest, w e d g e s h a p e d i n lateral o u t l i n e . To q u a n t i f y the differences
i n n e u r o c r a n i a l shape G r e e n w o o d (1980; 14, f i g . 1) d e f i n e d f o u r n e u r o c r a n i a l charac-
ters: v i z . N e u r o c r a n i a l W i d t h ( N W ) , O t i c D e p t h ( O t D ) , O r b i t a l D e p t h ( O D ) a n d
Preorbital Depth (PrOD).
Analysis
J u d g i n g f r o m d a t a i n species d e s c r i p t i o n s a n d i n e a r l i e r a t t e m p t s to u n r a v e l
p h y l e t i c l i n e s a m o n g s t the p i s c i v o r o u s species (e.g. f r o m G r e e n w o o d , 1962, 1967,
1974; G r e e n w o o d & G e e , 1969), the a l l o c a t i o n of v a r i o u s species to the genera of
G r e e n w o o d (1980) is at places inconsistent:
I. — Haplochromis decticostoma G r e e n w o o d & G e e , 1969, is p l a c e d b y G r e e n w o o d
(1980) i n Prognathochromis w i t h o u t a n y c o m m e n t e x c e p t 'see G r e e n w o o d & G e e
(1969: 55-7)'. O n the i n d i c a t e d page o n e reads: T h e s y n c r a n i u m of H. decticostoma is
v e r y s i m i l a r to that of H. spekii (Boulenger, 1906) i n most details' a n d : T h y l e t i c a l l y ,
H. decticostoma is a d e r i v a t i v e of the H. serranus stem, a n d p r o b a b l y f r o m a species
a n a t o m i c a l l y i n d i s t i n g u i s h a b l e f r o m H. spekii'. H o w e v e r , G r e e n w o o d (1980) places
Haplochromis serranus a n d H. spekii i n Harpagochromis.
II. — Haplochromis altigenis R e g a n , 1922, is p l a c e d b y G r e e n w o o d (1980) i n Harpa-
gochromis w i t h a reference to the r e d e s c r i p t i o n of the species i n G r e e n w o o d (1967: 60-
65). H o w e v e r , o n the i n d i c a t e d p a g e the a u t h o r states that: ' T h e n e u r o c r a n i u m of
Haplochromis altigenis resembles that of H. bayoni b u t is r e l a t i v e l y b r o a d e r i n the otic
r e g i o n . T h u s a l t h o u g h it s h o w s s o m e of the characters associated w i t h the "mento" -
t y p e it still retains the c u r v e d p r e o r b i t a l p r o f i l e , greater p r e o r b i t a l d e p t h a n d b r o a d
otic r e g i o n of the m o r e generalised n e u r o c r a n i u m . I n these characters it also resem-
b l e s H. pseudopellegrini', B o t h H. bayoni ( B o u l e n g e r , 1909)and H. pseudopellegrini
G r e e n w o o d , 1967 are allocated to Prognathochromis b y G r e e n w o o d (1980).
III. — Haplochromis xenostoma R e g a n , 1922, is p l a c e d b y G r e e n w o o d (1980) i n
Prognathochromis. I n the r e d e s c r i p t i o n of this species G r e e n w o o d (1967: 53) states:
T h e n e u r o c r a n i u m of H. xenostoma is s i m i l a r to that of H. victorianus b u t has a longer
p r e o r b i t a l face..; the n e u r o c r a n i o u m of the t w o species also differ i n that the supraoc-
c i p i t a l crest is relatively h i g h e r a n d m o r e p o i n t e d than i n H. victorianus'. A n d o n p .
55: 'the s k u l l of H. xenostoma is clearly of the "serranus"-type'. B o t h Haplochromis ser-
ranus a n d H. victorianus ( P e l l e g r i n , 1904) w e r e r e a l l o c a t e d to Harpagochromis b y
G r e e n w o o d (1980).
IV. — Haplochromis melichrous G r e e n w o o d & G e e , 1969, is p l a c e d b y G r e e n w o o d
(1980) i n the s u b g e n u s Prognathochromis (Tridentochromis). I n the d e s c r i p t i o n of H.
melichrous, G r e e n w o o d & G e e (1969, 26) state: ' T h e n e u r o c r a n i u m of H. melichrous
resembles that of H. victorianus: i n other w o r d s it is referable to the "serranus" g r o u p ' .
I n G r e e n w o o d (1974), i n the s e c t i o n o f species f e e d i n g o n b e n t h i c c r u s t á c e a , t h e
a u t h o r s t i l l w r i t e s 'Haplochromis melichrous d i f f e r s f r o m m e m b e r s o f t h e 'tridens'
g r o u p i n its n e u r o c r a n i a l m o r p h o l o g y , a n d i n these characters resembles m e m b e r s of
the "serranus" g r o u p p i s c i v o r e s ' . H o w e v e r , i n G r e e n w o o d (1980) Haplochromis ser-

ranus a n d H. victorianus are p l a c e d i n Harpagochromis.
V . — I n the d e s c r i p t i o n o f Haplochromis paraquiarti G r e e n w o o d (1967; 70) states:
' T h e n e u r o c r a n i u m of H. paraquiarti is i d e n t i c a l w i t h that of H. acidens. It differs
s o m e w h a t f r o m the p r e s u m e d generalised p i s c i v o r o u s s k u l l of H. quiarti a n d s h o w s
s o m e of the characters f o u n d i n the m o r e specialised type of H. prognathus. It i s , i n
fact almost i n t e r m e d i a t e b e t w e e n the t w o types.' I n 1980 Haplochromis paraquiarti a n d
H. prognathus are p l a c e d i n Prognathochromis, H. acidens G r e e n w o o d , 1967 is p l a c e d i n
Psammochromis a n d H. quiarti (Pellegrin, 1904)is p l a c e d i n Harpagochromis. T h e inter-
m e d i a t e p o s i t i o n of the n e u r o c r a n i a of Haplochromis paraquiarti a n d H. acidens is n o t
m e n t i o n e d a g a i n after 1967.
T h e e x a m p l e s I - V g i v e n above m a y i n d i c a t e that the d e l i m i t a t i o n b e t w e e n the
alleged taxa is n o t as sharp as o n e m i g h t expect f o r genera. T h e f o r e g o i n g inconsis-
Figs 3-6. Lateral, dorsal and ventral view of neurocrania of picivorous haplochromine species. Fig. 3.
Haplochromis bareli spec. nov. Fig. 4. H. dichrourus "Uganda". Fig. 5. H. dichrourus "Mwanza". Fig. 6. H.
pyrrhopteryx spec. nov. Scale equals 10 mm.
tensies m i g h t be e x p l a i n e d b y the fact that G r e e n w o o d ' s statements about s i m i l a r i t y

of n e u r o c r a n i a a n d the notes c o n c e r n i n g similarities i n p u b l i c a t i o n s before 1980 w e r e
based o n v i s u a l i m p r e s s i o n s o n l y Except for the p r e o r b i t a l face (in G r e e n w o o d , 1967:
50) G r e e n w o o d never d e f i n e d the characters he based his c o n c l u s i o n s o n i n terms of
m o r p h o m e t r i c measurements. E v e n i n G r e e n w o o d (1980), the shape of the supraoc-
c i p i t a l crest, a major character for separating Harpagochromis a n d Prognathochromis, i s
not d e f i n e d i n this w a y .
C l a s s i f y i n g n e w species
A t t e m p t i n g to classify p i s c i v o r o u s species f r o m the M w a n z a G u l f i n the genera
of G r e e n w o o d (1980) I m e a s u r e d the n e u r o c r a n i a (see figs. 3-6) u s i n g the d e f i n i t i o n s
of G r e e n w o o d (1980). It t u r n e d out to be i m p o s s i b l e to place s o m e n e w species o n
the basis of these m e a s u r e m e n t s i n either Harpagochromis o r Prognathochromis. F o r
instance i n Haplochromis bareli spec. nov. half of the range of S k u l l W i d t h lies i n the
r a n g e of Prognathocromis, the other half i n the range of Harpagochromis; the O t i c d e p t h
range entirely covers the range of Harpagochromis, b u t extends to part of the range of
Prognathochromis; O r b i t a l D e p t h lies i n the o v e r l a p p i n g area of Harpagochromis a n d
Prognatochromis, a n d P r e o r b i t a l D e p t h p a r l y falls i n the range of Prognathochromis
(fig. 7).
To e x c l u d e the p o s s i b i l i t y that the results o f m y m e a s u r i n g technique differed
f r o m those of G r e e n w o o d , I h a d t o c o m p a r e m y m e a s u r e m e n t s w i t h t h o s e o f
G r e e n w o o d . H o w e v e r , as G r e e n w o o d ' s (1980) p a p e r does not c o n t a i n a table w i t h
m e a s u r e m e n t s of i n d i v i d u a l s p e c i m e n s , the o n l y w a y to r e v e a l differences w a s to
c o m p a r e t h e t o t a l r a n g e s . G r e e n w o o d (1980) s t a t e d t h a t a l l t h e B M ( N H ) h a p -
l o c h r o m i n e m a t e r i a l f r o m the five lakes w e r e i n v o l v e d i n his r e v i s i o n . U s i n g the d e f i -
n i t i o n s of G r e e n w o o d (1980) I m e a s u r e d the 66 n e u r o c r a n i a f r o m the B M N H collec-
tions b e l o n g i n g to 44 p i s c i v o r o u s species (table 1). A l l m e a s u r e m e n t s w e r e t a k e n
w i t h T E S A D I G I T - C A L I I d i g i t a l calipers u n d e r a b i n o c u l a r microscope.
Table 1. Neurocranial mesurements of species placed in Harpagochromis, Prognathochromis and Psam-

mochromis by Greenwood (1980). All specimens from the B M N H collections. Measurements in mm..
Gen.: genus according to Greenwood (1980); Ps. = Psammochromis; P. = Prognathochromis; Ha. -
Harpagochromis; N L = Neurocranial Length; N W = Neurocranial Width; PrOD = Preorbital depth; O D
= Orbital Depth; OtD = Otic Depth; - = Damage on neurocranium makes accurate measurement
impossible.
NW PrOD OD OtD
No. Gen. Species NL NW NL PrOD N L OD NL OtD NL
1. Ps. acidens 31.9 16.8 50.7 7.7 24.1 10.6 33.2 14.0 43.8
2. Ha. altigenis 44.6 23.8 53.3 9.2 20.6 12.7 28.4 17.9 40.1
3. P. arcanus 31.6 14.3 45.2 5.8 28.1 8.9 28.1 13.2 41.7
4. P. argenteus 49.5 22.7 45.8 7.0 14.1 12.3 24.8 17.7 35.7
5. Ha. artaxerxes 19.3 10.0 51.8 4.3 22.2 6.0 31.0 8.3 43.0
6. P. bartoni 36.9 - 7.0 18.9 10.0 27.9 15.1 40.9
7. P. bayoni 37.1 19.4 52.2 5.9 15.9 9.1 24.5 14.3 38.5
8. Ha. cavifrons 32.6 17.9 54.9 5.8 17.7 8.3 254 12.9 39.5
" 40.2 - 7.3 18.1 9.5 23.6 17.0 42.2
9. P. crocopeplus 24.9 13.0 52.2 4.1 16.4 7.0 28.1 10.2 40.9
25.8 - 4.1 15.8 7.1 27.5 11.1 43.0
10. P. cryptogramma 26.4 12.0 45.4 5.0 18.9 7.1 26.8 11.1 42.0
11. P. decticostoma 52.3 28.6 54.6 10.1 19.1 14.1 26.9 21.5 41.1
12. P. dent ex 37.1 18.1 48.7 5.0 13.4 9.0 24.2 13.5 36.3
13. Ρ dichrourus 31.3 15.6 49.8 6.0 19.1 9.1 29.0 12.1 38.6
14 P. dolychorhynchus 30.2 14.3 48.0 4.6 15.2 8.3 27.4 12.2 40.3
15. P. estor 43.0 20.7 48.1 7.1 16.5 11.1 25.8 16.1 37.4
16. P. flavipinnis 30.6 16.9 55.2 5.9 19.2 8.9 29.0 13.2 43.1
17. P. gilberti 29.7 14.7 49.4 5.1 17.1 7.9 26.5 11.8 39.7
18. P. gowersi 40.4 17.8 44.0 6.7 16.5 11.9 29.4 14.8 36.6
19. Ha. guiarti 34.4 18.3 53.2 5.8 16.8 9.4 27.3 13.9 40.4
35.0 20.7 59.1 8.6 24.5 10.9 31.1 14.6 41.7
37.9 23.6 62.6 8.1 21.3 11.1 29.2 16.3 43.0
20. P. longirostris 30.9 14.0 45.3 4.1 13.2 7.4 23.9 12.0 38.8
34.9 18.0 51.5 5.7 16.3 8.3 23.7 12.3 35.2
35.7 16.6 46.6 5.9 16.5 8.9 24.9 13.0 36.4
21. P. macrognathus 44.2 20.6 46.6 5.5 12.4 9.3 21.0 17.2 38.9
43.1 19.0 44.0 6.3 14.6 10.1 23.4 15.0 34.8
22. Ha. maculipinna 38.8 22.1 56.9 8.4 21.6 11.9 30.6 17.0 43.8
23. P. mandibularis 34.9 15.6 44.6 4.8 13.7 8.4 24.0 13.1 37.5
24. P. mento 37.5 17.4 46.4 6.2 16.5 9.9 26.4 14.0 37.3
38.4 17.2 44.7 6.5 16.9 9.9 25.7 13.7 35.6
41.0 18.4 44.8 7.8 19.0 10.8 26.3 15.0 36.5
25. P. melichrous 25.5 14.8 58.0 4.9 19.2 7.2 28.2 11.2 43.9
26. Ha. michaeli 28.0 15.9 56.7 - - 10.1 36.0 12.2 43.5
35.9 19.6 54.5 6.5 18.1 9.1 25.3 14.9 41.5
27. P. nanoserranus 22.8 11.6 50.8 3.7 16.2 6.5 28.5 8.9 39.0
28. P. paraguiarti 30.2 15.7 51.9 5.8 19.2 8.8 29.1 13.1 43.3
29. P. pellegrini 21.0 10.7 50.9 3.1 14.7 5.4 25.7 8.0 38.3
30. Ha. plagiostoma 30.4 17.7 56.5 7.0 23.0 9.0 29.6 13.0 42.7
29.3 17.3 59.0 6.3 21.5 8.3 28.3 12.8 43.6
28.8 17.5 60.7 6.0 20.8 7.9 27.4 12.1 42.0
32.3 19.4 60.0 7.6 23.5 10.9 33.7 13.9 43.0
31.7 18.9 59.6 5.8 18.2 8.3 26.1 12.6 39.7
31. P. prognathus 23.2 11.1 47.8 4.5 19.3 6.8 29.3 9.3 40.0
33.8 16.1 47.6 5.2 15.3 9.5 28.1 13.6 40.2
34.5 16.6 48.1 6.2 17.9 10.0 28.9 13.8 40.0
36.2 - - 5.5 15.1 9.5 26.2 14.8 40.8
32. P. pseudopellegrini 31.9 - - 5.5 17.2 8.2 25.1 12.8 40.1
33. Ha. serranus 26.6 14.8 55.6 6.1 22.9 8.3 31.2 11.2 42.1
27.5 14.7 53.4 6.5 23.6 9.0 32.7 11.8 42.8
27.4 15.1 55.1 6.0 21.8 8.9 32.4 12.0 43.7
43.8 - - 8.8 20.0 11.7 26.7 18.0 41.0
34. Ha. spekii 36.9 20.1 54.4 7.8 21.1 11.4 30.8 16.0 43.3
38.2 20.3 53.1 6.7 17.5 10.9 28.5 16.8 43.9
35. Ha. squamipinnis 44.0 23.4 53.1 8.5 19.3 10.9 24.7 18.1 41.1
36. Ha. squamulatus 34.8 19.3 55.4 7.8 22.4 10.8 31.0 14.7 42.2
39.5 22.2 56.2 8.9 22.5 11.8 29.8 18.1 45.8
37. P. sulphurius 25.3 12.6 49.8 4.2 16.6 6.9 27.2 10.5 41.5
27.0 13.5 50.0 4.8 17.7 7.5 27.7 10.5 38.8
38. P. tridens 24.6 12.8 52.0 4.2 17.0 6.7 30.8 10.2 41.4
39. P. tyrianthinus 27.5 14.2 51.6 4.7 17.0 7.3 26.5 10.3 37.4
41. P. venator 35.1 17.1 48.7 5.8 16.5 8.9 25.3 13.3 37.8
42. Ha. victorianus 38.6 20.6 53.3 8.8 22.7 11.6 30.0 15.9 41.1
43. P. vittatus 23.4 11.8 50.4 4.2 17.9 6.7 28.6 8.9 38.0
44. P. xenostoma 45.5 22.4 49.2 7.6 16.7 11.3 24.8 18.8 41.3
Results a n d c o n c l u s i o n s
T h e r e are several discrepancies b e t w e e n the ranges g i v e n b y G r e e n w o o d (1980)
a n d the r a n g e s c a l c u l a t e d f r o m m y m e a s u r e m e n t s . I n fact o n l y o n e r a n g e (the
N W / N L r a t i o f o r Prognathochromis) is m o r e or less e q u a l (fig. 7). T h e differences
c o u l d be p a r t l y d u e to the fact that G r e e n w o o d ' s n e u r o c r a n i a l characters are d i f f i c u l t
to m e a s u r e accurately: O n l y the m e a s u r i n g p o i n t s of N e u r o c r a n i a l L e n g t h a n d S k u l l
W i d t h are (more o r less) f i x e d . H o w e v e r , O t i c D e p t h , w h i c h is m e a s u r e d between
p o i n t s o n c o n v e x surfaces, a n d especially P r e o r b i t a l D e p t h a n d O r b i t a l d e p t h , w h i c h
are m e a s u r e d i n the v e r t i c a l p l a n e f r o m i l l - d e f i n e d p o i n t s o n the n e u r o c r a n i u m to 'a
h o r i z o n t a l l i n e e x t e n d e d f r o m the v e n t r a l face of the p a r a s p h e n o i d ', are b o u n d to
g i v e results w i t h a r e l a t i v e l y large v a r i a t i o n .
I n o r d e r to s h o w that there is not just o v e r l a p of the extremes of the ranges of the
n e u r o c r a n i a l m e a s u r e m e n t s of the t w o genera, the m e a s u r e m e n t s of the i n d i v i d u a l
species are presented i n f i g . 8. T h i s figure clearly s h o w that there is i n d e e d a con-
s i d e r a b l e o v e r l a p i n the n e u r o c r a n i a l measurements w h i c h are s u p p o s e d to separate
Harpagochromis a n d Prognathochromis. T h e s e characters s e e m to f o r m c o n t i n u o u s
m o r p h o c l i n e s i n w h i c h a n y d i v i s i o n is arbitrary. Therefore, it is i m p o s s i b l e to m a i n -
t a i n the g e n e r a Harpagochromis a n d Prognathochromis as d e f i n e d b y G r e e n w o o d
(1980). G r e e n w o o d (1984:149), w r i t i n g about m o r p h o c l i n e s i n the p i s c i v o r o u s h a p l o -
c h r o m i n e s , states: O n e can trace changes i n relative s k u l l p r o p o r t i o n s , l e a d i n g b y d i f -
ferential g r o w t h of p a r t i c u l a r s k u l l regions, f r o m the g e n e r a l i z e d shape to one w h i c h
is elongate a n d s h a l l o w '. A s G r e e n w o o d (1980; 10) d e f i n e d the Harpagochromis n e u -
r o c r a n i u m as 'essentially of the generalised t y p e ' a n d the n e u r o c r a n i u m of Progna-
thochromis w a s s a i d to be 'elongate, slender a n d s h a l l o w ' ( G r e e n w o o d , 1980; 14) this
statement is not a p p l i c a b l e to o n l y one of the t w o genera. T h i s seems to i m p l y that
G r e e n w o o d (1984) realised that there is no clear m o r p h o l o g i c a l g a p b e t w e e n the n e u -
r o c r a n i a of the t w o genera.
T h e m a j o r i t y of p i s c i v o r o u s h a p l o c h r o m i n e c i c h l i d s are i m m e d i a t e l y r e c o g n i z -
able b y a r e l a t i v e l y large a n d slender body, a r e l a t i v e l y s m a l l eye, a deep cheek, a n d a
r e l a t i v e l y l o n g l o w e r j a w ( G r e e n w o o d , 1974; W i t t e & v a n O i j e n , 1990). H o w e v e r ,
m o r p h o l o g i c a l v a r i a t i o n a m o n g this most speciose t r o p h i c g r o u p of L a k e V i c t o r i a
h a p l o c h r o m i n e s is large ( G r e e n w o o d , 1974; v a n O i j e n , 1982): There are deep b o d i e d
species l i k e H. victorianus a n d H. bareli, s m a l l s i z e d species l i k e H. perrieri ( P e l l e g r i n ,
1909) a n d H. martini (Boulenger, 1906), species w i t h r e l a t i v e l y large eyes l i k e H. mac-
ulipinna ( P e l l e g r i n , 1912) a n d H. hoops G r e e n w o o d , 1967, a n d species w i t h a some-
w h a t s h o r t e r l o w e r jaw, l i k e H. guiarti. It is p o s s i b l e to d e v i d e the p i s c i v o r e s i n
g r o u p s e.g. o n the basis of b o d y f o r m or d e n t i t i o n . H o w e v e r , ' a m o n g the p i s c i v o r o u s
predators it is possible to detect series s h o w i n g a g r a d u a l , species b y species change-
o v e r f r o m a b i c u s p i d b i t i n g d e n t i t i o n to u n i c u s p i d g r a s p i n g d e n t i t i o n , ( G r e e n w o o d ,
1984: 149). T h e r e f o r e p r o b l e m s arise w h e n w e w a n t to d e l i m i t these g r o u p s . F o r
a l m o s t a n y character a series of species can be m a d e e x h i b i t i n g the r a n g e f r o m absent
to f u l l y d e v e l l o p e d ( G r e e n w o o d , 1974, 1984). It is this consistent lack of clear m o r -
p h o l o g i c a l gaps that m a k e s the L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d f a u n a so u n i q u e .
M a y b e , rather than m a k i n g artifical taxonomie separations i n m o r p h o c l i n e s w e
s h o u l d a c c e p t the fact that the e v o l u t i o n of the h a p l o c h r o m i n e c i c h l i d s i n L a k e
V i c t o r i a h a s n o t (yet?) p r o d u c e d g r o u p s of s p e c i e s t h a t d e s e r v e g e n e r i c s t a t u s .
%
H. bareli
H. dichourus 'Mwanza*
z
0
H. pyrrhopteryx
ranges : Greenwood, 1980 { 1

ranges : van Oijen {
<
0 Harpagochromis
!
X
1 Prognathochromis >
f
H . bareli 8
X
H. dichourus 'Mwanza'
O
H. pyrrhopteryx m
n
ranges: Greenwood, 1980 { n
PC
3
ranges : van Oijen { >
m
0
>
<
Fig. 7. Comparison of ranges of neurocranial measurements of species of Harpagochromis and Prognathochromis, as measured by Greenwood (1980) and by van 1
Oijen on skeletal material from the B M N H collection (see table 1). Measurements of specimens of H. bareli spec, nov., H. dichrourus "Mwanza" and H. pyrrho-
pteryx specnov. from the Mwanza Gulf are by van Oijen, and have not been included in the ranges. OtD = Otic Depth, O D = Orbital Depth, PrOD = Preorbital
>
Depth, N W = Neurocranial Width.
(Λ '
Φ 3
V
O. 7
οο
CO ·
-ΟΌ—ΟΟ
Ο Ο-ΟΌ
ο- -—Ο
(3D
72 23 24 25
OD/NL PrOD/NL
(Λ 2
ω 3
Q- 7
ο οοοο
34 35 36 37
OtD/NL NW/NL
Fig. 8. Neurocranial measurements of species placed by Greenwood (1980) in Harpagochromis,

Prognathochromis and Psammochromis, taken from neurocrania in the BM(NH) collection. Numbers on
vertical axis refer to species in table 1. OtD = Otic Depth, O D = Orbital Depth, PrOD = Preorbital
Depth, N W = Neurocranial Width. · = Prognathochromis (Greenwood, 1980), Ο = Harpagochromis
(Greenwood, 1980), • = Psammochromis (Greenwood, 1980).
P e r h a p s the d e f i n i t i o n of the genus Haplochromis s h o u l d be b r o a d e n e d rather t h a n

n a r r o w e d , to e n c o m p a s s a l l e n d e m i c L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s . T h e g a p
b e t w e e n Haplochromis a n d the m o n o t y p i c g e n e r a w a s a l r e a d y d i s p u t e d b y R e g a n
(1922: 160) w h o , i n h i s r e v i s i o n of the c i c h l i d fishes of L a k e V i c t o r i a stated: 'a f e w
extreme types are r e g a r d e d as generically distinct a l t h o u g h the close r e l a t i o n s h i p of
each to a species of Haplochromis is o b v i o u s / G r e e n w o o d (1974: 103), n o t l o n g before
his generic r e v i s i o n w r o t e : T h e generic status of Macropleurodus a n d Paralabidochro-
mis i s , I n o w t h i n k q u e s t i o n a b l e / T h e m o r p h o l o g i c a l differences b e t w e e n several
Haplochromis species are of the same m a g n i t u d e as those b e t w e e n certain Haplochro
mis species a n d Platytaeniodus degeni a n d Hoplotilapia retrodens (Witte, pers. c o m m . )
In m y v i e w G r e e n w o o d (1967: 32), d i s c u s s i n g the p r o b l e m of the generic d i v i s i o n
of the L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s , w a s r i g h t w h e n he stated: Ί d o n o t t h i n k
that the q u e s t i o n c a n be dealt w i t h u n t i l l the w h o l e L a k e V i c t o r i a species f l o c k has
been d e s c r i b e d / T i l l then, I w o u l d p r o p o s e to describe a l l n e w species of L a k e V i c
t o r i a h a p l o c h r o m i n e species i n the genus Haplochromis.
C r i t e r i a to d i s t i n g u i s h species
C r i t e r i a to d i s t i n g u i s h h a p l o c h r o m i n e species i n L a k e V i c t o r i a w e r e p r o p o s e d b y
v a n O i j e n et a l . (1981) a n d r e f i n e d b y H o o g e r h o u d & W i t t e (1981) a n d W i t t e & W i t t e -
M a a s (1987). A d u l t specimens are c o n s i d e r e d conspecific w h e n they share a u n i q u e
c o m b i n a t i o n of m o r p h o l o g i c a l (incl. coloration), e t h o l o g i c a l a n d e c o l o g i c a l charac
ters. S p e c i e s d e s c r i p t i o n of c i c h l i d s f r o m L a k e M a l a w i ( M a r s h et a l , 1981; L e w i s ,
1982; M a r s h , 1983; R i b b i n k et a l . , 1983) a n d L a k e T a n g a n y i k a ( Y a m a o k a , 1982, 1983)
g i v e the i m p r e s s i o n that i n f o r m a t i o n o n d e p t h d i s t r i b u t i o n , t e r r i t o r y size, f e e d i n g
b e h a v i o u r , f e e d i n g site u t i l i z a t i o n , agression a n d c o u r t s h i p collected b y u n d e r w a t e r
observations are almost i n d i s p e n s i b l e for getting a clear i n s i g h t i n t o the t a x o n o m y of
lacustrine c i c h l i d s .
I n d e e d i t is o n l y b y u n d e r w a t e r observations that direct evidence for r e p r o d u c t i v e
i s o l a t i o n u n d e r n a t u r a l c o n d i t i o n s , M a y r ' s (1942) criterion for species d e f i n i t i o n , c a n
be o b t a i n e d . H o w e v e r , because of the opacity of the water, u n d e r w a t e r observations
i n the M w a n z a G u l f are i m p o s s i b l e . T h u s , i n contrast to researchers w o r k i n g o n
c i c h l i d s i n the clear rift lakes, taxonomists d e a l i n g w i t h L a k e V i c t o r i a material h a v e to
r e l y m o r e o n the c l a s s i c a l t a x o n o m i c a l a p p r o a c h c o n c e n t r a t i n g o n m o r p h o l o g i c a l
differences. H o w e v e r , field data o n l i v e coloration w e r e often essential to trigger the
search for m o r p h o l o g i c a l differences (see H o o g e r h o u d & W i t t e , 1981). I f o l l o w Witte
& W i t t e - M a a s (1987) i n t a k i n g as a r u l e that because of its role i n the Specific M a t e
R e c o g n i t i o n S y s t e m , distinct differences i n m a l e coloration indicate different species.
I n a d d i t i o n to d a t a o n c o l o u r p a t t e r n s , d a t a c o n c e r n i n g b r e e d i n g p l a c e s a n d
b r e e d i n g seasons, d e d u c e d f r o m catch data, a n d ecological data i n f e r r e d f r o m
s t o m a c h contents investigations a n d samples f r o m f o o d o r g a n i s m s f r o m the preferred
habitat, are u s e d as m u c h as possible (e.g. H o o g e r h o u d et a l . , 1983; G o l d s c h m i d t et a l . ,
1990; G o l d s c h m i d t & W i t t e , 1990)).
Intraspecific v a r i a t i o n
W h e n c o m p a r i n g s p e c i m e n s of d e s c r i b e d species f r o m the M w a n z a G u l f w i t h
m a t e r i a l f r o m the n o r t h e r n part of the lake, m y m o r p h o l o g i c a l criteria for species d i s -
20 Z O O L O G I S C H E V E R H A N D E L I N G E N 272 (1991)
t i n c t i o n a p p e a r e d n a r r o w e r t h a n that of p r e v i o u s authors. M a t e r i a l u s e d for redes-

c r i p t i o n s p r o v e d p o l y s p e c i f i c a n d c o n t a i n e d specimens w h i c h c o u l d not be p l a c e d i n
a n y of the d e s c r i b e d species. I n a n u m b e r of cases these specimens c o u l d be p l a c e d i n
w h a t I c o n s i d e r e d u n d e s c r i b e d species f r o m the M w a n z a G u l f . A f t e r the m i s i d e n t i -
f i e d specimens w e r e r e m o v e d , m o r p h o m e t r i c ranges of the m a t e r i a l f r o m the n o r t h -
ern part of the l a k e still w e r e f o u n d to be larger t h a n those of conspecifics f r o m the
M w a n z a G u l f . T h i s i n spite of the fact that collections f r o m the M w a n z a G u l f for a
certain size r a n g e are larger. T h e w i d e r m o r p h o m e t r i c ranges of the species i n the
B M ( N H ) collections m a y be c a u s e d b y the fact that the B M ( N H ) collections of o n e
p a r t i c u l a r species are c o m p o s e d of samples f r o m different localities w h i c h are often
h u n d r e d s of k i l o m e t e r s a p a r t . G e o g r a p h i c a l v a r i a t i o n i n m o r p h o m e t r i c r a n g e s
a p p a r e n t l y is a c o m m o n feature of L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s (see W i t t e &
W i t t e - M a a s , 1987).
R e c e n t l y the first e x a m p l e s of intraspecific v a r i a t i o n i n L a k e V i c t o r i a h a p l o c h r o -
m i n e c i c h l i d s w e r e d e s c r i b e d b y W i t t e & W i t t e - M a a s (1987). W o r k i n g o n z o o p l a n k -
t i v o r o u s species collected f r o m different areas of L a k e V i c t o r i a , they n o t i c e d differ-
ences i n certain m o r p h o m e t r i c ranges a n d details of c o l o r a t i o n b e t w e e n a l l o p a t r i c
s p e c i m e n s b e l o n g i n g to one species. C i t i n g m a n y r e c e n t l y d e s c r i b e d e x a m p l e s of
p h e n o t y p i c p l a s t i c i t y i n c i c h l i d s , W i t t e & W i t t e - M a a s (1987) m a d e it p l a u s i b l e that
the differences b e t w e e n the a l l o p a t r i c f o r m s c o u l d be, for the greater part, a result of
i n f l u e n c e s of e n v i r o n m e n t a l c o n d i t i o n s o n the p h e n o t y p e , a n d c o n s e q u e n t l y des-
c r i b e d the f o r m s as p o p u l a t i o n s . Regrettably, i n s u f f i c i e n t m a t e r i a l f r o m d i f f e r e n t
areas is a v a i l a b l e to engage i n a s i m i l a r analysis for the species d e s c r i b e d b e l o w . Spe-
c i m e n s of H . dichrourus R e g a n , 1922, w e r e collected f r o m f o u r different areas a r o u n d
the lake, b u t f r o m three areas o n l y t w o to f o u r specimens w e r e collected. H o w e v e r ,
c o m p a r i s o n of the f o u r g r o u p s of specimens s h o w e d s m a l l b u t consistent differences
i n m o r p h o m e t r i c measurements a n d differences i n the shape of h e a d m a r k i n g s . A s it
is v e r y u n l i k e l y that a d d i t i o n a l m a t e r i a l of this species w i l l b e c o m e a v a i l a b l e w e
p r o b a b l y w i l l never be able to f i n d out w h e t h e r the v a r i a t i o n is c l i n a l or d i s c o n t i n u -
o u s . T h e r e f o r e the t a x o n o m i e status of these f o r m s w i l l a l w a y s be u n c e r t a i n . To
e m p h a s i z e the existence of g e o g r a p h i c v a r i a t i o n i n this species I h a v e , l i k e W i t t e &
W i t t e - M a a s (1987), a d a p t e d the suggestion of W i l s o n & B r o w n (1953), w h o advocat-
ed the usefulness of a d d i n g to the specific n a m e , the n a m e of the l o c a l i t y w h e r e the
p o p u l a t i o n is f o u n d . C o n s i d e r i n g the distance b e t w e e n v a r i o u s areas w h e r e the spec-
i m e n s of H. dichrourus w e r e collected a n d the ecological barriers w h i c h separate the
areas, the t a x o n o m i e status of p o p u l a t i o n for the g e o g r a p h i c f o r m s seems a m p l y jus-
tifiable. F o u r s p e c i m e n s of another species, H. pyrrhopteryx spec, nov., c o l l e c t e d i n
three different areas o u t s i d e the M w a n z a G u l f , d i d not d i f f e r m a r k e d l y f r o m their
M w a n z a G u l f c o n s p e c i f i c s . T h i s species i n h a b i t s deeper w a t e r s , w h i c h i n v a r i o u s
parts of the l a k e h a v e m o r e e c o l o g i c a l l y i m p o r t a n t factors i n c o m m o n t h a n h a v e the
s h a l l o w areas. P o s s i b l y the species has a m o r e c o n t i n u o u s d i s t r i b u t i o n .
S e x u a l d i m o r p h i s m i n m o r p h o l o g i c a l characters w a s o n l y f o u n d i n the m a x i m u m
size: i n a l l species, here d e s c r i b e d , for w h i c h a d u l t males a n d females are k n o w n ,
females reach a larger size t h a n males.
V A N OIJE N: PISCIVOROUS H A P L O C H R O M I N E CICHLIDAE O F L A K E VICTORIA 21
Species descriptions
H a p l o c h r o m i s b a r e l i spec. nov.
(figs 3,925,104, tables 24)
Haplochromis "black"; van Oijen, 1982: 340,349.
Material. — Holotype, cf, 100.5 mm, R M N H 29600, Mwanza Gulf, Tanzania, Lake Victoria, 7.X.1977,
HEST; Paratypes: 1 9, 69.1 mm + 3 <f<f, 89.093.0 mm, R M N H 2961215, Mwanza Gulf, Tanzania,
Lake Victoria, G C h . Anker & C D . N . Barel. Other para types from the Mwanza Gulf, Tanzania, Lake
Victoria collected by HE ST; 2 99, 772, 89.5 mm, R M N H 29651, 29616, 1977; 1 <f, 81.8 mm + U , 105.8
mm, M N H N 19871668, 4.xi.l977; 2 99, 98.4, 109.8 mm, R M N H 2965455, 18.xi.1977; 1 9, 106.2 mm,
R M N H 29617, 28.xi.1977; 1 <f, 103.0 mm B M N H 1987.2.4:18., 16.xii.1977; 1 9, 108.3 mm, R M N H
29619, 23.xii.1977; 1 9,118.2 mm, R M N H 29610, 6.1.1978; 1 o", 97.4 mm, R M N H 29611, 7.iv.l978; 1 9,
63.8 mm, R M N H 29620, 28.iv.1978; 1 <f, 64.0 mm, R M N H 29621, 5.V.1978; 1 o", 73 mm, R M N H 29622,
19.V.1978; 1 9, 103.5 mm, R M N H 29623, 26.V.1978; 1 9 114.1 mm, R M N H 29624, vi.1978; 1 9 Π5.2
f f
mm, R M N H 29625, 9.vi.l978; 1 <f, 109.1 m m + l f 116.0 mm, R M N H 2962829, 29.xii.1978; 1 f, 78.1
mm, A N S P 168240, 29.xii.1978; 1 <f, 103.0 mm, R M N H 29627, 29.xii.1978; 1 9, 119.8 mm, R M N H
29607,1978; 1 <S, 102.0 mm, R M N H 29609,19.Í.1979; 1 d\ 109.9 mm, R M N H 29632, 9.xi.l979; 1 9, 82.6
mm, ANSP 168239; 1 9, 44,5 mm, R M N H 29633,12.iii.1979; 1 o", 92.4 mm, R M N H 29634,14.iv.1979; 2
99, 82.0, 96.2 mm + 3 cf(f, 81.397.0 mm R M N H 2963539, 20.iv.1979; 1 9, 100.0 m m S.L., R M N H
30388, 20.ÍV.19978; 1 9, 52.8 mm, R M N H 29640, 25.iv 1979; 2 cfcf, 57.5, 92.00 mm, R M N H 2964142,
4.V.1979; 3 cftf, 78.194.6 mm, R M N H 2960103, 4.V.1979; 1 (f, 42.5 mm, R M N H 29643, 13.iii.1980; 1 f,
108.0 mm, R M N H 29644, 21.iv.1980; 1 9,125.0 mm, B M N H 1987.2.4:19, 22.iv.1980; 2 99, 124.5, 124.8,
R M N H 2964647, 23.iv.1980; 1 9, 41.1 mm, R M N H 29648, 16.V.1980; 1 <f, 97.0 mm, R M N H 29608,
22.Í.1980; 1 cf, 71.1 mm + 1 9, 72.0 mm, R M N H 2964950, l.vi.1984. Paratype from Speke Gulf: 1 9,
100 mm, R M N H 29626, 2.viii.l978. Other material: 2 99, 99.0,106.0 mm, R M N H 3038283,6.ix.l979.
E t y m o l o g y . — T h i s species is n a m e d i n h o n o u r of D r C . D . N . Barel, the initiator of

the H a p l o c h r o m i s E c o l o g y Project, w h o first collected specimens of this species. H i s
s t i m u l a t i n g e n t h u s i a s m a n d interest i n all aspects o f b i o l o g y e n g a g e d m a n y b i o l o g y
students i n c i c h l i d research. H i s research has c o n t r i b u t e d m u c h to o u r k n o w l e d g e of
the L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s .
D i a g n o s i s . — A s m a l l to m e d i u m s i z e d , r e l a t i v e l y d e e p b o d i e d , p i s c i v o r o u s spe
cies w i t h r e l a t i v e l y l a r g e eyes a n d s t r o n g l y r e c u r v e d u n i c u s p i d s teeth i n the o r a l
jaws. L i v e males are entirely black o n the v e n t r a l t w o t h i r d o f h e a d a n d b o d y . Snout
b r o w n i s h . D o r s a l part o f h e a d a n d b o d y , d a r k blueishgrey. L i v e females h a v e at least
the l o w e r half of the f l a n k black, b u t the v e n t r a l s i d e is b r i g h t w h i t e . Snout a n d h e a d
v e n t r a l to the eye g r e y i s h . D o r s a l part of h e a d a n d b o d y g o l d e n y e l l o w .
D e s c r i p t i o n . — (based o n 55 s p e c i m e n s ( i n c l u d i n g the h o l o t y p e ) 41125 m m stan
d a r d length).
H a b i t u s . — A r e l a t i v e l y d e e p b o d i e d species w i t h a rather g e n e r a l i z e d habitus.
D o r s a l h e a d o u t l i n e h a r d l y to s l i g h t l y i n t e r r u p t e d b y the p r e m a x i l l a r y p e d i c e l . Snout
s o m e w h a t b l u n t . C e p h a l i c lateral l i n e o p e n i n g s not e n l a r g e d ; lateral l i n e canals o n
the l a c h r y m a l not v i s i b l e . E y e circular a n d relatively large, p u p i l s u b c i r c u l a r w i t h a
distinct a p h a k i c aperture rostroventrally to the lens. M o u t h s l i g h t l y to m o d e r a t e l y
o b l i q u e . P r e m a x i l l a not e x p a n d e d m e d i a l l y . E x p o s e d part o f m a x i l l a s m a l l , the poste
r i o r tip r e a c h i n g a v e r t i c a l just rostral to the anterior b o r d e r of the eye lens. L i p s nor
m a l . L o w e r j a w i s o g n a t h o u s o r p r o g n a t h o u s ( m a i n l y i n larger specimens), s l i g h t l y
p r o t r u d i n g . R o s t r a l o u t l i n e of l o w e r j a w s l i g h t l y c o n v e x w i t h a s m o o t h l y r o u n d e d
m e n t a l area, r a r e l y w i t h a s m a l l m e n t a l p r o t u b e r a n c e . L a t e r a l sides o f l o w e r j a w
s l i g h t l y to m o d e r a t e l y o b l i q u e .
S c a l e s . — C h e e k , g i l l - c o v e r , n a p e a n d neck w i t h c y c l o i d scales; w e a k l y c t e n o i d
scales s o m e t i m e s present o n g i l l - c o v e r a n d nape. Scales o n the d o r s u m either a l l c y -
c l o i d o r a n a d m i x t u r e of c y c l o i d a n d a f e w w e a k l y c t e n o i d scales. S o m e c y c l o i d
scales m a y b e present o n the v e n t r a l part of the chest, o t h e r w i s e a l l b o d y scales cte-
n o i d . A g r a d u a l size t r a n s i t i o n b e t w e e n scales o f chest a n d the a d j o i n i n g parts o f the
b o d y . S m a l l elongate scales o n the p r o x i m a l half o f the c a u d a l f i n .
F i n s . — D o r s a l a n d a n a l f i n r e l a t i v e l y h i g h f o r a p i s c i v o r o u s species. Pectoral a n d
p e l v i c fins just o r n o t quite r e a c h i n g the anal f i n . D o r s a l a n d a n a l f i n n o t r e a c h i n g
base o f c a u d a l f i n . P e l v i c s w i t h first soft r a y s l i g h t l y p r o d u c e d . C a u d a l f i n o u t l i n e
truncate a n d s l i g h t l y emarginate.
G i l l - a p p a r a t u s . — E i g h t or n i n e g i l l rakers o n the l o w e r part o f the first g i l l arch;
the l o w e r m o s t 1 o r 2 r e d u c e d , the f o l l o w i n g 3 slender a n d s o m e w h a t c o n i c a l , a n d the
u p p e r 4 flattened w i t h s l i g h t l y e x p a n d e d c r o w n s . A p p r o x i m a t e l y 128 g i l l filaments
o n the first h e m i b r a n c h .
V i s c e r a . — Intestine l e n g t h varies f r o m 1.1-1.3 times the s t a n d a r d l e n g t h (n = 30).
F o l l o w i n g the d e f i n i t i o n s of Z i h l e r (1982) the arrangement of the d i g e s t i v e tracks of
a d u l t H. bareli is of t y p e D ( w i t h backflap).
O r a l t e e t h . — Shape. I n a d u l t specimens o v e r 77 m m S L , teeth of the outer r o w
are r a t h e r stout, c o n i c a l a n d a c u t e l y p o i n t e d u n i c u s p i d s , m o d e r a t e l y to s t r o n g l y
c u r v e d . S p e c i m e n s s m a l l e r than 77 m m S L h a v e a n a d m i x t u r e of e q u a l l y b i c u s p i d s
a n d u n i c u s p i d s , w i t h the u n i c u s p i d s d o m i n a t i n g . I n these specimens the teeth, espe-
c i a l l y the c r o w n s , are s o m e w h a t compressed. T h e u n i c u s p i d s m a y be s h o u l d e r e d , the
b i c u s p i d s w i t h the major cusp s h a r p l y p o i n t e d . Inner r o w s : I n larger specimens the
i n n e r r o w s i n b o t h j a w s consist m a i n l y of r o u n d e d , acutely p o i n t e d u n i c u s p i d s , s o m e
w e a k l y b i c u s p i d s a n d t r i c u s p i d s m a y also be f o u n d . B i c u s p i d a n d t r i c u s p i d teeth
m o r e n u m e r o u s i n s m a l l e r specimens. A l l i n n e r teeth m o d e r a t e l y c u r v e d . T o o t h size
g r a d u a l l y decreasing f r o m rostral to c a u d a l .
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d i n u p p e r j a w , s l i g h t l y
acute i n l o w e r jaw. T w o i n n e r r o w s i n b o t h jaws. Distance b e t w e e n outer r o w a n d
first i n n e r r o w s l i g h t l y larger t h a n the distance b e t w e e n the i n n e r r o w s .
Fig. 9. Haplochromis bareli spec. nov. Paratype, R M N H 29601. Scale equals 10 mm.
Figs 10-23. Haplochromis bareli spec. nov. Fig. 10. Premaxilla, lateral view. Fig. 11. Premaxilla, view per-
pendicular to dentigerous area. Fig. 12. Premaxilla, medial view of ascending arm. Fig. 13. Lower jaw,
lateral view. Fig. 14. Lower pharyngeal element, dorsal view. Fig. 15. Lower pharhyngeal element, lat-
eral view. Fig. 16. Lower pharhyngeal element, caudal view. Figs. 17-20. Premaxillary teeth, labial and
lateral view. Fig. 17. Fourth tooth of outer row. Fig. 18. Second tooth of outer row. Fig. 19. Third tooth
of first inner row. Fig. 20. Fourth tooth of first inner row. Figs. 21-23. Lower pharhyngeal teeth, lateral
and rostral view. Fig. 21. Seventh lateral tooth. Fig. 22. Fourth tooth of caudalmost row. Fig. 23. Second
tooth of caudalmost row. Scale of bony elements = 5 mm, scale of teeth = 1 mm.
— C o u n t s a n d setting. 52-62 teeth i n the outer r o w of the u p p e r jaw, 36-52 i n the

l o w e r jaw, the n u m b e r s s h o w i n g a p o s i t i v e correlation w i t h s t a n d a r d l e n g t h . O u t e r
r o w teeth are r e g u l a r l y set at a distance e q u a l to or s l i g h t l y less than the diameter of
the toothbase. O u t e r r o w o c c u p y i n g 9/10 of the p r e m a x i l l a a n d o n the d e n t a r y reach-
i n g the first t h i r d of the c o r o n o i d w i n g . Inner r o w s o c c u p y i n g 3/5 of the outer r o w
l e n g t h o n the p r e m a x i l l a a n d 3/4 o n the m a n d i b l e .
— I m p l a n t a t i o n . In u p p e r j a w outer r o w teeth erect ( m e d i a l teeth) to s l i g h t l y re-
c u m b e n t (caudal teeth). In l o w e r j a w outer r o w teeth s l i g h t l y p r o c u m b e n t (medially)
to erect ( c a u d a l l y ) . A l l teeth s l i g h t l y s u s p e n s o r i a d i n c l i n e d . I m p l a n t a t i o n of i n n e r
r o w teeth s t r o n g l y recumbent.
P h a r y n g e a l t e e t h . — C o u n t s . A b o u t 20 teeth i n the c a u d a l m o s t r o w . N i n e teeth i n
the m e d i a n series.
— Shape. M o s t p h a r y n g e a l teeth of the b e v e l l e d type, r e l a t i v e l y fine a n d acutely
p o i n t e d ; s o m e m e d i a l teeth of c a u d a l m o s t r o w a n d s o m e c a u d a l teeth of m e d i a n r o w
r e l a t i v e l y stout a n d h o o k e d , w i t h a b l u n t r o s t r o - d o r s a d directed major c u s p . Tooth
size i n c r e a s i n g f r o m rostral to c a u d a l a n d f r o m lateral to m e d i a l .
O s t e o l o g y . — T h e osteological descriptions are based o n skeletal elements of f i v e
dissected specimens, S L 97.4-119.8 m m . D e s c r i p t i o n of o r a l teeth based o n a l l t y p e
specimens.
— N e u r o c r a n i u m . C o m p a r e d w i t h the n e u r o c r a n i u m of a g e n e r a l i z e d c i c h l i d (H.
elegans (Barel et a l . , 1976)) that of H. bareli is b o t h b r o a d e r a n d m o r e slender, the otic
r e g i o n is less deep b u t the s u p r a o c c i p i t a l crest is deeper. T h e e t h m o v o m e r i n e b l o c k is
less d e c u r v e d a n d s l i g h t l y longer t h a n that of H. elegans. A s i n m o s t p i s c i v o r o u s c i c h -
l i d s f r o m L a k e V i c t o r i a the p h a r y n g e a l a p o p h y s i s is situated further c a u d a l l y as c o m -
p a r e d w i t h the g e n e r a l i z e d type, a n d the p o s t o r b i t a l process is b r o a d e n e d .
— O r a l jaws. P r e m a x i l l a ; dentigerous a r m l o n g e r t h a n a s c e n d i n g a r m , angle be-
t w e e n the a r m s 78-82°. B o t h a r m s r e l a t i v e l y b r o a d . V e n t r a l o u t l i n e of d e n t i g e r o u s
a r m a l m o s t straight. V e n t r a l part of the a s c e n d i n g a r m v e r y s l i g h t l y e x p a n d e d . M a n -
d i b l e r e l a t i v e l y stout, w i t h a rather steep c o r o n o i d process. L e n g t h / d e p t h ratio of
2.5. T h e tooth b e a r i n g part is s l i g h t l y less than half the j a w l e n g t h .
— L o w e r p h a r y n g e a l element. L o w e r p h a r h y n g e a l element rather shallow,
s l i g h t l y l o n g e r t h a n b r o a d ( l e n g t h / w i d t h = 1.05-1.17). D e n t i g e r o u s area s l i g h t l y
b r o a d e r t h a n l o n g ( l e n g t h / w i d t h = 0.85-1.00).
— Vertebrae. There are 29 vertebrae, c o m p r i s i n g 13 a b d o m i n a l a n d 16 c a u d a l ele-
m e n t s (n= 7).
C o l o r a t i o n . — L i v e quiescent males have a deep black c o l o u r b e l o w a n i r r e g u l a r
h o r i z o n t a l l i n e t h r o u g h the d o r s a l m a r g i n of the gillcover. T h e i r r e g u l a r shape of the
d o r s a l b o r d e r of the b l a c k area indicates that it is b u i l t u p of six or s e v e n b r o a d e n e d
v e r t i c a l bars. T h e cheek a n d snout m a y be s l i g h t l y lighter w i t h a d a r k b r o w n o r c o p -
per sheen. D o r s a l aspects of snout d a r k g o l d e n green. T h e d o r s a l , lighter part of the
b o d y d a r k grey w i t h a distinct b l u i s h f l u s h . F i n s : pectorals grey, p e l v i c s black, a n a l
b l a c k r o s t r a l l y a n d near the f i n base, the r e m a i n i n g part w h i t e w i t h a faint r e d f l u s h
rostrally. D o r s o - c a u d a l l y o n the a n a l t w o or three relatively large, o r a n g e - y e l l o w egg-
d u m m i e s b o r d e r e d b y a w h i t e i n n e r a n d a d a r k outer rim, the e g g - d u m m i e s some-
times p a r t l y f u s e d . D o r s a l f i n d u l l w h i t e , w i t h s m a l l , d a r k lappets a n d s m a l l d a r k r e d
spots b e t w e e n the rays. C a u d a l sooty p r o x i m a l l y , d u l l w h i t e distally, d a r k r e d spots
a n d streaks b e t w e e n the rays.
— S e x u a l l y active m a l e s . A l m o s t entirely b l a c k , the r e d spots b e t w e e n the f i n

r a y s b r i g h t e r t h a n i n quiescent males. C a u d a l f i n u s u a l l y w i t h a r e d f l u s h . E x c e p -
t i o n a l l y the entire c a u d a l a n d a n a l fins b r i g h t r e d . A b r o w n - c o p p e r y f l u s h o n the
h e a d a n d d o r s a l part of the b o d y .
— S e x u a l l y active females. V e n t r a l part of the h e a d grey w i t h a l i g h t e r v e n t r a l
side. G i l l - c o v e r d a r k grey to black. T h e b o d y w i t h a p a r t l y s i m i l a r b l a c k area as i n
males. T h e d o r s a l a n d c a u d a l extension of the b l a c k area v a r i a b l e . M o r e often t h a n i n
males the females h a v e the c a u d a l part of the black area b r o k e n u p i n b r o a d vertical
bars. D o r s a l m a r g i n of b l a c k area m o s t l y t o u c h i n g a h o r i z o n t a l t h r o u g h the d o r s a l
e d g e of the g i l l - c o v e r b u t i n s o m e specimens e x t e n d e d to the d o r s a l f i n base. C h e s t
a n d v e n t r a l s i d e b r i g h t w h i t e , c a u d a l part of c a u d a l p e d u n c l e l i g h t e r grey. D o r s a l
parts of b o d y a n d h e a d g r e y i s h - y e l l o w . F i n s : pectorals h y a l i n e , p e l v i c s w h i t e . A n a l
Fig. 24. Haplochromis bareli spec. nov. Live colours and markings of a sexually active male and ripe
female.
Table 2. Ranges of linear measurements of Haplochromis bareli spec. nov.
Standard length range in mm 41.0-57.5 63.3-78.1 81.0-89.5 90.8-98.4 100.0-109.1 114.1-125.0

Number of specimens 5 9 10 10 11 9
Body Depth %SL 30.7-33 8 33.7-37.3 33.3-37.9 33.4-36.7 34.2-36.1 35.3-37.8

Pectoral Fin Length %SL 23.5-29 2 27.0-31.1 28.1-30.4 28.2-31.9 27.4-32.7 27.6-32.1
Caudal Peduncle Length %SL 16.6-19.7 14.3-18.2 15.2-17.1 14.7-18.3 14.8-17.2 14.8-16.9
Caudal Peduncle Depth %SL 11.8-12.5 10.6-12.9 11.3-12.4 10.8-13.2 10.9-12.5 11.2-12.0 Ν
Caudal Fin Length %SL 24.4-26.5 23.5-27.2 23.1-25.1 23.1-25.9 21.9-24.8 22.5-24.7
Head Length %SL 34.8-37.7 36.3-37.9 35.6-38.1 35.7-38.2 36.0-38.0 36.9-38.5 8
Snout Length % HL 25.3-30.5 26.8-31.2 28.5-31.3 28.9-33.3 30.2-33.0 31.3-34.4
Snout Width % HL 29.5-32.8 31.5-34.7 30.1-34.8 30.2-34.2 29.7-34.7 29.3-34.6 8
Head Width % HL 43.1-45.1 43.9-45.9 44.3-48.8 44.7-46.6 42.7-46.5 42.7-48.1 χ
Interorbital Width % HL 19.0-22.5 20.6-24.4 22.8-26.9 22.7-24.7 22.8-26.5 24.5-28.1 <
m
Preorbital Width % HL 24.5-27.3 26.0-30.1 24.2-30.9 24.7-30.3 26.7-30.8 27.3-32.4
Χ
Lachrymal Width % HL 24.5-29.8 26.0-28.8 26.7-30.1 25.5-29.1 24.8-30.5 26.3-30.7 >
Preorbital Depth % HL 12.6-15.9 14.5-17.0 15.5-17.8 15.0-18.2 15.9-18.0 17.0-19.9 Ζ
Eye Length % HL 29.1-37.8 27.4-31 2 25.3-30.4 24.7-30.7 24.8-28.3 23.7-28.1 ο
Cheek Depth % HL 14 1-20.6 19.0-22 5 19.0-23.6 18.6-22.4 21.1-24.0 22.0-25.1 m
Ζ
Lower Jaw Length % HL 40.5-46.4 42.7-47.4 46.0-49.2 45.1-49.1 47.0-52.0 48.8-53.1 Ο
m
Lower Jaw Width % HL 18.7-22.4 18.6-22.2 17.6-23.5 19.7-23.2 19.7-23.7 19.7-25.9 Ζ
Ν>
Upper Jaw Length % HL 30.5-35.2 33.7-37.Z 33.1-39.2 36.6-38.8 36.0-40.9 37.8-42.5 ν]
Ν>
Premax. Pedicel Length % HL 25.1-26.9 24.1-27.7 24.9-28.1 25.6-27.8 26.2-30.0 26.9-29.8
νθ
Table 3. Means and standard deviations of linear measurements of Haplochromis bareli spec. nov.
VAN
Standard length range in mm 41.057.5 63.378.1 81.089.5 90.898.4 100.0109.1 114.1125 ζ

Number of specimens 5 9 10 10 11 9 O
ο
m
Ζ
Body Depth %SL 32.0±1.2 34.211.3 34.711.7 34.810.7 35.310.8 36.110.8 TJ
Pectoral Fin Length %SL 26.6±2.1 28.511.8 29.210.6 29.911.4 29.511.3 29.711.4
ÍN: PIS(
Caudal Peduncle Length %SL 18.2±1.1 16.811.0 16.210.6 16.011.7 15.710.7 15.910.7 <
<
Caudal Peduncle Depth %SL 12.210.5 12.Qt0.7 12.010.7 11.810.6 11.610.4 11.610.2
Caudal Fin Length %SL 25.5±0.8 24.811.4 24.210.9 24.410.9 23.110.9 23.510.5 §
Head Length %SL 37.111.2 37.310.4 36.710.6 37.110.8 37.010.6 38.010.4
αc
X
Snout Length % HL 27.0±1.8 29.211.5 29.710.8 31.0±1.3 31.210.9 33.110.9
Χ
Snout Width % HL 30.7±1.2 33.Q±1.0 32.711.4 32.311.2 33.312.0 33.112.0
%
Head Width % HL 44.1 ±0.8 44.411.0 44.910.7 45.210.6 45.111.2 45.912.1 Q
Interorbital Width % HL 20.111.2 22.511.1 24.411.2 23.610.7 24.811.2 25.4_1.0 R
Π
X
Preorbital Width % HL 26.111.1 26.9±1.4 27.812.2 28.111.3 28.511.2 28.710.6 Χ
Lachrymal Width % HL 25.911.2 27.410.7 28.Q±1.4 28.0±1.1 28.911.5 28.211.5 Ο
Preorbital Depth % HL 13.911.1 15.611.6 16.410.7 16.410.9 16.810.6 18.410.6 ζ
29.312.3 28.112.5 26.011.5 26.311.1 24.811.6 m
ROM] [NE C
Eye Length % HL 33.913.0

Cheek Depth % HL 16.612.3 20.Qtl.2 20.711.4 20.211.2 22.411.1 23.310.9 η
η
Lower Jaw Length % HL 43.512.4 45.111.4 47.011.0 47.111.2 49.711.4 51.111.6
η
X
r
Lower Jaw Width % HL 20.311.8 20.111.0 20.312.2 20.711.4 21.911.8 24.312.6 5
5
Upper Jaw Length % HL 33.311.6 35.310.9 36.611.6 37.910.6 38.211.3 40.211.6
Premax. Pedicel Length % HL 25.610.6 26.311.1 26.511.3 26.810.4 27.111.4 28.611.4 6
ο
τι
ο
τι
>
.AKE VI<
<
S
>
vi
w h i t e rostrally, m o r e h y a l i n e c a u d a l l y O n e or t w o s m a l l y e l l o w orange spots o n the

c a u d a l part. D o r s a l h y a l i n e , y e l l o w i s h basally. C a u d a l y e l l o w i s h p r o x i m a l l y , h y a l i n e
d i s t a l l y a n d b r i g h t y e l l o w ventrally.
— S e x u a l l y i n a c t i v e females a n d juveniles h a v e the same c o l o r a t i o n as the sexu-
a l l y active females a l t h o u g h the lighter v e n t r a l area is less contrasting a n d the y e l l o w
sheen o n the b o d y is d u l l e r a n d darker. T h e smallest juveniles (SL < 60 m m ) h a v e , o n
the f l a n k a n d the rostral part of the c a u d a l p e d u n c l e , six o r seven v e r t i c a l bars w h i c h
d o not e x t e n d o n t o the d o r s u m or v e n t r a l side.
— P r e s e r v e d c o l o r a t i o n of males a n d females are e q u a l l y d i s t i n c t i v e , as the c o n -
trast b e t w e e n the d o r s a l a n d v e n t r a l parts of the b o d y is increased b y p r e s e r v a t i o n .
T h u s , p r e s e r v e d m a l e s are d e e p b l a c k v e n t r a l l y , w i t h the d o r s a l p a r t of h e a d a n d
b o d y l i g h t g r e y i s h - b r o w n . T h e d o r s a l s i d e a s h a d e lighter. T h e c a u d a l p a r t of the
snout a n d a s m a l l area just c a u d a l to the eye l i g h t grey. O n the h e a d a s u p r a o r b i t a l
stripe, a n o s t r i l stripe a n d a b l o t c h just rostral to the d o r s a l f i n m a y be v i s i b l e . F i n s :
pectorals w h i t i s h , p e l vies a n d rostral part of a n a l black, c a u d a l part of a n a l h y a l i n e .
D o r s a l g r e y i s h , lappets darker. C a u d a l sooty p r o x i m a l l y , h y a l i n e distally. D a r k spots
b e t w e e n rays of d o r s a l a n d c a u d a l .
— P r e s e r v e d females have the h e a d d a r k e r g r e y - b r o w n , n a p e lighter, g i l l - c o v e r
a l m o s t black a n d the b o d y l i g h t y e l l o w i s h - b r o w n . O n f l a n k , b e l l y a n d rostral part of
the c a u d a l p e d u n c l e a black area is present, w h i c h m a y be m o r e or less d i v i d e d i n six
o r seven b r o a d v e r t i c a l bars. C h e s t a n d v e n t r a l side l i g h t y e l l o w i s h . A l l fins w h i t i s h -
hyaline.
D i s t r i b u t i o n . — H. bareli is o n l y k n o w n f r o m L a k e V i c t o r i a . S p e c i m e n s w e r e
caught i n the n o r t h e r n part a n d the entrance of the M w a n z a G u l f , a n d near N a f u b a
I s l a n d a n d i n M a g u B a y of the Speke G u l f (Tanzania).
E c o l o g y . — O c c u r r e n c e . In the p e r i o d f r o m 1977-1980 H. bareli w a s a c o m m o n p i s -
c i v o r o u s species i n the o p e n water of the M w a n z a G u l f . Since the e x p l o s i v e increase
of N i l e p e r c h p o p u l a t i o n i n the M w a n z a G u l f area, H. bareli has d i s a p p e a r e d f r o m the
catches.
— H a b i t a t . M o s t specimens w e r e caught i n a b o t t o m t r a w l o v e r m u d b o t t o m s
w i t h a d e p t h of 8-20 m . A f e w s p e c i m e n s w e r e c a p t u r e d at a d e p t h of 25 m . T h e
smallest specimens w e r e collected over m u d bottoms at a d e p t h of 2-4 m i n s h a l l o w
bays. G i l l n e t catches indicate that H. bareli lives near the b o t t o m .
— F o o d . Intestines of 36 specimens of H. bareli w e r e e x a m i n e d . T h i r t e e n of these
w e r e e m p t y , 18 c o n t a i n e d remains of h a p l o c h r o m i n e fishes, t w o c o n t a i n e d remains
of Rastrineobola argentea (Pellegrin, 1904), five contained Chaoborus L i c h t e n s t e i n , 1800,
l a r v a e , p u p a e a n d u n d e t e r m i n a b l e insects, a n d remains of the s h r i m p Caridina niloti-
ca ( R o u x , 1833) w e r e f o u n d i n o n l y one s p e c i m e n each. H. bareli specimens of 45 to 98
m m S L , w e r e f o u n d to p r e y o n h a p l o c h r o m i n e s of 9 to 16 m m , a n d specimens of 117
a n d 118 m m S L h a d p r e y e d o n h a p l o c h r o m i n e s of 20 to 30 m m . H . b a r e l i therefore is
a p r e d a t o r of j u v e n i l e (mostly postbuccal) h a p l o c h r o m i n e s . T h e Rastrineobola preys
w e r e of a m u c h larger size; a s p e c i m e n of H. bareli of 110 m m S L h a d ingested a spec-
i m e n of 50 m m (see v a n O i j e n , 1989 for an e x p l a n a t i o n of the l e n g t h difference be-
t w e e n the p r e y species).
— B r e e d i n g a n d g r o w t h . H. bareli m a t u r e s at a p p r o x i m a t e l y 80 m m S L . R i p e
males h a v e been caught f r o m J a n u a r y till M a y , spent females w e r e present i n catches
d u r i n g A p r i l a n d A u g u s t , b u t n o i n f o r m a t i o n is available o n the type of b r e e d i n g .
m V"0
H. bareli
* H. chrysogynaion
Mwanza G u l f
100 k m
Fig. 25. Catch localities of Haplochromis bareli spec. nov. and H. chrysogynaion spec. nov. in the southern
part of Lake Victoria.
Table 4. Angular and qualitative measurements, and counts of Haplochromis bareli spec. nov.
ranges mean ± st.dev.
Dorsal Head Profile Inclination 27°-43° 33.0° ±4°

Premaxillary Pedicel Inclination 32°-50° 41.0° ±4°
Snout Acuteness 65°-79° 73.2° ± 3°
Gape Inclination 30°-38° 34.6° ± 2°
Dorsal Head Profile (curvature) +

Premaxillary Pedicel Prominence (K+)/+
Lower Jaw Anterior Extension 0/+
Lateral Snout Outline 0/+
Mental Prominence 0/+
Lip Thickening 0
Premaxilla Beaked 0
Premaxilla Expanded 0
Maxillary Posterior Extension 0
Cephalic Lateral Line Pores: Width 0
Lateral Line Scales 30 (f= 2); 31 (f= 7); 32 (f= 17); 33 (f= 12)
Lateral Line - Dorsal Fin (Sc.rows) 5 (f= 10); 6 (f= 20); 7 (f= 8)
Pectoral - Pelvic Fin Bases (Sc.rows) 5 (f= 10); 6 (f= 18); 7 (f= 8); 8 (f= 2)
Cheek (Vertical Sc.rows) 3 (f= 6); 4 (f= 31); 5 (f= 1)
Dorsal Fin (Spines/Rays) XIV 9 (f= 1); XV8(f=l); XV9(f=25);

XV 10 (f= 6); XVI 8 (f= 2); XVI 9 (f= 2)
Anal Fin (Spines/Rays) III 8 (f= 19); III 9 (f= 19)
T h e r i g h t o v a r y g r o w s larger t h a n the left o n e . E g g s m e a s u r e d i n r i p e o v a r i e s o f

f e m a l e s o f 9 2 a n d 93 m m S L w e r e a p p r o x i m a t e l y 3.3x2.5 m m a n d 3.8x3.0 m m ,
respectively. F e m a l e s g r o w to a larger size t h a n males; the largest female w a s 125
m m S L , the largest m a l e 100 m m S L . G e n e r a l l y , h o w e v e r , the males are i n the r a n g e
85-95 m m S L , w h i l e females r e g u l a r l y reach a size o f 100 m m S L .
H a p l o c h r o m i s d i c h r o u r u s R e g a n , 1922
(figs 4 , 5 , 27-62, tables 5-9)
Paratilapia serranus (part): Boulenger, 1915: 334

Haplochromis dichrourus Regan, 1922:178, fig. 6; Greenwood & Barel, 1978:480-481, fig. 46
"Haplochromis" dichrourus; van Oijen et al., 1981:161.
Haplochromis dichrourus (part); Greenwood, 1967: 65-69, fig. 11 & 1974: 52.
Prognathochromis (P.) dichrourus (part); Greenwood: 1980:19.
Haplochromis pellegrini (part); Greenwood, 1962: 236-239.
D i a g n o s i s . A m e d i u m to m o d e r a t e l y large ( d e p e n d i n g o n the l o c a l i t y ; see b e l o w )

p i s c i v o r o u s species w i t h s t r o n g l y c u r v e d u n i c u s p i d teeth i n the outer r o w o f the o r a l
j a w s . L i v e c o l o r a t i o n o f b o t h sexes r a t h e r s i m i l a r ; f e m a l e s b a s i c a l l y c o l o u r e d as
males, h a v i n g black p e l v i c fins, c o l o u r e d m e d i a l fins (viz.; a n a l f i n b r i g h t r e d w i t h a n
o r a n g e e g g spot d o r s o - c a u d a l l y , a n d c a u d a l f i n b r i g h t r e d at least o n the v e n t r a l half)
a n d h e a d m a r k i n g s i n c l u d i n g a distinct l a c h r y m a l stripe. I n p r e s e r v e d specimens the
l i g h t l o w e r j a w contrasts s t r o n g l y w i t h the black i n t e r m a n d i b u l a r area a n d b r a n c h i o -
stegal m e m b r a n e .
N o t e s o n the B M ( N H ) collection of H. dichrourus

A p a r t f r o m t h e h o l o t y p e , G r e e n w o o d (1967) b a s e d h i s r e d e s c r i p t i o n o f H.
dichrourus o n s i x specimens (three f r o m U g a n d a a n d three f r o m Tanzania). O n e of
these ( B M ( N H ) 1 9 6 6 . 3 . 9 . 1 8 5 ) , a s e x u a l l y active m a l e c a u g h t o f f S o s w a I s l a n d (Tan-
z a n i a ) , i s n o w t h o u g h t to b e l o n g to a different species w h i c h is d e s c r i b e d b e l o w .
H o w e v e r , o n the basis o f this s p e c i m e n G r e e n w o o d (1967) d e s c r i b e d the l i v e m a l e
c o l o r a t i o n o f H. dichrourus. A s p e c i m e n f r o m Jinja ( B M ( N H ) 1966.3.9.186) d i v e r g e s
f r o m a l l other specimens i n its m o r e slender h e a d a n d b o d y shape a n d i n its v e r y
faint l a c h r y m a l stripe. A r a d i o g r a p h o f this s p e c i m e n r e v e a l e d that it has a n excep-
t i o n a l l y l o w s u p r a o c c i p i t a l crest a n d a h i g h n u m b e r of vertebrae. A s i t w a s i n c l u d e d
i n H. dichrourus o n the basis of its l i v e c o l o r a t i o n ( G r e e n w o o d , pers. c o m m . ) I consi-
d e r i t a n aberrant s p e c i m e n . A s p e c i m e n f r o m K a t e b o , U g a n d a ( B M ( N H ) 1966.3.
9.187), i.e. f r o m a l l specimens the o n e caught m o s t closely to the t y p e l o c a l i t y ( B u -
g a n g a , U g a n d a ) , is n o longer u n i m p a i r e d . Its h e a d has been dissected f o r the p r e p a -
r a t i o n o f skeletal m a t e r i a l . A p h o t o g r a p h of the w h o l e s p e c i m e n s h o w s that the h e a d
shape w a s c o m p a r a b l e to that of the h o l o t y p e . T h e p h o t o also s h o w s that the speci-
m e n h a d a short l a c h r y m a l stripe v e r y s i m i l a r to that of the h o l o t y p e i n the f i g u r e o f
R e g a n (1922)(see f i g . 26). Regrettably, measurements of this s p e c i m e n are n o t a v a i l -
able. W h i l e e x a m i n i n g the B M ( N H ) collections t w o m o r e specimens of H. dichrourus
w e r e f o u n d w h i c h w e r e not i n c l u d e d i n G r e e n w o o d ' s r e d e s c r i p t i o n ; a s p e c i m e n f r o m
G r a n t Bay, U g a n d a ( B M ( N H ) 1962.3.2.510) w a s f o u n d i n the c o l l e c t i o n o f H. pellegri-
ni R e g a n , 1922, a n d a s p e c i m e n f r o m Jinja w a s d i s c o v e r e d a m o n g s t u n c a t a l o g u e d
m a t e r i a l ( R M N H 30564).
H E S T m a t e r i a l of H. dichrourus came f r o m the M w a n z a G u l f area.
B a s e d o n differences i n m o r p h o l o g i c a l characters a n d p r e s e r v e d c o l o r a t i o n three
g e o g r a p h i c a l l y separate g r o u p s o f specimens c a n b e r e c o g n i z e d . T h e y w i l l b e de-
s c r i b e d as p o p u l a t i o n s of H. dichrourus. These groupes are: 1) Specimens f r o m U g a n -
d a ( i n c l u d i n g the h o l o t y p e ) 2) Specimens f r o m a beach N o r t h o f M a j i t a , T a n z a n i a , 3)
S p e c i m e n s f r o m the M w a n z a G u l f , Tanzania.
1. H a p l o c h r o m i s d i c h r o u r u s ' U g a n d a "
(figs 4 , 2 7 - 4 4 , 1 0 5 , tables 5-6).
Material.— Holotype, $, 113.9 mm, B M N H 1906.5.30.265, Lake Victoria near Buganga, Uganda,
E.Degen. Other material; 1 cf, 82.6 mm + 1 ex., 101.0 mm, B M N H 1966.3.9.188-9, Lake Victoria near
Karenia, Napoleon Gulf, Uganda, EAFRO; 1 cf, 91.1 mm, B M N H 1962.32.510, Lake Victoria, Grant
Bay, Uganda, EAFRO; 1$, 114.0 mm, R M N H 30564, Lake Victoria, near Jinja, Uganda, J. Kendall; 1
147.4 mm, B M N H 1966.3.9.186, Lake Victoria, off Golf Course near Jinja, Uganda, EAFRO; Skeletal
elements of ex. B M N H 1966.3.9.187, 124 mm (estimated from photograph), Lake Victoria near
Katebo, Uganda, EAFRO.
D e s c r i p t i o n . — Based o n the h o l o t y p e a n d f i v e other s p e c i m e n s , 82.6-147.4 m m

SL.
H a b i t u s . — A m e d i u m s i z e d , m o d e r a t e l y s l e n d e r species. D o r s a l h e a d p r o f i l e
g e n t l y c u r v e d f r o m d o r s a l f i n o r i g i n to just before the eye, w h e r e it is i n t e r r u p t e d b y
the p r o m i n e n t p r e m a x i l l a r y p e d i c e l t i p . P r e m a x i l l a s l i g h t l y e x p a n d e d m e d i a l l y i n
t w o specimens a n d s o m e w h a t b e a k e d i n the smallest s p e c i m e n . M o u t h m o d e r a t e l y
o b l i q u e , l i p s n o r m a l . T h e vertical t h r o u g h the c a u d a l tip o f the m a x i l l a r e a c h i n g o r
just p a s s i n g the rostral m a r g i n of the eye. L o w e r j a w a n d lateral snout o u t l i n e p r o -
gnathous. R o s t r a l o u t l i n e o f l o w e r j a w nearly straight, m e n t a l p r o m i n e n c e s m a l l o r
Fig. 26. Haplochromis dichrourus Regan, 1922; holotype, SL 113.9 mm. From Regan, 1922.
non-existent. L o w e r j a w sides s l i g h t l y o b l i q u e . R o s t r a l m a r g i n of v e r t i c a l a r m of pre-

o p e r c u l u m n e a r l y v e r t i c a l , the d o r s a l b o r d e r of the h o r i z o n t a l a r m h o r i z o n t a l o r
s l i g h t l y d e c l i n e d v e n t r a d . E y e n o r m a l , s l i g h t l y elongated i n r o s t r o - c a u d a l d i r e c t i o n .
P u p i l s i m i l a r l y e l o n g a t e d . C e p h a l i c lateral line pores not e n l a r g e d , the canals o n the
l a c h r y m a l not v i s i b l e .
S c a l e s . — C h e e k , gill-cover, d o r s a l h e a d surface a n d d o r s u m w i t h c y c l o i d scales.
F l a n k s a n d c a u d a l p e d u n c l e w i t h w e a k l y c t e n o i d scales. C h e s t a n d b e l l y w i t h a m i x -
ture of c y c l o i d (especially v e n t r a l l y ) a n d w e a k l y ctenoid scales. A g r a d u a l size transi-
t i o n b e t w e e n the r e l a t i v e l y s m a l l chest scales a n d the scales of the f l a n k . C a u d a l f i n
w i t h s m a l l elongate scales o n its p r o x i m a l half to 3/5.
F i n s . — P e c t o r a l a n d p e l v i c f i n s n o t q u i t e r e a c h i n g the o r i g i n o f the a n a l f i n .
D o r s a l a n d a n a l f i n not r e a c h i n g base of c a u d a l f i n . C a u d a l f i n o u t l i n e subtruncate,
s l i g h t l y emarginate.
G i l l a p p a r a t u s . — There are n i n e g i l l rakers o n the l o w e r half of the first g i l l arch
o n the left side. T h e s p e c i m e n of 91.1 m m S L has the l o w e r m o s t t w o r e d u c e d , n u m -
ber 3-5 s l e n d e r l y p o i n t e d , a n d 6-9 s l i g h t l y b r o a d e n e d w i t h a m o r e o r less b r o a d t i p .
T h e h o l o t y p e has the l o w e r m o s t f o u r r e d u c e d , c o n i c a l , the f i f t h e l o n g a t e c o n i c a l ,
n u m b e r six elongate a n d b r o a d e n e d a n d n u m b e r s 7-9 b r o a d e n e d w i t h a n e x p a n d e d
b i f i d o r t r i f i d h e a d . S p e c i m e n R M N H 30564 has 123 g i l l f i l a m e n t s o n the l a t e r a l
h e m i b r a n c h of the first g i l l arch f r o m the r i g h t side.
V i s c e r a . — Intestine l e n g t h of R M N H 30564 is 0.98 t i m e s its s t a n d a r d l e n g t h .
F o l l o w i n g the d e f i n i t i o n s of Z i h l e r (1982) the arrangement of the d i g e s t i v e tract of
this s p e c i m e n is of t y p e D ( w i t h backflap).
— O r a l teeth. A s the teeth are s t r o n g l y c u r v e d a n d d e e p l y e m b e d d e d i n the
m u c o s a , the tips of the teeth are barely v i s i b l e i n a lateral v i e w . H o w e v e r , i n the h o l o -
t y p e , p r o b a b l y d u e to s h r i n k a g e of the l i p tissue, they are v i s i b l e .
— Shape: N e a r l y a l l outer r o w teeth are acutely p o i n t e d , s t r o n g l y c u r v e d u n i c u s -
p i d s . R o s t r a l a n d lateral teeth r o u n d e d , postero-lateral teeth s o m e w h a t c o m p r e s s e d .
U p p e r j a w teeth g e n e r a l l y m o r e c u r v e d than teeth of the l o w e r jaw. In the h o l o t y p e
s o m e postero-lateral teeth are m o r e n e a r l y s h o u l d e r e d u n i c u s p i d s ; one of the lateral
l o w e r j a w teeth o n the r i g h t side is v e r y w e a k l y b i c u s p i d . G e n e r a l l y , the c o m p r e s s e d
teeth h a v e the c r o w n n a r r o w i n g m o r e abruptly, the sides of the c r o w n b e i n g some-
w h a t concave i n a l a b i a l v i e w . Tooth size g r a d u a l l y decreasing f r o m the s y m p h y s i s to
the c a u d a l e n d of the jaws. Inner r o w s w i t h a m i x t u r e of s l i g h t l y to moderately, c o m -
p r e s s e d s h o u l d e r e d u n i c u s p i d s a n d t r i c u s p i d s , the f o r m e r d o m i n a t i n g . A l l i n n e r
teeth are s t r o n g l y c u r v e d .
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r r o w covers near-
l y the w h o l e l e n g t h of the p r e m a x i l l a r y dentigerous a r m , the first i n n e r r o w about
2/3. T w o or three i n n e r r o w s i n the rostral quarter of the p r e m a x i l l a , decreasing to
zero i n the c a u d a l t h i r d . R o s t r a l l y a distinct gap b e t w e e n outer r o w a n d first i n n e r
row. I n l o w e r j a w the outer r o w e x t e n d i n g o n l y s l i g h t l y f u r t h e r c a u d a d t h a n the first
i n n e r r o w , just r e a c h i n g the c o r o n o i d w i n g . R o s t r a l l y a d i s t i n c t g a p b e t w e e n the
outer r o w a n d the first i n n e r r o w s .
— C o u n t s a n d setting: 55-62 teeth i n the outer r o w of the p r e m a x i l l a , 40-46 i n the
outer r o w of the l o w e r jaw. Teeth r e g u l a r l y set, neck-distance v a r y i n g b e t w e e n 1/2 to
n e a r l y e q u a l l i n g the n e c k w i d t h . Teeth c a u d a l l y i n b o t h jaws s l i g h t l y m o r e closely set.
A s m a l l m e d i a l g a p i n b o t h jaws.
Figs. 27-43. Haplochromis dichrourus "Uganda". Figs. 27 & 30. Premaxilla, lateral view. Figs. 28 & 31.
Premaxilla, view perpendicular to dentigerous area. Figs. 29 & 32. Medial view of premaxillary
ascending arm. Figs. 33 & 34. Right lower jaw, lateral view. Arrow in Fig. 33 indicates position of last
(22nd) outer row tooth. Fig. 35. Lower pharyngeal element, dorsal view. Fig. 36. Lower pharhyngeal
element, lateral view. Fig. 37. Lower pharhyngeal element, caudal view. Figs. 38-40. Premaxillary teeth,
labial and lateral views. Fig. 38. First tooth of outer row. Fig. 39. Seventh tooth of outer row. Fig. 40.
Fifth tooth of first inner row. Figs. 41-43. Lower parhyngeal teeth, loateral and rostral views. Fig. 41.
Seventh lateral tooth. Fig. 42. Fifth tooth caudalmost row. Fig. 43. First tooth of caudalmost row. Figs.
30-32 & 34 from specimen R M N H 30564, other figs, from B M N H 3.9.66.187. Scale of bony elements = 5
mm, scale of teeth = 1 mm.
Table 5. Linear and angular measurements of Haplochromis dichrourus Regan, 1922, and H. maisomei spec. nov. For convenience the ratios are multiplied by 100.
Haplochromis dichrourus Haplochromis maisomei
Locality Karenia Grant Bay Buganga Jinja Majita Maisome I. Standieri I.
Holotype Holotype
Museum BM(NH) BM(NH) BM(NH) RMNH BM(NH) BM(NH) RMNH RMNH RMNH
Reg.no. 1966.3.9.188-189 1962.3.2.510 1906.5.30.265 30564 1966.3.9.186 1966.3.9.183-184 30122 30121 30386
SLinmm 82.6 101.0 91.1 113.0 114.0 147.4 167.0 185.0 98.1 96.1 81.5
BD/SL 29.7 29.7 29.6 31.1 30.3 28.5 36.5 34.0 33.3 32.3 33.2
PFL/SL 23.8 22.7 24.3 24.8 25.0 22.3 24.9 23.4 25.2 23.7 24.6
CPL/SL 14.2 14.3 13.6 15.1 13.9 15.6 13.8 12.4 15.9 15.9 14.4
CPD/SL 10.8 11.7 12.0 11.0 10.7 10.9 12.3 11.4 11.9 12.1 12.2
CFL/SL 22.7 22.7 23.2 22.3 21.8 21.5 21.5 20.6 22.7 22.6 23.3
HL/SL 35.2 37.1 36.3 37.2 35.8 36.0 37.5 35.6 35.4 35.5 36.3
SnL/HL 31.9 35.4 32.3 35.1 34.4 35.3 39.7 40.0 33.9 32.1 33.1
SnW/HL 30.5 32.5 29.9 30.6 31.2 29.6 35.8 36.5 32.7 32-1 30.4
HW/HL 41.2 37.8 39.2 37.7 39.3 39.4 43.2 43.9 39.6 42.1 41.2
LOW/HL 17.5 18.1 18.4 19.2 19.5 17.6 21.5 22.5 19.6 19.8 19.2
POD/HL 23.3 24.0 24.1 22.5 24.6 21.6 25.5 25.7 25.3 25.7 27.7
IAW/HL 27.4 26.6 26.5 26.1 26.8 27.2 30.3 31.8 25.9 26.6 26.6
POW/HL 17.5 20.0 18.7 19.2 19.0 19.7 22.6 22.5 19.3 18.7 17.9
EyL/HL 24.3 22.4 24.7 20.6 22.1 23.1 18.6 18.1 25.6 26.3 23.6
ChD/HL 23.3 21.3 22.6 24.9 24.4 25.0 28.3 30.3 22.2 23-3 23.6
LJL/HL 51.5 48.5 51.0 49.4 51.1 53.9 50.2 52.2 47.8 49.1 45.9
LJW/HL 23.3 21.0 19.9 19.2 21.0 20.6 25.5 28.7 21.6 19-5 20.6
UJL/HL 40.8 40.2 41.6 38.2 41.0 41.7 44.9 45.1 35.6 35.6 35.8
PPL/HL 28.1 28.5 27.7 27.5 27.3 27.8 29.0 28.7 28.2 27.7 28.3
DHI 25° 30° 26° 32° 30° 28° 32° 28° 35° 35° 34°
PPI 32° 38° 32° 38° 38° 35° 38° 35° 35° 40° 41°
SnA 72° 76° 68° 73° 73° 73° 73° 74° 70° 72° 68°
GI 43° 43° 42° 39° 45° 45° 37° 39° 35° 34° 29°
VAN OIJEN: PISCIVOROUS HAPLOCHROMINE CICHLIDAE OF LAKE VICTORIA
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— I m p l a n t a t i o n . U p p e r jaw, outer r o w ; rostrally erect, laterally s o m e w h a t r e c u m -

bent. T h e lateral teeth i n c l i n i n g s o m e w h a t s u s p e n s o r i a d . L o w e r jaw, outer r o w ; erect.
T h e lateral teeth s o m e w h a t s u s p e n s o r i a d i n c l i n e d . In b o t h j a w s the i n n e r r o w teeth
i m p l a n t e d r e c u m b e n t to s t r o n g l y recumbent.
P h a r y n g e a l t e e t h . — C o u n t s : There are 21-24 teeth i n the c a u d a l m o s t transverse
r o w a n d 11-12 i n the t w o m e d i a n r o w s .
— Shape. M o s t p h a r y n g e a l teeth are r e l a t i v e l y fine a n d c o m p r e s s e d , a n d are of
the b e v e l l e d t y p e . T h e t w o or three c a u d a l m o s t teeth i n the t w o m e d i a n r o w s a n d
f o u r o r f i v e m e d i a l teeth i n the c a u d a l m o s t r o w of e a c h h a l f of the e l e m e n t are
e n l a r g e d , h o o k e d the b l u n t major cusps p o i n t i n g r o s t r o - d o r s a d . Tooth size increases
f r o m rostral to c a u d a l a n d f r o m lateral to m e d i a l .
— Osteology. T h e figures a n d d e s c r i p t i o n of the skeletal elements are based o n
B M ( N H ) 1966.3.9.187, S L c. 124 m m a n d R M N H 30564, S L 114 m m . A s the l o w e r p h a -
r y n g e a l element of the h o l o t y p e h a d already been dissected it c o u l d also be e x a m i n e d .
— N e u r o c r a n i u m . C o m p a r e d w i t h the n e u r o c r a n i u m of a g e n e r a l i z e d c i c h l i d (H.
elegans, Barel et a l . , 1976), the n e u r o c r a n i u m of H. dichrourus is less b r o a d a n d m o r e
d o r s o - v e n t r a l l y c o m p r e s s e d , b o t h the otic r e g i o n a n d the s u p r a o c c i p i t a l crest b e i n g
s h a l l o w e r . T h e e t h m o v o m e r i n e b l o c k is longer than i n H. elegans a n d less d e c u r v e d .
A s i n m o s t p i s c i v o r o u s c i c h l i d s f r o m L a k e V i c t o r i a the p h a r y n g e a l a p o p h y s i s is s i t u -
ated r e l a t i v e l y far caudally. T h e postorbital process is b r o a d .
— O r a l j a w s . P r e m a x i l l a d e n t i g e r o u s a r m l o n g e r (1.23-1.28 x) t h a n a s c e n d i n g
a r m . A n g l e b e t w e e n the t w o a r m s 79-84°. R o s t r a l part of d e n t i g e r o u s a r m of B M
( N H ) 1966.3.9.187 s l i g h t l y b e a k e d . Ventral o u t l i n e of dentigerous a r m concave. M a n -
d i b l e : A r e l a t i v e l y slender element, l e n g t h / d e p t h ratio 2.5-2.8. T h e c o r o n o i d process
i n s p e c i m e n B M ( N H ) 1966.2.3.187 is relatively short. Tooth b e a r i n g part of j a w slight-
l y m o r e t h a n half of the total j a w l e n g t h .
— L o w e r p h a r y n g e a l element. T h e l o w e r p h a r y n g e a l element longer than b r o a d
( L e n g t h / W i d t h = 1.1-1.2) a n d relatively shallow, the keel rather deep for a p i s c i v o r o u s
species. Dentigerous area o n l y slightly longer than b r o a d ( l e n g t h / w i d t h = 1.03-1.12).
— Vertebrae. There are 29-30 vertebrae, c o m p r i s i n g 13 a b d o m i n a l a n d 16-17 cau-
d a l elements. S p e c i m e n B M ( N H ) 1966.3.2.186 has 31 vertebrae.
C o l o r a t i o n . — L i t t l e i n f o r m a t i o n is available o n the c o l o r a t i o n of l i v e specimens.
W h e n R e g a n (1922) d e s c r i b e d H. dichrourus he n o t e d that the c o l o r a t i o n of the a n a l
f i n a n d the l o w e r half of the a n a l f i n w a s b r i g h t r e d . T h i s w a s sixteen years after the
s p e c i m e n h a d been caught. G r e e n w o o d (1974: 52) stated: " I n H. dichrourus, a l t h o u g h
the sexes are differently c o l o u r e d , the female is m o r e p o l y c h r o m a t i c t h a n the m a l e
(see G r e e n w o o d , 1967; these observations h a v e since been c o n f i r m e d a n d e x t e n d e d
b y a d d i t i o n a l s p e c i m e n s ) . " It s h o u l d be n o t e d that G r e e n w o o d (1967) e r r o n e o u s l y
b a s e d the d e s c r i p t i o n of l i v e m a l e c o l o r a t i o n of H. dichrourus o n a s p e c i m e n w h i c h
does not b e l o n g to this species (see p a g e 57).
— P r e s e r v e d c o l o r a t i o n . T h e h o l o t y p e , except for traces of h e a d m a r k i n g s , is n o w
a l m o s t c o m p l e t e l y colourless: H e a d a n d r o p e l i g h t b r o w n , g i l l cover w h i t i s h , b o d y
c r e a m y - w h i t e , area b e t w e e n l o w e r j a w s a n d g i l l - m e m b r a n e sooty. D o r s a l , a n a l a n d
p e c t o r a l fins c r e a m y - w h i t e . C a u d a l b r o w n i s h basally, g r a d u a l l y b e c o m i n g h y a l i n e
distally. P e l v i c s l i g h t basally, b l a c k i s h distally. N o trace of o c e l l i o n the anal. H e a d
m a r k i n g s : There is o n l y a v e r y faint bar left i n the r e g i o n w h e r e , i n the d r a w i n g of
R e g a n (1922), there is a b r o a d l a c h r y m a l stripe (fig. 26). A n o s t r i l stripe r u n s f r o m
each n o s t r i l r o s t r o - m e d i a d b u t it is i n t e r r u p t e d i n the area of the p r e m a x i l l a r y p e d i -
cel. Faint traces o f the starting p o i n t s o f the interorbital stripe near the eyes, a n d of
a n e x t e n d e d l a c h r y m a l stripe o n the eye a n d just a b o v e it. T h e area of the o p e r c u l a r
blotch is darkened.
T h e f o u r other s p e c i m e n s h a v e retained m o r e of their h e a d m a r k i n g s . I n their
preserved coloration n o differences are apparent between males a n d females.
G r o u n d c o l o u r of these specimens i s d a r k b r o w n i s h , g i l l - c o v e r silver-grey. A r e a be-
t w e e n the r i g h t a n d left side o f l o w e r jaw, a n d g i l l - m e m b r a n e black. T h e t w o speci-
m e n s f r o m K a r e n i a h a v e traces of three o r f o u r vertical b a n d s o n the f l a n k . H e a d -
m a r k i n g s : A distinct, b r o a d l a c h r y m a l stripe r u n n i n g f r o m the rostro-ventral m a r g i n
of the eye b e h i n d the c a u d a l tip o f the m a x i l l a to the h o r i z o n t a l a r m of the preoper-
c u l u m . T h e l a c h r y m a l stripe is extended d o r s a l l y a b o v e the eye. T h e n o s t r i l stripe is
c o n t i n u o u s i n o n l y t w o o f the specimens. O f the interorbital stripe o n l y the lateral
ends are present. A preopercular vertical stripe is present i n t w o specimens one of
w h i c h has also a b l a c k interopercular area.
F i n s . — d o r s a l , pectoral a n d c a u d a l b r o w n i s h . I n t w o s p e c i m e n s the v e n t r a l half
of the c a u d a l d i s t i n c t l y lighter than the d o r s a l part. A n a l f i n lighter b r o w n w i t h one
o r t w o d a r k - r i m m e d spots caudally. Pelvics black.
E c o l o g y . — Since s o f e w specimens are available a n d n o details are k n o w n about
the localities a n d techniques of capture, little can b e s a i d about the ecology o f these
specimens. D a t a f r o m s t o m a c h content investigations b y G r e e n w o o d (1967) m a k e i t
apparent that H. dichrourus " U g a n d a " is a p i s c i v o r o u s species. F r o m the presence of a
s m a l l j u v e n i l e ( ± 1 0 m m T L ) i n the m o u t h o f the h o l o t y p e , w h i c h I consider to b e a
spent female, it m a y be d e d u c e d that the species is a female m o u t h b r o o d e r .
2. H a p l o c h r o m i s d i c h r o u r u s " M a j i t a "
(figs 4 5 , 4 6 , 1 0 6 , tables 5-6).
Material.— 1 cr, 167 mm + 1 specimen (sex unknown)., 185 mm, B M N H 1966.3.9.183-4, Lake Victoria,
beach North of Majita, Tanzania, EAFRO.
D e s c r i p t i o n . — H a b i t u s . T h e s p e c i m e n s of H. dichrourus " M a j i t a " are r e l a t i v e l y

d e e p - b o d i e d a n d r e l a t i v e l y b r o a d . D o r s a l h e a d p r o f i l e gently c u r v e d f r o m o r i g i n o f
d o r s a l f i n to interorbital area, f r o m w h e r e the p r o f i l e is i n t e r r u p t e d b y the p r o m i n e n t
p r e m a x i l l a r y p e d i c e l . S n o u t s o m e w h a t b l u n t . M o u t h m o d e r a t e l y o b l i q u e . L i p s rather
b r o a d . P r e m a x i l l a not e x p a n d e d m e d i a l l y or b e a k e d . T h e e x p o s e d part of the m a x i l l a
s m a l l i n the smaller s p e c i m e n , rather large i n the larger s p e c i m e n ; u p p e r l i p t a p e r i n g
c a u d a d i n this s p e c i m e n . T h e vertical t h r o u g h the c a u d a l e n d of the m a x i l l a just rea-
c h i n g o r p a s s i n g the rostral eye m a r g i n . L o w e r j a w isognathous, s l i g h t l y p r o t r u d i n g .
R o s t r a l o u t l i n e of l o w e r j a w straight o r s l i g h t l y r o u n d e d , n o m e n t a l p r o m i n e n c e .
L a t e r a l sides o f l o w e r j a w relatively b r o a d , m o d e r a t e l y o b l i q u e . Vertical a r m of pre-
o p e r c u l u m n e a r l y v e r t i c a l , the h o r i z o n t a l a r m d e c l i n i n g s l i g h t l y v e n t r a d . E y e rela-
t i v e l y s m a l l , s l i g h t l y elongate, p u p i l (because o f p r e s e r v a t i o n o n l y f a i n t l y v i s i b l e )
s e e m s c i r c u l a r . C e p h a l i c l a t e r a l l i n e o p e n i n g s r e l a t i v e l y s m a l l , t h e c a n a l s o n the
lachrymal faintly visible.
S c a l e s . — A s i n U g a n d a specimens.
F i n s . — Pectoral shorter than p e l v i c f i n , the p e l v i c s just r e a c h i n g the a n a l f i n . I n
the s m a l l e r s p e c i m e n , posterior t i p of d o r s a l not r e a c h i n g the c a u d a l f i n . A n a l f i n of
s m a l l e r s p e c i m e n a n d d o r s a l a n d a n a l of the larger s p e c i m e n r e a c h i n g the base o f the

c a u d a l f i n . First p e l v i c f i n r a y p r o d u c e d i n b o t h specimens. C a u d a l f i n o u t l i n e w i t h
r o u n d e d angles; s l i g h t l y e m a r g i n a t e , p o s t e r i o r m a r g i n v e r y s l i g h t l y o b l i q u e i n the
smaller specimen.
G i l l r a k e r s . — E i g h t o r n i n e g i l l rakers o n the l o w e r p a r t o f the first g i l l a r c h of the
left s i d e . T h e l o w e r m o s t o n e r e d u c e d , c o n i c a l , t h e f o l l o w i n g three r a k e r s s h o r t ,
b r o a d l y f l a t t e n e d , a n d the u p p e r f o u r o r f i v e also r e l a t i v e l y s h o r t w i t h e x p a n d e d
Fig. 44. Catch localities of specimens of Haplochromis dichrourus Regan, H. maisomei spec, nov., and H.
kujunjui spec. nov. in Lake Victoria.
heads; s i m p l e , b i f i d o r t r i f i d .
V i s c e r a . — N o i n f o r m a t i o n is a v a i l a b l e o n the l e n g t h a n d c o n f i g u r a t i o n o f the
intestines.
O r a l t e e t h . — Shape: In b o t h specimens the teeth i n the outer r o w s o f b o t h j a w s
are a l l r e l a t i v e l y stout, r o u n d e d a n d acutely p o i n t e d u n i c u s p i d s . T h e w h o l e t o o t h
m o d e r a t e l y to s t r o n g l y c u r v e d , teeth of u p p e r j a w m o r e s t r o n g l y c u r v e d o v e r the d i s -
tal half. Teeth e m b e d d e d i n a thick m u c o s a thus g i v i n g the i m p r e s s i o n that they are
rather s m a l l . Tooth size g r a d u a l l y decreases f r o m rostral to c a u d a l . T h e teeth of the
i n n e r r o w s are s m a l l e r replicas of the outer r o w teeth. Teeth of first i n n e r r o w rela-
t i v e l y large.
— D e n t a l arcade a n d t o o t h b a n d : D e n t a l arcade r o u n d e d (to s o m e w h a t acute i n
l o w e r j a w ) . Distance b e t w e e n outer r o w a n d first i n n e r r o w s l i g h t l y larger t h a n d i s -
tance b e t w e e n i n n e r r o w s . There are three o r f o u r i n n e r r o w s r o s t r a l l y i n the l o w e r
j a w a n d f o u r i n n e r r o w s rostrally i n the u p p e r j a w decreasing to z e r o i n the c a u d a l
part of the jaws.
— O u t e r r o w o c c u p y i n g 4/5 of the p r e m a x i l l a r y dentigerous a r m a n d r e a c h i n g
the c o r o n o i d w i n g i n the l o w e r j a w A s m a l l m e d i a l g a p i n b o t h jaws.
— C o u n t s a n d setting. I n smaller s p e c i m e n a p p r o x i m a t e l y 66 teeth i n the outer
r o w of the u p p e r jaw, 76 i n the larger s p e c i m e n . L o w e r j a w of the smaller specimens
w i t h 49 teeth i n the outer r o w , the l o w e r j a w of the larger s p e c i m e n w i t h at least 42.
O u t e r r o w teeth r e g u l a r l y a n d rather closely set at a distance less t h a n the t o o t h d i a -
meter at its base.
— I m p l a n t a t i o n . O u t e r r o w teeth i n the u p p e r j a w erect ( m e d i a l l y ) to s l i g h t l y re-
c u m b e n t (caudally). O u t e r r o w teeth i n the l o w e r j a w s l i g h t l y p r o c u m b e n t ( m e d i a l l y )
to erect (laterally). Inner r o w teeth i n u p p e r j a w are r e c u m b e n t to s t r o n g l y r e c u m -
bent. I n the l o w e r j a w the teeth of the first i n n e r r o w erect, the other ones s l i g h t l y to
m o d e r a t e l y recumbent.
P h a r y n g e a l t e e t h . — C o u n t s : A b o u t 20 teeth i n the c a u d a l m o s t r o w a n d 12-14 i n
the m e d i a n series.
— Shape. M o s t teeth are r e l a t i v e l y stout, b l u n t l y p o i n t e d teeth of the b e v e l l e d
type. Three o r f o u r c a u d a l m o s t teeth i n the m e d i a n r o w a n d the three m e d i a l m o s t
teeth of the c a u d a l r o w are e n l a r g e d , h o o k e d teeth w i t h a b l u n t r o s t r o - d o r s a l directed
major c u s p . T o o t h size increases f r o m rostral to c a u d a l a n d f r o m lateral to m e d i a l .
O s t e o l o g y . — A s n o skeletal m a t e r i a l is available, little is k n o w n about the shape
of the skeletal elements. H o w e v e r , i n b o t h specimens the l o w e r p h a r y n g e a l element
h a d already been cut loose a n d c a n therefore be d e s c r i b e d .
— L o w e r p h a r y n g e a l e l e m e n t rather s h a l l o w , v e r y s l i g h t l y l o n g e r t h a n b r o a d
( l e n g t h / w i d t h = 1.05-1.11). D e n t i g e r o u s area as l o n g as b r o a d . H o r n s m a k i n g a n a n g -
le o f c. 3 2 ° w i t h the m e d i a n . Suture straight.
— Vertebrae. B o t h specimens h a v e 30 vertebrae c o m p r i s i n g 13 a b d o m i n a l a n d 17
c a u d a l elements.
C o l o r a t i o n . — There is n o i n f o r m a t i o n available o n the l i v e c o l o r a t i o n of the t w o
specimens.
— C o l o r a t i o n of preserved male(s?): H e a d a n d b o d y w i t h a d a r k b r o w n i s h
g r o u n d colour. C h e s t d a r k (blackish) ventrally, b e l l y black. D o r s a l h e a d area, interor-
b i t a l area, d o r s a l part of snout a n d p r e o p e r c u l a r h o r i z o n t a l a r m , i n t e r o p e r c u l u m a n d
sides of j a w s lighter t h a n s u r r o u n d i n g areas. L i g h t l o w e r j a w a n d p r e o p e r c u l a r area
Figs 45-46. H. dichrourus "Majita", lower pharhyngeal bone. Fig. 45. Dorsal view. Fig. 46. Caudal view.
Scale = 5 mm.
strongly contrasting w i t h black branchiostegal membrane a n d the black inter

m a n d i b u l a r area. H e a d m a r k i n g s : A distinct l a c h r y m a l stripe r u n n i n g f r o m the ros-
t r o - v e n t r a l m a r g i n o f the eye b e h i n d the e n d o f the m a x i l l a to the rostral e n d o f the
p r e o p e r c u l u m ; a n extension of the l a c h r y m a l stripe above the eye; a faint nape
b l o t c h ; a distinct b u t i n t e r r u p t e d interorbital stripe a n d a c o n t i n u o u s n o s t r i l stripe.
T h e n o s t r i l stripe is broader m e d i a l l y . R o s t r o - v e n t r a l l y to the t w o ends e q u a l l y d i s
tinct d a r k spots are present o n the snout. H o w e v e r , these spots are not connected w i t h
the n o s t r i l stripe. A n o p e r c u l a r b l o t c h is present. O n the b o d y of the smaller s p e c i m e n
a faint m i d l a t e r a l b a n d can be seen, whereas o n the b o d y of the larger s p e c i m e n traces
of 4 o r 5 vertical b a n d s are present. Fins: except for the black p e l v i c s , a l l fins w i t h a
u n i f o r m l y l i g h t b r o w n i s h colour. T h e d o r s a l part of the c a u d a l f i n o n l y s l i g h t l y d a r k e r
t h a n the v e n t r a l part. I n a s l i g h t l y d a r k e r area d o r s o - c a u d a l o n the anal f i n one l i g h t
ocellus i n the smaller s p e c i m e n a n d t w o l i g h t o c e l l i i n the larger s p e c i m e n . B o t h speci
m e n s w i t h s m a l l l i g h t spots b e t w e e n the last rays of the d o r s a l f i n .
E c o l o g y . — D e t a i l e d i n f o r m a t i o n o n the precise catch l o c a l i t y or habitat is l a c k i n g .
— F o o d . F o r h i s r e d e s c r i p t i o n o f H. dichrourus, G r e e n w o o d (1967) i n v e s t i g a t e d
the stomachs of specimens f r o m f o u r different localities i n c l u d i n g one f r o m M a j i t a .
A l l s p e c i m e n s w e r e f o u n d to h a v e c i c h l i d remains i n their stomachs.
— B r e e d i n g a n d G r o w t h . T h e 167 m m S L s p e c i m e n is a s e x u a l l y active m a l e . T h e
g o n a d s o f t h e o t h e r (185.0 m m S L ) s p e c i m e n h a v e b e e n r e m o v e d a l o n g w i t h t h e
intestines. T h e sex o f this s p e c i m e n therefore is i n d e t e r m i n a b l e .
3. Haplochromis dichrourus " M w a n z a "

(figs 5 , 2 1 - 2 8 , 8 3 , 1 0 7 , 1 0 8 , tables. 7-9).
Material.— Material from the Mwanza Gulf collected by G.Ch. Anker & C . D . N . Barel; 1 cf, 115.0 mm,
B M N H 1977.1.28.32, 9.vi.l975; 2 99, 104.0, 109.0 mm, R M N H 30061-62, 9.vi.l975; 1 cf, 95.5 mm,
R M N H 30063, 24.vi.1975. Material from the Mwanza Gulf collected by HEST; 1 cf, 98.0, R M N H
30064, 7.X.1977; 1 9, 136.8 mm, R M N H 30065,4.xi.l977; 1 9, 124.8 mm, R M N H 30066,18.xi.1977; 2 99,
111.7,122.5 mm, R M N H 30067-68, 8.xii.l977; 2 99,100.9,117.6 mm, R M N H 30069-70, 30.xii.1977; 1 9,
109.3 mm, R M N H 3 0 0 7 1 , 6 . Π 9 7 8 ; 1 cf, 107.0 mm, M N H N 1987-1667,5.V.1978; 1 9,131.5 mm, R M N H
30090, 19.V.1979; 6 99, 94.5-136.2 mm, R M N H 30073-78, 31.V.1978; 1 9, 94.1 mm, R M N H 30079,
21.vii.1978; 1 cf, 86.3 mm, R M N H 30080, ll.viii.1978; 1 9,132.0 mm R M N H 30060,18.viii.1978; 2 99,
128.0, 141.0 mm, R M N H 30081-82, 18.viii.1978; 2 cfcf, 95.1, 106 mm + 3 99, 114.2-126.8 mm, R M N H
30081-89, 1978; 1 9, 119.6 mm, ANSP 168242, 1978; 1 9, 107.5 mm. B M N H 1988.2.4.20, 5.V.1978; 1 cf,
117.5 mm, R M N H 30091, 19.Í.1979; 3 99, 111.7-137.0 mm, R M N H 30092-94, 4.V.1979; 1 juv., 70.5 mm,
R M N H 30095, 7.ix.l979; 1 9,108.9 mm, B M N H 1988 ?? , 5.iv.l980; 1 9,108.2 mm, M N H N 1987-1667,
21.iv.1980; 2 99,118.3-126.8 mm, R M N H 30098-99, 21.iv.1980; 2 99,111.8, 122.5 mm, R M N H 30100-01,
22.iv.1980; 1 9,121.8 mm, R M N H 30102, 23.iv.1980; 1 cf, 96.5 mm, R M N H 30103, 22.1.1981; 2 99,97.1-
122.2 mm, R M N H 30104-05, 26.1.1981; 2 cfcf, 96.3-98.0 mm, R M N H 30106-07, ll.xi.1981; 1 cf, 93.0
mm, R M N H 30108, 24.viii.1983; 2 cfcf, 88.9, 90.2 m m + 4 99, 86.9-119.1 mm, R M N H 30109-14,
21.ix.l983;l cf, 111.5 mm + 1 9,109.2 mm, R M N H 30116-17, 4.vi.l985.
D e s c r i p t i o n . — Based o n 57 specimens of 70.5-141.0 m m S L .

H a b i t u s . — B o d y m o d e r a t e l y slender. D o r s a l h e a d p r o f i l e s l i g h t l y c o n v e x , the
p r e m a x i l l a r y p e d i c e l s l i g h t l y to noticeably p r o m i n e n t . P r e m a x i l l a s l i g h t l y e x p a n d e d
m e d i a l l y , n o t b e a k e d . L i p s n o r m a l . A s m a l l part o f the m a x i l l a v i s i b l e c a u d a l l y a n d
d o r s o - c a u d a l l y of the p r e m a x i l l a . T h e v e r t i c a l t h r o u g h the c a u d a l t i p of the m a x i l l a
n o t r e a c h i n g t h e e y e i n m o s t s p e c i m e n s . M o u t h m o d e r a t e l y o b l i q u e , lateral s n o u t
o u t l i n e s l i g h t l y p r o g n a t h o u s , the l o w e r j a w p r o t r u d i n g . M e n t a l area r o u n d e d i n m o s t
specimens, i n s o m e w i t h a s m a l l p r o m i n e n c e . L a t e r a l sides of l o w e r j a w o n l y s l i g h t l y
oblique. H o r i z o n t a l a r m of preopercular h o r i z o n t a l , the vertical a r m vertical o r
i n c l i n i n g s l i g h t l y v e n t r a d rostrally. L a t e r a l l i n e canals a n d o p e n i n g s o n l a c h r y m a l n o t
e n l a r g e d . E y e a n d p u p i l subcircular to s l i g h t l y elongated i n a h o r i z o n t a l d i r e c t i o n .
S c a l e s . — A s i n U g a n d a specimens.
F i n s . — A s i n U g a n d a specimens.
G i l l a p p a r a t u s . — G i l l rakers h a v e been c o u n t e d i n 15 specimens. N u m b e r o f g i l l
rakers o n l o w e r part of the first g i l l arch o n r i g h t s i d e 8-9. L o w e r 3-4 r e d u c e d , c o n i -
c a l , n u m b e r s 4-5 elongate w i t h a c o n i c a l h e a d , a n d the u p p e r 3 o r 4 elongate a n d
e x p a n d e d w i t h b i f i d , t r i f i d o r m o r e c o m p l i c a t e d tips. A b o u t 132 g i l l filaments o n the
first h e m i b r a n c h .
V i s c e r a . — Intestine l e n g t h o f six e x a m i n e d specimens ranges f r o m 1.0-1.4 times
the s t a n d a r d l e n g t h . F o l l o w i n g the d e f i n i t i o n s of Z i h l e r (1982) the arrangement of the
d i g e s t i v e track o f a d u l t s o f M w a n z a G u l f s p e c i m e n s o f H. dichrourus is o f t y p e D
(with backflap).
O r a l t e e t h . — Shape. I n the outer r o w s a l l specimens, e v e n the smallest ones, have
o n l y slender, acutely p o i n t e d u n i c u s p i d s , r o u n d e d i n the rostral a n d lateral parts of
the j a w s , b u t c a u d o - l a t e r a l l y s o m e w h a t c o m p r e s s e d . I n t h e c o m p r e s s e d teeth the
sides o f t h e c r o w n a r e s o m e w h a t c o n c a v e i n l a b i a l v i e w . A l l teeth m o d e r a t e l y to
s t r o n g l y c u r v e d . U p p e r j a w teeth g e n e r a l l y m o r e s t r o n g l y c u r v e d t h a n l o w e r j a w
teeth. I n c o m p a r i s o n w i t h other p i s c i v o r e s the teeth are of a m o d e r a t e size. T h e size
of the teeth is n o t easily to d e t e r m i n e i n intact specimens as they are e m b e d d e d i n a
t h i c k m u c o s a . T o o t h size g r a d u a l l y decreasing f r o m rostral to c a u d a l . Inner r o w s i n
s m a l l e r specimens consist of a m i x t u r e of s l i g h t l y to m o d e r a t e l y c o m p r e s s e d s h o u l -
dered u n i c u s p i d s a n d tricuspids, the former d o m i n a t i n g . I n larger specimens a l l
teeth o f t h e i n n e r r o w s are r o u n d e d , slender, c o n i c a l , a c u t e l y p o i n t e d u n i c u s p i d s ,
s m a l l replicas of the outer r o w teeth. A l l i n n e r r o w teeth s t r o n g l y c u r v e d .
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r tooth r o w i n pre-
m a x i l l a covers w h o l e l e n g t h o f dentigerous a r m , i n n e r r o w s about t w o t h i r d s . T w o or
three i n n e r r o w s i n r o s t r a l p a r t o f p r e m a x i l l a , d e c r e a s i n g to z e r o i n c a u d a l p a r t .
R o s t r a l l y a distinct g a p b e t w e e n outer r o w a n d first i n n e r r o w . I n the l o w e r j a w the
outer r o w proceeds s l i g h t l y further c a u d a d than the inner r o w , a n d just reaches the

c o r o n o i d w i n g . T w o r o w s of i n n e r teeth i n the l o w e r j a w . R o s t r a l l y a d i s t i n c t g a p
b e t w e e n the outer r o w a n d the first i n n e r r o w .
— C o u n t s a n d setting. There are 54-64 teeth i n outer r o w of p r e m a x i l l a , a n d 36-44
i n outer r o w of l o w e r jaw. Teeth r e g u l a r l y a n d rather closely set, the neck-distance
v a r y i n g b e t w e e n less t h a n 1 /4 to n e a r l y e q u a l l i n g the neck diameter. A s m a l l m e d i a l
g a p i n b o t h jaws.
— I m p l a n t a t i o n . O u t e r r o w teeth i n u p p e r j a w erect to s l i g h t l y r e c u m b e n t rostral
ly, g r a d u a l l y m e r g i n g i n t o s t r o n g l y recumbent c a u d a l l y ; lateral teeth s o m e w h a t sus-
p e n d o r i a d i n c l i n e d . O u t e r r o w teeth i n l o w e r j a w erect r o s t r a l l y a n d laterally, s l i g h t l y
r e c u m b e n t c a u d o - l a t e r a l l y . L o w e r j a w teeth r e l a t i v e l y m o r e s u s p e n s o r i a d i n c l i n e d
t h a n teeth of u p p e r jaw. Inner r o w s ; i n b o t h jaws the i n n e r r o w teeth are m o d e r a t e l y
to s t r o n g l y recumbent.
P h a r y n g e a l teeth . — C o u n t s . There are 18-24 teeth i n the c a u d a l m o s t transverse
r o w a n d 10-12 i n the t w o m e d i a n r o w s .
— Shape. M o s t p h a r y n g e a l teeth are relatively fine a n d flattened a n d are of the
b e v e l l e d type. T h e t w o o r three c a u d a l m o s t teeth i n the t w o m e d i a n r o w s a n d about
f i v e m e d i a l teeth i n the c a u d a l m o s t r o w o n each side e n l a r g e d , stout, b e v e l l e d teeth
w i t h a large a n d b l u n t major c u s p p o i n t i n g r o s t r o - d o r s a d o r d o r s o - r o s t r a d . T o o t h
size i n c r e a s i n g f r o m rostral to c a u d a l a n d f r o m lateral to m e d i a l .
O s t e o l o g y . — O s t e o l o g i c a l descriptions are based o n skeletal elements of 3 speci
m e n s ( R M N H 30076, 30078 a n d 30079, S L 94.1-136.2 m m ) . D e s c r i p t i o n s of o r a l teeth
b a s e d o n a l l specimens.
— O r a l j a w s . P r e m a x i l l a d e n t i g e r o u s a r m l o n g e r (1.25x) t h a n a s c e n d i n g a r m s ,
angle b e t w e e n the arms 79-88°. R o s t r a l part of dentigerous area s l i g h t l y e x p a n d e d
r o s t r a d , v e n t r a l o u t l i n e of dentigerous area s l i g h t l y concave. L o w e r j a w m o d e r a t e l y
slender, l e n g t h / d e p t h ratio 2.3. L e n g t h of tooth-bearing part of the d e n t a r y is 0.4 χ
lower j a w length.
— L o w e r p h a r y n g e a l element r e l a t i v e l y s h a l l o w w i t h the keel area w e l l d e v e l
o p e d . T h e element is s l i g h t l y longer than b r o a d ( l e n g t h / w i d t h = 1.16), dentigerous
area as l o n g as b r o a d .
— Vertebrae. There are 29 vertebrae c o m p r i s i n g 13 a b d o m i n a l a n d 16 c a u d a l ele
ments (n = 8).
C o l o r a t i o n . — M w a n z a G u l f s p e c i m e n s of H. dichrourus c a u g h t b y H E S T h a v e
c o n f i r m e d the c o l o u r d e s c r i p t i o n for m a l e H. dichrourus as g i v e n i n G r e e n w o o d &
Barel (1978) a l t h o u g h the r e d f l u s h i n the d o r s a l f i n m a y be m u c h m o r e distinct t h a n
w a s p r e v i o u s l y d e s c r i b e d . T h e r e d c o l o u r s of the c a u d a l a n d a n a l fins are u s u a l l y
b r i g h t e r t h a n those of the d o r s a l f i n . Females of H. dichrourus are less p o l y c h r o m a t i c
a n d b r i g h t l y c o l o u r e d t h a n males, a l t h o u g h m u c h m o r e c o l o u r f u l than the females of
m o s t L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d species.
— L i v e c o l o u r s of s e x u a l l y active males. H e a d : L o w e r j a w a n d lateral aspects of
s n o u t green, cheek a n d v e n t r a l part of o p e r c u l u m d a r k green, v e n t r a l side of h e a d
a n d b r a n c h i o s t e g a l m e m b r a n e d a r k green to black. D o r s a l part of o p e r c u l u m , d o r s a l
part of s n o u t a n d d o r s a l h e a d surface d a r k b l o o d r e d . E y e : iris black, w i t h a t h i n s i l
v e r y i n n e r r i n g . M a r k i n g s : a b r o a d , v e n t r a l l y s o m e w h a t i n d i s t i n c t l a c h r y m a l stripe
r u n n i n g f r o m the r o s t r o - v e n t r a l m a r g i n of the eye b e h i n d the m o u t h c o r n e r to the
h o r i z o n t a l a r m of the p r e o p e r c u l u m . T h e l a c h r y m a l stripe is e x t e n d e d t h r o u g h a n d
Figs 47-59. Haplochromis dichrourus "Mwanza". Fig. 47. Premaxilla, lateral view. Fig. 48. Premaxilla,
view perpendicular to dentigerous area. Fig. 49. Premaxilla, medial view of ascending arm. Fig. 50.
Lower jaw, lateral view. Fig. 51. Lower pharyngeal element, dorsal view. Fig. 52. Lower pharhyngeal
labial and lateral view. Fig. 54. Second tooth of outer row. Fig. 55. Fourteenth tooth of outer row. Fig.
55. Fifth tooth of first inner row. Figs. 57-59. Lower pharhyngeal teeth, lateral and rostral view. Fig. 57.
Tenth lateral tooth. Fig. 58. Nineth medial tooth caudalmost row. Fig. 59. Second medial tooth of cau-
dalmost row. Scale of bony elements = 5 mm, scale of teeth = 1 mm.
a b o v e the e y e , the stripes of b o t h sides m e e t i n g m e d i a l l y . S n o u t c r o s s e d b y a b r o a d

n o s t r i l s t r i p e a n d a n e q u a l l y b r o a d i n t e r o r b i t a l s t r i p e r u n n i n g just d o r s a l l y to t h e p r e -
m a x i l l a r y p e d i c e l t i p . I n a l l b u t o n e s p e c i m e n the n o s t r i l s t r i p e l a t e r a l l y is e x t e n d e d
i n a r o s t r a l d i r e c t i o n g i v i n g this " s t r i p e " the " M " s h a p e w h i c h is c h a r a c t e r i s t i c f o r the
s p e c i m e n s f r o m the M w a n z a G u l f (figs. 28, 53). H a l f w a y b e t w e e n the d o r s a l m a r g i n
t
Table 7. Ranges of linear measurements of Haplochromis dichrourus "Mwanza" Regan, 1922. For convenience the ratios are multiplied by 100.
SLinmm 81.7-88.9 90.2-98.0 100.9-109.3 110.9-119.1 121.2-128.0 132.0-141.0

Ν 5 14 9 11 3 6
BD/SL 31.8-35.9 30.4-33.9 31.1-35.3 31.5-36.9 33.2-36.2 33.3-37.4

PFL/SL 25.6-29.3 25.1-29.4 26.4-28.8 255-29.8 25.6-28.5 26.7-28.0
CPL/SL 14.1-16.0 13.7-18.7 14.4-17.2 12.9-16.1 13.1-16.8 13.3-15.1
CPD/SL 11.5-12.3 10.8-12.4 10.9-13.3 10.4-12.5 11.1-12.6 11.2-12.6 Ν
CFL/SL 23.4-25.3 23.1-25.4 22.1-25.6 22.8-25.3 21.7-24.7 22.0-23.0
1
HL/SL 35.7-38.4 34.8-38.9 36.1-39.5 34.4-38.4 35.3-38.3 35.8-37.6 Γ"
8
Ο
SnL/HL 30.4-33.5 29.2-34.3 31.6-35.8 31.8-35.5 32.4-36.8 34.8-36.0
SnW/HL 28.3-33.8 28.1-33.8 29.9-33.5 30.1-34.8 30.0-35.0 31.8-35.7
Gisa
HW/HL 40.0-43.8 395-44.8 40.4-44.7 40.8-45.5 40.8-44.8 41.3^5.0

IOW/HL 19.8-21.0 18.7-22.3 18.7-22.0 20.2-23.8 20.0-23.4 21.7-23.7 <
POW/HL 24.6-26.8 24.3-27.1 25.3-28.1 25.3-28.5 25.0-27.7 24.5-28.5 73
m
LaW/HL 27.5-29.1 23.5-28.7 25.7-28.4 25.8-30.6 26.1-30.7 26.8-30.9 >
POD/HL 16.5-18.5 17.4-19.4 17.9-23.2 17.5-20.2 17.8-20.0 17.9-20.1
EyL/HL 21.5-26.3 20.9-25.6 21.4-24.1 20.5-26.1 21.4-25.6 19.4-22.4 m
r
ChD/HL 21.8-23.1 21.2-24.2 22.4-25.3 23.0-27.9 23.7-26.8 23.1-28.9 Ζ
LJL/HL 47.9-50.6 46.2-52.5 50.2-53.1 49.8-53.0 49.8-53.2 51.1-56.8 O
m
LJW/HL 19.8-23.3 18.6-23.6 19.5-23.8 20.9-26.8 21.2-26.2 21.4-25.1 Ζ
UJL/HL 37.5^0.3 36.6-41.8 37.7-415 38.9^11.7 38.3-42.3 39.6-45.3
PPL/HL 26.9-29.4 25.0-30.4 26.8-29.6 265-30.3 26.9-32.4 26.6-28.1
(1991
Table 8. Means and standard deviations of linear measurements of Haplochromis dichrourus "Mwanza" Regan, 1922. For convenience the ratios are multiplied by 100.
SL in mm 70.5 81.7-88.9 90.2-98.0 100.9-109.3 110.9-119.1 121.2-128. 132.0-141.0 ζ

Ν 1 5 14 9 11 13 6 g
m
Ζ
BD/SL 31.0 32.7±1.4 32.411.1 32.611.9 34.611.4 34.210.8 35.0115
PFL/SL 24.4 26.9±1.8 27.111.3 27.310.8 27.511.1 26.810.9 27.210.5 c/5
CPL/SL 16.7 15.5±0.9 14.911.1 15.210.8 14.410.8 14.711.0 14.310.6 η
CPD/SL 12.1 11.8±0.2 11510.4 11.610.7 11.810.6 11.610.4 11.910.5 <
ο
CFL/SL 24.6 24.410.5 24.310.7 23.711.0 23.910.7 23.510.8 22.610.2 ο
HL/SL 37.3 37.010.9 36.511.1 36.910.6 36.611.1 36.810.9 36.910.7 c
un
SnL/HL 30.4 31.711.5 32.111.2 33.011.7 34.111.1 34.311.3 35.110.4 DC
SnW/HL 30.2 30.611.9 31.211.4 31.811.9 33.111.7 33.311.6 32.4125
>
HW/HL 41.0 41.210.8 41.411.6 42.011.3 42.9±1.5 42.811.0 42.811.7 TJ
r
IOW/HL 19.3 20.210.6 20.710.9 20.811.1 22.011.0 21.810.8 22.310.9
POW/HL 25.8 25.910.8 25.511.1 26.410.9 26.112.1 26.410.7 26.711.3
R
LaW/HL 26.4 27.611.1 26.311.8 27.311.0 28.211.6 28.111.3 28.311.9
POD/HL 15.7 17.111.0 18.110.6 19.912.0 18.610.7 19.110.6 19.110.6
EyL/HL 26.2 26.410.6 25.0115 23.611.0 23.212.0 22.711.1 22.110.6 1
ChD/HL 20.9 22.310.9 22.810.8 24.011.0 24.811.4 25.310.9 26.412.0 η
LJL/HL 47.5 49.411.6 48.811.8 51.610.9 51.211.0 51.510.9 53.612.0 η
20.712.2 20.611.5 23.511.8 22.711.4 22511.0 r
LJW/HL 17.6 20.911.9 Ο
39.0±1.2 39.311.2 39.510.4 40.611.1 41.912.4 >
UJL/HL 37.6 38.511.1 m
PPL/HL 27.3 27.810.9 27.811.3 28.010.9 27.911.4 28511.3 27510.6 Ο
τι
r
>
<
g
2
>
Ol
of the o p e r c u l u m a n d the d o r s a l o u t l i n e of the n a p e a b r o a d n a p e b a n d r u n s m e d i a d .

T h e o p e r c u l a r b l o t c h is e x t e n d e d rostrad. B o d y : R o s t r a l part of f l a n k (above pectoral
f i n o r i g i n ) a n d rostral part of d o r s u m d a r k b l o o d r e d , the r e m a i n i n g part of f l a n k a n d
c a u d a l p e d u n c l e green, darkest ventrally. In s o m e s p e c i m e n s 4-6 faint v e r t i c a l b a n d s
are present o n the d o r s u m a n d d o r s a l part of the flank. Scales o n the centre of the
f l a n k w i t h a light-green iridescent f l u s h . F i n s : Pectoral h y a l i n e , p e l v i c s black. D o r s a l
g r e y i s h - h y a l i n e w i t h a d a r k r e d f l u s h rostrally, d a r k grey lappets, a n d d a r k r e d spots
b e t w e e n the rays. A n a l h y a l i n e , w i t h a d a r k r e d f l u s h rostrally a n d p r o x i m a l l y , a n d
w i t h t w o r o u n d d a r k o r a n g e spots w i t h a w h i t e i n n e r a n d grey outer m a r g i n d o r s o -
c a u d a l l y . C a u d a l f i n l i g h t r e d , m o r e intensely r e d ventrally. A grey f l u s h o n the d o r -
sal half of the f i n i n s o m e specimens.
— L i v e c o l o u r s of s e x u a l l y active females. H e a d : B l a c k ventrally, d a r k b l u e - g r e y
w i t h a r e d sheen dorsally. Scaled part of the cheek lighter i n s o m e specimens. L o w e r
j a w w i t h a b l u e sheen. A r e d sheen above the o p e r c u l u m . M a r k i n g s : In spite of the
Fig. 60. Haplochromis dichrourus "Mwanza". Live colours and markings of a sexually active male and a
ripe female.
v e r y d a r k h e a d c o l o u r s , a b r o a d l a c h r y m a l stripe, w h i c h is e x t e n d e d above the eye,

as w e l l as a n M s h a p e d n o s t r i l stripe, a n i n t e r o r b i t a l stripe a n d a n a p e b a n d c a n be
seen. B o d y : D a r k green to black w i t h a r e d sheen rostro-dorsally, F i n s : Pectoral h y a
l i n e , p e l v i c s black. D o r s a l g r e y i s h - h y a l i n e , r o s t r a l l y w i t h a r e d f l u s h , l a p p e t s d a r k
grey. S m a l l , d a r k r e d , r o u n d spots b e t w e e n the r a y s . A n a l d a r k r e d r o s t r a l l y a n d
p r o x i m a l l y , l i g h t e r c a u d a l l y . O n e o r t w o large, r o u n d , l i g h t o r a n g e o r r e d spots w i t h a
grey m a r g i n d o r s o - c a u d a l l y o n the f i n . C a u d a l d a r k grey w i t h a r e d f l u s h o n the d o r
sal half, r e d o n the v e n t r a l half.
— P r e s e r v e d c o l o r a t i o n . It is n o t p o s s i b l e to separate the sexes o n the basis of
their p r e s e r v e d c o l o r a t i o n since the c o l o u r pattern is the same f o r males a n d females.
G r o u n d c o l o u r b r o w n i s h . H e a d w i t h a distinct, b r o a d , l o n g l a c h r y m a l stripe extend
e d above the eye, a b r o a d characteristically M - s h a p e d n o s t r i l stripe, a b r o a d inter
r u p t e d i n t e r o r b i t a l s t r i p e a n d a b r o a d b u t i r r e g u l a r n a p e b a n d ( f i g . 108). A n a p e
b l o t c h m a y also be present. M o s t specimens also h a v e a distinct v e r t i c a l p r e o p e r c u l a r
b a n d . L o w e r j a w sometimes w i t h a d a r k m e n t a l area, o t h e r w i s e l i g h t l a t e r a l l y a n d
ventrally. B r a n c h i o s t e g a l m e m b r a n e b l a c k , g i l l - c o v e r a n d i n t e r o p e r c u l a r area d a r k .
B o d y u s u a l l y darkest ventrally, crossed d o r sally b y 5-8 v e r t i c a l stripes. A l l fins h y a
l i n e , b u t the c a u d a l grey dorsally. D a r k spots b e t w e e n the r a y s of the d o r s a l f i n , a n d
i n a f e w s p e c i m e n s , i n the d o r s a l half of the c a u d a l f i n . A n a l f i n i n males a l w a y s , i n
females o f t e n , w i t h o n e o r t w o large, l i g h t l y c o l o u r e d , d a r k r i m m e d , r o u n d spots
b e t w e e n the c a u d a l rays.
E c o l o g y . — O c c u r r e n c e . I n the n o r t h e r n part of the M w a n z a G u l f Haplochromis
dichrourus w a s a rare species. D e s p i t e the fact that w e w e r e k e e n to collect specimens
of this c o n s p i c u o u s l y c o l o u r e d species, three years of i n t e n s i v e s a m p l i n g a n d
i n s p e c t i n g m a n y t h o u s a n d s o f s p e c i m e n s i n the M w a n z a G u l f d i d n o t y i e l d m o r e
t h a n a p p r o x i m a t e l y 60 s p e c i m e n s . U p to 1987 t r a w l catches i n t h e s o u t h o f t h e
M w a n z a G u l f (at t h e e n t r a n c e of the S m i t h S o u n d ) s t i l l y i e l d e d s p e c i m e n s o f H.
dichrourus. A p p a r e n t l y this species s u r v i v e d the N i l e p e r c h l o n g e r t h a n H. bareli.
— H a b i t a t . A l l specimens except one, w e r e caught i n a b o t t o m t r a w l i n the o p e n
w a t e r of the M w a n z a G u l f , at m a n y different localities, w i t h b o t t o m d e p t h s r a n g i n g
f r o m 2-15 m . These catches h a d i n c o m m o n that they w e r e m a d e o v e r soft m u d bot
toms. O n e s p e c i m e n (SL 70.5 m m ) w a s caught i n the K i s e n d a B a y at a d e p t h b e t w e e n
1 a n d 2 m . H. dichrourus inhabits r e l a t i v e l y s h a l l o w a n d r e l a t i v e l y o p e n waters w i t h a
m u d bottom.
— F o o d . Stomachs a n d intestines of 13 specimens w e r e e x a m i n e d . T w o of these
w e r e empty, b u t a l l others c o n t a i n e d r e m a i n s of h a p l o c h r o m i n e c i c h l i d s . Specimens
of 56-81 m m c o n t a i n e d r e m a i n s of h a p l o c h r o m i n e s of 11-15 m m a n d i n specimens of
114-148 m m , p r e y l e n g t h r a n g e d f r o m 25-45 m m . I n t w o s m a l l e r s p e c i m e n s , apart
f r o m f i s h r e m a i n s , u n i d e n t i f i a b l e insect r e m a i n s a n d r e m a i n s of c h i r o n o m i d l a r v a e
w e r e f o u n d . H. dichrourus " M w a n z a " is a p r e d a t o r o f j u v e n i l e h a p l o c h r o m i n e s .
— B r e e d i n g a n d g r o w t h . N o i n f o r m a t i o n is available o n the t y p e of brood-care.
R i p e females w e r e c a u g h t i n January, M a y , June, A u g u s t , N o v e m b e r a n d December.
Size of ovaries a n d r i p e eggs is w i t h i n the range of other, e q u a l l y s i z e d p i s c i v o r o u s
species. T h e left o v a r y is a l w a y s larger than the r i g h t one. E g g sizes m e a s u r e d i n ripe
ovaries w e r e ± 3.2 χ 2.4 m m i n a female of 109 m m S L , a n d 3.6 χ 2.8 m m a n d 3.8 χ 3.0
m m i n females of 126 a n d 121.2 m m S L , respectively. R e m a r k a b l y , o n l y 10 of the 53
s p e c i m e n s c a u g h t b y H E S T are m a l e s . S e x u a l l y a c t i v e m a l e s w e r e c a u g h t o n l y i n
January a n d June. A l l specimens w i t h over 100 m m S L are mature. Females g r o w to a

larger size than males: the largest specimen, 141 m m S L , is a female; the largest male is
117 m m . T h e smallest sexually active m a l e is 88.9 m m , the smallest r i p e female 97.1 m m .
C o m p a r i s o n of the p o p u l a t i o n s
C o m p a r i s o n of the three a l l o p a t r i c g r o u p s o f specimens o f H. dichrourus is h a m -
p e r e d b y the fact that there is a difference i n size of the g r o u p s as w e l l as a difference
i n size of the specimens b e l o n g i n g to the g r o u p s . S t i l l , each g r o u p seems to h a v e its
o w n facies w i t h a p a r t i c u l a r h e a d a n d b o d y shape a n d p a t t e r n of h e a d m a r k i n g s .
T h e r e are also m i n o r differences i n dental characters. I m m e d i a t e l y a p p a r e n t is the
difference i n B o d y D e p t h b e t w e e n the g r o u p s . T h e specimens f r o m U g a n d a h a v e the
m o s t s h a l l o w b o d y , a n d the M w a n z a G u l f specimens the deepest b o d y (fig. 61). A n d
a l t h o u g h they seem to b e m o r e laterally c o m p r e s s e d the specimens of H. dichrourus
" M w a n z a " h a v e r e l a t i v e l y b r o a d e r heads (expressed i n r e l a t i v e l y h i g h e r measure-
m e n t s f o r P O W / H L a n d H W / H L ) . T h e M a j i t a s p e c i m e n s s e e m to h a v e a m o r e
r o u n d e d , " s m o o t h " p r o f i l e w i t h s l i g h t l y b r o a d e n e d lips. A s f o r the d e n t a l characters,
i n a l l g r o u p s the teeth i n the o r a l jaws are m o d e r a t e l y to s t r o n g l y c u r v e d u n i c u s p i d s ,
o n l y i n the h o l o t y p e s o m e s h o u l d e r e d u n i c u s p i d s a n d o n e u n e q u a l l y b i c u s p i d occur.
T h e specimens f r o m M a j i t a h a v e m o r e i n n e r r o w s than the specimens f r o m the other
g r o u p s ( P I R 4 v s 2-3, L I R 3-4 v s 2), b u t this m i g h t b e e x p l a i n e d b y their larger size.
T h e U g a n d a specimens h a v e m o r e teeth i n the outer r o w s of the o r a l jaws t h a n the
other g r o u p s . F i n a l l y there are slight differences i n i m p l a n t a t i o n .
There is a r e m a r k a b l e difference i n the shape of the h e a d m a r k i n g s b e t w e e n the
g r o u p s (fig. 62). I n t h e h o l o t y p e a n d the s p e c i m e n f r o m K a t e b o ( B M ( N H ) 1966.3.
9.187) there is a r e l a t i v e l y s h o r t , d i s t i n c t l a c h r y m a l s t r i p e w h i c h e n d s a b o v e t h e
m o u t h corner. T h e n o s t r i l stripe is i n t e r r u p t e d m e d i a l l y a n d of the i n t e r o r b i t a l stripe
o n l y the lateral ends are present. I n the other specimens f r o m U g a n d a the l a c h r y m a l
stripe is e x t e n d e d till the h o r i z o n t a l p r e o p e r c u l a r a r m . I n t w o specimens the n o s t r i l
stripe is c o n t i n u o u s .
In the s p e c i m e n f r o m M a j i t a the l a c h r y m a l stripe r u n s f r o m the e y e till the h o r i -
z o n t a l p r e o p e r c u l a r a r m , the i n t e r o r b i t a l stripe is distinct b u t i n t e r r u p t e d m e d i a l l y
a n d the n o s t r i l stripe is c o n t i n u o u s a n d b r o a d e r m e d i a l l y . R o s t r a l l y b e t w e e n the lat-
eral ends o f the n o s t r i l stripe a n d the l i p s a n isolated d a r k spot is present.
In the M w a n z a G u l f specimens the l o n g l a c h r y m a l stripe sometimes runs past the
h o r i z o n t a l p r e o p e r c u l a r a r m to the branchiostechal m e m b r a n e . A distinct, i n t e r r u p t e d
Fig. 61. Body outline of equally sized specimens of H. dichrourus "Mwanza" (continuous line) and H.
dichrourus "Uganda". Scale equals 10 mm.
V A N OIJEN: PISCIVOROUS H A P L O C H R O M I N E CICHLIDAE OF L A K E VICTORIA 49
Fig. 62. Head markings in A) Haplochromis dichrourus "Uganda", B) H. dichrourus "Majita", C) holotype
of H. dichrourus Regan, 1922 (shape of markings taken from fig. 6 of Regan, 1922, D)H.dichrourus
"Mwanza", E) H. maisomei spec. nov.
o r c o n t i n u o u s interorbital stripe a n d a n a p e b a n d are present. T h e n o s t r i l stripe has a

characteristic M - s h a p e . T h e lateral ends b e i n g extended rostrally to the l i p s .
A l t h o u g h it is k n o w n that e n v i r o n m e n t a l factors m a y h a v e a n effect o n certain
aspects o f the m o r p h o l o g y (Witte & W i t t e - M a a s , 1987), i t is n o t k n o w n if the s h a p e of
h e a d m a r k i n g s c a n change u n d e r the i n f l u e n c e o f e n v i r o n m e n t a l c o n d i t i o n s . T h e dif-
ferent h e a d m a r k i n g s m a y w e l l be genetically d e t e r m i n e d a n d c o u l d b e a reflection
of genetical i s o l a t i o n . It is l i k e l y that the h e a d m a r k i n g s are part o f the Specific M a t e
R e c o g i t i o n S y s t e m (Paterson, 1978; R i b b i n k et a l . , 1983) b u t it is n o t k n o w n w h e t h e r
the d i f f e r e n c e s i n h e a d m a r k i n g w o u l d p r e v e n t the m e m b e r s of the p o p u l a t i o n s of H.

dichrourus to m a t e a n d i n t e r b r e e d s h o u l d t h e y meet i n a n a t u r a l e n v i r o n m e n t .
Table 9. Angular and qualitative measurements, and counts of Haplochromis dichrourus "Mwanza '
Regan, 1922.
range mean and standard

deviations
Dorsal Head Inclination 26°-37° 31.1°±3.1°

Premaxillary Pedicel Inclination 28°-42° 35.9°±3.6°
Snout Acuteness 64°-78° 72.2°±3.2°
Gape Inclination 30°-42° 35.7°±3.1°
Dorsal Head Profile (curvature) 0(+)

Premaxilla Pedicel Prominence 0/+(+); mainly +
Lower Jaw: anterior extension +
Lateral Snout Outline 0/+; mainly +
Mental Prominence 0/+; mainly 0 and 0(+)
Lip Thickening 0
Premaxillar Beaked 0
Premaxillar Expanded 0/0(+); mainly 0
Maxillary Posterior Extension +
Cephalic lateral line Pores: Width 0
Lateral Line Scales 29 (f= 1); 30 (f= 3), 31 (f= 25); 32 (f= 11)
Lateral Line-Dorsal Fin (Sc.rows) 5 (f= 10); 6 (f= 28), (f= 2)
Pectoral Pelvic Fin Bases (Sc.rows) 5 (f= 3); 6 (f= 20), 7 (f= 15); 8 (f= 3)
Cheek (Vertical Sc.rows) 3 (f= 2); 4 (f= 21) 5 (f= 18)
Dorsal Fin (Spines/Rays) XIV 10 (f= 1); XV 9 (f= 14); X V 10 (f= 13); XVI9 (f= 12);
XVI10 (f= 1)
Anal Fin (Spines/Rays) III 8 (f= 3); III 9 (f= 21); III 10 (f= 5)
H a p l o c h r o m i s m a i s o m e i spec. nov.
(figs 6 1 E , 1 0 9 , tables 5, 6).
Material — Holotype, cf, 98.1 mm, R M N H 30122, Mikassa Bay, Maisome Island, Tanzania, Lake
Victoria, 20.vi.1985, HEST; paratype, $, 96.1 mm, R M N H 30121, Mikassa Bay, Maisome Island,
Tanzania, Lake Victoria, 20.vi.1985, HEST. Other material: juv. 81.5 mm, R M N H 30386, sand spit
South of Standieri Island, Tanzania, Lake Victoria, 5.XÜ.1984, HEST.
E t y m o l o g y . — T h e s p e c i f i c n a m e is d e d u c e d f r o m the n a m e of the i s l a n d i n a b a y
of w h i c h the t y p e s p e c i m e n s w e r e c a p t u r e d .
D i a g n o s i s . — A m e d i u m s i z e d p i s c i v o r o u s species w i t h s t r o n g l y r e c u r v e d u n i c u s -
p i d teeth i n the o u t e r r o w of the o r a l j a w s . F e m a l e c o l o r a t i o n c o n t a i n s e l e m e n t s u s u -
a l y r e s t r i c t e d to m a l e c o l o r a t i o n s u c h as b l a c k p e l v i c f i n s , a b r i g h t l y c o l o u r e d c a u d a l
a n d a n a l f i n , a n e g g d u m m y o n the a n a l f i n a n d d i s t i n c t h e a d m a r k i n g s . T h e m a l e is
y e l l o w i s h p i n k o n the cheek, the g i l l cover, the f l a n k a n d the c a u d a l p e d u n c l e . Its
h e a d m a r k i n g s i n c l u d e a d i s t i n c t l a c h r y m a l s t r i p e w h i c h is e x t e n d e d a b o v e the eye.
D e s c r i p t i o n . — Based o n the h o l o t y p e , a s e x u a l l y active m a l e 96.1 m m S L a n d one

p a r a t y p e , a n a d u l t r i p e n i n g female 98.1 m m S L .
H a b i t u s . — B o d y r e l a t i v e l y slender a n d m o d e r a t e l y laterally c o m p r e s s e d . D o r s a l
part of h e a d p r o f i l e convex, the l o w e r part s l i g h t l y concave (in the male) or straight
(in the female specimen). P r e m a x i l l a r y p e d i c e l h a r d l y (male) or d i s t i n c t l y (female)
i n t e r r u p t i n g t h e p r o f i l e . M o u t h s l i g h t l y to m o d e r a t e l y o b l i q u e . L i p s n o r m a l .
P r e m a x i l l a n o t e x p a n d e d m e d i a l l y o r b e a k e d . T h e e x p o s e d p a r t of the m a x i l l a s m a l l .
T h e v e r t i c a l t h r o u g h the c a u d a l t i p of the m a x i l l a p a s s i n g the rostral eye m a r g i n .
L o w e r j a w isognathous a n d s l i g h t l y p r o t r u d i n g . R o s t r a l o u t l i n e of l o w e r j a w r o u n d -
e d , n o m e n t a l p r o m i n e n c e . L a t e r a l sides of l o w e r j a w v e r y s l i g h t l y o b l i q u e .
H o r i z o n t a l a r m of p r e o p e r c u l u m i n female h o r i z o n t a l , i n m a l e v e n t r a d i n c l i n e d ros-
trally. T h e v e r t i c a l a r m v e r t i c a l i n female, r e c l i n i n g c a u d a d i n the male. E y e circular,
p u p i l s l i g h t l y e l o n g a t e d i n rostro-ventral d i r e c t i o n , a s m a l l a p h a k i c aperture rostro-
v e n t r a l l y to the lens. C e p h a l i c lateral l i n e o p e n i n g s r e l a t i v e l y s m a l l , the canals o n the
lachrymal faintly visible.
S c a l e s . — C h e e k , gill-cover, d o r s a l h e a d surface a n d greater part of d o r s u m w i t h
c y c l o i d scales. F l a n k , c a u d a l p a r t of d o r s u m a n d c a u d a l p e d u n c l e w i t h c t e n o i d
scales. C h e s t m a i n l y w i t h c y c l o i d scales, s o m e w e a k l y c t e n o i d scales l a t e r a l l y . A
g r a d u a l size t r a n s i t i o n b e t w e e n scales of chest a n d f l a n k . C a u d a l f i n scaled o n its
p r o x i m a l t w o - t h i r d part.
F i n s . — Pectorals s l i g h t l y longer than p e l v i c s , o n l y i n the m a l e just r e a c h i n g the
a n a l f i n . First p e l v i c f i n r a y s l i g h t l y p r o d u c e d . D o r s a l a n d a n a l f i n not r e a c h i n g base
of c a u d a l . C a u d a l f i n o u t l i n e subtruncate a n d straight.
G i l l a p p a r a t u s . — E i g h t g i l l rakers o n l o w e r p a r t of first g i l l arch o n r i g h t side.
T h e first one r e d u c e d , c o n i c a l , n u m b e r s 2-5 flattened, s i m p l y t a p e r i n g , 6 a n d 7 b r o a d -
e n e d , square t o p p e d a n d the last one m o r e r o u n d e d a n d t a p e r i n g distally. N u m b e r of
g i l l filaments o n the first h e m i b r a n c h 116 a n d 118.
V i s c e r a . — I n t e s t i n e l e n g t h of the t w o s p e c i m e n s e q u a l s the s t a n d a r d l e n g t h .
F o l l o w i n g the d e f i n i t i o n s of Z i h l e r (1982) the arrangement of the d i g e s t i v e track is of
t y p e D ( w i t h backflap).
O r a l t e e t h . — S h a p e . I n the o u t e r r o w of b o t h j a w s s l e n d e r , r o u n d e d , a c u t e l y
p o i n t e d u n i c u s p i d s d o m i n a t e , teeth o n the c a u d a l half of the d e n t a r y are s o m e w h a t
c o m p r e s s e d . In the l o w e r j a w of the female s p e c i m e n s o m e w e a k l y b i c u s p i d s a n d
s h o u l d e r e d u n i c u s p i d s are f o u n d laterally a n d caudally. A l l teeth are m o d e r a t e l y to
s t r o n g l y c u r v e d , i n u p p e r j a w teeth the c u r v a t u r e is stronger. T o o t h size g r a d u a l l y
decreasing c a u d a d . Inner r o w s consist of a m i x t u r e of m o d e r a t e l y c u r v e d , s l i g h t l y
compressed unicuspids and shouldered unicuspids.
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . A distinct m e d i a l g a p i n
the l o w e r jaw. T w o r o w s of inner teeth i n b o t h jaws. O u t e r t o o t h r o w seems to cover
w h o l e l e n g t h of dentigerous a r m of the p r e m a x i l l a , a n d seems to reach the c o r o n o i d
w i n g o n the l o w e r jaw.
— C o u n t s a n d setting. There are 49 a n d 53 teeth i n outer r o w of p r e m a x i l l a of the
m a l e a n d female s p e c i m e n , respectively; 30 a n d 32 i n the outer r o w of the l o w e r jaw.
Teeth are r e g u l a r l y set at a distance v a r y i n g b e t w e e n half to s l i g h t l y less t h a n the
d i a m e t e r of the toothbase.
— I m p l a n t a t i o n . O u t e r r o w of u p p e r j a w erect to s l i g h t l y r e c u m b e n t rostrally,
a n d s t r o n g l y r e c u m b e n t caudally. O u t e r r o w of l o w e r j a w erect to s l i g h t l y p r o c u m -
bent rostrally, s l i g h t l y r e c u m b e n t caudally. Inner r o w s : u p p e r j a w s t r o n g l y r e c u m -
bent, l o w e r j a w recumbent.
O s t e o l o g y . — T h e t w o M a i s o m e s p e c i m e n s h a v e n o t b e e n d i s s e c t e d to o b t a i n
skeletal m a t e r i a l .
— Vertebrae. There are 30 vertebrae i n b o t h specimens, c o m p r i s i n g 13 a b d o m i n a l
a n d 17 c a u d a l elements.
C o l o r a t i o n . — L i v e colours of a ripening m a l e : G r o u n d c o l o u r y e l l o w i s h - p i n k o n
c a u d a l p a r t of l o w e r jaw, cheek, gill-cover, f l a n k a n d c a u d a l p e d u n c l e . S n o u t , d o r s a l
area of h e a d , d o r s u m a n d v e n t r a l side l i g h t b l u e . L i p s l i g h t b l u i s h laterally, d a r k ros-
trally. V e n t r a l aspects of cheek, g i l l - c o v e r a n d branchiostegal m e m b r a n e bluish-grey.
V e n t r a l s i d e of chest a n d b e l l y sooty. H e a d m a r k i n g s : a d i s t i n c t b r o a d l a c h r y m a l
stripe e x t e n d e d above the eye, a n o s t r i l stripe, a n interorbital stripe, a n a p e b a n d , a
n a p e b l o t c h a n d a p r e o p e r c u l a r v e r t i c a l stripe. F i n s : D o r s a l l i g h t b l u e , w i t h a p i n k i s h
f l u s h p r o x i m a l l y , lappets d a r k grey. Pectorals h y a l i n e , p e l v i c s black. A n a l p i n k i s h - r e d
rostrally, t u r n i n g w h i t i s h d i s t a l l y o n the rostral half of the r a y e d part. C a u d a l part of
the a n a l h y a l i n e w i t h t w o b r i g h t orange egg d u m m i e s w i t h a t h i n w h i t e r i m . C a u d a l
f i n b r i g h t r e d o n its v e n t r a l half, d o r s a l half g r e y i s h - h y a l i n e w i t h d a r k streaks be-
t w e e n the rays.
— L i v e c o l o r a t i o n of a r i p e n i n g female: G r o u n d c o l o u r o n h e a d a n d b o d y d a r k
b l u e . A faint r e d d i s h f l u s h o n the n a p e a n d the area just above the gill-cover. L a t e r a l
parts of l i p s a n d l o w e r jaw, a n d h o r i z o n t a l a r m of p r e o p e r c u l u m l i g h t b l u e i s h c o n -
trasting w i t h b l a c k interopercular a n d i n t e r m a n d i b u l a r area a n d rostral part of the
branchiostegal membrane. H e a d m a r k i n g s : A distinct l a c h r y m a l stripe extended
a b o v e the eye, a fainter n o s t r i l stripe, lateral ends of the interorbital stripe a n d a large
o p e r c u l a r b l o t c h . F i n s : D o r s a l d a r k b l u e p r o x i m a l l y , l i g h t b l u e distally, lappets d a r k
grey. P e c t o r a l h y a l i n e . P e l v i c s black. A n a l d a r k r e d , c a u d a l m o s t area a r o u n d a b r i g h t
orange ocellus h y a l i n e . C a u d a l f i n b r i g h t r e d ventrally, the d o r s a l half g r e y s i h h y a -
l i n e w i t h d a r k streaks b e t w e e n the rays.
— P r e s e r v e d m a l e c o l o r a t i o n . G r o u n d c o l o u r of h e a d b r o w n i s h grey, lateral parts
of l i p s , c a u d o - l a t e r a l part of l o w e r j a w a n d rostral area of h o r i z o n t a l a r m of preoper-
c u l a r i v o r y , c o n t r a s t i n g w i t h d a r k g r e y to b l a c k i n t e r m a n d i b u l a r area a n d b r a n -
chiostegal m e m b r a n e . H e a d m a r k i n g : a distinct l a c h r y m a l stripe, e x t e n d e d above the
eye. T h e t w o extensions d o not meet m e d i a l l y . A v a g u e n o s t r i l stripe. O f the interor-
b i t a l stripe o n l y the lateral ends are present. T h e nape b a n d s m e d i a l l y m e r g e i n t o a
nape blotch. Opercular blotch vague. Body u n i f o r m l y brownish-grey. D o r s u m and
v e n t r a l side of b e l l y a n d chest s l i g h t l y darker, a s m a l l elongate area above the a n a l
f i n i v o r y . F i n s : p e c t o r a l a n d d o r s a l g r e y i s h , l a p p e t s d a r k grey. P e l v i c b l a c k . A n a l
i v o r y , c a u d a l l y w i t h t w o i v o r y spots i n a h y a l i n e area. C a u d a l g r e y i s h basally, i v o r y
o n the v e n t r a l half, grey w i t h d a r k streaks b e t w e e n the rays d o r s a l l y
— P r e s e r v e d female c o l o r a t i o n . H e a d : cheek, snout a n d d o r s a l h e a d surface grey-
b r o w n . L a t e r a l s i d e of l i p s a n d l o w e r j a w a n d h o r i z o n t a l a r m of p r e o p e r c u l u m i v o r y ,
c o n t r a s t i n g w i t h b l a c k i n t e r m a n d i b u l a r area, i n t e r o p e r c u l u m a n d b r a n c h o s t e g a l
m e m b r a n e . A distinct b l a c k l a c h r y m a l stripe r u n n i n g f r o m the rostro-ventral b o r d e r
of the eye b e h i n d the m a x i l l a to the cheek m a r g i n . Just c a u d a l to the l o w e r j a w a
b l a c k spot lies i n the extension of the l a c h r y m a l stripe w h i c h is also e x t e n d e d above
the eye. O n the d o r s a l snout surface a v a g u e n o s t r i l stripe a n d the lateral tips of a n
i n t e r o r b i t a l stripe c a n be seen. O p e r c u l a r b l o t c h large, m o r e d i s t i n c t t h a n i n m a l e .
B o d y d a r k grey r o s t r a l l y a n d dorsally, v e n t r a l s i d e of chest sooty. C a u d o - v e n t r a l part
of b o d y i v o r y . A v a g u e m i d l a t e r a l b a n d rostrally starting t h i n i n the grey area a n d

e n d i n g b r o a d l y o n the base of the c a u d a l f i n . F i n s : pectorals a n d d o r s a l grey, lappets
of d o r s a l d a r k e r grey. P e l v i c s black, anal i v o r y w i t h a d a r k r i m m e d i v o r y spot cau-
dally. C a u d a l g r e y i s h basally, i v o r y o n the v e n t r a l half, a n d grey w i t h d a r k streaks o n
its d o r s a l half.
D i s t r i b u t i o n . — H. maisomei is o n l y k n o w n f r o m L a k e V i c t o r i a . T h e t y p e speci-
m e n s w e r e caught i n the M i k a s s a B a y of M a i s o m e I s l a n d , a n d another s p e c i m e n w a s
caught s o u t h of S t a n d i e r i Island.
E c o l o g y . — H a b i t a t . B o t h type specimens w e r e caught i n a b o t t o m t r a w l o v e r a
m u d b o t t o m of 10-14 m d e p t h i n the relatively o p e n water. T h e t h i r d s p e c i m e n w a s
caught w i t h a b o t o m t r a w l , over s a n d i n relatively o p e n water w i t h a d e p t h of 4-7 m .
— F o o d . T h e d i g e s t i v e tract of b o t h t y p e s p e c i m e n s w a s e x a m i n e d . N o f o o d
r e m a i n s w e r e f o u n d i n t h e m a l e a n d the s t o m a c h o f the f e m a l e w a s also e m p t y .
H o w e v e r , i n the g u t of the female s p e c i m e n remains of a s m a l l h a p l o c h r o m i n e c i c h -
l i d , c. 20 m m S L a n d r e m a i n s of insect larvae w e r e f o u n d .
— B r e e d i n g a n d g r o w t h . T h e type specimens are m a t u r e , the s p e c i m e n of 81.5
m m S L is i m m a t u r e .
R e s e m b l i n g s p e c i e s . — Specimens of H. maisomei are easily m i s t a k e n f o r H. di-
chrourus because of their great s i m i l a r i t y i n d e n t i t i o n a n d c o l o u r pattern. C o m p a r e d
to specimens of H. dichrourus " U g a n d a " of a s i m i l a r size, specimens of H. maisomei
have a slightly deeper body, a slightly longer c a u d a l p e d u n c l e , a slightly larger
i n t e r o r b i t a l w i d t h a n d a s l i g h t l y larger eye. T h e d o r s a l h e a d i n c l i n a t i o n is steeper
a n d t h e m o u t h less o b l i q u e . S p e c i m e n s of H. maisomei h a v e i n c o m m o n w i t h the
h o l o t y p e of H. dichrourus the presence of some s h o u l d e r e d u n i c u s p i d s a n d u n e q u a l l y
b i c u s p i d s c a u d a l l y i n the o r a l jaws. T h e large size g a p b e t w e e n H. maisomei a n d the
specimens of H. dichrourus " M a j i t a " m a k e a c o m p a r i s o n d i f f i c u l t b u t it is clear that
the H. maisomei specimens h a v e a less convex d o r s a l h e a d p r o f i l e a n d a m u c h m o r e
laterally c o m p r e s s e d b o d y . C o m p a r e d to s i m i l a r l y s i z e d specimens of H. dichrourus
" M w a n z a " the specimens o f H. maisomei h a v e a less c o n v e x d o r s a l h e a d p r o f i l e a n d a
m o r e p r o n o u n c e d i n c u r v a t i o n above the eye, m a k i n g the area rostral to the e y e less
deep. D o r s a l h e a d i n c l i n a t i o n is r e l a t i v e l y steeper i n H. maisomei. T h e r e is a great
s i m i l a r i t y i n m o r p h o m e t r i c measurements; o n l y the ratio's f o r C F L / S L a n d U J L / H C
are s m a l l e r i n H. maisomei. Specimens of H. maisomei c a n be d i s t i n g u i s h e d f r o m speci-
m e n s of H. dichrourus " M w a n z a " b y the n u m b e r of g i l l - f i l a m e n t s v i z . 116-118 v s ca.
132. T h e l o w e r n u m b e r i n H. maisomei m i g h t be a correlated w i t h the m o r e o p e n a n d
c o n s e q u e n t l y better o x y g e n a t e d w a t e r i n w h i c h t h e s p e c i e s i s f o u n d . T h e m o s t
i m p o r t a n t difference b e t w e e n the species is the c o l o r a t i o n of the l i v e m a l e . M a l e s of
H. dichourus " M w a n z a " h a v e d a r k r e d areas o n the d o r s a l parts of h e a d a n d b o d y
w h i c h are n o t present i n the m a l e of H. maisomei. T h e m a l e c o l o r a t i o n of H. maisomei
is m u c h lighter t h a n that of H. dichrourus " M w a n z a " . T h e h e a d m a r k i n g s of H. mai-
somei differ f r o m a l l H. dichrourus specimens as the n o s t r i l stripe is v a q u e , o n l y the
lateral parts of the interorbital stripe are f a i n t l y v i s i b l e a n d the n a p e b a n d is absent
(fig. 28).
T h e specimens of H. maisomei demonstrate the i m p o r t a n c e of data o n l i v e colours
for the i d e n t i f i c a t i o n of h a p l o c h r o m i n e c i c h l i d s . If the data o n m a l e l i v e c o l o r a t i o n of
the specimens f r o m M a i s o m e i s l a n d h a d n o t been available, the specimens certainly
w o u l d h a v e been c o n s i d e r e d as representatives of another p o p u l a t i o n of H. dichrou-
rus. It is possible that the specimens of H. dichrourus f r o m U g a n d a o r M a j i t a differed
i n m a l e l i v e c o l o r a t i o n f r o m those of the M w a n z a G u l f . In that case they too s h o u l d

h a v e been c o n s i d e r e d separate species. W i t h the data n o w a v a i l a b l e it c a n o n l y be
c o n c l u d e d that the s p e c i m e n s f r o m the M w a n z a G u l f a n d t h o s e f r o m M a i s o m e
I s l a n d b e l o n g to different species. A s the h o l o t y p e of H. dichrourus has m o r e m o r p h o -
l o g i c a l c h a r a c t e r s i n c o m m o n w i t h the H. dichrourus-like s p e c i m e n s f r o m t h e
M w a n z a G u l f t h a n w i t h those of the M a i s o m e s p e c i m e n s , I c o n s i d e r the M w a n z a
G u l f specimens to be representatives of H. dichrourus.
H a p l o c h r o m i s k u j u n j u i spec. nov.
(fig. 101, tables 10,11)
Material.— Holotype, 9, 102.8 mm SL, R M N H 30466. Sand spit south of Standieri Island, Tanzania,
Lake Victoria, 5.VÍ.1985, HEST.
E t y m o l o g y . — T h e n a m e of the species refers to the n a m e of a l a n d m a r k ( K u j u n j u

p o i n t ) to the north-east of w h i c h the s p e c i m e n w a s caught.
D i a g n o s i s . — A m e d i u m s i z e d , slender, p i s c i v o r o u s species w i t h s t r o n g l y r e c u r v -
e d u n i c u s p i d teeth i n the o r a l jaws. F e m a l e l i v e c o l o r a t i o n ; w h o l e b o d y a n d h e a d
u n i f o r m l y b r o w n , m e d i a n fins r e d . A n a l f i n w i t h distinct, r e l a t i v e l y large, orange e g g
d u m m i e s . P e l v i c fins black.
D e s c r i p t i o n . — Based o n the h o l o t y p e only.
H a b i t u s . — A r e l a t i v e l y slender species w i t h a s p i n d l e s h a p e d b o d y . A s the pre-
s e r v e d specimens has its m o u t h o p e n , the u p p e r j a w is s l i g h t l y p r o t r u d e d a n d the
suspensória a b d u c t e d , the d e s c r i p t i o n of the headshape m a y be b i a s e d . D o r s a l h e a d
o u t l i n e s l i g h t l y c o n v e x w i t h a s m a l l i n c u r v a t i o n a b o v e the eye. T h e p r e m a x i l l a r y
p e d i c e l p r o b a b l y s l i g h t l y i n t e r r u p t i n g the d o r s a l h e a d p r o f i l e . S n o u t s o m e w h a t
b l u n t . E y e almost circular, p u p i l s o m e w h a t elongate i n the h o r i z o n t a l axis thus f o r m -
i n g a s m a l l o p h a k i c space rostral to the lens. M o u t h s l i g h t l y o b l i q u e . P r e m a x i l l a not
e x p a n d e d m e d i a l l y b u t l i p s seem s o m e w h a t b r o a d e n e d . Posterior t i p of the m a x i l l a
p r o b a b l y just not r e a c h i n g a v e r t i c a l t h r o u g h the anterior eye m a r g i n . L o w e r j a w
i s o g n a t h o u s , s l i g h t l y p r o t r u d i n g . R o s t r a l o u t l i n e of l o w e r j a w s l i g h t l y c o n v e x , m e n t a l
area s m o o t h l y r o u n d e d . V e n t r a l o u t l i n e or l o w e r j a w almost straight, l o w e r j a w sides
v e r y s l i g h t l y o b l i q u e . C e p h a l i c lateral line o p e n i n g s not e n l a r g e d .
S c a l e s . — C h e e k , g i l l - c o v e r , n a p e , neck, d o r s u m a n d v e n t r a l s i d e of chest w i t h
c y c l o i d scales. W e a k l y c t e n o i d scales o n lateral parts of chest. R e m a i n i n g parts of
b o d y w i t h c t e n o i d scales. A g r a d u a l size transition b e t w e e n chest a n d scales of flank.
S m a l l , elongate c y c l o i d scales present o n the p r o x i m a l t w o - t h i r d of the c a u d a l f i n .
F i n s . — P e l v i c fins not r e a c h i n g , pectoral fins just not r e a c h i n g a n a l f i n o r i g i n .
D o r s a l a n d a n a l fins not r e a c h i n g c a u d a l f i n base. P e l v i c fins w i t h first ray s l i g h t l y
p r o d u c e d . C a u d a l f i n o u t l i n e subtruncate, straight.
G i l l a p p a r a t u s . — E i g h t g i l l rakers o n the l o w e r part of the first g i l l arch; the l o w -
ermost t w o short, c o n i c a l , the f o l l o w i n g t w o elongated, c o n i c a l , a n d the r e m a i n i n g
f o u r flattened, the u p p e r m o s t t w o w i t h e x p a n d e d tips. A p p r o x i m a t e l y 125 g i l l f i l a -
m e n t s o n the first h e m i b r a n c h .
V i s c e r a . — Intestine l e n g t h is 1.1 times s t a n d a r d length. F o l l o w i n g the d e f i n i t i o n s
of Z i h l e r (1982) the arrangement of the digestive tract is of t y p e D ( w i t h b a c k f l a p ) .
O r a l t e e t h . — Shape: O u t e r r o w s i n b o t h jaws c o n t a i n r e l a t i v e l y slender u n i c u s -
p i d s , r o s t r a l l y circular i n cross section laterally a n d c a u d a l l y s o m e w h a t c o m p r e s s e d .

Several c o m p r e s s e d teeth h a v e a s h o u l d e r o n o n e side, f e w w i t h a v e r y s m a l l cusp.
Teeth i n the l o w e r j a w s o m e w h a t stouter t h a n those i n u p p e r j a w L o w e r j a w teeth
s l i g h t l y c u r v e d , those i n the u p p e r j a w m o d e r a t e l y c u r v e d . L a t e r a l teeth i n u p p e r j a w
s u s p e n s o r i a d i n c l i n e d . T o o t h size g r a d u a l l y decreasing f r o m rostral to c a u d a l . Inner
r o w s h a v e a m i c t u r e o f b i c u s p i d s a n d t r i c u s p i d teeth w i t h a protracted major c u s p , i n
the l o w e r j a w s o m e s h o u l d e r e d u n i c u s p i d s are also f o u n d . A l l i n n e r teeth are c o m -
pressed a n d m o d e r a t e l y c u r v e d .
— D e n t a l arcade a n d toothband. Dental arcade r o u n d e d . T w o inner r o w s i n
l o w e r jaw, three o r f o u r i n u p p e r jaw. Distance b e t w e e n outer r o w a n d first i n n e r r o w
normal.
— C o u n t s a n d setting. There are 60 teeth i n the outer r o w o f the u p p e r j a w a n d
46 i n the outer r o w of the l o w e r jaw. O u t e r r o w teeth are r e g u l a r l y set at a distance of
s l i g h t l y less t h a n the toothbase diameter.
— I m p l a n t a t i o n . O u t e r teeth i n u p p e r j a w erect (rostrally) to r e c u m b e n t (caudal-
l y ) , i n l o w e r j a w erect t o s l i g h t l y r e c u m b e n t . I n n e r teeth r e c u m b e n t to s t r o n g l y
r e c u m b e n t . T h e teeth are i n b e d d e d i n a thick layer of tissue w h i c h covers the p r o x i -
m a l 4/5 part of the outer r o w teeth a n d almost the entire inner teeth.
O s t e o l o g y . — T h e h o l o t y p e w a s n o t dissected to o b t a i n skeletal e l e m e n t s f o r a
detailed description.
— Vertebrae. There are 30 vertebrae, c o m p r i s i n g 13 a b d o m i n a l a n d 17 c a u d a l ele-
ments.
C o l o r a t i o n . — M a l e c o l o r a t i o n is u n k n o w n .
— T h e l i v e female has the d o r s a l part of h e a d a n d b o d y d a r k b r o w n . L i p s a n d
cheek are d a r k g r e y i s h . A faint l a c h r y m a l stripe, w h i c h is m o r e distinct dorsally, r u n s
f r o m the r o s t r o v e n t r a l m a r g i n of the e y e to the p r e o p e r c u l a r h o r i z o n t a l a r m . Ventral
p a r t of g i l l - c o v e r black. I n t e r m a n d i b u l a r area a n d branchiostechal m e m b r a n e cream
c o l o u r e d . C h e s t a n d v e n t r a l parts of b o d y a n d c a u d a l p e d u n c l e d a r k i s h grey. Ventral
s i d e w h i t e . D o r s a l f i n d a r k b r o w n to black basally, t u r n i n g lighter distally. L a p p e t s
a n d d i s t a l area of the r a y e d part of the d o r s a l r e d . Pectoral fins h y a l i n e . P e l v i c fins
b l a c k w i t h a s m a l l r e d area o n the m e d i a l l y part. A n a l f i n b r i g h t r e d except f o r the
h y a l i n e c a u d a l part i n w h i c h t w o b r i g h t orange spots are f o u n d . T h e rostal most, a n d
larger spot seems to b e s u r r o u n d e d b y a black r i n g . C a u d a l f i n d a r k b r o w n p r o x i m a l -
ly, b r i g h t r e d d i s t a l l y a n d a l o n g the v e n t r a l m a r g i n .
— T h e p r e s e r v e d female has a b r o w n g r o u n d c o l o u r . C h e s t a n d v e n t r a l s i d e are
l i g h t b r o w n i s h . L a c h r y m a l s t r i p e v e r y faint, a s m a l l o p e r c u l a r present. D o r s a l f i n
h y a l i n e w i t h p r o x i m a l l y d a r k e r areas b e t w e e n the spines a n d rays. Pectorals h y a l i n e ,
p e l v i c s d a r k i s h , A n a l h y a l i n e , w i t h b a s a l l y d a r k s areas b e t w e e n the rays a n d o n e rel-
a t i v e l y large w h i t e spot s u r r o u n d e d b y a d a r k r i n g near the c a u d a l m a r g i n . A n a l f i n
d a r k p r o x i m a l l y , h y a l i n e d i s t a l l y , w i t h d a r k stripes b e t w e e n the rays o f the d o r s a l
half.
Distribution.— O n l y k n o w n from Lake Victoria.
E c o l o g y . — H a b i t a t . Haplochromis kujunjui w a s caught w i t h a b o t t o m t r a w l over a
s a n d b o t t o m w i t h a d e p t h o f 5-8 m i n a r e l a t i v e l y e x p o s e d area.
— F o o d . E x a m i n a t i o n o f s t o m a c h a n d g u t contents s h o w e d that Haplochromis
kujunjui is a p i s c i v o r o u s species. T h e g u t w a s f i l l e d w i t h the r e m a i n s o f at least 10
v e r y s m a l l (postbuccal) h a p l o c h r o m i n e c i c h l i d s .
Table 10. Linear measurements of Haplochromis kujunjui spec. nov.
Holotype
R M N H reg. no. 30466
Standard Length 102.8

Body Depth %SL 51.3
Pectoral Fin Length %SL 24.4
Caudal Peduncle Length %SL 15.0
Caudal Peduncle Depth %SL 11.6
Caudal Fin Length %SL 22.3
Head Length %SL 35.9
Snout Length %HL 35.9
Snout Width %HL 29.7
Head Width %HL 42.7
Interorbital Width %HL 20.0
Preorbital Width %HL 26.4
Lachrymal Width %HL 27.0
Preorbital Depth %HL 18.3
Eye Length %HL 22.9
Cheek Depth %HL 21.0
Lower Jaw Length %HL 48.9
Lower Jaw Width %HL -
Upper Jaw Length %HL 37.5
Premax. Pedicel Length %HL 29.7
Table 11. Qualitative measurements and counts of Haplochromis kujunjui spec. nov.
Dorsal Head Profile (curvature) 0(+)

Premaxillary Pedicel Prominence 0(+)?*
Lower Jaw Extension 0(+)?
Lateral Snout Outline OW?
Mental Prominence 0
Lip Thickening 0
Premaxilla Beaked 0
Maxillary Posterior Extension 0
Lateral Line Scales 32

Lateral Line - Dorsal Fin (Sc.rows) 6
Pectoral - Pelvic Fin Bases (Sc.rows) 6
Cheek (Vertical Sc.rows) 5
Dorsal Fin (Spines/Rays) XVI9
Anal Fin (Spines/Rays) III 9
* ) ? = Measurements affected by protrusion.

V A N OIJE N: PISCIVOROUS H A P L O C H R O M I N E CICHLIDAE O F L A K E VICTORIA 57
— B r e e d i n g a n d g r o w t h . T h e h o l o t y p e is a m a t u r e , a l m o s t r i p e f e m a l e . T h e r i g h t
o v a r y i s larger.
R e s e m b l i n g s p e c i e s . — H. kujunjui is d i s t i n g u i s a b l e f r o m a l l o t h e r h a p l o c h r o m i n e
species of L a k e V i c t o r i a b y the f e m a l e c o l o r a t i o n . T h e species is treated w i t h the
dichrourus-complex b e c a u s e the f e m a l e c o l o r a t i o n g r e a t l y r e s e m b l e s that o f a m a l e ; it
has b l a c k p e l v i c f i n s (albeit w i t h a r e d m e d i a l area), b r i g t h l y c o l o u r e d a n a l a n d c a u
d a l f i n s , a n d t w o d i s t i n c t e g g d u m m i e s o n t h e a n a l f i n . H o w e v e r , this species does
n o t posess s i m a r l y s t r o n g l y c u r v e d teeth as H. dichrourus a n d H. maisomei.
H a p l o c h r o m i s p y r r h o p t e r y x spec. n o v .
(figs 6 , 6 3 7 8 , 111, tables. 1214)
Haplochromis dichrourus (part); Greenwood, 1967: 6569; specimen BM(NH) 1966.3.9.185.

Prognathochromis (P.) dichrourus (part); Greenwood, 1980:19.
Haplochromis "redback"; van Oijen, 1982: 341.
Material.— Holotype, cf, 154.3 mm, R M N H 30010, Mwanza Gulf, Lake Victoria, Tanzania, 25.viii,
HEST. Paratypes: 1 cf, 113.0 mm + 1 9, 105 mm, R M N H 3038485, Mwanza Gulf, Tanzania, Lake
Victoria, vi.1975, G.Ch. Anker & C . D . N . Barel; Other paratypes from the Mwanza Gulf, collected by
HEST; 2 99, 95.1, 99.5 mm, R M N H 3001112, 4.vi.l977; 2 99, 97.2, 112.9 mm, R M N H 3001314,
18.xi.1977; 1 9, 91.1 mm, R M N H 30015,16.xii.1977; 1 9,134.0 mm, R M N H 30016, Mwanza Gulf, L.V.,
Tanzania, 4.1.1978; 1 9, 107.0 mm, R M N H 30017, 27.U978; 1 9, 99.1 mm, R M N H 30018, Tanzania
27.1.1978, HE ST; 2 Cef, 107.0, 112.0 mm + 1 9, 192.5 mm, R M N H 3001921, 6.Ü.1978; 1 cf, 91.0 mm,
R M N H 30022, 10.iii.1978; 1 cf , 137.1 mm, R M N H 30023, 12.V.1978; 1 9 , 118.8 mm, R M N H 30024,
26.5.1978; 1 9, 107.4 m m , R M N H 30025, 19.vi.1978; 2 99, 87.6, 118.3 m m , R M N H 3002728,
ll.viii.1978; 1 9, 90.3 mm, R M N H 30029, 18.viii.1978; 3 cfcf, 84.9123.7 mm + 1 9, 103.5 mm, R M N H
3003033, 7.XÍ.1978; 1 cf, 75.4 mm, R M N H 30034, Butimba Bay, 27.xi.l9781; 9,94.0 mm, R M N H 30035,
15.xii.1978;. 1 cf, 107.0 mm, R M N H 30037, Nyegezi Bay, 23.X.1979; 1 9, 91.8 mm, R M N H 30382,
7.XÜ.1979; 1 9, 178.3 mm, R M N H 30038, 22.iv.1980; 3 cfcf, 112.5154.2 mm + 2 99, 95.9, 107.8 mm,
R M N H 3003943, 25.viii.1981; 1 cf, 142.2 mm, B M N H 1988.2.4:16, 25.viii.1981; 1 9, 69.2 mm, R M N H
30048, 6.vii,1982; 1 9,100.8 mm, R M N H 30045,19.X.1982;; 1 cf, 107.6 mm, R M N H 30047, 26.X.1982; 1
cf, 116.0 mm, R M N H 30049, 17.1.1983; 1 9, 133.1 mm, M N H N 19871669, 17.Í.1983; 1 9, 108.4 mm,
ANSP 168241, 1978; 1 cf, 115.8 mm, B M N H 1987.2.4.17,1978; 5 cfcf, 88.2135.9 + 3 99, 93.4150.8 mm,
R M N H 3005057, 1978. Paratypes from other localities collected by HEST; 1 9, 111.0 mm, R M N H
30046, Bukinga Bay, Tanzania, Lake Victoria, 22.X.1982; 1 9, 138.0 mm, R M N H 30026, off Nafuba
Island, Speke Gulf, Tanzania, Lake Victoria, 2.viii.l978; 1 9, 117.0 mm, R M N H 30036, off Ukerewe
Island, Tanzania, Lake Victoria, 6.vi.l979; 1 9, 77.2 mm, R M N H 30379, off Standieri Island, Tanzania,
Lake Victoria, 22.iii.1983; . Other material: 1 cf, 173.7 mm, B M N H 1966.3.9.185, off Soswa Island,
Tanzania, Lake Victoria, J.D. Kelsall; 2 cfcf, 123.0, 197.0 mm, R M N H 3038081, Kavirondo Gulf,
Kenia, Lake Victoria, 1976, R.S. Benda.
E t y m o l o g y . T h e s p e c i f i c l n a m e ( f r o m the G r e e k " π υ ρ ρ ο δ " = r e d d i s h , y e l l o w r e d ,

a n d " π τ ε ρ υ ε " = w i n g , f i n ) refers to the b r i g h t , o r a n g e r e d c o l o r a t i o n o f t h e m e d i a n
f i n s w h i c h is p r e s e n t i n l i v e s p e c i m e n s o f b o t h sexes, j u v e n i l e s as w e l l as a d u l t s .
D i a g n o s i s . A m o d e r a t e l y large a n d slender p i s c i v o r o u s species w i t h strongly
r e c u r v e d u n i c u s p i d teeth i n the o r a l j a w s . L i v e c o l o u r s f o r m a l e s a n d f e m a l e s are
black o n ventral parts of head a n d b o d y a n d orange dorsally. A l l m e d i a n fins orange.
M a l e s a n d females have black p e l v i c fins.
D e s c r i p t i o n . — B a s e d o n 49 s p e c i m e n s ( i n c l u d i n g t h e h o l o t y p e ) 75.4192.5 m m S L .
H a b i t u s . — B o d y m o d e r a t e l y e l o n g a t e , the h e a d h a v i n g a r a t h e r " h e a v y " appear
ance. D o r s a l h e a d p r o f i l e s l i g h t l y c u r v e d , f r o m n a p e to u p p e r l i p n e a r l y s t r a i g h t i n
Fig. 63. Haplochromis pyrrhopteryx spec, nov., para type ( R M N H 30023), SL 137.1 mm, lateral view. Scale
equals 10 mm.
s o m e specimens. P r e m a x i l l a r y p e d i c e l h a r d l y to m o d e r a t e l y i n t e r r u p t i n g the profile.

Snout somewhat blunt. M o u t h moderately oblique. Premaxilla not or slightly
e x p a n d e d m e d i a l l y . A n elongate area of the m a x i l l a e x p o s e d d o r s a l l y a n d c a u d a l l y to
the c a u d a l part of the p r e m a x i l l a . Posterior t i p o f the m a x i l l a just p a s s i n g o r n o t quite
r e a c h i n g a v e r t i c a l t h r o u g h t h e anterior eye m a r g i n . L i p s n o t t h i c k e n e d , l o w e r j a w
i s o g n a t h o u s o r s l i g h t l y p r o g n a t h o u s , t i p of the l o w e r j a w a l w a y s s l i g h t l y p r o t r u d i n g .
R o s t r a l o u t l i n e of l o w e r j a w s l i g h t l y convex. M e n t a l area r o u n d e d o r w i t h a s m a l l
m e n t a l p r o m i n e n c e . Sides o f l o w e r j a w s l i g h t l y to m o d e r a t e l y o b l i q u e . H o r i z o n t a l
a r m of preoperculum horizontal, vertical a r m vertical to slightly rostrad inclined.
C e p h a l i c lateral l i n e o p e n i n g s n o t e n l a r g e d , lateral l i n e canals o n the l a c h r y m a l n o t
v i s i b l e . E y e m o d e r a t e l y large a n d circular, p u p i l subcircular, a s m a l l a p h a k i c space
a l w a y s present r o s t r o - v e n t r a l l y to the lens.
S c a l e s . — C h e e k a n d n u c h a l area w i t h c y c l o i d scales o n l y , the g i l l cover w i t h m o r e
c y c l o i d scales t h a n w e a k l y c t e n o i d scales. Scales o n nape a n d b o d y c t e n o i d , those o n
n a p e a n d d o r s u m often w e a k l y c t e n o i d , chest w i t h rather s m a l l c t e n o i d scales. S m a l l
elongate scales o n the p r o x i m a l half of the c a u d a l f i n .
F i n s . — Pectoral a n d p e l v i c fins n o t quite r e a c h i n g the a n a l f i n . A n a l a n d d o r s a l
f i n n o t r e a c h i n g base o f c a u d a l . P e l v i c s w i t h the first soft r a y s l i g h t l y p r o d u c e d .
C a u d a l f i n o u t l i n e truncate a n d s l i g h t l y emarginate.
G i l l a p p a r a t u s . — E i g h t to ten g i l l rakers o n the l o w e r part of the first g i l l arch,
the l o w e r m o s t t w o o r three r e d u c e d , the next t w o o r three slender a n d p o i n t e d , a n d
the r e m a i n i n g three o r f o u r flattened w i t h the top r o u n d e d , b i f i d o r t r i f i d . There are
a p p r o x i m a t e l y 132 g i l l filaments o n the first h e m i b r a n c h .
V i s c e r a . — Intestine l e n g t h ranges f r o m 0.9 to 1.6 (mean 1.24) times the s t a n d a r d
l e n g t h (n= 13). F o l l o w i n g t h e d e f i n i t i o n s o f Z i h l e r (1982) t h e a r r a n g e m e n t o f t h e
d i g e s t i v e tracks of a d u l t s is of type D ( w i t h backflap).
O r a l t e e t h . — Shape. A l l specimens h a v e rather stout, r o u n d e d , s t r o n g l y c u r v e d
a n d acutely p o i n t e d u n i c u s p i d s teeth i n the outer r o w of b o t h jaws. I n s m a l l e r speci-
m e n s (90 m m ) a f e w w e a k l y b i c u s p i d s m a y be present caudally. C u r v a t u r e i s almost
c o n f i n e d to u p p e r h a l f of the teeth. Tooth size decreases g r a d u a l l y f r o m r o s t r a l to

c a u d a l . In the inner r o w s , s l i g h t l y c o m p r e s s e d u n i c u s p i d s d o m i n a t e o v e r w e a k l y t r i -
a n d b i c u s p i d s . A l l i n n e r r o w teeth are s t r o n g l y c u r v e d a n d s l i g h t l y c o m p r e s s e d .
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r r o w o c c u p i e s
9/10 of p r e m a x i l l a dentigerous a r m , a n d o n d e n t a r y does not reach c o r o n o i d w i n g .
B o t h j a w s h a v e t w o i n n e r r o w s . L e n g t h of i n n e r r o w o n p r e m a x i l l a 1/2 outer r o w
l e n g t h , o n d e n t a r y 4/5 of outer r o w l e n g t h . D i s t a n c e b e t w e e n outer r o w a n d first
i n n e r r o w o n l y s l i g h t l y larger than distance b e t w e e n i n n e r r o w s .
— C o u n t s a n d setting. There are 52-64 teeth i n outer r o w of the p r e m a x i l l a , a n d
36-56 teeth i n outer r o w of the l o w e r jaw. T h e n u m b e r s h o w s a p o s i t i v e correlation
w i t h s t a n d a r d length. O u t e r teeth s o m e w h a t i r r e g u l a r l y set at a distance of a half to
m o r e t h a n once the d i a m e t e r of the tooth base. T h e irregular setting m i g h t be d u e to
the fact that m a n y of the d e s c r i b e d specimens h a v e not yet reached a n a d u l t size.
— I m p l a n t a t i o n . In u p p e r j a w rostral teeth s l i g h t l y recumbent, c a u d a l teeth m o d -
erately r e c u m b e n t . Teeth i n l o w e r j a w are s l i g h t l y p r o c u m b e n t r o s t r a l l y a n d erect
c a u d a l l y . I n b o t h j a w s the teeth are s l i g h t l y s u s p e n s o r i a d i n c l i n e d . Teeth of i n n e r
r o w s s t r o n g l y r e c u m b e n t , second i n n e r r o w s i m p l a n t e d a l m o s t h o r i z o n t a l l y .
P h a r y n g e a l t e e t h . — C o u n t s . A p p r o x i m a t e l y 22 teeth i n c a u d a l m o s t transverse
r o w a n d 11 i n the t w o m e d i a n r o w s .
— Shape. Teeth of the c a u d a l m o s t r o w a n d 2 or 3 c a u d a l l y i n the m e d i a n r o w are
h o o k e d , the b l u n t major c u s p p o i n t i n g dorso-rostrad. O t h e r teeth are r e l a t i v e l y fine,
acutely p o i n t e d a n d b e v e l l e d . Tooth size increases f r o m rostral to c a u d a l a n d f r o m
lateral to m e d i a l .
O s t e o l o g y . — T h e o s t e o l o g i c a l d e s c r i p t i o n s of n e u r o c r a n i u m , p r e m a x i l l a , l o w e r
j a w a n d p h a r y n g e a l e l e m e n t are b a s e d o n f i v e s p e c i m e n s , R M N H 30033, 30017,
30019, 30022 a n d 30050, S L 94.0 to 133.0 m m . D e s c r i p t i o n of the o r a l teeth is b a s e d o n
a l l t y p e specimens.
— N e u r o c r a n i u m . C o m p a r e d to the n e u r o c r a n i u m of a g e n e r a l i z e d c i c h l i d , Ha-
plochromis elegans (see Barel et a l . , 1976), that of H. pyrrhopteryx is m o r e slender as it
has a s h a l l o w e r otic r e g i o n . T h e d e p t h of the s u p r a o c c i p i t a l crest is a p p r o x i m a t e l y
the same. T h e e t h m o v o m e r i n e b l o c k is relatively longer t h a n i n H. elegans a n d less
d e c u r v e d . T h e b r o a d p o s t o r b i t a l process is situated r e l a t i v e l y h i g h i n the n e u r o c r a n i -
u m . A s i n m o s t p i s c i v o r o u s c i c h l i d s f r o m L a k e V i c t o r i a the p h a r y n g e a l a p o p h y s i s is
situated far caudally.
— O r a l jaws. P r e m a x i l l a w i t h a s c e n d i n g a r m l o n g e r t h a n dentigerous a r m , angle
b e t w e e n the a r m s 75-81°. A s c e n d i n g a r m w i t h a s l i g h t l y c o n v e x o u t l i n e rostrally.
T r a n s i t i o n f r o m a s c e n d i n g a r m to relatively t h i n dentigerous a r m b r o a d . O u t l i n e of
t o o t h b e a r i n g side of dentigerous a r m straight to s l i g h t l y concave. M a n d i b l e w i t h rel-
a t i v e l y l o n g d e n t i g e r o u s p a r t w h i c h is o n l y s l i g h t l y less t h a n h a l f of l o w e r j a w
l e n g t h . D e n t i g e r o u s area s l i g h t l y d e c u r v e d . L e n g t h / d e p t h ratio of l o w e r j a w 2.3.
— L o w e r p h a r y n g e a l element (fig. 40) rather s h a l l o w , s l i g h t l y longer than b r o a d
( l e n g t h / w i d t h = 1.1). Dentigerous area slightly broader than l o n g ( l e n g t h / w i d t h = 0.9).
—Vertebrae: There are 29 or 30 vertebrae c o m p r i s i n g 13 a b d o m i n a l a n d 16 or 17
c a u d a l elements.
C o l o r a t i o n . — L i v e s e x u a l l y active males: E t h m o i d a l area a n d d o r s a l h e a d surface
b r i g h t o r a n g e - r e d , i n t e r r u p t e d b y d a r k a n d d i s t i n c t n o s t r i l a n d i n t e r o r b i t a l stripes
a n d a fainter s u p r a o r b i t a l b a n d , n a p e b a n d a n d n a p e b l o t c h . D o r s a l part of g i l l - c o v e r
Figs 64-76. Haplochromis pyrrhopteryx spec. nov. Fig. 64. Premaxilla, lateral view. Fig. 65. Premaxilla,
view perpendicular to dentigerous area. Fig. 66. Premaxilla, medial view of ascending arm. Fig. 67.
Lower jaw, lateral view. Fig. 68. Lower pharyngeal element, dorsal view. Fig. 69. Lower pharhyngeal
labial and lateral view. Fig. 71. First tooth of outer row. Fig. 72. Seventh tooth of outer row. Fig. 73.
Fifth tooth of 1st inner row. Figs. 74-76. Lower pharhyngeal teeth, lateral and rostral view. Fig. 74.
Seventh lateral tooth. Fig. 75. Sixth medial tooth of caudalmost row. Fig. 76. Second medial tooth cau-
dalmost row. Scale of bony elements = 5 mm, scale of teeth = 1 mm.
orange, a b l a c k o p e r c u l a r b l o t c h present. R e m a i n i n g lateral a n d v e n t r a l parts of h e a d

entirely black. B o d y : d o r s u m a n d f l a n k b r i g h t orange, m e r g i n g i n t o a d u l l e r orange
a n d f i n a l l y b l a c k c a u d a l l y , w h e r e iridescent green scales m a y be present. L o w e r part
of the f l a n k w i t h a y e l l o w sheen. Chest, central part of belly, v e n t r a l side a n d c a u d a l
peduncle black. Fins: pectoral hyaline, pelvics black. A n a l light r e d d i s h rostrally
turning into reddish-white caudally; two small yellow-orange eggdummies w i t h a
d a r k r i m d o r s a l l y o n the f i n . D o r s a l f i n l i g h t o r a n g e - r e d . S o m e d a r k e r r e d streaks
b e t w e e n the soft rays. C a u d a l f i n b r i g h t l i g h t r e d .
— Q u i e s c e n t males h a v e smaller a n d less b r i g t h orange areas o n the flank. T h e
o r a n g e - r e d f l u s h o n the m e d i a l fins is also r e d u c e d l e a v i n g the c a u d a l parts of the
fins h y a l i n e .
— L i v e f e m a l e s h a v e a c o l o r a t i o n c l o s e l y r e s e m b l i n g that of quiescent m a l e s :
H e a d b l a c k v e n t r a l l y . N a p e a n d neck sooty w i t h a n o v e r l y i n g o r a n g e f l u s h . Inter-
o r b i t a l area l i g h t orange, snout area orange b a r r e d b y the b l a c k n o s t r i l a n d interor-
b i t a l stripes. C r o s s b a n d s o n the d o r s a l h e a d surface faint. B o d y black, w i t h o v e r l y i n g
l i g h t o r a n g e f l u s h o n the rostral parts of d o r s u m a n d flank. F i n s : pectorals h y a l i n e ,
p e l v i c black, a n a l l i g h t y e l l o w - o r a n g e , b e c o m i n g a less intense y e l l o w - o r a n g e , almost
h y a l i n e d i s t a l l y , 1-4 o r a n g e spots o n the c a u d a l p a r t . D o r s a l l i g h t o r a n g e , d a r k e r
orange spots b e t w e e n the soft rays. C a u d a l l i g h t orange-red.
— Juveniles h a v e the same c o l o r a t i o n as females, b u t the orange f l u s h o n d o r s u m
a n d neck is v e r y faint.
— P r e s e r v e d m a l e s h a v e d o r s a l part of snout, i n t e r o r b i t a l area a n d n a p e l i g h t
i v o r y . N o s t r i l stripe distinct, e x t e n d e d o v e r the nostrils a n d , f r o m a p o i n t s l i g h t l y
m e d i a l of this, e x t e n d i n g rostrad to the u p p e r l i p . Interorbital stripes not r e a c h i n g the
eye m a r g i n a n d n o t m e e t i n g i n the m i d l i n e . S u p r a o r b i t a l b a n d rather faint, n a p e
b a n d b r o a d a n d distinct, nape b l o t c h faint. F r o m the eye a b r o a d black area stretches
across the l a c h r y m a l to the u p p e r jaw. C h e e k d a r k g r e y - b r o w n , g i l l - c o v e r darker, pre-
o p e r c u l u m black, l i p s d a r k i s h b r o w n , l o w e r j a w a n d v e n t r a l part of h e a d black. B o d y
g e n e r a l l y d a r k e r grey, lighter dorsally. Chest, b e l l y a n d v e n t r a l side black. F i n s : pec-
toral g r e y i s h , p e l v i c s black. A n a l , d o r s a l a n d c a u d a l v e r y l i g h t y e l l o w i s h .
— P r e s e r v e d females a n d juveniles. D o r s a l part of h e a d l i g h t y e l l o w i s h - g r e y . T h e
b a n d s i n this area b e i n g m u c h less distinct t h a n i n males (especially the b a n d o n the
nape). C h e e k d a r k b r o w n i s h , snout s l i g h t l y darker, r e m a i n i n g parts of h e a d black.
B o d y w i t h a s m a l l area of the d o r s a l part l i g h t b r o w n i s h a n d a larger d a r k b r o w n -
b l a c k area o n the v e n t r a l part. In the lighter part 4 vertical b a n d s m a y be v i s i b l e o n
the d o r s a l flank. F i n s : pectoral h y a l i n e , p e l v i c s black, a n a l , d o r s a l a n d c a u d a l l i g h t
yellowish.
D i s t r i b u t i o n . — H. pyrrhopteryx is o n l y k n o w n f r o m L a k e V i c t o r i a . T h e m a j o r i t y of
the specimens f r o m the H E S T collections w e r e c a p t u r e d i n the n o r t h e r n part of the
M w a n z a G u l f , b u t specimens w e r e also collected off U k e r e w e I s l a n d , a n d near N a f u -
b a Island i n the S p e k e G u l f . T h e s p e c i m e n B M ( N H ) 1967.3.9.185 f o r m e r l y i n c l u d e d i n
H. dichrourus w a s caught off S o s w a I s l a n d . T w o specimens c a m e f r o m the K a v i r o n d o
G u l f (Kenya). A l t h o u g h the areas are relatively far apart the m o r p h o m e t r i c measure-
ments a n d the c o l o r a t i o n of the specimens (coloration of the specimens f r o m the K a -
v i r o n d o G u l f is k n o w n f r o m p h o t o g r a p h s ) are v e r y similar.
E c o l o g y . — Occurrence. In the p e r i o d f r o m 1977-1980 H. pyrrhopteryx w a s a rare
species i n the M w a n z a G u l f area. Despite the fact that w e searched the catches for
Fig. 77. Haplochromis pyrrhopteryx spec. nov. Live colours and markings of a sexually active male and a
female.
s p e c i m e n s of this c o n s p i c u o u s l y c o l o u r e d species, w e collected o n l y 45 s p e c i m e n s

f r o m 1977 to 1980. Since the e x p l o s i v e increase of the N i l e p e r c h p o p u l a t i o n i n the
M w a n z a G u l f H. pyrrhopteryx has d i s a p p e a r e d f r o m the catches.
— H a b i t a t . Because of the f e w specimens e n c o u n t e r e d i n the catches it is n o t easy
to get a n i d e a of the p r e f e r r e d habitat of this species. It is r e m a r k a b l e that almost a l l
k n o w n s p e c i m e n s are i m m a t u r e . T h e smallest w e r e caught i n bays o v e r a m u d bot-
t o m at a d e p t h of 2-4 meters. T h e l e n g t h of the specimens seems to increase w i t h the
d e p t h of the w a t e r i n w h i c h they are caught. T h e m a j o r i t y of the s p e c i m e n s w e r e
c a u g h t i n a b o t t o m t r a w l o v e r m u d b o t t o m s w i t h a d e p t h of 10-16 m . T h e largest
s p e c i m e n w a s caught at the entrance to the M w a n z a G u l f at a d e p t h of 20 m . T h i s
suggests that large specimens n o r m a l l y l i v e i n deeper, e x p o s e d waters w i t h a m u d d y
b o t t o m a n d m a y c o m e to s h a l l o w waters o n l y to b r e e d .
— F o o d . T h i r t e e n intestines o f 23 e x a m i n e d s p e c i m e n s of H. pyrrhopteryx c o n -
t a i n e d r e m a i n s of s m a l l h a p l o c h r o m i n e c i c h l i d s . W h e r e the size of the p r e y c o u l d be
c a l c u l a t e d it w a s f o u n d that s p e c i m e n s of a b o u t 90 m m S L p r e y e d o n h a p l o c h r o -
m i n e s of 15-25 m m , a s p e c i m e n of 113 m m S L h a d f e d o n h a p l o c h r o m i n e s of 35 m m ,
a n d s p e c i m e n s of 136 a n d 142 m m S L f e d o n h a p l o c h r o m i n e s of respectively 40 a n d
60 m m . Therefore it seems that p r e y - s i z e increases w i t h p r e d a t o r size. I n t w o speci-
Fig. 78. Catch localities of Haplochromis pyrrhopteryx spec. no v., H. pellegrini Regan, 1922, and H.
harpakteridion spec. nov. in Lake Victoria.
Table 12. Ranges of linear measurements of Haplochromis pyrrhopteryx spec. nov. For convenience the ratios are multiplied by 100.
SL in mm 75.4-88.2 91.0-99.5 100.8-108.14 110.0-118.8 123.7-138.0 142.0-154.3

Ν 4 11 10 9 6 5
BD/SL 28.4-31.6 28.5-32.3 28.3-32.4 30.7-33.2 29.3-30.4 31.8-33.7

PFL/SL 24.2-30.5 26.9-31.8 25.7-30.6 23.4-29.5 26.6-31.1 24.1-29.9
CPL/SL 16.7-18.9 15.5-18.1 14.6-18.8 14.8-17.1 14.2-18.0 14.4-16.6
CPD/SL 10.8-11.9 10.0-11.9 10.5-11.8 10.7-11.5 10.6-11.3 10.8-11.8
CFL/SL 19.0-23.8 20.7-25.3 21.0-24.1 21.6-23.4 21.6-23.7 20.8-22.2
HL/SL 35.8-36.8 35.6-38.1 35.6-38.1 35.8-37.9 35.6-37.8 34.9-37.6
SnL/HL 29.2-32.3 29.4-33.6 31.6-35.1 32.7-36.1 33.1-36.9 33.6-36.2
SnW/HL 31.3-35.2 27.7-32.7 28.7-36.6 29.5-37.4 30.4-38.0 31.0-36.4
HW/HL 42.5-45.2 38.2-44.1 39.9-42.8 40.0-46.0 41.5-44.5 42.9-17.2
IOW/HL 20.9-22.7 20.8-23.0 22.0-24.0 22.0-24.3 22.2-23.9 23.3-24.0
POW/HL 25.4-27.5 23.4-27.7 25.0-26.9 25.1-27.5 25.4-27.4 27.5-28.7
LaW/HL 26.4-29.8 22.7-29.3 24.5-28.2 26.8-29.1 25.2-30.1 26.7-30.9
POD/HL 15.1-17.5 15.7-17.8 17.0-20.1 16.8-20.3 19.0-19.5 18.1-19.7
EyL/HL 23.0-25.2 21.2-26.5 20.7-25.5 20.2-26.0 19.8-22.0 20.0-22.5
ChD/HL 21.8-22.2 20.9-22.7 21.4-24.3 21.6-24.0 23.0-27.9 25.3-27.9
LJL/HL 44.8-47.7 44.8-49.7 47.7-50.7 46.8-51.6 48.7-50.1 50.0-54.7
LJW/HL 19.8-21.2 18.2-22.2 18.2-22.5 18.3-25.6 19.2-26.5 20.1-25.7
UJL/HL 34.6-38.7 34.8-38.3 35.3-38.8 34.4-39.7 39.2-41.7 40.3^2.2
PPL/HL 25.3-28.3 25.9-295 27.0-29.7 26.7-30.0 27.8-29.4 26.7-27.7
Table 13. Means and standard deviations of linear measurements of Haplochromis pyrrhopteryx spec. nov. For convenience the ratios are multiplied by 100. Ζ
— S
SL in mm 75.4-88.2 91.0-99.5 100.8-108.4 111.0-118.8 123.7-138.0 142.2-154.3 178.3 192.5 m
Ζ
Ν 4 11 11 9 6 5 1 1
30.310.7 32.811.5 32.4 32.9

η
BD/SL 29.7±1.6 30.211.2 30.311.4 31.711.0
PFL/SL 27.6±2.6 28.111.2 28.511.2 28.411.2 28.713.6 27.412.0 29.3 27.6 ο
CPL/SL 17.8±1.1 16.611.0 16.511.0 16.511.0 15.711.4 15.310.8 16.1 16.6 ο
CPD/SL 11.4±0.5 11.0t0.5 11.010.4 11.210.2 10.910.3 11.210.3 11.2 11.0 S
CFL/SL 21.6±1.9 22.911.4 22.311.0 22.510.6 22.510.8 21.710.5 21.2 19.7 oc
>
36.710.6 36.810.8 36.910.8 36.411.0 36.4 37.4 TJ
HL/SL 36.410.5 37.0±0.6
SnL/HL 31.011.6 31.611.2 33.411.3 33.411.3 34.211.1 35.010.9 37.2 39.8
33.112.7 34.412.2 34.312.3 38.6 38.2
R
SnW/HL 33.312.0 30.411.5 31.612.0
HW/HL 43.811.3 41.311.9 41.510.9 43.311.7 42.511.3 44.511.5 43.2 45.4
IOW/HL 21.911.0 21.510.6 21.610.6 22.610.6 22.710.7 23.610.4 23.5 25.5
POW/HL 26.211.0 25.411.3 24.412.2 26.510.8 26.310.8 27.910.6 26.6 29.4 1
UW/HL 27.511.5 25.811.6 26.111.1 27.911.0 28.612.5 29.512.0 27.8 32.8
POD/HL 16.411.2 16.610.8 17.910.8 18.0±0.7 19.010.6 18.910.7 20.0 22.0 η
23.411.9 22.111.6 21.210.9 20.4 20.5
η
EyL/HL 24.0±1.1 22.210.6 23.411.9 DC
r
ChD/HL 22.110.1 21.710.7 22.410.8 22.410.8 24.511.0 26.610.7 26.3 27.8
LJL/HL 45.811.5 47.0±1.8 48.811.3 48.811.3 49.610.5 52.512.2 54.1 58.6 5
LJW/HL 20.510.7 20.411.5 21.111.3 22.012.6 22.713.0 23.012.6 - 29.1 >
UJL/HL 36.711.7 37311.3 37.211.1 38.0tl.7 40.010.8 41.611.5 41.3 41.0 m
Ο
PPL/HL 26.911.2 27.611.0 27.811.0 27.911.0 28.610.5 27.210.7 28.1 29.1 τι
r
>
<
9
S
>
Si
Table 14. Angular and qualitative measurements, and counts of Haplochromis pyrrhopteryx spec. nov.
range means and standard

deviations
Dorsal Head Inclination 19°-30° 24.4°±6.3°

Premaxillary Pedicel Incl. 28°-40° 33.3°±5.4°
Snout Acuteness 64°-78° 70.9°±4.1°
Gape Inclination 32°-48° 39.0°±4.9°
Dorsal Head Profile (curvature) +

Premaxilla Pedicel Prominence 0(+)/+
Lower Jaw: Anterior Extension +/++
Lateral Snout Outline 0/+
Mental Prominence +
Lip Thickening 0(+)
Premaxilla Beaked 0
Maxillary Posterior Extension -/0
Lateral Line Scales 31 (f= 1); 32 (f= 12); 33 (f= 12); 34 (f= 6); 35 (f= 2)
Lateral Line - Dorsal Fin (Sc.rows) 5 (f= 2); 6 (f= 13); 7 (f= 13); 8 (f= 8)
Pectoral - Pelvic Fin (Sc.rows) 5 (f= 2); 6 (f= 7); 7 (f= 18); 8 (f= 4); 9 (f= 1)
Cheek (Vertical Sc.rows) 3 (f= 2); 4 (f= 14); 5 (f= 15)
Dorsal Fin (Spines/Rays) XIV 9 (f= 1); XV 9 (f= 5); XV 10 (f= 7); XVI 8 (f= 1);
XVI 9 (f= 14); XVI10 (f= 2); XVII 9 (f=l)
Anal Fin (Spines/Rays) III 8 (f=3); III 9 (f=22); III 10(f=l)
m e n s , 90 a n d 120 m m S L , r e m a i n s o f Rastrineobola argêntea w e r e f o u n d . I n b o t h speci-

m e n s the l e n g t h o f the p r e y w a s c. 50 m m . R e m a i n s of a s h r i m p s w e r e encountered
i n o n e s p e c i m e n , insect remains w e r e f o u n d i n t w o specimens a n d the intestines of 5
s p e c i m e n s w e r e empty.
— B r e e d i n g a n d g r o w t h . Specimens of H. pyrrhopteryx m a t u r e at a p p r o x i m a t e l y
135 m m S L . F e w a d u l t specimens h a v e been caught. T h e largest k n o w n s p e c i m e n is a
s e x u a l l y quiescent female o f 199 m m s t a n d a r d length. I n a l l females the r i g h t o v a r y
is larger t h a n the left. N o r i p e females w e r e caught, therefore the size of r i p e eggs i s
u n k n o w n . I n the m o u t h of one spent female, R M N H 30038, S L 178.3 m m , one e g g of
3x2.5 m m w a s f o u n d . H o w e v e r , i t is possible that this egg entered the m o u t h d u r i n g
capture o f the s p e c i m e n i n the trawl-net.
R e s e m b l i n g s p e c i e s . — T h e c o m b i n a t i o n of strongly r e c u r v e d u n i c u s p i d teeth a n d
the c o l o r a t i o n of H. pyrrhopteryx i m m e d i a t e l y sets the species apart f r o m a l l b u t one
of t h e L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s , v i z . H. dichrourus. H o w e v e r , the c o l -
o r a t i o n a l t h o u g h s i m i l a r i n s o m e respects, is d i s t i n c t e n o u g h t o separate t h e t w o
species at once. T h e m a i n c o l o u r s o f female H. dichrourus are b l u e - b l a c k a n d d a r k
r e d , a n d of males greenish-black a n d d a r k r e d . T h e i n t e n s i t y of the d o r s a l f i n c o l -
o r a t i o n is v a r i a b l e , b u t i t is u s u a l l y less b r i g h t r e d than the c a u d a l a n d anal fins. T h e
c a u d a l f i n i n f e m a l e s i s r e d o n l y o n i t s v e n t r a l half. I n H. pyrrhopteryx t h e m a i n
c o l o u r s f o r b o t h sexes a r e b l a c k a n d o r a n g e . A l l m e d i a n f i n s are l i g h t o r b r i g h t
o r a n g e e v e n i n j u v e n i l e s . P r e s e r v e d specimens of the t w o species can also b e separat-

e d b y their c o l o r a t i o n : the black c o l o u r of the v e n t r a l parts of the h e a d a n d b o d y o f
H. pyrrhopteryx are r e t a i n e d i n t h e p r e s e r v e d c o n d i t i o n , w h e r e a s p r e s e r v e d s p e c i -
m e n s o f H. dichrourus loose part of the d a r k v e n t r a l colour, the cheek a n d especially
the l o w e r j a w b e c o m i n g l i g h t e r , l e a v i n g t h e d a r k l a c h r y m a l s t r i p e a n d t h e b l a c k
i n t e r m a n d i b u l a r a r e a a n d b r a n c h o s t e g a l m e m b r a n e s t r o n g l y c o n t r a s t i n g . I n H.
pyrrhopteryx the cheeks a n d l o w e r jaws r e m a i n b l a c k a n d the l a c h r y m a l stripe is o n l y
f a i n t l y v i s i b l e . A n u m b e r o f m o r p h o m e t r i c features m a y b e u s e d to separate t h e
species. S m a l l differences are f o u n d i n the ranges of the ratio's for B o d y D e p t h / H e a d
L e n g t h , P r e o r b i t a l D e p t h / H e a d L e n g t h a n d Interorbital W i d t h / H e a d L e n g t h , w h i l e
the ranges of the ratio's o f S n o u t l e n g t h / H e a d L e n g t h , S n o u t W i d t h / H e a d L e n g t h ,
Cheek D e p t h / H e a d L e n g t h and L o w e r Jaw L e n g t h / H e a d L e n g t h s h o w almost n o
overlap.
W i t h H. bareli, H. pyrrhopteryx shares the s t r o n g l y c u r v e d teeth a n d the d a r k v e n -
tral b o d y c o l o r a t i o n of males. H o w e v e r , there are differences i n general b o d y shape
a n d i n m a n y m o r p h o m e t r i c measurements.
Haplochromis chrysogynaion spec. nov.

(figs 79-82,84-87,112, tables 15,16)
I n t r o d u c t i o n . — W h e n , i n the course of o u r investigations i n the s o u t h e r n p a r t of

L a k e V i c t o r i a w e c a u g h t g o l d c o l o u r e d s p e c i m e n s there w e r e t w o p o s s i b i l i t i e s f o r
t h e i r t a x o n o m i e status; either t h e y r e p r e s e n t e d c o l o u r p o l y m o r p h s o f a n e x i s t i n g
species (e.g. as d e s c r i b e d for Cichlasoma citrinellum i n N i c a r a g u a n L a k e s , see B a r l o w ,
1983; M c K a y e , 1980) o r they represented a n e w g o l d e n c i c h l i d species (a g o l d e n c i c h -
l i d , Pseudotropheus barlowi f r o m L a k e M a l a w i w a s d e s c r i b e d b y M c K a y & Staufer,
1986 a n d R i b b i n k et al, 1983). Because of their d i s t i n c t i v e tooth shape; v i z . s t r o n g l y
r e c u r v e d u n i c u s p i d s , t h e three specimens (all female) c l e a r l y b e l o n g e d to the
"dichrourus" g r o u p . C o m p a r i s o n o f the m o r p h o m e t r i c d a t a of the s p e c i m e n s w i t h
those o f s i m i l a r s i z e d specimens of the species o f the dichrourus g r o u p r e v e a l e d that
they d i f f e r f r o m H. pyrrhopteryx i n B D / S L , C h D / H L , P P L / H L a n d f r o m H. bareli i n
B D / S L , P F L / S L , I O W / H L , S n L / H L , P P L / H L ratio's. T h e m o r p h o m e t r i c measure-
ments o f the g o l d e n females fit i n the ranges of the M w a n z a G u l f p o p u l a t i o n of H.
dichrourus. H o w e v e r , closer e x a m i n a t i o n of the habitus o f the t w o g r o u p s revealed
differences i n b o d y a n d h e a d shape w h i c h are d i f f i c u l t to quantify. C o m p a r i s o n of
s o m e skeletal elements o f the largest g o l d e n female (91.3 m m ) w i t h those of a s i m i -
l a r l y s i z e d (93.1 m m ) s p e c i m e n of H. dichrourus f r o m the M w a n z a G u l f r e v e a l e d dif-
ferences i n t o o t h s h a p e a n d n u m b e r o f teeth (figs. 80, 83). Therefore I b e l i e v e t h e
g o l d e n f e m a l e s are representatives o f a d i f f e r e n t species. A l t h o u g h n o m a l e c o n -
s p e c i f i c s o f t h e f e m a l e s are k n o w n , i t seems a p p r o p r i a t e to d e s c r i b e h e r e a n e w
species b a s e d o n the females only.
Material.— Holotype; Ç, 89.0 mm, R M N H 30118, Mikassa Bay, Maisome Island, Tanzania, Lake
Victoria, 25.vi.1985, HEST. Paratypes: 1 9, 75.3 mm, R M N H 30119, Mikassa Bay, Maisome Island,
Tanzania, Lake Victoria, 25.vi.1985, HEST; 1 9, 91.3 mm, R M N H 30120, Mwanza Gulf, Tanzania, Lake
Victoria, 8.XÜ.1977, HEST.
E t y m o l o g y . — T h e n a m e c h r y s o g y n a i o n is c o m p o s e d of the G r e e k w o r d s f o r
g o l d e n ; " Χ ρ υ σ o " a n d " χ υ ν χ ι ο ν " ; the d i m i n u t i v e for female, a n d refers to the g o l d e n
l i v e c o l o r a t i o n of the females of this species.
D i a g n o s i s . — T h e f e m a l e s o f t h i s s p e c i e s are s e p a r a b l e f r o m a l l o t h e r h a p
l o c h r o m i n e c i c h l i d species b y their l i v e c o l o r a t i o n . P r e s e r v e d f e m a l e s d i f f e r f r o m
m o s t other p i s c i v o r o u s species i n h a v i n g s t r o n g l y r e c u r v e d u n i c u s p i d teeth. F r o m
other p i s c i v o r o u s species w h i c h h a v e s i m i l a r teeth they can be d i s t i n g u i s h e d b y their
u n i f o r m cream c o l o r a t i o n a n d lack of a l l m a r k i n g s .
D e s c r i p t i o n . — Based o n the h o l o t y p e a n d t w o paratypes 75.3-91.3 m m S L .
H a b i t u s . — A m o d e r a t e l y deep b o d i e d , r e l a t i v e l y s m a l l s i z e d species. D o r s a l h e a d
p r o f i l e n e a r l y straight, d i v i d e d i n t o t w o s l i g h t l y c o n v e x parts b y a slight i n c u r v a t i o n
a b o v e the eye . P r e m a x i l l a r y p e d i c e l v i s i b l e b u t s m o o t h l y m e r g i n g i n t o the h e a d p r o
file. C e p h a l i c lateral l i n e o p e n i n g s not e n l a r g e d , the canals o n the l a c h r y m a l barely
v i s i b l e . E y e circular, p u p i l s u b c i r c u l a r w i t h a s m a l l a p h a k i c aperture r o s t r o - v e n t r a l l y
to the lens. M o u t h m o d e r a t e l y o b l i q u e . P r e m a x i l l a n o t e x p a n d e d or b e a k e d . T h e
e x p o s e d part of the m a x i l l a s m a l l . Posterior tip of m a x i l l a r e a c h i n g a v e r t i c a l posteri
o r to the eye m a r g i n . L i p s n o r m a l . Jaws e q u a l anteriorly, l o w e r j a w tip s l i g h t l y p r o
t r u d i n g . R o s t r a l o u t l i n e of l o w e r j a w a n d m e n t a l area r o u n d e d . L o w e r j a w sides
s l i g h t l y to m o d e r a t e l y o b l i q u e .
S c a l e s . — C h e e k , g i l l - c o v e r a n d d o r s a l h e a d surface c o v e r e d w i t h c y c l o i d scales.
R o s t r a l part of d o r s u m w i t h a m i x t u r e of c y c l o i d a n d w e a k l y c t e n o i d scales, those o n
the c a u d a l p a r t c t e n o i d . C h e s t laterally w i t h c t e n o i d , v e n t r a l l y w i t h c y c l o i d scales. A
g r a d u a l size change b e t w e e n chest scales a n d scales of s u r r o u n d i n g areas. Scales o n
r e m a i n i n g p a r t of b o d y c t e n o i d . C a u d a l f i n s c a l e d w i t h elongate, w e a k l y c t e n o i d
scales o n its p r o x i m a l half to t w o thirds. N o basal scale sheath o n d o r s a l a n d a n a l f i n .
F i n s . — Pectorals r e a c h i n g a n a l i n one s p e c i m e n , p e l v i c s not r e a c h i n g a n a l . D o r s a l
a n d a n a l fins not r e a c h i n g base of c a u d a l f i n . First p e l v i c r a y s l i g h t l y p r o d u c e d . O u t
l i n e of c a u d a l f i n truncate or subtruncate a n d s l i g h t l y emarginate.
G i l l a p p a r a t u s . — E i g h t or n i n e g i l l rakers o n l o w e r part of first g i l l arch. L o w e r
m o s t t w o r e d u c e d , c o n i c a l , t h i r d a n d f o u r t h short a n d c o n i c a l or f o u r t h flattened
t a p e r i n g , f i f t h to seventh flattened w i t h b r o a d e n e d s i m p l e or b i f i d heads, last ones
less flattened, t a p e r i n g . O n the lateral h e m i b r a n c h of the first g i l l arch the n u m b e r of
g i l l f i l a m e n t s is about 110.
V i s c e r a . — Intestine length f r o m 1.0-1.3 times s t a n d a r d length. F o l l o w i n g the d e f i
n i t i o n s of Z i h l e r (1982) the c o n f i g u r a t i o n of the intestines is of type D ( w i t h backflap).
O r a l t e e t h . — S h a p e . In the o u t e r r o w of b o t h j a w s u n i c u s p i d teeth d o m i n a t e
w e a k l y b i c u s p i d teeth. T h e u n i c u s p i d s are s l e n d e r , r o u n d a n d a c u t e l y p o i n t e d .
Laterally, s o m e u n i c u s p i d s are s l i g h t l y c o m p r e s s e d d i s t a l l y a n d h a v e a s h o u l d e r o n
one side. A t the base of this s h o u l d e r a v e r y s m a l l cusp m a y be present. T h e i n n e r
r o w teeth i n the u p p e r j a w are a l l w e a k l y t r i c u s p i d s , i n the l o w e r j a w s h o u l d e r e d
u n i c u s p i d s d o m i n a t e w e a k l y b i c u s p i d s a n d t r i c u s p i d s . A l l t e e t h are s t r o n g l y
recurved.
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r r o w covers near
l y l e n g t h of dentigerous a r m of p r e m a x i l l a , i n n e r r o w s about 3/5 part. T w o or three
i n n e r r o w s of teeth i n rostral part of p r e m a x i l l a , decreasing to one at 1/2 i n n e r r o w
l e n g t h . T h e gap b e t w e e n the outer r o w a n d the first inner r o w rostrally is r e l a t i v e l y
s m a l l . O u t e r r o w of m a n d i b l e r e a c h i n g rostral part of c o r o n o i d w i n g . T h e longer of
Fig. 79. Haplochromis chrysogynaion spec. nov. Paratype R M N H 30120, SL 91.3 mm, lateral view. Scale
equals 10 mm.
Figs 80-82,84-87. Haplochromis chrysogynaion spec. nov. Fig. 83. Haplochromis dichrourus "Mwanza". Fig.
80, 83. Premaxilla, lateral view. Fig. 81. Premaxilla, perpendicular view of dentigerous area. Fig. 82.
Premaxilla ascending arm, medial view. Fig. 84. Lower jaw, lateral view. Fig. 85-87. Premaxillary teeth,
labial and lateral view. Fig. 85. Second tooth of outer row. Fig. 86. Thirteenth tooth of outer row. Fig.
87. Seventh tooth of 1st inner row. Scale of bony elements = 5 mm, scale of teeth = 1mm.
the t w o i n n e r r o w s occupies 3/4 of outer r o w l e n g t h a n d does not reach the c o r o n o i d

wing.
— C o u n t s a n d setting. 52-57 teeth i n outer r o w of u p p e r jaw, 38-46 i n outer r o w
of l o w e r jaw. Teeth r e g u l a r l y set i n b o t h j a w s , distance b e t w e e n t o o t h bases about
e q u a l to d i a m e t e r of toothbase. A s tooth size decreases c a u d a d the absolute distance
b e t w e e n the teeth also decreases. N o m e d i a l g a p i n either jaw.
— I m p l a n t a t i o n . U p p e r jaw, outer r o w teeth r o s t r a l l y erect to s l i g h t l y p r o c u m -
bent, rostro-laterally s l i g h t l y recumbent, c a u d a l l y m o d e r a t e l y recumbent. U p p e r j a w
i n n e r r o w teeth s t r o n g l y recumbent. L o w e r j a w outer r o w teeth erect rostrally, slight-
l y r e c u m b e n t caudally. L o w e r j a w inner r o w teeth s l i g h t l y to m o d e r a t e l y recumbent.
P h a r y n g e a l t e e t h . — C o u n t s : 24 teeth i n the c a u d a l transverse r o w a n d 10-11 i n
the t w o m e d i a n r o w s .
— Shape. Except for the teeth i n the c a d a l m o s t r o w , w h i c h are h o o k e d , a l l teeth
are of the b e v e l l e d type. Tooth size increases f r o m rostral to c a u d a l a n d f r o m lateral
to m e d i a l . T h e b e v e l l e d teeth are s m a l l , s l e n d e r a n d a c u t e l y p o i n t e d , the h o o k e d
teeth are s l i g h t l y larger a n d b l u n t . In one s p e c i m e n the tips of the h o o k e d teeth are
blackish.
O s t e o l o g y . — T h e l o w e r p h a r y n g e a l element w a s dissected i n three, the r i g h t pre-
m a x i l l a i n t w o s p e c i m e n s , the r i g h t l o w e r j a w w a s dissected i n one p a r a t y p e . Be-
cause of the f e w specimens available n o s p e c i m e n w a s dissected for o b t a i n i n g a n e u -
rocranium.
— O r a l j a w s . P r e m a x i l l a d e n t i g e r o u s a r m l o n g e r (1.1 x) t h a n a s c e n d i n g a r m ,
angle b e t w e e n the t w o arms 78°-80°. T r a n s i t i o n of dentigerous a n d a s c e n d i n g a r m
r e l a t i v e l y b r o a d . V e n t r a l o u t l i n e of d e n t i g e r o u s a r m s l i g h t l y c o n v e x . L o w e r j a w :
m o d e r a t e l y slender, l e n g t h / d e p t h ratio 2.3. L e n g t h of toothbearing part of the d e n -
tary 0.5 times l o w e r j a w l e n g t h .
— L o w e r p h a r y n g e a l element: a relatively s h a l l o w element, s l i g h t l y longer t h a n
b r o a d ( l e n g t h / w i d t h = 1.1). D e n t i g e r o u s area s l i g h t l y b r o a d e r t h a n l o n g ( l e n g t h /
w i d t h = 0.87-0.9).
— Vertebrae. There are 29 o r 30 vertebrae c o m p r i s i n g 13 a b d o m i n a l a n d 16 (f = 1)
o r 17 (f = 2) c a u d a l elements.
C o l o r a t i o n . — L i v e colours of m a t u r i n g a n d j u v e n i l e females. T i p of u p p e r l i p a n d
d o r s a l p a r t s of s n o u t , g i l l - c o v e r , h e a d , b o d y a n d c a u d a l p e d u n c l e d e e p y e l l o w -
o r a n g e (golden). M i d d l e part of f l a n k a n d c a u d a l p e d u n c l e y e l l o w i s h w i t h a n orange
f l u s h . V e n t r a l half of h e a d , branchiostegal m e m b r a n e , chest, b e l l y a n d v e n t r a l side
b r i g h t w h i t e , w i t h a s i l v e r sheen o n the cheek, v e n t r a l part of g i l l - c o v e r a n d v e n t r a l
side. (The s p e c i m e n f r o m the M w a n z a G u l f h a d a green a n d y e l l o w f l u s h o n the d o r -
sal p a r t of the g i l l cover a n d a green t i p o n the u p p e r l i p ) . Inner r i n g of eye g o l d e n .
F i n s : D o r s a l y e l l o w i s h p r o x i m a l l y , h y a l i n e distally, w h i t e streaks b e t w e e n the rays.
Pectorals y e l l o w i s h h y a l i n e , p e l v i c s h y a l i n e w i t h w h i t e rays. A n a l w h i t e b e t w e e n the
spines, h y a l i n e w i t h a y e l l o w i s h f l u s h b e t w e e n the rays, p r o x i m a l part of the rays
w h i t e . D o r s o - c a u d a l l y o n the a n a l f i n one or t w o s m a l l r o u n d , orange spots w i t h a
t h i n w h i t e r i m . C a u d a l f i n d a r k orange p r o x i m a l l y , h y a l i n e distally.
— P r e s e r v e d c o l o r a t i o n of m a t u r i n g a n d j u v e n i l e females. H e a d a n d b o d y almost
e n t i r e l y u n i f o r m l y cream c o l o u r e d . S n o u t a n d d o r s a l parts of h e a d a n d d o r s u m d a r k -
er g r e y i s h . A s m a l l faint orange area o n the d o r s a l part of the l a c h r y m a l is present i n
the M i k a s s a B a y specimens. A l l specimens h a v e a faint o p e r c u l a r b l o t c h . A l l fins h y a -
l i n e , s m a l l grey streaks b e t w e e n the rays of d o r s a l a n d c a u d a l f i n .
Table 15. Linear and angular measurements of Haplochromis chrysogynaion spec. nov.
Holotype
R M N H reg.no. 30119 30118 30120
Standard Length 75.3 89.0 91.3

Body Depth %SL 32.2 33.7 33.0
Pectoral Fin Length %SL 25.3 26.4 30.1
Caudal Peduncle Length %SL 16.6 16.7 16.5
Caudal Peduncle Depth %SL 11.8 11.5 11.5
Caudal Fin Length %SL 23.9 23.9 22.5
Head Length %SL 36.5 35.0 35.9
Snout Length %HL 32.2 31.7 30.7
Snout Width %HL 29.8 32.0 32.6
Head Width %HL 43.2 42.9 44.8
Interorbital Width %HL 21.0 19.8 21.3
Preorbital Width %HL 25.4 26.2 25.9
Lachrymal Width %HL 25.4 25.6 25.9
Preorbital Depth %HL 17.4 16.6 16.4
Eye Length %HL 26.1 25.6 25.3
Cheek Depth %HL 22.1 23.0 23.4
Lower Jaw Length %HL 44.0 46.4 46.4
Lower Jaw Width %HL 21.0 20.1 21.3
Upper Jaw Length %HL 36.3 36.8 36.5
Premax. Pedicel Length %HL 28.3 28.5 29.8
Dorsal Head Profile Inclination 30° 32° 32°

Premaxillary Pedicel Inclination 36° 38° 40°
Snout Acuteness 67° 69° 68°
Gape Inclination 35° 32° 32°
Table 16. Qualitative measurements and counts of Haplochromischrysogynaion spec.nov.
Dorsal Head Profile (curvature) 0

Premaxillary Pedicel Prominence 0(+)
Lower Jaw: anterior Extension +
Lateral Snout Outline 0
Mental Prominence 0
Lip Thickening 0
Premaxilla Beaked 0
Maxillary Posterior Extension +
Cephalic Lateral Lines Pores: width 0
Lateral Line Scales 31 (1); 32(1); 33(1);

Lateral Line - Dorsal Fin (Sc. rows) 5(1); 6(2)
Pectoral - Pelvic Fin Bases (Sc. rows) 5(1); 6(2)
Cheek (Vertical Sc. rows) 4(2); 5(1)
Dorsal Fin (Spines/Rays) XV 9(1); XVI 8 (1); XVI 9(1)
Anal Fin (Spines/Rays) III 8 (1); 1119(1); III 10(1)
D i s t r i b u t i o n . — Haplochromis chrysogynaion is o n l y k n o w n f r o m L a k e V i c t o r i a .
S p e c i m e n s w e r e collected at t w o localities o n l y ; i n the M i k a s s a B a y of M a i s o m e Is-
l a n d a n d i n the n o r t h e r n p a r t of the M w a n z a G u l f .
E c o l o g y . — H a b i t a t . T h e t w o catch localities o f the s p e c i m e n s of Haplochromis
chrysogynaion w e r e b o t h r e l a t i v e l y sheltered areas b o r d e r i n g o n o p e n water. T h e bot-
t o m at b o t h localities w a s m u d a n d the d e p t h b e t w e e n 12-15 m .
— F o o d . S t o m a c h a n d intestines of t w o specimens w e r e e x a m i n e d . I n the intes-
tines o f the smallest s p e c i m e n (75.3 m m ) r e m a i n s of a s h r i m p (Caridina nilotica) a n d
several Chaoborus larvae w e r e f o u n d . S t o m a c h a n d intestines o f the largest s p e c i m e n
(93.1 m m ) c o n t a i n e d r e m a i n s o f a s m a l l h a p l o c h r o m i n e c i c h l i d , s t a n d a r d l e n g t h
a b o u t 27 m m .
— B r e e d i n g & g r o w t h . T h e female of 75.4 m m S L is a j u v e n i l e , the female of 89.0
m m S L is r i p e n i n g ( p r o b a b l y m a t u r i n g ) , a n d the 91.3 m m S L female seems i m m a -
ture.
R e d e s c r i p t i o n of Haplochromis pellegrini R e g a n , 1922

G r e e n w o o d (1962) i n h i s r e d e s c r i p t i o n of Haplochromis pellegrini states that the
d e s c r i p t i o n w a s b a s e d o n t w e n t y - f i v e specimens (71-104 m m S L ) i n c l u d i n g the lecto-
a n d p a r a t y p e . H o w e v e r , i n the p a r a g r a p h o n f o o d he m e n t i o n s t h i r t y - f i v e e x a m i n e d
s p e c i m e n s , a n d i n the tabel ' s t u d y m a t e r i a l a n d d i s t r i b u t i o n records' he lists 38 speci-
m e n s . I n the B M ( N H ) jar w i t h the l i s t e d registration n u m b e r s I f o u n d n o labels i n d i -
c a t i n g w h i c h specimens w e r e u s e d f o r the r e d e s c r i p t i o n of m o r p h o m e t r i c a n d m e r i s -
tic characters. E x a m i n a t i o n o f t h e 36 E . A . F . R . O . s p e c i m e n s o f H. pellegrini i n t h e
B M ( N H ) c o l l e c t i o n r e v e a l e d that amongst these specimens w a s o n e s p e c i m e n of H.
dichrourus ( B M ( N H ) 1962.3.2.510) S L 91.1 m m (see page 31). Three m o r e specimens
d i v e r g e d f r o m the majority. O n e of these specimens, a female of 74.3 m m S L (from l o t
B M ( N H ) 1962.3.2.516-544), is a s p e c i m e n of H. perrieri ( P e l l e g r i n , 1904). T h e largest
s p e c i m e n of l o t B M ( N H ) 1962.3.2.516-544, a n i m m a t u r e female of 85 m m S L , also dif-
fers f r o m the m a j o r i t y o f specimens i n certain m o r p h o m e t r i c a n d s e m i q u a n t i t a t i v e
m e a s u r e m e n t s ; at present I a m n o t sure w h e r e to place this s p e c i m e n . A t h i r d speci-
m e n ( B M ( N H ) 1982.3.2.509) a s e x u a l l y active m a l e o f 85.7 m m S L , differs i n its d e n t i -
t i o n a n d p r e s e r v e d c o l o r a t i o n f r o m a l l other specimens G r e e n w o o d i d e n t i f i e d as H.
pellegrini, as w e l l as f r o m a l l other described species. T h e d e s c r i p t i o n of a n e w species
o n the basis o f this s i n g l e s p e c i m e n w o u l d be j u s t i f i e d , b u t I prefer to w a i t u n t i l other
m a t e r i a l has been e x a m i n e d . A f t e r G r e e n w o o d ' s (1962) r e d e s c r i p t i o n w a s m a d e , o n e
s p e c i m e n o f B M ( N H ) 1962.3.2.510-12 w a s dissected to o b t a i n i n f o r m a t i o n o n the n e u -
r o c r a n i a l shape . U n f o r t u n a t e l y n o p h o t o g r a p h s w e r e t a k e n p r i o r to the d i s s e c t i o n
a n d m o r p h o m e t r i c d a t a w e r e n o t p r e s e r v e d , so this s p e c i m e n c o u l d a l s o n o t b e
i n c l u d e d i n the f o l l o w i n g d e s c r i p t i o n .
T h e 31 r e m a i n i n g E . A . F . R . O . s p e c i m e n s i d e n t i f i e d b y G r e e n w o o d (1962) as H.
pellegrini differ f r o m the t y p e series of this species i n h e a d shape, m o r p h o m e t r i c a n d
m e r i s t i c characters, a n d d e n t i t i o n . I n m y o p i n i o n the types of H. pellegrini, w h i c h
h a v e a s t a n d a r d l e n g t h o f 103.8 a n d 100.0 m m , are b o t h i m m a t u r e , w h e r e a s a l l
E . A . F . R . O . s p e c i m e n s (68 to 84 m m S L ) are m a t u r e o r n e a r l y so. T h i s o b s e r v a t i o n
strengthens m y v i e w that the E . A . F . R . O . specimens b e l o n g to a separate t a x o n . F o r
this r e a s o n H. pellegrini is r e d e s c r i b e d (on the basis of its t y p e m a t e r i a l o n l y ) a n d the
r e m a i n i n g s p e c i m e n s are d e s c r i b e d as a n e w species. U n f o r t u n a t e l y , eight years of
c o l l e c t i n g b y H E S T i n the M w a n z a G u l f area d i d n o t y i e l d a n y a d d i t i o n a l specimens

of either species, so the descriptions are based o n the B M ( N H ) m a t e r i a l only.
Haplochromis pellegrini R e g a n , 1922

(figs 7 8 , 8 8 , 1 1 3 , tables 17,18)
Paratilapia prognatha Pellegrin (part); Boulenger, 1915:333.

Haplochromis pellegrini Regan, 1922:185, fig. 11.
Haplochromis pellegrini (part; types only); Greenwood, 1962:186-189, fig. 16 & 1974:, 89,91, fig. 58.
Prognathochromis (P.) pellegrini (part; types only); Greenwood, 1980: 20.
Material.— Lectotype, $, 103.8 mm, B M N H 1906.5.30.253, Lake Victoria, near Entebbe, Uganda, E.
Degen; Paralectotype; cf, 100.0 mm, B M N H 1906.5.30.254, Lake Victoria, near Entebbe, E. Degen.
D i a g n o s i s . — P r o b a b l y a m e d i u m s i z e d (both t y p e s p e c i m e n s are i m m a t u r e ) ,
m o d e r a t e l y slender, m o d e r a t e l y laterally c o m p r e s s e d p i s c i v o r o u s species w i t h slight-
l y c u r v e d u n i c u s p i d teeth i n the outer r o w of the o r a l jaws.
D e s c r i p t i o n . — B a s e d o n the lectotype, a s u b a d u l t female, a n d the paralectotype,
a n i m m a t u r e male.
H a b i t u s . — P r o b a b l y (both specimens b e i n g i m m a t u r e ) a m e d i u m s i z e d o r m o d -
erately large p i s c i v o r o u s species w i t h a rather slender a n d f a i r l y c o m p r e s s e d body.
Dorsal head profile slightly convex, premaxillary pedicel fairly prominent. Pre-
m a x i l l a s l i g h t l y e x p a n d e d m e d i a l l y i n b o t h s p e c i m e n s , s l i g h t l y b e a k e d i n the par-
alectotype. L i p s rather t h i n . A s m a l l part of the m a x i l l a v i s i b l e c a u d a l l y a n d d o r s o -
c a u d a l l y to the p r e m a x i l l a . T h e v e r t i c a l t h r o u g h the c a u d a l t i p o f the m a x i l l a n o t
r e a c h i n g the r o s t r a l e y e m a r g i n . M o u t h m o d e r a t e l y o b l i q u e . L a t e r a l s n o u t o u t l i n e
i s o g n a t h o u s i n the p a r a t y p e , s l i g h t l y p r o g n a t h o u s i n the lectotype, l o w e r j a w p r o -
t r u d i n g i n b o t h s p e c i m e n s . M e n t a l p r o m i n e n c e s m a l l . L a t e r a l sides o f l o w e r j a w s
m o d e r a t e l y o b l i q u e . H o r i z o n t a l a r m of p r e o p e r c u l u m s l i g h t l y d e c l i n i n g v e n t r a d , the
v e r t i c a l a r m s l i g h t l y r e c l i n i n g c a u d a d . L a t e r a l l i n e canals a n d o p e n i n g s o n l a c h r y m a l
n o r m a l . E y e subcircular, the p u p i l s l i g h t l y e l l i p s o i d o n a h o r i z o n t a l l e n g t h axis.
S c a l e s . — Scales of h e a d , rostral part o f d o r s u m a n d b e l l y c y c l o i d , chest w i t h a
m i x t u r e o f c y c l o i d a n d c t e n o i d scales. Scales o n r e m a i n d e r o f b o d y c t e n o i d . S m a l l
e l o n g a t e scales o n p r o x i m a l p a r t o f t h e c a u d a l f i n , those n e a r t h e m a r g i n b a r e l y
r e a c h i n g the second half of the f i n .
F i n s . — Pectoral a n d p e l v i c s just r e a c h i n g a n a l . D o r s a l a n d a n a l n o t r e a c h i n g base
of c a u d a l f i n . I n the lectotype the d o r s a l reaches further c a u d a d t h a n the a n a l , i n the
p a r a l e c t o t y p e the a n a l f i n extends s l i g h t l y further c a u d a d . C a u d a l f i n o u t l i n e s u b -
truncate, s l i g h t l y emarginate a n d s l i g h t l y o b l i q u e .
G i l l s . — Ten (lectotype) o r n i n e gill-rakers o n the l o w e r part of the first g i l l - a r c h of
the r i g h t side. T h e l o w e r three s m a l l a n d c o n i c a l , n u m b e r s f o u r a n d f i v e elongate,
a n d the u p p e r m o s t three o r f o u r elongate a n d b r o a d e n e d , the h e a d c o n i c a l o r b i f i d .
V i s c e r a . — T h e intestines a n d stomachs of b o t h specimens w e r e n o t r e m o v e d , s o
n o i n f o r m a t i o n is available about their l e n g t h a n d c o n f i g u r a t i o n .
O r a l t e e t h . — S h a p e . A l l outer r o w teeth i n b o t h j a w s r e l a t i v e l y s m a l l , s l i g h t l y
c u r v e d , acutely p o i n t e d u n i c u s p i d s . I n u p p e r j a w a n d r o s t r a l l y i n l o w e r j a w the teeth
are slender a n d r o u n d e d , laterally a n d posterio-laterally i n l o w e r j a w t h e teeth are
s o m e w h a t c o m p r e s s e d a n d s h o u l d e r e d . Inner r o w s consist of s l i g h t l y c u r v e d , rela-

t i v e l y l o n g , r o u n d e d u n i c u s p i d s rostrally, m i x e d w i t h s o m e w h a t c o m p r e s s e d , s h o u l -
d e r e d u n i c u s p i d s laterally. The size of the outer r o w teeth d i m i n i s h e s g r a d u a l l y , f r o m
rostral to c a u d a l .
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r r o w covers w h o l e
l e n g t h of p r e m a x i l l a r y d e n t i g e r o u s a r m , i n n e r r o w s a b o u t t w o t h i r d s . F o u r i n n e r
r o w s o n rostral part of p r e m a x i l l a decreasing to zero o n c a u d a l part. G a p b e t w e e n
outer r o w a n d i n n e r r o w distinct. In l o w e r j a w the outer r o w occupies rostral part of
c o r o n o i d w i n g . T w o to three i n n e r r o w s o n rostral part of l o w e r jaw. G a p b e t w e e n
outer r o w a n d first i n n e r r o w distinct.
— C o u n t s a n d setting. There are 65 a n d 62 teeth i n outer r o w of u p p e r jaw, a n d
40 a n d 38 i n outer r o w of l o w e r j a w for the lectotype a n d paralectotype, respectively.
T h e teeth are r e g u l a r l y set, neck distance i n u p p e r j a w a n d r o s t r a l l y i n l o w e r j a w
e q u a l to o r s l i g h t l y m o r e than neck w i d t h , decreasing to half neck w i d t h c a u d a l l y i n
the l o w e r jaw.
— I m p l a n t a t i o n . O u t e r r o w teeth recumbent i n u p p e r a n d l o w e r jaw. Inner r o w s
s l i g h t l y r e c u m b e n t to recumbent i n l o w e r jaw, recumbent to s t r o n g l y recumbent i n
u p p e r jaw.
P h a r y n g e a l t e e t h . — C o u n t s . T w e n t y teeth i n the c a u d a l m o s t transverse r o w a n d
9 or 10 i n the m e d i a n r o w s .
— Shape. In lateral v i e w , teeth of c a u d a l m o s t r o w e n l a r g e d , h o o k e d a n d rather
s h a r p , other teeth b e v e l l e d . Tooth size increases f r o m rostral to c a u d a l a n d f r o m lat-
eral to m e d i a l .
O s t e o l o g y . — A s o n l y the lectotype a n d paralectotype of this species are available
n o d e s c r i p t i o n can be g i v e n of skeletal elements. H o w e v e r , i n the lectotype the l o w e r
p h a r y n g e a l e l e m e n t h a d been disected p r e v i o u s l y , a n d is d e s c r i b e d b e l o w .
— L o w e r p h a r y n g e a l e l e m e n t : L o w e r p h a r y n g e a l e l e m e n t of the g e n e r a l i s e d
t y p e , s h a l l o w , s l i g h t l y longer than b r o a d ( l e n g t h / w i d t h = 1.2). D e n t i g e r o u s area as
l o n g as b r o a d .
— Vertebrae: B o t h type specimens have 29 vertebrae, c o m p r i s i n g 13 a b d o m i n a l
a n d 16 c a u d a l elements.
Fig. 88. Haplochomis pellegrini Regan, 1922. Holotype, BMNH.5.30.253, SL 103.8 mm. From Regan, 1922.
Table 17. Linear and angular measurements of Haplochromis pellegrini Regan, 1922 and H. harpakteridion
spec. nov.
Haplochromis pellegrini Haplochromis harpakteridion
Lectotype Paralecto- 30 specimens incl. Holotype

type range mean ± st.dev.
Standard Length 103.8 100.0 66.2-80.5 75.1+3.6

Body Depth %SL 30.0 31.0 29.5-34.3 31.5±1.1
Pectoral Fin Length %SL 23.3 20.6 9.7-23.7 21.7±1.0
Caudal Peduncle Length % SL 16.1 16.0 13.0-16.7 15.0±0.8
Caudal Peduncle Depth %SL 12.3 12.3 10.8-13.2 11.9±0.5
Caudal Fin Length %SL 24.2 25.1 21.5-24.5 22.9±0.7
Head Length %SL 34.8 36.3 35.4-38.6 36.8±0.6
Snout Length % HL 33.1 33.1 29.1-33.9 32.0±1.1
Snout Width % HL 30.1 30.1 27.9-33.4 29.8±0.5
Head Width % HL 37.2 36.2 37.2-42.9 40.4±1.2
Interorbital Width % HL 20.7 21.4 19.2-21.9 20.6±0.6
Preorbital Width % HL 26.5 25.3 23.4-27.3 25.2±0.8
Lachrymal Width % HL 24.5 25.3 24.2-28.8 27.0±1.2
Preorbital Depth % HL 19.8 20.6 15.7-20.0 17.0±3.0
Eye Length % HL 27.0 26.9 24.2-28.5 26.3±1.2
Check Depth % HL 22.6 23.6 20.0-23.8 21.6±1.0
Lower Jaw Length % HL 48.6 46.8 45.1-51.0 47.0±1.2
Lower Jaw Width % HL 16.5 15.4 16.8-23.5 19.3±1.9
Upper Jaw Length % HL 38.3 38.0 33.8-39.4 37.3±1.1
Premax. Pedicel Length % HL 25.4 25.6 27.9-32.0 29.511.1
Dorsal Head Profile Inclination 30° 30° 21°-30° 26.0° ± 3.0°

Premaxillary Pedicel Inclination 40° 35° 25° - 4 0 ° 33.0° ± 4.4°
Snout Acuteness 73° 62° 50° - 70° 65.7° ± 2.9°
Gape Inclination 40° 42° 31°-40° 36. 6° ± 3 . 1 °
C o l o r a t i o n . — C o l o r a t i o n o f p r e s e r v e d s p e c i m e n s . B o t h s p e c i m e n s are a l m o s t
c o m p l e t e l y b l e a c h e d b y their l o n g p r e s e r v a t i o n a n d n o differences c o u l d b e traced
b e t w e e n the m a l e a n d the female s p e c i m e n . H e a d , d o r s u m a n d d o r s a l p a r t o f c a u d a l
p e d u n c l e l i g h t b r o w n . G i l l - c o v e r , chest, belly, f l a n k a n d v e n t r a l p a r t of b o d y s i l v e r y -
w h i t e . P e l v i c , p e c t o r a l , d o r s a l a n d a n a l fins h y a l i n e , c a u d a l b r o w n i s h . Traces of d a r k
spots a n d stripes b e t w e e n the rays of d o r s a l a n d c a u d a l f i n . T h e three o c e l l i R e g a n
(1922) d e s c r i b e d o n the p o s t e r i o r part o f the a n a l f i n of one s p e c i m e n (paralectotype)
are n o w v e r y faint. A p a r t f r o m traces of a n o p e r c u l a r b l o t c h i n b o t h s p e c i m e n s there
are n o h e a d a n d b o d y m a r k i n g s .
D i s t r i b u t i o n . — H. pellegrini i s o n l y k n o w n f r o m L a k e V i c t o r i a . B o t h s p e c i m e n s
c o m e f r o m the same l o c a l i t y : Entebbe ( U g a n d a ) . N o details of capture are k n o w a n d
n o t h i n g is k n o w n about the habitat.
R e m a r k s . — A s n o a d u l t specimens are a v a i l a b l e a n d n o t h i n g is k n o w n about the
l i v e c o l o r a t i o n , s p e c i m e n s o f this species s h o u l d b e c o m p a r e d w i t h p r e s e r v e d j u v e -
niles o f other species. H e a d shape a n d d e n t i t i o n are rather s i m i l a r to that o f s o m e
s p e c i m e n s n o w i n c l u d e d i n H. prognathus b u t u n t i l this species is r e d e s c r i b e d n o t h i n g
Table 18. Qualitative measurements and counts of Haplochromis pellegrini Regan, 1922 and H. harpak-
teridion spec, no v.
Haplochromis pellegrini H. harpakteridion
Dorsal Head Profile (curvature) 0 <X+)

Premaxillary Pedicel Prominence + +/++
Lower Jaw: anterior Extension + +
Mental Prominence 0(+) 0
Lip Thickening 0 0
Premaxilla Beaked 0 0/+
Premaxilla Expanded 0 0/+
Maxillary Posterior Extension + +
Cephalic Lateral Line Pores: width 0 0
Lateral Line Scales 30 (f= 1); 31 (f= 7);

32 (f= 2); 33 (f= 2)
Lateral Line-Dorsal Fin (Sc. rows) 6 4 (f= 1); 5 (f= 13); 6 (f= 6)
Pectoral-Pelvic Fin (Sc. rows) 5 4 (f= 3); 5 (f= 10); 6 (f= 5)
Cheek (Vertical Sc. rows) 4; 5 3(f= l);4(f=17);5(f= 2)
Dorsal Fin (Spines/Rays) XIV 11 V 9 (f= 6); XV 10 (f= 6);
XVI 9 (f= 2); XVI10 (f= 2)
Anal Fin (Spines/Rays) III 9 III 8 (f= 5); III 9 (f= 9); III 10 (f=3)
precise can be s a i d about the r e l a t i o n s h i p o f these species. I n the absence o f skeletal

m a t e r i a l i t is d i f f i c u l t to establish the r e l a t i o n s h i p w i t h H. mento R e g a n , 1922, a n d H.
estor R e g a n , 1922, species w h i c h a c c o r d i n g to G r e e n w o o d (1962) h a v e a s i m i l a r b o d y
f o r m . T h e d e n t i t i o n i n H. pellegrini is s t r i k i n g l y d i f f e r e n t f r o m that i n these t w o
species. G r e e n w o o d ' s suggestion of a resemblance b e t w e e n H. pellegrini a n d H. per-
coides m u s t h a v e been b a s e d o n the E A F R O specimens d e s c r i b e d b e l o w .
Haplochromis harpakteridion spec. nov.

(figs 78,89-103,114, tables 17,18)
H. Pellegrini (part); Greenwood, 1962:186-189 & 1974: 90-91.

Prognathochromis (P.) pellegrini (part); Greenwood, 1980: 20.
Material.— Holotype, 9, 72.3 mm, B M N H 1962.3.2.516, Lake Victoria, near Karenia, near Jinja,
Uganda, E A F R O . Paratypes: 1 68.9 mm, B M N H 1962.3.2.512, Grant Bay, Lake Victoria, Uganda,
E A F R O ; 3 99, 74.0-80.5 mm, B M N H 1962.3.2.513-515, Beach in Buvuma Channal, Lake Victoria,
Uganda, E A F R O ; 9 <f<f, 72.9-80.0 mm + 17 99, 69.1-79.5 mm, B M N H 1962.3.2.517-542, Lake Victoria,
Karenia, near Jinja, Uganda, EAFRO.
E t y m o l o g y . T h e n a m e is d e r i v e d f r o m the d i m i n u t i v e o f the G r e e k απνακνπ (rob

ber) r e f e r r i n g to the p r e d a t o r y habits of this s m a l l species.
D i a g n o s i s . — A v e r y s m a l l , m o d e r a t e l y slender, p i s c i v o r o u s species w i t h m o d e r
ately c u r v e d teeth. L i v e females h a v e a u n i f o r m l y chocolate b r o w n c o l o u r o n h e a d
a n d b o d y . A l l fins are b l a c k i s h - b r o w n .
D e s c r i p t i o n . — B a s e d o n 31 specimens ( i n c l u d i n g the h o l o t y p e ) 66.2-80.5 m m S L .
Fig. 89. Haplochromis harpaktendion spec. nov. BM(NH) 1962.3.517, SL 78.6 mm, lateral view. Scale
equals 10 mm.
H a b i t u s . — A s m a l l , m o d e r a t e l y slender p i s c i v o r o u s species. D o r s a l h e a d p r o f i l e
straight to s l i g h t l y concave, d i v i d e d i n t w o s l i g h t l y c o n v e x areas v i z ; the n a p e , a n d
the part o f the h e a d f r o m the c o n c a v i t y above the eye to the snout t i p . P r e m a x i l l a r y
pedicel prominent, but i n most specimens smoothly m e r g i n g into the concavity
a b o v e the eye. L i p s n o r m a l , p r e m a x i l l a not e x p a n d e d m e d i a l l y o r b e a k e d . A relative-
l y s m a l l p a r t o f the m a x i l l a v i s i b l e c a u d a l l y to t h e p r e m a x i l l a r y d e n t i g e r o u s a r m .
Vertical t h r o u g h the p o s t e r i o r e n d of the m a x i l l a not r e a c h i n g the rostral eye m a r g i n .
M o u t h s l i g h t l y to m o d e r a t e l y o b l i q u e . N o m e n t a l p r o m i n e n c e . L a t e r a l s n o u t o u t l i n e
i s o g n a t h o u s . L o w e r j a w s l i g h t l y p r o t r u d i n g , the lateral sides m o d e r a t e l y o b l i q u e .
H o r i z o n t a l a r m of p r e o p e r c u l u m n e a r l y h o r i z o n t a l , the v e r t i c a l a r m r e c l i n i n g c a u -
d a d . C a n a l s a n d o p e n i n g s o f lateral l i n e system o n l a c h r y m a l s m a l l . E y e a n d p u p i l
s l i g h t l y e l o n g a t e d i n h o r i z o n t a l d i r e c t i o n , the p u p i l i n s o m e specimens a n a p h a k i c
space is v i s i b l e rostro-ventrally to the lens.
S c a l e s . — H e a d , rostral part of d o r s u m , chest a n d b e l l y w i t h c y c l o i d scales. Scales
o n r e m a i n i n g part o f the b o d y w e a k l y ctenoid. B o d y scales i m p l a n t e d relatively deep.
Size change b e t w e e n scales of chest a n d flank g r a d u a l . C a u d a l f i n w i t h s m a l l elongate
scales o n its basal half, the scales near the m a r g i n s e x t e n d i n g s o m e w h a t further.
F i n s . — E x c e p t for o n e s e x u a l l y active m a l e i n w h i c h t h e p e l v i c s just reach the
first a n a l s p i n e , pectoral a n d p e l v i c s d o not reach the anal. D o r s a l a n d a n a l f i n just
n o t r e a c h i n g c a u d a l f i n base. C a u d a l f i n o u t l i n e subtrunctate, s l i g h t l y e m a r g i n a t e
a n d slightly oblique.
G i l l r a k e r s . — G i l l rakers h a v e been c o u n t e d i n 12 specimens. N u m b e r o f rakers
o n l o w e r p a r t o f first g i l l arch o n the r i g h t side 8 o r 9. T h e l o w e r 3 s m a l l a n d c o n i c a l ,
n u m b e r s 4-6 e l o n g a t e a n d t h e u p p e r 2 o r 3 s l i g h t l y b r o a d e n e d , s o m e t i m e s w i t h a
s l i g h t l y e x p a n d e d b i f i d or t r i f i d h e a d .
V i s c e r a . — A s the stomachs a n d intestines of the d e s c r i b e d specimens h a d been
r e m o v e d p r e v i o u s l y , n o i n f o r m a t i o n is available o n their lengths a n d c o n f i g u r a t i o n .
O r a l t e e t h . — I n the p r e s e r v e d specimens o n l y a s m a l l p a r t o f the teeth is v i s i b l e
a b o v e the g u m s a n d l i p s ; the teeth therefore seem rather s m a l l . H o w e v e r , w h e n seen
i n a skeletal p r e p a r a t i o n they appear to be o f a n o r m a l size.
— Shape. O u t e r r o w i n b o t h j a w s w i t h a m i x t u r e of acutely p o i n t e d , s l i g h t l y t o
m o d e r a t e l y c u r v e d , u n e q u a l l y b i c u s p i d s , w e a k l y b i c u s p i d s a n d (rostrally) a f e w true
u n i c u s p i d s . T h e n u m b e r of u n i c u s p i d s teeth i n the u p p e r j a w is h i g h e r t h a n that i n

the l o w e r jaw. Rostrally, the teeth are s l i g h t l y c o m p r e s s e d , laterally a n d c a u d a l l y they
are m o d e r a t e l y c o m p r e s s e d . B i c u s p i d s h a v e a r o u n d e d n e c k a n d a c o m p r e s s e d ,
b a s a l l y e x p a n d e d c r o w n . M a j o r c u s p e q u i l a t e r a l , t i p bent s o m e w h a t s u s p e n s o r i a d .
M i n o r c u s p v e r y s m a l l , equilateral to isoscelene, c u s p g a p n a r r o w o r (mostly) n o n -
e x i s t e n t ^ T h e d i s s e c t e d s p e c i m e n o f H. "pellegrini" i n t h e B M ( N H ) c o l l e c t i o n is
e x c e p t i o n a l i n h a v i n g o n l y u n i c u s p i d teeth i n the outer r o w of b o t h jaws.) Inner r o w s
h a v e s m a l l , c o m p r e s s e d , t r i c u s p i d o r w e a k l y t r i c u s p i d teeth, the rostral ones w i t h a
r e l a t i v e l y l o n g m e d i a l cusp. Tooth size i n outer r o w greatest rostrally, decreasing i n
size c a u d a d w h e r e the teeth are less than half the size of the rostral ones.
— D e n t a l arcade a n d t o o t h b a n d . D e n t a l arcade r o u n d e d . O u t e r t o o t h r o w covers
w h o l e l e n g t h o f p r e m a x i l l a r y d e n t i g e r o u s a r m , i n n e r r o w s less t h a n t w o - t h i r d s .
Three (rarely four) i n n e r r o w s i n rostral part of p r e m a x i l l a , decreasing to z e r o cau-
d a l l y ; gap b e t w e e n outer r o w a n d first inner r o w relatively s m a l l rostrally, d i m i n i s h -
i n g c a u d a d . I n l o w e r j a w outer r o w extends to the rostral part of c o r o n o i d w i n g . T w o
i n n e r r o w s i n rostral part of l o w e r j a w decreasing to z e r o just before the c o r o n o i d
w i n g . G a p b e t w e e n outer r o w a n d first inner r o w of s i m i l a r size rostrally a n d lateral-
ly. Just before c o r o n o i d w i n g outer r o w bends m e d i a d c o n t i n u i n g the l i n e of the first
inner row.
— C o u n t s a n d setting. 55 - 58 teeth i n outer r o w of the p r e m a x i l l a , 40 - 48 i n the
l o w e r jaw. Teeth r e g u l a r l y set i n b o t h jaws. N e c k distance o f the teeth less t h a n 1/2 of
d i a m e t e r at toothbase, c r o w n distance laterally less t h a n 1/4 c r o w n w i d t h , c r o w n s
often almost t o u c h i n g each other. C a u d a l l y a n d laterally, teeth m o r e closely set t h a n
rostrally. N o m e d i a l gap i n the u p p e r jaw, a s m a l l one i n the l o w e r jaw.
— I m p l a n t a t i o n . O u t e r r o w teeth of p r e m a x i l l a s l i g h t l y p r o c u m b e n t rostrally,
erect rostro-laterally a n d laterally, a n d s l i g h t l y recumbent caudally. O u t e r r o w teeth
of m a n d i b l e erect r o s t r a l l y a n d laterally, c a u d a l m o s t teeth s l i g h t l y recumbent. Inner
r o w teeth of m a n d i b l e m o d e r a t e l y recumbent, i n n e r r o w teeth of p r e m a x i l l a s t r o n g l y
recumbent.
P h a r y n g e a l t e e t h . — C o u n t s . T w e n t y t w o teeth i n c a u d a l transverse r o w , 11 a n d
13 i n the t w o m e d i a n r o w s .
— Shape. Teeth of c a u d a l m o s t r o w , a n d t w o o r three c a u d a l teeth of the m e d i a n
r o w s h o o k e d a n d b l u n t ; other teeth b e v e l l e d . Tooth size increases f r o m rostral to cau-
d a l a n d f r o m lateral to m e d i a l .
O s t e o l o g y — T h e osteological d e s c r i p t i o n of the o r a l j a w s a n d the l o w e r p h a r y n -
geal e l e m e n t are based o n s p e c i m e n B M ( N H ) 1962.3.2.544, S L 77.4 m m . T h e d e s c r i p -
t i o n of the o r a l teeth is based o n a l l specimens. A s I a m n o t sure about the i d e n t i t y of
the B M ( N H ) skeletal m a t e r i a l l a b e l l e d H. pellegrini (oral teeth d i f f e r i n g f r o m a l l other
s p e c i m e n s o f H. harpakteridion) I h a v e not u s e d this m a t e r i a l for the d e s c r i p t i o n .
— O r a l jaws. P r e m a x i l l a . D e n t i g e r o u s a r m l o n g e r than a s c e n d i n g a r m , angle be-
t w e e n the arms 7 6 ° . V e n t r a l o u t l i n e of dentigerous a r m concave. V e n t r a l quarter of
a s c e n d i n g a r m e x p a n d e d r o s t r a d . M a n d i b l e : m o d e r a t e l y stout a n d short, l e n g t h /
d e p t h ratio 2.1. Tooth-bearing part of m a n d i b l e s l i g h t l y less t h a n half total m a n d i b l e
l e n g t h . C a u d a l m o s t teeth of outer r o w just r e a c h i n g c o r o n o i d w i n g .
— L o w e r p h a r y n g e a l element. L o w e r p h a r y n g e a l element of g e n e r a l i z e d t y p e ,
s l i g h t l y l o n g e r t h a n b r o a d ( l e n g t h / w i d t h = 1.1). D e n t i g e r o u s area s l i g h t l y l o n g e r
t h a n b r o a d ( l e n g t h / w i d t h = 1.05).
— Vertebrae. There are 28 (f= 5) o r 29 (f= 1) vertebrae, c o m p r i s i n g 12 (f= 4) o r 13

(f= 2) a b d o m i n a l a n d 15 (f= 1) or 16 (f= 5) c a u d a l elements.
C o l o r a t i o n . — G r e e n w o o d (1962) d e s c r i b e d the l i v e colours of female H. pellegri-
ni f r o m the E A F R O collections. A s the c o l o r a t i o n of the e r r o n e o u s l y i n c l u d e d female
of H. perrieri is k n o w n te be different (pers. obs.) a n d t w o of the three other errone-
o u s l y i n c l u d e d specimens are males, the data are b e l i e v e d to be a p p l i c a b l e to the fe-
males of H. harpakteridion.
— L i v e coloration of females: A c c o r d i n g to G r e e n w o o d (1962: 188) ' l i v e females
h a v e a d a r k chocolate-brown g r o u n d colour, lighter o n the chest a n d belly. A l l the fins
are b l a c k i s h - b r o w n . " T h e u n i f o r m l y d a r k coloration of the fins, especially of the p e l v i c
fins, is rare amongst females of the L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d species.
— L i v e c o l o r a t i o n of males is u n k n o w n .
— P r e s e r v e d c o l o r a t i o n of males. B o d y a n d h e a d u n i f o r m l y d a r k b r o w n , d o r s a l
h e a d surface a n d d o r s u m b l a c k i s h . G i l l - c o v e r a n d chest area w i t h a lighter b r o n z e
h u e . M a r k i n g s : a s m a l l o p e r c u l a r b l o t c h a n d a d i s t i n c t l a c h r y m a l stripe e x t e n d i n g
f r o m v e n t r a l e y e b o r d e r to b e l o w m o u t h corner. A l l fins d a r k b r o w n , the p e l v i c s
b l a c k l a t e r a l l y a n d distally, d a r k streaks b e t w e e n rays o f d o r s a l a n d c a u d a l . L a p p e t s
of d o r s a l f i n b l a c k rostrally. T w o s m a l l e g g spots w i t h a d a r k m a r g i n o n the c a u d a l
part of the a n a l f i n .
— P r e s e r v e d c o l o r a t i o n o f females. B o d y a n d h e a d u n i f o r m l y b r o w n , the d a r k -
ness of w h i c h differs (possibly as a result of preservation?) f r o m l i g h t b r o w n to v e r y
d a r k b r o w n . T w o b r o o d i n g females h a v e the lighter c o l o r a t i o n . G e n e r a l l y the o p e r c u -
l u m a n d chest area lighter than the s u r r o u n d i n g areas. A l l fins u n i f o r m l y b r o w n i s h ,
the p e l v i c s w i t h b l a c k tips a n d lateral sides i n a f e w specimens. M o s t specimens h a v e
a n o p e r c u l a r b l o t c h , a n d a l a c h r y m a l stripe, distinct o r faint, is present i n a l l speci-
mens.
D i s t r i b u t i o n . — Haplochromis harpakteridion is o n l y k n o w n from Lake Victoria. A l l
s p e c i m e n s w e r e collected b y w o r k e r s of E . A . F . R . O . at three localities situated rela-
t i v e l y closely together i n the n o r t h e r n part of the l a k e (i.e. G r a n t Bay, K a r e n i a , a n d
B u v u m a C h a n n e l ) . E x t e n s i v e s a m p l i n g i n the M w a n z a G u l f d i d n o t y i e l d a n y repre-
sentatives of this species, p o s s i b l y because its preferred habitat is rare i n the M w a n z a
Gulf.
E c o l o g y . — H a b i t a t . A c c o r d i n g to G r e e n w o o d (1962: 188) a l l specimens w e r e c o l -
l e c t e d at r e l a t i v e l y e x p o s e d s a n d y beaches n e a r d e n s e s t a n d s o f s u b m e r g e d a n d
emergent plants.
— F o o d . A c c o r d i n g to G r e e n w o o d (1962: 188) " O f t h e t h i r t y - f i v e s p e c i m e n s
e x a m i n e d , t w e n t y - f i v e h a d ingested m a t e r i a l i n the g u t . I n three fishes, this m a t e r i a l
w a s a n u n i d e n t i f i a b l e s l u d g e , i n o n e there w a s s l u d g e p l u s insect r e m a i n s , f i v e h a d
f e d o n l y o n insects ( p r i n c i p a l l y E p h e m e r o p t e r a n p u p a e a n d l a r v a l Diptera) o n e o n
insects a n d s m a l l fishes, a n d fifteen h a d f e d e x c l u s i v e l y o n fishes. T h e f i s h remains
w e r e v e r y f r a g m e n t a r y b u t i n most cases w e r e i d e n t i f i a b l e as c i c h l i d s . T h e size range
of the p r e y is 10-15 m m , that is i m m e d i a t e l y p o s t - l a r v a l fishes". T w o of the f o u r spec-
i m e n s w h i c h I r e m o v e d f r o m the g r o u p m e n t u i o n e d above are k n o w n to be p i s c i v o -
rous. J u d g i n g f r o m the m o r p h o l o g y , the other t w o m a y also be p i s c i v o r e s .
— B r e e d i n g a n d g r o w t h . H. harpakteridion i s a f e m a l e m o u t h b r o o d e r . L o t
B M ( N H ) 1962.3.2.513-515 contains a female of 80.5 m m S L w h i c h carries e m b r y o s (±
8 m m T L ) w i t h a large y o l k sac, a female of 72.8 m m S L c a r r y i n g eggs (length 3.5x2.7
Figs 90-103. Haplochromis harpakteridion spec. nov. Fig. 90. Premaxilla, lateral view Fig. 91. Premaxilla,
perpendicular view of dentigerous area. Fig. 92. Premaxilla, medial view of ascending arm. Fig. 93.
Lower jaw, lateral view. Fig. 94. Lower pharyngeal element, dorsal view. Fig. 95 Lower pharhyngeal
element, lateral view. Fig. 96. Lower pharhyngeal element, caudal view. Figs. 97-100. Premaxillary
teeth, labial and lateral view. Fig. 97. Second tooth of outer row. Fig. 98. Eighth tooth of outer row. Fig.
99. Thirteenth tooth of outer row. Fig. 100. Fifth tooth of first inner row. Figs. 101-103. Lower phahyn-
geal teeth, rostral and lateral view. Fig. 101. Tenth tooth of lateralmost row. Fig. 102. Second tooth of
formost caudal row. Fig. 103. Third medial tooth of caudalmost row. Scale = 1 mm.
m m ) , a n d a t h i r d female w i t h a d i s t e n d e d b u c c a l c a v i t y a n d spent ovaries w h i c h m a y

also h a v e been b r o o d i n g at the t i m e of capture. N o i n f o r m a t i o n is a v a i l a b l e o n the
date of capture. A l l e x a m i n e d specimens are adults o r s u b a d u l t s .
R e s e m b l i n g s p e c i e s . — N o t w i t h s t a n d i n g the s m a l l size of the adults, H. harpak-
teridion is i m m e d i a t e l y recognizable as a p i s c i v o r o u s species b y its slender b o d y , its
r e l a t i v e l y large h e a d , its l o n g l o w e r j a w a n d b y the shape of its teeth. F r o m the other
s m a l l p i s c i v o r e s , H. martini (Blgr., 1906), H. perrieri, H. percoides Blgr., 1915, a n d H.
nanoserranus G r e e n w o o d & Barel, 1976, it is i m m e d i a t e l y separable b y its b o d y shape
a n d c o l o u r pattern. A p a r t f r o m the smaller size of the a d u l t s H. harpakteridion can be
d i s t i n g u i s h e d f r o m H. pellegrini b y its m o r e c o n v e x d o r s a l h e a d p r o f i l e , its less
o b l i q u e m o u t h , its g e n e r a l l y m o r e p r o m i n e n t p r e m a x i l l a r y p e d i c e l , its r o u n d e d m e n -
tal area, the presence of a l a c h r y m a l stripe a n d b y its d e n t i t i o n . M o r p h o m e t r i c ratio
ranges o f L J W / H L a n d P P L / H L s h o w n o o v e r l a p a n d those of H W / H L , C F L / H L ,
S n L / H L , L a W / H L a n d P O D / H L s h o w o n l y v e r y slight o v e r l a p . T h e shape of the
h e a d is c o m p a r a b l e w i t h that of s o m e s p e c i m e n s n o w i n c l u d e d i n H. prognathus
( P e l l e g r i n , 1904).
C o n c l u d i n g remarks
W i t h the exception of H. pellegrini the species d e s c r i b e d here e x h i b i t s o m e charac-

ters r a r e l y f o u n d a m o n g s t t h e L a k e V i c t o r i a h a p l o c h r o m i n e c i c h l i d s . W h e r e a s
f e m a l e s o f m o s t species are d r a b l y c o l o u r e d , w i t h g r e y i s h s i l v e r o r s a n d y g r o u n d
c o l o u r s p r e d o m i n a t i n g ( G r e e n w o o d , 1974; p e r s . o b s . ) , f e m a l e s o f H. bareli, H.
dichrourus, H. maisomei, H. kujunjui, H. pyrrhopteryx, H. chrysogynaion, a n d H. harpak-
teridion h a v e distinct colours o n b o d y a n d fins. I n H. dichrourus a n d H. pyrrhopteryx
the c o l o r a t i o n o f the females greatly resembles that of the males. C o l o r a t i o n of l i v e
males of H. harpakteridion is u n k n o w n , b u t as p r e s e r v e d c o l o r a t i o n of the sexes does
not differ m u c h , females of this species m a y also resemble males i n l i v e c o l o r a t i o n . I n
H. bareli the resemblance of female a n d m a l e c o l o r a t i o n is n o t so s t r i k i n g a l t h o u g h
the b l a c k area of the lateral side i n female is v a r i a b l e : m o s t females h a v e o n l y the
v e n t r a l half o f the b o d y black b u t i n some specimens the b l a c k area stretched f r o m
the d o r s a l f i n base d o w n to the v e n t r a l side ( w h i c h a l w a y s remains b r i g h t w h i t e ) . A n
i m p o r t a n t aspect of m a l e c o l o r a t i o n - d a r k o r black p e l v i c fins - present i n females of
H. dichrourus, H. maisomei, H. kujunjui, H. pyrrhopteryx a n d H. harpakteridion, is n o t
f o u n d i n females of H. bareli a n d H. chrysonotus. T h e c o l o r a t i o n of H. dichrourus, H.
pyrrhopteryx a n d H. harpakteridion, a n d the males of H. bareli is also exceptional i n that
t h e b o d y is d a r k e s t v e n t r a l l y i n s t e a d o f c o u n t e r s h a d e d as i n m o s t o t h e r h a p -
l o c h r o m i n e species.
H. bareli, H. dichrourus, H. maisomei, H. pyrrhopteryx a n d H. chrysonotus a l l possess
s t r o n g l y c u r v e d u n i c u s p i d s teeth i n the outer r o w s o f the o r a l jaws. T h e teeth of H.
harpakteridion a n d H. kujunjui are s l i g h t l y less r e c u r v e d a n d those of H. harpakteridion
o n l y p a r t l y u n i c u s p i d . I n teeth r e c u r v a t i o n as w e l l as i n f e m a l e c o l o r a t i o n (male
m i m i c r y ; v a n O i j e n , i n prep.) H. pyrrhopteryx a n d H. dichrourus are the m o s t d i v e r -
gent species. H. bareli a n d H. chrysonotus are closer to these species i n d e n t i t i o n , w h i l e
H. harpakteridion (and p r o b a b l y H. kujunjui ) m i g h t be closer i n aberrant female c o l -
oration and headmarkings.
Acknowledgements
I wish to thank the Tanzanian gouvernment for permission to carry out held work from 7/1977 to
10/1980 and again in June 1985. For laboratory facilities, fishing gear, assistance and hospitality I
thank the officers in charge and staff of the Mwanza Fisheries Research Centre and the Freshwater
Fisheries Institute at Nyegezi. For hospitality and assistance on board of the M.V. Mdiria I am indebt
ed to Mr J. Kayungi and especially to Mr. Teunis Ras, who also gave assistance in many other ways.
Frans and E ls Witte, Rob Hoogerhoud, Pirn Wilhelm, Alois and Mohja are thanked for assistance in
collecting specimens. Special thanks are due to Kees Goudswaard, who collected specimens from
many localities in the years after I left Tanzania. For making available photographic material and Xray
facilities I thank the Morphology department of the Leiden University. I thank Schelte Zeilstra for
making many excellent Xray photographs, and E ugène van E sch and Henk Olivier for assistance in
making the black and white photographs. Adri 't Hooft is thanked for making slides during the '85
expedition. Thanks are also due to Hans Tegelaar for his assistence in making skeletal preparations
and to Eric Bosch for preparing figures 3 and 4. For permission to work in the BM(NH) collection I
thank Drs. P H . Greenwood and A . Wheeler. I am greatly indebted to Dr P H . Greenwood and Gordon
Howes for their personal interest, warm hospitality and facilities given me during many visits to 'the
Fish Section'. I thank the trustees of the Natural History Museum, London, for their permission to
reproduce figures from Regan, 1922. For comments on various drafts of the manuscript I am indebted
to Drs P H . Greenwood, C . D . N . Barel, F. Witte, R.J.C. Hoogerhoud, W. Wilhelm and Prof. Dr Ε.
Gittenberger. To Drs C . D . N . Barel and F. Witte I am indebted also for many stimulating discussions
throughout the preparation of this paper. Dr RT. van Straten is thanked for assistance in finding suit
able latinized Greek names, and Mrs J. Hammouda for typing various drafts of this paper. Fieldwork
(19771980) has been financially supported by the Netherlands Foundation for the advancement of
Tropical Research (WOTRO) grant W78129; the fieldtrip in 1985 was partly financed by the JanJoost
ter Pel wijk fund.
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Edited: E. Gittenberger & J.C. den Hartog
Fig. 104. Haplochromis bareli spec. nov. Holotype, R M N H 29600, dorsal, lateral and ventral view. Scale
equals 10 mm.
Fig. 105. Haplochromis dichrourus Regan. Holotype, BM(NH) 1906.5.30:265, dorsal, lateral and ventral
view. Scale equals 10 mm.
Fig. 106. Haplochromis dichrourus "Majita". BM(NH) 1966.3.9.183, SL 167.0 mm, dorsal, lateral and ven-
tral view. Scale equals 10 mm.
Fig. 107. Haplochromis dichrourus "Mwanza". R M N H 30060, SL 132.0 mm, dorsal, lateral and ventral
Fig. 108. Haplochromis dichrourus "Mwanza". R M N H 30088, dorsal view of head showing M-shaped
nasal band.
Fig. 109. Haplochromis maisomei spec. nov. Holotype, R M N H 30112, SL 96.1 mm, dorsal, lateral and
ventral view. Scale equals 10 mm.
Fig. 110. Haplochromis kujunjui spec nov. Holotype, R M N H 30466, SL 102.8 mm, dorsal, lateral and
Fig. 111. Haplochromis pyrrhopteryx spec. nov. Holotype, R M N H 30010, SL 154.3 mm, dorsal, lateral and
Fig. 112. Haplochromis chrysogynaion spec. nov. Holotype, R M N H 30118, SL 89.0 mm, dorsal and lateral
Fig. 113. Haplochromis pellegrini Regan. Lectotype, BM(NH) 1906.5.30.253, SL 103.8 mm, dorsal, lateral
and ventral view. Scale equals 10 mm.
Fig. 114. Haplochromis harpaktendion spec. nov. Holotype, BM(NH) 1962.3.2.516, SL 72.3 mm,
dorsal, lateral and ventral view. Scale equals 10 mm.

Systematic revision of piscivorous cichlids of Lake Victoria

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Systematic revision of piscivorous cichlids of Lake Victoria

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A systematic revision of the piscivorous haplochromine Cichlidae

(Pisces: Teleostei) of Lake Victoria (East Africa). Part I

c i c h l i d fishes (11 f a m i l i e s a n d c. 50 species), not o n l y i n n u m b e r of species b u t also i n

Material, methods a n d techniques

T h e major part of the m a t e r i a l d e s c r i b e d b e l o w w a s collected f r o m t r a w l catches

a n d W i t t e & W i t t e - M a a s (1981). T w o n e w measurements a n d a q u a l i t a t i v e character

G r e e n w o o d ' s r e v i s i o n of the genus Haplochromis: i n t r o d u c t i o n

ferences are consistent w i t h the p o s s i b i l i t y that the h a p l o c h r o m i n e species flock arose

I m p l i c a t i o n s of the revisions for t a x o n o m y

c o m b i n e s characters of three genera; it is intermediate b e t w e e n Astatotilopia, Labro-

G r e e n w o o d ' s generic classification of the p i s c i v o r o u s species: i n t r o d u c t i o n

the "serranus" g r o u p p i s c i v o r e s ' . H o w e v e r , i n G r e e n w o o d (1980) Haplochromis ser-

tensies m i g h t be e x p l a i n e d b y the fact that G r e e n w o o d ' s statements about s i m i l a r i t y

Table 1. Neurocranial mesurements of species placed in Harpagochromis, Prognathochromis and Psam-

ranges : Greenwood, 1980 { 1

Fig. 8. Neurocranial measurements of species placed by Greenwood (1980) in Harpagochromis,

P e r h a p s the d e f i n i t i o n of the genus Haplochromis s h o u l d be b r o a d e n e d rather t h a n

t i n c t i o n a p p e a r e d n a r r o w e r t h a n that of p r e v i o u s authors. M a t e r i a l u s e d for redes-

Haplochromis "black"; van Oijen, 1982: 340,349.

E t y m o l o g y . — T h i s species is n a m e d i n h o n o u r of D r C . D . N . Barel, the initiator of

— C o u n t s a n d setting. 52-62 teeth i n the outer r o w of the u p p e r jaw, 36-52 i n the

— S e x u a l l y active m a l e s . A l m o s t entirely b l a c k , the r e d spots b e t w e e n the f i n

Standard length range in mm 41.0-57.5 63.3-78.1 81.0-89.5 90.8-98.4 100.0-109.1 114.1-125.0

Body Depth %SL 30.7-33 8 33.7-37.3 33.3-37.9 33.4-36.7 34.2-36.1 35.3-37.8

Standard length range in mm 41.0­57.5 63.3­78.1 81.0­89.5 90.8­98.4 100.0­109.1 114.1­125 ζ

Eye Length % HL 33.913.0

w h i t e rostrally, m o r e h y a l i n e c a u d a l l y O n e or t w o s m a l l y e l l o w orange spots o n the

ranges mean ± st.dev.

Dorsal Head Profile Inclination 27°-43° 33.0° ±4°

Dorsal Head Profile (curvature) +

Dorsal Fin (Spines/Rays) XIV 9 (f= 1); XV8(f=l); XV9(f=25);

T h e r i g h t o v a r y g r o w s larger t h a n the left o n e . E g g s m e a s u r e d i n r i p e o v a r i e s o f

Paratilapia serranus (part): Boulenger, 1915: 334

D i a g n o s i s . A m e d i u m to m o d e r a t e l y large ( d e p e n d i n g o n the l o c a l i t y ; see b e l o w )

N o t e s o n the B M ( N H ) collection of H. dichrourus

D e s c r i p t i o n . — Based o n the h o l o t y p e a n d f i v e other s p e c i m e n s , 82.6-147.4 m m

non-existent. L o w e r j a w sides s l i g h t l y o b l i q u e . R o s t r a l m a r g i n of v e r t i c a l a r m of pre-

Haplochromis dichrourus Haplochromis maisomei

Locality Karenia Grant Bay Buganga Jinja Majita Maisome I. Standieri I.

— I m p l a n t a t i o n . U p p e r jaw, outer r o w ; rostrally erect, laterally s o m e w h a t r e c u m -

D e s c r i p t i o n . — H a b i t u s . T h e s p e c i m e n s of H. dichrourus " M a j i t a " are r e l a t i v e l y

s m a l l e r s p e c i m e n a n d d o r s a l a n d a n a l of the larger s p e c i m e n r e a c h i n g the base o f the

strongly contrasting w i t h black branchiostegal membrane a n d the black inter­

3. Haplochromis dichrourus " M w a n z a "

D e s c r i p t i o n . — Based o n 57 specimens of 70.5-141.0 m m S L .

outer r o w proceeds s l i g h t l y further c a u d a d than the inner r o w , a n d just reaches the

a b o v e the e y e , the stripes of b o t h sides m e e t i n g m e d i a l l y . S n o u t c r o s s e d b y a b r o a d

SLinmm 81.7-88.9 90.2-98.0 100.9-109.3 110.9-119.1 121.2-128.0 132.0-141.0

BD/SL 31.8-35.9 30.4-33.9 31.1-35.3 31.5-36.9 33.2-36.2 33.3-37.4

HW/HL 40.0-43.8 395-44.8 40.4-44.7 40.8-45.5 40.8-44.8 41.3^5.0

SL in mm 70.5 81.7-88.9 90.2-98.0 100.9-109.3 110.9-119.1 121.2-128. 132.0-141.0 ζ

of the o p e r c u l u m a n d the d o r s a l o u t l i n e of the n a p e a b r o a d n a p e b a n d r u n s m e d i a d .

v e r y d a r k h e a d c o l o u r s , a b r o a d l a c h r y m a l stripe, w h i c h is e x t e n d e d above the eye,

January a n d June. A l l specimens w i t h over 100 m m S L are mature. Females g r o w to a

o r c o n t i n u o u s interorbital stripe a n d a n a p e b a n d are present. T h e n o s t r i l stripe has a

the d i f f e r e n c e s i n h e a d m a r k i n g w o u l d p r e v e n t the m e m b e r s of the p o p u l a t i o n s of H.

range mean and standard

Dorsal Head Inclination 26°-37° 31.1°±3.1°

Dorsal Head Profile (curvature) 0(+)

D e s c r i p t i o n . — Based o n the h o l o t y p e , a s e x u a l l y active m a l e 96.1 m m S L a n d one

of b o d y i v o r y . A v a g u e m i d l a t e r a l b a n d rostrally starting t h i n i n the grey area a n d

i n m a l e l i v e c o l o r a t i o n f r o m those of the M w a n z a G u l f . In that case they too s h o u l d

Standard length range in mm 41.057.5 63.378.1 81.089.5 90.898.4 100.0109.1 114.1125 ζ

strongly contrasting w i t h black branchiostegal membrane a n d the black inter

Haplochromis dichrourus (part); Greenwood, 1967: 6569; specimen BM(NH) 1966.3.9.185.

E t y m o l o g y . T h e s p e c i f i c l n a m e ( f r o m the G r e e k " π υ ρ ρ ο δ " = r e d d i s h , y e l l o w r e d ,

E t y m o l o g y . T h e n a m e is d e r i v e d f r o m the d i m i n u t i v e o f the G r e e k απνακνπ (rob