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Neolamprologus devosi sp. n., a new riverine lamprologine cichlid (Teleostei, Cichlidae) from the lower Malagarasi River, Tanzania

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Neolamprologus devosi sp. n., a new species of riverine lamprologine cichlid is described from the lower Malagarasi River, Tanzania. The new species is tentatively placed within the poorly defined genus Neolamprologus, though generic reassignment may be necessary once ongoing work on the phylo-geny and classification of lamprologines is completed. Notably, Neolamprologus devosi sp. n. does not present characters diagnostic of Congo (ex-Zaire) River lamprologines, and contrary to previous suggestions appears to be phylogenetically distinct from those taxa. This raises questions regarding the number of putative riverine colonizations of lamprologines from a lacustrine source.
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Accepted: 24 November 2003; published: 1 December 2003 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2003 Magnolia Press
Zootaxa 373: 111 (2003)
www.mapress.com/zootaxa/
Neolamprologus devosi sp. n., a new riverine lamprologine cichlid
(Teleostei, Cichlidae) from the lower Malagarasi River, Tanzania
ROBERT SCHELLY1,2, MELANIE L.J. STIASSNY1 & LOTHAR SEEGERS3
1. Department of Ichthyology, American Museum of Natural History, 79th Street at Central Park West, New
York, NY 10024, USA. Email: schelly@amnh.org and mljs@amnh.org
2. Department of Ecology, Evolution and Environmental Biology, and Center for Environmental
Research and Conservation, Columbia University, New York, USA.
3. Hubertusweg 11, D 46535 Dinslaken, Germany. Email: L.Seegers@t-online.de
Abstract
Neolamprologus devosi sp. n., a new species of riverine lamprologine cichlid is described from the
lower Malagarasi River, Tanzania. The new species is tentatively placed within the poorly defined
genus Neolamprologus, though generic reassignment may be necessary once ongoing work on the
phylo-geny and classification of lamprologines is completed. Notably, Neolamprologus devosi sp.
n. does not present characters diagnostic of Congo (ex-Zaire) River lamprologines, and contrary to
previous suggestions appears to be phylogenetically distinct from those taxa. This raises questions
regarding the number of putative riverine colonizations of lamprologines from a lacustrine source.
Key words: Neolamprologus, new riverine species, lamprologines
Introduction
De Vos et al. (2001) provide an up-to-date summary of current knowledge of the ichthyo-
fauna of the Malagarasi River, the largest affluent of Lake Tanganyika in Burundi and Tan-
zania. They note the mixed origin of the system’s fishes, and estimate that, in addition to
Congolese and Nilotic elements, about 15% of the fauna is comprised of Malagarasi
endemics—a tally which includes a number of undescribed species. Among these is a
small lamprologine cichlid, which De Vos et al. posit is closely related to Lamprologus
mocquardi Pellegrin, 1903 from the Congo River basin. However, results from a revision
of the riverine species of Lamprologus of the Congo (ex-Zaire) basin including Lamprolo-
gus mocquardi are at odds with that assessment (Schelly & Stiassny, submitted). The Mal-
agarasi lamprologine does not belong to the clade of riverine Lamprologus, and instead is
tentatively placed within the genus Neolamprologus and described herein.
SCHELLY ET AL.
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ZOOTAXA Materials and methods
Counts and measurements follow Barel et al. (1977). All measurements of bilaterally
paired structures were taken on the left side with digital calipers with an accuracy of +/-
0.05 mm. Clearing and staining of bones and cartilages follows Dingerkus & Uhler (1977).
Water parameters were taken in the field using a WTW pH 96-B microprocessor pH meter,
a WTW OXI 96-B microprocessor Oximeter, and a Bischof L 17 conductimeter, German
hardness was measured using a DUPLA test-kit.
Abbreviations are: AMNH, American Museum of Natural History, New York; MRAC,
Musée Royal de l’Afrique Centrale, Tervuren; SL, standard length; HL, head length; BD,
body depth.
The following sources were consulted for comparative counts and measurements of all
known species of Neolamprologus: Boulenger (1915), Büscher (1989, 1991, 1992a,
1992b, 1993, 1995a, 1995b, 1997), Konings (1998), Poll (1956, 1978, 1986), Staeck
(1980), Staeck and Seegers (1986).
Note on generic assignment. The generic-level classification of lamprologine cichlids is
in need of thorough revision (Stiassny, 1997). Pending the results of a wide-ranging phy-
logenetic analysis (Schelly, in prep.), the new species described herein is provisionally
placed in the genus Neolamprologus Colombe & Allgayer, 1985. In so doing we follow
the phenetic classification of Poll (1986) who placed within the genus Neolamprologus all
lamprologines in which the first pelvic ray is the longest in the fin (in contrast to the condi-
tion in Lamprologus sensu Poll, in which the second or third pelvic ray is the longest), and
which lack the defining characters of Chalinochromis, Julidochromis, Telmatochromis,
Altolamprologus, and Lepidiolamprologus. Although it is probable that the genus Neolam-
prologus, as it is presently constituted, is non-monophyletic we have chosen this generic
assignment as a pragmatic approach to facilitate species description. It is highly probable
that a generic reassignment will be necessary once a fuller understanding of the generic
limits and relationships amongst lamprologine clades is gained.
Neolamprologus devosi, new species
(Fig. 1)
Holotype. AMNH 233614, male, 56.9 mm SL, a few kilometers upstream from Ilgala,
lower course of Malagarasi River, 5°11'59"S 29°49'59"E, Tanzania, L. Seegers, 02/07/94.
Paratypes. Total of 24 specimens, 23.0-57.1 mm SL; AMNH 233615, male, 57.1 mm
SL, a few kilometers upstream from Ilgala, lower course of Malagarasi River, 5°11'59"S
29°49'59"E, Tanzania, L. Seegers, 02/07/94. —MRAC 93-152-P-1385-1394, 23.0-35.6
mm SL, delta Malagarasi, right arm, Tanzania, 5°13’S 29°48’E, L. DeVos, 20/08/1993. —
MRAC 96-031-P-0528-0537, 28.1-53.3 mm SL (of which 2 specimens are cleared and
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double stained), delta Malagarasi, right arm, Tanzania, 5°13’S 29°48’E, L. De Vos, 20/08/
1993.
Differential diagnosis. Neolamprologus devosi is distinguished from the following
congeners in having fewer than 40 scales in the longitudinal series: N. furcifer, N. christyi,
N. prochilus, N. pleuromaculatus, N. hecqui, N. meeli, N. boulengeri, and N. bifasciatus.
With 34–35 lateral line scales, N. devosi is further distinguished from N. variostigma,
which has 39–40 lateral line scales. A low gill raker count (5–6) on the lower limb of the
first arch distinguishes N. devosi from the following taxa that have 8 or more rakers: N.
brevis, N. longicaudatus, N. moorii, N. sexfasciatus, N. toae, N. leleupi, and N. calliurus.
The presence of 5–6 anal fin spines distinguishes N. devosi from both N. niger and N. fas-
ciatus, which have 8 or more anal spines, and from N. tetracanthus, which has only 4 anal
spines. The caudal fin of Neolamprologus devosi is enlarged, rounded, and paddle shaped,
distinguishing it from the following species with emarginate caudal fins, often also with
the outer caudal rays produced and filamentous: N. mondabu, N. leloupi, N. caudopuncta-
tus, N. savoryi, N. falcicula, N. gracilis, N. crassus, N. marunguensis, N. pulcher, N. bri-
chardi, N. buescheri, N. splendens, N. olivaceous, and N. helianthus. Lacking their
striking coloration and large molariform lower pharyngeal jaw teeth, N. devosi is distin-
guished from N. tretocephalus and N. modestus; similarly it lacks the numerous vertical
bars of N. similis, N. cylindricus, and N. multifasciatus, the solid orange coloration of N.
longior, and the solid black coloration of N. schreyeni. With a body depth of 22.5–25.9%
SL, N. devosi is shallower-bodied than N. mustax (28.3–33.4% SL), N. wauthioni (28.6–
29.4% SL), N. petricola (30.3–35.7% SL), and N. obscurus (25.6–34.8% SL), which is
further distinguished in having 7–8 anal spines compared to 5–6 in N. devosi. Neolampro-
logus devosi has fewer vertebrae (31–32) than N. nigriventris and N. pectoralis, which
have 34 or more vertebrae, and finally, N. devosi lacks the extremely elongate pelvic fins
characteristic of N. ventralis.
Description: Counts and measurements for the type series are given in Table 1. A rel-
atively gracile, elongate species (BD 22.5–25.9, mean 23.9% SL), bearing a superficial
resemblance to Lamprologus mocquardi from the Congo River, but differing from that
taxon in the possession of a pelvic fin with the first ray longer than the second rather than
vice versa as in L. mocquardi, and in the absence of tubular elements in the infraorbital
series (see below). Greatest body depth at about base of second or third dorsal spine. Head
length 32.5–35.0, mean 33.8% SL. Dorsal head profile slightly interrupted by prominent
premaxillary pedicel then smoothly rounded to the dorsal fin origin. Dorsal body profile
convex, curving gently downward along the length of the dorsal fin base to the caudal
peduncle; ventral body profile somewhat rounded and posteriorly curving gently upward
just anterior to caudal peduncle.
Fins. Dorsal fin with XVII–XIX (mode XIX) spines and 8–9 (mode 8) soft rays. Anal
fin with V–VI (mode VI) spines and 6–7 (mode 7) soft rays. Spines in both fins gradually
increase in length posteriorly. Dorsal and anal fins with tapering filamentous extensions
SCHELLY ET AL.
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ZOOTAXA reaching to about the middle of the caudal fin in larger individuals, and just beyond the
caudal fin base in juveniles. Caudal fin enlarged, rounded, and paddle-shaped, with 14
branched rays. Pectoral fins short, not reaching a vertical through the anus, with 13 rays
reaching to fin margin. First ray is the longest in the pelvic fin, and this is produced in
larger specimens and reaches just beyond the second anal fin spine, and to the level of the
anus in juveniles.
Teeth (Fig.2). Lower jaw slightly prognathous, with both outer and inner row teeth
pointed unicuspids in both jaws. A series of 4–8 enlarged, recurved, procumbent canines
situated anteriorly in the jaws with the largest teeth furthest from the symphysis. Posterior
to the enlarged procumbent canines is a single row of slightly enlarged canines extending
almost the entire length of the dentigerous arm of the dentary and premaxilla. Inner teeth
in 3 rows of tightly-packed, small, recurved caniniform teeth tapering by mid-jaw into a
single row.
TABLE 1. Morphometric and meristic data for Neolamprologus devosi sp. n. (Numbers in paren-
thesis refer to number of specimens).
Character holotype N mean min max SD
Standard length (mm)
PERCENT OF STANDARD LENGTH
Body depth
Head length
Caudal peduncle depth
Caudal peduncle length
Anal fin base length
Dorsal fin base length
Pelvic fin length
Caudal fin length
Pectoral fin length
Pre-dorsal distance
Pre-anal distance
Pre-pectoral distance
Pre-pelvic distance
PERCENT OF HEAD LENGTH
Lower jaw length
Eye diameter
Snout length
Interorbital width
56.9
26.5
32.7
10.8
13.9
22.5
60.7
25.6
25.7
21.8
31.5
61.6
34.8
36.4
41.4
27.1
36.4
21.1
24
17
17
17
17
17
17
17
17
17
17
17
17
17
17
17
17
17
38.2
24.1
33.7
10.1
14.1
22.8
56.2
24.8
25.9
21.9
33.1
63.4
36.0
38.0
36.5
26.1
31.4
16.8
23.0
22.5
32.5
8.5
12.6
20.4
53.6
21.7
22.4
18.5
31.5
60.2
34.2
35.8
29.0
23.7
28.1
12.3
57.1
26.5
35.0
11.2
16.9
28.4
60.7
32.0
28.4
23.9
35.0
69.3
39.0
41.2
41.8
27.9
36.4
21.1
1.28
0.87
0.79
1.25
2.17
2.30
2.64
1.51
1.55
1.01
2.24
1.39
1.81
3.53
1.21
2.65
2.38
COUNTS
Lateral line scales
Dorsal fin
Anal fin
Gill rakers
Vertebrae
35
XIX, 9
VI, 7
5,1,3
14+18
34 (12), 35 (5)
XVII 8 (1), XVIII 8 (4), XVIII 9 (3), XIX 8 (4), XIX 9 (5)
V 7 (3), VI 6 (4), VI 7 (10)
5,1,2 (1), 5,1,3 (8), 6,1,2 (1), 6,1,3 (4), 6,1,4 (3)
14+17 (2), 14+18 (14)
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FIGURE 1. Holotype of Neolamprologus devosi sp.n., AMNH 233614, Male, 56.9 mm SL, a few
kilometers upstream from Ilgala, lower course of Malagarasi River, 5°11'59"S 29°49'59"E, Tanza-
nia.
FIGURE 2. Neolamprologus devosi sp. n., MRAC 96-301-P-0528-0537, 44.8 mm SL. Isolated
buccal jaws in lateral view.
Gill rakers (Fig. 3c). Relatively short, blunt and non-denticulate. 8–11 (mode 9) gill
rakers along the outer row of the first gill arch. No rakers present on the hypobranchial, 5–
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ZOOTAXA 6 (mode 5) along the ceratobranchial, one raker in the angle of the arch, and 2–4 (mode 3)
epibranchial rakers.
Lower pharyngeal jaw (Fig. 4). Lower pharyngeal jaw wider than long, with straight,
or very slightly interdigitating, ventral suture. Usually 20–24 teeth in the posterior tooth
row. Median row teeth slightly enlarged and with a robust hook, lateral teeth slender and
either beveled or bluntly hooked.
FIGURE 3. Neolamprologus devosi sp. n., MRAC 96-301-P-0528-0537, 44.8 mm SL, lateral view
of a. neurocranium; b. right lachrymal; c. first gill arch and rakers. Arrow indicates position of the
median coronal pore.
Scales. Flank scales large, ctenoid and regularly imbricating. Pored lateral line scales
34–35. Upper branch of lateral line never overlapping lower branch, which is short, con-
sisting of only 4–6 pored scales. Cheek and chest naked, belly with small scales. Nape
naked to level of 6th dorsal spine. A few scattered scales on the opercle. Dorsal and anal
fins scaleless. Small scales over proximal half of caudal fin.
Vertebrae. Vertebral count 31–32; 14+18 (13), 14+17 (2).
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Additional osteology. Infraorbital series composed of a broad plate-like lachrymal
which is perforated by five large pores of the sensory canal system (Fig. 3b), there are no
tubular infraorbitals adjacent to the lachrymal, and no dermosphenotic is present. Supra-
neural usually absent (Fig. 3a), but in four individuals a very small weakly ossified supra-
neural is present. Neurocranium is somewhat dorsoventrally compressed and the
supraoccipital crest is low, as is the frontal ridge, which extends anteriorly to the median
coronal pore (Fig 3a).
FIGURE 4. Neolamprologus devosi sp. n., MRAC 96-301-P-0528-0537, Male, 55.3 mm SL,
lower pharyngeal jaw in a. dorsal view, b. ventral view, c. caudal view.
Coloration (Fig. 5). In life, base body coloration light beige brown fading to pale yel-
low on the belly and along the dorsum, and with a bright yellow patch under the eye. Four
or five dusky vertical bars are often present along the body from nape to caudal peduncle.
Individual body scales have dark pigment around the free scale margins, creating a reticu-
late pattern of thin, oblique bands of pigment that present the appearance of “chain mail”
or a “chain link fence”. A broad lachrymal stripe more-or-less covers the lachrymal bone,
and a dark horizontal band runs from the eye over the dorsal cheek and merges with a
scaleless opercular spot. The lower lip and opercle are suffused with a pale blue flush.
Dorsal fin with a yellow marginal band and a black submarginal band. The interspinous
and soft ray membranes of the dorsal and anal fins with a strong pattern of yellow banding
and merging spotting. Caudal fin also banded and spotted. Caudal fin with a yellow mar-
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ZOOTAXA ginal band and black submargin; ventral portion of fin has a bluish flush. Preserved color-
ation, beige brown with banding and scale pigmentation usually visible.
Viscera and diet. Gut short and straight, intestinal length 60–70% SL. Contents
included some detritus, freshwater shrimps, ostracods, and the soft-bodies of snails. As no
crushed snail shells were found in the stomach or intestines this perhaps indicates that N.
devosi is “picking” the snails from shells rather than crushing the shells to obtain the flesh.
Reproductive biology. Nothing is currently known of the reproductive biology of this
species.
FIGURE 5. Holotype of Neolamprologus devosi sp. n., AMNH 233614, male, 56.9 mm SL, a few
kilometers up river from Ilagala, lower course of Malagarasi River, 5°11’59”S 29°49’59”E. Live
coloration shortly after capture. The specimen was held for some months in an aquarium prior to
preservation.
Distribution and habitat. Currently N. devosi is known only from two localities in the
Malagarasi River basin: from the delta and in the river a few kilometers upstream from
Ilgala, which itself is situated some kilometers to the east of the Malagarasi delta on the
northern bank of the Malagarasi River (Fig. 6). It is noteworthy that when one of us (L.S.)
collected in the Malagarasi delta (12/1991) at the same locality from where most paratypes
were collected by L. De Vos (8/1993), no specimens of N. devosi were seen. This might be
due to the fact that there is a small rainy season in December-January with relatively high
water stands whereas in July the dry season is ongoing and water levels are low. However,
the possibility that N. devosi undergoes seasonal migration is raised.
Water parameters taken in December 1991 in the delta were: air temperature, 28.5°C,
water temperature, 26.0°C, 7.8 mg/l O2, conductivity, 185 microsiemens/cm-1, pH 8.33,
German hardness, 5° DH. The visibility of the water was very low. These measures differ
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markedly from those taken in December 1991 near Kigoma on Lake Tanganyika where
conductivity was 450 microsiemens/cm-1, pH, 9.17 and German hardness was 12° DH.
The lower conductivity, pH and German hardness clearly indicate the riverine conditions
that prevail in the delta as opposed to the lake proper.
FIGURE 6. Collection localities of Neolamprologus devosi sp. n. in lower Malagarasi River, Tan-
zania.
Etymology. Named in memory of our friend and colleague Luc (Tuur) De Vos who
dedicated so much of his career to expanding our knowledge of the fishes of East and Cen-
tral Africa. His sudden and untimely death is a great loss to our community.
Remarks. De Vos et al. (2001) suggest that the presence of a lamprologine cichlid in
the lower Malagarasi River with close phylogenetic affinities with certain of the Congo-
lese Lamprologus casts doubt on the hypothesis, advanced by Sturmbauer et al. (1994),
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ZOOTAXA that the Congolese riverine lamprologines (or at least some of them) are phylogenetically
nested within the Lake Tanganyika flock and represent a secondary invasion of the Congo
River system. They reason that given such a scenario, it is highly unlikely that one would
find two closely related riverine species both to the west and to the east of the lake. Ongo-
ing studies, however, indicate that the resemblance between Neolamprologus devosi and
Lamprologus mocquardi is superficial, and that no close phylogenetic relationship exists
between them (Schelly & Stiassny, submitted) and the question of the origin/s of the river-
ine Lamprologus radiation remains an outstanding issue.
Acknowlegements
Our thanks to B. Brown, D. Rodriguez, and R. Arrindell (AMNH) for assistance with col-
lections. P. Bwathondi (Tanzania Fisheries Research Institute, TAFIRI, Dar es Salaam),
T.W. Maembe (Fisheries Division, Ministry of Tourism, Natural Resources and Environ-
ment of Tanzania, Dar es Salaam) and A.M. Nikundiwe (Department of Zoology, Univer-
sity of Dar es Salaam, Tanzania) provided and/or supported permissions to collect and
export live and preserved fishes. We thank Jos Snoeks (MRAC) for loan of material. The
Axelrod Research Curatorship provided ongoing support to MLJS. RS is supported by an
AMNH-Columbia University Graduate Fellowship, and L.S. is grateful to COSTECH (the
Tanzania Commission for Science and Technology, Dar es Salaam) for permission to con-
duct research in Tanzania.
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the major substrate spawning lineage of cichild [sic] fishes from Lake Tanganyika in Eastern
Africa. Molecular Biology and Evolution, 11, 691–703.
... Some of the most intriguing of these questions are whether ancestral lamprologines arose in rivers or lakes, and the number of transitions between the two modalities in the history of the group. Molecular analyses that included some of the Congo River species of Lamprologus (e.g., Sturmbauer et al., 1994;Salzburger et al., 2002) apparently support Regan's (1920Regan's ( , 1922 contention that lamprol-3 Additionally, Neolamprologus devosi from the Malagarasi River on the eastern shore of Lake Tanganyika has been newly described (Schelly et al., 2003), bringing the total number of exclusively fluviatile lamprologines in the greater Congo basin to nine. Neolamprologus devosi is not a member of the Congo River Lamprologus clade (Schelly et al., 2003), and is therefore excluded from this treatment. ...
... Molecular analyses that included some of the Congo River species of Lamprologus (e.g., Sturmbauer et al., 1994;Salzburger et al., 2002) apparently support Regan's (1920Regan's ( , 1922 contention that lamprol-3 Additionally, Neolamprologus devosi from the Malagarasi River on the eastern shore of Lake Tanganyika has been newly described (Schelly et al., 2003), bringing the total number of exclusively fluviatile lamprologines in the greater Congo basin to nine. Neolamprologus devosi is not a member of the Congo River Lamprologus clade (Schelly et al., 2003), and is therefore excluded from this treatment. ogines first diversified in Lake Tanganyika and only secondarily invaded the Congo River system. ...
... If the lake is accepted as a barrier to riverine taxa, this would suggest that fluviatile lamprologines predate the formation of the lake, and that lake formation fragmented their range. However, a closer examination of this species (described as Neolamprologus devosi; Schelly et al., 2003), has revealed that any resemblance to L. mocquardi is merely superficial, while its true affinities lie with certain Lake Tanganyika endemics. The relevant characters in Neolamprologus devosi are derived and shared with members of the lacustrine radiation. ...
... members of the tribe Lamprologini, which are also represented in rivers across East and Central Africa [18][19][20], and species of the tribe Haplochromini, which are distributed in rivers and lakes all over Africa, but reach their highest levels of diversity in Lakes Victoria and Malawi (e.g., [18,[21][22][23][24]). ...
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Background: The diversification process known as the Lake Tanganyika Radiation has given rise to the most speciose clade of African cichlids. Almost all cichlid species found in the lakes Tanganyika, Malawi and Victoria, comprising a total of 12-16 tribes, belong to this clade. Strikingly, all the species in the latter two lakes are members of the tribe Haplochromini, whose origin remains unclear. The 'out of Tanganyika' hypothesis argues that the Haplochromini emerged simultaneously with other cichlid tribes and lineages in Lake Tanganyika, presumably about 5-6 million years ago (MYA), and that their presence in the lakes Malawi and Victoria and elsewhere in Africa today is due to later migrations. In contrast, the 'melting pot Tanganyika hypothesis' postulates that Haplochromini emerged in Africa prior to the formation of Lake Tanganyika, and that their divergence could have begun about 17 MYA. Haplochromine fossils could potentially resolve this debate, but such fossils are extremely rare. Results: Here we present a new fossil haplochromine from the upper Miocene site Waril (9-10 million years) in Central Kenya. Comparative morphology, supported by Micro-CT imaging, reveals that it bears a unique combination of characters relating to dentition, cranial bones, caudal skeleton and meristic traits. Its most prominent feature is the presence of exclusively unicuspid teeth, with canines in the outer tooth row. †Warilochromis unicuspidatus gen. et sp. nov. shares this combination of characters solely with members of the Haplochromini and its lacrimal morphology indicates a possible relation to the riverine genus Pseudocrenilabrus. Due to its fang-like dentition and non-fusiform body, †W. unicuspidatus gen. et sp. nov. might have employed either a sit-and-pursue or sit-and-wait hunting strategy, which has not been reported for any other fossil haplochromine cichlid. Conclusions: The age of the fossil (9-10 MYA) is incompatible with the 'out of Tanganyika' hypothesis, which postulates that the divergence of the Haplochromini began only 5-6 MYA. The presence of this fossil in an upper Miocene palaeolake in the Central Kenya Rift, as well as its predatory lifestyle, indicate that Haplochromini were already an important component of freshwater drainages in East Africa at that time.
... Members of the genus Neolamprologus (49 described species, all but one endemic to Lake Tanganyika) are highly diverse, and the genus is probably not monophyletic (Poll, 1986;Schelly et al., 2003;Aibara et al., 2005). Within the genus Neolamprologus, 10 described species are characterised by a lunate tail with long filaments, uncommon among lamprologines. ...
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Two new cichlid species, Neolamprologus walteri sp. nov. and N. chitamwebwai sp. nov., from the Bangwe peninsula (Tanzania), on the east coast of Lake Tanganyika, are described. Both species belong to the N. savoryi species complex, characterised by a lunate tail with long filaments. Neolamprologus chitamwebwai and N. walteri are highly stenotopic species, restricted to rocky habitat. They occur sympatrically with each other and with two other species of the complex but occupy different microhabitats. Neolamprologus walteri lives in large numbers in sheltered areas with rubble substrate whereas N. chitamwebwai occurs in much lower numbers on more exposed parts of the rocky shores, with large boulders, lower sedimentation rates, coarser sediment, and higher visibility. The two new species although closely related, show clear ecological differentiation. Evidence of ecological differentiation between closely related sympatric cichlids is rare in literature. Closely related sympatric cichlids in the large African lakes (with literature mainly referring to Lake Victoria and Lake Malawi) have been suggested to differ little in diet and habitat use, which has encouraged the idea that cichlid species can coexist without niche partitioning. Our paper provides a different perspective with cichlids from Lake Tanganyika.
... The clade of the species-rich tribe Lamprologini Poll, 1986 (more than 90 species belonging to eight genera are currently treated as valid) diverged close to the root of MVhL (Nishida, 1991;Kocher et al., 1995;Salzburger et al., 2002;Clabaut et al., 2005;Day et al., 2008;Meyer et al., 2015;Weiss et al., 2015;McGee et al., 2016). Most Lamprologini species are endemic to Lake Tanganyika, while nine inhabit the Congo River and one the Malagarasi River (Schelly et al., 2003;Schelly and Stiassny, 2004;Tougas and Stiassny, 2014). The Congo River is connected to Lake Tanganyika via Lukuga River, and Malagarasi River flows into the lake. ...
Article
The huge monophyletic group of the East African cichlid radiations (EAR) consists of thousands of species belonging to 12-14 tribes; the number of tribes differs among studies. Many studies have inferred phylogenies of EAR tribes using various genetic markers. However, these phylogenies partly contradict one another and can have weak statistic support. In this study, we conducted maximum-likelihood (ML) phylogenetic analyses using restriction site-associated DNA (RAD) sequences and propose a new robust phylogenetic hypothesis among Lake Tanganyika cichlid fishes, which cover most EAR tribes. Data matrices can vary in size and contents depending on the strategies used to process RAD sequences. Therefore, we prepared 23 data matrices with various processing strategies. The ML phylogenies inferred from 15 large matrices (2.0×10(6) to 1.1×10(7) base pairs) resolved every tribe as a monophyletic group with 100% bootstrap support and shared the same topology regarding relationships among the tribes. Most nodes among the tribes were supported by 100% bootstrap values, and the bootstrap support for the other node varied among the 15 ML trees from 70% to 100%. These robust ML trees differ partly in topology from those in earlier studies, and these phylogenetic relationships have important implications for the tribal classification of EAR.
... The remainder, about 15 % of this ichthyofauna, is composed of Malagarazi endemics. These include nine undescribed species reported by De Vos et al. (2001) i. e. two small 'Barbus', a first one described below and a second one similar to 'B.' quadrilineatus and 'B.' lineomaculatus, the status of which still needs further confirmation and is currently under study; one Labeobarbus; one Leptoglanis; one Chiloglanis; two Mastacembelus; one 'Haplochromis'; and one Lamprologus species subsequently described as Neolamprologus devosi by Schelly et al. (2003). David (1937), Poll (1946) and Greenwood (1962) reported the presence of 'B.' eutaenia in the Malagarazi River basin based on 22 specimens (MRAC 47257-47278: three of which are now BMNH 1962.7.26:77-79) collected by A. Lestrade in the "rivière Malagarazi et ses affluents en territoire de Bururi" (Burundi). ...
Article
'Barbus' devosi, new species, is described from the Malagarazi River drainage, the main affluent of Lake Tanganyika. It belongs to the group of small African 'Barbus' with a strongly ossified and serrated last unbranched dorsal-fin ray and a blackish mid-lateral band, that extends from the tip of the snout to the caudal-fin base, or onto the mid-central part of the caudal fin. 'Barbus' devosi resembles 'B.' eutaenia and 'B.' niiolepis, to which it has formerly been erroneously attributed. It can be distinguished in having a higher number of gill rakers on the ceratobranchial of the first gill arch (6-9 vs. 2-5), shorter anterior (11-19 vs. 20-35% HL for 'B.' eutaenia and 2037% HL for 'B.' miolepis) and posterior (15-27 vs. 22-44% HL for 'B.' eutaenia and 27-46% HL for 'B/ miolepis) barbels, and the absence of a sheath of enlarged, prominent, scales at the base of the dorsal fin.
... The cichlid tribe Lamprologini forms a major component of the endemic fish fauna of Lake Tanganyika, with about 80 species distributed in seven (Poll, 1986), eight, or possibly nine (Stiassny, 1997) genera. An additional eight species are known only from the Congo River, and one spe-cies only from the Malagarasi River (Schelly & Stiassny, 2004;Schelly et al., 2003). Several of the endemic lacustrine lamprologin species are known from very few specimens and/or few localities. ...
Article
Neolamprologus timidus, new species, is described from Ulwile Island and adjacent localities on the Tanzanian coast of Lake Tanganyika. The species was observed or collected along about 100 km of coastline from Kolwe Point, Cape Mpimbwe, south to Kisi Island. It is distinguished from the most similar species, N. furcifer, by presence of scales on most of the cheek, long pectoral fin and pelvic fin with the second ray longer than the first. Neolamprologus timidus is sympatric with N. furcifer at Kolwe Point and south to Kampempa Point, and at Lupita and Ulwile Islands south to Kisi Island, but N. furcifer is otherwise absent from the range of N. timidus. Two morphologically distinct forms are recognized in N. furcifer. Samples of N. furcifer from Ulwile Island and slightly more southern localities possess a caudal fin with rounded lobes and long middle rays, appearing only slightly emarginate. Samples of N. furcifer from Udachi and nearby localities possess pointed caudal-fin lobes with greatly elongated streamers, similar to N. timidus and to N. furcifer from other parts of Lake Tanganyika, including the type specimens from the southern part of the lake. The variation in caudal-fin shape may be an expression of character displacement as it occurs in the area of sympatry between N. timidus and N. furcifer. Mitochondrial DNA sequences are nearly identical in samples of N. furcifer with pointed or rounded caudal fin. A molecular phylogenetic analysis of a large set of lamprologin cichlids using two mitochondrial genes corroborates earlier analyses and places N. furcifer and N. timidus in different clades with different species of Neolamprologus, Julidochromis, Chalinochromis, and Telmatochromis despite sharing a unique combination of fin and body shape, and colour pattern. A 4648 base-pair multiloci analysis of a smaller number of species using fragments of three mitochondrial and two nuclear genes resolves N. furcifer and N. timidus in sister clades, but the N. timidus clade also includes Telmatochromis brachygnathus, and N. furcifer is sister species of Chalinochromis brichardi.
... One sample of Orthochromis stormsi (NRM 45081) was obtained from near Nyunzu far downstream of the mouth of the Niemba River, and this species is widespread in the Congo River basin (Greenwood & Kullander, 1994). The Malagarasi River also has a species, Neolamprologus devosi, which belongs to the lake species flock, but has not been found outside the lower Malagarasi River (Schelly et al., 2003). It then would be a case of speciation outside the lake like the Tanganicodus species at Niemba ...
Article
The Lukuga River is a large permanent river intermittently serving as the only effluent of Lake Tanganyika. For at least the first one hundred km its water is almost pure lake water. Seventy-seven species of fish were collected from six localities along the Lukuga River. Species of cichlids, cyprinids, and clupeids otherwise known only from Lake Tanganyika were identified from rapids in the Lukuga River at Niemba, 100 km from the lake, whereas downstream localities represent a Congo River fish fauna. Cichlid species from Niemba include special-ized algal browsers that also occur in the lake (Simochromis babaulti, S. diagramma) and one invertebrate picker representing a new species of a genus (Tanganicodus) otherwise only known from the lake. Other fish species from Niemba include an abundant species of clupeid, Stolothrissa tanganicae, otherwise only known from Lake Tangan-yika that has a pelagic mode of life in the lake. These species demonstrate that their adaptations are not neces-sarily dependent upon the lake habitat. Other endemic taxa occurring at Niemba are known to frequent vegetat-ed shore habitats or river mouths similar to the conditions at the entrance of the Lukuga, viz. Chelaethiops minutus (Cyprinidae), Lates mariae (Latidae), Mastacembelus cunningtoni (Mastacembelidae), Astatotilapia burtoni, Ctenochromis horei, Telmatochromis dhonti, and Tylochromis polylepis (Cichlidae). The Lukuga frequently did not serve as an ef-fluent due to weed masses and sand bars building up at the exit, and low water levels of Lake Tanganyika. Since the Lukuga River does not have a permanent connection with Lake Tanganyika we hypothesize that its lake fishes are not temporary occurrences due to occasional spills from the lake. Apparently the Lukuga maintains a stable fish community that is independent of the lake, and may depend more on the flow of the major tributary, the left bank tributary Niemba River, than on outflow from Lake Tanganyika.
... If the lake is accepted as a barrier to riverine taxa, this would suggest that fluviatile lamprologines predate the formation of the lake, and that lake formation fragmented their range. However, a closer examination of this species (described as Neolamprologus devosi; Schelly et al., 2003), has revealed that any resemblance to L. mocquardi is merely superficial , while its true affinities lie with certain Lake Tanganyika endemics. The relevant characters in Neolamprologus devosi are derived and shared with members of the lacustrine radiation. ...
Article
The Congo River Lamprologus are revised and two new species are described. Lamprologus teugelsi, n.sp., from Malebo Pool and the lower Congo River rapids, and L. tigripictilis, n.sp., from the lower Congo River rapids, are readily distinguished from the remaining Congo River Lamprologus based on counts, measurements, osteology, and color pattern. Monophyly of the Congo River Lamprologus species is tentatively accepted and a key to the group is provided. All available collection localities for re-identified Lamprologus material from the Congo River are plotted for each species. Maps of collection localities reveal large distributional voids, suggesting that Congo River lamprologine diversity remains incompletely sampled.
Article
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The species-flocks of cichlid fishes in the East African Great Lakes Victoria, Malawi and Tanganyika constitute the most diverse extant adaptive radiations in vertebrates. Lake Tanganyika, the oldest of the lakes, harbors the morphologically and genetically most diverse assemblage of cichlids and contains the highest number of endemic cichlid genera of all African lakes. Based on morphological grounds, the Tanganyikan cichlid species have been grouped into 12 to 16 distinct lineages, so-called tribes. While the monophyly of most of the tribes is well established, the phylogenetic relationships among the tribes remain largely elusive. Here, we present a new tribal level phylogenetic hypothesis for the cichlid fishes of Lake Tanganyika that is based on the so far largest set of nuclear markers and a total alignment length of close to 18 kb. Using next-generation amplicon sequencing with the 454 pyrosequencing technology, we compiled a dataset consisting of 42 nuclear loci in 45 East African cichlid species, which we subjected to maximum likelihood and Bayesian inference phylogenetic analyses. We analyzed the entire concatenated dataset and each marker individually, and performed a Bayesian concordance analysis and gene tree discordance tests. Overall, we find strong support for a position of the Oreochromini, Boulengerochromini, Bathybatini and Trematocarini outside of a clade combining the substrate spawning Lamprologini and the mouthbrooding tribes of the ‘H-lineage’, which are both strongly supported to be monophyletic. The Eretmodini are firmly placed within the ‘H-lineage’, as sister-group to the most species-rich tribe of cichlids, the Haplochromini. The phylogenetic relationships at the base of the ‘H-lineage’ received less support, which is likely due to high speciation rates in the early phase of the radiation. Discordance among gene trees and marker sets further suggests the occurrence of past hybridization and/or incomplete lineage sorting in the cichlid fishes of Lake Tanganyika.
Article
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The phylogenetic relationship of the speciose and ecologically diverse lamprologine cichlids of Africa are investigated utilising a range of anatomical data. The monophyly of the group is firmly established and the basal position of Variabilichromis moorii, an hypothesis originally inferred from molecular data is supported. Within the lamprologine clade a large monophyletic subclade, the so-called 'ossified group', is recognised. As membership within the 'ossified group' includes species currently arrayed among three lamprologine 'genera', the present study poses a challenge to current taxonomic convention and highlights the need for continued study. The relationship of the lamprologine clade to other African cichlid lineages is investigated and while anatomical evidence is far from compelling, a case is made for a sistergroup relationship with the Zairean genus, Teleogramma and a more distal relationship to the west African / Zairean genus, Steatocranus, is tentatively posited.
Article
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Lake Tanganyika harbors the oldest, morphologically and behaviorally most diverse flock of cichlid species. While the cichlids in Lakes Malawi and Victoria breed their eggs exclusively by buccal incubation (termed "mouthbrooding"), the Tanganyikan cichlid fauna comprise mouthbrooding and substrate-spawning lineages (fish spawn on rocks, and never orally incubate eggs or wrigglers). The substrate-spawning tribe Lamprologini appears to occupy a key position that might allow one to elucidate the origin of the Tanganyika flock, because five riverine (therefore nonendemic) species from the Zaire River system have been assigned to this tribe, in addition to the lake's endemic species, which make up almost 50% of all 171 species known from this lake (Poll 1986). From 16 species (18 individuals) of the tribe Lamprologini, a 402-bp segment of the mitochondrial cytochrome b gene was sequenced, and, from 25 lamprologine species (35 individuals), sequences from the mitochondrial control region were obtained. To place the Lamprologini into a larger phylogenetic framework, orthologous sequences were obtained from eight nonlamprologine Tanganyikan cichlid species (13 individuals). The Lamprologini are monophyletic, and a clade of six Tanganyikan lineages of mouthbrooders, representing five tribes (Poll 1986), appears to be their sister group. Comparisons of sequence divergences of the control region indicate that the Lamprologini may be older than the endemic Tanganyikan tribe Ectodini, and short basal branches might suggest a rapid formation of lineages at an early stage of the Tanganyika radiation. It is interesting that three analyzed riverine members of the tribe form a monophyletic group; however, they are not the most ancestral branch of the Lamprologini. This might indicate that they are derived from an endemic lamprologine ancestor that left Lake Tanganyika by entering the Zaire River system. These riverine species may not have seeded the Tanganyikan radiation, as currently thought, but may have recently recolonized the river after a long period of isolation, as soon as the lake was connected to the Zaire River again about 2 Mya. Neolamprologus moorii, endemic to Lake Tanganyika, appears to represent the most basal clade of the Lamprologini. Complex breeding behavior, involving the usage of gastropod shells and associated with dwarfism, is likely to have evolved in parallel in several lineages among the Lamprologini. The tribe Lamprologini may be in need of revision, since several genera appear to be polyphyletic.
Article
The haplochromine Cichlidae endemic to Lake Victoria are recently estimated to comprise approximately 200 species. Morphologically these species vary within narrow limits but ecologically they demonstrate a wide and continuous spectrum of adaptations. Except for the papyrus swamps, the fish-fauna of each habitat is dominated by a particular community of haplochromine species. Besides restrictions to certain habitats most species have limited food-regimes. Together these diets comprise all food categories available (possibly with exception of zooplankton). Morphologically similar species realizing such a wide scala of adaptations are ideal objects for comparative biological research. For this reason haplochromine cichlids have been recently introduced into ethology, ontogeny, physiology, functional morphology and-naturally-ecology. Another recent interest in these fishes concerns their potential economic importance: the overwhelming quantity with which haplochromine cichlids (still) occur, together with an improved fishing gear, make these relatively small fishes a source for fishmeal and human consumption. Regrettably no ecological survey preceded the planning for these extensive fisheries. A major cause for this omission are the difficulties experienced in identifying haplochromine species. These difficulties result partly from the overall morphological similarity and partly from the fact that only half the estimated number of species has been described so far. The importance of identification for both fishery-biology, ecology and other biological research has been the main reason for writing this paper which introduces the techniques applied in the taxonomy of the haplochromine Cichlidae from Lake Victoria. As this taxonomy is mainly based on morphological characters, the paper consequently provides basic information on the anatomy and external aspects of these cichlids. The subtitle refers to the fact that all currently valid species have been described (or revised) by GREENWOOD in a long series of papers mainly intended for the use of taxonomists. The present introduction has been explicitly written for students unfamiliar with the morphology and taxonomy of haplochromine fishes. After a short review of the problems related to haplochromine taxonomy, there
Article
Preparation of small vertebrates cleared after alcian blue staining of cartilage is facilitated by trypsin digestion. Specimens are fixed in formation, washed, skinned, and eviscerated. After staining in a solution of alcian blue in acetic acid-alcohol for 24-48 hours, they are transferred to water through graded alcohols. Excess alcian blue is removed over a period of up to three weeks by changes every 2-3 days of 1% trypsin in approximately one-third-saturated sodium borate. Bony tissues may be stained after this in a solution of alizarin red S in 0.5% KOH. Specimens are bleached if necessary and dehydrated through graded KOH-glycerine mixtures for storage in glycerine. Since alcohol treatment in addition to formalin fixation does not affect results with this method, it should be useful to researchers who want to study the cartilage or cartilaginous skeletons in museum specimens, which are routinely fixed in formalin and stored in alcohol.
Ein neuer Cichlide vom Ostufer des Tanganjikasees: Lamprologus leleupi longior n. ssp. (Pisces, Cichlidae). Revue de zoologie africaine
  • W Staeck
Staeck, W. (1980) Ein neuer Cichlide vom Ostufer des Tanganjikasees: Lamprologus leleupi longior n. ssp. (Pisces, Cichlidae). Revue de zoologie africaine, 94, 11-14.
Ein neuer Cichlide von der zairischen Küste des Tanganyikasees: Neolamprologus variostigma n. sp. Die Aquarien-und Terrarien Zeitschrift
  • H H Büscher
Büscher, H.H. (1995b) Ein neuer Cichlide von der zairischen Küste des Tanganyikasees: Neolamprologus variostigma n. sp. Die Aquarien-und Terrarien Zeitschrift (DATZ), 48, 794–797.