Accepted: 24 November 2003; published: 1 December 2003 1
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2003 Magnolia Press
Zootaxa 373: 1–11 (2003)
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: firstname.lastname@example.org and email@example.com
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
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
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.
2 © 2003 Magnolia Press
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,
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
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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A NEW NEOLAMPROLOGUS
double stained), delta Malagarasi, right arm, Tanzania, 5°13’S 29°48’E, L. De Vos, 20/08/
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.
4 © 2003 Magnolia Press
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
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
Caudal peduncle depth
Caudal peduncle length
Anal fin base length
Dorsal fin base length
Pelvic fin length
Caudal fin length
Pectoral fin length
PERCENT OF HEAD LENGTH
Lower jaw length
Lateral line scales
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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A NEW NEOLAMPROLOGUS
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-
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–
SCHELLY ET AL.
6 © 2003 Magnolia Press
ZOOTAXA 6 (mode 5) along the ceratobranchial, one raker in the angle of the arch, and 2–4 (mode 3)
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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A NEW NEOLAMPROLOGUS
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-
SCHELLY ET AL.
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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
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
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
© 2003 Magnolia Press 9
A NEW NEOLAMPROLOGUS
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-
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),
SCHELLY ET AL.
10 © 2003 Magnolia Press
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.
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.
Barel, C.D.N., van Oijen, M.J.P., Witte, F. & Witte-Maas, E.L.M. (1977) An introduction to the tax-
onomy and morphology of the haplochromine Cichlidae from Lake Victoria. A manual to
Greenwood’s revision papers. Netherlands Journal of Zoology, 27, 333–389.
Boulenger, G.A. (1915) Catalogue of the Freshwater Fishes of Africa in the British Museum (Natu-
ral History). Vol. III. British Museum (Natural History), London, 526 pp.
Büscher, H.H. (1989) Ein neuer Tanganjika-Cichlide aus Zaire Neolamprologus marunguensis n.
sp. (Cichlidae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 42, 739–743.
Büscher, H.H. (1991) Ein neuer Tanganjikasee-Cichlide aus Zaire Neolamprologus pectoralis n. sp.
(Cichlidae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 44, 788–792.
Büscher, H.H. (1992a) Ein neuer Cichlide aus dem Tanganjikasee Neolamprologus similis n. sp.
Die Aquarien- und Terrarien Zeitschrift (DATZ), 45, 520–525.
Büscher, H.H. (1992b) Neolamprologus nigriventris n. sp.: ein neuer Tanganjikasee-Cichlide
(Cichlidae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 45, 778–783.
Büscher, H.H. (1993) Neolamprologus bifasciatus n. sp.: ein neuer Tanganjikasee-Cichlide (Cichl-
idae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 46, 385–389.
Büscher, H.H. (1995a) Ein neuer Cichlide aus dem Tanganjikasee Neolamprologus ventralis n. sp.
(Cichlidae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 48, 379–382.
© 2003 Magnolia Press 11
A NEW NEOLAMPROLOGUS
Büscher, H.H. (1995b) Ein neuer Cichlide von der zairischen Küste des Tanganyikasees: Neolam-
prologus variostigma n. sp. Die Aquarien- und Terrarien Zeitschrift (DATZ), 48, 794–797.
Büscher, H.H. (1997) Ein neuer Cichlide aus dem Tanganjikasee: Neolamprologus helianthus
(Cichlidae, Lamprologini). Die Aquarien- und Terrarien Zeitschrift (DATZ), 50, 701–706.
Colombe, J. & Allgayer, R. (1985) Description de Variabilichromis, Neolamprologus et Paleolam-
prologus genres nouveaux du Lac Tanganyika, avec redescription des genres Lamprologus
Schilthuis, 1891 et Lepidiolamprologus Pellegrin, 1904. Revue Française Cichlidophiles,
De Vos, L., Seegers, L., Taverne, L. & Thys van den Audenaerde, D. (2001) L’ichtyofaune du
bassin de la Malagarasi (Système du Lac Tanganyika): une synthèse de la connaissance actu-
elle. Annales Musée Royal de l’Afrique Centrale, Sciences Zoologiques, 285, 117–135.
Dingerkus, G. & Uhler, L.D. (1977) Enzyme clearing of alcian blue stained whole small vertebrates
for demonstration of cartilage. Stain Technology, 52, 229–232.
Konings, A. (1998) Tanganyika cichlids in their natural habitat. Cichlid Press, El Paso, Texas, 272
Poll, M. (1956) Poissons Cichlidae. Résultats scientifiques de l’exploration hydrobiologique du Lac
Tanganika (1946–1947). Institut Royal des sciences naturelles de Belgique, 3, (5B), 1–619.
Poll, M. (1978) Contribution à la connaissance du genre Lamprologus SCHTH. Description de qua-
tre espèces nouvelles, réhabilitation de Lamprologus mondabu et synopsis remanié des espèces
du lac Tanganika. Bulletin de la Classe des sciences, Académie royale de Belgique, (5 série),
Poll, M. (1986) Classification des Cichlidae du lac Tanganika. Tribus, genres et espèces. Académie
Royale de Belgique Mémoires de la Classe des Sciences, (2 série), 45, 1–163.
Staeck, W. (1980) Ein neuer Cichlide vom Ostufer des Tanganjikasees: Lamprologus leleupi lon-
gior n. ssp. (Pisces, Cichlidae). Revue de zoologie africaine, 94, 11–14.
Staeck, W. & Seegers, L. (1986) Neolamprologus cylindricus spec. nov. Beschreibung eines neuen
Tanganjikacichliden aus dem südlichen Tansania und dem östlichen Sambia. Die Aquarien-
und Terrarien Zeitschrift (DATZ), 39, 448–451.
Stiassny, M.L.J. (1997) A phylogenetic overview of the lamprologine cichlids of Africa (Teleostei,
Cichlidae): a morphological perspective. South African Journal of Science, 93, 513–523.
Sturmbauer, C., Verheyen, E. & Meyer, A. (1994) Mitochondrial phylogeny of the Lamprologini,
the major substrate spawning lineage of cichild [sic] fishes from Lake Tanganyika in Eastern
Africa. Molecular Biology and Evolution, 11, 691–703.