Accepted by J. Friel: 8 Jul. 2009; published: 31 Jul 2009
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 2173: 41–48 (2009)
A new dwarf cichlid genus and species (Teleostei, Cichlidae)
from Guinea, West Africa
University of Vienna, Department of Evolutionary Biology, Althanstrasse 14, A-1090 Vienna, Austria.
Enigmatochromis lucanusi, a new cichlid genus and species, is described from Guinea (West Africa). It is a member of
the chromidotilapiine cichlid clade, and differs from other genera within the group in a combination of morphological
characters and coloration patterns; e.g., twelve circumpeduncular scales; three tubular infraorbital bones in addition to
the lachrymal bone; upper lateral-line separated from the dorsal-fin base; first ray of pelvic fin in adult females of equal
length or longer than second ray; sexual dimorphism well developed; small juveniles with 3–4 rows of irregular dark dots
on body; and breeding coloration of females.
Key words: Enigmatochromis, chromidotilapiine
Enigmatochromis lucanusi, un nouveau genre et espèce de cichlidé, est décrit de Guinée (Afrique de l’Ouest). C’est un
membre du clade des cichlidés chromidotilapiines, et il diffère des autres genres du groupe par une combinaison de
plusieurs caractères morphologiques et patrons de coloration; e.g., 12 écailles autour du pédoncule caudal; trois os
infraorbitaires tubulaires en plus de l’os lacrymal; ligne latérale supérieure séparée de la base de la nageoire dorsale;
premier rayon de la nageoire pelvienne des femelles adultes de longueur égale ou plus long que le deuxième rayon;
dimorphisme sexuel bien développé; juvéniles de petite taille avec 3–4 rangées de marques noires irrégulières sur le
corps; et livrée nuptiale des femelles.
In 2004, a small cichlid species from Guinea was introduced into the aquarium hobby by several commercial
importers in Canada, the U.S.A. and Europe. This new cichlid clearly was a member of the chromidotilapiine
cichlid assemblage sensu Greenwood (1987) as it possessed the characteristic combination of features of this
group: (1) a typical, visor-like and well circumscribed projection (“hanging pad”) of the pharyngeal tissues
situated anterior to the upper pharyngeal bones of each side; (2) no microbranchiospines; (3) outer-row jaw
teeth unicuspid; (4) some anterolaterally positioned teeth in the outer tooth row of the lower jaw, curved
directly posteriorly and not buccally as they are in all other tooth rows; (5) tuberculate gill rakers on the first
ceratobranchial. Females possessed coloration patterns unknown from any other species within this
assemblage, e.g., a bright blue dorsal-fin in females. The species was initially called Pelvicachromis sp. “blue
fin”, and was thought to be closely related to P. roloffi. Further morphological and behavioral observations
however revealed differences from the genus Pelvicachromis.
In 2006, Canadian importer and aquarist Oliver Lucanus collected specimens of this species in Guinea at a
single locality near the village of Fria, and provided the author with a detailed description of the habitat (pers.
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comm.). It is the aim of this paper to describe this new genus and species based on several collections all from
Material and methods
External counts and measurements follow Barel et al. (1977). All measurements were taken on the left side of
the specimens with digital calipers with an accuracy of +/- 0.1 mm.
Radiographs were produced for the AMNH material for vertebrae counts. The clearing and staining
protocol to examine bones and cartilage followed Dingerkus & Uhler (1977). Definitions of genera described
before 1987 follow all diagnoses as given in Greenwood (1987). Description of coloration was based on both
wild caught and tank raised specimens. Abbreviations used include: AMNH, American Museum of Natural
History, New York; MNHN, Muséum National d´Histoire Naturelle, Paris; MRAC, Musée Royal de l´Afrique
Centrale, Tervuren; NMW, Naturhistorisches Museum, Wien; ZSM, Zoologische Staatssammlung, München;
SL, standard length; HL, head length; and undet., undetermined gender.
FIGURE 1. Enigmatochromis lucanusi, NMW 95063, holotype, male, 44.7 mm SL; Guinea: Small creek near the
village of Fria, north of Conakry. Scale bar = 10 mm.
Enigmatochromis new genus
(Figs. 1 & 2.; Table 1)
Type species. Enigmatochromis lucanusi, new species by original designation.
Diagnosis. The new genus is distinguished from all other chromidotilapiine genera by a unique combination
of characters. It posseses twelve scales around the caudal peduncle vs. (13 or 14 scales) in Limbochromis
Greenwood 1987, (14–16 scales) in Chromidotilapia, Boulenger 1898, and (16 scales) in Benitochromis
Lamboj 2001, Pelvicachromis Thys van den Audenaerde 1968 and Thysochromis Daget 1988. Among the
remaining chromidotilapiine genera with twelve circumpenducular scales this new genus is further
distinguished from Congochromis Stiassny & Schliewen 2007 and Nanochromis Pellegrin 1904 by: an
infraorbital series containing a lachrymal and three additional tubular bones, and a gap between the 2
tubular infraorbitals (vs. lachrymal and one tubular bone), plus the lateral line is clearly separated from the
dorsal-fin base (vs. posterior part contiguous with the dorsal-fin base). It is distinguished from Divandu
Lamboj & Snoeks 2000 by: an infraorbital series with a lachrymal and three additional tubular bones and a
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NEW DWARF CICHLID FROM GUINEA, WEST AFRICA
gap between the 2
tubular infraorbitals (vs. four tubular bones), only four openings of the
laterosensory system in the lachrymal bone (vs. five), the first ray of pelvic fin in adult females is of equal
length or longer than second ray of this fin (vs. first ray always longer), being a pair-bonding cave breeder (vs.
a mouthbrooder), and by well developed sexual dichromatism (vs. weakly developed). Finally it is
distinguished from Parananochromis Greenwood 1987 by: an infraorbital series with a lachrymal and three
additional tubular bones with a gap between the 2
tubular infraorbitals (vs. four tubular bones in some
species of Parananochromis), juveniles with 3 or 4 rows of irregular dark brown to black dots on body (vs.
maximum of 2 rows), and the first ray of pelvic fin in adult females of equal length or longer than the second
ray of this fin (vs. second ray slightly longer or of equal length).
Etymology. Enigma from enigmatic, refers to the somewhat intermediate characters between
Pelvicachromis and Parananochromis in possessing pigmentation similarities to the first, but anatomical
similarities to the second genus; and chromis – a common ending for African cichlid fishes. Gender
FIGURE 2. Photographs showing coloration of Enigmatochromis lucanusi, nov. sp.: A) male, wild collected specimen
in aquaria, ~40 mm SL, not preserved, displaying submissive coloration; B) male, wild collected specimen in aquaria,
~42 mm SL, not preserved, displaying aggressive coloration; C) female, wild collected specimen in aquaria, ~38 mm SL,
not preserved, displaying submissive coloration; D) female, wild collected specimen in aquaria, ~39 mm SL, not
preserved, displaying aggressive coloration; E) female, wild collected specimen in aquaria, ~38 mm SL, with juveniles at
the age of approx. one week after swimming free and size of less than 12 mm TL, not preserved; F) aquarium bred
juveniles with the age of app. one week after swimming free, size less than 12 mm TL, not preserved.
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TABLE 1. Morphometrics and meristics of the holotype and 25 syntopic paratypes of Enigmatochromis lucanusi.
Holotype Mean SD Range
SL 44.7 38.2 30.8–45.3
% of SL
Body depth 35.5 32.4 2.9 27.2–36.6
Head length 34.21 33.3 0.9 31.9–35.2
Caudal-peduncle length 12.5 11.8 0.9 9.5–13.7
Caudal-peduncle depth 16.8 16.5 0.9 14.6–18.2
Length of dorsal-fin base 61.9 60.8 1.9 56.9–66.3
Length of anal-fin base 21.1 19.5 1.3 17.3–22.3
Predorsal distance 31.0 28.8 0.9 27.4–31.0
Preanal distance 69.7 70.0 1.8 66.6–73.7
Prepectoral distance 33.6 34.5 1.6 30.9–38.4
Prepelvic distance 37.3 37.1 1.7 32.4–39.9
% of HL
Head depth 62.2 64.8 3.4 57.0–71.3
Snout length 30.5 27.2 2.9 21.3–33.1
Eye diameter 29.5 32.1 1.7 29.2–34.9
Postorbital distance 40.1 40.7 1.7 37.7–43.8
Interorbital distance 27.8 26.6 1.3 24.3–29.9
Cheek depth 31.9 30.8 2.8 25.8–35.7
Lower jaw length 33.7 35.2 3.1 29.5–40.9
Preorbital distance 18.7 20.2 2.2 17.4–24.0
% of caudal peduncle depth
Caudal peduncle length 74.4 71.7 7.0 58.7–86.3
Meristics Median Range
Upper lateral-line scales 17 18 16–20
Lower lateral-line scales 7 6 5–8
Total lateral-line scales 26 26 24–28
Circumpeduncular scales 12 12
Dorsal-fin spines 17 18 17 or 18
Dorsal-fin rays 8 7 7–9
Anal-fin spines 3 3 3
Anal-fin rays 6 7 6 or 7
Pectoral-fin rays 12 11 11 or 12
Gill rakers on lower limb of first arch 8 9 8–12
Total gill rakers on first arch 15 15 14–17
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NEW DWARF CICHLID FROM GUINEA, WEST AFRICA
Enigmatochromis lucanusi, new species
(Figs. 1 & 2; Table 1)
Pelvicachromis sp. “Blue Fin” (Lamboj 2005)
Holotype. NMW 95063, male, 44.7 mm SL; Guinea: Foto River near the town of Fria, north of Conakry,
Konkoure River system. 10° 20.074’ N, 13° 43.431’ W, collected by W. M. Traore, November 2007.
Paratypes. NMW 95064, 2 males, 1 female, 32.9–45.3 mm SL, same collection data as holotype. NMW
95065, 2 males, 2 females, 1 undet., 32.8–40.6 mm SL, same locality as holotype, but collected in January
2004; 1 female, 1 undet. used for C&S after taking counts and measurements. NMW 95066, 5 males, 4
females, 32.8–44.3 mm SL, same locality as holotype but collected in February 2006, 2 males and 2 females
used for C&S after taking counts and measurements. AMNH 238830, 2 males, 2 females 30.8–44.4 mm SL,
same locality as holotype, but collected in January 2004. MRAC 2008-015-P-1-4, 2 males, 2 females,
33.1–43.9 mm SL, same locality as holotype but collected in January 2008.
None-types. In addition, 8 pairs of wild collected fishes and their F1-descendents were used for
behavioural observations, but not included in morphological examinations or type series or other collections.
Diagnosis. Same as for genus.
Description. Measurements and meristic counts for holotype and 25 syntopic paratypes are presented in
Small cichlid species with body moderately gracile, dorsal head profile smoothly rounded and short snout.
Ventral body wall gently rounded to caudal peduncle. Caudal fin rounded. Caudal peduncle short, always
deeper than long. Sexual dimorphism well-developed with males usually 10–15% larger than females, with
soft dorsal and anal fin rays more elongated. First ray of pelvic fin always longest in males, in females of
equal length or longer than second ray. In males, tips of pelvic fin reaching origin of anal fin.
Osteology and dentition. Infraorbital bones series with lachrymal and three additional tubular bones and
a gap between 2
tubular infraorbital; lachrymal with four openings of laterosensory system. 23–25
total vertebrae (12–14 abdominal and 11 or 12 caudal).
Premaxilla with one or two, dentary with one to three rows of regularly set unicuspid teeth. Anteriorly in
the lower jaw a few teeth oriented posteriorly, not buccally. Lower pharyngeal bone triangular, with unicuspid
to weakly bicuspid teeth on lateral parts of this bone, and asymmetric bicuspid teeth in the central field.
Gill rakers on first gill arch. Eight to twelve tuberculate gill rakers on ceratobranchials, 4–7 pointed gill rakers
on epibranchials. Well-developed hanging pad on roof of the pharynx.
Scales. Cycloid, one or two rows of scales on cheek; four horizontal rows on opercle. Dark spot on outer
edge of opercle unscaled. Chest-scales smaller than body scales, four or five scales between pectoral and
pelvic fins. Upper lateral-line separated from dorsal-fin base anteriorly by three scales, at the 8
by one or one and a half scales, and at last pored scale by no or a half scale. End of upper lateral-line never
overlapping lower lateral-line, and separated from beginning of lower lateral line by one to four rows of
scales. About ¼ of caudal fin covered with scales, and all other fins unscaled.
Coloration. Live specimens of both sexes (Fig. 2): Head and body pale brown to greyish brown. Dorsum
somewhat darker than ventral parts of body. Dark scaleless spot on the outer edge of opercle, with a light blue
margin on posterior edge. Upper lip brownish to yellow-brown, lower lip greyish to brown. Throat and ventral
parts of branchiostegal membranes yellow, more prominent in dominant specimens. Dark longitudinal stripe
sometimes visible on sides (in stressed, submissive, breeding and guarding individuals), at about height of
lower lateral-line, from posterior edge of eye to end of caudal peduncle, but not extending onto caudal fin.
Dark stripe from anterior edge of the eye to the middle of upper lip. Upper edge of eye golden-yellow.
Lachrymal bluish, with a red to reddish horizontal stripe from angle of the mouth to anterior edge of
preopercular. Pectoral fin clear.
Male specific coloration. Dorsal fin pale yellow, with red margin, followed by a thin white submargin,
morer prominent in anterior portion of this fin. Upper half of caudal fin yellow, lower half reddish. Soft ray
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parts of dorsal fin and upper half of caudal fin with some to numerous black dots in individual variation. Base
of anal fin yellowish, rest of this fin reddish to violet. Anterior edge of pelvic fins yellow, first soft ray dark
grey to black, other parts of this fin pale reddish to pale violet. Body scales with pale greyish margins. Most
ventral parts of opercle, suborpercle, interopercle and chest are yellow.
Female specific coloration. Spiny portion of dorsal fin iridescent blue, sometimes extending to first one
to two soft rays, with a red margin, followed by a thin dark to black submargin. Rest of fin clear. One or two
black dots in most posterior part of iridescent blue coloration in many individuals. Caudal fin clear to very
pale reddish, without patterns or marks. Anal-fin base and most anterior parts bluish, rest of the fin clear to
pale reddish. Anterior edge of pelvic fins black, as first soft ray dark, other parts of this fin reddish with thin
blue vertical lines in some specimens. Lower parts of opercle, subopercle and interopercle are blue, also most
anterior parts of chest. Flanks and belly region, up to about upper lateral line or higher in some individuals, are
Juveniles of both sexes (before getting adult coloration) exhibit a pattern of three or four rows of irregular
dark spots on brown coloration, up to about 10–12 mm SL. With increasing size, sex-specific coloration is
Preserved specimen coloration General coloration of both sexes: Head and body brown, darker dorsally.
Cheek, throat, pre-pelvic and pre-pectoral regions of flanks and chest pale light brown. Lips greyish. Dark
spot on outer edge of opercle. A dark longitudinal stripe visible in some individuals, reaching from posterior
edge of eye to end of caudal peduncle, but not extending into the caudal fin. Dark bar from anterior edge of
eye to angle of mouth.
Male specific coloration. The general preserved coloration of adult males is shown in the holotype (Fig.
1). Anterior portion of dorsal fin greyish. Posterior portion of dorsal fin and upper parts of caudal fin
yellowish to greyish, with numerous dark spots. Lower parts of caudal fin pale grey. Anal fin dusky grey to
dark grey anteriorly, posterior portion pale greyish to pale reddish. Pelvic fins dark grey to reddish grey, and
pectoral fins clear.
Female specific coloration. Belly dark grey to reddish grey. Anterior portion of dorsal fin bluish grey to
dark grey, posterior portion of this fin clear with no up to two black dots. Caudal fin clear without any
markings. Anal fin base dark grey, sometimes with a pale reddish flush, outer parts of fin clear. Pelvic fins
reddish to reddish brown, with dark to black anterior edge, and pectoral fins clear.
Breeding behaviour. In aquaria, this species is a monogamous, pair bonding, cave spawner. Eggs are
guarded by both sexes, but more intensively by the female. Hatching occurs after three days post-spawn.
Larvae are normally deposited on the bottom of the cave, rarely in other caves nearby the original cave.
Juveniles are free swimming 8 or 9 days post-hatching, and are guarded by both parents for about 5 to 6
weeks. Breeding and guarding individuals of both sexes regularly exhibit more aggressive and intensive
coloration. The dark, longitudinal stripe that is typical for breeding and guarding specimens of both sexes in
many other cave breeders of the chromidotilapiine lineage (e.g., Pelvicachromis, Congochromis, and
Parananochromis) is prominently visible in males, but is more rarely and weakly visible in females. In this
character, females of Enigmatochromis differ from females of Pelvicachromis, Congochromis and
Parananochromis, where a prominent dark, longitudinal stripe is typical for females (and only rarely for
males) during the first 2 to 4 weeks when guarding fry.
Distribution. This species is thus far only known from type locality, the Foto River, what is a small
savannah river near the bauxite mining town of Fria in coastal Guinea, north of Conakry, 10° 20.074’ N, 13°
Habitat. Detailed habit data for the type locality was provided by O. Lucanus. The river was 3–6 meters
wide, and during the dry season only 30–90 cm deep with water visibility ~1.2 m. Water parameters measured
during the dry season in February, 2006 were: General hardness 0, Carbonate hardness 0, pH 5.8, and
temperature 24° C.
This site was ringed by dense forest and often fallen trees obstructing access to the water in many places.
Margins of the river were densely overgrown by Anubias lanceolata, and exposed boulders and fallen trees
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NEW DWARF CICHLID FROM GUINEA, WEST AFRICA
have large clusters of Bolibitis heudelotti and other ferns (Fig. 3). The river substrate consisted of fine gravel
with a few larger rocks and boulders. Dense growths of Vallisneria were found where sunlight penetrated
through the trees.
Enigmatochromis lucanusi occured syntopically at this site with Pelvicachromis humilis.
Etymology. The species name is dedicated to the collector, aquarist and friend Oliver Lucanus.
FIGURE 3. Type locality of Enigmatochromis lucanusi, Guinea: Foto River, near the town of Fria, north of Conakry.
10°20.074’ N, 13° 43.431’ W. Photo by O. Lucanus.
This new genus and species clearly differs from Pelvicachromis, in having 12 circumpeduncular scales vs. 16,
despite the overall similarities in basic coloration patterns or body shape (Lamboj, 2005). Within
chromidotilapiine cichlids, 12 circumpeduncular scales are present in the genera Divandu, Congochromis,
Nanochromis and Parananochromis, but several other characters clearly separate the first three of these
genera from Enigmatochromis (see diagnosis of the genus for details). This leaves Parananochromis as the
chromidotilapine most morphologically similar to Enigmatochromis. However Enigmatochromis can still be
readily distinguished from Parananochromis by a much higher degree of colour dimorphism between the
sexes, a juvenile colour pattern of three or four rows of dark dots on a brownish body (vs. only two rows), and
the lack of a prominent, dark longitudinal stripe in females when guarding young fry (vs. the presence of
Additionally, the distribution of the new genus is clearly disjunct from that of all Parananochromis
species (i.e., Southern Cameroon, Rio Muni region of Equatorial Guinea, and Northern Gabon). First
indications from unpublished molecular data do not support a close relationship of the new genus to
Parananochromis. Instead the new genus seems to be more closely related to Pelvicachromis, and this
hypothesis will be evaluated in upcoming works on chromidotilapiine cichlids.
Comparative material examined.
Congochromis dimidiatus: MNHN 92-120, holotype, Central African Republic, Bangui. Congochromis
squamiceps: BMNH 1902.4.14.11, holotype, Democratic Republic of the Congo, Congo system: Lindi River.
MRAC 49244-29251, Democratic Republic of the Congo, Congo system: Kunungu. MRAC 78-19-P-265-
268, Democratic Republic of the Congo, Congo system: Riviere Iteli. Divandu albimarginatus: MRAC 95-
125-P-0569, holotype, Congo (Brazzaville): Mpoulou. Nanochromis consortus: MCZ 50552, paratypes,
Democratic Republic of the Congo: Zaire River. Nanochromis minor: MCZ 50342, paratype, Democratic
Republic of the Congo: Zaire River. Nanochromis nudiceps: MRAC 1045, holotype, Democratic Republic of
the Congo: Lac Leopold II. BMNH 1822.214.171.124, paralectotype, Democratic Republic of the Congo: Stanley
LAMBOJ 48 · Zootaxa 2173 © 2009 Magnolia Press
Pool. Nanochromis parilus: MCZ 50309, paratypes, Democratic Republic of the Congo: Zaire River. MCZ
50475, paratypes, Democratic Republic of the Congo: Zaire River. Nanochromis splendens: MCZ 50311,
paratypes, Democratic Republic of the Congo: Zaire River. MCZ 50553, paratypes, Democratic Republic of
the Congo: Zaire River. Nanochromis transvestitus: MRAC 81-14-P-1-10, paratypes, Democratic Republic
of the Congo: Lac Mai Ndombe. Parananochromis axelrodi:
AMNH 230714, holotype, Gabon: Ivindo
system. Parananochromis brevirostris: AMNH 232536, holotype, Gabon: Ivindo system. Parananochromis
caudifasciatus: BMNH 1904.7.1.244-249, syntypes, Cameroon: Nyong River system. Parananochromis
gabonicus: BMNH 19126.96.36.199, holotype, Gabon: Ogowe River system. Parananochromis longirostris:
BMNH 1903.7.28.77-83, syntypes, Cameroon: Kribi River. Parananochromis ornatus: MRAC A2-011-P-
10, holotype, Gabon: Ivindo system. Pelvicachromis humilis: BMNH 1915.4.13:44, holotype, Sierra Leone:
North Sherbo District. MRAC 154802, Sierra Leone: Kasewe Forest. Pelvicachromis pulcher: BMNH
1901.1.28.13-20, syntypes, Nigeria: Ethiop River. Pelvicachromis roloffi: All from Sierra Leone, Kwabolake,
Sherbo District: MRAC 73399, holotype. MRAC 733400, allotype. MRAC 73401-402, paratypes.
Pelvicachromis taeniatus: BMNH 1901.1.28.21, holotype, Nigeria: Ethiop River. BMNH 19188.8.131.52-28 +
BMNH 1902.11.12.164-165 (syntypes of P. kribensis), Cameroon: Kribi River.
I thank the following persons and institutions for help in my work, loan of material or discussion on the
manuscript: M. Stiassny, R. Schelly (AMNH), J. Maclaine (BMNH), J. Snoeks, M. Parrent (MRAC), U.
Schliewen (ZSW), E. Mikschi (NMW). Also thanks to two anonymous reviewers for their valuable
comments. Very special thanks to O. Lucanus (Montreal, Canada), T. Orso (Vernon, USA) and M. Hakansson
(Gothenburg, Sweden), for donating material and to O. Lucanus for information about the type locality and
distribution of the new genus and species.
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