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209
Accepted: 28 May 2003; published: 6 June 2003 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2003 Magnolia Press
Zootaxa 209: 1-19 (2003)
www.mapress.com/zootaxa/
Three new Parananochromis species (Teleostei, Cichlidae) from
Gabon and Cameroon, Central Africa
ANTON LAMBOJ * & MELANIE L.J. STIASSNY§
* Institut für Zoologie der Universität Wien, Abteilung für Evolutionsbiologie, Althanstrasse 14, A - 1090
Wien, Austria. E-mail: anton.lamboj@univie.ac.at
§Department of Ichthyology, American Museum of Natural History, 79th Street at Central Park West, New
York, New York 10024, USA. Email: mljs@amnh.org
Abstract
Parananochromis axelrodi,P. brevirostris and P. ornatus, three new cichlid species, are described
from Gabon and southern Cameroon. Parananochromis axelrodi is a deep-bodied species differing
from congeners in a combination of morphometric and meristic characters. Parananochromis brevi-
rostris and P. ornatus are distinguished by the presence of four elements in the infraorbital series
versus five elements in congeners. Parananochromis brevirostris differs from P. ornatus in lacking
chest squamation and in the presence of a weakly-developed pharyngeal pad (versus well-devel-
oped pad and scaled chest in P. or na tu s).
Key words:Parananochromis, new species, South Cameroon, Gabon
Introduction
The cichlid genus Parananochromis was erected by Greenwood (1987) to accommodate
three species formerly placed in Pelmatochromis and Nanochromis. These species were
P.longirostris (Boulenger, 1903) designated as the type of Parananochromis,P. caud if as -
ciatus (Boulenger, 1913), and P.gabonicus (Trewavas, 1975). Although the precise rela-
tionships of this taxon remain problematical our current understanding is that Para-
nanochromis belongs to the chromidotilapiine lineage. This assemblage currently com-
prises seven genera united by the possession of several derived anatomical features,
including the presence of a visor-like pad on the roof of the buccopharynx just anterior to
the upper pharyngeal bones, tuberculate gill-rakers on the outer row of the first gill arch,
and the absence of microbranchiospines on the second to fourth gill arches (Greenwood
1987; Stiassny 1991). Parananochromis are readily distinguished from the chromidotilapi-
LAMBOJ & STIASSNY
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209
ZOOTAXA ine genera Thysochromis,Chromidotilapia,Pelvicachromis,andBenitochromis by the
possession of 12 circumpeduncular scales (versus 15-16), from Divandu by the presence
of 4 openings of the laterosensory system in the first infraorbital bone (versus 5 in
Divandu), and from Nanochromis (the only other chromidotilapiine genus with a Haplo-
chromis-type rather than a Tilapia-type of neurocranial apophysis) by the configuration of
the infraorbital series and lateral line.
Shortly after Greenwood´s (1987) paper was published two Parananochromis species,
unassignable to any described species, were collected from localities in the Ivindo River, a
large black water tributary of the Ogowe River, Gabon. One was collected in the vicinity
of Belinga (Numrich & Wunderlich, 1988), and the second from Makokou (Lamboj, 1988).
Aquarists raised both species and information on behaviour and reproductive biology has
since appeared in the aquarium hobbyist literature (e.g., Lamboj 1999a, Linke & Staeck
2002) under the names Parananochromis sp. “Belinga” and Parananochromis sp.
“Makokou”. Between 1995 and 2002, as part of an ongoing effort to document the ichthy-
ofauna of the Lower Guinean ichthyofaunal province (Cameroon, Rio Muni, Gabon and
Congo-Brazzaville), numerous additional collections were made throughout the region,
and examination of these as well as historical collections have led to a better understand-
ing of the distribution of these two undescribed species. Additionally, recent collections in
the region of Makokou, and historical material from a nearby locality, have revealed the
presence there of a third undescribed Parananochromis species.
In the past five years numerous new cichlid taxa have been described from the Lower
Guinean icthyofaunal province (see e.g., Lamboj 1999b, 2001, 2002, 2003; Lamboj &
Snoeks, 2000; Schliewen & Stiassny, 2003), and many more remain to be described.
Among these are the three Parananochromis species mentioned above, which we herein
describe.
Material and methods
Counts and measurements follow Barel et al. (1977). All measurements of bilaterally
paired structures were taken on the left side with digital callipers with an accuracy of +/-
0.03 mm.
Clearing and staining of bones and cartilage follows Dingerkus & Uhler (1977).
Abbreviations are: AMNH, American Museum of Natural History, New York;
BMNH, Natural History Museum, London; CU, Cornell University, Ithaca; IRET, Institut
de Recherche en Ecologie Tropicale, Libreville; MHNG, Muséum d’Histoire Naturelle,
Genève; MRAC, Musée Royal de l´Afrique Centrale, Tervuren; NMW, Naturhistorisches
Museum, Wien; SL, standard length; HL, head length; BD, body depth.
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PARANANOCHROMIS
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ZOOTAXA
Parananochromis axelrodi, new species
(Fig. 1)
Holotype. AMNH 230714, male, 79.2 mm SL; Gabon: Ivindo system; mouth of Bale
creek into Ivindo. Just inside mouth of Bale creek in forest, 0°30´53.9”N, 12°48´21.2”E,
J.P. Sullivan et al., Jan 1998.
Paratypes. Total of 8 specimens, 25.6-91.2 mm SL; AMNH 233351, 1 undet., 25.6
mm SL; Gabon: Ivindo River system, lower Bale creek, 0°30´57”N, 12°48´17.8”E, M.L.J.
Stiassny et al., Jan 1998. – AMNH 230665, 1 male, 53.7 mm SL; Gabon: Ivindo system;
gill net across mouth of Bale creek into Ivindo, 0°30´54.5”N, 12°48´20.5”E, E. Vreven
and C. Hopkins., Jan 1998. – CU 87044, 1 undet., 35.0 mm SL, Gabon: Ivindo system,
small creek mouth emptying into Ivindo River, across from IRET field station,
0°30´15.4”N, 12°48´52.4”E, M.L.J. Stiassny et al., Jan 1998. – AMNH 233350, 3 undet.,
25.7-51.0 mm SL, 1 spec. 51.0 mm SL cleared and stained, Gabon: Ivindo system, small
creek about 200 m downstream of IRET field station (on opposite side of river),
0°30´34”N, 12°48´33”E, M.L.J.Stiassny and E. Vreven., Feb. 1998. - MRAC-A2-046-P-
1, 1 male, 49.5 mm SL, Gabon: Ivindo system, small creek about 200 m downstream of
IRET field station (on opposite side of river), 0°30´34”N, 12°48´33”E, M.L.J.Stiassny
and E. Vreven, Feb. 1998. – MHNG 2203.047, 1 male, 91.2 mm SL, Gabon: Ivindo sys-
tem, Bras mort of the Djouah, upstream of Mvaddi, J. Géry, Sept. 1964.
Differential Diagnosis. Parananochromis axelrodi is readily distinguished from
P.brevirostris and P. ornatus by the possession of 4 (versus 3) tubular infraorbital bones
and a total of 15-18 gill rakers on the outer row of the first gill arch (versus 8-12 in P.
brevirostis and 10-13 in P. ornatus), from P. caudifasciatus and P. gabonicus by number
of gill rakers along outer row of first gill arch (15-18 versus 10-14 in P. caudifasciatus and
11-14 in P. gabonicus), and by its deep-body (BD 35.4-42.2% SL versus 27.0-36.0% in P.
gabonicus and 29.7-36.2% in P.caudifasciatus), and from P. longirostris by body depth
(BD 35.4-42.1% SL versus 31.1-35.3% in P. longirostris), and by its deep caudal peduncle
(length as % of depth 57.6-86.5% versus 88.5-105.6% in P. longirostris).
FIGURE 1. Holotype of Parananochromis axelrodi, AMNH 230714, male, 79.2 mm SL; Gabon:
Ivindo River system; mouth of Bale creek into Ivindo. Just inside mouth of Bale creek in forest.
LAMBOJ & STIASSNY
4© 2003 Magnolia Press
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ZOOTAXA Description. Measurements and meristic counts for holotype and 8 paratypes are given
in Table 1.
TABLE 1. Morphometrics and meristics of the holotype and 8 paratypes of Parananochromis axel-
rodi.
holotype mean SD max. min.
Standard length 53.0 42.5 25.6 - 79.2
% of standard length
Body depth 42.1 38.1 2.3 35.4 - 42.1
Head length 35.7 35.7 1.5 33.0 - 37.2
Caudal peduncle length 11.4 12.4 1.2 10.2 - 13.5
Caudal peduncle depth 17.7 16.7 0.9 15.6 - 17.8
Length of dorsal fin base 60.1 58.3 2.7 55.5 - 62.9
Length of anal fin base 19.4 18.5 1.1 17.0 - 20.0
Predorsal distance 31.2 31.0 2.0 29.2 - 34.7
Preanal distance 72.3 71.0 2.7 67.8 - 75.4
Prepectoral distance 36.8 37.2 2.1 34.4 - 39.9
Prepelvic distance 42.6 40.3 1.4 37.7 - 42.6
% of head length
Head depth 70.6 65.2 3.2 61.3 - 70.6
Snout length 35.9 29.3 5.0 24.6 - 35.9
Eye diameter 26.3 31.0 2.7 26.3 - 33.7
Postorbital distance 37.8 39.7 2.8 36.0 - 43.2
Interorbital distance 24.9 23.6 2.3 19.5 - 26.8
Cheek depth 30.9 27.6 2.4 24.6 - 30.9
Lower jaw length 31.4 28.1 5.7 19.0 - 35.4
Preorbital depth 19.8 19.8 1.3 18.1 - 22.2
% of caudal peduncle depth
Caudal peduncle length 64.5 74.3 10.1 57.5 - 86.5
Meristics median
Upper lateral-line series 17 18 17 - 19
Lower lateral-line series 9 7 6 - 9
Total lateral-line series 27 27 26 - 27
Circumpeduncular scales 12 12 12
Spines dorsal fin 15 15 15
Rays dorsal fin 11 11 11
Rays anal fin 8 8 7 - 8
Rays pectoral fin 13 13 12 - 13
Gill rakers on lower limb of first arch 12 10 9 - 12
Total gill rakers on first arch 18 16 15 - 18
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PARANANOCHROMIS
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A robust, deep-bodied species (BD 35.4-42.2% SL, 40.0-42.2% in specimens over 50
mm SL). The four largest specimens are males (91.2, 79.2, 53.7, 49.5 mm SL) with mini-
mal development of the testes; the remainder are immatures of indeterminate sex. Cur-
rently we have limited information on sexual dichromatism in the species. First ray of
pelvic fin is the longest in the fin and is produced in large males reaching to, or overlap-
ping the anal fin spines. Caudal fin is rounded.
Osteology and dentition. Infraorbital series (Fig. 2a) first infraorbital (commonly
termed the lachrymal) is plate-like and followed by four tubular elements. There is a small
gap between the fourth and fifth infraorbitals. Infraorbital 1 has four openings of the lat-
erosensory system. Twenty five vertebrae, of which 13 are precaudal and 12 caudal. Pre-
maxilla and dentary with 1-3 rows of small, acutely pointed, unicuspid teeth. Outer row
teeth are slightly larger than those of inner rows. Lower pharyngeal boneis narrowly trian-
gular, with numerous, slender, shouldered unicuspid teeth on lateral parts of the bone and
larger asymmetric bicuspid teeth in the central field.
FIGURE 2. Infraorbital series of a) Paranaochromis axelrodi, AMNH 233350 (51.0 mm SL), b)
Parananochromis brevirostris, AMNH 230707 (41.2 mm SL), and c) Parananochromis ornatus,
AMNH 233349 (46.8 mm SL). Position of openings of the laterosensory system on the first infraor-
bital bones indicated with black dots.
Gill rakers on first gill arch. Lower limb with 9-12 tuberculate gill rakers, 5-6 elongate
gill rakers on the epibranchial. No microbranchiospines are present. A well-developed
hanging pad is present on roof of the pharynx.
LAMBOJ & STIASSNY
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ZOOTAXA
FIGURE 3. Geographical distribution of P. axelrodi, P. brevirostris, and P. ornatus.
Squamation. Cycloid, 2-3 rows of scales on the cheek, 3-4 horizontal rows on the
opercle. Naked dark spot on the outer edge of opercle. Chest scales are smaller than scales
on flanks.
Upper lateral line separated from dorsal-fin base at its highest point (8th pored scale)
by 2 scales, at last pored scale by ½-1 scales. No overlap between the end of upper lateral
line and lower lateral line. Caudal fin scaled basally for about one third of its length; other
fins are unscaled.
Coloration. Living specimens: Base body coloration is pale bronze brown with a dark
spot on outer edge of opercle, no other body markings are evident. Dorsal, anal and caudal
fin membranes with numerous rows of dark maculae (MLJS, pers. field obs.).
Preserved specimens (Fig. 1): Coloration of the head and body is pale to dark brown
and darker dorsallythan ventrally. A midlateral stripe is variously developed, passing from
the eye through the opercular spot onto the anterior third of the body. The midlateral stripe
is interrupted from mid-body to the caudal peduncle, and does not extend over the caudal
Ogowe
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PARANANOCHROMIS
209
ZOOTAXA
fin. Four-5 vertical bars are variously developed from the dorsum to somewhat below the
midlateral stripe. The unpaired fins are dusky grey to brownish. The soft-dorsal, soft-anal
and caudal fin membranes have numerous rows of dark maculae.
Breeding behaviour. No information available.
Distribution (Fig. 3). Currently known only from the Ivindo River system in the
regions of Makokou and Belinga, Central Gabon.
Etymology. Named for Herbert Axelrod in recognition of his generous support of ich-
thyological research and exploration.
Remarks. Despite considerable collecting efforts in the Ivindo River (Géry, 1965; pers.
obs.), particularly in the region of Makokou, very few specimens of P. axelrodi have so far
been collected, and the species is currently known only from the nine specimens of the
type series. These have all been collected in small forest creeks and streams (marigots),
but not in the main channel of the Ivindo River.
Parananochromis brevirostris, new species
(Figs. 4-9)
Nanochromis cf. dimidiatus – Géry, 1965: 381
Parananochromis sp. “Makokou” – Lamboj, 1999a: 126; Linke & Staeck, 2002: 151.
Holotype. AMNH 232536, male, 42.5 mm SL; Gabon: Ivindo system, Small, shallow
creek ca. 200m downstream of IRET station (on opposite side of river), 0°30´34”N,
12°48´33” E, M.L.J. Stiassny & E. Vreven, Jan 1998.
Paratypes. Total of 57 specimens, 21.2-51.2 mm SL, AMNH 230720, 15 males, 6
females, 2 undet. 31.4-42.2 mm SL, Gabon: Ivindo system, Small creek ca. 200m down-
stream of IRET station (on opposite side of river), 0°30´34”N, 12°48´33” E, M.L.J.
Stiassny & E. Vreven, Jan 1998. – AMNH 230707, 4 undet, cleared and stained, 32.0-41.2
mm SL, Gabon: Ivindo system, small creek mouth emptying into Ivindo River, across
from IRET field station, 0°30’15.4”N, 12°48’52.4”E, M.L.J. Stiassny et al., Jan. 1998. –
MRAC 73-02-P-2147-150, 1 male, 2 females, 1 undet., 21.2-47.4 mm SL, Gabon: Ivindo
system, route Makokou-Loloa, 3rd marigot, D.F.E.Thys van den Audenaerde, Nov 1964. –
MRAC 93-085-P-0334-0340, 3 males, 4 females, 30.0-45.4 mm SL, Cameroon: Ntem sys-
tem, riv. Lé, downstream of Abang-Minkoo, 2°20´N, 11°26´E, A. Kamden, May 1993. –
MRAC A2-11-P-15, 1 male, 46.2 mm SL, Gabon: Ivindo system, tributary of riv. Aboy,
route Makokou-Okondja, 0° 32´N, 12°56´E, A. Lamboj, R. Guggenbühl, P. Sewer & A.
Weissenbacher, Jul 1995. – MRAC A2-11-P-16, 1 female, 33.0 mm SL, Gabon: Ivindo
system, little creek on route Makokou-Ovan, 7km from Makokou, 0°34´N, 12°45´E, A.
Lamboj, R. Guggenbühl, P. Sewer & A. Weissenbacher, Jul 1995. – MRAC A2-11-P-17, 1
undet., 37.8 mm SL, Gabon: Ivindo system, Ngong-Bissoga, route Makokou-Ovan, at vil-
lage Adoue, 0°31´N, 12°33´E, A. Lamboj, R. Guggenbühl, P. Sewer & A. Weissenbacher,
LAMBOJ & STIASSNY
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ZOOTAXA Jul 1995. – MRAC A2-011-P-18, 1 male, 51.2 mm SL, Gabon: Ivindo system, tributary of
riv. Aboy, route Makokou-Okondja, 0° 32´N, 12°56´E, A. Lamboj, R. Guggenbühl, P.
Sewer & A. Weissenbacher, Jul 1995. – BMNH 2002.8.8.4-6, 2 males, 1 undet., 38.4-48.6
mm SL, Gabon: Ivindo system, tributary of riv. Aboy, route Makokou-Okondja, 0° 32´N,
12°56´E, A. Lamboj, R. Guggenbühl, P. Sewer & A. Weissenbacher, Jul 1995. – NMW
94630, 3 females, 31.6-49.8 mm SL, Gabon: Ivindo system, riv. Zaidie, on bridge of road
Makokou-La Scierie-Belinga, 0°44´N, 13°09´E, A. Lamboj, R. Guggenbühl, P. Sewer &
A. Weissenbacher, Jul 1995. – MHNG 2640.27, 1 male, 1 undet, 29.4-40.9 mm SL, Djad-
die system, marigot in the Djaddie, Mekambo, J.Gery, Aug 1964. – MHNG 2640.28, 2
undet. 31.9-37.9 mm SL, Marigot, Djaddie at Mekambo, J.Gery, Aug 1964. - CU 87042 , 2
males, 3 females, 33.2-40.4 mm SL, Gabon: Ivindo system, Small creek ca. 200m down-
stream of IRET station (on opposite side of river), 0°30´34”N, 12°48´33” E, M.L.J.
Stiassny & E. Vreven, Jan. 1998.
Differential Diagnosis. Parananochromis brevirostris is unique among congeners in
the absence of scales on the chest and weakly developed, or absent, pharyngeal hanging
pad. It is also readily distinguished from all congeners except P. orna tu s by the possession
of 4 (versus 5) infraorbital bones.
Description. Measurements and meristic counts for holotype and 50 paratypes are
giveninTable2.
Thisisadiminutivespecies,thelargestspecimenisamaturemale(51.2mmSL),and
the largest female is 38.2 mm SL with ripening eggs in its ovaries. Sexual dimorphism is
well developed. Males are usually 20-30 % larger than females, with soft dorsal and anal
fin rays more produced. The first ray of pelvic fin is usually the longest in the fin of both
sexes, and is produced in males; in females the second ray may occasionally be slightly
longer than first. The tips of pelvic fins in large males usually reach the anus. Lappets of
spiny dorsal fin are elongated and pronounced, particularly so in males. The caudal fin is
rounded in both sexes. Snout is short and rounded (length 19.2-31.3% HL), and the body
gracile even in the largest specimens (depth 22.7-31.6% SL).
Osteology and dentition. Infraorbital series (Fig. 2b) with a plate-like first infraorbital
followed by three 3 tubular elements. There is a large gap between the third infraorbital
and a small fourth infraorbital. Infraorbital 1 with four openings of the laterosensory sys-
tem. Usually 25 vertebrae, 12 precaudal and 13 caudal (26 vertebrae, 12 precaudal and 14
caudal in two individuals). Premaxilla and dentary with (2) 3-5 rows of numerous, acutely
cusped, unicuspid teeth with little size difference between the teeth of inner and outer rows
(Fig. 5). Lower pharyngeal bone narrowly triangular, with numerous, slender, shouldered
unicuspid teeth on lateral parts of the bone and larger asymmetric bicuspid teeth in the
central field.
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ZOOTAXA
TABLE 2. Morphometrics and meristics of the holotype and 50 paratypes of Parananochromis
brevirostris.
holotype mean SD max. min.
Standard length 42.5 37.3 21.2 - 51.2
% of standard length
Body depth 25.2 27.4 1.8 22.7 - 31.6
Head length 29.2 32.2 1.5 28.5 - 35.8
Caudal peduncle length 14.8 13.7 1.1 10.6 - 16.3
Caudal peduncle depth 13.7 13.7 0.7 12.4 - 15.4
Length of dorsal fin base 57.6 56.2 1.9 52.2 - 59.6
Length of anal fin base 17.3 17.5 1.0 14.8 - 19.5
Predorsal distance 29.4 30.1 1.9 25.9 - 35.4
Preanal distance 68.3 69.3 2.4 64.9 - 78.1
Prepectoral distance 32.6 35.8 1.8 32.6 - 41.4
Prepelvic distance 35.4 38.1 2.1 31.7 - 43.7
% of head length
Head depth 58.9 56.5 4.0 47.7 - 64.4
Snout length 23.3 25.1 2.6 19.2 - 31.3
Eye diameter 33.9 32.6 1.8 27.7 - 37.2
Postorbital distance 42.8 42.6 1.8 38.6 - 46.6
Interorbital distance 18.4 18.3 1.2 15.4 - 21.6
Cheek depth 25.2 22.8 1.8 18.7 - 25.6
Lower jaw length 34.7 29.7 3.4 21.4 - 34.7
Preorbital depth 16.9 15.1 1.8 11.8 - 19.1
% of caudal peduncle depth
Caudal peduncle length 108.6 100.2 8.9 78.7 - 119.3
Meristics median
Upper lateral-line series 17 17 16 - 20
Lower lateral-line series 5 6 4 - 8
Total lateral-line series 26 26 23 - 29
Circumpeduncular scales 12 12 12
Spines dorsal fin 15 15 14 - 16
Rays doral fin 9 9 8 - 10
Spines anal fin 3 3 3
Rays anal fin 7 7 6 - 8
Rays pectoral fin 12 12 11 - 13
Gill rakers on lower limb of first arch 7 7 5 - 8
Total gill rakers on first arch 11 10 8 - 12
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ZOOTAXA
FIGURE 4. Holotype of Parananochromis brevirostris; AMNH 230720, male, 42.5 mm SL;
Gabon: Ivindo River system, Small creek ca. 200m downstream of IRET station.
FIGURE 5. Parananochromis brevirostris, premaxillary dentition (SEM-Photo: R. Riehl)
Gill rakers on first gill arch. Lower limb with 5-8 tuberculate gill rakers, 2-4 pointed
gill rakers on the epibranchial. No microbranchiospines are present. The hanging pad on
roof of the pharynx is poorly developed or absent.
Squamation. Cycloid, cheek usually naked, occasionally with a single scale row, 3-4
horizontal scale rows on the opercle. Dark spot on outer edge of opercle is unscaled. Chest
is naked. Upper lateral line is separated from dorsal-fin base at its highest point (8th pored
scale) by 1-1½ scales, at last pored scale by 0-½ scales. No overlap between the end of the
upper lateral line and the lower lateral line. Caudal fin scaled basally for about one third of
its length; the other fins are unscaled.
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ZOOTAXA
Coloration. Living specimens (Figs. 6-9): Base body coloration is grey to light brown,
darker dorsally than ventrally. Dark spot on the outer edge of opercle is extended anteri-
orly and posteriorly into a midlateral stripe. In males the soft dorsal fin and upper margin
of the caudal fin have a narrow white border, in females these borders are red. In both
sexes these fins have a thin black submarginal band. Soft-dorsal, caudal and posterior parts
of the soft-anal fins in both sexes are clear or grey, with rows of dark maculae, which are
always more marked in males than in females – females sometimes with immaculate fins.
Anterior parts of the anal fin are violet to wine red in males and pale yellow or clear in
females. Males with reddish coloration in the lower lobe of the caudal fin. Pelvic fins are
pale yellow to reddish, with a dark anterior leading edge; all colorations are more marked
in males than in females. Pectoral fins are clear. Sometimes a dark horizontal stripe is vis-
ible, passing from the snout through the eye, and extending onto the first third of the cau-
dal fin. Occasionally this stripe is replaced by a row of black spots or blotches. Well-
marked interorbital, nostril, and lachrymal stripes are present on the head. Ripe females
have a rosy to orange or violet belly. Lips are light grey to light brown. Body scales with
dark margins on the whole body in males, in females such markings are restricted to the
dorsal parts of the body.
FIGURES 6-9. Parananochromis brevirostris (specimens not preserved). 6, male, same locality as
NMW 94630; 7, male, from the Menguengne river near Ndjole, Gabon, Ogowe River system; 8,
female, same locality as NMW 94630; 9, female, from the Menguengne river near Ndjole, Gabon,
Ogowe River system.
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ZOOTAXA Preserved specimens (Fig. 4): Coloration of the head and body are brown, with the
upper half darker than the ventral parts. A dark, midlateral stripe, is continuous from the
eye (merging with the dark opercular spot) and extending onto the first third of the caudal
fin. Unpaired fins are dusky grey or brownish. The soft-dorsal, soft-anal, and caudal fin
membranes have rows of dark maculae.
Breeding behaviour. In aquaria a pair-bonding, cave-breeding species. For further
information on breeding biology see Lamboj, (1999a, as Parananochromis sp.
“Makokou”).
Distribution (Fig. 3). Currently the species is known from scattered localities in the
Ivindo and Ntem systems in eastern Gabon and southeastern Cameroon, and from the
Ogowe system in the region of Ndjole in Western Gabon.
Etymology. From the Latin brevis, short and rostrum, beak or snout, in reference to
the short, rounded snout of the species.
Remarks. This diminutive species is found in abundance in the many small forest
creeks and streams (marigots) of the Ivindo River system and is also recorded from the
River Lé, a small affluent of the Ntem in similar habitats. Specimens from the Men-
guengne River (Ogowe system) near the town of Ndjole in western Gabon have been col-
lected and raised in aquaria (Figs 7,9), but not yet preserved. Individuals from the vicinity
of Makokou were misidentified by Géry (1965) as Nanochromis cf. dimidiatus.
Parananochromis ornatus, new species
(Figs 10-12)
Parananochromis sp. “Belinga” – Lamboj, 1999a: 126; Linke & Staeck, 2002: 149.
Holotype. MRAC A2-011-P-10, male, 51.0 mm SL; Gabon: Ivindo system, small creek on
route Makokou-Ovan, 7km from Makokou, 0°34´N, 12°45´E, A. Lamboj, R. Guggenbühl,
P. Sewer & A. Weissenbacher, Jul 1995.
FIGURE 10. Holotype of Parananochromis ornatus, MRAC A2-011-P-10, male, 51.0 mm SL;
Gabon: Ivindo River system, small creek on route Makokou-Ovan, 7km from Makokou.
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Paratypes. Total of 18 specimens, 27.3 – 52.0mm SL, MRAC A2-011-P-11-14, 1
male, 3 females, 31.0-45.2 mm SL, Gabon: Ivindo system, tributary of riv. Aboy, route
Makokou-Okondja, 0° 32´N, 12°56´E, A. Lamboj, R. Guggenbühl, P. Sewer & A. Weis-
senbacher, Jul 1995. – NMW 94632, 1 male, 1 undet., 27.3-30.04 mm SL, Gabon: Ivindo
system, tributary of riv. Aboy, route Makokou-Okondja, 0° 32´N, 12°56´E, A. Lamboj, R.
Guggenbühl, P. Sewer & A. Weissenbacher, Jul 1995. – AMNH 230704, 1 male, 3
females, 29.5-39.1 mm SL, Gabon: Ogowe system, Okondja, creek south of Okondja
about 1 km south of 98-045 site 2, Muddy water. M.L.J. Stiassny et al., Jan. 1998. –
AMNH 232113, 1 male, 1 female, 37.7-46.9 mm SL, Gabon: Ivindo system, Minkebe
Gold Camp forest, 01°44´N, 12°48´E, S.A.Lahm, May 2000 – AMNH 233349, 2 males, 2
females, 30.2-52.0 mm SL, 3 cleared and stained, 30.2-46.8 mm SL, Gabon: Ivindo sys-
tem, Minkebe Gold Camp forest, 0°43’N, 12°49’E, S.A. Lahm, May 2000 - CU 87043, 2
males 46.4-37.6 mm SL, Gabon: Ivindo system, Minkebe Gold Camp forest, 01°44´N,
12°48´E, S.A.Lahm, May 2000.
Differential Diagnosis. Parananochromis ornatus is readily distinguished from all
congeners except P. brevirostris by the possession of 4 (versus 5) infraorbital bones. From
brevirostris it differs in the possession of a scaled chest (naked in brevirostris), a well-
developed pharyngeal pad, and usually 26 vertebrae (versus 25 in brevirostris).
Description. Measurements and meristic counts for holotype and 18 paratypes are
giveninTable3.
The largest specimen in the type series is a mature male, 52.0 mm SL, but males regu-
larly attains sizes of up to 70.0 mm SL in aquaria. Sexual dimorphism is well developed.
Males are usually 20-30 % bigger than females and the soft dorsal and anal rays are pro-
duced. In adults of both sexes, the first ray of pelvic fin is usually the longest, but occa-
sionally the second ray is slightly longer than the first in females. The tips of pelvic fins in
larger males reach to or overlap the anus. Caudal fin is normally rounded in both sexes;
sometimes in males the upper lobe has a slight prolongation. The snout is rounded or
somewhat acute.
Osteology and dentition. Infraorbital series (Fig. 2c) with plate-like first infraorbital
followed by 3 tubular elements. A small gap between the third and fourth infraorbitals.
Infraorbital 1 with four openings of the laterosensory system. Usually 26 vertebrae with 13
precaudal and 13 caudal (25 vertebrae, 13 precaudal and 12 caudal in two individuals, 27
vertebrae, 13 precaudal and 14 caudal in one individual). Premaxilla and dentary with 2-3
(rarely 4) rows of acutely cusped, unicuspid teeth. Outer row teeth are slightly larger than
those of the inner rows. Lower pharyngeal bone is narrowly triangular, with numerous,
slender, shouldered unicuspid teeth on the lateral parts of the bone and larger asymmetric
bicuspid teeth in the central field.
Gill rakers on first gill arch. Lower limb with 7-9 tuberculate gill rakers, 2-4 pointed
gill rakers on the epibranchial. No microbranchiospines are present. A well-developed
hanging pad is present on the pharynx roof.
LAMBOJ & STIASSNY
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ZOOTAXA TABLE 3. Morphometrics and meristics of the holotype and 18 paratypes of Parananochromis
ornatus.
holotype mean SD max. min.
Standard length 51.0 37.8 27.3 - 52.0
% of standard length
Body depth 29.6 27.9 2.1 24.1 - 33.0
Head length 33.4 33.7 1.4 31.6 - 36.0
Caudal peduncle length 13.7 13.5 1.1 11.2 - 14.8
Caudal peduncle depth 13.4 13.1 0.6 12.0 - 14.1
Length of dorsal fin base 54.7 55.2 2.5 50.3 - 59.3
Length of anal fin base 17.0 17.4 0.9 16.0 - 19.3
Predorsal distance 32.6 30.9 1.2 28.5 - 32.9
Preanal distance 66.1 68.8 2.2 65.2 - 73.6
Prepectoral distance 37.0 36.4 1.4 34.1 - 38.8
Prepelvic distance 37.5 37.9 1.6 35.3 - 40.3
% of head length
Head depth 60.7 58.2 4.3 48.8 - 63.7
Snout length 33.7 29.0 2.3 26.3 - 33.7
Eye diameter 27.2 30.2 1.4 27.2 - 32.7
Postorbital distance 39.1 40.8 2.0 36.8 - 44.0
Interorbital distance 19.1 19.6 1.4 16.9 - 22.8
Cheek depth 31.5 25.9 1.9 23.0 - 31.5
Lower jaw length 37.9 34.7 3.9 27.3 - 41.1
Preorbital depth 20.2 17.7 2.1 14.8 - 21.1
% of caudal peduncle depth
Caudal peduncle length 102.0 102.9 8.8 80.2 - 119.3
Meristics median
Upper lateral-line series 20 19 16 - 20
Lower lateral-line series 7 7 6 - 7
Total lateral-line series 27 28 26 - 29
Circumpeduncular scales 12 12 12
Spines dorsal fin 15 16 15 - 16
Rays doral fin 9 9 9 - 10
Rays anal fin 8 7 7 - 8
Rays pectoral fin 12 12 12 - 13
Gill rakers on lower part of first arch 8 8 7 - 9
Total gill rakers on first arch 13 13 10 - 13
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Squamation. Cycloid, cheek usually with 2-3 scale rows but occasionally only a single
row, 3 horizontal scale rows on the opercle. Dark unscaled spot on outer edge of opercle is
well-developed. Chest scales are very small and deeply embedded, often with no free edge
exposed, usually with 3-5 scales between pectoral and pelvic fin insertion. Upper lateral
line separated from dorsal-fin base at its highest point (8th pored scale) by 1½-2 scales, at
last pored scale by 0-½ scales. No overlap between the end of upper lateral line and lower
lateral line. Caudal fin scaled basally for about one quarter of length; other fins are
unscaled.
Coloration. Living specimens (Figs 11-12): Body greyish to brownish, darker dorsally
than ventrally with a dark spot on the outer edge of the opercle. Dorsal fin and upper part
of the caudal fin with a thin white margin, sometimes also with a thin red submargin.
Anterior parts of the dorsal fin yellowish brown, anterior parts of the anal fin yellowish
red. Posterior parts of the soft-dorsal fin, the upper 2/3 of caudal and the posterior parts of
the soft- anal fin membranes clear or bluish, always with rows of reddish maculae in
males, sometimes also in females but then fewer in number and intensity. Pelvic fins clear
or reddish, with dark anterior edges; coloration is more intense in males than in females.
Pectoral fins are clear. Normally a black midlateral band is visible, passing from the fore-
head, through the eye, and extending onto the first third of the caudal fin. A lachrymal
stripe is well developed. Occasionally the midlateral band is indistinct or absent. The
upper edge of the eye is red or yellow. A black band on the uppermost part of the back is
sometimes present. This band may also be dispersed and blotchy or completely absent.
Ripe females with a rosy belly. Lips are grey-brown. Dark margins on the body scales are
particularly marked in males.
FIGURES 11-12. Parananochromis ornatus (specimen not preserved). 11, male, same locality as
NMW 94632; 12, female, same locality as NMW 94632.
FIG. 12
LAMBOJ & STIASSNY
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ZOOTAXA Preserved specimens (Fig. 10): Base body coloration is brown or dark grey, with the
upper half darker than the ventral half. A midlateral stripe passing from the snout to the
middle of the body (merging with the dark spot on the outer edge of the opercle) is usually
visible. Unpaired fins are dusky grey to brownish. Soft-dorsal, soft-anal and caudal fin
membranes always with rows of dark maculae in males, sometimes also in females but
then less prominently marked.
Breeding behaviour. In aquaria a pair bonding, cave-breeding species. For further
information on breeding biology see Lamboj, (1999a, as Parananochromis sp. “Belinga”).
Distribution (Fig. 3). The Ivindo system in Eastern Gabon and the Ogowe system in
the vicinity of Okondja.
Etymology. From the Latin ornatus, handsome or splendid, in reference to the attrac-
tive coloration of the species.
Remarks. At the locality where the holotype was collected (Ivindo River system, small
creek on route Makokou-Ovan, 7km from Makokou), P. ornatu s was collected with speci-
mens of P. brevirostris and P. longirostris. Thys van den Audenaerde (1968) mentions a
taxon, Pelmatochromis sp. aff. caudifasciatus, as being widespread in the Ogowe system
and also in the periphery of the central Congo. His brief description of this taxon
(1968:368), which is presented as the “link” to Nanochromis,isprobablyP. o rn at us
although none of the material which Thys van den Audenaerde based his comments on has
been located.
Discussion. The combination of the possession of four openings of the sensory canal
system in the lachrymal (versus 5), 12 circumpeduncular scales (versus 14-16), and a Hap-
lochromis-type (versus a Tilapia-type) of neurocranial apophysis, characterizes the two
genera, Parananochromis and Nanochromis. Greenwood (1987) considered the latter two
features as derived within the chromidotilapiine lineage and therefore as evidence of a sis-
tergroup relationship between the two genera. The distinction between Nanochromis and
most of the species currently included in Parananochromis was discussed by Thys van den
Audenaerde (1968), and later by Greenwood (1987) who diagnosed Nanochromis on the
basis of a number of derived features (e.g., two bones in the infraorbital series, at least the
last 3 or 4 pored scales of the upper lateral line contiguous with dorsal fin base, and a low
supraoccipital crest). However, no such diagnosis was presented for Parananochromis,
which remains uncharacterised by uniquely derived features and possibly therefore a para-
phyletic assemblage. In the course of the present study we have found no evidence that
could be used to support the monophyly of Parananochromis. In fact two of the new taxa
described herein (P. brevirostris and P. or nat us ) display an elevation of the upper lateral
line (albeit not to the extent found in Nanochromis), a reduction of infraorbital elements
from five to four (Nanochromis has a single tubular element), and the possession of a low
crest on the supraoccipital. In these features P. brevirostris and P. or nat us display a trend
towards the derived Nanochromis condition.
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Final resolution of the phylogenetic status of the genus Parananochromis (and indeed
the other chromidotilapiine genera currently lacking phylogenetic diagnoses) must await a
thoroughgoing phylogenetic analysis, a task that is beyond the scope of the current paper.
For the time being, and as a pragmatic approach for the purposes of species description,
we have assigned the three new species to Parananochromis, while alerting readers to the
possibility that generic reassignment may be necessary once a fuller understanding of the
generic limits and relationships among chromidotilapiines is gained.
Comparative material
Parananochromis caudifasciatus: All from Cameroon: BMNH 1904.7.1.244-249, syn-
types, 2 males, 3 females, 1 undet., 21.94-79.9 mm SL, Nyong system, small streams near
Nyong river, G.L.Bates. – BMNH 1908.7.28.185-190, syntypes, 7 undet., 34.5-71.4 mm
SL, Ja river at Bitye, G.L.Bates. – BMNH 1909.7.9.79-80, syntypes, 2 females 52.2+63.8
mm SL, Bumba river at Assobun, G.L.Bates. – BMNH 1909.7.9.81-82, syntypes, 2 undet.,
72.3+73.8 mm SL, Ja river at Efayong, G.L.Bates. – BMNH 1913.10.29.28-30, syntypes, 2
males, 1 undet., 26.1-82.0 mm SL, Nyong river, G.L.Bates. – BMNH1914.5.27.16, syn-
type, 1 undet., 58.1 mm SL, Nyong river at Akondinga, G.L.Bates. – MRAC 73-03-P-11-
17, 2 males, 5 undet., 25.9-63.1 mm SL, riv. Nyong, environ de Nbalmayo d´Ekombitye,
N. Leontont 1965. – MRAC 76-14-P-681-684, 4 males, 43.1-60.0 mm SL, Mieri, riv.
Doume, 04°14´N, 13°58´E M. Louette & F. Duylaert, Jul 1976.
Parananochromis gabonicus: All from Gabon: BMNH 1967.10.12.57, holotype,
female, 46.6 mm SL; Medouneu road from Mitzic to Medouneu, 15 km from Mitzic, Cam-
bridge expedition, 1957. – CU 80737, 9 males, 3 females, 39.2-74.2 mm SL, Woleu-Ntem,
swampy stream crossing road, 1°32´N, 11°45´E, J.P.Friel, S.Lavone & J.P.Sullivan, Aug
1999. – NMW 94628, 2 males, 2 undet., 33.6-44.8 mm SL, Road Mitzic-Sam, little creek
5 km from Mitzic, 0°49´N, 11°29´E, A. Lamboj, R. Guggenbühl, P. Sewer & A. Weissen-
bacher, Aug 1995.
Paranaochromis longirostris: BMNH 1903.7.28.77-83, syntypes, 7 undet., 63.8-87-1
mm SL; Cameroon : Kribi river at Kribi, G.L.Bates. – MRAC 16881, 1 males, 102.6 mm
SL, cleared and stained, Cameroon : Ntem. – MRAC 173288-289, 2 females, 65.1+73.5
mm SL, Rio Muni: Riv. N´Bo, affl. du Ntem, B.Roman, Aug 1967. – MRAC73-18-P-
3359-364, 72.5-97.0 mm SL, Cameroon: riv. Kom, afll. Ntem, D.E.F.Thys van den Aude-
naerde, Apr 1973. – NMW 94629, 1 male, 2 females, 1 undet., Gabon: Riv. Ngnabegle at
village of Ebiegne, 0°35´N, 12°43´E, A. Lamboj, R. Guggenbühl, P. Sewer & A. Weissen-
bacher, Jul 1995.
Nanochromis squamiceps: AMNH 6079, 2 males, 2 females, 3 undet., 30.0-50.0mm
SL, Stanleyville, Congo, Lang-Chapin Expedition, May 1915.
Nanochromis dimidiatus: AMNH 8150, 2 females, 29.5-27.0mm SL, Ubangui River at
Bangui, A. Baudon, Oct. 1919 – MRAC 89-43-P-3413-414, Riviere Kalukuluka, left bank
Km 70 ancienne route Kisangani-Ubundu, Zaire, L. DeVos-A. Vandelannoote, Feb. 1989.
LAMBOJ & STIASSNY
18 © 2003 Magnolia Press
209
ZOOTAXA Acknowledgements
Our thanks to the following persons and institutions for help with loan of material, assis-
tance with collecting permits and in-country logistics, and for helpful discussion: G.G.Teu-
gels, J. Snoeks, M. Parrent (MRAC), D. Rodriguez, B. Brown (AMNH), C. Hopkins, John
Friel (CU), E. Mikschi, R. Wellendorf (NMW), O. Crimmen and A. Gill (BMNH), H.
Bourobou-Bourobou, F. Obame, P. Posso (Institut de Recherche en Ecologie Tropicale,
Libreville), D. Altmann, H. Kratochvil, G. Okorn, A. Weissenbacher (University of
Vienna), R. Riehl (University of Düsseldorf), F. Panholzer (Graz), M. Hasselmann (Ber-
lin), J. Maclaine (BMNH), R. Guggenbühl (St. Gallen), P. Sewer (Zürich), F. Bitter
(Geeste), U. Schliewen (Munich). Thanks also to Marisol Rosa-Shapiro (student intern,
Hunter College High School, New York) for help with assembling specimen data. Funds
from the Axelrod Reseach Curatorship supported a research visit of AL to the AMNH, and
provide ongoing support to MLJS.
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