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The species-level taxonomy of all extant members of the family Polypteridae is revised. Two genera are recognised: Polypterus and the monotypic Erpetoichthys. Thirteen species of Polypterus are regarded as valid: P. bichir (type species), P. ansorgii, P. congicus, P. delhezi, P. endlicherii, P. mokelembembe, P. ornatipinnis, P. palmas, P. polli, P. retropinnis, P. senegalus, P. teugelsi and P. weeksii. Polypterus lapradei and P. bichir katangae are considered junior synonyms of P. bichir. Polypterus senegalus meridionalis is synonymized with P. senegalus, and P. buettikoferi and P. retropinnis lowei are regarded as junior synonyms of P. palmas. Lectotypes of P. ansorgii, P. congicus and P. delhezi are designated. Distribution maps for each species are compiled and a key for all species of Polypteridae is provided.
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Ichthyological Exploration of Freshwaters/IEF-1094/pp. 1-96 Published 23 July 2019
LSID: http://zoobank.org/urn:lsid:zoobank.org:pub:9FEAF636-6AA5-49B0-BEA4-2CCD75EBE13E
DOI: http://doi.org/10.23788/IEF-1094
Ichthyol. Explor. Freshwaters – ISSN 0936-9902 (print)
© 2019 by Verlag Dr. Friedrich Pfeil, München, Germany www.pfeil-verlag.de
Revision of the extant Polypteridae
(Actinopterygii: Cladistia)
Timo Moritz* and Ralf Britz**
The species-level taxonomy of all extant members of the family Polypteridae is revised. Two genera are recog-
nised: Polypterus and the monotypic Erpetoichthys. Thirteen species of Polypterus are regarded as valid: P. bichir
(type species), P. ansorgii, P. congicus, P. delhezi, P. endlicherii, P. mokelembembe, P. ornatipinnis, P. palmas, P. polli,
P. retropinnis, P. senegalus, P. teugelsi and P. weeksii. Polypterus lapradei and P. bichir katangae are considered junior
synonyms of P. bichir. Polypterus senegalus meridionalis is synonymized with P. senegalus, and P. buettikoferi and P. re-
tropinnis lowei are regarded as junior synonyms of P. palmas. Lectotypes of P. ansorgii, P. congicus and P. delhezi are
designated. Distribution maps for each species are compiled and a key for all species of Polypteridae is provided.
* Deutsches Meeresmuseum, Katharinenberg 14-20, 18439 Stralsund, Germany; and Institute of Systematic
Zoology and Evolutionary Biology, Friedrich-Schiller-University Jena, Erbertstr. 1, 07743 Jena, Germany.
E-mail: timo.moritz@outlook.com (corresponding author)
** Department of Life Sciences, The Natural History Museum, Cromwell Road, London, SW7 5BD, United
Kingdom. E-mail: r.britz@nhm.ac.uk
Introduction
Since their discovery, polypterids continue to puz-
zle scientists. Due to their phylogenetic position as
the most basal extant actinopterygians (Patterson,
1982; Min & Schultze, 2001), polypterids have at-
tracted a lot of scientific attention (see Gosse, 1984
for literature until 1983). Polypterids show several
character states that are considered primitive for
actinopterygians or even bony fishes, e. g. a pair
of true lungs, a body armour of thick ganoid
scales (Sire, 1995), a holoblastic egg development,
(Kerr, 1907; Bartsch et al., 1997) and functional
spiracles (Magid, 1966). They also show many
autapomorphies most of them not known from
any other fish group, fossil or extant, like separate
dorsal finlets, cartilaginously preformed epicen-
tral intermusculars that articulate with the ganoid
scales distally (Britz & Bartsch, 2003), a highly-
specialized anatomy of the pectoral fin (Jessen,
1973), a reduced number of gill arches (Britz &
Johnson, 2003), convergent in some anguilliforms
(Nelson, 1966), and a unique sexually dimorphic
anal fin associated with a unique mating behavior
(Holden, 1971; Britz & Bartsch, 1998).
Despite the numerous papers that have fo-
cussed on bichirs in the past, the species-level
taxonomy of polypterids is still poorly studied.
The number of nominal valid species of poly-
pterids is inconsistently cited as 9 species and 5
sub-species (= 14 taxonomic units in Poll, 1941a),
9 species and 6 sub-species (= 15 taxonomic units
in Gosse, 1984), 18 species of which 5 supposedly
represent undescribed species (Schäfer, 2004), or
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Moritz & Britz: Revision of Polypterus
10-15 species (Britz, 2004). The major reason for
the inconsistency in species numbers is the un-
resolved nomenclatural status of several species-
group names. The goal of the present study is a
species-level revision of the extant Polypteridae
in order to provide a comprehensive taxonomic
framework for future studies on extant members
of this family.
Material and methods
Studied specimens are listed under each species.
The material is divided into two categories: Mate-
rial examined contains all specimens for which
meristic information and/or measurements were
incorporated into the study. Additional mate-
rial contains lots, which have been verified for
their species identity and of which their locality
information has been used for the compilation of
distribution maps.
Measurements used are listed below and
illustrated in Figure 1. All measurements were
taken point to point with digital callipers (CD-
20DCX Absolute Digimatic, Mitutoyo) to the
nearest 0.1 mm for all distances up to 160 mm.
Any measurements above this value were taken
with rulers to the nearest mm. In the case of dis-
torted specimens we used a non-elastic cord to
record the distance between landmarks and then
measured this distance on a ruler.
The following abbreviations have been used
for morphometric and meristic data: AFB, length
of anal-fin base; AFL, length of longest ray in
anal fin; AR, number of anal-fin rays; BD, body
depth at tip of pectoral fin; BW, body width at
tip of pectoral fin; CPL, caudal-peduncle length:
distance between end of anal fin and ventral
origin of caudal fin; D1L, length of first dorsal-
fin spine; D1W, width of first dorsal-fin spine
at its base; D2L, length of second dorsal-fin
spine; DCR, number of soft rays in confluent
dorsal and caudal fin; DFL, length of spinous
part of dorsal fin: distance between base of first
dorsal-fin spine and base of first dorsal-fin soft
ray; DisPD, distance from distal tip of adducted
pectoral fin to first finlet (if first finlet is positioned
in front of tip of pectoral fin, measurement will
have negative value); DS, number of dorsal-fin
spines, including separate finlets and posterior-
most spine connected to soft-rayed portion of
dorsal fin; EyD, eye diameter; GuL, gular length:
distance between tip of lower jaw along mid-
ventral line and caudal most extension of both
gularia; HL, head length: distance between tip
of snout and posterior border of opercle (not in-
EyD
prePL
NOD
preVL
preAL
AFL
GuL
SnL PoL
HL
SnPL
preDL
SL
DFL
TL
DistPD
AFB
BD CPL
IOW HW D1W IPW
PBW
D1L
D2L
AFB
BD CPL
IOW HW D1W IPW
PBW
D1L
D2L
EyD
prePL
NOD
preVL
preAL
AFL
GuL
SnL PoL
HL
SnPL
preDL
SL
DFL
TL
DistPD
AFB
BD CPL
IOW HW D1W IPW
PBW
D1L
D2L
AFB
BD CPL
IOW HW D1W IPW
PBW
D1L
D2L
Fig. 1. Representation of measurements used in this study. For abbreviations see text. Based on the figure of
Polypterus senegalus from Boulenger (1907).
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Ichthyol. Explor. Freshwaters, IEF-1094
cluding fleshy opercular membrane); HW, head
width at anterior border of opercle; IoW, inter-
orbital width: distance between upper margins
of orbitae; IPW, inter-pectoral width: distance
between ventral bases of left and right pectoral
fins measured in ventral view; NoD, nostril
distance at base of nasal tentacle; PBW, width of
pectoral-fin base directly at insertion; PoL, post-
orbital length: distance from posterior border of
orbit to posterior border of opercle; PR, number
of pectoral-fin rays; preAL, pre-anal length:
distance between tip of snout and origin of anal
fin; preDL, pre-dorsal length: distance between
tip of snout and first dorsal-fin spine; prePL,
pre-pectoral length: distance between tip of snout
and base of pectoral fin at insertion; preVL, pre-
pelvic length: distance between tip of snout and
base of pelvic fin at insertion; PvR, number of
pelvic-fin rays; ScB, scales around body at tip of
pectoral fin; ScpD, pre-dorsal scales; ScL, scales
in longitudinal series; ScpP, pre-pelvic scales
(including interpelvic scale); SL, standard length:
distance between tip of snout and posterior most
body scale; SnL, snout length: distance between
tip of snout and anterior margin of orbit; SnPL,
extension of pectoral fin: distance between tip of
snout and posterior end of adducted pectoral fin
placed flat on the body; TL, total length: from tip
of snout to tip of caudal fin; TVer, total number of
vertebrae, only vertebrae with an ossified centrum
are counted here, so that the actual first vertebra
which has no centrum (Britz & Johnson, 2010) is
not counted.
In addition to meristic and morphometric dif-
ferences species of the Polypteridae can also be
distinguished by their color pattern. The recurring
elements of the color pattern are (Fig. 2): pectoral-
fin base blotch: a well-defined dark brown to
black blotch laterally in the distal portion of the
pectoral-fin base; dorsal blotches: a series of black
markings, several scale rows in width, usually
reaching across dorsum to other side forming
saddle-like blotches; lateral blotches: a series
of midlateral dark markings extending several
scale rows in width; dorsolateral bars: formed
by confluence of dorsal and lateral blotches, ei-
ther directed obliquely anteriorly (when dorsal
blotch is confluent with preceding lateral blotch),
or directed obliquely posteriorly (when dorsal
blotch is confluent with succeeding lateral blotch);
sometimes expressed as x/y-shaped bars (when
dorsal blotches confluent with both preceding
and succeeding lateral blotches); ventrolateral
markings: dark brown to black markings along
the ventrolateral sides of the body, often expressed
as series of dark spots, one scale wide and form-
ing irregular interrupted lines (as in some color
morphs of P. bichir or in P. ansorgii) or irregular
speckling of ventrolateral body; sometimes also
forming larger blotches that may be confluent
with lateral blotches (as in P. endlicherii); subor-
bital stripe: an oblique dark brown to black stripe
below the eye extending from the posterior base
of the nasal tentacle to the margin of the jaw ar-
ticulation or sometimes expressed only in part of
this distance (as in the postorbital area in P. polli)
and/or expressed as a line of separate spots rather
than as a continuous stripe (as in P. retropinnis);
lip markings: brown to black markings of varying
shape and size, can be expressed as vermicula-
tions on both lips, as peppering with tiny spots,
as a series of bar-like markings or a combination
of all; dorsal and lateral head markings: brown
to black or light beige markings of varying shape
and size, can be expressed as vermiculations, spots
or blotches.
Distribution data of all checked museum
specimens were critically evaluated and, when
found to be lacking precision, were given ap-
proximated GPS coordinates. Locality information
that was considered too general, such as River
Nile, Senegal or Congo was not included.
dorsal head markings
lip markings
suborbital stripe
pectoral-fin base blotch lateral blotches ventrolateral markings
dorsal blotches
dorsolateral bars
lateral head markings
Fig. 2. Overview of principal color patterns in Polypterus.
4
The distribution data were complemented by
information obtained by TM during several field
trips. In addition, museum and internet databases,
i. e. Fishbase (Froese & Pauly, 2016) and fishnet2
(www.fishnet2.net), were checked for distribution
data, but this information was critically evaluated
and questionable records were not included. Last-
ly, numerous published sources of distributional
information were evaluated, [e. g. Pellegrin (1900),
Blanc & Daget (1957), Paugy & Bénech (1989), Ro-
man (1966), Moritz (2010), Neumann et al. (2016)].
Locality records are grouped into four categories
(1) type locality, (2) verified records, i. e. the spe-
cies identity of the respective specimens has been
checked and confirmed in this study, (3) other
records, i. e. records from literature and database,
for which specimens could not be checked, but
for which locality records are plausible, and (4)
questionable records; i. e. records that could not
be verified but that seem to be based on correct
identifications, but are considered questionable in
the context of the other records. Doubtful records,
i. e. records that are almost certainly based on
misidentifications have been excluded from the
maps, but discussed for each respective species, to
produce the most comprehensive, up to date dis-
tributional information for the different species.
Data were collected in tables from which maps
were created with the freeware QGIS 2.12.3 (www.
qgis.org) using freely available shape files from
FAO, as e. g. Rivers of Africa and Inland Water
Bodies of Africa (www.fao.org/geonetwork).
Simple diagrams and box-plots were created
directly in table calculation software EXCEL
(Microsoft Corp.) or CALC (OpenOffice 4.1.1,
www.openoffic.org); statistical calculations such
as Principal Components Analyses (PCAs) were
conducted in PAST 3.11 (Hammer et al., 2001).
Graphics including figures were either created
or at least finalized with the software GIMP 2.8
(www.gimp.org). For Polypterus palmas several
characters were analyzed and displayed accord-
ing to the river system where the specimens were
collected, following a North to South sequence of
the respective deltas of the basins.
The following institutional abbreviations have
been used: BMNH, The Natural History Muse-
um, London; DMM, Deutsches Meeresmuseum,
Stralsund; IRSNB, Institut Royal des Sciences Na-
turelles de Belgique; MNHN, Muséum national
dHistoire naturelle, Paris; MRAC, Musée Royal
de lAfrique Centrale, Tervuren; NMSZ, National
Museums Scotland, Edinburgh; NMW, Naturhis-
torisches Museum, Vienna; RMNH, Naturalis
Biodiversity Centre, Leiden; USNM, National
Museum of Natural History, Washington, D.C.;
ZSM, Bavarian State Collection of Zoology, Mu-
nich.
Family Polypteridae Bonaparte, 1835
Polypterini Bonaparte, 1835: 8.
Type genus. Polypterus Lacépède, 1803 (see re-
marks under Polypterus below).
Diagnosis. Polypterids are distinguished from
all other living actinopterygians by the presence
of complete ganoid scales covering the entire
body, a unique skeletal anatomy of the pectoral
fin, first part of dorsal fin separated into finlets,
i. e. sharp spines with secondary supporting rays
and attached membrane, as well as paired lungs
with ventral connection to oesophagus, a modi-
fied anal fin in males, adhesive eggs and larval
stages with external opercular gills.
Descriptive synopsis. Two types of body shapes
among polypterids: elongated, cylindrical in
Polypterus (body depth 7.0-16.5 % SL) and an-
guilliform in Erpetoichthys (body depth 2.9-5.2 %
SL); body in both types round in cross section at
anterior third and laterally flattened at posterior
third. Head in cross-section round or moder-
ately to strongly dorso-ventrally flattened. Body
completely covered with fully developed ganoid
scales consisting of three layers: bone, dentin and
ganoin (Goodrich, 1907; Sire, 1995). Anterior part
of dorsal fin separated into 5-18 finlets, i. e. single
spines with sharp bilaterally paired distal tips
with posterior fin membrane supported by soft
rays originating from posterior margin of spine
perpendicular to spine body. Posterior part of
dorsal fin confluent with caudal fin. Pectoral-fin
skeleton with unique pro-, meso- and meta-
pterygium between pectoral girdle and pectoral
radials. Sexually dimorphic anal fin: fin of males
greatly enlarged mainly as result of anteropos-
terior widening of anal-fin rays; number of
anal-fin rays on average only slightly higher in
males of Polypterus, but clearly higher in males
of Erpetoichthys.
Mouth large, with posterior angle situated
behind vertical through posterior margin of orbit;
lips thick, supported by labial cartilage. Anterior
Moritz & Britz: Revision of Polypterus
5
nostril extended as tube well beyond tip of snout.
Paired gular plates present. Interopercle absent.
Series of spiracular plates between lateral margin
of skull roof and opercular series, with some of
them covering the functional spiraculum (Bartsch,
1997; Claeson et al., 2007). Dermohyal present.
Maxilla immovably attached to skull (akinetic
upper jaw). Fifth gill arch absent and lower phar-
yngeal tooth plates associated with fourth cerato-
branchial (Britz & Johnson, 2003); pseudobranch
absent. Juveniles with well-developed external
opercular gills. Mesocoracoid absent; clavicles
well developed. Unique epicentral intermuscular
bones with cartilaginous tip articulating with
lateral line scales (Britz & Bartsch, 2003). Intestine
with spiral valve. Caudal fin rounded or pointed
(especially in juveniles) confluent with posterior
dorsal fin; posterior most vertebrae smaller than
preceding ones.
Biology. For most species information on ecology
and life history is scarce. From the available and
published information it seems that species are
quite similar to each other in various aspects. All
are predominantly nocturnal and strictly fresh-
water species. They are predators, apparently
mostly guided by their sense of smell (Pfeiffer,
1968, 1969), feeding on aquatic and terrestrial
invertebrates and fish, sometimes amphibians,
with prey size mainly depending on the size of
the mouth cleft of the respective specimen. Some
plant material has occasionally been reported
from stomach contents (e. g. Daget, 1948), but was
likely ingested accidentally. Polypterus species
are able to prey on fishes of more than half their
size (pers. obs. TM). Prey can be sucked in with
force and speed by enlarging the mouth cavity
and projecting the cartilage-supported labial folds
to form a funnel (Bartsch, 1997). All polypterids
show sexual dimorphism in their anal fin, which
is important for courtship: during spawning, the
male forms a cup with its anal fin into which
the female releases her eggs. Eggs are then ferti-
lized in this cup before being scattered by rapid
lateral movements of the caudal and anal fins of
the male (Bartsch & Britz, 1996; Britz & Bartsch,
1998). The anal fin of males does not function
as a copulatory organ as initially hypothesized
by Harrington (1899). In nature, spawning takes
place in (densely) vegetated areas, usually flood
plains (Budgett, 1900; Daget, 1948). The eggs are
relatively large and adhesive (Arnoult, 1961, 1964;
Bartsch & Britz, 1996) with a single micropyle
(Britz & Bartsch, 1998). The larval development
was described in detail for P. senegalus (Kerr, 1907;
Bartsch et al., 1997; Schugardt & Kirschbaum,
2006), and for P. ornatipinnis and E. calabaricus
(Britz & Bartsch, 1998). When larvae start to
feed in captivity, they wait for prey rather than
actively search for it (Bartsch, 1997). Juveniles
have external gills (Boulenger, 1902b), which may
persist for a long time up to even larger sizes in
some species (e. g. large external gills were still
present in a 335 mm SL specimen of P. weeksii
[MRAC A7-033-P-0028], a 334 mm SL specimen
of P. bichir [BMNH 1900.6.28.9], a 315 mm SL
specimen of P. congicus [IRSNB 9960], and in
a 279 mm SL specimen of P. endlicherii [NMW
63218]). In captivity larvae and juveniles initially
show fast growth up to a certain, species-specific,
size but subsequently growth rates slow down
(Daget et al., 1965). At least in some species sexual
maturity may be reached in one year (Svensson,
1933). The lungs of bichirs are paired and well
developed and vascularized and atmospheric air
is regularly taken in for respiration. Air uptake
may be realized in two different ways: inhaling
air via the mouth, or via the spiracles (Budgett,
1900; Bartsch, 1997).
Remarks. The oldest available family group name
for polypterids dates back to Bonaparte (1835: 8)
who listed Subfamilia 36. Polypterini in his
Prodromus Systematis Ichthyologiae.
Genus Erpetoichthys Smith, 1865
Erpetoichthys Smith, 1865a: 2.
Calamichthys Smith, 1866a: 113.
Calamoichthys Smith, 1866a: 112.
Type species. Erpetoichthys calabaricus Smith, 1865.
Diagnosis. Erpetoichthys is distinguished from
all other members of the family by absence
of pelvic fin and subopercle (vs. presence in
Polypterus), body depth 2.9-5.2 % SL (vs. 7.0-16.5
in Polypterus), 109-115 vertebrae (vs. fewer than
67 in Polypterus), short head (7.2-10.5 % SL vs.
14.4-26.8 in Polypterus), and 16-21 pectoral fin-
rays (vs. 24-48 in Polypterus).
Descriptive synopsis. Body greatly elongated,
anguilliform with 109-115 vertebrae and 102-115
scales in lateral series. Sexual dimorphism in anal
Ichthyol. Explor. Freshwaters, IEF-1094
6
fin much pronounced: males with 11-14 (mean
13) and female with 8-13 (mean 10) anal-fin rays;
each ray in mature males widened along its an-
teroposterior axis. For meristic and morphometric
data refer to Tables 1-2.
Etymology. Erpetoichthys is derived from the
Greek ρπετ)+, reptile or snake; and θς, fish.
Remarks. Confusion about the valid name of
the genus arose from multiple descriptions by
Smith (1865a-b; 1866a-c) and from a subsequent
change of the generic name by Smith one year later
(1866a, c), because he erroneously considered the
original name Erpetoichthys to be preoccupied. De-
tails on this nomenclatural issue were discussed
by Swinney & Heppell (1982) and Rizzato &
Bockmann (2017) with contradicting results. We
follow Swinney & Heppells (1982) view and
consider the name Erpetoichthys calabaricus to be
available from Smith (1865a). This is in contrast
to the recent conclusion by Rizzato & Bockmann
(2017) which rests solely on their interpretation of
Art. 8.1.1 of the Internacional Code of Zoological
Nomenclature (ICZN, 1999; hereafter treated as
the Code). Rizzato & Bockmann (2017) argued
that publication of Smiths (1865a) name in the
newspaper Daily Review was not issued for
the purpose of providing a public and permanent
scientific record as the Art. 8.1.1. requires for a
name to be available. The authors argue in detail,
that First, and most importantly, the report was
clearly intended solely to publicize a scientific
breakthrough for the general public of Edinburgh,
not to name the fish according to formal scientific
standards. While Rizzatto & Bockmann (2017)
present this argument as a fact, we consider this
to be speculation, as the intention of the author
is unknowable. Rizzatto & Bockmanns (2017)
main conclusion that Smiths (1865a) name is not
available centers very much around the intention
of a given author and they mention intention,
intend, intentionally, unintentionally 21 times
in their lengthy paper. However, according to
the Code it is not the intention that is decisive.
While it cannot be objectively decided whether
Smiths (1865a) first publication of the name was
issued for the purpose of providing a public
and permanent scientific record, we argue that
publication of the name Erpetoichthys calabaricus in
the Daily Review in 1865, which is still available
today, in conjunction with a diagnosis, makes
this name available, as concluded previously by
Swinney & Heppell (1982). We therefore reject
Rizzatto & Bockmanns (2017) conclusion and
retain Erpetoichthys calabaricus as the valid name
for the rope eel. Furthermore, following Rizzatto
& Bockmanns (2017) rationale would also greatly
destabilize nomenclature generally, as numerous
nomenclatural acts in the 1800s and early 1900s
were published in minutes and proceedings of
scientific sessions before the more comprehensive,
actual paper was published. Fricke et al. (2018)
also decided not follow the conclusions of Rizzato
& Bockmann (2018).
Erpetoichthys calabaricus Smith, 1865
(Figs. 3-5; Tables 1-2)
Erpetoichthys calabaricus Smith, 1865a: 2.
Erpetoichthys robbianus Smith, 1865b: 2.
Polypterus erpetoideus Smith, 1865a: 2.
Material examined. 32 specimens. Ouém é Basi n:
RMNH 24896, 1, 187 mm SL; Benin: Cotonou, L. B.
Holthuis & H. Hoestland, 1963. niger Delta: RMNH
10327, 1, 313 mm SL; Nigeria: Warri, Diergaarde Rot-
terdam, 20 Nov 1918. – RMNH 8858, 2, 293-318 mm
SL; Nigeria: Warri, J. Büttikofer, Oct 1912. – BMNH
1977.11.8.1-4, 4, 262-300 mm SL; Nigeria: Lagos: En-
ergo project site near Lagos, D. Thomas. – MNHN
2008-2103, 9, 252-307 mm SL; Nigeria: Port Harcourt
[market]; T. Roberts, 5 Nov 1987. CrOss river Basin:
MNHN 0000-4599, 2, 223-301 mm SL; Nigeria: Old
Calabar, J. A. Smith, before 1867. – NMW 22831, 1,
147 mm SL; Nigeria: Old Calabar, 1874. – NMW 22832,
1, 162 mm SL; Nigeria: Old Calabar, 1874. – NMW 50282,
1, 343 mm SL; Nigeria: Old Calabar. – NMW 63194, 1,
205 mm SL; Nigeria: Old Calabar, 1874. – NMW 63195,
2, 247-320 mm SL; Nigeria: Old Calabar, 1877. limBe
river Basin: MNHN 1900-0218, 1, 180 mm SL; Cam-
eroon: Victoria, Lennier. sanaga river Basin: MNHN
1978-0732, 1, 313 mm SL; Cameroon: Sanaga River at
Mouanko, Depierre, 1978. – MRAC 73-002-P-0001-0003,
3, 271-286 mm SL; Cameroon: Sanaga River at Edea
[below waterfalls], D. F. E. Thys van den Audenaerde,
15-16 Oct 1964. – MRAC 75-005-P-4758, 1, 305 mm SL;
Cameroon: km 5 on the road Douala to Edea, D. F. E.
Thys van den Audenaerde, 21 Oct 1966. WOuri river
Basin: MNHN 1928-0080, 1, 321 mm SL; Cameroun:
Vouri [= Wouri] River at Nono, Monod.
Additional material. Ouémé Basin: DMM IE/10401, 5,
Benin: Zou: Lokoli swamp forest, 7°03'40" N 2°15'32" E;
T. Moritz, 16 Jan 2005. iguiDi river Basin: DMM
IE/14378, 2, Benin: Iguidi River at Igolo, 6°39'18" N
2°42'43" E; T. Moritz & V. von Vietinghoff, 11 May 2005.
– DMM IE/14388, 1, Benin: Iguidi River at Tchakou,
Moritz & Britz: Revision of Polypterus
7
Fig. 3. Erpetoichthys calabaricus, NMW 63195:1, 320 mm SL, male, non-type, Nigeria: Old Calabar [type locality];
1877.
a
b
cc
Fig. 4. Erpetoichthys calabaricus, a-b, ~280 mm SL; Nigeria: aquarium import; c-d, ~180 mm SL; Benin: Ouémé
Basin: Lokoli swamp forest.
6°28'27" N 2°42'48" E; T. Moritz, 22 Mar 2007. CrOss
river Basin: MNHN 1988-0253, 1, Nigeria, Cross River
at Uyo, King, 1987.
Diagnosis. Same as presented for the genus.
Descriptive synopsis. Meristic and morpho-
metric data in Tables 1-2. Body anguilliform
(body depth 18-34 times in SL); body circular
in cross section along most of body; but later-
ally compressed in posterior most part towards
Ichthyol. Explor. Freshwaters, IEF-1094
8
caudal peduncle. Mouth subterminal with upper
jaw strongly projecting beyond lower jaw; head
only slightly depressed. Maximum observed size
343 mm SL (355 mm TL). Occasionally reported
maximum sizes of up to 900 mm (e. g. Terofal,
1970; Gosse, 1984) are erroneous.
Coloration. In life: dorsal half of body brown-
yellowish with borders of vertical scale rows
darker; no additional markings (Fig. 4). Ventral
half of body yellowish-orange. No clear border
between upper and lower half of flank colora-
tion. Head coloration same as that on body, but
nasal tentacles lighter; suborbital stripe absent.
Table 2. Meristic characters of Erpetoichthys calabaricus. Numbers in brackets indicate number of specimens with
the respective value.
all material
Dorsal-fin spines 9 (8), 10 (11), 11 (9)
Dorsal- and caudal-fin rays 10 (1), 12 (2), 13 (7), 14 (10), 15 (6), 16 (1)
Pectoral-fin rays 16 (3), 17 (3), 18 (6), 19 (10), 20 (4), 21 (1)
Anal-fin rays 8 (3), 9 (4), 10 (3), 11 (6), 12 (4), 13 (4), 14 (3)
Scales in longitudinal series 102 (1), 104 (1), 105 (1), 106 (3), 107 (5), 108 (1), 109 (8), 110 (4), 111 (1), 112 (1), 113 (1),
114 (1)
Scales around body 27 (3), 28 (3), 29 (5), 30 (10), 31 (5), 32 (1), 33 (1)
Pre-dorsal scales 44 (1), 48 (1), 49 (5), 50 (2), 51 (5), 52 (2), 53 (4), 54 (2), 55 (2), 56 (1), 57 (1), 58 (1), 59 (1)
Total number of vertebrae 109 (1), 110 (1), 112 (2), 113 (1), 115 (1)
Table 1. Morphometric characters of Erpetoichthys calabaricus. SD: standard deviation
range mean SD n
Standard length [mm] 147-343 266.5 51.2 28
Total length [mm] 171-355 281.2 47.5 27
In percent of standard length
Body depth 2.9-5.2 4.2 0.7 28
Body width 3.5-5.5 4.6 0.5 28
Head length 7.2-10.5 8.1 0.7 28
Pre-dorsal length 41.0-54.2 47.5 3.1 28
Dorsal-fin length 34.9-53.9 47.8 4.0 28
Pectoral to dorsal fin 28.6-40.7 34.6 2.8 26
Pre-pectoral length 7.8-10.7 8.8 0.7 28
Pectoral-fin extension 10.6-19.3 12.8 1.6 26
Inter-pectoral width 2.8-4.1 3.6 0.3 28
Pectoral-fin base width 0.8-1.5 1.2 0.2 28
Pre-anal length 93.0-95.1 94.1 0.5 28
Anal-fin ray length 3.6-5.5 4.6 0.5 27
Anal-fin base length 1.2-3.6 2.8 0.5 27
Caudal-peduncle length 0.8-1.6 1.1 0.2 27
In percent of head length
Head width 49.1-68.0 58.5 5.7 28
Interorbital width 21.6-28.5 25.4 1.8 28
Nostril distance 11.2-19.6 15.3 2.2 28
Snout length 21.9-26.9 24.8 1.3 28
Eye diameter 8.8-16.3 12.0 1.6 28
Postorbital length 60.4-65.2 63.3 1.2 28
Gular length 54.5-63.3 59.0 2.1 28
Length of first spine 8.8-22.5 17.0 2.9 28
Length of second spine 6.7-25.9 17.6 3.9 28
Width : length of first spine 24.3-51.2 35.2 7.1 28
Length of second : first spine 70.2-143.2 103.9 19.0 28
Moritz & Britz: Revision of Polypterus
9
Iris orange, brownish or grey; lips of upper and
lower jaw yellowish-orange without any mark-
ings. Pectoral fin yellowish-orange, at its proximal
part with black blotch extending onto distalmost
part of pectoral-fin base, leaving most of pectoral-
fin base without markings. Soft part of dorsal fin
and caudal fin brown, grey or olive with yellow-
orange fin membranes. Dorsal-fin spines brown,
grey or olive, their distal part orange; membrane
of finlets transparent or slightly yellowish with
only very few small dark spots.
In preservative: dorsal half of body and head
brown or brownish-grey (Fig. 3); ventral half of
body and head including lips and nasal tenta-
cles pale beige or brownish, sometimes slightly
orange-brown.
Distribution. From the Ouémé Basin in Benin,
across the Niger Delta in Nigeria to the Sanaga
River in Cameroon along West Africas coastal
basins (Fig. 4). A single record from the Chiloango
River (= Rio Hi) in the Cabinda region of Angola
mentioned in Boulenger (1909: 18) seems ques-
tionable; the species has not been reported since
from that area (Boulenger, 1912; Pellegrin, 1928;
Daget & Stauch, 1968).
Biology. Prefers small streams and swamps
(Smith, 1866a). Feeds on terrestrial insects, e. g.
termites (Smith, 1866c), insect larvae and aquatic
invertebrates. Reproduction in captivity and
early development were described by Britz &
Bartsch (1998).
Etymology. Smith (1865a) used calabaricus to refer
to . . . the locality [Old Calabar, Nigeria] where
it was discovered.
Remarks. Originally two syntypes were present,
but both were lost (see Swinney & Heppell, 1982).
The original description was based on two poorly
preserved specimens from Old Calabar, Calabar
River and adjacent water bodies, Nigeria, col-
lected by A. Robb on an unspecified date. There
are two specimens in the Paris Museum, MNHN
0000-4599, which are labelled as TYPES, and
listed as paratypes in the museums on-line
collection database (https://science.mnhn.fr/
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 5. Records of Erpetoichthys calabaricus. , type locality; , verified records; , other records; , question-
able records.
Ichthyol. Explor. Freshwaters, IEF-1094
10
19
17
15
13
11
9
7
5
69
64
59
54
49
55
50
45
40
35
30
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
DSScLScB
Fig. 6. Box-and-whisker plots of selected meristic data of Polypterus species. DS, number of dorsal spines;
ScL, number of scales in lateral series; ScB, number of scales around body; ScpD, number of pre-dorsal scales;
ScpP, number of pre-pelvic scales; P.ans, P. ansorgii (n = 7); P.bic, P. bichir (n = 139); P.con, P. congicus (n = 66);
P.end, P. endlicherii (n = 64); P.del, P. delhezi (n = 44); P.mok, P. mokelembembe (n = 26); P.orn, P. ornatipinnis (n = 63);
P.pal, P. palmas (n = 122); P.pol, P. polli (n = 85); P.ret, P. retropinnis (n = 46); P.sen, P. senegalus (n = 226);
P.teu, P. teugelsi (n = 4); P.wek, P. weeksii (n = 45).
institution/mnhn/collection/ic/item/search/
form). Smith sent these specimens to the MNHN
and Duméril (1870) used them for his account of
the species in his Histoire naturelle des poissons.
Bertin (1940) finally listed the MNHN specimens
as paratypes referring to the redescription of
Duméril (1870). Although the specimens were sent
by Smith and come from the type locality they are
not part of the original type series. Smith (1865a)
had only two males, for his species description,
while the two specimens in the MNHN are male
and female; furthermore, the MNHN collection
database provides 1867 as the collection date for
these specimens, thus two years after the descrip-
tion of the species.
Polypterus erpetoideus was proposed as another
name for the species by Smith and reported in the
same newspaper article which served as original
description. This name was not used subsequently
and Swinney & Heppell (1982), as first revisers,
Moritz & Britz: Revision of Polypterus
11
Ichthyol. Explor. Freshwaters, IEF-1094
40
35
30
25
20
15
10
55
50
45
40
35
30
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ScpDScpP
Fig. 6. (continued).
selected Erpetoichthys calabaricus as the valid name
for the taxon.
Smiths (1865b) description of Erpetoichthys
robbianus was based on specimens with a small,
narrower anal fin, which is the condition in fe-
males of the sexually dimorphic anal fin (Traquair,
1866a-b; Smith, 1866a-c) in polypterids. Follow-
ing Swinney & Heppell (1982) NMSZ 1867.18.1;
NMSZ 1979.006.002 and BMNH 1908.1.27.2 pre-
sumably are part of the type series of E robbianus.
Herman et al. (1990) additionally listed NMSZ
1885.052 as part of this type series.
Genus Polypterus Lacèpede, 1803
Polyptere Geoffroy [Saint Hilaire], 1802a: 12, Plate
V.
Polypterus Lacépède, 1803: 340.
Type species. Polypterus bichir (Geoffroy, 1802)
by monotypy.
Diagnosis. Distinguished from Erpetoichthys by
presence of pelvic fin, presence of subopercle,
more pectoral-fin rays (24-48 vs. 16-21) and
comparatively shorter body (body depth 6-12
times in SL vs. 18-29 times) with fewer vertebrae
(67 or fewer vs. 109-115).
Descriptive synopsis. Body elongated to highly
elongated and eel-like, circular in cross section,
but laterally compressed in the postabdominal
part of the body. Polypterus is characterized by
50-70 scales in the lateral series, 10-38 pre-dorsal
scales, 31-54 scales around the body, 34-53 pre-
pelvic scales, 5-18 dorsal-fin spines, 13-24 soft
rays in confluent soft dorsal plus caudal fin, 24-
48 pectoral-fin rays; and pelvic fin present, with
7-17 rays. Figures 6 and 7 summarize selected
meristic and morphometric characters for the
genus. Sexual dimorphism in number of anal-fin
rays less pronounced than in Erpetoichthys: males
on average with one additional ray compared
to females; equally conspicuous enlargement of
anal fin in mature males through antero-posterior
widening of individual anal-fin rays.
Biology. See remarks for Polypteridae.
12
70
60
50
40
30
20
10
40
30
20
10
0
10
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
preDLdistPD
65
60
55
50
45
40
35
30
25
DFL
Fig. 7. Box-and-whisker plots for selected morphometric characters of Polypterus species. preDL, pre-dorsal
length; DFL, dorsal-fin length; distPD, distance between end of pectoral fin and origin of dorsal fin; HL, head
length; IOW, inter-orbital width; SnL, snout length; for other abbreviations and number of specimens see Figure 6.
Etymology. Polypterus is derived from the Greek
πλς – many and πτερ)+ – wing, apparently re-
ferring to the anterior part of the dorsal fin which
comprises multiple separate finlets.
Remarks. The genus name was originally spelled
Polyptère by Geoffroy (1802a). Lacépède (1803)
used this original name but also provided a
Latinized version. Only a few years later also
Geoffroy (1809) used both names, the original
French spelling and the Latinised version in his
Histoire naturelle des Poissons du Nil.
Eschmeyer (1998) originally listed the type
species as Polypterus bichir Geoffroy St. Hilaire
but changed it in the on-line edition of the Catalog
of Fishes (Fricke et al., 2018) to Polypterus bichir
Lacépède, 1803, arguing that the generic name
was not Latinised in Geoffroys (1802a) descrip-
tion. Holly (1933) had come to this conclusion and
Schäfer (2004) shared this same view. Likewise,
Shaw (1804) regarded Geoffroys (1802a) name
as not valid, and even introduced a new species-
name, i. e. P. niloticus, but using at the same time
the Latinised genus of the name given by Geof-
Moritz & Britz: Revision of Polypterus
13
Ichthyol. Explor. Freshwaters, IEF-1094
Fig. 7. (continued).
26
24
22
20
18
16
14
28
26
24
22
20
18
16
14
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
P.ans P.bic P.con P.end P.del P.mok P.orn P.pal P.pol P.ret P.sen P.teu P.wek
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
ı ı ı ı ı ı ı ı ı ı ı ı ı ı
HLSnL
35
30
25
20
15
IOW
froy. The argumentation of the above mentioned
authors, however, is not in congruence with Art.
11.3 of the Code, which clearly states that a name
may be a word in or derived from any language.
Thus Polyptere (without apostrophe following
Arts. 27 and 32.5.2 of the Code) Geoffroy is an
available name and a senior synonym of Polypterus
Lacépède. However, because Polyptere has not
been used as a valid genus name since 1899 (Art.
23.9.1.1 of the Code) we declare Polyptere Geof-
froy, 1802 a nomen oblitum under Art. 23.9.2 of the
Code and the name Polypterus Lacépède a nomen
protectum because the latter has been used in at
least 25 works in the last 50 years by at least 10 dif-
ferent authors (Art. 23.9.1.2 of the Code) (Banister
& Bailey, 1979; Daget & Desoutter, 1983; Gosse,
1984, 1988, 1990, 2003; Bailey, 1994; Hanssens
et al., 1995; Bartsch & Britz, 1996; Seegers, 1996;
Bartsch, 1997; Britz & Bartsch, 1998, 2003; Britz
& Johnson, 2003; Britz, 2004; Lalèyè et al., 2004;
Schäfer, 2004; Poplin & Dutheil, 2005; Schliewen
& Schäfer, 2006; Schugardt & Kirschbaum, 2006;
Claeson et al., 2007; Claeson & Hagadorn, 2008;
Suzuki et al., 2010; Cuervo et al., 2012; Trape, 2013;
Near et al., 2014; Neumann et al., 2016; Grandstaff
et al., 2017; Moritz, 2017).
Thirteen valid species of Polypterus are herein
recognised. Morphologically two groups can be
14
a
b
c
a
b
c
Fig. 9. Changes in head shape during growth in Poly-
pterus endlicherii. a, DMM IE/10399, 102 mm SL; Benin:
Pendjari National Park; b, DMM IE/10393, 150 mm SL;
Benin: Pendjari National Park; c, BMNH 1949.10.20.1,
272 mm SL; Ghana: Volta Basin.
Moritz & Britz: Revision of Polypterus
121 2
3
ab
Fig. 8. Anteriormost scale row running continuously from dorsal to ventral mid-line is a, third row in Polypterus
ansorgii, and b, second row in P. senegalus.
distinguished: the large species group (P. ansor-
gii, P. bichir, P. congicus and P. endlicherii), and the
small species group, including all the remaining
members of the genus. Available molecular data
(Suzuki et al., 2010; Near et al., 2014) support the
monophyly of the large species, a taxon we call
herein the Polypterus bichir group. The small
species apparently do not form a monophyletic
group. The typical character states of the P. bichir
group seem to be rather derived conditions, e. g.
high number of dorsal-fin spines, low number
or pre-dorsal scales (Fig. 6), the third scale row
being the first to continue from dorsal to ventral
midline (vs. second scale row being the first to
continue all the way to the ventral midline; Fig. 8).
Our taxonomic study offers no obvious character
evidence that would support additional clades
among the small polypterid species group sup-
ported by molecular data.
Accurate identification of polypterids is often
complicated due to the similarity in their general
appearance, but also because of changes in body
shape during growth. One example is the project-
ing lower jaw of specimens of Polypterus bichir,
P. congicus and P. endlicherii. This character is often
used in identification keys (e. g. Boulenger, 1898;
Pellegrin, 1923a; Holly, 1933; Poll, 1941a; Gosse,
1990, 2003). In juveniles of these species, however,
jaws are still equal and the mouth is in a terminal
position. This has occasionally resulted in the er-
roneous identifications of juvenile P. endlicherii or
P. bichir as P. ansorgii in collections (e. g. AMNH
215310, USNM 339728) or in publications (e. g., La-
lèyè et al., 2004). Polypterus ansorgii is the smallest
species of the P. bichir group with jaws remaining
almost equal in size throughout life. In general
the head shape of members of the P. bichir group
changes during growth becoming flatter and thus
wider in appearance; this is most pronounced
in P. endlicherii (Fig. 9). A similar flattening of
the head can be seen to a smaller extent in large
specimens of P. weeksii.
Furthermore, the finlet membranes attached
to each dorsal-fin spine are supported by second
order dorsal-fin rays (Fig. 10a), the number of
which is only of limited, if any, diagnostic value
because during growth finlets acquire more of
these secondary rays (Fig. 10b; Sewertzoff, 1924).
As a result, adults of large species of the P. bichir
group seem to have more of these second order
15
a b
Fig. 10. Dorsal-fin spines and secondary rays in live Polypterus delhezi, lateral view. a, juvenile of 72 mm SL; b,
adult 240 mm SL.
dorsal-finlet rays than their juveniles or than
adults of other species.
The position of the first dorsal-fin spine in
relation to the posterior margin of the pectoral fin
is often used (e. g. Gosse, 1990, 2003) to distinguish
between the large and small species, i. e. the
P. bichir group and the remaining Polypterus spe-
cies. Although helpful to identify certain species,
this character does not allow a clear distinction be-
tween the two groups: occasionally in specimens
of P. senegalus the first dorsal-fin spine is situated
directly above the posterior tip of the pectoral fin
and some P. bichir specimens show a clear gap
between the vertical through the posterior margin
of the adducted pectoral fin and that of the base
of the first dorsal-fin spine (Fig. 8). Unusual in
this regard is P. delhezi which usually has its first
dorsal-fin spine anterior to the posterior margin of
the pectoral fin, a condition typical of the P. bichir
group, although other characters are consistent
with its inclusion among the small Polypterus
species. The commonly used number of dorsal-fin
spines and number of scales to characterize species
are often overlapping between species, but there
are still a number of useful meristic and morpho-
metric characters to distinguish poly pterid species
(Fig. 6-7). In general, certain polypterid species
have a somewhat similar morphology and are
best distinguished by differences in coloration.
Polypterus ansorgii Boulenger, 1910
(Figs. 11-13, Tables 3-4)
Polypterus ansorgii Boulenger, 1910: 424.
Material examined. 7 specimens. type speCimens:
BMNH 1910.9.13.4, lectotype (Fig. 11; present designa-
tion), 169 mm SL; likely juvenile female, Guinea-Bissau:
Corobal River at Tchitoli, W. J. Ansorge, Jun 1909. –
NMW 63252: 1-2, 2, paralectotypes, 147-160 mm SL;
collected with the lectotype. géBa river Basin: MRAC
73-005-P-0031, 1, 180 mm SL; Senegal: Geba River Basin:
Kanyanga River at Kounkane, 12°55' N 14°06' W; D. F.
E. Thys van den Audenaerde, 6 Apr 1966. – MRAC
73-005-P-0032-0033, 2, 179-210 mm SL; Senegal: Geba
River Basin: Kanyanga River at Kounkane, 12°55' N
14°06' W; D. F. E. Thys van den Audenaerde, 8 Apr
1966. CasamanCe river Basin: MRAC 73-005-P-0030, 1,
210 mm SL; Senegal: Casamance River at Kolda, 12°53' N
14°57' W; D. F. E. Thys van den Audenaerde, 8 Apr 1966.
Diagnosis. Distinguished from all Polypterus spe-
cies, except P. bichir, P. congicus and P. endlicherii
by 13-15 dorsal finlets (vs. 5-12) and the second
dorsal scale row not reaching ventral midline
(vs. reaching ventral midline). Distinguished
from P. congicus and P. endlicherii by smaller
interorbital width (15.7-18.5 % HL vs. 17.9-
27.4 %) and by coloration consisting of more or
less regular, rectangular lateral blotches usually
not confluent with dorsal saddle blotches (vs.
no separate regular rectangular lateral blotches,
but series of bars formed by confluence of dorsal
blotches with lateral blotches in P. congicus and
with lateral blotches and ventrolateral markings
in P. endlicherii). Polypterus ansorgii differs from
P. bichir by fewer scales in lateral series (54-56
vs. 55-70), fewer scales around body (43-45 vs.
43-54) and smaller adult size (maximum recorded
size 313 mm SL vs. 750); in specimens between
150 and 200 mm SL, external gills usually com-
pletely reduced in P. ansorgii (vs. present in most
specimens of P. bichir). Specimens smaller than
147 mm SL of P. ansorgii have not been available
for this study and it remains unclear, whether
it would be possible to distinguish them from
equally small specimens of P. bichir based on
external characters.
Ichthyol. Explor. Freshwaters, IEF-1094
16
Descriptive synopsis. Meristic and morphomet-
ric data in Tables 3-4. Body moderately elongated.
Jaws terminal, in larger specimens (> 200 mm
SL) lower jaw slightly projecting beyond upper
jaw. Head flattened. Likely the smallest species
among the Polypterus bichir group with the largest
specimens in collections measuring 210 mm SL
(246 mm TL), and unpreserved aquarium speci-
men reaching 313 mm SL (371 mm TL).
Table 3. Morphometric characters of Polypterus ansorgii. Range, mean and standard deviation (SD) include values
of lectotype. Negative values under Pectoral to dorsal fin refer to dorsal-fin origin in front of pectoral-fin end.
lectotype range mean SD n
Standard length [mm] 169.0 147-210 178.0 22.1 7
Total length [mm] 205.0 178-246 211.3 21.3 7
In percent of standard length
Body depth 11.6 10.8-12.3 11.6 0.5 7
Body width 12.9 11.2-12.9 12.4 0.6 7
Head length 23.3 22.6-24.9 23.5 0.8 7
Pre-dorsal length 30.1 30.1-33.5 31.9 1.2 7
Dorsal-fin length 59.9 52.0-59.9 56.1 2.6 7
Pectoral to dorsal fin -6.0 -6.3--3.1 -4.5 1.2 7
Pre-pectoral length 22.9 22.9-25.0 23.8 0.7 7
Pectoral-fin extension 35.7 35.4-38.2 37.0 1.1 7
Inter-pectoral width 8.6 7.7-9.6 8.9 0.6 7
Pectoral-fin base width 3.3 3.3-3.6 3.5 0.1 7
Pre-pelvic length 70.2 69.8-72.1 71.1 0.9 7
Pre-anal length 85.9 84.1-86.0 85.3 0.8 7
Anal-fin ray length 14.2 12.3-14.6 13.5 0.8 7
Anal-fin base length 5.0 4.5-5.6 5.2 0.4 7
Caudal-peduncle length 3.6 2.8-3.7 3.3 0.3 7
In percent of head length
Head width 56.0 49.8-62.3 55.2 4.9 7
Interorbital width 17.3 15.7-18.5 17.4 0.9 7
Nostril distance 12.2 10.6-12.2 11.7 0.6 7
Snout length 19.5 17.2-21.2 19.5 1.6 7
Eye diameter 12.6 11.1-13.5 12.3 0.9 7
Postorbital length 67.5 65.1-68.5 66.4 1.2 7
Gular length 70.3 53.9-70.3 59.5 5.2 7
Length of first spine 19.7 19.4-22.5 21.0 1.1 7
Length of second spine 23.8 23.1-25.5 23.9 0.8 7
Width : length of first spine 23.1 16.7-23.1 19.7 2.2 7
Length of second : first spine 120.7 103.3-122.8 114.3 7.2 7
Table 4. Meristic characters of Polypterus ansorgii. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of lectotype.
lectotype all material
Dorsal-fin spines 15 13 (5), 14 (1), 15 (1)
Dorsal- and caudal-fin rays 17 17 (2), 19 (5)
Pectoral-fin rays 39 35 (1), 37 (1), 39 (2), 40 (1), 42 (2)
Pelvic-fin rays 10 10 (2), 11 (3), 12 (2)
Anal-fin rays 11 11 (2), 12 (2), 13 (2), 14 (1)
Scales in longitudinal series 56 54 (1), 55 (4), 56 (2)
Scales around body 44 43 (1), 44 (4), 45 (2)
Pre-dorsal scales 11 11 (1), 12 (2), 13 (4)
Pre-pelvic scales 40 40 (1), 41 (5), 42 (1)
Total number of vertebrae 57 57 (1)
Moritz & Britz: Revision of Polypterus
17
Coloration. In life (based on photographs of two
imported specimens of ca. 300 mm SL): dorsal
half of body grey or brown with dark grey to dark
brown dorsal saddle blotches; along flanks dark,
more or less regular, rectangular lateral blotches in
longitudinal series; separated from dorsal bars by
narrow light stripe (Fig. 12). Ventral half of body
whitish to cream with ventrolateral markings
expressed as interrupted dark line below lateral
blotches. Head dark grey dorsally, becoming
lighter laterally and whitish ventrally; irregular
dark spots all over head. Iris light grey with dark
grey irregular pattern. Coloration of upper lips
not differing from dorsal head coloration; lower
lips light beige with large dark spots. Pectoral,
pelvic and confluent dorsal-caudal fin yellowish;
small number of dark spots in pelvic and anal
fins, spots more numerous in soft part of dorsal
fin and in caudal fin; spots arranged in regular
rows on pectoral fin. Dorsal-fin spines dark grey
on anterior face, more yellowish on lateral and
posterior faces; finlet membrane transparent to
slightly greyish.
In preservative: all studied preserved speci-
mens considerably depigmented: no blotches or
spots on lower jaw, head or lower half of flanks.
Dorsal saddle blotches and rectangular blotches
along flank, however, appearing more prominent
than in life.
Distribution. Known only from three river ba-
sins in Guinea-Bissau and the Senegal: Corobal
River, Géba River and Casamance River (Fig. 13).
Records from eastern Niger basin (MRAC 92-014-
P-0001-0002) are based on confusion with P. bichir
(see below). Dagets (1954: 57) record of P. ansorgii
from the Upper Niger basin was subsequently
regarded by Daget & Iltis (1965) as based on an
atypical P. bichir lapradii.
Biology. At least in captivity not different in
behaviour from other species of P. bichir-complex:
voracious predator, predominantly active during
night.
Etymology. Named after Dr. W. J. Ansorge who
collected type specimens.
Remarks. Three specimens were mentioned in
the original description. They are syntypes and are
currently housed in the collections of BMNH and
NMW. We hereby designate BMNH 1910.9.13.4
(Fig. 11) as the lectotype of P. ansorgii.
Fig. 11. Polypterus ansorgii, BMNH 1910.9.13.4, 169 mm SL, juvenile or female, lectotype (present designation);
Guinea Bissau: Corobal River at Tchitoli; W. J. Ansorge; Jun 1909.
Ichthyol. Explor. Freshwaters, IEF-1094
18
Previous confusion of P. ansorgii with other
Polypterus species was probably caused by the sig-
nificant variation in coloration in P. bichir across
its range, and by changes in head shape during
growth in large Polypterus species. The jaws are
of equal length in juveniles in those species that
have a projecting lower jaw in adults, a condition,
which may have led to confusion with P. ansorgii.
Several keys list the equal length of the jaws as
a distinguishing character of P. ansorgii from the
other large species (e. g. Daget & Iltis, 1965). Ad-
ditional records of P. ansorgii from Lake Kainji,
Ogun River (AMNH 215310, USNM 339728)
and Ouémé River (Lalèyè et al., 2004) represent
misidentifications.
a
b
c
a
b
c
Fig. 12. Polypterus ansorgii, aquarium import from Guinea Bissau. a, 313 mm SL, male with light coloration; b, same
specimen with darker coloration; c, detail of head.
Moritz & Britz: Revision of Polypterus
19
Polypterus bichir (Geoffroy, 1802)
(Figs. 14-20, Tables 5-6)
Polyptere bichir Geoffroy [Saint-Hilaire], 1802a: 98.
Polypterus bichir katangae Poll, 1941a: 145.
Polypterus bichir lapradei Steindachner, 1869: 103.
Polypterus niloticus Shaw, 1804: 122.
Material examined. 148 specimens. MNHN 0000-5761,
lectotype (Fig. 14), 585 mm SL; Egypt: Nile River, E.
Geoffroy Saint-Hilaire, 1799. – MNHN 0000-5806, 1,
paralectotype, 549 mm SL; dry mounted specimen,
Egypt: Nile River, E. Geoffroy Saint-Hilaire, 1799. sen-
egal: NMW 63244, 1, 464 mm SL; syntype of P. lapradei,
Senegal: Taoué close to Richar-Toll [= Touey-Marigot],
16°31' N 15°41' W; F. Steindachner, Nov 1868 [but jar
labelled with March 1869, likely referring to date of
registration] (Fig. 15). – NMW 63242, 1, 182 mm SL;
syntype of P. lapradei, Senegal: Podor, ca. 16°39' N
14°57' W; F. Steindachner, Nov 1868 [but jar labelled
with March 1869, likely referring to date of registration].
– NMW 63243:1-2, 2, 166-190 mm SL; syntypes of
P. lapradei, Senegal: Dagana, ca. 16°31' N 14°57' W; F.
Steindachner, Nov 1868 [but jar labelled with March
1869, likely referring to date of registration]. – BMNH
1900.6.28.1, 1, 536 mm SL; Senegal: St. Louis, M. Delhez.
– BMNH 1900.6.28.2-3, 2, 625-675 mm SL; Senegal:
Kaedi, M. Delhez. – BMNH 1900.6.28.4-7, 4, 284-364 mm
SL; Senegal: Kaedi, M. Delhez. – BMNH 1900.6.28.9, 1,
334 mm SL; Senegal: Kaedi, M. Delhez. – MRAC 140104,
1, 308 mm SL; Senegal: Kaedi, M. Delhez, around 1900.
– MRAC 73-005-P-0024, 1, 226 mm SL; Senegal: Djeuss
[St. Louis] River at Bouben, D. F. E. Thys van den
Audenaerde, 8 Mar 1966. – MRAC 73-005-P-0027-0028,
2, 377-399 mm SL; Senegal: Lampsar River [in Senegal
River Delta] at Ross Bethio, D. F. E. Thys van den
Audenaerde, 10 Mar 1966. – MRAC 73-005-P-0029, 1,
494 mm SL; Senegal: Casamance River at Kolda, D. F.
E. Thys van den Audenaerde, 3 Apr 1966. – NMW 63053,
1, 750 mm SL; Senegal, Steindachner. – MNHN 1907-
0250, 1, 338 mm SL; Senegal, Gruvel. gamBia river
Basin: BMNH 1901.7.17.1, 1, 387 mm SL; Gambia, J. S.
Budgett. – BMNH 1905.11.13.10, 1, 326 mm SL; Gambia,
Capt. Vipan. – MNHN 1980-1603, 1, 293 mm SL; Sen-
egal: Gambia River at Wassadou, D. Paugy. geBa
river: BMNH 1912.4.1.2-3, 2, 525-630 mm SL; Guinea-
Bissau: Geba River at Bafata, W. Ansorge. vOlta river
Basin: MRAC 140927-140929, 2, 198-213 mm SL; Bur-
kina Faso: Kou River [tributary of Mouhoun], 4 Jun
1964, B. Roman. – MRAC 143517-143518, 2, 194-220 mm
SL; Burkina Faso: north of Bobo-Dioulasso, B. Roman.
niger river anD triButaries: BMNH 1884.6.9.9-10, 2,
510-517 mm SL; Nigeria: Niger River, Esq. W. Forbes.
– BMNH 1884.6.9.14-18, 5, 257-330 mm SL; Nigeria:
Niger River, Esq. W. Forbes. – BMNH 1907.5.3.1, 259 mm
SL; Nigeria: Benue River north of Ibi, Capt. Vipan. –
BMNH 1928.7.3.1, 1, 255 mm SL; Nigeria: Kiyawa
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 13. Records of Polypterus ansorgii. , type locality; , verified records.
Ichthyol. Explor. Freshwaters, IEF-1094
20
River at Katagum, Esp. L. Lloyd. – BMNH 1953.4.28.126,
1, 503 mm SL; Nigeria: Hadeija-River, E. Trewavas. –
BMNH 1969.3.25.10, 1, 315 mm SL; Nigeria: Sokoto
River, M. H. Holden, Jun 1957. – BMNH 1902.11.10.28-
30, 2, 80-87 mm SL; Nigeria: Assay, W. Ansorge. –
BMNH 1902.11.10.27, 1, 161 mm SL; Nigeria: Abo, W.
Ansorge. – BMNH 1913.12.5.1, 1, 89 mm SL; Nigeria:
Anambra River, W. Crocombe. – MNHN 1925-0172, 1,
242 mm SL; Guinea: Kouroussa, Thomas. – MNHN
1961-0012, 4, 205-295 mm SL; Mali: Diafarabé, J. Daget,
22 May 1951. – MNHN 1961-0013, 2, 160-175 mm SL;
Mali: Diafarabé, J. Daget, 8 Apr 1950. – MNHN 1961-
0014, 1, 116.6 mm SL; Mali: Diafarabé, J. Daget, 27 Oct
1949. – MNHN 1961-1130, 1, 224 mm SL; Guinea: Niger
River at Faranah, Raimbault, 28 Aug 1958. – MNHN
1992-0769, 4, 210-233 mm SL; Guinea: Banie [= Niger]
River at Sanassiya, D. Paugy and R. Bigorne, 2 Mar
1989. – MRAC 72790-72793, 4, 179-253 mm SL; Mali:
Diafarabé, J. Daget, Nov 1946. – MRAC 73-012-P-0007-
0008, 2, 344-448 mm SL; Mali, Niger River at Mopti, H.
Matthes, 17 Sep 1969. – MRAC 92-014-P-0001, 1, 253 mm
SL; Nigeria: Orashi River at Odieke, C. B. Powell, 15
Nov 1991. – MRAC 92-014-P-0002, 1, 264 mm SL; Nige-
ria: Isemu Lake, C. B. Powell, 1-9 Nov 1991. lake ChaD
anD triButaries: BMNH 1928.7.4.1, 1, 361 mm SL; Chad:
Chari River below junction of Lodgone River, C.
Markham. – BMNH 1930.8.7.1-2, 2, 517-554 mm SL;
Chad: Chari River at Logone, J. Glover. – MNHN 1904-
0066, 1, 470 mm SL; Chad: Chari River, Chevalier and
Decorse. – MNHN 1928-0069, 1, 351 mm SL; Cameroon:
El Beid River at Afade, J. Monod. – MNHN 1959-0390,
1, 187 mm SL; Cameroon: El Beid; J. Blache and A.
Stauch, 5 Oct 1954. – MRAC 66466, 1, 405 mm SL;
Cameroon: Benoue River at Lere, J. Monod. – MRAC
153839-153840, 2, 416-486 mm SL; Chad: Mandoul
River at Bahr-Sara, A. Sutter, Mar 1965. – MRAC 73-
015-P-0034, 1, 336 mm SL; Cameroon: Logone River at
Guémé; W. Verheyen et al., 26 Nov 1970. – MRAC 73-
015-P-0035, 1, 608 mm SL; Cameroon: Logone River at
Guémé; W. Verheyen et al., 26 Nov 1970. – MRAC 73-
015-P-0036-0038, 2, 448-503 mm SL; Cameroon: Logone
River at Guémé, W. Verheyen et al., 26 Nov 1970. –
MRAC 73-015-P-0044-0046, 3, 211-377 mm SL; Came-
roon: Benoué River at Garoua, W. Verheyen et al., 7 Dec
1970. – MRAC 73-015-P-0047, 1, 269 mm SL; Cameroon:
Benoué River at Garoua, W. Verheyen et al., 8 Dec 1970.
nile Basin: BMNH 1850.7.29.1, 1, 517 mm SL; Egypt:
Nile, E. Rüppell. – BMNH 1899.4.28.1, 1, 550 mm SL;
Egypt: lower Nile River, Harrington and Harrington.
– BMNH 1899.4.28.2, 1, 466 mm SL; Egypt: lower Nile
River, Harrington and Harrington. – BMNH 1900.9.22.37,
1, 512 mm SL; Sudan: Bahr-el-Jebel, Capt. S. Fowler. –
BMNH 1906.3.3.1-3, 2, 261-299 mm SL; Egypt: west
Damietta, J. Mitchell. – BMNH 1907.12.2.117, 1, 261 mm
SL; Egypt: Nile River near Cairo; W. L. S. Loat, 15 Mar
1899. – BMNH 1907.12.2.118, 1, 667 mm SL; Sudan:
White Nile at Kawa, W. L. S. Loat. – BMNH 1907.12.2.119-
121, 3, 255-273 mm SL; Sudan: White Nile at Goz Abou
Gumah, W. L. S. Loat, 1 May 1901. – BMNH 1907.12.2.123-
124, 2, 203-214 mm SL; Sudan: White Nile at Ghab-el-
Aish, W. L. S. Loat. – BMNH 1907.12.2.125-134, 11,
252-554 mm SL; Sudan: White Nile at Fashoda, W. L.
S. Loat. – BMNH 1907.12.2.135, 1, 655 mm SL; Sudan:
White Nile near Kerro; W. L. S. Loat. – BMNH
1907.12.2.136, 1, 615 mm SL; Sudan: Lako No; W. L. S.
Loat. – MNHN 0000-2609, 2, 482-520 mm SL; Nile; Hit-
torf and Ehrenberg; 1844. – MNHN 0000-5762, 1, 583 mm
SL; Egypte: Nile; Clotbey; 1856. – MNHN 0000-5763, 1,
451 mm SL; Nile; Lenormand. – MRAC 140103, 1,
244 mm SL; Sudan: White Nile at Goz Abou Gumah,
W. L. S. Loat; 1907. – NMW 30297, 2, 426-429 mm SL;
Sudan: Bahr-el-Gazal; Marno, 1882. – NMW 30299, 1,
386 mm SL; Nile, Steindachner. – NMW 3302, 1, 443 mm
SL; Nile, Kotschy, 1842. – NMW 63209, 3, 245-267 mm
SL; Sudan: Khartoum, Drasche, 1876. – NMW 63210, 1,
465 mm SL; Nile, Steindachner, Nov 1910. – NMW 63211,
2, 321-357 mm SL; Nile; Steindachner, Oct 1910. – NMW
63212, 1, 364 mm SL; Sudan: Bahr-el-Seraf, Marno, 1881.
– NMW 63213, 1, 518 mm SL; Sudan: Bahr-el-Ghazal,
Steindachner, 1882. – NMW 63214, 1, 495 mm SL; Sudan:
Zuflüsse des oberen Nil [affluents of the Upper Nile],
Marno, 1881. – NMW 63215, 1, 497 mm SL; Sudan: White
Nile at Khor Attac, Werner, 1907. – NMW 63216, 1,
457 mm SL; Nile, Steindachner, Oct 1910. – NMW 63217,
1, 212 mm SL; Nile, Steindachner, 2 Nov 1910. – NMW
63219, 1, 434 mm SL; Sudan, Steindachner, Aug 1910.
– NMW 77728, 1, 528 mm SL; Sudan: Melut, 10°27' N
32°13' E; W. Sixl, 1981. – NMW 79061, 2, 485-572 mm
SL; Sudan: Melut, 10°27' N 32°13' E; W. Sixl, 1983. – NMW
9584, 1, 340 mm SL; side branch of Nile, Steindachner
and Marno, 1981. – ZSM 34546, 1, 330 mm SL; Sudan:
Omdourman [fishmarket], N. Pöllath, 14 Feb 2006.
katanga regiOn: MRAC 20671, 1, 389 mm SL; holotype
of P. bichir katangae, Democratic Republic of Congo:
Nyonga: Lake Upemba, G.-F. de Witte, 8-12 May 1925.
– MRAC 49319, 1, 109.5 mm SL; paratype of P. bichir
katangae, Democratic Republic of Congo: Lualaba Rivert
at Bukuma, P. Brien, Jun 1937. – MRAC 66561, 1,
134.5 mm SL; paratype of P. bichir katangae, Demo-
cratic Republic of Congo: Lualaba Rivert at Bukuma, P.
Brien, 23-26 Jun 1937. – MRAC 66562, 1, 149.2 mm SL;
paratype of P. bichir katangae, Democratic Republic of
Congo: Lualaba Rivert at Bukuma, P. Brien, 23-26 Jun
1937. – MRAC 66563, 1, 123.6 mm SL; paratype of
P. bichir katangae, Democratic Republic of Congo: Lu-
alaba Rivert at Bukuma, P. Brien, 23-26 Jun 1937
(Fig. 16). – IRSNB 68, 1, 415 mm SL; paratype of P. bichir
katangae, Democratic Republic of Congo: Lualaba River
at Maka, H. J. Bredo, 1 Feb 1939. – MRAC 178671, 1,
52.7 mm HL [only head preserved], paratype of P. bichir
katangae, Democratic Republic of Congo: Lake Kazi-
baziba, W. de Smet, 17 Nov 1956. – MRAC 178672, 1,
45.6 mm HL [only head preserved], paratype of P. bichir
katangae, Democratic Republic of Congo: Make, 50 km
from Bukuma, W. de Smet, 27 Dec 1955. – MRAC 178673,
1, 51.4 mm HL [only head preserved], paratype of
P. bichir katangae, Democratic Republic of Congo: Make,
50 km from Bukuma, W. de Smet, 27 Dec 1955. – MRAC
Moritz & Britz: Revision of Polypterus
21
69702, 1, 445 mm SL; Democratic Republic of Congo:
Lualaba River at Maka, M. Poll, 25 Jun 1947. – MRAC
79-01-P-0003, 1, 396 mm SL; Democratic Republic of
Congo: Lake Upemba: Mabwe, F. de Witte, 7-9 Sep
1947. WithOut lOCality Data: NMW 50276, 1, 585 mm
SL.
Additional material. nile Basin: DMM IE/10412, 2,
Sudan: Khartoum [central fish market]; T. Moritz et
al., Apr 2008.
Diagnosis. Polypterus bichir is distinguished from
all other members of the genus except P. ansorgii,
P. endlicherii and P. congicus, by 13 or more dorsal-
fin spines (vs. 12 or fewer) and the third dorsal
scale row being the first to reach ventral midline
(vs. second dorsal scale row reaching ventral
midline). Polypterus bichir is distinguished from
P. endlicherii and P. congicus by dorsal saddle-like
blotches, when present not confluent with, but
separate from lateral blotches (vs. dorsal and
lateral blotches confluent and always present).
When present, dorsal saddle blotches are more
numerous in P. bichir, with 9 or more such blotches
(vs. 6-8 in P. congicus and 4-6 in P. endlicherii), and
wide, i. e. at least 3 scale rows wide in P. bichir
(vs. less than 3 scale rows wide in P. congicus).
Polypterus bichir is distinguished from P. ansorgii
by more scales around the body (43-54 vs. 43-45)
and a larger maximum adult size (750 mm SL vs.
313). Most specimens can be distinguished from
P. ansorgii, P. congicus and P. endlicherii by 61 or
more scales in lateral series (55-70 [lower quartile
61] vs. 54-60 [upper quartile 60]).
Descriptive synopsis. Meristic and morphomet-
ric data in Tables 5-6. Body elongated with 55-70
scales and 57-67 vertebrae. Jaws terminal only in
juveniles; in large specimens lower jaw project-
ing beyond upper jaw. In large specimens head
and anterior body region often slightly dorso-
ventrally flattened (but never to the degree as in
large P. endlicherii and P. congicus). Size up to at
least 750 mm SL (820 mm TL).
Coloration. In life: coloration highly variable
(Fig. 17). Dorsal half of body brownish, light
brown, light grey or yellowish with dark brown
or grey markings varying from very faint to
conspicuous and pronounced: dorsal and lateral
blotches expressed only from about midlength,
more anteriorly replaced by two dark brown
longitudinal bands separated by lighter narrow
longitudinal stripe. Ventral half of body uniform
beige with no dark ventrolateral markings or with
few to many markings formed by aggregated dark
scales in anterior two-thirds of flanks, forming
irregular to regular lines; on posterior part of
ventrolateral body sometimes with several ventro-
lateral blotches in confluence with lateral blotches
forming irregular network of blotch lines. Dorsal
half of head light brown, grey or yellowish with
dark marbling of varying density. Ventral half of
head and cheek lighter with numerous dark spots
of variable sizes; often spotting extending down
onto gular plates. Lip markings formed by small
blotches. Iris yellow or orange with irregular dark
spots. Nasal tentacles showing same uniform or
marbled coloration as dorsal half of head. Also
markings on head very variable ranging from
virtually absent to much pronounced.
Pectoral fin yellow with dark brown elongate
spot-like markings arranged in several transversal
more or less regular bands; pectoral-fin base with
numerous large dark spots arranged to form a
blotch varying in expression from almost absent to
pronounced. Pelvic fin yellowish transparent with
or without dark spots arranged in more or less
regular bands. Soft part of dorsal fin and conflu-
ent caudal fin brown, yellowish or grey with dark
markings frequently expressed as densely set dark
spots loosely arranged in stripes. Dorsal-fin spines
grey, dark grey or brown with light or yellowish
spots arranged at approximately the same level as
in succeeding spines; similar pattern continuing
onto first and second order secondary fin rays
originating from spines; finlet-membrane grey-
transparent, usually without pattern, sometimes
with some dark marbling.
Different intensity of coloration not restricted
to any geographic areas, except the intensely
colored specimens from Koliba River. Plain
colored and more intensely pigmented specimens
present in same river basins at same time.
In preservative: all areas which are light
in life become pale in preservative; dark color
markings less pronounced; fine marbling on head
sometimes absent; dark spots on lower half of
head and body less intense. Dry paralectotype
(MNHN 0000-5806) without any color pattern;
ethanol preserved lectotype (MNHN 0000-5761)
only with faint color markings.
Distribution. Widely distributed in the major
river basins of the Nilo-Sahelo-Sudan region: Nile,
Chad, Niger, Volta and Senegal (Fig. 18). Addi-
tionally in few West African coastal basins: Gam-
Ichthyol. Explor. Freshwaters, IEF-1094
22
Fig. 14. Polypterus bichir, MNHN 0000-5761, 585 mm SL, male, lectotype; Egypt: Nile; E. Geoffroy Saint-Hilaire;
1799.
Fig. 15. Polypterus bichir, NMW 63244, 464 mm SL, male, syntype of P. lapradei; Taoué close to Richar-Toll [= Touey-
Marigot]; F. Steindachner; Nov 1868.
Moritz & Britz: Revision of Polypterus
23
bia, Casamanche and Geba River. Also distributed
in the Katanga area of the Congo basin, mainly
the Lualaba River. Occurrences of P. bichir in Lake
Turkana (e. g. Pellegrin, 1914; Holly, 1933; Poll,
1941a; Reitzer et al., 1972; Gosse, 1990; Seegers et
al., 2003) and Ouémé River basin (Gosse, 1990) are
questionable, as they are not supported either by
museum specimens or photographs.
Biology. Apparently most active at night, active
swimmer and not essentially a bottom-liver or a
mudfish (Harrington, 1899), living on muddy
substrates (Chad: Pellegrin, 1914). Polypterus bichir
prefers deeper depressions of muddy river beds
(Nile: Harrington, 1899) and feeds on small fish
of all types and on crustaceans (Chad: Pellegrin,
1914). Reproduction takes place during flood
season (Senegal: Reitzer et al., 1972) and mature
fishes enter flood plains for spawning (Gambia:
Budgett, 1900; Niger: Daget, 1954).
Etymology. The name refers to the local Egyptian
name, bichir, for the species (Geoffroy, 1802a-b).
Remarks. Some confusion has arisen about
which of three publications represents the origi-
nal description of this species (Geoffroy 1802a-b;
Lacépède, 1803). Geoffroys publication in the
Bulletin des Sciences (1802a) was published
between 21st of March and 20th of April in 1802.
This predates his publication in the Annales du
Muséum dHistoire naturelle (Geoffroy, 1802b),
which was not published before 22nd of September
of the same year. Thus Geoffroy (1802a) made the
species name available, with the original spelling
as Polyptère bichir (which needs to be amended
to Polyptere bichir in accordance with Art. 33.3 of
the Code). One year later, Lacépède (1803: 340)
used the Latinised version, Polypterus bichir, in his
publication and this work that has been cited by
some authors, though incorrectly, as the descrip-
tion of the species (e. g. Holly, 1933; Schäfer, 2004;
Claeson & Hagadorn, 2008; Grandstaff et al., 2012;
Fricke et al., 2018). See also above under remarks
for the genus Polypterus.
Geoffroy Saint-Hilaire (1802a) did not specify
any type material and did not mention on how
Fig. 16. Polypterus bichir, MRAC 66563, 124 mm SL, juvenile, paratype of P. bichir katangae; Democratic Republic
of Congo: Lualaba Rivert at Bukuma; P. Brien; 23-26 Jun 1937.
Ichthyol. Explor. Freshwaters, IEF-1094
24
Table 5. Morphometric characters of Polypterus bichir. Range, mean and standard deviation (SD) include values
of lectotype. Negative values under Pectoral to dorsal fin refer to dorsal-fin origin in front of pectoral-fin end.
lectotype range mean SD n
Standard length [mm] 585.0 109.5-750 364.0 140.9 139
Total length [mm] 660.0 131.9-820 407.7 155.5 131
In percent of standard length
Body depth 10.1 7.1-14.5 11.0 1.3 137
Body width 11.6 8.2-14.1 11.9 1.0 137
Head length 19.2 17.8-26.9 21.4 1.9 139
Pre-dorsal length 28.0 25.3-37.7 30.2 2.0 138
Dorsal-fin length 59.8 46.8-62.5 56.5 3.0 138
Pectoral to dorsal fin 4.0 -8.4-4.00 -3.6 1.7 135
Pre-pectoral length 19.6 16.7-26.6 21.6 1.8 137
Pectoral-fin extension 31.6 27.5-41.6 34.0 2.7 135
Inter-pectoral width 7.0 5.9-10.4 7.7 0.9 136
Pectoral-fin base width 3.1 1.8-3.9 2.9 0.4 137
Pre-pelvic length 70.3 64.8-73.6 70.0 1.5 138
Pre-anal length 85.1 80.6-88.3 85.0 1.4 138
Anal-fin ray length 11.4 7.3-16.3 12.3 1.6 136
Anal-fin base length 6.3 3.2-7.6 5.3 0.7 136
Caudal-peduncle length 4.1 2.2-4.8 3.4 0.5 136
In percent of head length
Head width 51.6 38.9-63.6 50.9 3.8 142
Interorbital width 17.8 15.6-24.8 19.0 1.7 141
Nostril distance 12.2 8.0-15.4 12.2 1.2 141
Snout length 19.0 14.6-22.3 19.5 1.4 141
Eye diameter 8.7 7.4-15.2 10.1 1.6 142
Postorbital length 72.5 62.3-74.2 69.6 2.1 142
Gular length 50.4 48.6-75.0 61.4 6.5 142
Length of first spine 22.7 11.0-28.5 22.4 3.0 133
Length of second spine 18.3 13.4-30.4 24.3 2.6 134
Width : length of first spine 15.2 12.7-36.7 17.8 3.8 133
Length of second : first spine 80.6 63.8-215.5 110.5 17.5 132
Table 6. Meristic characters of Polypterus bichir. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of lectotype.
lectotype all material
Dorsal-fin spines 16 13 (17), 14 (36), 15 (30), 16 (37), 17 (16), 18 (2)
Dorsal- and caudal-fin rays 19 18 (1), 19 (9), 20 (39), 21 (65), 22 (22), 23 (2), 24 (1)
Pectoral-fin rays 41 34 (1), 35 (4), 36 (1), 37 (13), 38 (15), 39 (21), 40 (25), 41 (25), 42 (18), 43 (8),
44 (3), 45 (1), 46 (1), 48 (1)
Pelvic-fin rays 13 10 (8), 11 (23), 12 (42), 13 (45), 14 (18), 15 (3)
Anal-fin rays 16 9 (1), 10 (3), 11 (11), 12 (28), 13 (44), 14 (33), 15 (16), 16 (1)
Scales in longitudinal series 67 55 (1), 56 (1), 58 (3), 59 (11), 60 (11), 61 (11), 62 (17), 63 (14), 64 (16), 65 (21),
66 (23), 67 (6), 68 (2), 70 (1)
Scales around body 44 43 (6), 44 (7), 45 (15), 46 (22), 47 (26), 48 (18), 49 (11), 50 (16), 51 (8), 52 (5),
53 (2), 54 (2)
Pre-dorsal scales 13 10 (2), 11 (6), 12 (23), 13 (39), 14 (35), 15 (25), 16 (3), 17 (4)
Pre-pelvic scales 48 38 (1), 39 (1), 40 (5), 41 (2), 42 (5), 43 (10), 44 (15), 45 (20), 46 (23), 47 (19),
48 (16), 49 (15), 50 (5), 51 (1)
Total number of vertebrae 57 (4), 58 (2), 59 (1), 61 (3), 62 (3), 63 (3), 64 (4), 65 (2), 66 (6), 67 (2)
Moritz & Britz: Revision of Polypterus
25
a
b
c
d
Fig. 17. Polypterus bichir. a-b, 351 mm SL, aquarium import, Koliba-coloration, Guinea; c, Senegal: Nikologo-
Koba National Park (photo by M. Reichard); d, 340 mm SL; Sudan: Khartoum [central fish market] (photo by N.
Pöllath).
many specimens he based his description. His
remark that P. bichir has 16, 17 or 18 finlets indi-
cates that he had at least three specimens. In the
MNHN there are only two specimens preserved
from Geoffroys collection dated to 1799: MNHN
0000-5761 (Fig. 14) and MNHN 0000-5806. Both
were originally stored in ethanol, but the latter
specimen was made into a dry specimen in 1810,
according to Bertin (1940) who listed these two
specimens as holotype and paratype, respectively.
This view was followed by Gosse (1984). Subse-
quently, Eschmeyer (1998) designated MNHN
Ichthyol. Explor. Freshwaters, IEF-1094
26
0000-5761 as lectotype thus rendering MNHN
0000-5806 a paralectotype.
In 1869, Steindachner published the descrip-
tion of a new bichir from the Senegal, Polypterus
lapradei. He noted that the essential difference
between P. lapradei and P. bichir is the shape of
the head, whereas counts of dorsal-fin spines
and scales show an overlapping gradient: 14-15
vs. 15-17 dorsal-fin spines, 60-62 vs. 64 scales
in lateral row, 13-14 vs. 12-13 pre-dorsal scales
(values given in Steindachner, 1869).
Steindachner (1870) mentioned four speci-
mens, which he used for his description. All
four specimens are deposited at the Naturhis-
torisches Museum in Vienna and catalogued as
three objects with numbers NMW 63242, NMW
3243 (2 specimens) and NMW 63244 (Fig. 15).
Regarding P. lapradei, Eschmeyer (1998: 870) noted
Syntypes: (3) NMW 63053 (1)., thus indicating
the presence of four syntypes, although he only
provided the catalog number for one of them.
This specimen, NMW 63053, however, was not
mentioned in the original description, as it ex-
ceeds the maximal length given by Steindachner
(1870) and therefore cannot be regarded as part
of the type series. One syntype (NMW 63244) was
photographed by F. Schäfer (2004) and referred to
as holotype with the catalogue number NMW
32244 in the text, a lapsus for NMW 63244, as
evidenced by the photographed label.
Polypterus lapradei had been regarded as a
valid species until Poll (1941a-b, 1942) referred to
it as a subspecies of P. bichir. Since the beginning
of the 1970s this view has become widely ac-
cepted (Gosse, 1984). In his keys for West African
polypterid species, Gosse (1990, 2003) employed
the following criteria to distinguish both subspe-
cies: number of scales in a longitudinal series and
number of dorsal-fin spines, but noted that both
meristic characters show overlap. He also stated
and illustrated disjunct distribution areas for the
two subspecies: the Benoué River and river basins
west of it for P. bichir lapradei, and the Chad basin
and river basins east of it for P. bichir bichir (Gosse,
1990, 2003). Schäfer (2004) in his popular book on
bichirs treated P. lapradei again as a valid species,
and distinguished it from P. bichir by details of
the color pattern: P. lapradei was considered to
possess usually clearly marked longitudinal
stripes in the anterior region of body (vs. absence
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 18. Records of Polypterus bichir. , type locality; , verified records; , other records; , questionable
records.
Moritz & Britz: Revision of Polypterus
27
-3.0 -2.5 -2.0 -1.5 -1.0 -0.5 0.5 1.0 1.5
PC1
PC2
-2.4
-1.6
-0.8
0.8
1.6
2.4
3.2
4.0
-3.0 -2.5 -2.0 -1.5 -1.0 -0.5 0.5 1.0 1.5
-2.4
-1.6
-0.8
0.8
1.6
2.4
3.2
4.0
PC1
PC2
a
b
Fig. 19. Scatterplots of first and second components (PC 1 and PC 2) of Polypterus bichir specimens throughout its
distributional range. a, meristic characters: scales in longitudinal series, pre-dorsal scales, pre-pelvic scales,
dorsal-fin spines, pectoral-fin rays, pelvic-fin rays, anal-fin rays, soft rays in confluent dorsal and caudal fin;
b, meristic characters as in Figure 19a, plus morphometric characters: body depth, body width, head length, head
width, pre-dorsal length, length of spinous part of dorsal fin, distance from distal tip of adducted pectoral fin to
first finlet, pre-pectoral length, extension of pectoral fin, inter-pectoral width, width of pectoral-fin base, pre-
pelvic length, pre-anal length, length of longest ray in anal fin, length of anal-fin base, caudal-peduncle length,
inter-orbital width, nostril distance, eye diameter, post-orbital length. All data are normalized. , , West Africa
[, Western coastal basins; , Senegal basin; , Volta basin; , Niger basin, without Benoue]; , , Chad and Benoué
[, Benoué basin; , Chad basin]; , , Nile basin; , , Katanga region.
of such stripes) and a uniformly plain colored
area above the anal fin (vs. a pattern consisting
of light blotches on a darker background). In our
view the color pattern of P. bichir is more variable
throughout the distribution area (see above).
Furthermore, the aquarium-specimens regarded
by Schäfer (2004) as P. lapradei originated from
Nigeria, and the one regarded as P. bichir came
Ichthyol. Explor. Freshwaters, IEF-1094
28
from the upper Niger in Guinea. These putative
origins of his P. bichir and P. lapradei individuals
do not fit the distribution patterns published by
Gosse (1990, 2003).
A second subspecies, Polypterus bichir katangae
(Fig. 16), was described by Poll (1941a) from the
Katanga region in the Congo basin based on a
holotype and five paratypes. He distinguished
this sub-species from P. bichir bichir by fewer
dorsal-fin spines (12-14 vs. 14-18), fewer scales in
a lateral series (58-60 vs. 63-70) and fewer scales
around the body (44-46 vs. 46-54; values given
by Poll, 1941a). For his comparison Poll (1941a)
used only the values provided by Boulenger (1907)
for P. bichir and studied just a single additional
non-Katanga specimen of P. bichir, an individual
from Cameroon.
Our analysis of the meristic and morphometric
data collected for this study revealed no clear
distinction between the supposed subspecies of
P. bichir (Fig. 19) and show great overlap between
the West African specimens and those with a Nile,
Chad or Katanga origin. Exclusion of all West
African specimens from the analysis results in a
distinction between Nile plus Chad specimens
on the one hand and the Katanga specimens on
the other hand (Fig. 19a). The PCA analysis dem-
onstrates that distant populations within a large
distribution area can become distinguishable from
each other. But, if sampling covers the whole area
the values merge and form a continuous range.
Plotting several meristic data grouped according
to their geographical origin on box-and-whisker-
plots (Fig. 20) shows some trends for P. bichir: the
Katanga
W.Africa
Chad
Nile
12 13 14 15 16 17 18 19
Katanga
W.Africa
Chad
Nile
54 56 58 60 62 6 4 66 68 70 72
ScL
DS
Katanga
W.Africa
Chad
Nile
9 10 11 12 13 14 15 16 17 18
ScpD
Katanga
W.Africa
Chad
Nile
36 38 40 42 44 46 48 50 52
ScpP
Fig. 20. Box-and-whisker plots of selected meristic data of P. bichir from throughout its distributional range,
grouped into geographical regions. Nile (n = 54); Lake Chad, including Chari and Benoué (n = 17); West Africa,
from Niger on to the West (n = 58); Katanga region (n = 8). DS, dorsal-fin spines; ScL, scales in longitudinal series;
ScpD, pre-dorsal scales; ScpP, pre-pelvic scales.
Moritz & Britz: Revision of Polypterus
29
more west they occur individuals from the Nile
across the Chad basin to West Africa tend to have
fewer scales in the lateral line, fewer dorsal-fin
spines and fewer pre-pelvic scales; specimens
from the Katanga region also show low values
in these parameters similar to the West African
individuals (Fig. 20). Despite the geographical
trend in the number of dorsal-fin spines there is
no such trend in the number of pre-dorsal scales
(Fig. 20).
At first glance the color pattern appears to fol-
low a similar geographical trend: specimens are
more pigmented and contrasted the more west
they were collected. This trend is culminating in
the coloration of the population from the Corubal
River [= Koliba River] along the border of Guinea-
Bissau and Guinea from which conspicuously pig-
mented specimens (Fig. 17 a,b) enter the aquarium
trade under the trade name P. lapradei Koliba
or P. sp. Koliba. Specimens from Katanga also
show a more pronounced, contrasted coloration
(Fig. 16). Specimens collected in the Lower Niger,
Chad area and Nile basin (Fig. 17d) on the other
hand are often much more faintly pigmented, and
faint specimens are also common in rivers of West
Africa. Yet, conspicuously colored specimens
are not just restricted to the Corubal River and
Katanga region, but have also been collected in
the Niger River close to its delta (e. g. MRAC 92-
014-P-0001, MRAC 92-014-P-0002) and from the
Nile north of Cairo (e. g. BMNH 1907.12.2.117).
Based on analysis of our data we synonymise
P. lapradei and P. bichir katangae with P. bichir and
only recognize a single species, however, with
geographical variation in meristic counts. The
cause of the sometimes striking differences in
coloration remains unclear.
Recently, Otero et al. (2006) described a well-
preserved almost complete fossil of Polypterus as a
new species, P. faraou. Meristic and morphometric
characters of the fossil P. faraou (values taken
from Otero et al., 2006), however, fall completely
within the ranges we report here for P. bichir.
Thus, there is little reason to consider the fossil
P. faraou distinct from P. bichir. Polypterus faraou
may potentially be conspecific with P. bichir, a
conclusion that needs to be further tested.
Polypterus congicus Boulenger, 1898
(Figs. 21-23, Tables 7-8)
Polypterus congicus Boulenger, 1898: 418.
Material examined. 66 specimens. type speCimens:
BMNH 1897.9.30.28, lectotype (Fig. 21, present des-
ignation), male, 656 mm SL; Democratic Republic of
Congo: Stainley Falls. – BMNH 1899.2.20.33, 1, para-
lectotype, 167 mm SL; juvenile; Democratic Republic
of Congo: New Antwerp [Makanza]. – MRAC 79, 1,
paralectotype, male, 580 mm SL; Democratic Republic
of Congo: Mayanga, Wilventh, 1896. – MRAC 119, 1,
paralectotype, 205 mm SL; juvenile, Democratic Re-
public of Congo: Leopoldville [= Kinshasa], Wilventh,
1896. COngO river: BMNH 1899.6.27.1, 1, 579 mm SL;
Democratic Republic of Congo: Upper Congo. – BMNH
1899.6.28.1, 1, 202 mm SL; Democratic Republic of
Congo: Brazaville, M. Meuse. – BMNH 1900.6.23.14,
192 mm SL; Democratic Republic of Congo: Bolobo, G.
Grenfell. – BMNH 1919.9.10.1, 1, 270 mm SL; Democratic
Republic of Congo: Stanley Falls. – BMNH 1919.9.10.1,
1, 265 mm SL; Democratic Republic of Congo: Luvua
River at Ankoro, K. Banister, 1974/1975. – IRSNB 9960,
1, 315 mm SL; Democratic Republic of Congo: Congo
River at Yangambi, A. Hulot, Dec 1949. – IRSNB 100015,
1, 471 mm SL; Democratic Republic of Congo: Congo
River at Yangambi, A. Hulot, 1948. – MNHN 1886-0292,
1, 485 mm SL; Democratic Republic of Congo: Congo
River at Nangtchou; S. de Brazza. – MRAC 2243, 1,
271 mm SL; Democratic Republic of Congo: Stanleyville,
C. Christy, before 1913. – MRAC 19804, 1, 145.9 mm SL;
Democratic Republic of Congo: Stanleyville, Richard,
1929. – MRAC 43966, 1, 225 mm SL; Democratic Republic
of Congo: Kinshasa, A. Tinant, 1933. – MRAC 103366,
1, 260 mm SL; Democratic Republic of Congo: Stanley
Pool, A. Werner, Jun 1955. – MRAC 103993-103996, 4,
239-274 mm SL; Democratic Republic of Congo: Stanley
Pool, A. Werner, Jun 1955. – MRAC 115689, 1, 205 mm
SL; Democratic Republic of Congo: Stanley Pool, Brien
and Poll, 6 Apr 1956. – MRAC 115692, 1, 724 mm SL;
Democratic Republic of Congo: Stanley Pool, Brien and
Poll, Sep 1957. – MRAC 73-022-P-0012, 1, 237 mm SL;
Democratic Republic of Congo: Stanley Pool: marai
Limgundu, J. Mandeville, 9 Apr 1958. – MRAC 73-022-
P-0013-0015, 3, 194-245 mm SL; Democratic Republic of
Congo: Stanley Pool, J. Mandeville, 9 Apr 1958. – MRAC
A7-033-P-0029, 1, 361 mm SL; Democratic Republic
of Congo: Malebo Pool: Mipongo island, 4°16'19" S
15°30'41" E; Hanssens et al., 11 Sep 2007. – MRAC
A7-033-P-0030, 1, 257 mm SL; Democratic Republic
of Congo: Malebo Pool: Mipongo island, 4°16'19" S
15°30'41" E; Hanssens et al., 11 Sep 2007. – RMNH 20873,
1, 207 mm SL; Democratic Republic of Congo: Stanley
Pool, A. Werner, Jul 1955. – ZSM 38691, 1, 505 mm SL:
Democratic Republic of Congo: Malbeo Pool: Kenkole
[fishmarket], U. Schliewen et al., 29 Jul 2008. kasai
river anD triButaries: MRAC 66950, 1, 374 mm SL;
Democratic Republic of Congo: Kasai River: region de
Ichthyol. Explor. Freshwaters, IEF-1094
30
Mushie, Vleeschouwers, Oct 1945. – MRAC 67073, 2,
453-503 mm SL; Democratic Republic of Congo: Kasai
River: region de Mushie, Vleeschouwers, Oct 1945. Ou-
Bangui river: MRAC 58071, 1, 216 mm SL; Democratic
Republic of Congo: Oubangui River at Yakoma, Rosy,
1938. – MRAC P-182255, 1, 354 mm SL; Democratic
Republic of Congo: Dungu River at Gangala na Bodio,
C. dElzius, 1954/1955. – MRAC P-179888-179889,
2, 386-412 mm SL; Democratic Republic of Congo:
Dungu River at Gangala na Bodio, M. Poll, Nov 1956.
Table 8. Meristic characters of Polypterus congicus. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of lectotype.
lectotype all material
Dorsal-fin spines 14 11 (1), 12 (2), 13 (13), 14 (39), 15 (11), 16 (1)
Dorsal- and caudal-fin rays 19 17 (1), 18 (4), 19 (37), 20 (21), 21 (3)
Pectoral-fin rays 43 37 (2), 38 (2), 39 (6), 40 (6), 41 (10), 42 (12), 43 (14), 44 (8), 45 (7), 46 (1)
Pelvic-fin rays 13 10 (1), 11 (3), 12 (15), 13 (23), 14 (21), 15 (3), 16 (1)
Anal-fin rays 16 10 (2), 11 (2), 12 (13), 13 (17), 14 (16), 15 (12), 16 (5)
Scales in longitudinal series 56 54 (2), 55 (10), 56 (24), 57 (17), 58 (9), 59 (4), 60 (1)
Scales around body 49 41 (1), 42 (1), 44 (4), 45 (3), 46 (14), 47 (14), 48 (11), 49 (8), 50 (10), 51 (1)
Pre-dorsal scales 14 11 (5), 12 (22), 13 (28), 14 (10), 15 (2)
Pre-pelvic scales 42 39 (1), 40 (3), 41 (15), 42 (18), 43 (18), 44 (8), 45 (4)
Total number of vertebrae 55 52 (1), 53 (1), 55 (3), 57 (1)
Table 7. Morphometric characters of Polypterus congicus. Range, mean and standard deviation (SD) include values
of lectotype. Negative values under Pectoral to dorsal fin refer to dorsal-fin origin in front of pectoral-fin end.
lectotype range mean SD n
Standard length [mm] 656.0 145.9-788 405.5 168.6 66
Total length [mm] 740.0 174.8-876 478.0 189.6 58
In percent of standard length
Body depth 12.3 9.2-16.6 11.8 1.6 66
Body width 14.9 10.8-17.1 14.1 1.3 66
Head length 20.9 20.0-26.1 22.6 1.5 66
Pre-dorsal length 32.3 27.2-36.3 31.5 1.8 66
Dorsal-fin length 56.9 49.3-64.7 55.9 3.0 66
Pectoral to dorsal fin -3.2 -8.4--1.0 -4.6 1.7 66
Pre-pectoral length 20.6 19.5-26.8 23.2 1.6 66
Pectoral-fin extension 34.8 31.6-43.2 36.6 2.8 66
Inter-pectoral width 9.0 6.8-12.1 9.5 0.9 65
Pectoral-fin base width 3.7 2.8-4.3 3.5 0.3 66
Pre-pelvic length 70.7 64.5-73.1 70.2 1.4 66
Pre-anal length 85.8 82.6-90.6 85.4 1.2 66
Anal-fin ray length 11.3 10.3-17.4 13.4 1.5 66
Anal-fin base length 6.1 3.1-7.4 5.7 0.8 66
Caudal-peduncle length 3.7 2.3-4.9 3.5 0.5 66
In percent of head length
Head width 62.4 50.5-67.2 57.7 3.3 67
Interorbital width 24.9 18.3-27.4 21.5 1.7 67
Nostril distance 15.9 10.0-16.6 13.8 1.4 67
Snout length 22.0 19.4-24.5 21.9 1.0 67
Eye diameter 7.9 6.7-13.1 9.3 1.3 67
Postorbital length 70.5 63.3-72.0 68.6 1.9 67
Gular length 72.7 52.3-72.3 61.6 6.0 67
Length of first spine 17.5 17.1-27.2 22.0 2.3 65
Length of second spine 18.6 19.7-29.1 23.6 2.2 65
Width : length of first spine 26.0 13.1-27.3 18.2 3.4 65
Length of second : first spine 106.7 91.5-128.5 108.0 7.6 65
Moritz & Britz: Revision of Polypterus
31
lake tanganyika anD triButaries: BMNH 1906.9.6.1-
4, 4, 465-577 mm SL; Democratic Republic of Congo:
Lake Tanganyika at Kalambo, J. E. S. Moore. – BMNH
1906.9.8.1, 1, 540 mm SL; Zambia: Lake Tanganyika
at Kituta, W. Cunnington. – BMNH 1936.6.15.18, 1,
601 mm SL; Lake Tanganyika, C. Christy. – BMNH
1936.6.15.19-21, 3, 381-603 mm SL; Lake Tanganyika,
C. Christy. – BMNH 1936.6.15.23-26, 6, 579-693 mm SL;
Lake Tanganyika, C. Christy. – BMNH 2005.4.22.1-2,
2, 576 mm SL [of one specimen only head preserved],
Zambia: Lake Tanganyika: mouth of Lufubu River,
8°33'40" S 30°43'24" E; H. Buescher, 10 Nov 2004. – IRSNB
9443, 1, 600 mm SL; Tanzania: delta of Malagarasi River,
20 May 1947. – MRAC 88875, 1, 788 mm SL; Democratic
Republic of Congo: Albertville [= Kalemie] [fishmarket],
M. Poll, 1 Feb 1944. – MRAC 190227, 1, 375 mm SL;
Zambia: Lake Tanganyika: Chinyika bay: Kasaba, H.
Matthes, 20-23 Apr 1967. – MRAC 93-152-P-0323, 1,
251 mm SL; Tanzania: delta of Malagarasi River, L.
de Vos, 21 Aug 1993. katanga: BMNH 1975.6.20.1,
1, 265 mm SL; Democratic Republic of Congo: Luvua
River at Akoro, K. Banister, 1974/1975. – MRAC 3205,
1, 523 mm SL; Democratic Republic of Congo: Sankuru
River at Kondue, Luja, before 1913. – MRAC 3206, 1,
470 mm SL; Democratic Republic of Congo: Sankuru
River at Kondue, Luja, before 1913. – MRAC 3207, 1,
329 mm SL; Democratic Republic of Congo: Sankuru
River at Kondue, Luja, before 1913. – MRAC 3210, 1,
584 mm SL; Democratic Republic of Congo: Sankuru
River at Kondue, Luja, before 1913. – MRAC 34775,
1, 360 mm SL; Democratic Republic of Congo: Luvua
River at Kiambi, G. F. de Witte, 18 Apr-3 May 1931.
– MRAC 35030, 1, 351 mm SL; Democratic Republic of
Congo: Luvua River at Kiambi, G. F. de Witte, 4-20
May 1931. – MRAC 49320, 1, 625 mm SL; Democratic
Republic of Congo: Lualaba River at Maka, P. Brien, Jul
1937. – MRAC 66560, 1, Democratic Republic of Congo:
Lualaba River at Maka, P. Brien, Jul 1934. – MRAC
69701, 1, 688 mm SL; Democratic Republic of Congo:
Lualaba River at Maka, M. Toll, 9 Apr 1958. – MRAC
90-047-P-0088, 1, 274 mm SL; Democratic Republic of
Congo: Lualaba River at km 47 on road Kisangani-Wanie
Rukula, L. de Vos, 28 May 1990.
Fig. 21. Polypterus congicus, BMNH 1897.9.30.28, 656 mm SL, male, lectotype (present designation); Democratic
Republic of Congo: Stainley Falls.
Ichthyol. Explor. Freshwaters, IEF-1094
32
Additional material. COngO river: MRAC 72-022-P-
0010, 1, Democratic Republic of Congo: Stanley Pool, J.
Mandeville, 10 Jan 1955. OuBangui river: MRAC 1306,
1, juvenile, Democratic Republic of Congo: Oubangui
River at Banzyville [= Mobayi-Mbongo], Royanne, 1901.
– MRAC 58057-58070, 14, Democratic Republic of Con-
go: Oubangui River at Yakoma, Rosy, 1939. katanga:
IRSNB 4870, 1, Democratic Republic of Congo: Lualaba
River at Maka, J. H. Bredo, 1 Feb 1939. – MRAC 30547,
[only remnants], Democratic Republic of Congo: Luvua
River at Kiambi, 1930. – MRAC 178712, 1, Democratic
Republic of Congo: Kikondja, W. de Smet, 14 Apr 1956.
Diagnosis. Polypterus congicus is distinguished
from all other members of the genus except P. an-
sorgii, P. bichir and P. endlicherii, by third dorsal
scale row being the first to reach ventral midline
(vs. second). Polypterus congicus is distinguished
from P. ansorgii, and P. bichir by 6-8 (rarely 9)
dorsolateral bars formed by the confluence of
dorsal and lateral blotches (vs. dorsal and lateral
blotches separated, not forming continuous bars)
and from P. endlicherii by more numerous (6-8,
rarely 9 vs. 4-6) and narrower bars (less than 3
scale rows wide vs. wider than 3 scale rows).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 7-8. A large species of
the P. bichir-group reaching at least 788 mm SL
(876 mm TL). Body moderately elongated. Jaws
terminal in juveniles only; in large specimens
lower jaw projecting beyond upper jaw. Anterior
body region and head becoming dorso-ventrally
flattened with increasing size.
Coloration. In life: dorsal half of body grey to
brown with slender dark grey to black dorsolateral
bars; on anterior body bars directed obliquely
posteriorly, on posterior body obliquely anteriorly
(Fig. 22). Ventral half of body light, white or yel-
lowish, sometimes with few irregular dark grey
spots arranged approximately in a horizontal line
as ventrolateral markings (much less conspicuous
than in P. endlicherii). Dorsal half of head including
upper jaw brown or yellowish-brown with dense
irregular dark brown to black spots and marbling;
cheeks lighter. Ventral half of head with fewer but
larger dark spots; lower jaw light with dark spots
usually forming several vertical lines. Iris yellow
to orange with dense brown marbling. Pectoral,
a
b
a
b
Fig 22. Polypterus congicus. a-b, aquarium import (photos: Schuzo Nakano/Aqualog).
Moritz & Britz: Revision of Polypterus
33
pelvic and anal fins yellowish with fine, densely
set, dark brown spots and marbling (usually not
expressed as lines as in P. endlicherii). Base of pec-
toral fin with dark spots and blotches concentrated
in distal part. Soft dorsal fin and confluent caudal
fin hyaline to grey with light yellowish spots, rays
darker. Dorsal-fin spines grey or brownish with
dark brown irregular pattern; finlet membrane
translucent with color pattern similar to that of
soft dorsal fin.
A conspicuously colored specimen from Lake
Tanganyika was depicted by Schäfer (2004: 61-63).
The specimen had strongly contrasted color mark-
ings on body and regularly, radially arranged
dark markings on roof of head and opercular
region. Schäfer (2004) suggested that this might
represent a new species, by labelling it Polypterus
sp. aff. congicus. We see, however, no reason to
regard it as anything other than a strongly and
particularly colored P. congicus.
In preservative: color pattern similar to life
coloration, but yellow and yellowish markings in
life now white, and black markings in life dark
grey (Fig. 21).
Distribution. Polypterus congicus is widely dis-
tributed in the Congo River basin, and tributaries
and southern part of Lake Tanganyika (Fig. 23).
Biology. Within the Congo basin the species
seems to be restricted to the main river and
neighbouring water bodies and it has been re-
ported to enter the flooded forest during high
water levels (Gosse, 1965). In the southern part
of Lake Tanganyika the species inhabits areas
with clear water and sandy substrate with large
boulders (Büscher, 2004). Interestingly, so far only
large specimens have been recorded from Lake
Tanganyika; it remains unclear in which areas
breeding and juvenile growth take place (Büscher,
2004). Polypterus congicus feeds on bottom dwell-
ing organisms (Gosse, 1965).
Etymology. Not explicitly stated, but apparently
an allusion to the type locality (Congo).
Remarks. Boulengers (1898) original description
was based on four syntypes. One specimen which
can be clearly assigned to the original type series
(based on collection date, collection place, number
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 23. Records of Polypterus congicus. , type locality; , verified records.
Ichthyol. Explor. Freshwaters, IEF-1094
34
of dorsal-fin spines and number of lateral line
scales) is: BMNH 1897.9.30.28 (Fig. 21). Fricke et
al. (2018) incorrectly listed this specimen as holo-
type. We designate herein BMNH 1897.9.30.28
as lectotype of P. congicus. Three additional
specimens from New Antwerp, Leopoldville and
Manayanga were mentioned by Boulenger (1898)
and these paralectotypes are most likely: BMNH
1899.2.20.33, MRAC 79 and MRAC 119.
Polypterus delhezi Boulenger, 1899
(Figs. 24-26, Tables 9-10)
Polypterus delhezi Boulenger, 1899: 61, Pl. XXX,
Fig. 2.
Material examined. 44 specimens. type speCimens:
MRAC 422, lectotype (Fig. 24, present designation),
male, 314 mm SL; Democratic Republic of Congo: Lake
Leopold at Kutu, P. Delhez. – BMNH 1899.2.20.17, 1,
paralectotype, female, 270 mm SL; Democratic Republic
of Congo: New Antwerp [= Makanza], Wilvert and Wa-
genaar. COngO river: BMNH 1975.8.15.67, 1, 241 mm SL;
Democratic Republic of Congo: Congo River, A. Fraser-
Brunner. – MNHN 1886-0294, 1, 311 mm SL; Democratic
Republic of Congo: Nangtchou, S. de Brazza. – MRAC
103987, 1, 260 mm SL; Democratic Republic of Congo:
Stanley Pool, A. Werner, Jun 1955. – MRAC 154587, 1,
165.4 mm SL; Democratic Republic of Congo: Stanley
Pool, P. Brichard, 1966. – MRAC 177711, 1, 277 mm
SL; Democratic Republic of Congo: Stanley Pool, J. van
Orshoven, 5 May 1964. – MRAC 177717, 1, 251 mm SL;
Democratic Republic of Congo: Bolobo; NKele, 1958.
– MRAC 73-022-P-0008-0009, 2, 250-285 mm SL; Demo-
cratic Republic of Congo: Stanley Pool, J. Mandeville,
5 Sep 1958. – MRAC 88-025-P-0642, 1, 135.0 mm SL;
Democratic Republic of Congo: Congo River at Bumba,
L. de Vos, 30 Jan 1988. – MRAC 91-013-P-0640-0642, 3,
214-295 mm SL; Democratic Republic of Congo: N' Sele
River [affluent of Congo River] close to Kinshasa, 4°08' S
16°40' E; M. Tshibwabwa Sinaseli, 1985/1987. – MRAC
A7-0033-P-0031-0033, 3, 226-234 mm SL; Democratic
Republic of Congo: Canal Kisanga at Inga, 5°27'55" S
13°34'58" E; Hanssens et al., 29 Aug 2007. – RMNH 20874,
1, 190 mm SL; Democratic Republic of Congo: Stanley
Pool, A. Werner, Jul 1955. kasai river: MRAC 67060,
1, 280 mm SL; Democratic Republic of Congo: Fimi
River at Bungi, Vleeschouwers, 22 Oct 1945. – MRAC
68750-68754, 4, 173-250 mm SL; Democratic Republic
of Congo: Fimi River at Mushie, Vleeschouwers, 21 Sep
1946. – MRAC 138889, 1, 278 mm; Democratic Republic
of Congo: Mushie, M. Creeter, 1957. – MRAC 66832, 1,
258 mm SL; Democratic Republic of Congo: Kasai River
at Fimi, Vleeschouwers, Sep 1941. – MRAC 66852, 1,
260 mm SL; Democratic Republic of Congo: Kasai River
at Fimi, Vleeschouwers, Sep 1941. lake tumBa: IRSNB
4871, 1, 350 mm SL; Democratic Republic of Congo: Lake
Tumba at NKosso Norma, G. de Loneux, 31 Dez 1938.
– MRAC 14688, 1, 102.4 mm SL; Democratic Republic
of Congo: Lake Tumba at Bikoro, D. Schouteden, 1921.
– MRAC 14689, 1, 99.5 mm SL; Democratic Republic
of Congo: Lake Tumba at Bikoro, D. Schouteden, 1921.
– MRAC 57751-57752, 2, 340-348 mm SL; Democratic
Republic of Congo: Lake Tumba, G. de Loneux, 1928.
– MRAC 14690; Democratic Republic of Congo: Lake
Tumba, G. de Loneux, 1928. likOuala river: MNHN
1962-0349, 9, 60.8-111.6 mm SL; Democratic Republic
Fig. 24. Polypterus delhezi, MRAC 422, 314 mm SL, male, lectotype (present designation); Democratic Republic of
Congo: Lake Leopold at Kutu; P. Delhez.
Moritz & Britz: Revision of Polypterus
35
of Congo: Likouala River at Mossaka, Stauch, 24 Feb
1961. OuBangui river: MRAC 179773, 1, 247 mm SL;
Democratic Republic of Congo: Ngiri River [affluent
of Oubangui River], P. van Leynseele, 1970. ruki river
anD triButaries: MRAC 80642, 1, 329 mm SL; Demo-
cratic Republic of Congo: Tshuapa River at Bokuma, P.
Lootens, 20 Sep 1952. – MRAC 73-022-P-003, 1, 158.4 mm
SL; Democratic Republic of Congo: Tshuapa River at
Boende, R. Philippe, Jun 1956. aquarium speCimen:
NMW 90940, 1, 217 mm SL; Tiergarten Schönbrunn.
Additional material. COngO river: MRAC 39459-
39461, 3, Democratic Republic of Congo: Leopoldville
[= Kinshasa], A. Tinant, 1933. – MRAC 43978, 1, Demo-
cratic Republic of Congo: Leopoldville [= Kinshasa], A.
Tinant, 1935. – MRAC 47995, 1, Democratic Republic
of Congo: Leopoldville [= Kinshasa], A. Tinant, 1937.
– MRAC 67462, 1, Democratic Republic of Congo:
Leopoldville [= Kinshasa], Bureau. – MRAC 68034, 1,
Democratic Republic of Congo: Leopoldville [= Kin-
shasa], De Vier, Aug 1945. – MRAC 102572-102576,
5, Democratic Republic of Congo: Kingabwa, J. Man-
deville, Aug-Sep 1955. – MRAC 115693, 1, Democratic
Republic of Congo: Stanley Pool, Brien and Poll, 4 May
1956. – MRAC 115694, 1, Democratic Republic of Congo:
Stanley Pool, Brien and Poll, 23 Jul 1956. – MRAC 115695,
1, Democratic Republic of Congo: Stanley Pool, Brien
and Poll, 26 Sep 1957. – MRAC 115696, 1, Democratic
Republic of Congo: Stanley Pool, Brien and Poll, 4 Oct
1957. – MRAC 115697-115701, 5, Democratic Republic of
Congo: Stanley Pool, Brien and Poll, Sep 1957. – MRAC
118720, 1, Democratic Republic of Congo: Stanley Pool,
P. Brichard, 1956. – MRAC 140106-140108, 3, Democratic
Republic of Congo: environs of Leopoldville [= Kinsha-
sa], van de Weyer, 1960. – MRAC 177721, 1, Democratic
Republic of Congo: Bolobo, NKele, 1956. – MRAC 73-
a
b
c
a
b
c
Fig. 25. Polypterus delhezi, aquarium specimens. a, 240 mm SL, adult; b, 72 mm SL, juvenile; c, early juvenile
(photo c: Schuzo Nakano/Aqualog).
Ichthyol. Explor. Freshwaters, IEF-1094
36
33-P-1, 1, Democratic Republic of Congo: environs of
Leopoldville [= Kinshasa], R. Henrion, 1971. – MRAC
73-022-P-0001-0002, 2, Democratic Republic of Congo:
Stanley Pool: Mikunga passage, J. Mandeville, 29 Oct
1957. – MRAC 73-022-P-0003, 1, Democratic Republic of
Congo: Stanley Pool: Nsele River, J. Mandeville, 29 Oct
1957. – MRAC 73-022-P-0004, 1, Democratic Republic of
Congo: Stanley Pool: Ndjiili River, J. Mandeville, 30 Oct
1957. – MRAC 73-022-P-0006, 1, Democratic Republic
of Congo: Stanley Pool, J. Mandeville, 8 Nov 1957.
Table 10. Meristic characters of Polypterus delhezi. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of lectotype.
lectotype all material
Dorsal-fin spines 10 10 (3), 11 (33), 12 (8)
Dorsal- and caudal-fin rays 16 15 (7), 16 (34), 17 (3)
Pectoral-fin rays 39 31 (1), 32 (1), 33 (3), 34 (11), 35 (9), 36 (3), 37 (6), 38 (7), 39 (3)
Pelvic-fin rays 10 9 (9), 10 (24), 11 (10), 12 (1)
Anal-fin rays 12 9 (4), 10 (8), 11 (9), 12 (11), 13 (9), 14 (2), 15 (1)
Scales in longitudinal series 55 54 (4), 55 (19), 56 (17), 57 (4)
Scales around body 37 35 (1), 36 (6), 37 (9), 38 (9), 39 (11), 40 (6), 41 (2)
Pre-dorsal scales 17 13 (7), 14 (24), 15 (8), 16 (3), 17 (2)
Pre-pelvic scales 37 35 (1), 36 (8), 37 (16), 38 (15), 39 (3)
Total number of vertebrae 54 (1), 55 (2)
Table 9. Morphometric characters of Polypterus delhezi. Range, mean and standard deviation (SD) include values
of lectotype. Negative values under Pectoral to dorsal fin refer to dorsal-fin origin in front of pectoral-fin end.
lectotype range mean SD n
Standard length [mm] 314.0 60.8-350 212.9 87.2 44
Total length [mm] 339.0 77.9-382 235.6 89.8 40
In percent of standard length
Body depth 10.4 8.5-14.2 11.6 1.1 44
Body width 10.8 9.9-13.2 11.7 0.8 44
Head length 17.0 16.2-23.4 19.0 2.0 44
Pre-dorsal length 34.0 28.0-38.2 32.5 2.1 44
Dorsal-fin length 56.0 47.1-61.7 56.0 3.1 44
Pectoral to dorsal fin 2.4 -3.8-3.3 -0.3 1.5 44
Pre-pectoral length 17.2 15.4-23.2 19.4 1.8 44
Pectoral-fin extension 30.3 28.4-38.0 32.9 2.7 44
Inter-pectoral width 5.8 5.8-9.7 7.6 0.9 44
Pectoral-fin base width 2.9 2.6-3.7 3.1 0.3 44
Pre-pelvic length 65.6 62.5-67.6 65.0 1.1 44
Pre-anal length 84.7 81.8-87.8 84.4 1.2 44
Anal-fin ray length 9.0 7.9-15.2 11.9 1.8 44
Anal-fin base length 6.2 3.9-7.0 5.2 0.7 44
Caudal-peduncle length 3.6 2.2-5.0 3.5 0.6 44
In percent of head length
Head width 53.2 45.3-63.7 56.7 3.6 44
Interorbital width 21.8 18.1-24.5 21.5 1.3 44
Nostril distance 13.5 9.6-17.1 13.2 1.5 44
Snout length 19.2 18.0-24.3 21.4 1.4 44
Eye diameter 12.3 10.8-18.0 13.3 1.8 44
Postorbital length 66.9 57.5-68.9 64.8 2.6 44
Gular length 58.0 56.0-69.5 60.1 2.7 44
Length of first spine 34.2 17.9-35.9 26.0 3.8 44
Length of second spine 35.2 21.0-40.4 29.0 3.4 43
Width : length of first spine 16.8 15.4-35.1 22.4 3.9 43
Length of second : first spine 103.0 89.2-142.1 112.8 11.8 42
Moritz & Britz: Revision of Polypterus
37
kasai river: MRAC 2804, 1, Democratic Republic of
Congo: Bokala, D. J. Maes. – MRAC 2805, 1, Democratic
Republic of Congo: Bokala, D. J. Maes. lake tumBa:
MRAC 14686, 1, Democratic Republic of Congo: Lake
Tumba at Tondu, D. Schouteden, 1921. – MRAC 14687,
1, Democratic Republic of Congo: Lake Tumba at Tondu,
D. Schouteden, 1921. – MRAC 14690, 1, Democratic Re-
public of Congo: Lake Tumba at Bikoro, D. Schouteden,
1921. – MRAC 14691, 1, Democratic Republic of Congo:
Lake Tumba at Bikoro, D. Schouteden, 1921. ruki river
anD triButaries: IRSNB 21048, 1, Democratic Republic
of Congo: Ruki River, Kiss, Feb 1980. – MRAC 74871,
1, Democratic Republic of Congo: Bokungu, L. Dupuis,
1950. – MRAC 177776, 1, Democratic Republic of Congo:
Boende, P. Brichard, 1969.
Diagnosis. Polypterus delhezi is distinguished
from species of the P. bichir group by its second
dorsal scale row reaching ventral midline (vs.
third scale row) and lateral line scales with cen-
tral pores and entire posterior margin (vs. with
incision at posterior margin). Polypterus delhezi
is distinguished from all remaining Polypterus
species except P. senegalus by low number of pre-
dorsal scales (13-17 vs. 20-37). It is distinguished
from P. senegalus by the presence of a series of
vertical dark brown to black dorsolateral bars
(vs. absence) and by more dorsally located eyes
(lower margin of orbit visible in dorsal view vs.
not visible). Furthermore, Polypterus delhezi differs
from P. senegalus in usually having more dorsal-
fin spines (10-12, median 11 vs. 8-11, median 9)
and smaller interorbital width (18.1-24.5 % HL
vs. 22.4-30.7).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 9-10. Body moderately
elongated. Mouth terminal, upper jaw usually not
projecting beyond lower jaw; in rare cases upper
jaw very slightly projecting. Eyes in dorso-lateral
position. First dorsal-fin spine at about vertical
through posterior margin of pectoral fin. Size up
to at least 350 mm SL (382 mm TL).
Coloration. In life: dorsal half of body light
grey with 5-9 black, vertical or oblique dorsolat-
eral bars of approximately two scale rows width
formed by the confluence of dorsal with preceding
(rarely succeeding) lateral blotches; bars some-
times with light or yellowish margin; confluence
frequently incomplete resulting in separate dorsal
and lateral blotches (Fig. 25). Ventral half of body
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 26. Records of Polypterus delhezi. , type locality; , verified records.
Ichthyol. Explor. Freshwaters, IEF-1094
38
whitish or cream, sometimes with ventrolateral
markings expressed as longitudinal stripe and/or
blotches surrounding lighter yellowish blotches,
especially in juveniles (Fig. 25c). Dorsal half of
head, iris and upper lip grey with irregular dark
grey to black marbling; ventral half of head and
cheeks white, with suborbital stripe expressed as
line of dark spots and extending posteriorly to
dark spots and blotches on lower half of opercular
region. Frequently with dark line formed by series
of spots ventral and parallel to suborbital stripe;
lower lip and jaw whitish with distinct black spots.
Pectoral fin with dark spots arranged in two or
three narrow rows; distal part of pectoral-fin base
with dark markings, not forming a distinct blotch.
Pelvic fin with dark spots sometimes arranged
in thin rows. Otherwise pectoral and pelvic fins
whitish, yellowish or translucent. Dorsal, caudal
and anal fins dark grey with light, yellowish,
irregular pattern; distal parts of fins sometimes
darker. Dorsal-fin spines grey with dark grey to
black markings at mid-level and in distal parts;
finlet-membrane transparent with dark markings,
often a dark irregular area at mid-level.
In preservative: dorsal half of body light
brown to beige; bars conspicuous, black or dark
brown; color pattern on head, including subor-
bital stripe less distinct than in life; ventrolateral
markings less distinct (Fig. 24).
Distribution. Restricted to the major rivers in the
central and lower Congo basin (Fig. 26).
Biology. Life history largely unknown.
Etymology. Named after Paul Delhez who col-
lected the lectotype.
Remarks. Boulenger (1899) mentioned two speci-
mens in the original description of Polypterus
delhezi, without designating a holotype. Fricke et
al. (2018) stated that Boulenger (1909) provided a
lectotype designation in his catalogue of African
fishes. There, however, Boulenger (1909: 14) only
mentioned one type deposited in MRAC and
another type being part of the BMNH collection,
clearly regarding both specimens as syntypes.
We designate herein MRAC 422, the specimen
illustrated by Boulenger (1899, pl. XXX, fig. 2;
1909, fig. 9), as the lectotype of P. delhezi (Fig. 24).
Polypterus endlicherii Heckel, 1848
(Figs. 27-31, Tables 11-12)
Polypterus endlicherii Heckel, 1848: 310, Pl. 22,
fig. 1.
Material examined. 64 specimens. type spe Cimen :
NMW 91244, holotype (Fig. 27), 684 mm SL; dry speci-
men; Sudan: Nile River at Khartoum, [likely] Kotschy.
COastal river Basins Of Côte DivOire anD ghana:
BMNH 1923.3.2.2-3, 2, 136-211 mm SL; Côte dIvoire:
Bandama River, W. Lowe. – BMNH 1934.8.31.1, 1,
167 mm SL; Ghana: Ashanti Forest: Ejura, W. Lowe.
– BMNH 1969.4.28.10, 1, 296 mm SL; Ghana: Afram
River at Aframoso, W. R. Smith, 5 Sep 1952. – MNHN
1894-0392, 1, 206 mm SL; Côte dIvoire: Bandama
River at Tiassale, Pobeuin. – MNHN 1894-0393, 1,
192 mm SL; Côte dIvoire: Bandama River at Tiassale,
Pobeuin. – MNHN 1960-0177, 193 mm SL; Burkina
Faso: Yanon River [tributary of Comoé], Arnoult and
dAubenton. – MNHN 1991-0439, 1, 344 mm SL; Côte
dIvoire: Bandama River at Lamto, Lamotte, Jul 1970.
– MRAC 74-014-P-0003, 1, 449 mm SL; Côte dIvoire:
Lake Kossou, S. Kyeli, 17 Dec 1973. – MRAC 79-036-
P-0001, 1, 261 mm SL; Côte dIvoire: Lagune Ebrié at
Layo, D. F. E. Thys van den Audenaerde, 20 Oct 1979.
– MRAC 80-019-P-0001, 1, 388 mm SL; Côte dIvoire:
Sassandra River at Soubré, G. Teugels and L Risch,
24-25 Apr 1980. – MRAC 85-029-P-0001, 1, 281 mm
SL; Côte dIvoire: Boubo River at Babokon, G. Teugels,
8-9 May 1985. – MRAC 85-029-P-0002, 1, 285 mm SL;
Côte dIvoire: Boubo River at Babokon, G. Teugels,
9-10 May 1985. – MRAC 85-055-P-001, 1, 188 mm SL;
Burkina Faso/Côte dIvoire [border]: Léreba River at
Léreba, G. Teugels, 24 Nov 1985. – MRAC 86 013-P-0001,
201 mm SL; Côte dIvoire: Bandama River at Tiassale,
G. Teugels, 3-4 Apr 1986. – MRAC 87-018-P-0001,
1, 188 mm SL; Burkina Faso/Côte dIvoire [border]:
Léreba River at Léreba, G. Teugels, 29 Jan 1987. lake
ChaD anD triButaries: MNHN 1919-0074, 1, 175 mm
SL; Central African Republic: Gribingui River, Baudon.
– MNHN 1928-0071, 1, 234 mm SL; Chad: Mayo Kebbi:
Lere, Monod. – MNHN 1938-0032, 1, 213 mm SL; Chad:
Logone River at Lai, Thomas. – MRAC 73-026-P-0006, 1,
209 mm SL; Chad: south of Hollom: Tofoul, 11°10'20" N
15°03'10" E; M. Y. Brandily, 24 Aug 1969. Ouémé river
Basin: MRAC 73-001-P-0006, 1, 262 mm SL; Benin:
Porto Novo, P. de Kimpe, 15 Nov 1963. vOlta river
Basin: BMNH 1949.10.20.1, 1, 272 mm SL; Ghana: Volo,
Buxton. – MNHN 2007-0246, 1, 344 mm SL; Togo: Oti
River at Mango, C. Lévêque, 3 Nov 1982. – MRAC
140923-140924, 2, 183.1-206 mm SL; Burkina Faso: Kou
River [tributary of Mouhoun], B. Roman, 4 Jun 1964.
niger river anD triButaries: BMNH 1902.11.10.31, 1,
139.2 mm SL; Nigeria: Abo, S. W. J. Ansorge, 20 Oct 1901.
– BMNH 1913.12.5.2-3, 2, 96.4-129.7 mm SL; Nigeria
Anambra River, W. Crocombe. – BMNH 1936.11.24.1,
1, 241 mm SL; Nigeria: Kaduna River, J. Welman. –
Moritz & Britz: Revision of Polypterus
39
BMNH 1959.8.18.79, 1, 153.1 mm SL; Nigeria: Lokoja,
Maclaren, 1 Nov 1948. – MNHN 1961-0015, 1, 276 mm
SL; Mali: Niager River at Diafarabe, J. Daget, 8 Apr 1950.
– MNHN 1961-0016, 1, 205 mm SL; Mali: Niager River at
Diafarabe, J. Daget, 8 Apr 1950. – MNHN 1962-0444, 1,
197 mm SL; Cameroon: Benoue River at Mayo-Rey, A.
Stauch, May 1960. – MNHN 1986-0051, 3, 226-294 mm
SL; Guinea: Milo River at Boussoule, C. Lévêque, Feb
2006. – MRAC 140111, 1, 219 mm SL; Cameroon: Benoue
River, A. Stauch. nile river Basin: BMNH 1865.11.15.4,
Table 12. Meristic characters of Polypterus endlicherii. Numbers in brackets indicate number of specimens with
the respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 12 11 (5), 12 (34), 13 (22), 14 (3)
Dorsal- and caudal-fin rays (damaged) 18 (12), 19 (33), 20 (15), 21 (1), 22 (1)
Pectoral-fin rays 40 35 (1), 38 (4), 39 (9), 40 (7), 41 (7), 42 (16), 43 (5), 44 (8), 45 (5), 46 (2)
Pelvic-fin rays 14 11 (3), 12 (13), 13 (17), 14 (17), 15 (8), 16 (3), 17 (3)
Anal-fin rays 13 9 (1), 10 (1), 11 (2), 12 (16), 13 (24), 14 (16), 15 (3)
Scales in longitudinal series 55 53 (3), 54 (9), 55 (18), 56 (24), 57 (6), 58 (1), 59 (1), 60 (1)
Scales around body 43 39 (4), 40 (2), 41 (8), 42 (13), 43 (11), 44 (12), 45 (9), 46 (3), 47 (1), 48 (1)
Pre-dorsal scales 13 12 (18), 13 (28), 14 (10), 15 (7), 16 (1)
Pre-pelvic scales 40 39 (1), 40 (11), 41 (16), 42 (13), 43 (14), 44 (6), 45 (3)
Total number of vertebrae 53 (1), 54 (4), 55 (1), 56 (2)
Table 11. Morphometric characters of Polypterus endlicherii. Range, mean and standard deviation (SD) include values
of holotype. Negative values under Pectoral to dorsal fin refer to dorsal-fin origin in front of pectoral-fin end.
holotype range mean SD n
Standard length [mm] 684.0 96.4-780 324.5 169.4 64
Total length [mm] (damaged) 115.9-764 356.1 170.6 59
In percent of standard length
Body depth 10.6 8.6-16.0 11.2 1.6 64
Body width 15.1 11.9-16.2 14.1 1.1 64
Head length 20.8 18.4-26.4 22.4 1.8 64
Pre-dorsal length 30.0 27.4-47.5 32.1 3.0 64
Dorsal-fin length 59.8 49.5-62.8 56.2 3.4 64
Pectoral to dorsal fin -2.0 -7.7-0.0 -3.9 1.8 63
Pre-pectoral length 20.1 19.4-27.4 22.8 1.9 64
Pectoral-fin extension 32.2 31.4-43.2 36.3 2.8 64
Inter-pectoral width 9.1 8.0-12.2 9.6 0.9 64
Pectoral-fin base width 3.6 3.0-4.5 3.8 0.3 64
Pre-pelvic length 67.3 66-73 70.1 1.6 64
Pre-anal length 88.2 82.6-89.9 85.6 1.5 64
Anal-fin ray length 12.0 9.9-14.9 12.4 1.3 63
Anal-fin base length 6.4 3.2-7.3 5.3 0.7 64
Caudal-peduncle length 2.8 1.5-5.4 3.7 0.7 64
In percent of head length
Head width 57.9 51.1-71.7 59.5 4.1 64
Interorbital width 21.7 17.9-26.7 21.8 2.1 64
Nostril distance 11.4 10.9-16.8 13.4 1.2 64
Snout length 15.9 14.9-22.9 19.8 1.6 64
Eye diameter 7.6 7.0-13.2 9.7 1.5 64
Postorbital length 68.7 64.1-74.4 69.1 2.2 64
Gular length 53.0 50.9-73.8 60.3 5.4 64
Length of first spine 22.2 11.0-28.6 22.0 3.5 63
Length of second spine 22.6 18.5-29.7 25.0 2.4 62
Width : length of first spine 21.5 12.8-44.8 21.5 5.6 63
Length of second : first spine 101.6 85.7-269.8 117.0 25.6 61
Ichthyol. Explor. Freshwaters, IEF-1094
40
1, 335 mm SL; Egypt: Nile River below cataracts, Peth-
erick. – BMNH 1907.12.2.137, 1, 230 mm SL; Sudan:
near Goz Shebesha, W. L. S. Loat, 3 Jan 1901. – BMNH
1907.12.2.138, 1, 611 mm SL; Sudan: White Nile near
Kawa, W. L. S. Loat, 5 Jan 1901. – BMNH 1907.12.2.139,
1, 565 mm SL; Sudan: White Nile at Goz Abu Gumah,
W. L. S. Loat. – BMNH 1907.12.2.140, 1, 574 mm SL;
Sudan: White Nile at Fashoda, W. L. S. Loat, 28 Mar
1901. – BMNH 1907.12.2.141, 1, 525 mm SL; Sudan: Lake
No, W. L. S. Loat, 15 Feb 1901. – BMNH 1907.12.2.142, 1,
444 mm SL; Sudan: White Nile at Gondokoro, W. L. S.
Loat, 11 Mar 1902. – BMNH 1907.12.2.143-144, 2, 248-
652 mm SL; Sudan: Omdurman: Sheduah, W. L. S. Loat,
31 Dec 1900. – MNHN 2001-2180, 1, 224 mm SL; Sudan:
aab
Fig. 27. Polypterus endlicherii, NMW 91244, 684 mm SL, female, holotype; Sudan: Nile River at Khartoum; [likely]
Kotschy.
Fig. 28. Polypterus endlicherii, view of predorsal region. a, NMW 91244, holotype; b, illustration of holotype
(Fenzl et al., 1849); arrow points to scale anomaly.
Moritz & Britz: Revision of Polypterus
41
White Nile: Jebel Aulia reservoire at Kowa, T. Roberts.
– NMW 10954, 1, 304 mm SL; Nil, Steindachner, Nov
1910 (Fig. 29). – NMW 50278, 1, 340 mm SL; Sudan: Bahr
el Seraf, Steindachner, 1881. – NMW 63023, 1, 704 mm
SL; Sudan: Bahr el Arab, Marno, 1881. – NMW 63024, 1,
705 mm SL; Sudan: White Nile, Kotschy, 1852. – NMW
63054, 1, 698 mm SL; Sudan: Bahr el Ghazal, [likely]
Marno, 1881. – NMW 63218, 1, 279 mm SL; Sudan:
Khartoum, Drasche and Steindachner [?], 1876. – NMW
63237, 2, 270-343 mm SL; Sudan, Steindachner, 1910. –
NMW 63239, 1, 367 mm SL; Sudan, Steindachner, 1910.
– NMW 63240, 2, 381-388 mm SL; Sudan, Steindachner,
Aug 1910. – NMW 63241, 1, 550 mm SL; Sudan: White
Nile at Ed Dueim, Werner, 1907. – NMW 91233, 1,
780 mm SL; [likely] Sudan: Khartoum, [likely] Reitz,
[likely] 1851. – ZSM 35162, 1, 254 mm SL; Sudan: White
Nile at Kosti, 13°10'17" N 32°40'07" E; D. Neumann, 13
Jan 2007. – ZSM 39631, 1, 558 mm SL; Sudan: Omdour-
man [fishmarket], T. Moritz et al., Apr 2008. aquarium
traDe: ZSM 34041, 1, 216 mm SL; Guinea: [likely] Niger
River at Faranah and Kobikoro, F. Schäfer, 2004.
Additional material. COastal river Basins Of Côte
DivOire anD ghana: MRAC 140109, 1, Côte dIvoire:
Bandama River, Lowe. niger river Basin: MRAC 91-100-
P-0004, 1, Nigeria: Nun River at Kaiama, 5°08' N 6°18' E;
C. B. Powell, 12-13 Sep 1991. vOlta river Basin: DMM
IE/10393, 1, Benin: Pendjari National Park: Pendjari
River at Pont Arly, 11°28'05" N 1°34'01" E; T. Moritz,
25 Nov 2002. – DMM IE/10399, 1, Benin: Pendjari Na-
tional Park: Mare Hippotrague, 11°23'14" N 1°22'15" E;
T. Moritz, 14 Apr 2007. – DMM IE/10400, 3, Benin:
Pendjari National Park: Pendjari River at Hotel Pendjari,
11°24'33" N 1°35'22" E; T. Moritz, Apr 2007 . – MRAC
93-127-P-0001, 1, Ghana: Daka River [tributary of Lake
Volta] at Dindu, 8°10' N 0°20' W; P. C. Goudswaard, 16
Apr 1993. – MRAC 93-127-P-0002, 1, Ghana: White Volta
at Kope, 8°45' N 1°28' W; P. C. Goudswaard, 24 Sep 1992.
Diagnosis. Polypterus endlicherii is distinguished
from all other members of the genus except P. an-
sorgii, P. bichir and P. congicus, by third dorsal
scale row being the first to reach ventral midline
(vs. second). Furthermore, Polypterus endlicherii
is distinguished from all Polypterus species ex-
cept P. bichir and P. congicus by short pre-dorsal
length of only 21.7-27.4 % SL (vs. 27.4-47.5).
Polypterus endlicherii is distinguished from P. an-
sorgii, P. bichir and P. congicus by its unique color
pattern: few (4-5, rarely 6), broad (more than 3
scale rows wide), dark transverse bars extending
from dorsum to middle of flank by confluence of
dorsal and lateral blotches sometimes including
ventrolateral markings (vs. dorsal and lateral
blotches distinct, not confluent and forming bars
in P. bichir and P. ansorgii, or 6-8 narrow bars in
P. congicus).
Descriptive synopsis. Meristic and morphomet-
ric data in Tables 11-12. Large species, up to at
least 780 mm SL. Body moderately elongated.
Jaws terminal in juveniles but in larger specimens
lower jaw projecting beyond upper jaw. Anterior
Fig. 29. Polypterus endlicherii, NMW 10954, 304 mm SL, female; Nile River.
Ichthyol. Explor. Freshwaters, IEF-1094
42
body region and head becoming more dorso-
ventrally flattened with increasing size (Fig. 9).
Coloration. In life: dorsal half of body grey to
brownish-grey with broad dark-grey saddle-
like dorsolateral bars; bars more than three scale
rows wide, frequently also confluent with ven-
trolateral markings; bars less regular in caudal
area (Fig. 30). Ventral half of body light, whitish,
with ventrolateral markings usually expressed as
irregular dark-grey to black blotches sometimes
confluent with wide bars; occasionally with an
additional longitudinal line further ventrally.
Dorsal half of head including upper jaw dark
grey with dark brown to black spots. Lower half
of head including lower jaw light, whitish, with
dark spots on lateral side of head and lower jaw,
but not on gular plate; spots sometimes arranged
in more or less regular lines. Iris and skin around
eye with same coloration as head. Pectoral and
a
b
c
Fig. 30. Polypterus endlicherii. a, large specimen ~400 mm SL; Côte dIvoire: Comoé National Park: Comoé River;
b-c, ~100 mm SL, juvenile; Benin: Pendjari National Park: Pendjari River.
Moritz & Britz: Revision of Polypterus
43
pelvic fins light, whitish, with dark spots and
small blotches; pelvic fin usually with no or faint
darker markings; pattern on pectoral fin arranged
in 3-5 more or less regular lines. Base of pectoral
fin with irregular spots and small blotches, more
concentrated in distal portion; pectoral-fin base
blotch also present; well developed blotch also on
medial side of pectoral fin base. Dorsal, caudal
and anal fins grey with dark irregular mark-
ings. Dorsal-fin spines lighter than dorsal part
of body, with few dark spots; finlet membranes
translucent with irregular grey color markings,
more pronounced in distal part; coloration and
markings of finlet membranes similar to those of
soft dorsal fin, caudal fin and anal fin.
In preservative: color pattern similar to that
in live specimens, but generally less contrasted
(Fig. 29).
Two specimens from the Niger River at Diafa-
rabé, Mali, (MNHN 1961-0015, MNHN 1961-0016)
with unusual color pattern resembling that of
P. congicus, however, consisting of narrow rather
than broad bars and with ventrolateral markings
expressed as irregular blotches on ventral half of
flanks.
Distribution. Widely distributed in Nile, Chad
and Niger basins; and in coastal rivers of Côte
dIvoire, i. e. Sassandra River, Boubo River, Ban-
dama River, Ebrié Lagoon and Comoé River, and
Volta Basin; a single record from coastal lagoon
of Ouémé River (Fig. 31).
Etymology. Named after Stephan Ladislaus
Endlicher (1804-1849), an Austrian botanist and
colleague of Heckel.
Remarks. The exact date of the original descrip-
tion of P. endlicherii remains somehow obscure,
as the publication of Heckel is labelled as 1846-
1849. Kottelat (2013: 492) convincingly argued
that the most likely actual year of publication
of this article is 1848, which we follow here. The
corresponding figure for the original description
was not published in Heckel (1848), but in Fenzl
et al. (1849: plate 22).
There is some confusion about the holotype
of P. endlicherii. Heckel (1848) did not refer to any
collection number, but the description clearly
refers to a single specimen. About 50 years later,
Steindachner, then curator at the Vienna museum,
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 31. Records of Polypterus endlicherii. , type locality; , verified records; , other records.
Ichthyol. Explor. Freshwaters, IEF-1094
44
added Polypterus endlicheri Heck. Typ. to the la-
bel of NMW 63024, a spirit specimen of about the
same size of that in the original description. This
specimen was subsequently referred to by several
authors (Eschmeyer, 1998; Schäfer, 2004; Fricke
et al., 2018) as the holotype. In the original de-
scription, however, Heckel (1848) provided some
additional comments on the holotype, stating
among other things that it is a dry specimen with
12 dorsal-fin spines. Both these features clearly
exclude NMW 63024 from being the holotype.
There are two dry specimens of P. endlicherii in the
Vienna collection, NMW 91233 and NMW 92144,
both with 12 dorsal-fin spines. Only NMW 92144
(Fig. 27), however, has the matching size and its
color pattern corresponds quite well to that of the
drawing accompanying the original description.
In addition, the shape of the spiracular bones
on the skull roof of NMW 92144 matches that of
the same bones in the drawing of the holotype
(Fig. 28), with the exception of the second bone on
the right side, which is divided in three parts in
NMW 92144, but illustrated as a single bone. The
number and arrangement of the small spiracular
bones and the color pattern are quite variable in
the species and thus provide a useful source to
identify the holotype. Finally, a scale anomaly in
the 10th and 11th pre-dorsal scale row of NMW
92144 matches a similar anomaly in the draw-
ing of the holotype: the third scale of the tenth
row on the left side and the second scale of the
eleventh row seem to be fused (Fig. 28b; Fenzl et
al., 1849). This, in our opinon, demonstrates that
NMW 91244 is the holotype of P. endlicherii. It is
the specimen collected by Kotschy in Sudan, likely
in Khartoum, and sold to the NMW as Polypterus
bichir in 1840 (acquisition book 1840.XI.17). The
second dry specimen of P. endlicherii in Vienna,
NMW 91233, is very likely a specimen collected
on 30 Nov 1851 in Khartoum by C. Reitz (Reitz,
1852) and has no type status.
Polypterus mokelembembe
Schliewen & Schäfer, 2006
(Figs. 32-34, Tables 13-14)
Polypterus mokelembembe Schliewen & Schäfer,
2006: 24, fig. 1a.
Material examined. 32 specimens. type speCimens:
MRAC 87921, holotype (Fig. 32), female, 166 mm
SL; Democratic Republic of Congo: Maringa River
at Wamba, 0°11' N 22°28' E; H. Bertels, 8 Nov 1952. –
Fig. 32. Polypterus mokelembembe, MRAC 87921, 166 mm SL, female, holotype; Democratic Republic of Congo:
Maringa River; H. Bertels; 8 Nov 1952.
Moritz & Britz: Revision of Polypterus
45
MRAC 101372, 1, paratype, female, 221 mm SL; Demo-
cratic Republic of Congo: Bokuma, Hulstaert, 28 Aug
1955. – MRAC 79224-79225, 2, paratypes, 71-78 mm
SL; Democratic Republic of Congo: Bokuma, Lootens,
1-15 Jul 1952. – MRAC 72434-72435, 2, paratypes, 81-
172 mm SL; Democratic Republic of Congo: surround-
ing of Leopoldville. – BMNH 1907.12.3.4, 1, paratype
of P. mokelembembe and paralectotype of P. retropinnis,
male, 192 mm SL; Democratic Republic of Congo: Alima
River at Diele, S. de Brazza. – MNHN 1886-0297, 1, male,
paratype of P. mokelembembe and paralectotype of P. re-
tropinnis, 207 mm SL; Democratic Republic of Congo:
Alima River at Leketi, S. de Brazza. lefini river: MRAC
A4-046-P-0716, 1, 207.1 mm SL; Democratic Republic of
Congo: Itsotso River 5 km upstream of confluence with
Lefini River. – MRAC unreg., 1, 179 mm SL; Democratic
Republic of Congo: Lefini River, 17 Sep 2007. – MRAC
unreg., 6, 175.4-195 mm SL; Democratic Republic of
Congo: Lefini River, 8 Sep 2007. – MRAC unreg., 1,
218 mm SL; Democratic Republic of Congo: Lefini River,
15 Sep 2007. – MRAC unreg., 9, 127.3-244 mm SL; Demo-
cratic Republic of Congo: small tributary of Lefini River,
2 Sep 2008. – MRAC unreg., 1, 181 mm SL; Democratic
Republic of Congo: Lefini River, 17 Sep 2007. – MRAC
unreg., 1, 214 mm SL; Democratic Republic of Congo:
Lefini River, 19 Sep 2007. – MRAC unreg., 1, 261 mm
SL; Democratic Republic of Congo: Lefini River, 30 Apr
2007. aquarium traDe: ZSM 33571, 3, 194-232 mm SL;
Democratic Republic of Congo: exported via Kinshasa.
a
b
c
c
Fig. 33. Polypterus mokelembembe, aquarium import from Congo. a, 232 mm SL, female; b, head of specimen in a;
c, 149 mm SL, male.
Ichthyol. Explor. Freshwaters, IEF-1094
46
Table 14. Meristic characters of Polypterus mokelembembe. Numbers in brackets indicate number of specimens
with the respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 76 (7), 7 (17), 8 (3)
Dorsal- and caudal-fin rays 15 13 (1), 15 (16), 16 (10)
Pectoral-fin rays 27 24 (1), 25 (3), 26 (3), 27 (8), 28 (6), 29 (2), 30 (4)
Pelvic-fin rays 98 (3), 9 (14), 10 (9), 11 (1)
Anal-fin rays 9 (4), 10 (11), 11 (2), 12 (5), 13 (4)
Scales in longitudinal series 58 57 (2), 58 (8), 59 (12), 60 (4), 61 (1)
Scales around body 36 31 (1), 32 (4), 33 (3), 34 (5), 35 (9), 36 (5)
Pre-dorsal scales 33 30 (2), 31 (2), 33 (5), 34 (2), 35 (10), 36 (2), 37 (4)
Pre-pelvic scales 43 43 (1), 44 (3), 45 (4), 46 (11), 47 (6), 48 (2)
Total number of vertebrae 58 (2), 59 (3), 60 (1), 61 (1)
Table 13. Morphometric characters of Polypterus mokelembembe. Range, mean and standard deviation (SD) include
values of holotype.
holotype range mean SD n
Standard length [mm] 166.1 125.5-261 194.5 30.2 27
Total length [mm] 142.7-280 214.0 31.8 26
In percent of standard length
Body depth 10.0 9.5-12.8 11.2 0.9 27
Body width 9.9-12.4 11.0 0.6 26
Head length 16.9 14.8-16.9 15.8 0.6 27
Pre-dorsal length 59.2 52.8-66.3 59.8 3.2 27
Dorsal-fin length 32.5 26.8-40.0 32.3 3.2 27
Pectoral to dorsal fin 29.7 23.3-38.4 31.2 3.4 27
Pre-pectoral length 16.0 15.5-17.6 16.6 0.5 27
Pectoral-fin extension 26.8-30.6 28.5 1.0 26
Inter-pectoral width 6.1-8.0 7.0 0.5 26
Pectoral-fin base width 2.7-3.4 3.1 0.2 26
Pre-pelvic length 70.3 70.3-76.2 74.0 1.5 27
Pre-anal length 87.7 84-89 86.2 1.2 27
Anal-fin ray length 8.8-12.7 10.9 1.2 26
Anal-fin base length 4.2-7.5 6.0 0.9 26
Caudal-peduncle length 2.2-3.8 2.8 0.4 26
In percent of head length
Head width 65.2 56.6-71.2 62.4 3.3 27
Interorbital width 40.9 23.3-30.8 27.8 1.9 27
Nostril distance 20.1 11.1-16.2 13.7 1.5 27
Snout length 40.9 20.7-25.8 23.9 1.1 27
Eye diameter 25.6 12.6-16.3 14.3 0.9 27
Postorbital length 61.4-69.5 63.5 1.7 26
Gular length 53.5-65.9 58.0 2.9 26
Length of first spine 19.8-32.5 26.6 3.0 26
Length of second spine 25.9-34.5 30.8 2.6 26
Width : length of first spine 23.6-37.4 23.6 3.9 26
Length of second : first spine 98.4-133.7 133.7 9.2 26
Diagnosis. Polypterus mokelembembe is distin-
guished from all other species of Polypterus except
P. palmas, P. retropinnis and P. teugelsi by the
number of pre-dorsal scales (30-37 vs. 11-29); it
is distinguished from P. teugelsi and from all other
congeners except P. palmas, P. retropinnis, P. sene-
galus and P. weeksii by the number of pectoral-fin
rays (24-30 vs. 31-48). Polypterus mokelembembe is
Moritz & Britz: Revision of Polypterus
47
distinguished from P. teugelsi by fewer scales in
lateral series (57-61 vs. 64-65) and fewer pectoral-
fin rays (24-30 vs. 40-43). Polypterus mokelembembe
is distinguished from P. palmas and P. retropinnis
by a distinct continuous dark suborbital stripe
strongly contrasted against a plain background
(vs. suborbital stripe absent or hardly distinguish-
able from background coloration in P. palmas; and
a series of small spots rather than a continuous
suborbital stripe in P. retropinnis), dorsal half of
head with uniform grey or with slightly marbled
coloration (vs. dorsal half of head with dark grey
or brown marbling); margins of opercular and
spiracular bones clearly marked in dark grey or
almost black (vs. respective margins hardly or
not at all contrasted by coloration), and iris pure
orange without any dark speckles or stripes (vs.
grey with dark grey radial pattern in P. retropin-
nis, and white to yellow with numerous dark
spots in P. palmas, respectively). Furthermore,
P. mokelembembe differs from P. palmas in having
more pre-dorsal scales (30-37, median 35 vs.
21-31, median 26, respectively), fewer pectoral-
fin rays (24-30, median 27 vs. 30-38, median 35,
respectively) a distinct dark blotch on distal part
of pectoral-fin base (vs. absence of such blotch).
Polypterus mokelembembe differs from P. retropin-
nis by the number of pectoral-fin rays (24-30,
median 27 vs. 28-36, median 32, respectively),
color pattern of iris (uniform vs. with radial pat-
tern, respectively) and first dorsal-fin spine width
(7.2-9.8 % HL vs. 4.6-7.5, respectively).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 13-14. A small species,
maximum size 261 mm SL (280 mm TL). Body
moderately elongated with dorsal fin positioned
further posteriorly. Upper jaw slightly projecting
beyond lower jaw.
Coloration. In life: dorsal half of body light grey
to brown with 5-7 large dark brown irregular
dorsolateral saddle-like bars, sometimes conflu-
ent with blotch-like ventrolateral markings and
then extending onto upper part of ventral half of
flank. Ventral half of body light: white or yellow-
ish (Fig. 33). Border between coloration of upper
and lower half of flank not conspicuous. Dorsal
half of head uniform grey or with slight dark
grey marbling on light grey background; margins
of opercular and spiracular bones marked as
prominent black lines; lower half of cheek white;
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 34. Records of Polypterus mokelembembe. , type locality; , verified records; , other records.
Ichthyol. Explor. Freshwaters, IEF-1094
48
suborbital stripe distinct, expressed as black line
(Fig. 33b). Iris uniform orange without (in life
sometimes with very few) prominent markings.
Upper lip whitish, lower lip white with dark
brown dorsal margin and faint black line along
ventral margin. Tube of anterior nostril grey at
base, black at tip. Side of head below opercle and
gular plate uniformly whitish without blotches or
pattern, except for suborbital stripe.
Pectoral fin light beige with prominent distinct
dark lines becoming less distinct distally and
sometimes with scattered black spots; prominent
well-defined black blotch on distal part of lateral
pectoral-fin base. Pelvic fin white or yellowish
pale without any darker markings. Anal fin white
or yellowish pale with or without few black ir-
regular markings especially in distal part of fin.
Dorsal and caudal fin dark grey or brownish with
equally distributed, densely set, light yellowish
spots often forming lines. Dorsal-fin spine grey,
light grey or yellowish, with or without dark
markings in proximal part; finlet membrane
transparent grey with dark irregular black spot
in middle of fin membrane.
In preservative: black dorsolateral saddle bar
pattern less prominent; marbling on head absent;
ventral half of body cream to light beige (Fig. 32).
Iris uniformly grey without dark markings. Sub-
orbital stripe and pectoral-fin base blotch distinct.
Distribution. Central Congo basin, upstream of
Kinshasa up to Maringa River (Fig. 34). Not yet
recorded from Kasai, Oubangi or Congo above
confluence with Maringa River, as well as Congo
below Kinshasa.
Etymology. Named after the mythological crea-
ture Mokele-mbembe in reference to the archaic
nature of polypteriforms (Schliewen & Schaefer,
2006: 30). A noun in apposition.
Remarks. The type series of Polypterus retropin-
nis was originally composed of three syntypes and
actually contained representatives of two distinct
species. Schliewen & Schäfer (2006) designated
MNHN 1886-0295 as lectotype of P. retropinnis
The paralectotypes (BMNH 1907.12.3.4, MNHN
1886-0297) have been designated as paratypes of
Polypterus mokelembembe. Thus these specimens of
P. mokelembembe are at the same time paratypes of
P. mokelembembe and paralectotypes of P. retropin-
nis (see Schliewen & Schäfer, 2006).
The diagnostic difference in internostril dis-
tance between P. mokelembembe and P. retropinnis
reported by Schliewen & Schäfer (2006: 11.6-
13.7 % HL vs. 14.3-18.0) was not confirmed in
this study. Rather the two species differed in the
mean of this morphometric character only (11.1-
16.2 % HL, mean 13.7 vs. 12.7-20.5, mean 16.6).
Polypterus ornatipinnis Boulenger, 1902
(Figs. 35-37, Tables 15-16)
Polypterus ornatipinnis Boulenger, 1902a: 23, Pl. 7,
fig. 1.
Material examined. 63 specimens. type spe Cimen :
BMNH 1901.12.21.1, holotype (Fig. 35), female, 346 mm
SL; Democratic Republic of the Congo: Monsembe,
J. Weeks. COngO river anD its smaller triButaries:
IRSNB 14456, 1, 210 mm SL; Democratic Republic of
Congo: Libenge: Isato, Cremer and Neumann, 16 Dec
1947. – MNHN 1886-2093, 1, 451 mm SL; Democratic Re-
public of Congo: Congo River at Nganchou, S. de Brazza.
– MRAC 78112, 1, 323 mm SL; Democratic Republic of
Congo: Leopoldville [= Kinshasa], M. H. Pierret, 1951.
– MRAC A6-007-P-1145, 1, 285 mm SL; Democratic Re-
public of Congo: Lantusi River [tributary of Luala River]
at Kimpese, 4°51'14" S 13°50'46" E; 19 Sep 2005. – ZSM
35716, 1, 75.4 mm SL; Democratic Republic of Congo:
Congo River at Inga rapids, 5°27'19" S 13°35'24" E; U.
Schliewen and R. C. Schelly, 14 Jul 2005. – ZSM 37515,
1, 122.3 mm SL; Democratic Republic of Congo: Maiko
River at Lubutu road bridge, 0°11'11" N 25°31'47" E; U.
Schliewen, 18 Jul 2008. – ZSM 37703, 1, 176.8 mm SL;
Democratic Republic of Congo: Congo River upriver
of Maluku, 4°02'14" N 15°35'08" E; J. Schwarzer et al.,
12 Aug 2008. aruWimi river: BMNH 1920.5.20.1,
1, 339 mm SL; Democratic Republic of Congo: Ituri:
Avakubi, Bequart. – MRAC 2872, 1, 344 mm SL; Demo-
cratic Republic of Congo: Aruvimi River at Avakubi,
D. Christy. – MRAC 7107, 1, 413 mm SL; Democratic
Republic of Congo: Bafwasende, D. Christy. – MRAC
7115, 1, 332 mm SL; Democratic Republic of Congo:
Bosabangi, D. Christy. – MRAC 14575, 1, 348 mm
SL; Democratic Republic of Congo: Aruvimi River at
Avakubi, Bequart, 1 Jan 1914. itimBiri river: MRAC
49272, 1, 140.6 mm SL; Democratic Republic of Congo:
Buta, R. F. Hutsebout, 1937. – MRAC 75089, 1, 154.9 mm
SL; Democratic Republic of Congo: Ibembo, R. F. Hut-
sebout, 1950. kasai riv er anD tri Butaries: MRAC
14680, 1, 165 mm SL; Democratic Republic of Congo:
NGome River at NGombe, D. Schouteden. – MRAC
101935, 1, 245 mm SL; Democratic Republic of Congo:
Kasai River, P. Chiry, Sep 1955. – MRAC 119199-119201,
2, 308-379 mm SL; Democratic Republic of Congo:
Gungu, P. Toye, 1958. – MRAC 157787, 1, 234 mm SL;
Democratic Republic of Congo: Lumbebe River at Muita
Moritz & Britz: Revision of Polypterus
49
Luembe, Petchkowsky, Sep 1948. – MRAC 157788,
1, 465 mm SL; Angola: Luachimo River at Calonda,
8°23' S 20°32' E; E. Assuda, Jun 1959. – MRAC 157789, 1,
262 mm SL; Angola: Kasai River, 7°22' S 21°50' E; A. L.
Ferreira, 16 Jun 1961. – MRAC 78-006-P-001, 1, 544 mm
SL; Angola: Luembe River at Lumbomba, A de Ballot
Machado, Sep 1972. – MRAC 78-006-P-002, 1, 375 mm
SL; Angola: Luembe River at Lumbomba, A de Ballot
Machado, Sep 1972. – MRAC 89-055-P-0001, 1, 175 mm
SL; Democratic Republic of Congo: Kwilu River close
to Kisandji, L. Risch, 3 Apr 1988. lake eDWarD Basin:
MRAC 20651, 1, 287 mm SL; Democratic Republic of
Congo: plaine of Ruindi River, Flamand, 1928. linDi
river: BMNH 1919.9.10.2, 1, 422 mm SL; Democratic
Republic of Congo: Lindi River at Bafwasende, C.
Christy. – BMNH 1919.9.10.3, 1, 354 mm SL; Democratic
Republic of Congo: Lindi River at Bosabangi, C. Christy.
– MRAC 2866, 1, 490 mm SL; Democratic Republic of
Congo: Lindi River at Bafwasende, C. Christy. lualaBa
river anD triButaries: BMNH 1975.6.20.2, 1, 322 mm
SL; Democratic Republic of Congo: Lufira River at
Kaswabilenga, K. Banister. – BMNH 1975.6.20.3-4, 2,
345-422 mm SL; Democratic Republic of Congo: Lufira
River at Kasonga, K. Banister. – BMNH 1975.6.20.5,
1, 254 mm SL; Democratic Republic of Congo: Lufira
River at Kaswabilenga, K. Banister, 1974. – BMNH
1976.10.12.298, 1, 411 mm SL; Democratic Republic of
Congo: Lufira River at Kaswabilenga, K. Banister. –
IRSNB 20339, 1, 453 mm SL; Democratic Republic of
Congo: Lufira River at Kaswabilenga, G.-F. de Witte, 13-
27 Sep 1947. – IRSNB 20340, 1, 329 mm SL; Democratic
Republic of Congo: Lufira River at the base of mount
Sambwe, G.-F. de Witte, 10-14 Aug 1947. – MRAC
36114, 1, 156.4 mm SL; Democratic Republic of Congo:
Luvua Rifer at Kiambi, F. de Witte, 4-20 May 1931. –
MRAC 71465; Democratic Republic of Congo: Sombe
14 km upstream of Kiabo, B. Dewit, Apr 1948. – MRAC
178706, 1, 240 mm SL; Democratic Republic of Congo:
Lufira River at Kiubo, W. de Smet, 4 Mar 1956. – MRAC
178707, 1, 156.2 mm SL; Democratic Republic of Congo:
Lufira River at Kiubo, W. de Smet, 4 Mar 1956. – MRAC
79-0001-P-0005, 1, 578 mm SL; Democratic Republic of
Congo: Lufira River at Kaswabilenga, G.-F. de Witte,
13-27 Sep 1947. – MRAC 79-0001-P-0011, 1, 508 mm SL;
Democratic Republic of Congo: Kilwezi River [tributary
of Lufira River], G.-F. de Witte, 26-28 Jul 1948. – MRAC
79-0001-P-0012, 1, 354 mm SL; Democratic Republic of
Congo: Kilwezi River [tributary of Lufira River], G.-F. de
Witte, 26-28 Jul 1948. malagarazi river (triButary tO
lake tanganjika): IRSNB 8601, 1, 433 mm SL; Tanzania:
Malagarzi River, 25 Feb 1947. – MRAC 88877, 1, 379 mm
SL; Tanzania: Malagarzi River, M. Poll, 21 May 1947.
– MRAC 93-152-P-0324, 1, 194 mm SL; Tanzania: delta
of Malagarzi River, 5°13' S 29°48' E; L. de Vos, 20 Aug
1993. OuBangui river anD triButaries: MRAC 7373,
1, 292 mm SL; Democratic Republic of Congo: Poko,
D. Christy. – MRAC 7380, 1, 306 mm SL; Democratic
Republic of Congo: Poko, D. Christy, before 1919. –
MRAC 7388, 1, 270 mm SL; Democratic Republic of
Congo: Avakubi, D. Christy. – MRAC 59423, 1, 359 mm
SL; Democratic Republic of Congo: Bambesa, Vrydayh,
13 Mar 1939. – MRAC 77988, 1, 344 mm SL; Democratic
Republic of Congo: Dungu River at km 999 of royal
road Congo-Nile, Vleeschouwers, 1 Nov 1950. – MRAC
82-013-P-0421, 1, 365 mm SL; Democratic Republic of
Congo: Ouaka River downstream of Ngouakoubo River,
J. Kempeneers and L. de Vos, 24 Jan 1982. – MRAC
82-021-P-001, 1, 364 mm SL; Central African Republic:
Baidou River [tributary of Ouaka], J. P. Marquet, 1 May
1982. –MNHN 2008-2087, 2, 198-221 mm SL; Demo-
cratic Republic of Congo: Oubangui River at Binmbe
island; T. Roberts; 27 Feb 1988. – MNHN 2008-2090, 1,
123.4 mm SL; Democratic Republic of Congo: rapids
of Bembe 74 km upstream from Bangui, T. Roberts,
1 Feb 1988. – MNHN 2008-2094, 2, 205-232 mm SL;
Democratic Republic of Congo: Oubangui River at
Satema, T. Roberts, 25 Jan 1988. – MNHN 2008-2101,
2, 151.2-132.0 mm SL; Democratic Republic of Congo:
Oubangui River near Palombo 61-64 km upriver from
Bangui, T. Roberts, 2-5 Feb 1988. – MNHN 2008-2110, 1,
120.1 mm SL; Democratic Republic of Congo: Mboumou
about 10 km southwest of Rafai, T. Roberts, 1 Feb 1988.
– ZSM A-0218, 1, 99.5 mm SL; Democratic Republic of
Congo: Zobia, R. P. van Woensel, 1940-1945. sangah
river anD triButaries: MNHN 1977-0307, 1, 105.0 mm
SL; Cameroon: Ngoko-River, Depierre. – ZSM 30063,
1, 168.1 mm SL; Cameroon: Dja River between rapids
of Chollet and Nki falls, U. Schliewen; 26-28 Jul 1999.
Additional material. COngO river anD its smaller
triButaries: MRAC 43969, 1, Democratic Republic of
Congo: Leopoldville [= Kinshasa], A. Tinant, 1935. –
MRAC 20657, 1, Democratic Republic of Congo: Kim-
pese, D. Stukx. – MRAC 87-042-P-0002, 1, Democratic
Republic of Congo: Kisangani: Tshopo River 20 km
before hydroelectric dam, L. de Vos et al., 31 Jul 1987. –
MRAC 87- 042-P-0765, 1, Democratic Republic of Congo:
Tshopo River on km 122 of route Kisangani- Bafwaboli. –
MRAC 88-025-P-0647, 1, Democratic Republic of Congo:
Kisangani: Tshopo River above hydroelectric dam, L.
de Vos et al., Oct 1987. itimBiri river: IRSNB 15963, 1,
Democratic Republic of Congo: Itimbiri River at Ibembo,
H. Simeons, 16 Apr 1940. – MRAC 61059, 1, Democratic
Republic of Congo: Buta, R. F. Hutsebaut, 1949. – MRAC
63157, 1, Democratic Republic of Congo: Buta, R. F.
Hutsebaut, 1949. – MRAC 68386, 1, Democratic Republic
of Congo: Buta, R. F. Hutsebaut, 1949. – MRAC 71732,
1, Democratic Republic of Congo: Buta, R. F. Hutse-
baut, 1949. – MRAC 72725, 1, Democratic Republic of
Congo: Buta, R. F. Hutsebaut, 1949. kasai river anD
triButaries: MRAC 1332, 1, Democratic Republic of
Congo: Lusambo. – MRAC 14685, Moakechi River at Ka-
maiembi, D. Schouteden. – MRAC 68749, 1, Democratic
Republic of Congo: Mushie, Vleeschouwers. – MRAC
94688, 1, Democratic Republic of Congo: Lulua River at
Luebo, B. Worth, May 1954. – MRAC 177710, 1, Demo-
cratic Republic of Congo: Lulua River at Luluabourg, A.
de Roo, 11 Nov 1964. lualaBa river anD triButaries:
Ichthyol. Explor. Freshwaters, IEF-1094
50
IRSNB 20552, 1, Democratic Republic of Congo: Lufira
River close to Lake Upemba, G.-F. de Witte, 31 Oct
1947. – MRAC 314, 1, Democratic Republic of Congo:
Lofoi, Lemaire. – MRAC 30551-30552, 2, Democratic
Republic of Congo: Kiambi, P. Gerard, 1930. – MRAC
182229, 1, Democratic Republic of Congo: Dungu River
at Gangala na Bodio, M. Poll, Nov 1956. OuBangui river
anD triButaries: IRSNB 14687, 1, Democratic Republic
Table 15. Morphometric characters of Polypterus ornatipinnis. Range, mean and standard deviation (SD) include
values of holotype.
holotype range mean SD n
Standard length [mm] 346.0 75.4-578 291.4 120.6 63
Total length [mm] 370.0 86.1-613 314.0 125.6 62
In percent of standard length
Body depth 9.3 8.0-13.3 10.7 1.2 63
Body width 11.2 9.9-12.9 11.6 0.7 63
Head length 16.8 15.3-22.0 18.5 1.3 63
Pre-dorsal length 41.2 39.5-48.6 43.7 2.4 63
Dorsal-fin length 50.6 40.1-54.1 46.6 2.8 62
Pectoral to dorsal fin 11.9 7.2-17.2 11.8 2.3 63
Pre-pectoral length 17.2 14.7-23.4 18.9 1.6 63
Pectoral-fin extension 29.3 25.9-37.3 31.9 2.2 63
Inter-pectoral width 6.7 5.6-8.9 7.5 0.7 61
Pectoral-fin base width 3.1 2.6-3.9 3.4 0.3 63
Pre-pelvic length 68.5 65.7-72.0 68.9 1.2 63
Pre-anal length 87.0 82.3-89.1 87.1 1.1 63
Anal-fin ray length 10.5 9.1-13.2 11.2 1.0 63
Anal-fin base length 4.4 4.0-7.2 5.5 0.8 63
Caudal-peduncle length 2.6 1.5-3.8 2.7 0.4 63
In percent of head length
Head width 63.9 55.8-72.9 64.2 3.6 63
Interorbital width 30.2 24.1-35.8 29.6 2.6 63
Nostril distance 17.1 9.9-20.1 15.8 1.6 63
Snout length 24.2 19.5-28.3 24.5 1.7 63
Eye diameter 10.3 9.1-16.7 11.7 1.8 63
Postorbital length 64.7 59.5-69.1 64.9 1.9 63
Gular length 70.1 52.7-73.2 59.5 5.3 63
Length of first spine 24.8 15.7-31.8 22.5 3.2 63
Length of second spine 26.4 18.2-32.7 25.5 3.1 63
Width : length of first spine 24.8 17.1-36.6 26.1 3.8 63
Length of second : first spine 106.4 90.9-161 113.8 10.8 63
Table 16. Meristic characters of Polypterus ornatipinnis. Numbers in brackets indicate number of specimens with
the respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 11 8 (1), 9 (18), 10 (32), 11 (11), 12 (1)
Dorsal- and caudal-fin rays 15 14 (1), 15 (8), 16 (50), 17 (4)
Pectoral-fin rays 37 31 (2), 32 (4), 33 (6), 34 (16), 35 (14), 36 (12), 37 (7), 38 (1), 39 (1)
Pelvic-fin rays 14 9 (1), 10 (3), 11 (13), 12 (25), 13 (18), 14 (3)
Anal-fin rays 17 9 (2), 10 (2), 11 (5), 12 (15), 13 (22), 14 (11), 15 (5), 17 (1)
Scales in longitudinal series 62 57 (1), 58 (1), 60 (7), 61 (17), 62 (21), 63 (12), 64 (4)
Scales around body 40 39 (3), 40 (10), 41 (10), 42 (19), 43 (10), 44 (8), 45 (2), 48 (1)
Pre-dorsal scales 24 22 (3), 23 (10), 24 (17), 25 (17), 26 (11), 27 (4), 29 (1)
Pre-pelvic scales 45 42 (2), 43 (10), 44 (19), 45 (13), 46 (13), 47 (4), 48 (1), 49 (1)
Total number of vertebrae 59 59 (1), 62 (1)
Moritz & Britz: Revision of Polypterus
51
of Congo: Oubangui River at Motenge-Boma, Cremer
and Neumann, 2 Jan 1948. – MRAC 23613, 1, Democratic
Republic of Congo: Nyangara, D. Schouteden. – MRAC
15563-15566, 4, Democratic Republic of Congo: Faradje,
Young-Chapin. – MRAC 166244, 1, Democratic Republic
of Congo: Oubangui River at Isato, R. Cremer and M.
Neumann, 16 Nov 1947. ruki river: IRSNB 21049, 1,
Democratic Republic of Congo: Yonda River 10 km
downstream of Mbandaka, Kiss, Feb 1980. sangah
river anD triButaries: DMM IE/5527, 1, juvenile,
Cameroon: Dja River, T. Moritz and J. Schwarzer, Feb
2008. – DMM IE/10407, 1, Cameroon: Boumba River
[tributary of Dja River], T. Moritz and J. Schwarzer, 28
Jan 2008. – DMM IE/10408, 1, Cameroon, Dja River:
Nki falls, T. Moritz & J. Schwarzer, 9 Feb 2008. – DMM
IE/10409, 1, Cameroon, Dja River: downstream of Nki
falls, T. Moritz & J. Schwarzer, 9 Feb 2008.
Diagnosis. Polypterus ornatipinnis is distin-
guished from all other Polypterus species by its
unique color pattern consisting of black color
markings expressed as numerous black spots
and lines, often confluent or contiguous forming
irregular lines and networks on head and extend-
ing down to lower jaw and including gular plates
(vs. a distinct color pattern). Polypterus ornatipin-
nis is distinguished from P. ansorgii, P. bichir,
P. congicus, P. endlicherii and P. delhezi by more
numerous pre-dorsal scales (22-29 vs. 10-17),
from P. mokelembembe by fewer scales in pre-dorsal
row (22-29 vs. 30-37), and from P. teugelsi by the
number of pectoral-fin rays (31-39 vs. 40-43, re-
spectively). Polypterus ornatipinnis is distinguished
from P. palmas and P. retropinnis by more scales
around body (39-48 vs. 31-38, rarely 39 in P. ret-
ropinnis). Polypterus ornatipinnis is distinguished
from P. polli by more pre-pelvic scales (42-49 vs.
34-40, respectively). Polypterus ornatipinnis differs
from P. senegalus by more numerous pre-dorsal
scales (22-29, median 25 vs. 15-23, median 17),
more scales around body (39-48, median 42 vs.
32-44, median 36) and more pre-pelvic scales (42-
49, median 45 vs. 35-43, median 38). In meristic
and morphometric characters it is very similar
to P. weeksii from which it is distinguished by
its coloration as follows: suborbital stripe incon-
spicuous (vs. conspicuous), distinct pectoral-fin
base blotch absent (vs. present), network of light
beige to whitish spots on a dark brown to black
background (vs. more or less distinct bars), black
spots and lines on gular plates (vs. gular plates
uniform beige to light brown). Furthermore,
P. ornatipinnis is distinguishable from P. weeksii
by a shorter head (15.3-22.0 % SL, median 18.5
vs. 18.3-24.9, median 21.2).
Fig. 35. Polypterus ornatipinnis, BMNH 1901.12.21.1, 346 mm SL, female, holotype; Democratic Republic of the
Congo: Monsembe; J. Weeks.
Ichthyol. Explor. Freshwaters, IEF-1094
52
a
b
c
d
e
Moritz & Britz: Revision of Polypterus
53
Descriptive synopsis. Meristic and morpho-
metric data in Tables 15-16. Large species, at
least 578 mm SL (613 mm TL). Body moderately
elongated. Upper jaw slightly projecting beyond
lower jaw.
Coloration. In life: dorsal half of body black
with numerous densely set whitish or yellowish
roughly circular blotches and interspersed spots
(Fig. 36). Ventral half of body white or yellowish;
no clear demarcation between dorsal and ventral
coloration. Dorsal and lateral parts of head black
with irregular light whitish or yellowish densely
set irregular spots sometimes like marbling;
slightly larger spots only on cheeks; lips black
with light spots giving a striped appearance;
nasal barbels patterned like head. Coloration of
ventral half of head comparable to that of dorsal
half, but lighter with whitish background and
black marbling. Color pattern of head continuing
onto medial and lateral bases of pectoral fin and
external gills (when present).
Pectoral fin yellowish, in proximal part with
one or two irregular bands; distal part yellowish
with black spots (Fig. 36b). Pectoral-fin base whit-
ish with irregular black lines or dots. Pelvic and
anal fins yellowish with black markings surround-
ing yellow spots; often with black distal margin.
Dorsal fin including finlet membranes and caudal
fin black with yellowish spots sometimes form-
ing yellowish lines. Dorsal-fin spine black with
yellow proximal and subdistal bands, continuing
into black finlet membrane. Color pattern present
from juvenile stage on (Fig. 36c).
In preservative: all yellowish light areas pale
white; otherwise similar to coloration in life, but
color pattern on gular plates often less prominent
than in life.
Distribution. Polypterus ornatipinnis is widely
distributed all across Congo basin; few records
from delta of Malagarazi River (Lake Tanganyika)
and single record from affluent (Ruindi River
plane) of Lake Edward (Fig. 37).
Fig. 36. Polypterus ornatipinnis. a-b, 207 mm SL, aquar-
ium import from Democratic Republic of Congo;
c, 38 mm SL, small juvenile; Cameroon: Dja River at
Nki falls; d, 55 mm SL, juvenile; Cameroon: Dja River
at Nki falls; e, 135 mm SL; Cameroon: Boumba River
[tributary of Dja River].
/
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 37. Records of Polypterus ornatipinnis. , type locality; , verified records; , other records.
Ichthyol. Explor. Freshwaters, IEF-1094
54
Biology. The species is often kept in aquaria;
spawning and larval development was described
by Britz & Bartsch (1998).
Etymology. The specific name ornatipinnis ap-
parently refers to the ornate coloration of its fins.
Remarks. Polypterus ornatipinnis potentially oc-
curs also in Lake Rukwa (see Seegers, 1996).
Brichard (1978) suggested that P. ornatipinnis
may have already been present in the Malagarazi
River when it was still a tributary of the Congo,
before formation of Lake Tanganyika. Poll (1941a)
doubted the correctness of the information on the
label for the Ruindi specimen and excluded P. or-
natipinnis from his list of Lake Albert fish species
(Poll, 1939). It is surprising that this species has
not been recorded yet from the Ruki River and
its tributaries, e. g. Tshuapa River, at the center
of the Congo basin, with the exception of a single
specimen, which was caught close to the conflu-
ence of Ruki and Congo River.
Polypterus palmas Ayres, 1850
(Figs. 38-46, Tables 17-18)
Polypterus palmas Ayres, 1850: 241, Pl. 6.
Polypterus buettikoferi Steindachner, 1891: 179.
New synonym.
Polypterus lowei Boulenger, 1911: 377.
Material examined. 135 specimens. CasamanCe river:
MRAC 73-005-P-0034, 1, 278 mm SL; Senegal: Casa-
mance River at Kolda, D. F. E. van den Audenaerde,
29-31 Mar 1966. – MRAC 73-005-P-0035, 1, 242 mm
SL; Senegal: Casamance River at Kolda, D. F. E. van
den Audenaerde, 5 Apr 1966. Cavally river: MNHN
1987-1167, 1, 254 mm SL; Côte dIvoire: Cavally River
at Saibly near Toule-Pleu, K. Traore and G. Teugels,
16 May 1986. – MRAC 140113, 1, 153.6 mm SL; Liberia:
environs of Monrovia, D. Bout. – MRAC 73-005-P-0017-
0022, 6, 136.0-239 mm SL; Côte dIvoire: Cavally River
at Flanpleu, D. F. E. Thys van den Audenaerde, 20-24
Jul 1966. – MRAC 73-010-P-0001-0005, 5, 255-269 mm
SL; Côte dIvoire: Cavally River at Sahibili, D. F. E. Thys
van den Audenaerde, 18 Mar 1969. – MRAC 73-010-P-
0006-0007, 2, 227-233 mm SL; Côte dIvoire: Cavally
River at Sahibili, D. F. E. Thys van den Audenaerde,
19-20 Mar 1969. – MRAC 73-010-P-0008-0009, 2, 263-
299 mm SL; Côte dIvoire: Cavally River at Sahibili, D.
F. E. Thys van den Audenaerde, 20 Mar 1969. – MRAC
80-036-P-0009, 1, 88.0 mm SL; Liberia: Cavally basin at
Pahlwoken, M. Louette and P. Rigaux, 21 Mar 1980. –
MRAC 80-036-P-0010, 1, 140.2 mm SL; Liberia: Cavally
basin at Pahlwoken, M. Louette and P. Rigaux, 22 Mar
1980. – MRAC 80-036-P-0016, 1, 220 mm SL; Liberia:
Cavally basin at Zleh Town, M. Louette and P. Rigaux,
31 Mar 1980. – MRAC 80-036-P-0017-18, 2, 213-220 mm
SL; Liberia: Cavally basin at Zleh Town, M. Louette
and P. Rigaux, 1 Apr 1980. COruBal river [= kOliBa
river]: BMNH 1910.11.28.1-2, 2, 85.2-66.1 mm SL;
Guinea-Bissau: Corubal River at Tchitole, J. Weeks. –
NMW 63245, 1, 78.0 mm SL; Guinea-Bissau: Corubal
River at Tchitole, Steindachner. fatala Basin: MNHN
2000-5138, 8, 64.5-299 mm SL; Guinea: tributary of Fatala
River, R. Bigorne, 11 May 1993. – MNHN 2000-5139, 2,
138.6-164.5 mm SL; Guinea: tributary of Fatala River,
R. Bigorne, 11 May 1993. kOgOn river: MNHN 1990-
0460, 1, 329 mm SL; Guinea: Kogon River at Wendou-
Dorou, D. Paugy and R. Bigorne, Mar 1989. – MNHN
2000-5131, 2, 247-292 mm SL; Guinea: Kogon River at
Kogon, R. Bigorne, 13 Apr 1992. – MNHN 2000-5132, 3,
313-367 mm SL; Guinea: Kogon River at Kogon, R. Big-
orne, 12 Apr 1992. – MNHN 2000-5135, 5, 187-367 mm
SL; Guinea: Kogon River at Samba Sombe, R. Bigorne,
13 Apr 1992. – MNHN 2000-5136, 3, 295-348 mm SL;
Guinea: Kogon River at Samba Sombe, R. Bigorne, 13
Apr 1992. – MRAC 92-059-P-0010-0020, 6, 259-353 mm
SL; Guinea: Kogon River at Kogon, 11°22' N 13°55' W; G.
Teugels et al., 10 Apr 1992. – NMW 63246, 1, 78.2 mm
SL; Guinea: [likely] Kogon River [labelled as Kongo],
Steindachner, 1909. kOlente river: MNHN 1961-0017,
3, 99.6-130.0 mm SL; Guinea: Kolente River at Seguea,
J. Daget, Feb 1958. – MNHN 1991-0295, 1, 263 mm SL;
Guinea: Kolente River at Kolente, G. Teugels, 15 May
1987. – MNHN 2000-5133, 1, 147.1 mm SL; Guinea: Bal-
isso River, R. Bigorne, Mar 1992. – MNHN 2000-5134,
9, 169-284 mm SL; Guinea: Kolente River, R. Bigorne,
21 Nov 1991. – MNHN 2000-5137, 3, 84.1-98.7 mm SL;
Guinea: Balisso River at Yembere, R. Bigorne, Mar
1992. kOnkOure river: MNHN 1935-0196, 1, 215 mm
SL; Guinea: Mayonkoure River at Kindia, Waterlot.
– MNHN 1961-0018, 107.5 mm SL; Guinea: Labe, J.
Daget. – MNHN 2002-0790, 1, 261 mm SL; Guinea:
Konkoure River at Badi Baki, R. Bigorne, 7 May 1992.
mafa river: NMW 63207, 2, 76.3-83.9 mm SL; paralec-
totypes of P. buettikoferi, Liberia: Mafa River at Buluma,
1893. – RMNH 5332, 5, 74.7-104.1 mm SL; Liberia:
Mafa River, A. F. Demery, before 1889. – RMNH 5371,
2, 108.5-179 mm SL; paralectotype of P. buettikoferi,
Liberia: Buluma: swamps of fishermen lake, J. Büttikofer
and J. A. Sala, 1881 (Fig. 40). nanna Cru river: BMNH
1911.5.31.1-4, 4, 122.4-198 mm SL; Liberia: Nanna Cru,
Janson (Fig. 41). nipOue (= CestOs) river: MNHN 1963-
0239, 1, 253 mm SL; Côte dIvoire: Boan River at Danane,
J. Daget, Jul 1963. – MNHN 1979-0190, 1, 246 mm SL;
Côte dIvoire: Boan River at Danane, 7°21' N 8°10' W;
C. Lévêque. – MNHN 2000-0859, 7, 207-243 mm SL;
Côte dIvoire: Nipoué River at Toyebli, 6°37' N 8°29' W;
R. Bigorne, 20 Mar 1988. – MRAC 73-005-P-0023, 1,
146.8 mm SL; Côte dIvoire: Nipoué River at Toyebli,
D. F. E. Thys van den Audenaerde, Jul 1969. – MRAC
73-010-P-0010-0013, 4, 232-277 mm SL; Côte dIvoire:
Moritz & Britz: Revision of Polypterus
55
Nipoué River at Toyebli, D. F. E. Thys van den Aude-
naerde, 18 Mar 1969. – MRAC 73-010-P-0014-0015, 2,
208-227 mm SL; Côte dIvoire: Nipoué River at Toyebli,
D. F. E. Thys van den Audenaerde, 20 Mar 1969 (Fig. 39).
– MRAC 80-036-P-0002-0004, 3, 88.4-232 mm SL; Libe-
ria: south of L.T.C. [?]: Nipoué River, M. Louette and
P. Rigaux, 29 Feb 1980. – MRAC 80-036-P-0005-0006, 2,
190.4-241 mm SL; Liberia: south of L.T.C. [?]: Nipoué
River, M. Louette and P. Rigaux, 3 Mar 1980. – MRAC
85-029-P-0005-0006, 2, 270-272 mm SL; Côte dIvoire:
Table 17. Morphometric characters of Polypterus palmas.
range mean SD n
Standard length [mm] 64.5-367 205.7 76.1 122
Total length [mm] 78.4-411 233.4 82.0 120
In percent of standard length
Body depth 8.8-13.8 10.9 1.1 122
Body width 8.5-12.8 11.1 0.9 122
Head length 15.1-22.6 17.6 1.7 122
Pre-dorsal length 40.4-56.8 48.8 4.1 122
Dorsal-fin length 33.2-50.3 41.7 4.1 122
Pectoral to dorsal fin 10.4-29.4 18.3 4.6 122
Pre-pectoral length 15.2-23.8 18.1 1.8 122
Pectoral-fin extension 25.0-37.2 30.2 2.4 122
Inter-pectoral width 5.1-10.3 7.2 0.9 122
Pectoral-fin base width 1.7-3.9 3.2 0.3 122
Pre-pelvic length 66.3-74.3 70.5 1.3 122
Pre-anal length 79.9-90.0 86.9 1.4 122
Anal-fin ray length 8.7-14.4 11.3 1.1 122
Anal-fin base length 3.7-7.6 5.4 1.0 122
Caudal-peduncle length 1.4-4.3 2.7 0.5 122
In percent of head length
Head width 51.1-71.2 62.0 4.2 122
Interorbital width 22.7-32.1 28.3 2.0 121
Nostril distance 9.3-21.3 16.3 2.1 120
Snout length 19.1-27.2 23.3 1.7 121
Eye diameter 9.5-18.0 12.5 1.9 121
Postorbital length 60.2-70.7 64.7 1.8 121
Gular length 52.5-72.4 59.8 4.0 121
Length of first spine 13.9-31.4 23.5 3.5 121
Length of second spine 15.2-34.1 25.9 3.7 121
Width : length of first spine 18.6-39.1 27.8 4.1 121
Length of second : first spine 73.6-143.1 110.9 12.2 121
Table 18. Meristic characters of Polypterus palmas. Numbers in brackets indicate number of specimens with the
respective value.
all material
Dorsal-fin spines 7 (27), 8 (62), 9 (28), 10 (5)
Dorsal- and caudal-fin rays 14 (4), 15 (37), 16 (73), 17 (8)
Pectoral-fin rays 30 (2), 31 (4), 32 (4), 33 (19), 34 (21), 35 (30), 36 (25), 37 (11), 38 (6)
Pelvic-fin rays 8 (20), 9 (41), 10 (45), 11 (13), 12 (3)
Anal-fin rays 7 (1), 8 (1), 9 (6), 10 (19), 11 (17), 12 (26), 13 (31), 14 (17), 15 (4)
Scales in longitudinal series 50 (1), 53 (2), 54 (2), 55 (5), 56 (42), 57 (48), 58 (20), 59 (2)
Scales around body 31 (1), 33 (7), 34 (30), 35 (25), 36 (34), 37 (13), 38 (12)
Pre-dorsal scales 21 (1), 22 (6), 23 (13), 24 (15), 25 (23), 26 (15), 27 (12), 28 (7), 29 (9), 30 (11), 31 (10)
Pre-pelvic scales 39 (5), 40 (10), 41 (34), 42 (39), 43 (26), 44 (5), 45 (3)
Total number of vertebrae 53 (4), 54 (4), 55 (14), 56 (19), 57 (12), 58 (2)
Ichthyol. Explor. Freshwaters, IEF-1094
56
Nipoué River at Binhouye, 6°47' N 8°19' W; G. Teugels,
28-29 Apr 1985. – MRAC 86-018-P-0001-0002, 2, 220-
244 mm SL; Côte dIvoire: Nipoué River at Binhouye,
K. Traore, 14 May 1986. riO nunez: MRAC 92-059-P-
0001-0009, 9, 176.3-300 mm SL; 11°00' N 14°19' W; G.
Teugels et al., 8 Apr 1992. saint paul river: MNHN
1940-0069, 1, 216 mm SL; Liberia: Saint Paul River at
Monrovia, Bouet. sangWin river: BMNH 1969.11.19.7,
1, 74.5 mm SL; Liberia: Grand Gedeh County, A. Fraser-
Brunner. seWa anD mOa rivers: BMNH 1915.5.21.1, 1,
202 mm SL; Sierra Leone: Pejehun, N. Thomas. – BMNH
1968.9.18.1, 1, 200 mm SL; Sierra Leone: Kwarko Lake,
T. S. Jones. – MRAC 73-010-P-0018, 1, 251 mm SL; Sierra
Leone: Waanje River at Pujehun, D. E. F. Thys van den
Audenaerde, 16 Apr 1969. – RMNH 32818, 1, 187 mm
SL; paralectotype of P. buettikoferi, Sierra Leone: Sulima
River at Juring, A. F. Demery, before 1890. – RMNH 5396,
1, 238 mm SL; lectotype of P. buettikoferi, Sierra Leone:
Sulima River at Juring, A. F. Demery, before 1890. sierra
leOne (WithOut further lOCality): BMNH 1866.1.28.3,
1, 240 mm SL; Sierra Leone, Stevens. – BMNH 1911.4.6.1,
1, 122.4 mm SL; Sierra Leone. unClear lOCality: NMW
63208, 1, 188 mm SL; paralectotype of P. buettikoferi,
[jar labelled with Buluma × Sulymah, thus originat-
ing either from Liberia: Mahfa River at Buluma or
Sierra Leone: Moa River at Sulymah = Sulima], 1893.
– MNHN 1933-0061, 99.8 mm SL; [labelled as Koba-
River, Guinea – the Niokolo-Koba River, however, is
located in Senegal], Laborey.
Additional material. Cavally river: DMM IE/10395,
1, Côte dIvoire: Tai National Park: small tributary of
Audrenisrou River, K. Mody, 1 May 2000. – MRAC
80-036-P-0025, 1, Liberia: Cavally basin at Zleh Town,
M. Louette and P. Rigaux, 2 Apr 1980. granD Cess:
MRAC 80- 036-P-0008, 1, Liberia: Grand Cess at Juok-
woke, M. Louette and P. Rigaux, 20 Mar 1980. kOgOn
river: MRAC 92-059-P-0021, 1, Guinea: small tributary
of Kogon River at Pabole, 11°21' N 14°08' W; G. Teugels
et al., 12 Apr 1992. – MRAC 92-059-P-0022-0025, 4,
Fig. 38. Polypterus palmas, illustration of holotype from original description (Ayres, 1850).
Fig. 39. Polypterus palmas, MRAC 73-010-P-0014-0015, 227 mm SL, male, non type; Côte dIvoire: Nipoué River
at Toyebli; D. F. E. Thys van den Audenaerde; 20 Mar 1969.
Moritz & Britz: Revision of Polypterus
57
Guinea: Kogon River at Samba Sobe, 11°30' N 13°59' W;
G. Teugels et al., 13 Apr 1992. – MRAC 92-059-P-0026-
0054, 28, Guinea: Nguelodi River [tributary of Kogon
River] close to Kogon, 11°22' N 13°55' W; G. Teugels et al.,
12 Apr 1992. nipOue (= CestOs) river: MRAC 80-036-
P-0001, 1, Liberia: Duzei, M. Louette and P. Rigaux, 29
Feb 1980. saint jOhn river: MRAC 80-036-P-0011-0015,
5, Liberia: Duo Down, M. Louette and P. Rigaux, 27 Mar
1980. sangWin river: MRAC 80-036-P-0019-0023, 5, Li-
beria: Sangwin basin at Jusohn, M. Louette and P. Rigaux,
1 Apr 1980. – MRAC 80-036-P-0024, 1, Liberia: Sangwin
basin at Chesla, M. Louette and P. Rigaux, 2 Apr 1980.
Diagnosis. Polypterus palmas is distinguished
from P. ansorgii, P. bichir, P. congicus, P. endlicherii
and P. delhezi by more pre-dorsal scales (21-31 vs.
12-17). Polypterus palmas is distinguished from
P. teugelsi by fewer scales in lateral series (50-59
vs. 64-65). Polypterus palmas is distinguished
from P. ornatipinnis and P. weeksii by fewer scales
around body (31-38 vs. 39-51). Polypterus palmas
is distinguished from P. mokelembembe by absence
of distinct suborbital stripe (vs. presence), absence
of dark blotch on pectoral-fin base (vs. pres-
ence) and more pectoral rays (30-35 vs. 24-30).
Polypterus palmas is distinguished from P. sene-
galus by dark irregular anastomosing markings
on body formed by confluence of dorsal with
lateral blotches and ventrolateral markings (vs.
uniform, no dark markings on flanks) and more
pre-dorsal scales (21-29, median: 26 vs. 15-23,
median 17). Polypterus palmas is distinguished
from P. retropinnis by complete absence of distinct
pectoral-fin base blotch (vs. presence of indistinct
blotch), irregularly marbled iris (vs. iris with
regular dark radial stripes) and yellow marbling
or spots on head and dorsal half of body (vs.
light grey spots or marbling). Polypterus palmas
is distinguished from P. polli by combination of
number of dorsal-fin spines (7-10 vs. 5-8) and
pre-pelvic scales (39-45 vs. 34-40) (Fig. 46): speci-
mens with overlapping count of 7 or 8 dorsal-fin
spines distinguishable by 39-45 pre-pelvic scales
in P. palmas (vs. 35-38 in P. polli).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 17-18. Maximum size at
least 367 mm SL (411 mm TL). Body moderately
elongated with dorsal fin positioned far back.
Upper jaw projecting beyond lower jaw.
Coloration. In life: dorsal half of body light grey
to brown with dorsal and lateral blotches conflu-
ent forming anastomosing network of 5-9 irregu-
lar darker bars running from mid-dorsal area in
Fig. 40. Polypterus palmas, RMNH 5371, 179 mm SL, male, paralectotype of P. buettikoferi; Liberia: Buluma: swamps
of fishermen lake; J. Büttikofer & J. A. Sala; 1881.
Ichthyol. Explor. Freshwaters, IEF-1094
58
caudo-ventral direction; numerous, often densely
set, lighter, usually yellowish spots, surrounded
by these dark markings; number and size of lighter
spots as well as intensity and conspicuousness of
dark anastomosing bars varies with geographical
origin (Fig. 42). Ventral half of body uniformly
white, sometimes yellowish; clear demarcation
between dorsal and ventral coloration of body.
Dorsal half of head light grey to brown with
irregular yellow marbling or yellow spots; size,
amount and detailed pattern of this yellow mark-
ings varying with geographical origin. Lower half
of head uniform white or yellowish as lower half
of body; suborbital stripe poorly developed: not
continuous and expressed as thin marbled line or
sometimes absent. Iris, upper lip and nasal bar-
bel same coloration as neighbouring head areas;
same color pattern also on lower lip in specimens
from southern part of distribution, but absent in
specimens from northern part of distribution.
Pectoral fin densely covered with black and
yellow spots; base of pectoral fin with or without
large greyish area, often covered with yellow
spots, but not forming distinct blotch. Pelvic fin
yellowish with no or little grey spots. Anal fin
pigmented like pelvic fin, but with more grey
spots. Soft part of dorsal fin and caudal fin dark
grey to black with dense yellow spotting. Finlet
membranes with few yellow spots but predomi-
nantly black, more conspicuous in specimens from
northern part of distribution, in which black col-
oration extends onto dorsal-fin spines; specimens
from the south of distribution having black area of
finlet membranes less defined and not extending
onto spines. Differences in color pattern hugely
variable and not strictly geographically correlated.
In preservative: all yellowish color patterns
in life pale white or brownish; light spots and
stripes in dorsal part of body less conspicuous
(Figs. 39-41).
Fig. 41. Polypterus palmas, BMNH 1911.5.31.1-4, 198 mm SL, male, syntype of P. lowei; Liberia: Nanna Cru; Janson.
Moritz & Britz: Revision of Polypterus
59
Distribution. Upper Guinea region (Fig. 43):
coastal river basins from Casamance River in
Senegal to Cavally basin in Côte dIvoire.
Etymology. Apparently named after the type
locality at Cape Palmas in Liberia close to the
border of Côte dIvoire.
Remarks. The original description was based
on a single specimen collected at Cape Palmas,
Liberia by Perkins (Fig. 38). This holotype was
originally deposited in the Boston Museum. The
respective collection was later transferred to the
Museum of Comparative Zoology at Harvard, but
the Polypterus palmas type was never received and
is probably lost (Hanssens et al., 1995).
Steindachner frequently published short notes
on new species descriptions predating his more
detailed descriptions. He likely intended to do
the same with his description on P. buettikoferi
(original spelling büttikoferi, mandatory change
to buettikoferi following Art. 32.5.2.1 of the Code),
a species named honouring Johann Büttikofer
(1850-1927) a Swiss zoologist, who, during two
expeditions (1879-1882, 1886-1887), collected
zoological specimens in Liberia, among them the
type series of P. buettikoferi. Soon after publishing
his note (Steindachner, 1891), Steindachner (1895)
must have been convinced that his P. buettikoferi
was a junior synonym of Polypterus palmas Ay-
res, 1850, and thus a more detailed description
of P. buettikoferi was never published. The type
localities provided in Steindachners note (1891)
are Buluma (Liberia), the Mahfa River (also Mafa
and Marfa, Liberia) and Juring (Sierra Leone).
Hanssens et al. (1995: 703) designated RMNH 5396
a
b
b
c d
Fig. 42. Polypterus palmas. a-b, 190 mm SL, aquarium import from Guinea; c-d, ~299 mm SL; Côte dIvoire: Tai
National Park: small tributary of Audrenisrou River. Images c-d with sides reversed (Photos c-d by K. Mody).
Ichthyol. Explor. Freshwaters, IEF-1094
60
as lectotype of P. buettikoferi stating that Steindach-
ners description was performed . . . on the base
of nine specimens . . .. Steindachner, however,
never noted how many specimens he studied
for his description. In addition to the material
recognized as part of the type series by Hanssens
et al. (1995), an additional five specimens collected
by Büttikofer from the locations provided in
Steindachners (1891) description had been sent
from the RMNH to Steindachner in Vienna, who
registered at least four of them in 1893 in NMW.
These specimens should be regarded as original
syntypes, now paralectotypes, of P. buettikoferi.
The four specimens where originally registered
as NMW 63207 and NMW 63208, however the
latter contains only a single specimen suggesting
loss of the fourth NMW paralectotype.
Boulenger (1911) described Polypterus lowei
from the Nanna Kru River in Liberia. Four syn-
types are known (BMNH 1911.5.31.1-4, Fig. 41).
The respective specimens, however, cannot be
distinguished from P. palmas and we therefore,
like Gosse (1988), regard P. lowei as a junior syno-
nym of P. palmas. Other reports of P. lowei from
Africa west of Ghana (e. g. Pellegrin, 1923a; Holly,
1933) also most likely refer to P. palmas. Specimens
identified as P. lowei and originating from Cen-
tral Africa (e. g. Steindachner, 1912; Holly, 1930,
1933) may represent P. polli, or, though less likely,
P. retropinnis.
Hanssens et al. (1995) treated P. buettikoferi as a
valid subspecies of P. palmas based on differences
in the number of pre-dorsal scales and of dorsal-
fin spines, with both characters, however, show-
ing substantial overlap and differing only in their
medians. As a further difference, these authors
cited the geographical distribution: Polypterus
buettikoferi was claimed to occur in the coastal riv-
ers from Senegal to the St. Paul River in Northern
Liberia and P. palmas was stated to be restricted
to the Southern half of Liberia and the western
Ivory Coast. Hanssens et al. (1995) considered
Gosses (1988) P. polli to represent a subspecies of
P. palmas restricted to the Congo basin. These same
authors also claimed that Congolese P. palmas
polli differed from the Upper Guinean P. palmas
palmas in the number of dorsal-fin spines and of
scales in a longitudinal series. Again, differences
were only found for the medians of these ranges,
which showed substantial overlap. Hanssens et
al.s (1995) PCA included five meristic characters
and was able to distinguish specimens from the
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 43. Records of Polypterus palmas. , type locality; , verified records; , other records.
Moritz & Britz: Revision of Polypterus
61
Congo basin (P. palmas polli) from Upper Guinean
specimens (P. palmas palmas). Both subspecies
recorded from Upper Guinea, P. palmas palmas
and P. palmas buettikoferi, however could not be
separated by a PCA (Hanssens et al. 1995, see
their figures 6 and 7). Following Hanssens et al.
(1995), most subsequent authors treated P. palmas
palmas, P. palmas buettikoferi and P. palmas polli as
three valid subspecies (e. g. Britz, 2004; Schliewen
& Schäfer, 2006). Schäfer (2004) regarded all
three subspecies as valid species based mainly
on differences in coloration: the dorsal color
pattern of P. palmas was claimed to be more
irregular, whereas P. buettikoferi would show
regular herring-bone markings. However, the
live specimens of P. palmas which Schäfer used
for his comparison originated from an area in
Guinea, in which according to Hanssens et al.
(1995) only P. buettikoferi is expected to occur.
While the regular herring-bone color mark-
ings are indeed present in some of the juvenile
syntypes of P. buettikoferi (e. g. NMW 63207: 2),
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
10
12
14
16
18
20
22
24
26
28
30
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
40
42
44
46
48
50
52
54
56
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
20
22
24
26
28
30
32
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
6
7
8
9
10
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
DFL in % SL DisPD in % SL preDL in % SL ScpD DS
River system
32
30
28
26
24
22
20
52
50
48
46
44
42
40
38
36
34
32
30
28
26
24
22
20
18
16
14
12
10
56
54
52
50
48
46
44
42
40
10
9
8
7
6
Fig. 44. Meristic and morphometric characters with geographical trend for Polypterus palmas. River systems are
ordered from North to South referring to their estuary: 1, Casamance (n = 2); 2, Corubal (n = 3); 3, Kogon (n = 16);
4, Rio Nunez (n = 9); 5, Fatala (n = 10); 6, Konkouré (n = 3); 7, Kolente (n = 8); 8, Sewa and Moa estuary (n = 3); 9, Moa
(n = 2); 10, Mafa (n = 10); 11, St. Paul (n = 2); 12, Nippoué/Cestos (n = 23); 13, Sehnkwehn (n = 1); 14, Dugbe (n = 4);
15, Cavally (n = 15); dotted line indicates trend line. DisPD, distance from distal tip of adducted pectoral fin to
first finlet; DFL, length of spinous part of dorsal fin; DS, number of dorsal-fin spines; preDL, pre-dorsal
length; ScpD, pre-dorsal scales.
Ichthyol. Explor. Freshwaters, IEF-1094
62
66
68
70
72
74
01 2 3 4 5 6 7 8 9 10 11 12 13 14 15
15
16
17
18
19
20
21
22
23
24
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
30
32
34
36
38
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
30
32
34
36
38
40
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
50
52
54
56
58
60
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
ScLScBPRprePL in % SLpreVL in % SL
River system
1 2 345678910 11 12 13 14 15
60
58
56
54
52
50
40
38
36
34
32
30
38
36
34
32
30
24
23
22
21
20
19
18
17
16
15
74
72
70
68
66
Fig. 45. Selected meristic and morphometric characters without geographical trend for Polypterus palmas. River
systems are ordered from North to South referring to their estuary: 1, Casamance (n = 2); 2, Corubal (n = 3); 3, Kogon
(n = 16); 4, Rio Nunez (n = 9); 5, Fatala (n = 10); 6, Konkouré (n = 3); 7, Kolente (n = 8); 8, Sewa and Moa estuary
(n = 3); 9, Moa (n = 2); 10, Mafa (n = 10); 11, St. Paul (n = 2); 12, Nippoué/Cestos (n = 23); 13, Sehnkwehn (n = 1);
14, Dugbe (n = 4); 15, Cavally (n = 15); dotted line indicates trend line. prePL, pre-pectoral length; preVL, pre-
pelvic length; PR, number of pectoral fin rays; ScB, scales around body; ScL, scales in longitudinal series.
others show a more irregular pattern (e. g. NMW
63207: 1) or such a pattern is entirely absent (e. g.
NMW 63208). Polypterus polli, on the other hand,
always exhibits a prominent dark suborbital stripe
(Schäfer, 2004).
The herein reported morphometric and mer-
istic characters from Upper Guinean specimens
of Polypterus palmas s. l. allows to divide the
characters in two groups (Figs. 44-45): some vary
across the entire range and show no geographical
trend (Fig. 45); others show geographically cor-
related variation, but are largely overlapping or
very similar in their ranges (Fig. 44). We found
that number of dorsal-fin spines and length of the
dorsal fin itself both decrease from north to south.
Number of pre-dorsal scales, pre-dorsal distance
and distance between pectoral fin and dorsal fin
all increase in the same direction (Fig. 44). There
is no discontinuity in this north-south clinal vari-
ation that would point to a separation between
different taxa. Most values, however, do not show
such a trend and are quite equally distributed
between and within groups of specimens in each
river (Fig. 45). Two specimens do not fit with the
Moritz & Britz: Revision of Polypterus
63
reported geographical trends. They originate
from the far north of the distribution area, the
Casamance River, and have values that resemble
much more those from specimens from the very
south (Fig. 44, rivers system 1). We are unable to
evaluate the significance of these differences, be-
cause only two specimens from this location were
available for our study. In general, however, we
did not find any evidence in our morphological
analysis that would suggest the existence of two
separate taxa within P. palmas s. l. in the Upper
Guinean region.
The situation is different for the specimens
from the Congo region. Hanssens et al. (1995)
provided a PCA separating specimens from the
Congo basin from those of Upper Guinea (see their
Fig. 7). Based on our data we can confirm the dis-
tinction of these two taxa, when dorsal-fin spine
number is plotted against number of pre-dorsal
scales or pre-pelvic scales (Fig. 46). In the first case
there is a single shared data point between both
taxa, 7 spines and 25 pre-dorsal scales (Fig. 46a).
Dorsal-fin spine number in combination with
pre-pelvic scales, however, shows no overlap
(Fig. 46b) and allows a clear distinction.
The melanophore color pattern of specimens
from the two regions is principally very similar,
but the dark dorsal blotches in Upper Guinea
specimens are brownish forming an irregular
marbled pattern, often with broad stripes of more
than two scales width. In the Congo specimens
they are grey or black, usually arranged in bars of
up to two scales width. In addition the suborbital
stripe is narrow in Upper Guinea specimens and
often incomplete (interrupted by light areas),
but is usually wider, more conspicuous, and
uniformly dark grey in the Congo specimens. In
direct comparison there is also a slight difference
in head shape in lateral view, appearing rounded
and blunt in Upper Guinea specimens but less
rounded and more dorsoventrally flattened in
specimens from the Congo area.
Genetic data support this distinction and the
existence of two species, P. palmas and P. polli: the
Congo specimens (P. polli) were resolved as sister
group of P. delhezi and, both together, in a sister
group relationship with P. senegalus (Suzuki et
al., 2010; Near et al., 2014). The Upper Guinean
specimens (P. palmas, for which only four speci-
mens from aquarium trade were used, thus likely
representing only a small part of the distribution
area or even a single location) are placed either as
the sister taxon of the group containing P. delhezi,
P. senegalus and P. polli (Near et al., 2014) or as
the sister group of the Lower Guinean species
P. teugelsi (Suzuki et al., 2010).
In summary, analysis of our data demonstrates
the existence of two readily distinguishable spe-
cies formerly subsumed under P. palmas s. l.:
Polypterus palmas distributed in the Upper Guinea
region, and Polypterus polli in the Congo basin.
Polypterus buettikoferi cannot be distinguished
from P. palmas and has therefore to be regarded
as a junior synonym of P. palmas. The unusual
looking Polypterus sp. Golddust bichir of Schäfer
(2004) imported from the Killy River in Guinea
may be a color morph of P. palmas but needs to
be further investigated.
ScpD ScpP
DS
DS
4
5
6
7
8
9
10
18 20 22 24 26 28 30 32
4
5
6
7
8
9
10
32 34 36 38 40 42 44 46
a b
Fig. 46. Scatterplots of Polypterus palmas (%, n = 118) and P. polli ( , n = 85). a, number of dorsal-fin spines (DS)
against number of pre-dorsal scales (ScpD); b, number of dorsal-fin spines (DS) against number of pre-ventral
scales (ScpP). One symbol may represent several specimens.
Ichthyol. Explor. Freshwaters, IEF-1094
64
Polypterus polli Gosse, 1988
(Figs. 47-49, Tables 19-20)
Polypterus polli Gosse, 1988: 241, fig. 1.
Material examined. 85 specimens. type speCimens:
MRAC 47667, holotype (Fig. 47), 261 mm SL; Democratic
Republic of Congo: Ruki River at Eala, J. Ghesquière,
Nov 1936. – IRSNB 798, 1, paratype, 170 mm SL;
Democratic Republic of Congo: Mongala River at Binga
Moke, A. Collart, 26 Sep 1927. – MRAC 1371-1372, 2,
paratypes, 251-239 mm SL; Democratic Republic of
Congo: Ruki River at Eala, M. Laurent, 1905. – MRAC
2799, 1, paratype, 253 mm SL; Democratic Republic
of Congo: Kwamouth, De Maes. – MRAC 3209, 1,
paratype, 209 mm SL; Democratic Republic of Congo:
Sankuru River at Kondue, E. Luja, 1914. – MRAC
3211-3213, 3, paratypes, 197-257 mm SL; Democratic
Republic of Congo: Sankuru River at Kondue, E. Luja,
1914. – MRAC 3216-3219, 4, paratypes, 192-235 mm
SL; Democratic Republic of Congo: Sankuru River at
Kondue, E. Luja, 1914. – MRAC 14676, 1, paratype,
230 mm SL; Democratic Republic of Congo: Congo River
at Mogende, H. Schoueteden, 1925. – MRAC 14684, 1,
paratype, 92.9 mm SL; Democratic Republic of Congo:
Kasai River at Basongo, H. Schoueteden, 1925. – MRAC
14694, 1, paratype, 110.9 mm SL; Democratic Republic
of Congo: Congo River at Ikengo, H. Schoueteden, 1925.
– MRAC 30037, 1, paratype, 237 mm SL; Democratic
Republic of Congo: Congo River at Lisala, H. Schoue-
teden, Jun 1926. – MRAC 41649, 1, paratype, 209 mm
SL; Democratic Republic of Congo: Congo River at
Kinshasa, A. Tinant, 1934. – MRAC 56291, 1, paratype,
248 mm SL; Democratic Republic of Congo: Congo
River at Vista, E. Dartevelle, 1937. COngO river anD
smaller triButaries: BMNH 1887.1.13.12, 1, 175.4 mm
SL; Democratic Republic of Congo: Lower Congo,
Hans. – BMNH 1896.3.9.28-30, 3, 69.4-76.9 mm SL;
Democratic Republic of Congo: Congo River 50 miles
south of Mangala, J. Weeks. – BMNH 1897.9.30.29, 1,
260 mm SL; Democratic Republic of Congo: Stanley
Falls, 0°30' N 25°12' E; W. Bentley. – BMNH 1898.7.8.38,
1, 121.9 mm SL; Democratic Republic of Congo: Mon-
sembe, J. Weeks. – BMNH 1899.6.27.2, 1, 212 mm SL;
Democratic Republic of Congo: Upper Congo River.
– BMNH 1901.12.21.3, 1, 264 mm SL; Democratic Re-
public of Congo: Monsembe, J. Weeks. – IRSNB 14423,
1, 231 mm SL; Democratic Republic of Congo: Kalamu
River at Boma, S. Lefevere, 20 Sep 1955. – IRSNB 9958, 1,
275 mm SL; Democratic Republic of Congo: Yangambi:
falaise de Yaosuka, A. Hullot, 5 Sep 1948. – MNHN
1900-0197, 1, 113.1 mm SL; Democratic Republic of
Congo, Degeorgis. – MNHN 1958-0026, 1, 224 mm SL;
Democratic Republic of Congo: Stanley Pool, M. Poll et
al., Sep 1957. – MNHN 1958-0027, 1, 80.3 mm SL; Demo-
cratic Republic of Congo: Stanley Pool, M. Poll et al., Mar
1956. – MNHN 1958-0028, 1, 111.3 mm SL; Democratic
Republic of Congo: Stanley Pool, M. Poll et al., Mar 1956.
– MRAC 14693, 1, 108.4 mm SL; Democratic Republic
of Congo: River Mongende, D. Schouteden. – MRAC
19743, 1, 199 mm SL; Democratic Republic of Congo:
Boende, C. Gerard, Aug 1928. – MRAC 56590, 1, 208 mm
SL; Democratic Republic of Congo: Kunungu: Ndva
village, D. Schouteden, 1938. – MRAC 79463-79466, 3,
175-228 mm SL; Democratic Republic of Congo: Ebale
River at Botanakasa, B. NKele, 1952. – MRAC 79884,
Democratic Republic of Congo: Inkenge, R. P. Lootens,
20 Feb 1952. – MRAC 98803, 1, 141.1 mm SL; Democratic
Republic of Congo: Umpuelo River opposite of Boma,
I. Mesmaekers, Mai 1955. – MRAC 98482-98483, 2,
186-256 mm SL; Angola: small tributary of Umpuelo
River opposite of Boma, I. Mesmaekers, 1955. – MRAC
98814-98817, 4, 258-289 mm SL; Democratic Republic of
Congo: Umpuelo River opposite of Boma, I. Mesmaek-
ers, Jun 1955. – MRAC 99520, 1, 191 mm SL; Democratic
Republic of Congo: Umpuelo River opposite of Boma, I.
Mesmaekers, Jul 1955. – MRAC 115711, 1, 93.0 mm SL;
Democratic Republic of Congo: Stanley Pool, Brien and
Poll, 4 May 1956. – MRAC 115715, 1, 220 mm SL; Demo-
cratic Republic of Congo: Stanley Pool at maluku, Brien
and Poll, 4 Oct 1957. – MRAC 119119, 1, 267 mm SL;
Democratic Republic of Congo: Luki, Wagemans, 1957.
– MRAC 154588, 1, 212 mm SL; Democratic Republic of
Congo: Stanley Pool, P. Brichard, 1966. – MRAC 177705,
1, 241 mm SL; Democratic Republic of Congo: Stanley
Pool, P. Brichard, 1967. – MRAC 177715-177716, 2,
165-210 mm SL; Democratic Republic of Congo: Bolobo,
NKele, 1956. – MRAC 177718-177720, 3, 167-211 mm
SL; Democratic Republic of Congo: Bolobo, NKele, 20
Feb 1958. – MRAC 73-022-P-0026, 1, 213 mm SL; Demo-
cratic Republic of Congo: Stanley Pool, J. Mandeville,
23 Apr 1958. – MRAC 99515-99516, 2, Democratic
Republic of Congo: Umpuelo River opposite of Boma,
I. Mesmaekers, Sep 1955. – MRAC A6-007-P-1484, 1,
248 mm SL; Democratic Republic of Congo: Lukunga
River at Lukunga, 5°49'10" S 12°52'53" E; 9 Sep 2005.
itimBiri river: IRSNB 15949, 1, 267 mm SL; Democratic
Republic of Congo: Ibembo: Uele, H. Simeons, 10 Dec
1949. – MRAC 75695, 1, 83.5 mm SL; Democratic Re-
public of Congo: Ibembo: Uele, F. Hutsebaut, Dec 1950.
kasai river anD triButaries: MRAC 3208, 1, 241 mm
SL; Democratic Republic of Congo: Sankuru River at
Kondue, E. Luja, 1914. – MRAC 3214, 1, 230 mm SL;
Democratic Republic of Congo: Sankuru River at Kon-
due, E. Luja, 1914. – MRAC 3215, 1, 219 mm SL; Demo-
cratic Republic of Congo: Sankuru River at Kondue, E.
Luja, 1914. – MRAC 51633, 1, 103.8 mm SL; Democratic
Republic of Congo: Sankuru River at Inkongo, Wilson,
1937. likOuala river: MNHN 1962-0037, 2, 112.5-
122.7 mm SL; Democratic Republic of Congo: Likouala
aux Herbes at Bakouango, A. Stauch, 27 Feb 1961. –
MNHN 1962-0348, 5, 75.2-86.7 mm SL; Democratic
Republic of Congo: Likouala aux Herbes at Bakouango,
A. Stauch, 27 Feb 1961. OuBangui river: IRSNB 14457,
1, 200 mm SL; Democratic Republic of Congo: Oubangui
River at Isato, Cremer and Neumann, 16 Nov 1947. ruki
river anD triButaries: MRAC 131230, 1, 183.2 mm SL;
Moritz & Britz: Revision of Polypterus
65
Democratic Republic of Congo: Tshuapa River at Marais
Boteli, H. Matthes, 13 Sep 1959. – MRAC 75630-75631,
2, 236-253 mm SL; Democratic Republic of Congo:
Tshuapa River at Bokungu, L. Dupuis, 1950. – MRAC
79882, 1, 216 mm SL; Democratic Republic of Congo:
Bokuma, R. P. Lootens, 3 Nov 1951. – MRAC 98822,
1, 244 mm SL; Democratic Republic of Congo: Tsh-
uapa River at Boende, R. Philippe, Jan 1959. – MRAC
177714, 1, 221 mm SL; Democratic Republic of Congo:
Tshuapa River, P. Lootens, 1960-1964. sangah river
Table 19. Morphometric characters of Polypterus polli. Range, mean and standard deviation (SD) include values
of holotype.
holotype range mean SD n
Standard length [mm] 261.0 63-315 193.9 65.0 85
Total length [mm] 284.0 77.5-352 218.1 70.4 83
In percent of standard length
Body depth 11.6 9.6-14.6 12.3 1.1 85
Body width 12.3 10.1-14.5 12.4 0.8 85
Head length 17.0 15.8-23.1 18.6 1.7 85
Pre-dorsal length 52.1 41.3-57.6 49.9 3.1 85
Dorsal-fin length 37.0 30.3-44.2 38.4 2.9 85
Pectoral to dorsal fin 20.1 6.2-22.6 16.0 3.3 85
Pre-pectoral length 16.7 15.6-23.1 18.6 1.7 85
Pectoral-fin extension 30.6 28.7-40.5 33.3 2.5 85
Inter-pectoral width 7.3 6.1-9.8 7.8 0.9 85
Pectoral-fin base width 3.3 2.7-4.0 3.3 0.3 85
Pre-pelvic length 68.6 64.0-71.3 68.4 1.3 85
Pre-anal length 86.2 81.6-87.7 85.2 1.3 85
Anal-fin ray length 11.3 9.3-15.1 12.3 1.3 85
Anal-fin base length 5.5 4.2-7.7 5.6 0.8 85
Caudal-peduncle length 3.9 1.9-5.0 3.2 0.6 85
In percent of head length
Head width 61.3 54.6-72.4 62.0 4.2 85
Interorbital width 27.1 22.2-29.8 27.0 1.4 84
Nostril distance 13.5 10.9-19.7 15.0 1.9 85
Snout length 25.2 19.5-28.0 23.8 1.5 85
Eye diameter 14.2 11.4-20.3 14.2 1.9 85
Postorbital length 62.4 57.5-67.3 62.4 1.8 85
Gular length 54.4 49.3-72.3 57.6 4.7 85
Length of first spine 19.9 17.1-36.8 24.2 3.4 83
Length of second spine 31.3 20.8-39.3 28.8 3.4 84
Width : length of first spine 35.0 16.8-47.2 26.9 4.7 83
Length of second : first spine 157.4 91.2-208.8 120.4 17.6 83
Table 20. Meristic characters of Polypterus polli. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 65 (5), 6 (64), 7 (15), 8 (1)
Dorsal- and caudal-fin rays 15 14 (4), 15 (49), 16 (32)
Pectoral-fin rays 39 31 (1), 32 (2), 33 (7), 34 (7), 35 (18), 36 (28), 37 (10), 38 (4), 39 (7), 40 (1)
Pelvic-fin rays 11 8 (2), 9 (11), 10 (54), 11 (13), 12 (5)
Anal-fin rays 13 8 (1), 9 (3), 10 (13), 11 (17), 12 (24), 13 (23), 14 (3), 15 (1)
Scales in longitudinal series 52 51 (5), 52 (9), 53 (26), 54 (35), 55 (8), 56 (1), 57 (1)
Scales around body 37 32 (3), 33 (1), 34 (8), 35 (15), 36 (26), 37 (13), 38 (13), 39 (3), 40 (2), 41 (1)
Pre-dorsal scales 26 20 (1), 21 (1), 22 (5), 23 (7), 24 (20), 25 (13), 26 (19), 27 (14), 28 (3), 29 (2)
Pre-pelvic scales 36 34 (1), 35 (7), 36 (26), 37 (32), 38 (16), 39 (2), 40 (1)
Total number of vertebrae 52 50 (2), 51 (3), 52 (6), 53 (6), 54 (2), 56 (2)
Ichthyol. Explor. Freshwaters, IEF-1094
66
Fig. 47. Polypterus polli, MRAC 47667, 261 mm SL, holotype; Democratic Republic of Congo: Ruki River at Eala;
J. Ghesquière; Nov 1936; (lateral view right side reversed).
anD triButaries: MNHN 1977-0380, 2, 123.3-262 mm
SL; Cameroon: Ngoko [= Dja] River, Depierre. – NMW
63250, 1, 245 mm SL; Cameroon: Dja River: Molundu
district, Haberer, 7 Sep 1912. – ZSM 30062, 1, 260 mm
SL; Cameroon: Lobeke close to Djalumbe savanna and
Lake Lobeke, U. Schliewen, 18-19 Jul 1999.
Additional material. COngO river anD smaller triBu-
taries: IRSNB 7537, 1, Democratic Republic of Congo:
Boende River at Yangambi, G. Gilbert, 6 Apr 1946. –
IRSNB 7541, 2, Democratic Republic of Congo: Boende
River at Yangambi, G. Gilbert, 3 Jul 1946. – MRAC 66565,
1, Democratic Republic of Congo, P. Gerard. – MRAC
14683, 1, Democratic Republic of Congo: Kinshasa, D.
Schouteden. – MRAC 14692, 1, Democratic Republic
of Congo: River Mongende, D. Schouteden. – MRAC
15508-15511, 4, Democratic Republic of Congo: Zambi,
Lang and Chapin. – MRAC 48658-48659, 2, Democratic
Republic of Congo: region des Momgangi: rivers Ley,
Yole, Mo and Meba, Ghenne. – MRAC 100403, 1, Demo-
cratic Republic of Congo: environs de Leopoldville, van
de Weyer, 1954. – MRAC 115707-115710, 4, Democratic
Republic of Congo: Stanley Pool, Brien et al., 1 Mar
1996. – MRAC 115712, 1, Democratic Republic of Congo:
Stanley Pool, Brien et al., 23 Mar 1956. – MRAC 115713,
1, Democratic Republic of Congo: Stanley Pool: NDjiili
island, Brien et al., 26 Sep 1956. – MRAC 131417-131418,
2, Democratic Republic of Congo: Yaekama River, J. P.
Gosse, 10 Dec 1953. – MRAC 73-022-P-0027-0030, 4,
Democratic Republic of Congo: Stanley Pool, J. Mandev-
ille, 10 Jan 1955. – MRAC 73-022-P-0031-0033, 4, Demo-
cratic Republic of Congo: Stanley Pool, J. Mandeville, 9
May 1958. – MRAC 88-001-P-0001-0002, 2, Democratic
Republic of Congo: swamp close to Kingabwa, P. Brien,
17 Sep 1957. – MRAC 88-001-P-0003-0004, 2, Democratic
Republic of Congo: Stanley Pool, P. Brien et al., 2 Mar
1956. – MRAC 88-025- P-0644-0646, 3, Democratic Re-
public of Congo: Bumba River, L. de Vos, 29 Jan 1988.
– MRAC 88- 025-P-0648-0660, 13, Democratic Republic
of Congo: Bumba River, L. de Vos, 30 Jan 1988. itimBiri
river: MRAC 23535, 1, Democratic Republic of Congo:
Koteli, D. Schouteden, 17 Jan 1925. – MRAC 73943, 1,
Democratic Republic of Congo: Ibembo: Uele, F. Hut-
sebaut, 1950. – MRAC 78981, 1, Democratic Republic
of Congo: Ibembo: Uele, F. Hutsebaut, Feb 1952. Ou-
Bangui river: IRSNB 14688, 2, Democratic Republic of
Congo: Oubangui River at Kala, Cremer and Neumann,
16 Dec 1947. – MRAC 166245-166246, 2, Democratic
Republic of Congo: Oubangui River at Kala, Cremer
and Neumann, 16 Dec 1947. ruki river anD triButar-
ies: MRAC 177728-177729, 2, Democratic Republic of
Congo: Boende, P. Lootens, 1967. sangah river anD
triButaries: DMM IE/10410, 3, Cameroon: Sangah
River at Libongo, T. Moritz and J. Schwarzer, Feb 2008.
Diagnosis. Polypterus polli is distinguished from
P. ansorgii, P. bichir, P. congicus, P. endlicherii and
P. delhezi by more pre-dorsal scales (20-29 vs.
12-17). Polypterus polli is distinguished from
P. mokelembembe, P. ornatipinnis, P. retropinnis,
P. teugelsi and P. weeksii by fewer pre-pelvic scales
Moritz & Britz: Revision of Polypterus
67
(34-40 vs. 41-52). Polypterus polli is distinguished
from P. senegalus by presence of dark markings
on body and fins (vs. no dark markings, uniform
coloration) and fewer dorsal-fin spines (5-8,
median: 6 vs. 8-11, median: 9). Polypterus polli
is distinguished from P. palmas by combination
of fewer dorsal-fin spines (5-8 vs. 7-10) and
pre-pelvic scales (Fig. 42B): specimens with 7 or
8 dorsal-fin spines with 35-38 pre-pelvic scales
(vs. 39-45).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 19-20. Maximum size at
least 367 mm SL (411 mm TL). Body moderately
elongated with dorsal fin positioned far back.
Upper jaw projecting beyond lower jaw.
Coloration. In life: dorsal half of body light
grey to brownish with dark grey dorsolateral
bars originating at about equal distances on mid-
dorsal line running in caudo-ventral direction,
often forming y- or x-shaped markings (Fig. 48).
Ventral half of body uniform white or cream with
a clear demarcation between dorsal and ventral
body coloration. Dorsal half of head light grey
to brownish with irregular dark grey marbling
extending onto iris and nasal tentacles. Lower
half of head white or cream. Lips usually white
with dark grey marbling (rarely upper lips grey
with dark grey marbling). Suborbital stripe dark
grey, thin, but prominent.
Pectoral fin yellow with fine dark grey speck-
ling; dark pectoral-fin base blotch present and
well defined. Pelvic and anal fins yellow with
only few dark markings. Soft dorsal and caudal
fin dark grey with yellow spots. Dorsal-fin spines
yellowish with dark markings at base and also
often in middle part; finlet membrane light grey
to translucent with dark grey mark in distal part,
covered by orange spots.
In preservative: all yellowish markings in life
pale white or brownish; body and fin markings
a
b
Fig. 48. Polypterus polli, aquarium import from Congo. a, 270 mm SL, male, total; b, 275 mm SL, female, head.
Ichthyol. Explor. Freshwaters, IEF-1094
68
less dark and therefore less contrasted than in
life; marbling on dorsal half of head absent, leav-
ing uniform greyish to brown coloration; nasal
tentacle dark brown to black (Fig. 47).
Distribution. Congo River basin (Fig 49): Congo
River from its mouth upstream to Kisangani; in
northern central Congo basin known from sev-
eral tributaries; in southern central Congo basin
recorded only from the Kasai up to confluence
with Lulua; no records from southern tributar-
ies of Kasai.
Biology. The species seems to prefer calm and
muddy areas of the river (Gosse, 1965).
Etymology. Named after Max Fernand Leon Poll,
a French ichthyologist, who first recognized this
as a different species.
Remarks. See remarks under P. palmas.
Polypterus retropinnis Vaillant, 1886
(Figs. 50-52, Tables 21-22)
Polypterus retropinnis Vaillant in Rivière, 1886: 17.
Material examined: 46 specimens. type spe Cimen :
MNHN 1886-0295, lectotype (Fig. 50), female, 201 mm
SL; Democratic Republic of Congo: Alima River,
before 1886. COngO river anD smaller triButaries:
IRSNB 7535, 1, 131.6 mm SL; Democratic Republic
of Congo: Stanleyville [= Kinshasa]: Lobilo River,
G. Gilbert, 1 Nov 1947. – IRSNB 8113, 1, 230 mm SL;
Democratic Republic of Congo: environs de Bolongo,
R. P. Schoenbroodt, 1 Nov-8 Dec 1950. – IRSNB 15253,
2, 158.6-222 mm SL; Democratic Republic of Congo:
district Stanleyville [= Kinshasa]: marais Yona, J. P.
Gosse, 4 Jun 1960. – MNHN 1931-0136, 1, 168.3 mm
SL; Democratic Republic of Congo, Baudon. – MRAC
19952, 1, 257 mm SL; Democratic Republic of Congo:
Stanleyville [= Kinshasa], 0°30' N 25°12' E, E, P. Richard,
Jan 1930. – MRAC 46203, 1, 184.6 mm SL; Democratic
Republic of Congo: Karawa, Wallin, 1936. – MRAC
56591, 1, 194 mm SL; Democratic Republic of Congo:
Kunungu, 2°06' S 16°26' E; H. Schouteden, 10 Aug 1938.
– MRAC 68533, 1, 178 mm SL; Democratic Republic
of Congo: Boonde River at Yangambi, 1945. – MRAC
68583-68584, 2, 168.3-244 mm SL; Democratic Republic
of Congo: Boonde River at Yangambi, 1945. – MRAC
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 49. Records of Polypterus polli. , type locality; , verified records.
Moritz & Britz: Revision of Polypterus
69
73431, 1, 151.5 mm SL; Democratic Republic of Congo:
environs of Stanleyville [= Kinshasa], 0°30' N 25°12' E; J.
K. Miller, 1949. – MRAC 79459-79462, 4, 192-198 mm SL;
Democratic Republic of Congo: Ebala River at Botanan-
kasa, 2°16' S 16°15' E; B. NKele, 1952. – MRAC 119966, 1,
234 mm SL; Democratic Republic of Congo: Yangambi:
Lake Yandja, A. Hulot, 1 Apr 1949. – MRAC 90-047-P-
0087, 1, 250 mm SL; Democratic Republic of Congo:
Romée River on km 30 of road Kisangani-Opala, 0°20' N
25°05' E; L. de Vos, 11 Mar 1990. – MRAC P-131419, 1,
163.0 mm SL; Democratic Republic of Congo: Yaekama
River: marais Baulu, J. P. Gosse, 10 Dec 1953. itimBiri
river: MRAC 29825, 1, 320 mm SL; Democratic Republic
of Congo: Koteli, H. Schouteden, Dec 1924. kasai anD
triButaries: MRAC 101269-101270, 2, 147.2-240 mm
SL; Democratic Republic of Congo: Edumbe River
[tributary of Sankuru], G. Vandebroeck, 1955. – MRAC
44897, 1, 138.2 mm SL; Democratic Republic of Congo:
region de Djuma: Luie River, Rots, 1955. – ZSM 29729,
1, 241 mm SL; Democratic Republic of Congo: Band-
undu province: Lui Kotale, U. Schliewen, 23-27 Aug
2002. mamBili river: MNHN 1964-0253, 1, 293 mm
SL; Democratic Republic of Congo: Odzala: Mambili,
Decarpentries and Villiers, Oct 1963. mOngala river:
MRAC P-56389-56390, 2, 76.3-111.5 mm SL; Democratic
Republic of Congo: Mongala River at Lipanga, J. Deheyn,
13 Feb 1938. OuBangui river: MNHN 1962-0336, 1,
187 mm SL; Democratic Republic of Congo: Oubangui
River, A. Stauch. – MRAC 1146, 1, 117 mm SL; Demo-
cratic Republic of Congo: Oubangui River at Banzyville,
Royaux, 1901. OgOOue river (gaBOn): MNHN 1930-
0001, 7, 69.5-126.0 mm SL; Gabon: Ogooue River at
Franceville, Baudon. – MRAC A0-049-P-0037-0038, 1,
265 mm SL [a second specimen heavily damaged and
not measured], Gabon: Franceville Mapia, D. Adriaens,
26 Sep 2000. – MRAC A0-049-P-0041, 1, 320 mm SL;
Gabon: Franceville Mapia, D. Adriaens et al., 15 Feb
2001. ruki river anD triButaries: MRAC 79880-79881,
1, 245 mm SL; Democratic Republic of Congo: Bukuma,
R. P. Lootens, 3 Nov 1951. sangah river anD triButar-
ies: NMW 63247, 2, 255-269 mm SL; Cameroon: Dja
River, Haberer, 7 Sep 1912. – NMW 63248, 1, 255 mm
SL; Cameroon: Dja River, Haberer, 28 Aug 1912. – NMW
63249, 1, 263 mm SL; Cameroon: Dja River, Haberer, 7
Sep 1912. – NMW 63250, 1, 263 mm SL; Cameroon: Dja
River, Haberer, 7 Sep 1912. – ZSM 29991, 1, 236 mm
SL; Cameroon: Sangha basin: Djalumbe [tributary to
Lobeke], U. Schliewen, 17-19 Jul 1999.
Additional material. COngO river anD smaller triBu-
taries: IRSNB 9956, 4, Democratic Republic of Congo:
Boonde river at Yangambi, A. Hulot, 1 Feb 1947. – IRSNB
9957, 1, Democratic Republic of Congo: Baomba-
Bosambila, A. Hulot, 31 Jul 1947. – IRSNB 15251, 3,
Democratic Republic of Congo: district Stanleyville
[= Kinshasa]: marais Bekwa, J. P. Gosse, 4 Jun 1960. –
IRSNB 15253, 7, Democratic Republic of Congo: district
Stanleyville [= Kinshasa]: marais Yona, J. P. Gosse, 4
Jun 1960. – MRAC 29661, 1, Democratic Republic of
Congo: Kunungu, 2°06' S 16°26', E, H. Schouteden,
1931. – MRAC 30559-30561, 3, Democratic Republic of
Congo: Kunungu, 2°06' S 16°26' E; H. Schouteden, 1931.
– MRAC 38567-38568, 2, Democratic Republic of Congo:
Kunungu, 2°06' S 16°26' E; H. Schouteden, 1932. – MRAC
48717, 1, Democratic Republic of Congo: Kunungu,
2°06' S 16°26' E; H. Schouteden, 1937. – MRAC 56586-
56588, 3, Democratic Republic of Congo: Kunungu,
2°06' S 16°26' E; H. Schouteden, 10 Aug 1938. – MRAC
67789-67790, 2, Democratic Republic of Congo: Keseki
village close to Kwamouth, H. Schouteden, 1946. –
MRAC 67996, 1, Democratic Republic of Congo: area of
Ndva, H. Schouteden, 1946. – MRAC 68629, 1, Demo-
cratic Republic of Congo: Lubilo River at Yangambi, 9
Feb-21 Mar 1946. – MRAC 119967-119969, 3, Demo-
cratic Republic of Congo, Boonde River at Yangambi,
A. Hulot, 1 Feb 1949. – MRAC P-131420, 1, Democratic
Republic of Congo: Lilanda River at Yaekama, J. P.
Gosse, 1 Sep 1952. itimBiri river: MRAC 75090, 1,
Democratic Republic of Congo: Ibembo, J. Hutsebaut,
1950. OuBangui river: MRAC 1288, 1, Democratic Re-
public of Congo: Oubangui River at Banzyville, before
1940. ruki river anD triButaries: MRAC 14695-14696,
2, Democratic Republic of Congo: Congo River at
Ikengo, H. Schouteden. – MRAC 79883, 1, Democratic
Republic of Congo: Inkenge, P. Lootens, 20 Feb 1952. –
MRAC 153730, Democratic Republic of Congo: region
of dIkela, P. Lootens, Jul 1956. – MRAC 177724-177727,
4, Democratic Republic of Congo: Boende, P. Lootens,
1967. – MRAC 77-025-P-0002, 1, Democratic Republic of
Congo: Lona River, J. Lambert, 19 May 1958.
Diagnosis. Polypterus retropinnis is distinguished
from P. ansorgii, P. bichir, P. congicus, P. endlicherii,
P. delhezi, P. senegalus and P. weeksii by more pre-
dorsal scales (27-34 vs. 10-26). Polypterus retropin-
nis is also distinguished from P. ansorgii, P. bichir,
P. congicus, P. endlicherii, and P. delhezi by fewer
dorsal-fin spines (6-9 vs. 10-18). Furthermore, it
is distinguished from P. weeksii by fewer scales
around body (33-39 vs. 42-51). Polypterus ret-
ropinnis is distinguished from P. teugelsi by fewer
scales in lateral series (56-60 vs. 64-65) and fewer
pre-pelvic scales (41-46 vs. 47-52). Polypterus
retropinnis can be distinguished from P. polli by
more pre-pelvic scales (41-46 vs. 34-40) and
usually more scales in lateral series (56-60, me-
dian 58 vs. 51-57, median 54). It is distinguished
from P. ornatipinnis by a larger distance between
the posterior margin of the pectoral fin and the
anterior origin of the dorsal fin (19.4-30.1 vs. 7.2-
17.2 %) and by fewer scales around body (33-39
vs. 39-42). Polypterus retropinnis can be readily
distinguished from P. palmas by a distinct black
pectoral-fin base blotch (vs. poorly-defined grey
area, often covered with yellowish dots), head
Ichthyol. Explor. Freshwaters, IEF-1094
70
and dorsal half of body grey (vs. yellow or with
yellowish marbling and/or spots), and presence of
prominent dark radial stripes on iris (vs. irregular
marbling). Furthermore, it differs from P. palmas
by more pre-dorsal scales (27-34, median 31 vs.
21-31, median 26) and can be distinguished from
P. mokelembembe by an incomplete suborbital
stripe, consisting of a row of individual spots
Table 21. Morphometric characters of Polypterus retropinnis. Range, mean and standard deviation (SD) include
values of lectotype.
lectotype range mean SD N
Standard length [mm] 201.0 69.5-320 189.9 70.5 46
Total length [mm] 78.9-351 227.7 71.2 37
In percent of standard length
Body depth 9.3 8.5-12.9 10.5 1.0 46
Body width 10.9 9.9-13.1 11.3 0.8 46
Head length 17.6 16.3-20.8 18.3 1.1 46
Pre-dorsal length 57.8 49.9-61.4 56.0 2.8 46
Dorsal-fin length 34.7 28.6-40.2 34.9 2.9 46
Pectoral to dorsal fin 30.0 19.4-30.1 24.8 2.9 46
Pre-pectoral length 18.2 16.0-21.7 18.8 1.3 46
Pectoral-fin extension 29.0 27.3-34.1 31.0 1.7 46
Inter-pectoral width 7.3 6.2-10.1 7.8 0.7 46
Pectoral-fin base width 3.0 2.7-3.7 3.3 0.2 46
Pre-pelvic length 72.1 68.3-76.6 73.0 1.6 46
Pre-anal length 88.6 83.3-91.8 88.2 1.8 46
Anal-fin ray length 9.2 7.9-12.6 10.7 1.0 46
Anal-fin base length 3.8 3.3-7.1 4.9 0.9 46
Caudal-peduncle length 2.8 1.3-4.1 2.4 0.5 46
In percent of head length
Head width 63.4 56.7-69.3 63.4 2.3 46
Interorbital width 30.5 25.3-31.2 28.7 1.6 46
Nostril distance 17.3 12.7-20.5 16.6 1.7 45
Snout length 22.1 16.2-26.4 23.0 1.8 46
Eye diameter 11.6 9.8-14.4 12.3 1.3 46
Postorbital length 65.6 61.1-69.0 65.2 1.7 46
Gular length 59.6 55.3-67.1 59.9 2.3 46
Length of first spine 19.5 12.9-28.6 18.5 2.8 46
Length of second spine 20.5 11.7-26.6 20.6 2.8 45
Width : length of first spine 23.8 23.8-57.0 32.0 5.7 46
Length of second : first spine 105.5 54.8-162.2 112.3 17.0 45
Table 22. Meristic characters of Polypterus retropinnis. Numbers in brackets indicate number of specimens with
the respective value. All material includes values of lectotype.
lectotype all material
Dorsal-fin spines 86 (1), 7 (18), 8 (26), 9 (1)
Dorsal- and caudal-fin rays 15 14 (4), 15 (36), 16 (5), 17 (1)
Pectoral-fin rays 32 28 (2), 29 (4), 30 (1), 31 (5), 32 (13), 33 (7), 34 (6), 35 (4), 36 (4)
Pelvic-fin rays 97 (1), 8 (6), 9 (17), 10 (15), 11 (5), 12 (2)
Anal-fin rays 11 8 (2), 9 (4), 10 (4), 11 (9), 12 (15), 13 (7), 14 (5)
Scales in longitudinal series 58 56 (2), 57 (20), 58 (16), 59 (7), 60 (1)
Scales around body 34 33 (4), 34 (10), 35 (9), 36 (13), 37 (6), 38 (2), 39 (1)
Pre-dorsal scales 34 27 (1), 28 (2), 29 (4), 30 (7), 31 (10), 32 (11), 33 (6), 34 (5)
Pre-pelvic scales 45 41 (1), 42 (3), 43 (9), 44 (21), 45 (8), 46 (4)
Total number of vertebrae 56 (1), 58 (3)
Moritz & Britz: Revision of Polypterus
71
(vs. distinct, continuous suborbital stripe), dorsal
half of head with dark marbling (vs. plain grey,
but with dark lines following sutures between
opercular and spiracular bones), and grey iris
with prominent dark radial stripes (vs. uni-
form red iris). Polypterus retropinnis differs from
P. mokelembembe by more pectoral fin rays (28-36,
median 32 vs. 24-30, median 27) and more slender
first dorsal-fin spine (4.6-7.5 % HL vs. 7.2-9.8).
a
b
Fig. 50. Polypterus retropinnis, MNHN 1886-0295, 201 mm SL, female, lectotype; Democratic Republic of Congo:
Alima River.
Fig. 51. Polypterus retropinnis, 225 mm SL, aquarium import from Democratic Republic of Congo. a, total; b, head.
Ichthyol. Explor. Freshwaters, IEF-1094
72
Descriptive synopsis. Meristic and morphomet-
ric data in Tables 21-22. Body moderately elon-
gated with dorsal fin positioned far back. Upper
jaw only projecting slightly beyond lower jaw.
Maximum recorded size 320 mm SL (351 mm TL).
Coloration. In life: dorsal half of body light grey
to grey with dark grey to black dorsolateral bars,
bifurcated caudo-ventrally and rostro-ventrally,
often confluent with preceding and subsequent
bifurcated bars resulting in a grid-like pattern,
especially in the post-anal portion of the body
(Fig. 51a). Ventral half of body white or yel-
lowish, with clear demarcation between dorsal
and ventral half of body coloration. Dorsal half
of head grey to olive-grey with dark spots and
speckles, some bone sutures of skull roof lined in
black. Ventral half of head whitish; cheeks and
lower part of opercular area with grey or brown
speckles. Suborbital stripe present, dark, almost
black, but frequently interrupted, not continuous,
and often slightly curved, not in straight line. Iris
white or light grey with prominent, radially ar-
ranged lines of dark spots; similar pattern on skin
surrounding eye (Fig. 51b). Nasal tentacle light
grey with dark speckles; upper lip and dorsal
half of lower lip white with black spots, more
prominent on lower lip.
Pectoral fin yellowish with densely set grey
spots, resulting in a striped appearance; lateral
pectoral-fin base with a dark grey blotch. Pelvic
fin yellow with grey spots; anal fin yellow with
black spots or stripes especially in anterior por-
tion. Soft part of dorsal fin and confluent caudal fin
yellow with black markings, arranged regularly
resulting either in a striped pattern or appearing
as yellow spots on black background; pattern
coarser in proximal and finer in distal portions
of fin. Dorsal-fin spines greyish or brownish with
dark marbling, sometimes with few small yellow-
ish speckles on proximal part. Finlet membranes
translucent and grey with a prominent black
blotch in rostro-ventral half and with yellow and
black specks in distal portion.
In preservative: color pattern similar to live
coloration, but light parts yellowish and dorsal
half of body less contrasted, i. e. light grey areas
in life appearing darker, and dark grey mark-
ings lighter; spots and speckles on dorsal half of
head absent (Fig. 50). Typical radially arranged
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 52. Records of Polypterus retropinnis. , type locality; , verified records; , other records.
Moritz & Britz: Revision of Polypterus
73
iris markings and interrupted cheek-stripe still
visible, the latter often more pronounced than in
life.
Distribution. Central Congo basin (Fig. 52): main
river and tributaries (e. g. Sangha, Oubangui, Sa-
longa and Kasai) from Kinshasa to Kisangani. In
Kasai basin not further south than 4°14' S. Outside
Congo basin known from Ogoué basin.
Biology. The species seems to prefer smaller
rivers, swamps and smaller lakes (Gosse, 1965).
Etymology. The name apparently refers to the
more posterior position of the first dorsal-fin
spine.
Remarks. Thirteen years before the detailed
description of Vaillant (1899) the name Polypterus
retropinnis with reference to Vaillant was already
made available in Rivière (1886). This short
note (Rivière, 1886) did not list the number of
specimens on which the description was based,
but Vaillant (1899) subsequently listed three
specimens which were present at an exhibition
in 1886. Thus the type series consists of these
three specimens, all of which are preserved in
the Paris museum. The type series, however,
includes specimens belonging to two different
species, which led Schliewen & Schäfer (2006)
to designate MNHN 1886-0295 as lectotype of
Polypterus retropinnis (Fig. 50). The paralectotypes
BMNH 1907.12.3.4 and MNHN 1886-0297 are
specimens of P. mokelembembe.
Polypterus senegalus Cuvier, 1829
(Figs. 53-58, Tables 23-24)
Polypterus senegalus Cuvier, 1829: 330.
Polypterus senegalus meridionalis Poll, 1941a: 146.
Material examined. 262 specimens. type speCimen:
MNHN 0000-5765, holotype (Fig. 54), female, 151.0 mm
SL; Senegal, Jubelin. senegal river: BMNH 1860.6.19.5,
1, 211 mm SL; Senegal, Cuming. – BMNH 1900.6.28.10-
12, 3, 211-308 mm SL; Senegal: Kaedi, M. Delhez. –
BMNH 1978.12.5.2, 1, 233 mm SL; Senegal: Lake Guiens,
D. Dorfman, 16 Feb 1978. gamBia river Basin: BMNH
1862.5.20.1-3, 3, 161-169 mm SL; Gambia, Dalton. –
BMNH 1901.7.17.2, 1, 303 mm SL; Gambia, J. S. Budgett.
– MNHN 1980-1604, 1, 230 mm SL; Senegal: Niereko
River [tributary of Gambia] at Wassadou, C. Lévêque
and D. Paugy, Apr 1980. vOl ta Basin: BMNH
1969.3.17.1-3, 3, 134-154 mm SL; Ghana, F. Irvine, 1 Jan
1938. – BMNH 2005.7.26.6, 1, 247 mm SL; Benin: Pendjari
River at Pendjari National Park, T. Moritz, 10 May 2003.
– BMNH 2006.4.17.40, 1, 228 mm SL; Burkina Faso:
Mouhoun River, T. Moritz, Dec 2003. – BMNH
2006.4.17.5, 1, 209 mm SL; Burkina Faso: Dissin, T.
Moritz, 22 May 2003. – MNHN 1961-0009, 2, 130.3-
200 mm SL; Burkina Faso: Mare aux Hippos, J. Daget,
22 Nov 1956. – MNHN 1984-0576, 4, 174.1-185.3 mm
SL; Ghana: Kume River at Wamale, C. Lévêque, Feb
1984. – MNHN 1987-0646, 3, 182-192 mm SL; Burkina
Faso: Lake Bam, C. Lévêque, Feb 1984. – MNHN 2004-
0180, 2, 167.1-171.4 mm SL; Ghana: Volta Rivert at
Kpandu, Y. Fermon, 25 Nov 2001. – MNHN 2004-0181,
3, 156-172 mm SL; Ghana: Volta Rivert at Kpandu, Y.
Fermon, 25 Nov 2001. – MNHN 2004-0184, 1, 180.1 mm
SL; Ghana: Volta Rivert at Kpandu, Y. Fermon, 25 Nov
2001. – MNHN 2004-0185, 1, 156 mm SL; Ghana: Volta
Rivert at Kpandu, Y. Fermon, 25 Nov 2001. – NMW
63221, 1, 222 mm SL; Ghana: Volta River, Fric, 1881.
tOgO: MNHN 1987-0647, 1, 189 mm SL; Togo: Sio
River at Kati, C. Lévêque, 1985. – MNHN 2002-0527, 2,
104-225 mm SL; Togo: Oti River at Mango, D. Paugy,
3 Nov 1982. Côte DivOire : MNHN 1908-0004, 1,
103.1 mm SL; Côte dIvoire, Bouet. niger river anD
triButaries: BMNH 1884.6.9.1-6, 6, 115-254 mm SL;
Nigeria: Niger River, W. Forbes. – BMNH 1902.11.10.32-
37, 6, 42-97 mm SL; Nigeria: Abo. – BMNH 1902.11.10.38-
40, 3, 60 mm SL [only largest specimen measured],
Nigeria: Assay, W. Ansorge. – BMNH 1904.1.20.1, 1,
247 mm SL; Nigeria: Mureji, J. S. Budgett. – BMNH
1904.1.20.4, 1, 68 mm SL; Nigeria: Niger Delta: Assay,
J. S. Budgett. – BMNH 1907.5.3.2, 1, 215 mm SL; Nigeria:
Benoue River north of Ibi, H. L. N. Traill. – BMNH
1913.12.5.4-13, 16, 58-93 mm SL; Nigeria: Anambra
River. – BMNH 1928.7.3.2-3, 2, 163-191 mm SL; Nigeria:
Kiyawa River near Katagum, L. Lloyd. – BMNH
1930.3.22.1, 1, 210 mm SL; Nigeria: Kiyawa River near
Katagum, L. Lloyd. – BMNH 1953.4.28.125, 1, 218 mm
SL; Nigeri: Mouth of Oni River in Lekki Lagoon, E.
Trewavas. – BMNH 1953.4.28.127, 1, 206 mm SL; Nigeri:
Lekki Lagoon, E. Trewavas. – BMNH 1959.8.18.78, 1,
111 mm SL; Nigeria: Lagos: Adboyi, P. I. R. Maclaren,
24 Feb 1948. – BMNH 1969.3.26.61, 1, 195 mm SL; Ni-
geria: Niger River, R. Whitehead. – BMNH 1982.4.13.6,
1, 124.1 mm SL; Nigeria: Bahindi about 15 km from
confluence of Sokoro and Niger River, G. McGregor-
Reid. – BMNH 1982.4.13.7-9, 3, 105.6-157.7 mm SL;
Nigeria: Sokoro River at Bunza, G. McGregor-Reid. –
BMNH 1982.4.13.10-11, 3, 79.7-93.6 mm SL; Nigeria:
Godomo near Arungu, G. McGregor-Reid, 11 Nov 1980.
– BMNH 1982.4.13.13-14, 2, 63.6-115.0 mm SL; Nigeria:
Birni Kebbi near pumping station, G. McGregor-Reid,
4 Apr 1981. – BMNH 1982.4.13.503, 1, 157 mm SL; Ni-
geria: Sokoro River at causeway bridge, G. McGregor-
Reid. – BMNH 2006.4.18.13, 1, 75 mm SL; Benin: Niger
River east of Malanville, T. Moritz. – MNHN 1912-0150,
1, 179 mm SL; Guinea: Niger River at Kouroussa, Joyeux.
– MNHN 1921-0143, 1, 228 mm SL; Guinea: Niger
Ichthyol. Explor. Freshwaters, IEF-1094
74
River, Fertelle. – MNHN 1932-0212, 1, 109.0 mm SL;
Burkina Faso: Fada NGourma, Alluaud and Chap-
puis. – MNHN 1961-0005, 1, 229 mm SL; Mali: Niger
River at Diafarabé, J. Daget, 8 Apr 1950. – MNHN
1961-0011, 1, 193 mm SL; Mali: Niger River at Kokry, J.
Daget, 20 Oct 1949. – MNHN 1979-0474, 1, 92.0 mm SL;
Mali: Niger River at Kotaka, Blanc and Daubenton, 1954.
– MNHN 1992-0770, 3, 228-248 mm SL; Guinea: Bani a
Sanassiya, D. Paugy and R. Bigorne, 2 Mar 1989. – NMW
7988, 1, 206 mm SL; Niger: Lake Dadin-Kowa at Deba,
Weidholz, Feb 1917. – NMW 7989, 1, 163.5 mm SL;
Niger: Lake Dadin-Kowa at Deba, Weidholz, Feb 1917.
lake ChaD anD triButaries: BMNH 1928.7.4.2, 1,
283 mm SL; Chad: Chari River below junction of
Lodgone River, C. Markham. – MNHN 1904-0068, 1,
283 mm SL; Chad: Chari River at Sarh, 9°08' N 18°22' E;
Chevalier and Decorse. – MNHN 1909-0391, 1, 188 mm
SL; Chad: Lake Chad, Tilho. – MNHN 1909-0392, 1,
145.4 mm SL; Chad: Lake Chad, Tilho. - MNHN 1909-
0393, 1, 136.6 mm SL; Chad: Lake Chad, Tilho. – MNHN
1909-0394, 1, 67.2 mm SL; Chad: Lake Chad, Tilho. –
MNHN 1919-0075, 1, 289 mm SL; Central African Re-
public: Gribingui River [tributary of Chari River],
Baudon. – MNHN 1919-0076, 1, 104.4 mm SL; Central
African Republic: Gribingui River [tributary of Chari
River], Baudon. – MNHN 1919-0077, 1, 136.6 mm SL;
Central African Republic: Gribingui River [tributary of
Chari River], Baudon. – MNHN 1919-0078, 1, 100.1 mm
SL; Central African Republic: Gribingui River [tributary
of Chari River], Baudon. – MNHN 1928-0072, 1, 255 mm
SL; Chad: Mayo Kebbi: Lere, Monod. – MNHN 1928-
0073, 1, 229 mm SL; Chad: Mayo Kebbi: Lere, Monod.
– MNHN 1928-0075, 1, 275 mm SL; Chad: Mayo Kebbi:
Lere, Monod. – MNHN 1928-0076, 1, 274 mm SL; Chad:
Mayo Kebbi: Lere, Monod. – MNHN 1928-0077, 1,
118.4 mm SL; Chad: Logone River at Yagoua, Monod.
– MNHN 1928-0078, 1, 188 mm SL; Chad: Logone
River at Yagoua, Monod. – MNHN 1940-0020, 1,
144.0 mm SL; Chad: Liro River, Ganay. – MNHN 1979-
0473, 1, 197 mm SL; Chad: Chari River, Monod, 1926.
nile river Basin: BMNH 1900.9.22.40-41, 2, 233-254 mm
SL; Sudan: Bahr-el-Jebel, Capt. S. Flower. – BMNH
1905.10.16.8-10, 3, 120.2-285 mm SL; Ethiopia: Baro
River at Polkom, P. Zaphiro, 4 Apr 1905. – BMNH
1907.12.2.145-148, 4, 130-169 mm SL; Sudan: White Nile
at Goz Abu Gumah, W. L. S. Loat. – BMNH 1907.12.2.149-
150, 2, 141-173 mm SL; Sudan: White Nile at Kaka, W.
L. S. Loat, 17 Apr 1901. – BMNH 1907.12.2.151-157, 6,
139.1-218 mm SL; Sudan: White Nile at Gharb-el-Aish,
W. L. S. Loat, 5-11 Apr 1901. – BMNH 1907.12.12.161-
175, 19, 127.9-370 mm SL; Sudan: White Nile at Fashoda,
W. L. S. Loat, 18-29 Mar 1901. – BMNH 1907.12.2.176,
1, 273 mm SL; Sudan: White Nile at Tewfikych, W. L.
S. Loat. – BMNH 1907.12.2.177-178, 2, 301-312 mm SL;
Sudan: White Nile at Tonga, W. L. S. Loat, 24-25 Jan
1901. – BMNH 1907.12.2.179, 1, 52.5 mm SL; Sudan:
Lake No, W. L. S. Loat. – BMNH 1907.12.2.181-182, 2,
198-294 mm SL; Sudan: mouth of Lake No, W. L. S.
Loat, 04 Mar 1901. – BMNH 1907.12.2.183, 1, 232 mm
SL; Sudan: Bahr-el-Jebel at Kerro, W. L. S. Loat, 26 Feb
1901. – BMNH 1907.12.2.184-190, 5, 221-315 mm SL;
Sudan: White Nile at Gondokorro, W. L. S. Loat, 16 Feb
- 18 Mar 1902. – BMNH 1969.1.31.119, 1, 61.7 mm SL;
Sudan: White Nile, Hamblyn. – MNHN 0000-5764, 2,
285-300 mm SL; Sudan: White Nile, Darnaud, 1843. –
NMW 50273, 1, 225 mm SL; Sudan: Khartoum, Marno,
1881. – NMW 50274, 1, 262 mm SL; Sudan: Khartoum,
Marno, 1881. – NMW 50275, 1, 395 mm SL; Sudan:
Khartoum, Marno, 1881. – NMW 63222, 1, 399 mm SL;
Sudan: Zuflüsse des oberen Nil [affluents of the Up-
per Nile], Marno, 1881. – NMW 63223, 1, 361 mm SL;
Sudan: Zuflüsse des oberen Nil [affluents of the Up-
per Nile], Marno, 1881. – NMW 63225, 2, 216-235 mm
SL; Sudan: White Nile at Gondokoro, Kmuncke, 1912.
– NMW 63226, 4, 231-263 mm SL; Sudan: White Nile
at Gondokoro, Kmuncke, 1912. – NMW 63227, 1, 281 mm
SL; Sudan: Zuflüsse des oberen Nil [affluents of the
Upper Nile], Marno, 1881. – NMW 63228, 1, 265 mm
SL; Sudan: Zuflüsse des oberen Nil [affluents of the
Upper Nile], Marno, 1881. – NMW 63229, 5, 170-228 mm
SL; Sudan: Zuflüsse des oberen Nil [affluents of the
Upper Nile], Marno, 1881. – NMW 63230, 1, 254 mm
SL; Sudan: White Nile at Khor Attac, Werner. – NMW
63231, 4, 229-240 mm SL; Sudan: White Nile at Tonga,
Werner, Apr 1917. – NMW 63232, 5, 181-260 mm SL;
Sudan: Khartoum, Marno, 1874. – NMW 63234, 1,
239 mm SL; Sudan: Bahr-el-Arab, Marno, Nov 1880. –
NMW 63235, 4, 214-406 mm SL; Sudan: Bahr-el-Seraf,
Marno, 1881. – NMW 63236, 4, 74.9-114.2 mm SL; Sudan:
Dagana, Steindachner, 1869. – NMW 63251, 3, 254-
293 mm SL; Sudan: Dagana, Steindachner, 1869. – NMW
63266, 2, 255-262 mm SL; Sudan: White Nile at Gon-
dokoro, Kmuncke, 1912. – ZSM 345345, 2, 251-280 mm
SL; Sudan: Omdourman [fishmarket], N. Pöllath, 14 Feb
2006. – ZSM 35173, 5, 237-260 mm SL; Sudan: Omdour-
man [fishmarket], D. Neumann, 24 Jan 2007. – ZSM
35188, 1, 178.8 mm SL; Sudan: White Nile at Kosti,
13°10'26" N 32°40'10" E; D. Neumann, 13 Jan 2007. – ZSM
35216, 1, 123.7 mm SL; Sudan: White Nile at Kosti,
13°10'26" N 32°40'10" E; D. Neumann, 13 Jan 2007. – ZSM
35248, 1, 95.7 mm SL; Sudan: White Nile at Kosti,
13°10'26" N 32°40'10" E; D. Neumann, 13 Jan 2007. lake
alBert: BMNH 1904.1.19.2, 1, 338 mm SL; Uganda:
Lake Albert at Butiaba, J. S. Budgett. – IRSNB 12659, 1,
189 mm SL; Democratic Republic of Congo: Lake Albert
at Kaseni, 24 May 1953. – IRSNB 12660, 1, 225 mm SL;
Democratic Republic of Congo: Lake Albert at Kaseni,
5 Oct 1953. lake turkana: BMNH 1908.1.20.1-3, 3,
342-377 mm SL; Ethiopia: Lake Turkana at Galeba, P.
Zaphiro. – BMNH 1973.5.21.245, 1, 234 mm SL; Kenya:
Lake Turkana at Todenyang, A. Hopson. – BMNH
1981.2.17.1253, 1, 239 mm SL; Kenya: Lake Turkana at
Todenyang, A. Hopson, 29 Oct 1970. – MNHN 1933-
0073, 1, 247 mm SL; Ethiopia: Lake Turkana: Omo
Delta, 2 Feb 1933. – MNHN 1933-0074, 1, 203 mm SL;
Ethiopia: Lake Turkana: Omo Delta, 2 Feb 1933. ka-
tanga regiOn: BMNH 1975.6.20.6-7, 2, 221-221 mm
SL; Democratic Republic of Congo: Lugue River, K.
Moritz & Britz: Revision of Polypterus
75
Banister. – BMNH 1975.6.20.8, 1, 255 mm SL; Demo-
cratic Republic of Congo: Lugue River 1 km upstream
from junction with Congo River, K. Banister. – BMNH
1975.6.20.10, 1, 215 mm SL; Democratic Republic of
Congo: Lulaba River 4 km north of Mulongo, K. Ban-
ister. – BMNH 1975.6.20.11, 1, 224 mm SL; Democratic
Republic of Congo: Lake Kinsale, K. Banister. – BMNH
1975.6.20.12, 1, 217 mm SL; Democratic Republic of
Congo: Lake Zunbambe at Malemba, K. Banister. –
BMNH 1975.6.20.13-14, 2, 214-220 mm SL; Democratic
Table 23. Morphometric characters of Polypterus senegalus. Range, mean and standard deviation (SD) include
values of holotype.
holotype range mean SD N
Standard length [mm] 151.0 61.7-406 218.6 71.3 226
Total length [mm] 164.0 73-450 244.2 78.2 219
In percent of standard length
Body depth 10.8 8.2-14.0 10.8 1.1 223
Body width 10.4 8.5-14.6 10.8 0.8 223
Head length 17.6 14.4-22.6 16.6 1.6 226
Pre-dorsal length 35.1 30.0-45.3 35.0 2.6 225
Dorsal-fin length 56.6 44.7-61.8 55.6 2.8 226
Pectoral to dorsal fin 4.5 0.0-11.5 5.0 2.3 224
Pre-pectoral length 17.4 14.7-22.8 17.2 1.5 226
Pectoral-fin extension 30.0 23.6-38.6 29.8 2.3 224
Inter-pectoral width 6.3 4.1-11.1 6.1 0.9 223
Pectoral-fin base width 3.1 2.3-4.0 2.9 0.3 223
Pre-pelvic length 66.2 61.7-71.2 67.5 1.5 222
Pre-anal length 83.5 77.7-90.0 85.3 1.6 226
Anal-fin ray length 10.0 7.9-15.3 11.0 1.1 221
Anal-fin base length 4.6 3.7-7.4 5.4 0.8 226
Caudal-peduncle length 2.9 1.5-4.3 2.7 0.5 225
In percent of head length
Head width 53.2 48.6-66.8 57.8 3.7 226
Interorbital width 26.2 22.4-30.7 25.9 1.3 226
Nostril distance 14.9 8.5-19.0 14.0 1.8 225
Snout length 19.8 16.0-26.5 20.7 1.5 226
Eye diameter 16.4 9.8-20.8 14.4 1.9 226
Postorbital length 63.0 55.6-69.4 64.5 2.2 226
Gular length 56.8 50.1-74.1 63.0 6.4 225
Length of first spine 27.1 14.2-38.6 27.5 4.0 222
Length of second spine 34.0 19.1-43.1 31.4 4.1 224
Width : length of first spine 23.8 17.9-43.8 27.3 4.3 222
Length of second : first spine 125.2 66.7-210.3 114.2 14.4 221
Table 24. Meristic characters of Polypterus senegalus. Numbers in brackets indicate number of specimens with
the respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 10 8 (7), 9 (129), 10 (86), 11 (4)
Dorsal- and caudal-fin rays 17 14 (3), 15 (14), 16 (77), 17 (117), 18 (9), 19 (4), 20 (1)
Pectoral-fin rays 37 28 (1), 31 (4), 32 (9), 33 (19), 34 (54), 35 (49), 36 (44), 37 (25), 38 (15), 39 (5), 41 (1)
Pelvic-fin rays 98 (8), 9 (60), 10 (83), 11 (50), 12 (22), 13 (3)
Anal-fin rays 12 9 (6), 10 (30), 11 (49), 12 (53), 13 (47), 14 (25), 15 (9), 16 (3)
Scales in longitudinal series 57 53 (1), 54 (4), 55 (14), 56 (41), 57 (66), 58 (68), 59 (25), 60 (4)
Scales around body 36 32 (1), 33 (8), 34 (40), 35 (57), 36 (59), 37 (41), 38 (9), 39 (4), 40 (3), 44 (1)
Pre-dorsal scales 17 15 (13), 16 (41), 17 (63), 18 (52), 19 (41), 20 (8), 21 (3), 22 (1), 23 (1)
Pre-pelvic scales 36 35 (2), 36 (12), 37 (36), 38 (63), 39 (62), 40 (33), 41 (13), 42 (1), 43 (1)
Total number of vertebrae 53 (2), 54 (4), 55 (11), 56 (23), 57 (29), 58 (34), 59 (15), 60 (3), 61 (1)
Ichthyol. Explor. Freshwaters, IEF-1094
76
Republic of Congo: Lake Tomphwe at Muyumba, K.
Banister. – IRSNB 4869, 1, 233 mm SL; Democratic Re-
public of Congo: Lualaba River at Maka, H. J. Bredo, 1
Feb 1939. – IRSNB 9959, 1, 173 mm SL; Democratic
Republic of Congo: Lake Lukushi, A. Hulot, 9 Jan 1947.
– IRSNB 20336, 1, 263 mm SL; Democratic Republic of
Congo: Lake Upemba at Mabwe, G.-F. Witte, 7-9 Sep
1947. – IRSNB 20337, 2, 261-271 mm SL; Democratic
Republic of Congo: Lake Upemba at Mabwe, G.-F. Witte,
10-14 Aug 1947. – MNHN 1927-0342, 1, 300 mm SL;
Democratic Republic of Congo: Tonga, Babault. – MRAC
119970, 1, 213 mm SL; Democratic Republic of Congo:
Lukushi River, A. Hulot, Apr 1949. – MRAC 20655, 1,
303 mm SL; Democratic Republic of Congo: Nyonga,
G. F. de Witte, 1-7 May 1925. – MRAC 20656, 1, 226 mm
SL; Democratic Republic of Congo: Nyonga, G. F. de
Witte, 13-20 May 1925. – MRAC 34706, 1, 256 mm SL;
Democratic Republic of Congo: Kiambi, G. F. de Witte,
4-20 May 1931. – MRAC 35634-35635, 2, 169-197 mm
SL; Democratic Republic of Congo: Kiambi, G. F. de
Witte, 4-20 May 1931. – MRAC 69918, 1, 229 mm SL;
Democratic Republic of Congo: swamp close to Luala-
ba River at Kabalo, M. Poll, 6 Jul 1947. – MRAC 70084,
1, 250 mm SL; Democratic Republic of Congo: Lualaba
River at Kindu, M. Poll, 6 Jul 1947. – MRAC 72697, 1,
264 mm SL; Democratic Republic of Congo: Lake Ka-
bamba, 16-17 Jul 1948. – MRAC 81070, 1, 310 mm SL;
Democratic Republic of Congo: Lake Lukushi at Mano-
no, 22 Mar 1947. – MRAC 81071, 1, 219 mm SL; Demo-
cratic Republic of Congo: Lake Lukushi at Manono, Apr
1947. – MRAC 81072, 1, 187 mm SL; Democratic Repub-
lic of Congo: Lake Lukushi at Manono, 9 Jan 1947. –
MRAC 178673-178676, 2, 191-223 mm SL; Democratic
Republic of Congo: Kikondja, W. de Smet, 4-5 Apr 1956.
– MRAC 79-001-P-0021, 1, 165 mm SL; Democratic
Republic of Congo: Lake Upemba at Mabwe, G. F. de
Witte, 17-19 Nov 1948. – MRAC 79-001-P-0024, 1,
165 mm SL; Democratic Republic of Congo: Lake Up-
emba at Mabwe, G. F. de Witte, 2 Dec 1948. aquarium
traDe: BMNH 2004.6.3.6, 1, 236 mm SL; no data. – ZSM
34028, 1, 153.4 mm SL; Democratic Republic of Congo:
export via Kinshasa without exact location, F. Schäfer,
2004.
Additional material. vOlta Basin: DMM IE/10390,
4, Burkina Faso: Navrikpé Lake at Dissin 10°58'55" N
2°56'39" W, T. Moritz Jan 2004. – DMM IE/10391, 1,
Benin: Pendjari National Park: Pendjari River at Hotel
Pendjari, 11°24'33" N 1°35'22" E; T. Moritz, 20 May 2007.
– DMM IE/10392, 1, Benin: Pendjari National Park:
Pendjari River at Pont Arly, 11°28'05" N 1°34'01" E; T.
Moritz, 25 Nov 2002. – DMM IE/10394, 4, Benin: Pendjari
National Park, T. Moritz, Jan 2004. – DMM IE/10397,
4, Burkina Faso: Navrikpé Lake at Dissin, 10°58'55" N
2°56'39" W; T. Moritz, Jan 2005. – DMM IE/10398, 6,
Burkina Faso: Navrikpé Lake at Dissin, 10°58'55" N
2°56'39" W; T. Moritz, Jan 2004. – DMM IE/10402, 1,
Benin: Pendjari National Park: small floodpool close to
Pendjari River, 11°27'36" N 1°28'57" E; T. Moritz, 2 Jan
2004. – DMM IE/10403, 1, Benin: Pendjari National Park,
T. Moritz, 2004. – DMM IE/10413, 4, Benin: Pendjari
National Park: Mare Diwouni, 11°25'34" N 1°34'55" E;
T. Moritz, 2007. – MNHN 2003-0613, 6, Burkina Faso:
Kadiogo: Barrage le Baskoy, Y. Fermon, Aug 1996. –
MNHN 2003-0625, 6, Ghana: Lake Volta at Akasombo,
Y. Fermon, 1997. COastal rivers Of tOgO, Benin anD
nigeria: DMM IE/9243, 1, Benin, Ouémé River at
Adjohoun, T. Moritz, 20 Nov 2003. – IRSNB 20869,
1, Benin: Porto Novo, de Kimpe, 1 Oct 1963. – IRSNB
20870, 1, Benin: Porto Novo, de Kimpe, 4 Sep 1964. –
MNHN 1984-0508, 2, Benin: Ouémé River, Doussou.
niger river anD triButaries: DMM IE/10240, 1, Benin:
Niger River at Malanville, D. Bierbach, 21 Sep 2008. –
DMM IE/10244, 3, Benin: Niger River at Malanville,
D. Bierbach, 2 Jun 2008. – DMM IE/10246, 1, Benin:
Niger River at Malanville, D. Bierbach, 6 Sep 2008. –
DMM IE/10257, 1, Benin: Niger River at Malanville,
D. Bierbach, 30 Jul 2008. – MNHN 1961-0006, 13, Mali:
Niger River at Diafarabé, J. Daget, 8 Apr 1950. – MNHN
1961-0010, 5, Mali: Niger River at Diafarabé, J. Daget,
Nov 1952. – MNHN 1962-0445, 1, Cameroon: Vendou
Ouainero River at Polbomi, A. Stauch. lake ChaD anD
triButaries: MNHN 1959-0448, 4, Chad: Chari Delta,
Blanche et al., 26 Jul 1955. – MNHN 1959-0449, 1, Chad:
Mayo Kebbi at Tikem, J. Blanche and A. Stauch, 26
Jul 1955. – MNHN 1959-0450, 1, Chad: Chari Delta, J.
Blanche et al., 23 Jul 1955. – MNHN 1959-0451, 5, Chad:
Chari Delta, J. Blanche et al., 23 Jul 1955. – MNHN 1959-
0452, 1, Chad: Mayo Kebbi at Lere, J. Blanche et al., Jan
1958. – MNHN 1959- 0453, 1, Chad: Boulou River at El
Obeid, J. Blanche, May 1954. – MNHN 1979-0472, 1,
Chad: Mayo Kebbi at Fianga, Monod, 1926. – MRAC
A6-020-P-0056, 1, Chad: Gara plain, Wildekamp et al.,
6 Dec 2005. nile river Basin: DMM IE/6385, 1, Sudan:
White Nile River at Kosti, 13°10'21" N 32°40'23" E; T.
Moritz and V. von Vietinghoff, 23-24 Jan 2006. – DMM
IE/6386, 1, Sudan: White Nile River at Kosti, 13°10'21" N
32°40'23" E; T. Moritz and V. von Vietinghoff, 23-24
Jan 2006. – DMM IE/6387, 2, Sudan: White Nile River
at Kosti, 13°10'21" N 32°40'23" E; T. Moritz and V. von
Vietinghoff, 23-24 Jan 2006. – DMM IE/6388, 3, Sudan:
White Nile River at Kosti, 13°10'21" N 32°40'23" E; T.
Moritz and V. von Vietinghoff, 23-24 Jan 2006. – DMM
IE/9370, 5, Sudan: White Nile River at Kosti, 13°10'21" N
32°40'23" E; T. Moritz and V. von Vietinghoff, 24 Jan
2006. – DMM IE/9371, 3, Sudan: White Nile River at
Kosti, 13°10'21" N 32°40'23" E; T. Moritz and V. von
Vietinghoff, 24 Jan 2006. – DMM IE/9372, 2, Sudan:
White Nile River at Kosti, 13°10'21" N 32°40'23" E; T.
Moritz and V. von Vietinghoff, 24 Jan 2006. – DMM
IE/10404, 2, Sudan: Khartoum [central fish market], T.
Moritz et al., Apr 2008. – DMM IE/10405, 1, Sudan: Wad
al Saki [via central fish market Khartoum], T. Moritz et
al., Apr 2008. – DMM IE/10406, 4, Sudan: Getina [via
central fish market Khartoum], T. Moritz et al., Apr
2008. – DMM IE/10689, 25, Sudan: White Nile River at
Kosti, 13°10'29" N 32°40'15" E; T. Moritz et al., 15 Apr
2016. – DMM IE/10690, 21, Sudan: White Nile River at
Moritz & Britz: Revision of Polypterus
77
Kosti, 13°10'29" N 32°40'15" E; T. Moritz et al., 15 Apr
2016. – DMM IE/10691, 16, Sudan: White Nile River at
Kosti, 13°10'29" N 32°40'15" E; T. Moritz et al., 15 Apr
2016. – DMM IE/10741, 5, Sudan: White Nile River at
Aba Island, 13°11'22" N 32°40'16" E; T. Moritz et al., 16
Apr 2016. – DMM IE/14166, 1, Sudan: White Nile River
at Al Jebelein, 12°34'59" N 32°48'12" E; T. Moritz et al.,
24 Mar 2018. – DMM IE/14174, 1, Sudan: White Nile
River at Kosti, 13°10'29" N 32°40'15" E; T. Moritz et al.,
22 Mar 2018. – DMM IE/14225, 1, Sudan: White Nile
River at Aba Island, 13°11'22" N 32°40'16" E; T. Moritz
et al., 23 Mar 2018. lake alBert: IRSNB 12656, 1,
Democratic Republic of Congo: Lake Albert at Kaseni,
22 Feb 1953. – IRSNB 12657, 11, Democratic Republic
of Congo: Lake Albert at Kaseni, 7 May 1953. – IRSNB
12658, 12, Democratic Republic of Congo: Lake Albert
at Kaseni, 16 May 1953. kOttO river (nOrthern COngO
Basin): MRAC 82-021-P-0002, 1, Central African Re-
public: Kotto River at Bria, J. P. Marquet, 14 May 1982.
katanga regiOn: IRSNB 20338, 2, Democratic Republic
of Congo: Lake Upemba at Mabwe, G.-F. Witte, 27-30
Nov 1948. – MRAC 23547, 1, Democratic Republic of
Congo: Kabalo, D. Schouteden, Mar 1926. – MRAC
29705, 1, Democratic Republic of Congo: Nyonga, G.
F. de Witte, May 1925. – MRAC 30089, 1, Democratic
Republic of Congo: Nyonga, G. F. de Witte, May 1925. –
MRAC 30548-30550, 3, Democratic Republic of Congo:
Kiambi, T. Gerard, 1930. – MRAC 35461, 1, Democratic
Republic of Congo: Kiambi, G. F. de Witte, 4-20 May
1931. – MRAC 49321, 1, Democratic Republic of Congo:
Lovoi River at Kikondjia, P. Brien, Aug 1937. – MRAC
49342, 1, Democratic Republic of Congo: Bukama, P.
Brien, 12 May 1937. – MRAC 49343, 1, Democratic
Republic of Congo: Bukama, P. Brien, 12 May 1937.
Diagnosis. Polypterus senegalus is distinguished
from species of the P. bichir group by its second
dorsal scale row reaching the ventral midline
(vs. third scale row) and lateral line scales with
central pores and entire posterior margin (vs.
with incision at posterior margin). Polypterus
senegalus is distinguished from all other conge-
ners by uniform brownish to grey coloration on
dorsal half of body without any additional darker
markings in adult, but with dark horizontal bands
in larvae and small juveniles. Furthermore, it is
distinguished from P. mokelembembe, P. retropinnis
and P. teugelsi by fewer pre-dorsal scales (15-23
vs. 27-37); most specimens of P. senegalus are also
distinguished from all remaining small species
except P. delhezi by fewer pre-dorsal scales (20
or fewer vs. 21 or more, respectively). Polypterus
senegalus differs from P. ornatipinnis and P. weeksii
by fewer scales around body (32-38, rarely 39-40
[a single specimen out of 223 with 44] vs. 39-48 in
P. ornatipinnis and 42-51 in P. weeksii). Polypterus
senegalus is distinguished from P. polli by a longer
dorsal fin (44.7-61.8 % SL vs. 30.3-44.2, respec-
tively). Polypterus senegalus differs from P. palmas
by shorter distance between pectoral-fin margin
and first dorsal-fin spine (approximately on
same level: 0-11.5 % SL, mean 5.0 vs. 10.4-29.4,
mean 18.3). Polypterus senegalus is most similar to
P. delhezi, but differing by a greater interorbital
width (22-31 % HL, mean 26 vs. 18-24, mean 21).
a
b
Fig. 53. First illustrations of Polypterus senegalus. a, from Cuvier & Valenciennes, 1836; b, from Guichenot, 1839.
Both illustrations likely depict MNHN-0000-5765.
Ichthyol. Explor. Freshwaters, IEF-1094
78
Descriptive synopsis. Meristic and morpho-
metric data in Tables 23-24. Body moderately
elongated. Jaws sub-terminal. Eyes in dorso-
lateral position. Maximum recorded size 406 mm
SL (450 mm TL), but usually below 300 mm SL
(see remarks).
Coloration. In life: dorsal half of body uniform
brown (Figs. 55-56): from light brown or yellow-
ish or olive brown to intensive brown; sometimes
with very fine lighter marbling. Ventral half
of body uniform whitish, yellowish or cream.
Coloration of dorsal and ventral half merging
without clear demarcation. Lateral-line pores
often marked by black pigment; usually a row of
few small dark spots in dorsal half of body start-
ing midway between lateral line and mid-dorsal
lateral line, caudally approaching dorsal-fin base.
Dorsal half of head and nasal tentacle show same
coloration as dorsal half of body, but often with
fine, light brown or yellowish marbling; lower half
of head like lower half of body, but with distinct
demarcation between dorsal and ventral halves
(Fig. 55c); suborbital stripe present, but light
brown to greyish. Iris dark yellowish or orange
with many very small dark speckles. Lip of upper
jaw and upper half of lower jaw uniform brown,
only slightly lighter than dorsal half of body.
Pectoral fin translucent, yellowish to brown-
ish; darker blotch on lateral pectoral-fin base more
or less distinct, but never sharply demarcated.
Pelvic, anal, caudal and dorsal fins including
finlet-membranes transparent, slightly yellowish
or brownish, sometimes with very small darker
speckling; dorsal-fin spines and rays more in-
tensely colored, usually matching body coloration.
Juveniles with different color pattern (Fig.
56a-b): dorsal half of body with three dark grey
to black horizontal stripes on lighter background;
dorsal half of head and body close to dorsal mid-
line with yellowish marbling; lateral side of head
with dark stripe from snout tip through eye to
opercular series; suborbital stripe distinct and
extended onto opercular region, often continuing
on body to base of pectoral fin. Dorsal, caudal and
anal fins often with yellowish or light marbling.
Some medium-sized specimens from White Nile
at Kosti (Fig. 56c) with very faint remnants of
juvenile stripes
In preservative: very similar to life, but ventral
half faint whitish or yellowish and coloration of
dorsal half less intense (Fig. 54).
Distribution. Polypterus senegalus is widely dis-
tributed from Gambia and Senegal Rivers over Ni-
ger and Chad basin to the Nile (Sudd, White Nile
Fig. 54. Polypterus senegalus, MNHN 0000-5765, 151 mm SL, female, holotype; Senegal; Jubelin.
Moritz & Britz: Revision of Polypterus
79
and tributaries, north to Khartoum); Volta basin
and several coastal rivers from Togo to western
Nigeria (Fig. 57). It is also present in Lake Albert
and Lake Turkana [= Lake Rudolf] including Omo
River. A relict population is known from Lake
Boukou, Norther Chad (Trape, 2013). It is also
present in the Katanga region in southern Congo
basin. A single specimens has been collected in
each, the central Congo basin downstream of
Kisangani (Gosse, 1965) and in northern Congo
basin from the Kotto River, a tributary of the
Oubangui River (MRAC 82-021-P-0002).
Biology. Reproduction takes place during high
water on floodplains or smaller water bodies
(Gambia: Budgett, 1900; Niger: Daget, 1954;
Senegal: Reitzer et al., 1972).
Etymology. The name apparently refers to type
locality: Senegal.
Remarks. The description of P. senegalus is prob-
ably one of the shortest descriptions in systematic
ichthyology: Il y en a une espèce à seize dorsales,
découverte dans le Nil par M. Geoffroy (Polypterus
bichir.), . . .; et une autre du Sénégal, qui na que
douze dorsales sur le dos, P. senegalus, n. Leur
a
b
c
a
b
c
Fig. 55. Polypterus senegalus, aquarium import from Nigeria. a-c, 180 mm SL, male; b, 200 mm SL, female.
Ichthyol. Explor. Freshwaters, IEF-1094
80
chair est bonne à manger. [There is one species
with 16 dorsals discovered in the Nile by M. Geof-
froy (Polypterus bichir.) . . ., and another one from
the Senegal, which has 12 dorsals on the back,
Polypterus senegalus, n. Their flesh is good for eat-
ing.] (Cuvier, 1829: 330). A specimen in the Paris
museum, MNHN 0000-5765, has been regarded as
the holotype (Bertin, 1940; Fricke et al., 2018). This
specimen, however, possesses only 10 dorsal-fin
spines, as already noted by Duméril (1870: 395),
resulting in 9 or 10 finlets (10 if one includes the
last finlet connected to the remaining soft dorsal
fin). Apparently, no type material was mentioned
in the original description, nor was there any ref-
erence to an illustration (Cuvier, 1829). Plate 105
(Valenciennes, 1836) in the atlas accompanying
the third edition of Le Règne Animal (Cuvier,
1836) illustrates P. senegalus (Fig. 53a). The same
specimen is probably figured (Fig. 53b) in a more
detailed description by Guichenot (1839) and
likely represents MNHN 0000-5765 (Fig. 54).
This specimen seems to be the only specimen of
P. senegalus in the MNHN old enough to have
potentially served as the basis for the original
description. However, the discrepancy between
the 12 dorsals mentioned in the description and
the 10 dorsal-fin spines in the specimen is still
unaccounted for. Duméril (1870: 395) referred to
this conflict, however, could not find an explana-
tion why Cuvier (1829) mentioned 12 instead of
10 dorsal-fin spines. Most of the 226 specimens of
P. senegalus analysed during our study have either
9 or 10 spines. Only 4 specimens have 11, but we
did not find a single specimen with 12 dorsal-fin
spines. The same is true for Dagets (1954) study
of 559 specimens of P. senegalus, of which only
four specimens had 11 dorsal-fin spines and none
had 12 (Daget, 1954). Therefore, we suspect that
Cuviers description concerning the number of
dorsal-fin spines may be erroneous and MNHN
0000-5765 is indeed the holotype of the species.
The name Polypterus arnaudii was used by
Duméril (1870: 394, pl. 23), but on the illustrating
plate it is mentioned: Polypterus senegalus (sous
le nom de Pol. Arnaudii). Thus, this name is not
available because it was published as a synonym
of P. senegalus (following Art. 11.5 of the Code)
by Dumèril (1870).
a b
c d
e
Fig. 56. Polypterus senegalus. a-b, 45 mm SL, small juvenile; Sudan: White Nile at Kosti; c-d, 78 mm SL, juvenile;
Benin: Niger River at Malanville; e, 110 mm SL, young adult; Sudan: White Nile at Kosti.
Moritz & Britz: Revision of Polypterus
81
Steindachner (1881) mentioned a specimen
of 500 mm length and Banister & Bailey (1979)
reported specimens up to 700 mm SL from the
Katanga area, which were not preserved. Given
the smaller maximum recorded size of P. senegalus
it is possible that these records rather refer to
specimens of the sympatric species P. bichir.
Despite its wide distribution, closely matching
that of Polypterus bichir, there is surprisingly little
geographic variation. A comparison of meristic
characters among P. senegalus specimens did not
show any clear difference when grouped accord-
ing to major biogeographical areas, i. e. West
Africa, Chad basin, Nile basin and Katanga area
in the southern part of the Congo basin (Fig. 58).
Even specimens from presumably more isolated
populations like those of Lake Albert or Lake
Turkana do not show any noticeable differences
from other specimens of P. senegalus, yet the
available museum material from these areas is
very limited. There is no diagnostic, or otherwise
conspicuous, difference between P. senegalus
specimens from the Katanga area and those
from elsewhere in Africa. Specimens originating
from the Katanga area have been described as a
separate subspecies, P. senegalus meridionalis (Poll,
1941a). In Polls (1941a) key, where P. senegalus
meridionalis is first mentioned, there is indeed
no difference at all between P. s. senegalus and
P. s. meridionalis: all scale counts for P. s. meridio-
nalis are entirely within the range of the same
counts provided for P. s. senegalus. Poll (1941a)
cited differences between the two subspecies in
mean values of these scale counts (Poll, 1941a),
but this could not be confirmed in our study. The
only additional characters cited by Poll (1941a,
1942) to distinguish the two subspecies are small
differences in the size up to which the external
larval gills persist and their length. Poll (1941a,
1942) compared his Katanga specimens only with
specimens of one sample from the Gambia River
in these characters and did not consider whether
length or persistence of external gills may rather
be determined by external factors. Already Daget
(1948) raised concerns about the validity of P. sene-
galus meridionalis and recommended a revision
of the sub-species in P. senegalus. In summary,
we were unable to find any character in which
P. senegalus specimens from the Katanga area
differ from those of other parts of Africa and
therefore place P. s. meridionalis in synonymy with
P. senegalus.
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 57. Records of Polypterus senegalus. , type locality; , verified records; , other records.
Ichthyol. Explor. Freshwaters, IEF-1094
82
7
8
9
10
11
12
14
15
16
17
18
19
20
21
22
23
ScpD
52
53
54
55
56
57
58
59
60
ScL
31
33
35
37
39
41
43
45
ScB
ScpP
34
35
36
37
38
39
40
41
42
43
DS
PR
AR
DCR
27
29
31
33
35
37
39
41
8
9
10
11
12
13
14
15
16
13
14
15
16
17
18
19
20
WA Chad Nile L. Alb L. Tur Kata WA Chad Nile L. Alb L. Tur Kata
Fig. 58. Box-and-whisker plots for selected meristic characters of Polypterus senegalus from different areas: WA, West
Africa (n = 76); Chad, Chad basin (n = 22); Nile, Nile basin (n = 91); L. Alb, Lake Albert (n = 3); L. Tur, Lake Tur-
kana (n = 6); Kata, Katanga region of Congo basin (n = 32). AR, anal-fin rays; DCR, soft rays in confluent dorsal
and caudal fin; DS, dorsal-fin spines; PR, pectoral-fin rays; ScB, scales around body; ScL, scales in longitudinal
series; ScpD, pre-dorsal scales; ScpP, pre-pelvic scales.
Polypterus teugelsi Britz, 2004
(Figs. 59-61, Tables 25-26)
Polypterus teugelsi Britz, 2004: 180, fig. 1.
Material examined. 13 specimens. USNM 303913,
holotype (Fig. 59), male, 397 mm SL; Cameroon: Manyu:
Cross River system downstream of Mamfe at Bapuo
River junction with Cross, 5°55'30" N 9°09'00" W; G. M.
Reid, 20 Feb 1988. – USNM 375961, 5, paratypes, 88.3-
241 mm SL [smallest specimen not measured in detail];
Cameroon: Manyu: Cross River system downstream of
Mamfe at Bapuo River junction with Cross, 5°55'30" N
9°09'00" W; G. M. Reid, 20 Feb 1988. – USNM 303773,
3, paratypes, 113-127 mm SL; Cameroon: Manyu:
Cross River system downstream of Mamfe at Mam
River junction, 5°50'30" N 9°14'50" E; G. M. Reid, 16
Feb 1988. – USNM 303799, 1, paratype, 107 mm SL;
Cameroon: Manyu: Cross River system downstream
of Mamfe, stream junction with Cross River, 5°51'25" N
9°12'50" E; G. M. Reid, 17 Feb 1988. – USNM 303820,
Moritz & Britz: Revision of Polypterus
83
3, paratypes, 23-64 mm SL [not measured in detail];
Cameroon: Manyu: Cross River system downstream of
Mamfe at Akinyam River junction, 5°50'30" N 9°14'50" E;
G. M. Reid, 17 Feb 1988. – MRAC P-73040-0005-0007,
3, paratypes, 119-163 mm SL; Cameroon: Cross River,
J. Grimshaw, 18 Nov 1970. – MRAC P-73029-0342, 1,
paratype, 415 mm SL; Cameroon: Cross River at Mamfe,
D. F. E. Thys van den Audenaerde, 23 Apr 1970.
Diagnosis. Polypterus teugelsi is distinguished
from all other species of Polypterus, except P. or-
natipinnis, by combination of the following char-
acters: 7-9 dorsal-fin spines and 28-33 pre-dorsal
scales. Polypterus teugelsi is distinguished from
P. ornatipinnis by having 40-43 pectoral rays (vs.
31-39) and a distinct coloration of gular plates
(gular plates white vs. black with whitish or
yellowish marbling) and by fewer scales around
body (37-40 vs. 39-48).
Descriptive synopsis. Meristic and morphomet-
ric data in Tables 25-26. Body much elongated (in
comparison with other Polypterus species); ante-
rior body almost round in diameter, in posterior
third laterally compressed. Upper jaw slightly
projecting beyond lower jaw. Head slightly
depressed. Maximum recorded size 415 mm SL
(467 mm TL).
Coloration. In life: dorsal half of body beige to
brown-orange with irregular dark brown anasto-
mosing pattern formed by confluent dorsal and
lateral blotches (Fig. 60). Ventral half of body uni-
form whitish-yellowish to light orange. Distinct
border between dorsal and ventral half of body
coloration, less distinct in postanal area. Dorsal
half of head like body, but ventrally lighter and
set apart from uniform white gular plates; cheek
with irregular spot-like markings forming faint
suborbital stripe. Iris orange; skin around eye not
different from remaining head coloration. Lips of
Fig. 59. Polypterus teugelsi, USNM 303913, 397 mm SL, male, holotype; Cameroon: Manyu: Cross River system;
G. M. Reid; 20 Feb 1988.
Ichthyol. Explor. Freshwaters, IEF-1094
84
a
a
b
Fig. 60. Polypterus teugelsi, 430 mm SL, aquarium import. a, total; b, head.
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 61. Records of Polypterus teugelsi. , type locality; , verified records.
Moritz & Britz: Revision of Polypterus
85
upper and lower jaw light beige with fine dark
brown marbling (Fig. 60b).
Pectoral, pelvic and anal fins beige to brown-
orange with fine, dark brown spots sometimes
arranged in rows. Lateral pectoral-fin base whitish
proximally and brownish distally; medial pecto-
ral-fin base with brownish markings arranged in
rows. Soft parts of dorsal and caudal fins brown-
Table 25. Morphometric characters of Polypterus teugelsi. Range, mean and standard deviation (SD) include
values of holotype.
holotype range mean SD n
Standard length [mm] 397.0 107.3-415 187.5 105.8 13
Total length [mm] 439.0 13.3-467 236.4 148.0 12
In percent of standard length
Body depth 10.1 8.9-12.5 10.5 1.0 13
Body width 9.8-11.3 10.6 0.6 4
Head length 15.1 15.1-19.6 18.1 1.6 13
Pre-dorsal length 46.7 46.7-54.8 51.0 2.6 13
Dorsal-fin length 42.6 34.6-43.9 39.6 2.8 13
Pectoral to dorsal fin 22.4 16.8-25.2 21.6 2.5 13
Pre-pectoral length 15.5 15.2-20.8 18.4 2.0 13
Pectoral-fin extension 26.5-31.5 30.1 2.4 4
Inter-pectoral width 5.6-7.6 6.9 0.9 4
Pectoral-fin base width 2.8-3.1 3.0 0.2 4
Pre-pelvic length 66.7 66.7-72.9 70.6 1.6 13
Pre-anal length 83.3 79.7-87.7 85.3 2.1 13
Anal-fin ray length 10.5-13.1 11.6 1.2 4
Anal-fin base length 4.6-6.0 5.0 0.6 4
Caudal-peduncle length 2.6-3.5 3.0 0.5 4
In percent of head length
Head width 65.4 56.0-65.8 60.7 3.0 13
Interorbital width 29.6 22.9-30.6 26.2 2.3 13
Nostril distance 14.3-18.7 17.1 2.1 4
Snout length 24.6 22.6-28.1 25.1 1.6 13
Eye diameter 8.9 8.9-14.1 12.2 1.7 13
Postorbital length 63.4-65.6 64.4 1.0 4
Gular length 55.6-60.6 57.7 2.4 4
Length of first spine 16.9-22.7 19.7 3.0 4
Length of second spine 21.0-24.5 22.5 1.7 4
Width : length of first spine 23.3-28.8 23.3 2.6 4
Length of second : first spine 107-125.8 125.8 9.3 4
Table 26. Meristic characters of Polypterus teugelsi. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 87 (1), 8 (11), 9 (1)
Dorsal- and caudal-fin rays 15 (1), 16 (1), 17 (2)
Pectoral-fin rays 40 40 (4), 41 (1), 42 (7), 43 (1)
Pelvic-fin rays 12 11 (4), 12 (7), 13 (2)
Anal-fin rays 11 (1), 13 (2), 15 (1)
Scales in longitudinal series 65 64 (6), 65 (7)
Scales around body 38 37 (3), 38 (8), 39 (1), 40 (1)
Pre-dorsal scales 28 (1), 29 (1), 30 (5), 31 (3), 32 (2), 33 (1)
Pre-pelvic scales 49 47 (1), 48 (2), 49 (8), 50 (1), 52 (1)
Total number of vertebrae 64 63 (6), 64 (3), 65 (1)
Ichthyol. Explor. Freshwaters, IEF-1094
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orange with dark brown to black irregular mark-
ings. Dorsal-fin spines orange-brown with two
dark oblique bars; finlet membranes dark brown
to black, posterior portion often translucent, with
irregular, light brown-orange spots.
In preservative: all orange-brown or light
orange markings in life white or light beige; less
contrasted than in life (Fig. 59); darker markings
on head much fainter; dark blotch on lateral
pectoral-fin base pale brownish.
Distribution. Endemic to the Upper Cross River
system (Fig. 61).
Biology. A black-water species found in densely
forested areas (Teugels et al., 1992).
Etymology. Named in honour of Guy G. Teugels
(1954-2003), a Belgian ichthyologist.
Polypterus weeksii Boulenger, 1898
(Figs. 62-64, Tables 27-28)
Polypterus weeksii Boulenger, 1898: 419.
Polypterus schoutedeni Pellegrin, 1923b: 296.
Material examined. 45 specimens. type spe Cimen :
BMNH 1898.7.8.87, holotype (Fig. 62), 135 mm SL;
Democratic Republic of Congo: Monsembe, J. H.
Weeks. COngO river anD small triButaries: BMNH
1901.12.21.2, 1, 344 mm SL; Democratic Republic of
Congo: Monsembe, J. H. Weeks. – MNHN 1923-0102,
1, 132 mm SL; holotype of P. schoutedeni, Democratic
Republic of Congo: Mogende, H. Schouteden. – MRAC
103988, 1, 219 mm SL; Democratic Republic of Congo:
Stanley Pool, A. Werner, Jun 1955. – MRAC 115716, 1,
135 mm SL; Democratic Republic of Congo: Stanley
Pool, Brien and Poll, 10 Jan 1955. – MRAC 119965, 1,
355 mm SL; Democratic Republic of Congo: Lubilaye
River, A. Hulot, 29 Oct 1949. – MRAC 15023, 1, 120.0 mm
SL; paratype of P. schoutedeni, Democratic Republic
of Congo: Mogende, H. Schouteden, 1922. – MRAC
38084, 1, 162 mm SL; Democratic Republic of Congo:
Kunungu, H. Schouteden, 1932. – MRAC 55612, 1,
488 mm SL; Democratic Republic of Congo: Leopol-
dville [= Kinshasa], A. Tinant, 1937. – MRAC 56592, 1,
216 mm SL; Democratic Republic of Congo: Kunungu:
NDuva, H. Schouteden, 10 Aug 1938. – MRAC 66719,
1, 175 mm SL; Democratic Republic of Congo: region
of Tshela: Mayumba, Vleeschouwers. – MRAC 72-022-
P-0034, 1, 311 mm SL; Democratic Republic of Congo:
Stanley Pool: swamps of Limgundu, J. Mandeville, 9
Apr 1958. – MRAC 72-022-P-0035, 1, 281 mm SL; Demo-
cratic Republic of Congo: Stanley Pool, J. Mandeville,
16 Apr 1958. – MRAC 72-022-P-0036, 1, 260 mm SL;
Democratic Republic of Congo: Stanley Pool, J. Man-
deville, 9 May 1958. – MRAC A7-031-P-0495, 1, 531 mm
SL; Democratic Republic of Congo: Léfini basin: Liya
river [tributary of Lessio River], 3°15'30" S 15°28'48" E,
I. Zamba et al., 18 Sep 2007. – MRAC A7-033-P-0028,
1, 335 mm SL; Democratic Republic of Congo: Malebo
Pool: N' Sele, 54°15'26" S 15°34'55" E; Hanssens et al., 4
Sept 2007. – MRAC P-178982-178986, 3, 257-422 mm SL;
Democratic Republic of Congo: Stanley Pool at King-
abwa, Brien et al., 4 Aug 1958. – MRAC unregistered,
1, 373 mm SL; Democratic Republic of Congo: Lefini
River 2.4 km downstream of camp Malina, 2°59'14" S
15°11'45" E; 26 Apr 2007. kasai river anD triButaries:
MRAC 15024, 1, 115.5 mm SL; paratype of P. schoutedeni,
Democratic Republic of Congo: Kasai River at Basongo,
H. Schouteden, 1921. – MRAC 67071, 1, 311 mm SL;
Democratic Republic of Congo: Fimi River at Gantoko,
Vleeschouwers, 1945. – MRAC A4-046-P-0772, 1, 359 mm
SL; Republic of Congo: Itsotso River at left shore of
Louna River, about 5 km upstream of confluence with
Léfini River, 3°03' S 15°49' E; Mamonekene et al., 16 Sep
2004. – MRAC A5-014-P0008, 1, 353 mm SL; Republic of
Congo: Itsotso River at left shore of Louna River, about
5 km upstream of confluence with Léfini River, 3°03' S
15°49' E; Mamonekene et al., 16 Sep 2004. likOuala
aux herBes: MNHN 1962-0334, 2, 82.6-133.2 mm SL;
Democratic Republic of Congo: Likouala aux Herbes at
Boloko, A. Stauch, 28 Feb 1961. – MNHN 1962-0335, 2,
104-142mm SL; Democratic Republic of Congo: Likouala
aux Herbes at Boloko, A. Stauch, 27 Feb 1961. – MRAC
14677, 1, 175 mm SL; Democratic Republic of Congo:
Ikingo River, H. Schouteden, 1921. – MRAC 14678, 1,
168.9 mm SL; Democratic Republic of Congo: Ikingo
River, H. Schouteden, 1921. lOmami river: MRAC 89-
43-P-0270, 1, 214 mm SL; Democratic Republic of Congo:
Lobaye River on the road Yaogama-Yaeti, L. de Vos and
A. Vandelannoote, 20 Feb 1989. lulOnga river: MRAC
80367, 1, 459 mm SL; Democratic Republic of Congo:
Bosekele-Zolu close to Basankusu, J. van Vynckt, Sep
1952. OuBangui river: MRAC 179890, 1, 318 mm SL;
Democratic Republic of Congo: Dungu River at Gangala
na Bodio, M. Poll, Nov 1956. – MRAC 20652, 1, 419 mm
SL; Democratic Republic of Congo: Dungu River at
Faradje, H. Schouteden, 1925. – MRAC 75789, 1, 563 mm
SL; Democratic Republic of Congo: Oubangui River at
Bosobolo, D. Vachauder, 1950. – MRAC 84-032-P-0001,
1, 407 mm SL; Central African Republic: Ngouakoubo
River 60 km southeast of Bambari, J. P. Marquet, Feb
1983. ruki river anD triButaries: MRAC 14679, 1,
438 mm SL; Democratic Republic of Congo: Ruki River
at Eala, H. Schouteden, 1921. – MRAC 15017, 1, 275 mm
SL; Democratic Republic of Congo: Ruki River at Eala,
von Oye, 26 Jan 1923. – MRAC 15019, 1, 346 mm SL;
Democratic Republic of Congo: Ruki River at Eala,
von Oye, 17 Apr 1923. – MRAC 46169, 1, 336 mm SL;
Democratic Republic of Congo: Ruki River at Eala, J.
Ghesquiere, 22 Apr 1936. – MRAC 101370-101371, 1,
169 mm SL [a second specimen in the lot heavily dam-
aged and therefore not measured]; Democratic Republic
Moritz & Britz: Revision of Polypterus
87
of Congo: Flandria, P. Hulstaert, 23 May 1955. – MRAC
73-023-P-0002, 1, 321 mm SL; Democratic Republic of
Congo: Tshupa River at Boende, R. Philippe, Jun 1956.
– MRAC 73-023-P-0004, 1, 132.0 mm SL; Democratic
Republic of Congo: Tshupa River at Boende, R. Philippe,
Jul 1956. – MRAC 96526, 1, 232 mm SL; Democratic Re-
public of Congo: Tshupa River at Bokuma, P. Lootens,
1954. aquarium traDe: ZSM 34028:2, 1, 146 mm SL;
Democratic Republic of Congo [imported via Kinshasa
without exact location], F. Schäfer, 2004.
Table 28. Meristic characters of Polypterus weeksii. Numbers in brackets indicate number of specimens with the
respective value. All material includes values of holotype.
holotype all material
Dorsal-fin spines 99 (10), 10 (25), 11 (10)
Dorsal- and caudal-fin rays 16 15 (8), 16 (31), 17 (4), 18 (2)
Pectoral-fin rays 37 30 (1), 32 (5), 33 (1), 34 (6), 35 (13), 36 (5), 37 (10), 38 (3), 39 (1)
Pelvic-fin rays 11 10 (5), 11 (18), 12 (15), 13 (6), 14 (1)
Anal-fin rays 13 9 (2), 10 (4), 11 (7), 12 (10), 13 (16), 14 (4), 15 (2)
Scales in longitudinal series 58 55 (1), 57 (7), 58 (12), 59 (13), 60 (12)
Scales around body 46 42 (4), 43 (6), 44 (6), 45 (4), 46 (9), 47 (5), 48 (5), 49 (2), 50 (3), 51 (1)
Pre-dorsal scales 24 20 (6), 21 (11), 22 (14), 23 (6), 24 (7), 26 (1)
Pre-pelvic scales 45 41 (2), 42 (6), 43 (14), 44 (14), 45 (8), 46 (1)
Total number of vertebrae 57 (1); 58 (1)
Table 27. Morphometric characters of Polypterus weeksii. Range, mean and standard deviation (SD) include
values of holotype.
holotype range mean SD n
Standard length [mm] 135.1 82.6-563 272.9 124.9 45
Total length [mm] 154.6 102.7-612 307.1 138.9 42
In percent of standard length
Body depth 10.3 9.1-14.7 11.9 1.2 45
Body width 11.2 9.6-15.4 12.5 1.1 45
Head length 21.9 18.3-24.9 21.1 1.7 45
Pre-dorsal length 47.6 38.7-49.6 43.3 2.6 45
Dorsal-fin length 41.5 41.1-55.7 47.3 3.0 45
Pectoral to dorsal fin 9.6 2.2-13.3 7.3 2.5 45
Pre-pectoral length 23.0 18.5-24.4 21.6 1.6 45
Pectoral-fin extension 37.9 28.8-43.2 35.8 3.2 45
Inter-pectoral width 9.3 6.8-10.7 8.9 0.9 45
Pectoral-fin base width 2.9 2.9-4.3 3.6 0.4 45
Pre-pelvic length 71.7 68.2-76.0 71.2 1.6 45
Pre-anal length 87.2 82.3-91.3 87.1 1.4 45
Anal-fin ray length 12.5 8.9-15.1 12.1 1.6 45
Anal-fin base length 4.6 4.0-7.8 5.5 0.9 45
Caudal-peduncle length 3.6 2.0-4.0 2.9 0.5 45
In percent of head length
Head width 57.5 53.6-69.4 62.4 4.2 45
Interorbital width 25.1 23.0-32.3 28.1 2.4 45
Nostril distance 22.4 15.2-22.4 17.5 1.6 45
Snout length 23.6 21.9-27.6 24.6 1.3 45
Eye diameter 11.2 8.8-14.1 10.8 1.2 45
Postorbital length 65.0 61.2-70.5 65.3 2.0 45
Gular length 67.5 50.5-67.6 58.0 3.5 45
Length of first spine 22.9 10.1-25.9 20.3 3.0 45
Length of second spine 24.4 17.2-29.1 22.8 2.4 45
Width : length of first spine 24.1 20-48 25.7 4.5 45
Length of second : first spine 106.4 77.6-229.3 114.6 21.3 45
Ichthyol. Explor. Freshwaters, IEF-1094
88
Additional material. itimBiri river: MRAC 88-025-P-
0643, 1, Democratic Republic of Congo: Kagola River
[tributary of Itimbiri] at Aketi, L. de Vos; 18-19 Jan 1988.
ruki river anD triButaries: MRAC 74870, 1, Democratic
Republic of Congo: Bokungu, L. Dupuis, 1950. – MRAC
80785, 1, Democratic Republic of Congo: Bamania, P.
Hulstaert, 1952. – MRAC 99708, 1, Democratic Republic
of Congo: Momboyo River at Loselinga, P. Hulstaert,
5 May 1955. – MRAC 104722, 1, Democratic Republic
of Congo: Ruki River, P. Hulstaert, Jan 1956. – MRAC
177722-177723, 2, Democratic Republic of Congo:
Boende, P. Lootens, 1967.
Diagnosis. Polypterus weeksii is distinguished
from P. ansorgii, P. bichir, P. congicus, P. endlicherii
and P. delhezi by more pre-dorsal scales (20-26 vs.
17 or fewer). Polypterus weeksii is distinguished
from P. mokelembembe, P. palmas, P. polli, P. retro-
pinnis and P. teugelsi by more scales around body
(42-51 vs. 31-41). Most specimens of P. weeksii
differ from P. senegalus also by more scales around
body (41-51, median 46 vs. 32-44, median 36). It
can also be distinguished from P. senegalus by
color pattern (series of dark dorsolateral bars on
dorsal half of body vs. dorsolateral bars or dorsal
and lateral blotches absent, uniform brownish
to greyish in adults or with horizontal stripes
in small juveniles and larvae). In meristic and
morphometric characters P. weeksii is most similar
to P. ornatipinnis, but clearly distinguished by its
color pattern (dorsal half of body grey with dark
bars vs. black with numerous white or yellowish
spots and blotches; lower half of cheek and gular
plates uniform beige without darker markings
except suborbital stripe vs. lower half of cheek
and gular plates black with whitish or yellowish
marbling; pectoral fin without conspicuous bands
vs. with one or two black bands).
Descriptive synopsis. Meristic and morpho-
metric data in Tables 27-28. Body moderately
elongated. In larger specimens head slightly more
dorsoventrally depressed and broader. Upper jaw
slightly projecting beyond lower jaw. Maximum
recorded size 273 mm SL (307 mm TL).
Coloration. In life: dorsal half of body uniform
light grey with slightly darker grey marbling or
grey with 8-10 dark grey to black oblique dorso-
lateral bars, often bifurcated ventrally and some-
times confluent with preceding and subsequent
bars; bars usually only 1-2 scale rows wide, some
individuals, however, with bars up to 4 scale rows
wide (Fig. 63). Sometimes in anterior half of body
with small additional yellowish speckles. Ventral
half of body uniform white or cream, with very
clear demarcation between dorsal and ventral half
of body coloration. Dorsal half of head light grey
to grey with light or yellowish marbling and/
or yellowish speckles. Lower half of cheeks and
ventral half of head uniform white to cream. Iris
yellowish-orange with fine, irregular, dark spots;
sometimes arranged in radial lines resembling
Fig. 62. Polypterus weeksi, BMNH 1898.7.8.87, 135 mm SL, female, holotype; Democratic Republic of Congo: Mon-
sembe; J. H. Weeks.
Moritz & Britz: Revision of Polypterus
89
b
a
c
d
a
c
d
Fig. 63. Polypterus weeksii, aquarium imports. a-b, 125 mm SL, juvenile; c, larger specimen; d, small juvenile
(photos c-d: Schuzo Nakano/Aqualog).
Ichthyol. Explor. Freshwaters, IEF-1094
90
pattern in P. retropinnis. Nasal tentacle and upper
lip same color as dorsal half of head; dorsal half of
lower lip with dark grey spots. Suborbital stripe
present, but usually with single light yellow spots
interrupting the clear dark stripe.
Pectoral fin yellowish with fine grey speckling,
only sometimes expressed as concentric lines;
lateral pectoral-fin base with dark grey blotch.
Pelvic fin yellowish with fine grey speckling,
sometimes with dark marbling. Anal fin colora-
tion like pelvic fin, but often darker. Soft parts of
dorsal and caudal fins greyish to dark grey with
whitish or yellowish more or less distinct spots.
Dorsal-fin spines yellowish-grey with one or
two dark, almost black, bands; finlet membranes
transparent grey with prominent dark blotch
continuous with proximal dark band on spine;
sometimes with additional smaller translucent
spots in distal part of finlet membranes.
In preservative: light grey areas in life pale
grey or brown; bars dark brown (Fig. 62); fine mar-
bling on head largely absent; ventral half of body
and head pale white or yellowish; suborbital stripe
more distinct than in life; pectoral fin brownish;
blotch on lateral pectoral-fin base more distinct;
pelvic fin transparent to whitish or pale grey.
Distribution. In the Congo basin present in
Congo River from mouth upstream to Kisangani;
Oubangui basin upstream to Uele River; Lulonga,
Salonga; Kasai upstream to confluence with Lulua
(Fig. 64). Not recorded from southern tributaries
of Kasai and also absent from Sangha River.
Biology. For general life history refer to the
genus, but largely unknown.
Etymology. The species was named after Rev. J.
H. Weeks who secured the type specimen for the
British Museum.
Remarks. Images of specimens labelled as Poly-
pterus cf. weeksii from Lobeke River (erroneously
referred to as Losche river) and P. weeksii from
Boulou River were published by Schäfer (2004:
159, 161). Both images do not allow precise spe-
cies identifications. The Boulou River specimen
was not preserved. The specimen from the Lobeke
River is now deposited in the ZSM collection
(ZSM 30062) and was identified as P. polli during
our study. Poll (1933) reported P. weeksii from
the Lofoi River, but the later corrected his initial
identification to P. ornatipinnis (Poll, 1941a: 162)
30°N
20°N
10°N
10°S
30°N
20°N
10°N
10°S
10°W 10°E 20°E 30°E
10°W 10°E 20°E 30°E
Fig. 64. Records of Polypterus weeksii. , type locality; , verified records; , other records.
Moritz & Britz: Revision of Polypterus
91
and confirmed that P. weeksii is not known from
the Katanga region.
Discussion
Our revision of extant polypterids based on the
study of more than 1200 specimens, of which 997
specimens have been measured in detail, recog-
nizes 13 valid species of the genus Polypterus and
a single species of Erpetoichthys. Their interrela-
tionships are still poorly known, but the results
of molecular studies (Suzuki et al., 2010; Near et
al., 2014) seem to suggest that at least the so-called
large species, i. e. P. ansorgii, P. bichir, P. congicus
and P. endlicherii form a monophyletic group.
The hypothesized relationships of the remaining
species of Polypterus based on molecular data
remain unresolved and a morphological analysis
will certainly contribute important information
to the question of polypterid interrelationships.
With the exception of the Nile, polypterids
are restricted in their distribution to sub-Saharan
Africa and have their southernmost distributional
limit in the southern tributaries of the Congo ba-
sin. The Congo basin has the highest number of
polypterid species with a total of nine out of the
13 species, of which seven are restricted to this
basin. Interestingly some species are restricted
to small areas, thus representing local endemics,
such as P. ansorgii and P. teugelsi, but other species
are widely distributed across large parts of Africa
(e. g. P. bichir and P. senegalus). Despite its huge
range, Polypterus senegalus shows little geographi-
cal variation. Polypterus bichir and P. palmas, on
the other hand, exhibit remarkable variation along
their distributional ranges. In P. palmas variations
among populations follow a north-south gradient
along coastal basins of West Africa; in P. bichir
the variation rather follows an east-west gradi-
ent from the Nile, across the Tchad basin to West
Africa. For this latter species it seems remarkable
that specimens from the distributionally isolated
Katanga area resemble specimens from West
African river systems more closely in meristic
characters than specimens from the Nile basin.
For many species, not only information about
their distribution is incomplete but ecological or
general biological data also remain scarce or are
even absent. At the same time, it seems that at
least the populations of the larger species from
the P. bichir group are highly impacted by human
activities, like fishing (Moritz, 2017) or habitat
alteration. Based on the patchy information we
have, it is hard to estimate, whether certain species
are declining and would be in need of protective
measures, as reliable information on population
trends for the different species is missing or is
based on subjective impressions obtained from
fishermen. Other indications often refer only to
single points in their distribution area, e. g. the
report of P. bichir being now regionally extinct
from Egypt (Garcia et al., 2010).
Our study is only a small step towards a better
understanding of bichirs, but we hope it will be a
starting point for more detailed studies on their
life history, distribution, ecology, morphology
and evolution.
Key to the species of Polypteridae
1 Pelvic fins present; 50-70 scales in lateral
line; 10-37 predorsal scales (Polypterus).
....................................................................... 2
Pelvic fins absent; 102-114 scales in lateral
line; 44-59 predorsal scales [coastal West
Africa] (Erpetoichthys).
................................................... E. calabaricus
2 – Third transverse scale row continuing to
ventral midline of body (Fig. 8a); base of
first dorsal finlet in front of vertical through
posterior margin of pectoral fin; 11-18
dorsal-fin spines; tip of first dorsal-fin spine,
when depressed clearly exceeding base of
subsequent spine.
....................................................................... 3
Second transverse scale row continuing to
ventral midline of body (Fig. 8b); base of
first dorsal finlet (usually) behind vertical
through posterior margin of pectoral fin;
5-12 dorsal-fin spines; first dorsal-fin spine,
when depressed, (usually) not reaching base
of subsequent spine.
....................................................................... 6
3 Body with series of black, clearly marked,
continuous, dorsolateral bars.
....................................................................... 4
Body with or without dark markings, but
never with dorsolateral bars.
....................................................................... 5
Ichthyol. Explor. Freshwaters, IEF-1094
92
4 – 4-6 (usually 4-5) saddle like dorsolateral
bars, at least 4-5 scale rows wide [West
Africa, Chad Basin, Nile].
...................................... Polypterus endlicherii
– 6-9 (usually 6-8) dorsolateral bars, 3 or
fewer scale rows wide [Congo basin and
Lake Tanganyika].
......................................... Polypterus congicus
5 – 54-56 scales in lateral line; 43-45 scales
around body [Corubal River].
.......................................... Polypterus ansorgii
– 55-70 (usually more than 58) scales in lat-
eral line; 43-54 scales around body [Nile
Basin, Chad Basin, major river basins of
West Africa (including Corubal River),
Katanga].
............................................. Polypterus bichir
6 Uniformly colored with no dark markings
on flanks (small juveniles with dark brown
horizontal bands) [Nile Basin, Chad Basin,
West Africa, Katanga].
....................................... Polypterus senegalus
With dark dorsolateral bars or anastomosing
dark markings.
....................................................................... 7
7 – 13-17 pre-dorsal scales [Congo Basin].
............................................ Polypterus delhezi
20 or more pre-dorsal scales.
....................................................................... 8
8 – Conspicuous, black and light (white or yel-
low) color pattern across body and unpaired
fins including ventral half of flank and
gular plate; pectoral fin with distinct con-
centric stripes [Congo Basin].
................................... Polypterus ornatipinnis
Color pattern different; pectoral fins without
markings or only faint stripes, gular plate
uniform light, without dark markings.
....................................................................... 9
9 – 63-65 scales in lateral line [Cross River].
.......................................... Polypterus teugelsi
61 or fewer scales in lateral line.
..................................................................... 10
10 – 42-51 scales around body [Congo Basin].
........................................... Polypterus weeksii
41 or fewer scales around body.
..................................................................... 11
11 – 5-8 (usually 6) dorsal-fin spines; specimens
with 7 or 8 dorsal-fin spines with 35-38
pre-pelvic scales.
................................................ Polypterus polli
– 7-10 dorsal-fin spines; specimens with 7 or
8 dorsal-fin spines with more than 38 pre-
pelvic scales.
..................................................................... 12
12 – Distal part of pectoral-fin base with prom-
inent dark blotch.
..................................................................... 13
– Distal part of pectoral-fin base greyish with
irregular dark pattern, but never with
prominent dark blotch [Upper Guinea re-
gion].
........................................... Polypterus palmas
13 Lower half of cheek uniformly white, but
with pronounced dark suborbital stripe; iris
uniformly red in life; 24-30 pectoral-fin rays
[Congo Basin].
............................... Polypterus mokelembembe
– Lower half of cheek with irregular fine
greyish mottling; suborbital stripe expressed
as series of spots; iris grey in life, with radi-
ally arranged dark spots or short lines;
28-36 pectoral-fin rays [Congo Basin, Low-
er Guinea region].
..................................... Polypterus retropinnis
Acknowledgements
This study profited from financial support of a Marie
Curie Inter European Fellowship (project number
219268) of TM and two scholarships from SYNTHESYS
(AT-TAF-3803, GB-TAF-2925). We are grateful to the
curators and collection managers of various museum
collections which provided various help and support
and who remained patient even when loans had to be
extended again; special thanks go to James Maclaine
(BMNH), Ulrich Schliewen and Dirk Neumann (ZSM),
Emmanuel Vreven and Miguel Parrent (MRAC), Ernst
Mickschi (NMW), Ronald de Ruiter (RMNH) and Patrice
Pruvost (MNHN).
Furthermore, we thank Frank Schäfer (aqualog),
Karsten Mody, Martin Reichard, Tobias Musschoot,
Nadja Pöllath and Dirk Neumann for providing some
of the images used in this manuscript. Special thanks
go to the colleagues in various countries who have
supported field trips by helping with organisation of
the trips, with permits, fieldwork, and the preparation
and processing of specimens. For the Sudan trips TM
thanks Ali El-Tahir Sharaf El-Din, Dirk Neumann, Nadja
Moritz & Britz: Revision of Polypterus
93
Pöllath and Vivica von Vietinghoff; for the Benin and
Burkina Faso expeditions: Philippe Laléyé, Diafarou A.
Tiomoko, Adjima Thiombiano, Brice Sinsin, K. Eduard
Linsenmair, Gisbert Dreyer, Konrad Vielhauer, Vivica
von Vieting hoff, Kevin W. Conway, Oussmane Swa-
dogo and Kongo; for fieldwork in Cameroon: Ulrich
Schliewen, Julia Schwarzer, Linus Makazi, Lennard
Usongo and S. Mbakwa; and for the collecting trips to
the Ivory Coast: K. Eduard Linsenmair, Minnattallah
Boutros and Karsten Mody. Further thanks go to the
many students of TM helping in keeping polypterids in
aquaria over many years and to Harald Benke (DMM)
granting TM the freedom and time for research. We
thank our families for their patience during the addi-
tional long working hours needed to finish this paper.
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Received 28 June 2018
Revised 1 October 2018
Accepted 5 May 2019
Moritz & Britz: Revision of Polypterus
... All these fossils equate to a total of six genera that comprise 17 species, of which two are Polypterus species [2,4,[6][7][8]. Since pinnules are an exclusive characteristic of Polypteriformes [1,21,22], fossil pinnules are confidently assigned to polypteriform fish. Conversely, diagnostic values for distinguishing species appear fragile, and only two critical works regarding the validity of pinnules as taxonomic units in extant fish have been published so far. ...
... First, three features differed in Meunier and Gayet's paper (their features 8, 13, and 19 in [5] Appendix 2) among the now known invalid subspecies of P. bichir [22]. This indicates that these three features have no specific value. ...
... The fossil record of extinct taxa mainly consists of Late Cretaceous materials from Nigerian (In Becetem), Sudanese (Wadi Milk Formation), and Moroccan (Kem Kem beds) localities, with certain species being identified from multiple localities (e.g., [7,8]). This may indicate that some species had a large geographical distribution and coexisted with each other and with species with a narrower distribution, as observed with extant taxa [22]. However, since it is possible that several nominal taxa represent the same species, geographical co-occurrence could also be used to discuss such a hypothesis. ...
Article
Full-text available
Pinnules are the peculiar, impaired spines that carry each of the numerous finlets that constitute the dorsal fins of polypterid fishes (Cladistia and Polypteriformes). Previous studies, including a recent detailed paper on the comparative analysis of the morphology of pinnules in most of the extant species (genera Polypterus and Erpetoichthys), suggest that they display unique characteristics that allow for species identification. Since most of the polypterid fossil records are composed of scales that lack specific characteristics and isolated pinnules, this work aims to test their taxonomic relevance before comparing the fossil pinnule morphologies across the fossil records in order to evaluate polypterid paleodiversity. Therefore, we describe the intra-individual and intra-specific morphological variations of the pinnules in the extant species Polypterus bichir. Furthermore, we compared it with the various morphologies described in the Polypteridae family. We report intra-individual variability related to the anteroposterior axis of the dorsal fin. We also report morphological differences in the pinnules among specimens that overlap those anticipated among different species, concluding that the pinnule morphology cannot support taxonomic purposes in polypterid fishes in their current state.
... Much less is known about the physiology, behavior, and functional development of sensory systems in these ancient fishes, whose earliest fossils date back to the mid-Jurassic. The bichir species living nowadays (14 species) inhabit the water bodies of tropical Africa and the Nile River basin (Moritz and Britz, 2019b). ...
... For most species, sucrose has an indifferent taste, but stimulates food intake only in fish, which consume much plant food, or the latter makes up a significant proportion of the diet (Kasumyan and Morsy, 1997;Kasumyan and Nikolaeva, 2002;Vinogradskaya et al., 2017). Bichirs, including gray bichir and saddle bichir, feed on animal food, such as fish, aquatic larvae and nymphs of insects (Insecta), crustaceans (Crustacea), various worms (Hickley and Bailey, 1987;Dankwa, 2004;Raji et al., 2004;Ayoade et al., 2018;Moritz and Britz, 2019b). In the rainy season, terrestrial insects (termites (Termitidae) and ants (Formicidae)) may dominate in the diet of bichirs, these food objects enter the water due to flooding of territories adjacent to water bodies (Dankwa, 2004;Ayoade et al., 2018). ...
... In the rainy season, terrestrial insects (termites (Termitidae) and ants (Formicidae)) may dominate in the diet of bichirs, these food objects enter the water due to flooding of territories adjacent to water bodies (Dankwa, 2004;Ayoade et al., 2018). Fragments and seeds of aquatic plants are also often found in the digestive tract of fish, but it is believed that they are accidentally captured by fish along with the main food (Moritz and Britz, 2019b). The taste aversiveness to sucrose exhibited by bichirs is in good agreement with the general idea that sucrose taste preference is inherent in herbivorous ani-mals (Harborne, 1993). ...
... Currently, P. bichir is described as a single species with the subspecies Polypterus bichir bichir and Polypterus bichir lapradei no longer considered valid (Moritz and Britz 2019). This sinking of subspecies is a consequence of the high degree of morphological similarity (Britz and David Johnson 2003;Britz and Johnson 2010). ...
Article
Transposable elements (TEs) can make up more than 50% of any given vertebrate's genome, with substantial variability in TE composition among lineages. TE variation is often linked to changes in gene regulation, genome size, and speciation. However, the role that genome duplication events have played in generating abrupt shifts in the composition of the mobilome over macroevolutionary timescales remains unclear. We investigated the degree to which the teleost genome duplication (TGD) shaped the diversification trajectory of the teleost mobilome. We integrate a new high coverage genome of Polypterus bichir with data from over 100 publicly available actinopterygian genomes to assess the macroevolutionary implications of genome duplication events on TE evolution in teleosts. Our results provide no evidence for a substantial shift in mobilome composition following the TGD event. Instead, the diversity of the teleost mobilome appears to have been shaped by a history of lineage specific shifts in composition that are not correlated with commonly evoked drivers of diversification such as body size, water column usage, or latitude. Collectively, these results provide additional evidence for an emerging perspective that TGD did not catalyze bursts of diversification and innovation in the actinopterygian mobilome.
... Intact fish. The Senegal bichir is distributed throughout almost all of equatorial Africa from Gambia and Senegal in the west to the Nile River basin in the east (Moritz and Britz, 2019). In rivers and standing water bodies in the habitat areas of the species (Upper Volta), the water temperature varies throughout the year from 22 to 30°C, and the temperature also changes during the day (Pekkola, 1919;Arnoult, 1964). ...
Article
Full-text available
For the first time, the locomotor activity of the intact and visually deprived Senegal bichir Polypterus senegalus was assessed at different water temperatures (20, 25, 30 and 34°C). Using the open field method, it was shown that in intact fish, with increasing temperature, locomotor activity increases (most rapidly in the range of 20–25°C) and reaches a maximum at a temperature of 30°C, which can be close to the temperature optimum (or correspond to it) for the Senegal bichir. In visually deprived fish, locomotor activity is maximum at 20°C and decreases monotonically with increasing temperature; all indicators of locomotor activity (frequency of crossing test lines; time spent for the test line crossing; distance covered by the fish, swimming speed) vary in visually deprived fish weaker than in intact ones. The discovered differences in the behavior of intact and visually deprived fish indicate the presence of a functional relationship between vision and locomotor activity in evolutionarily ancient Cladistia.
... Intact fish. The Senegal bichir is distributed throughout almost all of equatorial Africa from Gambia and Senegal in the west to the Nile River basin in the east (Moritz and Britz, 2019). In rivers and standing water bodies in the habitat areas of the species (Upper Volta), the water temperature varies throughout the year from 22 to 30°C, and the temperature also changes during the day (Pekkola, 1919;Arnoult, 1964). ...
... These values overlap with our measurements. When the mean values of the four specimens studied here were compared, which ranged from 8.71 to 9.28 μm, they specifically overlapped with four taxa from the literature, including two invalid subspecies of P. bichir(Moritz & Britz, 2019) ...
Article
Full-text available
Many actinopterygian fish groups, including fossil and extant polypteriforms and lepisosteiforms, fossil halecomorphs, and some basal teleosts, have stout bony scales covered by layers of ganoin—an enamel layer ornamented with minute tubercles. Ganoid scales preserve well as disarticulated remains and notably constitute most of the fossil record for polypteriform in both South America and Africa. Based on two variables (tubercle size and distance between tubercles), some authors reported that the ganoin tubercle ornamentation in these scales is constant within a species and differs between species and allows distinguishing species or at least groups of species. However, despite its promising potential for assessing polypteriform paleodiversity, this tool has remained unused, probably because the variables are not well defined, and intraspecific variation does not seem to have been considered. To address this gap, we aimed to test the intraspecific and intra‐individual variation in the ornamentation of ganoid scales in the type species Polypterus bichir. We propose three different parameters to describe the tubercle ornamentation: the distance between contiguous tubercles centers, their density, and their relative spatial organization. With these parameters, we investigate the variation in ganoin ornamentation among four specimens and across different regions of the body. Our results show that the distribution of the tubercles is highly variable within a same species, regardless of the body region, and sometimes even between different sectors of a same scale. Moreover, the variation observed in P. bichir overlaps with the distribution described in the literature for several extant and fossil species. Thus far, the ornamentation of ganoid scales is not a reliable diagnostical feature for polypterids.
... If needed, the field and laboratory identifications made at BEZHU (DR Congo) were subsequently verified at the RMCA where voucher specimens were deposited (collection numbers: RMCA 2012-031; RMCA 2015-005; RMCA 2016-003; RMCA 2016-025; RMCA 2018-018; and RMCA 2021-020). The publications used for the identification of the studied specimens are the checklists of the fishes of the region or the revisions of the genera and families present in the UNP [11,16,[18][19][20][21][22][23][24][25][26][27]. In addition, existing historical collections from the UNP, mainly housed at the RMCA, the Royal Belgian Institute of Natural Sciences (RBINS, Brussels, Belgium), and the Natural History Museum (NHM, London, United Kingdom), have been verified and, when necessary, re-identified. ...
Article
Full-text available
An annotated checklist of the ichthyofauna of the Upemba National Park, draining part of the Upper Lualaba basin and situated in the southern part of the Democratic Republic of the Congo, is presented, based on a literature review, a re-examination of museum collections, and a study of recent collections (2012–2020). In total, 247 native and 1 introduced species, Heterotis niloticus, are reported. The native species belong to 78 genera, 26 families, and 15 orders. Of these, 45 species (18%) are endemic to the park, 35 species (14%) await formal description, and 5 taxa (2%) need further study to clarify their status. With 51 species, the Cyprinidae is by far the most species-rich family, followed by the Mormyridae (26), Mochokidae (26), Alestidae (18), Distichodontidae (18), Amphiliidae (17), and Cichlidae (16). The remaining families are represented by less than 15 species. Comments about the species distribution and the fish fauna shared with adjacent ecoregions are provided. Although the park provides some protection for the fish species living within its borders by limiting human access to the core zone, the annex and buffer zones are both subject to strong anthropogenic pressure. These observations underscore the need for the implementation and further elaboration of fish-related preservation guidelines and plans to enable better protection/conservation of the park’s ichthyofauna.
Preprint
Full-text available
Transposable elements (TEs) can make up more than 50% of any given vertebrate's genome, with substantial variability in TE composition among lineages. TE variation is often linked to changes in gene regulation, genome size, and speciation. However, the role that genome duplication events have played in generating abrupt shifts in the composition of the mobilome over macroevolutionary timescales remains unclear. We investigated the degree to which the teleost genome duplication (TGD) shaped the diversification trajectory of the ray-finned fish mobilome. We integrate a new high coverage genome of Polypterus bichir with data from over 100 publicly available actinopterygian genomes to assess the macroevolutionary implications of genome duplication events on TE evolution. Our results provide no evidence for a substantial shift in mobilome composition following the TGD event. Instead, the diversity of the actinopterygian mobilome appears to have been shaped by a history of lineage specific shifts in composition that are not correlated with commonly evoked drivers of diversification such as body size, water column usage, or latitude. Collectively, these results provide a new perspective on the early diversification of the actinopterygian mobilome and suggest that historic ploidy events may not necessarily catalyze bursts of TE diversification and innovation.
Article
Full-text available
We readdress the controversy about the valid generic name to be applied to the African Reedfish, a species from a monotypic genus that, along with the eleven valid species of Polypterus Lacepède, 1803, comprises the known extant diversity of the order Polypteriformes. The initial conflict was established due to the inadequate replacement of the name Erpetoichthys, wrongly assumed preoccupied, by Calamoichthys, combined with the desynchronization between the sequence in which Smith’s accounts with descriptions and nomenclatural acts about the Reedfish were written and submitted for publication, and the sequence in which they were actually published/distributed. The controversy seemed to be settled in the 1980s by the finding of an earlier report published in an Edinburgh’s newspaper in 1865, in which the name Erpetoichthys was used prior to all scientific accounts by Smith. However, we demonstrate that this report cannot be considered to contain a valid nomenclatural act according to the regulations of the International Code of Zoological Nomenclature. Therefore, we undertook a detailed study to reconstruct the sequence of publication of Smith’s accounts on the Reedfish, whose correct dates of publication/distribution had not been properly established yet, to settle down once and for all the dilemma about the precedence of these names. Our conclusion is that Calamoichthys Smith, 1866a is the valid generic name to be applied to the Reedfish, and Erpetoichthys Smith, 1866a, its junior synonym, represents a name published in synonymy but later made available by Smith himself. We use the nomenclatural example of the Reedfish, as well as other cases from the literature, to draw attention to the fact that, in agreement with Article 8.1.1 of the Code, zoological names are available only when there is an unequivocal intention by their authors to scientifically describe them, even if other requirements of the Code are met. When this Article is not met in a given situation, the name is considered unavailable and an available one should be set in place for the taxon, or a new name should be proposed.
Book
Full-text available
The Catalog of Fishes covers more than 61,700 species and subspecies, over 11,000 genera and subgenera, and includes in excess of 34,000 bibliographic references. Entries for species, for example, consist of species/subspecies name, genus, author, date, publication, pages, figures, type locality, location of type specimen(s), current status (with references), family/subfamily, and important publication, taxonomic, or nomenclatural notes. Nearly all original descriptions have been examined, and much effort has gone into determining the location of type specimens. Online version: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp
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Full-text available
There are 3108 valid and named native fish species in the inland waters of Southeast Asia between the Irrawaddy and Red River drainages, the small coastal drainages between the Red River and Hainan, the whole Indochinese Peninsula, Andaman and Nicobar Islands, Indonesia (excluding Papua Province, Waigeo, Aru [but Kai is included]), and the Philippines. They belong to 137 families. Their taxonomy and nomenclature are reviewed. The original descriptions of all 7047 recorded species-group names and 1980 genus-group names have been checked in the original works for correct spelling, types, type locality and bibliographic references. The bibliography includes about 4700 titles. Synonymies are given, based on published information as well as unpublished observations. The names of 49 introduced species and 347 extralimital taxa cited in the discussions have also been checked. The original descriptions of all species not present in the covered area but cited as type species of genera have been checked for availability, authorship, date and correct spelling. The availability of some family-group names has been checked when there was suspicion of possible nomenclatural problems. Bibliographic notes include new informations on the dates of publication of works by, among others, Bleeker, Bloch, Heckel and Steindachner and discussion of authorship of names in various works.
Book
French zoologist and naturalist Georges Cuvier (1769–1832), one of the most eminent scientific figures of the early nineteenth century, is best known for laying the foundations of comparative anatomy and palaeontology. He spent his lifetime studying the anatomy of animals, and broke new ground by comparing living and fossil specimens - many he uncovered himself. However, Cuvier always opposed evolutionary theories and was during his day the foremost proponent of catastrophism, a doctrine contending that geological changes were caused by sudden cataclysms. He received universal acclaim when he published his monumental Le règne animal, which made significant advances over the Linnaean taxonomic system of classification and arranged animals into four large groups. The sixteen-volume English translation and expansion, The Animal Kingdom (1827–35), is also reissued in the Cambridge Library Collection. First published in 1817, Volume 4 of the original version covers zoophytes and concludes with several beautiful plates.
Article
48 species from 24 families have been recorded in Iguidi River, a small forest stream East of Porto Novo, Benin. The most diverse families in this stream are the Mormyridae and Cichlidae with each six species and the cyprinodontiforms (Poecilidae and Notobranchiidae) with eight species. Two species are new records for Benin, i.e. Arnoldichthys spilopterus and Schube brevianalis; Gnathonemus petersii just recently recorded for Benin, was also found in this study. Pictures showing live colouration of most species are provided. Iguidi River is special by uniting fish species known from the Niger delta region and the Ouémé basin.
Article
Seventeen species and sub-species of fishes belonging to four families (Cyprinidae, Clariidae, Aplocheilidae, Cichlidae) were known to occur in perennial bodies of water in the Sahara desert. The study of fishes collected in Lake Boukou near Ounianga Serir (Borkou, northern Chad) shows, for the first time, the occurrence in the Sahara desert of relict populations of Polypterus senegalus (Polypteridae) and Poropanchax normani (Poeciliidae). The Cichlidae Tilapia zilli was also collected in this lake. With these new records, the relict fish fauna currently known in lakes and gueltas of the Borkou plateaus comprises six species. In the Ennedi Mountains, where the specific status of Barbus populations was unclear, B. macrops was collected in Bachikere guelta. The toad Amietophrynus regularis was collected in Ounianga Kebir.
Article
The olfactory organ of the Polypteridae (Brachiopterygii) was studied histologically. It differs fundamentally from the olfactory organ of the Teleostei (Actinopterygii) and resembles that of Latimeria (Crossopterygii). The size of the olfactory organ indicates that the Polypteridae are macrosmatic.