ArticlePDF Available

Revision of the extant Polypteridae (Actinopterygii: Cladistia)

July 2019
Ichthyological Exploration of Freshwaters

DOI:10.23788/IEF-1094

Authors:

Timo Moritz

Deutsches Meeresmuseum

Ralf Britz

Senckenberg Natural History Collections Dresden

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.

Overview of principal color patterns in Polypterus.

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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.

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Records of Polypterus ansorgii. , type locality; , verified records.

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Polypterus bichir, MNHN 0000-5761, 585 mm SL, male, lectotype; Egypt: Nile; E. Geoffroy Saint-Hilaire; 1799.

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+38

Polypterus bichir, NMW 63244, 464 mm SL, male, syntype of P. lapradei; Taoué close to Richar-Toll [= ToueyMarigot]; F. Steindachner; Nov 1868.

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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)

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

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

SnPL

preDL

DFL

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

SnPL

preDL

DFL

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).

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.

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

d’Histoire naturelle, Paris; MRAC, Musée Royal

de l’Afrique 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

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

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 & Heppell’s (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 Smith’s (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 & Bockmann’s (2017)

main conclusion that Smith’s (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

Smith’s (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 & Bockmann’s (2017) conclusion and

retain Erpetoichthys calabaricus as the valid name

for the rope eel. Furthermore, following Rizzatto

& Bockmann’s (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

Fig. 3. Erpetoichthys calabaricus, NMW 63195:1, 320 mm SL, male, non-type, Nigeria: Old Calabar [type locality];

1877.

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

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

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 Africa’s 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 museum’s on-line

collection database (https://science.mnhn.fr/

30°N

20°N

10°N

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 5. Records of Erpetoichthys calabaricus. , type locality; , verified records; , other records; , question-

able records.

Ichthyol. Explor. Freshwaters, IEF-1094

–

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

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

Ichthyol. Explor. Freshwaters, IEF-1094

–

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

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.

Smith’s (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

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.

–

–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

ı ı ı ı ı ı ı ı ı ı ı ı ı ı

preDLdistPD

–

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 Geoffroy’s (1802a) descrip-

tion. Holly (1933) had come to this conclusion and

Schäfer (2004) shared this same view. Likewise,

Shaw (1804) regarded Geoffroy’s (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

Ichthyol. Explor. Freshwaters, IEF-1094

Fig. 7. (continued).

–

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

ı ı ı ı ı ı ı ı ı ı ı ı ı ı

HLSnL

–

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

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

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

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

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

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). Daget’s (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

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.

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 13. Records of Polypterus ansorgii. , type locality; , verified records.

Ichthyol. Explor. Freshwaters, IEF-1094

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

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

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

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). Geoffroy’s 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 d’Histoire 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

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

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 Geoffroy’s 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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 18. Records of Polypterus bichir. , type locality; , verified records; , other records; , questionable

records.

Moritz & Britz: Revision of Polypterus

-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

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

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

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

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

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. d’Elzius, 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

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

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,

Fig 22. Polypterus congicus. a-b, aquarium import (photos: Schuzo Nakano/Aqualog).

Moritz & Britz: Revision of Polypterus

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. Boulenger’s (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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 23. Records of Polypterus congicus. , type locality; , verified records.

Ichthyol. Explor. Freshwaters, IEF-1094

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; N’Kele, 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 N’Kosso 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

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, N’Kele, 1956. – MRAC 73-

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

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 26. Records of Polypterus delhezi. , type locality; , verified records.

Ichthyol. Explor. Freshwaters, IEF-1094

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 D’ivOire anD ghana:

BMNH 1923.3.2.2-3, 2, 136-211 mm SL; Côte d’Ivoire:

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 d’Ivoire: Bandama

River at Tiassale, Pobeuin. – MNHN 1894-0393, 1,

192 mm SL; Côte d’Ivoire: Bandama River at Tiassale,

Pobeuin. – MNHN 1960-0177, 193 mm SL; Burkina

Faso: Yanon River [tributary of Comoé], Arnoult and

d’Aubenton. – MNHN 1991-0439, 1, 344 mm SL; Côte

d’Ivoire: Bandama River at Lamto, Lamotte, Jul 1970.

– MRAC 74-014-P-0003, 1, 449 mm SL; Côte d’Ivoire:

Lake Kossou, S. Kyeli, 17 Dec 1973. – MRAC 79-036-

P-0001, 1, 261 mm SL; Côte d’Ivoire: Lagune Ebrié at

Layo, D. F. E. Thys van den Audenaerde, 20 Oct 1979.

– MRAC 80-019-P-0001, 1, 388 mm SL; Côte d’Ivoire:

Sassandra River at Soubré, G. Teugels and L Risch,

24-25 Apr 1980. – MRAC 85-029-P-0001, 1, 281 mm

SL; Côte d’Ivoire: Boubo River at Babokon, G. Teugels,

8-9 May 1985. – MRAC 85-029-P-0002, 1, 285 mm SL;

Côte d’Ivoire: Boubo River at Babokon, G. Teugels,

9-10 May 1985. – MRAC 85-055-P-001, 1, 188 mm SL;

Burkina Faso/Côte d’Ivoire [border]: Léreba River at

Léreba, G. Teugels, 24 Nov 1985. – MRAC 86 013-P-0001,

201 mm SL; Côte d’Ivoire: Bandama River at Tiassale,

G. Teugels, 3-4 Apr 1986. – MRAC 87-018-P-0001,

1, 188 mm SL; Burkina Faso/Côte d’Ivoire [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

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

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

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

D’ivOire anD ghana: MRAC 140109, 1, Côte d’Ivoire:

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

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

Fig. 30. Polypterus endlicherii. a, large specimen ~400 mm SL; Côte d’Ivoire: Comoé National Park: Comoé River;

b-c, ~100 mm SL, juvenile; Benin: Pendjari National Park: Pendjari River.

Moritz & Britz: Revision of Polypterus

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

d’Ivoire, 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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 31. Records of Polypterus endlicherii. , type locality; , verified records; , other records.

Ichthyol. Explor. Freshwaters, IEF-1094

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

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.

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

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 34. Records of Polypterus mokelembembe. , type locality; , verified records; , other records.

Ichthyol. Explor. Freshwaters, IEF-1094

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:

N’Gome River at N’Gombe, 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

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

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

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

Moritz & Britz: Revision of Polypterus

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 37. Records of Polypterus ornatipinnis. , type locality; , verified records; , other records.

Ichthyol. Explor. Freshwaters, IEF-1094

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 d’Ivoire: 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 d’Ivoire: 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 d’Ivoire: 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 d’Ivoire: 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 d’Ivoire: 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 d’Ivoire: Boan River at Danane,

J. Daget, Jul 1963. – MNHN 1979-0190, 1, 246 mm SL;

Côte d’Ivoire: Boan River at Danane, 7°21' N 8°10' W;

C. Lévêque. – MNHN 2000-0859, 7, 207-243 mm SL;

Côte d’Ivoire: 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 d’Ivoire: 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 d’Ivoire:

Moritz & Britz: Revision of Polypterus

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 d’Ivoire: 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 d’Ivoire:

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

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 d’Ivoire: 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 d’Ivoire: 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 d’Ivoire: Nipoué River

at Toyebli; D. F. E. Thys van den Audenaerde; 20 Mar 1969.

Moritz & Britz: Revision of Polypterus

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

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

Distribution. Upper Guinea region (Fig. 43):

coastal river basins from Casamance River in

Senegal to Cavally basin in Côte d’Ivoire.

Etymology. Apparently named after the type

locality at Cape Palmas in Liberia close to the

border of Côte d’Ivoire.

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 Steindachner’s 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

c d

Fig. 42. Polypterus palmas. a-b, 190 mm SL, aquarium import from Guinea; c-d, ~299 mm SL; Côte d’Ivoire: 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

as lectotype of P. buettikoferi stating that Steindach-

ner’s 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

Steindachner’s (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

Gosse’s (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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 43. Records of Polypterus palmas. , type locality; , verified records; , other records.

Moritz & Britz: Revision of Polypterus

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

DFL in % SL DisPD in % SL preDL in % SL ScpD DS

River system

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

01 2 3 4 5 6 7 8 9 10 11 12 13 14 15

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

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

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

18 20 22 24 26 28 30 32

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

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. N’Kele, 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,

N’Kele, 1956. – MRAC 177718-177720, 3, 167-211 mm

SL; Democratic Republic of Congo: Bolobo, N’Kele, 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

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

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: N’Djiili

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

(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

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 49. Records of Polypterus polli. , type locality; , verified records.

Moritz & Britz: Revision of Polypterus

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. N’Kele, 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 d’Ikela, 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

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

(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).

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 52. Records of Polypterus retropinnis. , type locality; , verified records; , other records.

Moritz & Britz: Revision of Polypterus

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 D’ivOire : MNHN 1908-0004, 1,

103.1 mm SL; Côte d’Ivoire, 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

River, Fertelle. – MNHN 1932-0212, 1, 109.0 mm SL;

Burkina Faso: Fada N’Gourma, 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

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

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

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).

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

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

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 n’a que

douze dorsales sur le dos, P. senegalus, n. Leur

Fig. 55. Polypterus senegalus, aquarium import from Nigeria. a-c, 180 mm SL, male; b, 200 mm SL, female.

Ichthyol. Explor. Freshwaters, IEF-1094

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 Daget’s (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

Cuvier’s 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

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

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 Poll’s (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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 57. Records of Polypterus senegalus. , type locality; , verified records; , other records.

Ichthyol. Explor. Freshwaters, IEF-1094

ScpD

ScL

ScB

ScpP

DCR

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

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

Fig. 60. Polypterus teugelsi, 430 mm SL, aquarium import. a, total; b, head.

30°N

20°N

10°N

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 61. Records of Polypterus teugelsi. , type locality; , verified records.

Moritz & Britz: Revision of Polypterus

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

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:

N’Duva, 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

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

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

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

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

0°

10°S

30°N

20°N

10°N

0°

10°S

10°W 10°E 20°E 30°E0°

Fig. 64. Records of Polypterus weeksii. , type locality; , verified records; , other records.

Moritz & Britz: Revision of Polypterus

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

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

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.

Literature cited

Arnoult, J. 1961. Premières observations en captivité sur

un poisson de la famille de Polypteridae: Polypterus

senegalus senegalus (Cuvier) : comportement sexuel

et ponte provoquée. Bulletin du Muséum National

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Article

Dec 2024

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.

Locomotor Activity of the Intact and Visually Deprived Senegal Bichir Polypterus senegalus (Cladistia) at Different Water Temperatures

Article

Full-text available

Oct 2024

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.

Locomotor Activity of the Intact and Visually Deprived Senegal Bichir Polypterus senegalus (Cladistia) at Different Water Temperatures

Article

Full-text available

Jun 2024

Taxonomical use of scale ornamentation: Challenges by intraspecific and intra‐individual variations in four adult specimens of Polypterus bichir

Article

Full-text available

Apr 2024
J FISH BIOL

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.

The Upemba National Park (Upper Congo Basin, DR Congo): An Updated Checklist Confirming Its Status as an African Fish Biodiversity Hotspot

Article

Full-text available

Aug 2023
Diversity

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.

Phylogenetic Classification of Living and Fossil Ray-Finned Fishes (Actinopterygii)

Article

Apr 2024

Investigating the impact of whole genome duplication on transposable element evolution in ray-finned fishes

Preprint

Full-text available

Dec 2023

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.

The Bichir Handbook, Joshua Pickett, Published by Independent Publishing Network, London, UK, 138 pp., price £24.95, ISBN: 978‐1‐78972‐770‐8

Article

Full-text available

May 2021
J FISH BIOL

Timo Moritz

Unraveling a 150 years old controversy: Calamoichthys Smith, 1866 is the valid name for the African Reedfish (Cladistii: Polypteriformes), with comments about the availability of involuntarily proposed zoological names

Article

Full-text available

Mar 2017

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.

Catalog of Fishes: Genera, Species, References

Book

Full-text available

Apr 2016

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

The fishes of the inland waters of Southeast Asia: A catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries

Article

Full-text available

Nov 2013
RAFFLES B ZOOL

Maurice Kottelat

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.

Le règne animal distribué d'après son organisation: Pour servir de base à l'histoire naturelle des animaux et d'introduction à l'anatomie comparée

Book

Nov 2015

Georges Cuvier

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.

Freshwater fishes from the Gambia river (British West Africa). Results from the Swedish expedition 1931

Article

G.S.O. Svensson

General zoology or systematic natural history

Article

G. Shaw

On the reproduction and early development of Erpetoichthys calabaricus, Polypterus senegalus, and P. ornatipinnis (Actinopterygii: Polypteridae)

Article

Dec 1998
ICHTHYOL EXPLOR FRES

Fishes of Iguidi River—A small forest stream in South-East Benin

Article

Mar 2010
ICHTHYOL EXPLOR FRES

Timo Moritz

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.

A study of the relict fish fauna of northern Chad, with the first records of a polypterid and a poeciliid in the Sahara desert

Article

Nov 2013
CR BIOL

Sébastien Trape

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.

Das Geruchsorgan der Polypteridae (Pisces, Brachiopterygii)

Article

Mar 1968

Wolfgang Pfeiffer

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.

Revision of the extant Polypteridae (Actinopterygii: Cladistia)

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