Aquat. Living Resour., 1990, 3, 265-282
Phylogenetic relationships and classification of western
palaearctic species of the genus Barbus (Osteichthyes,
Cyprinidae)
Ignacio Doadrio
Museo Nacional de Clencias Naturales, Jojé C;uliérre~
Abascal 2,28006 Madrid, $pain.
Keceivcd Fetnuary 23, 1990;arcrpted July 5 , ~ g y o .
Doadrio 1. Aquat. Living Resour., 1990, 3, 265-282.
Abstract
The relationships of 25 different populations of the genus Barbus S. str. have been studied using
different parsimony grouping methods and similarity indexes. The results indicate that Barbus S. str.
is composed of two different groups. They are provisionally placed into two different subgenera
(Barbus and Lucioharbus) in order to make the taxonomic work easier. The European species
( B . barbus, B. plebejus, B. ciscaucasicus, B. cyclolepis, B. petenyi, B. haasi and B. meridionalis) are
included in the subgenus Barbus. Within this group there are two evolutionary lines, one formed by
central European species and the other by Mediterranean species which live in mountain fluvial
courses. The subgenus Luciobarbus is constituted by Iberian, northern African and Asian species. The
relationships between them are not satisfactorily resolved. Two northern African species (B. nasus and
B. magniatlantis) are not included in any subgenus, as their phylogenetic relationships are not clear
enough. The fossil record of this genus has been reviewed and new remains from the middle Miocene
of Spain are reported as being the earliest known fossils of the genus Barbus. The first known fossils
of Luciobarbus date from the Upper Miocene in Spain, Africa and Asia. The dispersion of Barbus in
two main periods is postulated, one of them during the main Alpine orogenic phase at the end of the
Oligocene, the other during the Upper Miocene transgression. The first one maybe favoured Barbus
and the second Luciobarbus. Barriers that might have fragmented the different populations of the
genus Barbus have been evaluated and a comprehensive biogeographic model, which minimizes the
dispersion events, is proposed.
Keywords :Evolution, fossil record, taxonomy, biosystematics, biogeography, Barbus.
Les relations phylogénétiques et la classification d'espèces paléarctiques occidentales du genre Barbus
(Osteichthyes, Cypnnidae).
Résumé
Les relations de 25 populations différentes du genre Barbus S . str, à l'aide de différentes méthodes
de groupement et d'indices de similitude, ont été étudiées. Les résultats indiquent que Barbus S . str se
compose de deux groupes différents. Ceux-ci sont classés provisoirement en deux sous-genres différents
(Barbus et Luciobarbus) afin de faciliter le travail de taxonomie. Les espèces européennes (B. barbus,
B. plebejus, B. ciscaucusicus, B. c~~clolepis,
B. petenyi, B. haasi et B. meridionalis) sont incluses dans le
sous-genre Barbus. Dans ce groupe ont été trouvées deux lignées évolutives, l'une formée par des
espèces d'Europe centrale, l'autre par des espèces méditerranéennes qui habitent des cours d'eau de
montagne. Le sous-genre Luciobarbus se compose d'espèces ibériques, nord-africaines et asiatiques.
Les liens qui existent entre eux ne sont pas constatés de façon satisfaisante. Deux espèces nordafricaines (B. nasus et B. magniatlantis ne sont incluses dans aucun sous-genre, puisque leur connexion
phylogénétique n'est pas certaine. Le témoin fossile de ce genre a été réexaminé et de nouveaux
gisements du miocène-moyen en Espagne sont présentés comme étant les plus vieux fossiles connus
Aquat. Living Resour. 90103 265 18 $ 3.8010 IFREMER -Gauthier-Villars
1. Doadrio
du genre Burhu.r. Les premiers fossilcs dc Luciohurhus datcnt du miocène-supérieur en Espagne, en
Afriquc et en Asie. On suppose que la dispersion des Burhus s'est effectuée en dcux périodes majeures:
l'une pcndant la phase alpinc d'orogenèse importante A la fin de l'oligocène, l'autre pendant la
transgression du miocène-supéricur. La première aurait favorisé l'évolution des Barhu.~et la secondc,
dcs Luc,iohurhus. Des obstacles ont pu diviser Ics différentes populations d u genre Burhu.~qui ont été
évalués et on propose un modèle biogéographiquc complet lequel réduit au minimum les cléments de
dispersion.
Mots-clés : kvolution, poisson fossilc, taxonomie, biosystématiquc, biogèographic, Bc1rhu.r.
INTRODUCTION
MA'I'ERIAL AND ME'I'HODS
The genus Burhu.~Cuvier & Cloquet, 1816 shows a
great apparent diversity with approximately 800 spccies (Howes, 1987). Howcver, this diversity scems to
be due to a poor understanding of the gcneric limits
within the barbelled cyprinids with the subsequcnt
addition of species with a doubtful generic allocation.
For this reason, the taxa grouped in the gcnuï Uurhus
show very different morphological fcatures and origins.
A comprehensivc study of such a hetcrogeneous
group is cumbersomc and, as Myers (1961) indicated,
the investigation of its relationships should derive
from a separate detailed analysis of its main evolutionary lines. Following this scheme we propose a
phylogenetic study of certain Palaearctic species that
constitute so-callcd Burhus sensu stricto (Pellegrin,
1921; Myers, 1961; Banarescu, 1973, 1977), which
have been formally defined on the basis of synapomorphic characters by Howes (1987).
Up to now, the relationships in this group have
been based mainly on external morphological characters (corporal proportions, scale and ray number,
etc.), and rather few interna1 ones (numbcr of pharyngeal teeth and gill rakers). The methodologies formerly used for morphological data are variable.
Either somc data are selectcd and evolution is inferred
from them (Karaman, 1971), or taxa are grouped on
the basis of total similarity (Almaça, 1985). In both
cases the results arc contradictory and give rise to a
lack of consistcncy in classification. For example,
Karaman (1971) establishes two main groups: thc first
includes B. cupito and B. meridionulis and the second
the remaining specics, while Almaça (1984) proposes
scven different groups (namely bocagei, cycIoIepis,
xanthoperus, microcephalus, figuigensis, magniatlantis and miliaris). Following the latter classification,
B. capito would belong to the bocagei group and
B. meridionulis would be considered a mcmber of the
cyclolepis group.
We intend to perform a phylogenetic study based
on the analysis of synapomorphies, without u priori
relation of characters, using data from osteology, and
making use of several parsimony criteria to derive a
more stable classification of the group.
Twenty-five populations of Barhu.~S. str. and a
probe species of the subgenus Luheohurl7u.~(Barbus
frilschi) have been studied. Thc material is listed in
Appendix 1. Due to observcd differences, populations
of B. seluleri from thc Guadiana, Guadalquivir and
Scgura rivers have been considered separately. The
same has bccn done in the case of the Atlantic and
Mediterranean populations of B. cu//en.ris. A possible
hybrid population of B. mcridionu1i.s and B. huu.si has
also been analyzed.
The characters uscd are mainly ostcological ones,
which show a lower interspecific variability than thc
external morphological characters. Polarity of characters has bccn established according to the outgroup
criterion. The resultant polarity is also indicated
according to the ontogenetic criterion; it is only considered in ordcr to corroborate a certain polarity, or
to indicatc that it might be uncertain. On the othcr
hand, the difficulties inhcrcnt in this criterion arc as
in the outgroup method (Queiroz, 1985).
The outgroup cmployed is the gcnus Opsuriichthys.
It is a basal Leuciscinae which, together with Zacco,
has bcen considered to bc thc most primitive one
(Hcnsel, 1970; Fink and Fink, 1981). Even whcn this
last assumption is doubtful it is generally accepted
that 0psariichthy.s is a more primitive genus than
Burbus.
The rclationships among taxa were establishcd by
parsimony using the Wagner method. These results
are compared with those produced by the CaminSokal method (Felsenstein, 1982). The probability
of the branches being monophyletic has also been
estimatcd by a Bootstrap mcthod. In al1 the analyscs
the program Phylip version 3 (Felsenstein) was used.
Other programs such as Paup and Hcnnig 86 can
also be used to yicld the pliylogenetic trees, but Phylip
was the only which allowed a comparison with the
results obtained through Wagner and Camin Sokal.
In any case, the Wagner trees obtained through the
various programs are quite similar, not differing in
more than two or three evolutionary steps.
Option A, in which the outgroup is codified as al1
zeros, has bcen used for al1 the analyses. A Nelson
Aquat. Living
Resour.
Phylogenetic relationships of species of the genus Barbus
consensus tree has also been used. A similarity analysis was performed using the NTSYS program package
with a Jaccard index and the UPGMA clustering
method.
Since there is a substantial literature concerning
the anatomy of the genus Barbus (Gonçalvez, 1923;
Vandervalle, 1977; Rojo and Ramos, 1982; Doadrio,
1984; Howes, 1987), no further study of anatomy was
undertaken. Therefore only the characters used, their
States and polarities are reported (Appendix 2). The
distribution of character-states for each species is
shown in tuble 1.
267
only character 47 (quadrate height) is uniquely derived.
Burhu.~S. str. is composed of two main branches:
the first branch includes the European species (with
the exception of the Iberian species); the second
branch includes species of Asia (Middle East), northern Africa and the Iberian Peninsula. Two African
species (B. nu.c.u.c. and B. magniut1unti.c.) seem to be
more related to the European group than to the
Ibero-African. However, this is supported by only
one character uniquely derived (the inclination of the
anterior edge of the cleithrum) so its position cannot
Table 1. - Distribution of character-states in the different taxa considered.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 2û 21 22 23 24 25 262728 29 30 31 32 33 34 35 36 37 33 39 4041 42 43 4445 46 47 48 49 50 51 52 53 54 55 56
0 0 0 0 0 1 0 0 0 2 1 0 1 0 0 0 0 1 0 0 0 0 1 0 0 1 2 1 0 0 0 1 0 1 1 0 0 1 0 0 1 0 0 0 1 2 0 1 0 0 0 0 0 0 0 2
0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 2 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 1 1 0 2
hagei
0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 2 0 1 0 1 1 0 2
yirmis
0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 2 1 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 0 1 1 0 2
wlateril
0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 3 0 1 0 1 1 0 2
wlaten2
00000000111 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 3 0 1 0 1 1 0 2
sclatwi3
00000001111 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 1 0 0 0 0 0 1 0 1 0 3 0 1 0 0 1 0 2
0 1 0 0 0 0 0 1 1 1 1 1 0 0 0 2 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 1 1 1 1 0 0 0 1 0 0 0 0 0 0 1 0 3 0 2 0 1 1 0 2
coniza
micraephalusOOOOOOOO111 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 1 1 1 1 1 3 0 1 0 1 1 0 2
0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 3 0 1 0 1 1 0 2
callmsisl
callmsis2 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 1 0 1 0 1 1 3 0 1 0 1 1 0 2
mnilmsis 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 1 0 0 0 0 1 0 1 1 1 1 3 0 1 0 1 1 0 2
mllaryi
0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 0 1 0 0 0 0 0 0 1 1 1 1 3 0 1 0 0 1 0 2
lqiceps
0 0 0 0 0 0 2 1 1 1 1 0 0 0 0 1 1 1 1 0 0 0 0 0 2 0 1 1 1 0 0 1 0 1 3 1 0 0 1 1 0 0 0 0 0 0 1 0 3 0 1 0 0 1 1 2
caito
0 0 0 0 0 0 0 1 1 1 1 1 0 0 1 1 1 1 1 0 0 0 0 1 2 0 1 1 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 3 0 1 0 0 1 1 2
brachpphalusOOOOOOO1111OOO1111 1 0 0 0 0 0 2 0 1 0 0 0 1 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 1 0 3 0 1 0 0 1 1 0
~ i a t 1 a n t i s 0 0 1 0 0 0 1 0 1 0 0 11 0 0 1 1 0 0 0 1 1 0 0 2 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 3 0 1 0 1 1 1 2
~lolepis 1 0 0 0 1 2 2 0 1 0 0 1 0 1 0 1 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 0 0 0 1 0 1 1 3 0 1 0 0 0 1 2
neridianalis 1 0 0 0 1 2 0 0 1 0 0 1 0 1 0 1 0 0 0 1 0 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 2
haasil
1 0 0 0 1 2 0 0 1 0 0 1 0 1 0 1 0 0 0 1 0 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 0 1 0 0 1
haasi2
1 0 0 0 1 2 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 O l O O 1
pdenyi
1 0 0 0 1 2 0 0 1 0 0 1 0 1 0 1 0 0 0 0 1 1 0 0 1 0 0 0 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 1 2
ciwaurasicus1000102110010101 1 0 0 1 1 1 0 0 1 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 3 1 3 0 0 0 1 2
plebejus
1 0 0 1 1 0 2 1 1 0 0 1 0 1 0 1 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 2 0 3 0 0 0 1 2
S&
1 0 0 1 1 0 2 1 1 0 0 1 0 1 0 1 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 3 0 3 0 0 0 1 2
'I~SUS
1 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 1 0 0 0 1 1 0 0 1 0 0 1 0 0 0 1 O1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 3 0 1 1 1 0 1 1
fritwhii
graellsii
RESULTS
Forty-seven equally parsimonious trees were identified with the Wagner method. This is due to the
presence of a high number of homoplasies. In figure 1,
a Nelson consensus tree can be seen. The data used
do not contradict the hypothesis of the existence
of a well differentiated group within the genus Burbus s. str. (Howes, 1987), which includes Palaearctic
species. This group shows many differences from the
probe species (B. fritschi) that we have included as a
representative of the subgenus Labeobarbus. However,
Vol. 3, no 4 - 1990
be established with certainty. In fact, when B. nasus
and B. magniatlantis are excluded from the analysis
the results are more consistent.
The first group, or European group, shares four
synapomorphic characters (21, 22, 49, 55) and seven
if B. magniatlantis and B. nusus are excluded from
the analysis (1, 5, 13, 21, 22, 49, 55). The characters
uniquely derived are: narrow ethmoid; a large prevomer extending over almost the whole anterior part of
the parasphenoid; frontal with a sharp and concave
anterior border; the anterior edge of the cleithrum
inclined.
Nine synapomorph features are shared by the Tbero-African and Asiatic group (8, 10, 11, 18, 19, 25,
3 5 , 4 6 a n d 54)and 11 (8, 10, 11, 18, 19,25, 27,35,
J. Doadrio
fritschi
I
I
,
.
mayiatlantis
nasus
petenyi
meridionalis
haasi 2
haasi 1
cyclolepis
ciscaucasicus
pl e b e j u s
barbus
"il
brachycephalus
longicepe
capito
sclateri 3
coniiza
4 'J
callensis 2
callensis 1
1
1
-microcechalus
~.
sclateri 2
sclateri 1
,-pallaryi
pCïqW
moulouyensis
1 \ bocagei
guiraonis
I 35 l
graellsii
bigure 1.- Phylogenctic rclationships among spccies of the gcnus
Burhu.~cstimated from a Wagncr parsimony method. Autapomorphier are not indicated.
36, 46 and 54) if B. ma,qniatlunti.s and B. na.su.s are
excluded. The characters uniquely derived are: contact
of the exoccipital and the pterotic; high medial process of the urohyal; narrow exoccipital apophysis of
the pterotic; eresccnt shaped masticatory surface of
the pharyngeal teeth; strong fourth pharyngeal tooth;
basal plate of the pharyngeal bone extended and wide
4th and 5th infraorbitals.
Most of the spccies of Ibero-African group, specifically the Iberian (B. hocugei, B. grucllsii, B. sclutcri,
B. guiraonis, B. comiza, B. microccphalus) and north
African (B. cullensis, B. moulouyensi.~, B. pallaryi)
show quite well developed nuptial tubercles in contrast to the European group. However, in our study
this character has been excluded owing to the lack of
information about the Asian species. Some authors
(e. g. Günther, 1868) have considercd this character
in classifying the group, but without mention of its
presence in Ïberian species.
The relationships among B. nasus, B. magniutlantis
and the European group are doubtful. They might
result from convergence or from the absence in the
analysis of other non Palaearctic species which might
constitute a different group to which they belong.
Further studies using other characters and more species might throw light on these relationships which
create many difficulties in our evolutionary and biogeographic analyses.
Within the "European group" there arc two
groups. The first includes B. barbus, B. plehejus and
B. ci.scuucu.sicus.These species live in Central Europe,
ltaly and the Caucasus and share the following synapomorphic characters: a wide parasphenoid and a
long lachrymal. The second group consists of a collection of mediterranean species which have spread over
Central Europe to a certain extent via the Danube
and live in mountain fluvial courses. The osteology
of B. peloponnesius of Greece could not be studied,
but its description (Kollcr, 1926) suggests it is close
to this group. Two synapomorphie characters that
support this cluster are the large extension of the
media1 region of the parasphcnoid and a convex lower
branch of the media1 region of the parasphenoid and
a convex lower branch of the pharyngeal bone.
B. cyclolipis is a problematic species and its inclusion
within the mediterranean species is supported by only
a single character (6).
The ensemble of Iberian and North African species
does not show clear relationships. Basic revisionary
studies of many of them have not been performed
yet, and it is difficult to establish the limits of many
of the species, as for example the North African
B. callensis.
Two groups could be differentiated: that composed
of Asiatic species B. longiceps, B. cupito and
B. brachycephalus and some Iberian (B. comiza and
B. sclateri 3) and B. callensis 2 from north Africa.
The most important problem arising from this analysis of phylogenetic relationships is that the characters
considercd discriminate main groups, but not specific
relationships, particularly in the Ibero-African group.
A detailed study of morphological and genetic charactcrs of the numerous taxa living in northern Africa is
needed, as well as the analysis of their phylogenetic
relationships.
The cvolution of the group presented in this paper
and the hypotheses previously reported by former
authors are not compatible. By and large, we refute
the hypothesis defended by many authors of a close
relationship between the Iberian populations and
B. burhu.~of northern Europe (Almaça, 1967, 1976;
Banarescu, 1960; Berg, 1932; Lozano Rey, 1935).
CLASSIFICATION AND TAXONOMIC
REMARKS
The above mentioncd results imply the need for a
new partition of the genus. In our opinion, the former
classification is not very useful due to the great number of species included. The analysis of reduced
groups improves the consistency of the classification.
According to Hennigian methodology (1 966) there
are two strictly monophyletic groups. However only
the Iberian, African and near Asia Barbus group is
very consistent as its species share a large number of
Aquat. Living Rcsour.
269
Phylogenetic relationships of species of the genus Barbus
characters. The European group, including two species of north Africa, is probably not strictly monophyletic. Only when both north African species are removed from the Europcan group d e s this latter show a
greater stability and consistcncy. We therefore consider the genus Barbus s. str. divided into two different subgenera: the nominal subgenus Barbus Cuvier
and Cloquet, 1816, provisionally composed of
B. barbus, B. plrhejus, B. ciscaucasicus, B. cyclolepis,
B. meridionulis, B. hau.si, B. petenyi, and the subgenus
Luciobarbus Heckel, 1843, which would now include
northern African, Asian and Iberian species.
Luciobarbus was originally referred to a group of
Western-Asiatic species. The osteology of these species has not been presently studied; for this reason
the attribution of the Ibero-African group to this
subgenus is only provisional, in order to facilitate
biogeographical and taxonomic study. However, the
available descriptions concerning the western-Asiatic
species (see Almaça, 1983b) indicate a high number
of common characters. A new subgenus could be
described for the northern African, Asian and Iberian
group, but forma1 description of any new subgenus
without a complctc study of the whole group might
only increase the already chaotic nomenclature of the
genus Barbus.
Another subgenus, Aspiobarbus Berg, 1932 is available for the Iberian, northern African and Asiatic
group, but there is a lack of an adequate definition
of it and only Ladiges and Vogt (1965) have used it.
The re-description of the genus Lucioharbus is as
follows:
Luciobarbus Heckel, 1843.
Synonym: Aspiobarbus Berg, 1932.
Type species: Lucioburhu,s xanthopterus Heckel,
1843.
Diagnosis: The species of this subgenus share the
following synapomorphic characters: 1. The exoccipita1 contacts the pterotic largely. 2. High medial process of the urohyal. 3. Narrow exoccipital apophysis
of the pterotic. 4. Wide 4th and 5th infraorbitals.
Description: Other characters shared by al1 the
species are: 1. Anterior border of the cleithrum perpendicular to the lower limb. 2. Wide basal plate of
the urohyal. 3. Tendency for a decreased pharyngeal
teeth number in adult specimens; the fourth tooth of
the external row becomes strong. 4. In adults crescent
shaped masticatory surface to teeth. 5. Convex anterior border of the opercular. 6 . Wide ethmoid.
7. Small prevomer with a long and sharply pointed
top. 8. Upper limb of the cleithrum elongated and
narrowed. 9. Large pharyngeal cartilage. 10. Basal
plate of the pharyngeal bone extended downwards.
The species studied within this subgenus are listed
below: B. bocagei, B. graellsii, B. guiraonis, B. sclateri, B. microcephalus, B. comizu, B. callensis, B. moufouyensis, B. pallaryi, B. brachycephalus, B. capito and
B-longiceps. The material studied is listed in
Vol. 3, no 4 - 1990
Appendix 1. Osteology of Barbus xanthopterus has
been partially studied.
In order to prove the stability of the classification
suggested, different criteria can be used.
The Camin-Sokal criterion, which does not admit
any reversion (see Felsestein, 1982 for more details),
yields a closer relationship between Labeobarbus and
Lucioburbu.s than between Barbus and Luciobarbus ( j ï g . 2).
petenyi
meridionalis
1,5,6,14
haasi 2
haasi 1
9.16.22,25,48,49.52,57
::
32 57
.
28.50.54
-,
1
plebe jus
barbus
ciscaucasicus
cyclolepis
nasus
magniatlantis
brachycephalus
pallaryi
capito
longiceps
moulouyensis
callensis 2
comiza
sclateri 3
9.16.17.1
12
10,11,18
'
p
l
I
128.47%49.54.57
sclateri 1
callensis 1
bodagei
microcephalus
sclateri 2
guiraonis
graellsii
fritschi
Figure 2.- Phylogenetic rclatioiiships arnong specics of the gcnub
Barbus estimated frorn the Camin-Sokal parsirnony method. Autapomorphies are not indicated.
Applying a Bootstrap for 20 repetitions, we obtain
a percentage of 80% of Luciobarbus, Barbus and
Labeobarbus subgenera being monophyletic and 90%
if the northern African species B. nasus and
B. magniat1anti.s are eliminated.
InfilFure 3, both groups are well defined when using
the phenetic methods based on a Jaccard index and
the UPGMA clustering method. Even though we use
a different concept that admits paraphyletic groups
and a "convex group" classification is made, the
dendrogram does not justify the existence of other
groups. From the phenetic results we could infer
that the subgenus Luciobarbus is more similar to
Labeobarbus than to Barbus.
These data when analyzed by the Camin-Sokal and
UPGMA methods, supported the status of Barbus
and Lucioharbus but not the monophyly of
Barbus S. str.
1. Doadrio
fritschi
graellsii
guiraoni s
bocagei
sclateri 1
callensis 1
r
l
L
sclateri 2
microcephalus
callensis 2
moulouyensis
h
pallaryi
sclateri 3
comiza
L
.capitO
longiceps
brachycephalus
magniatlantis
cyclolepis
ciscaucasicus
plebejus
barbus
L
nasus
rneridionalis
haasi 1
haasi 2
petenyi
Figure 3.- Phylogenctic rclationships among spccics of the genus Burhus eatimated from the Jaccard index and UPGMA cluslering method.
As for more particular aspects of the differcnt taxa,
Our classification has the following taxonomic implications:
- B. ciscuucusicus: Kamaran (1971) included it in
B. plebejus; this synonymization has not been
reviewed and is still used by many authors (see Illies,
1978). According to Our phylogcnetic hypothesis, if
we were to dcfend B. ciscaucasicu.~as a subspecics of
B. plebejus, we would have to consider B. p1eheju.s
as a subspecies of B. burhus, since therc are fewer
distinctive characters between both species. At the
moment, we think there are sufficient characters to
consider the three taxa specitïcally distinct.
- B. haasi: it has been included in B. plehejus by
Almaça (1982) and has been synonymized with
B. capito (Karaman, 1971) or B. hocagei (Almaça,
1971). However, Doadrio (1987) has considcred it as
a well differentiated species. Our study indicates that
it is quite close to B. meridionalis, but we consider it
a distinct species. The population of the Ripoll River
(Besos basin, Catalonia), analyzed separately,
seems to be a hybrid of B. meridionalis and B. huusi
as indicated by biochemical data (Machordom et al.,
1990). Our osteological results seem to corroborate
this. Some populations of B. rneridionalis in the
Besos basin (Doadrio et al., 1988) are identified as
these hybrids.
- B. petenyi: aftcr Koller's revicw (1926) it was
considered to be a subspecies of B. meridionalis. Karaman (1971) however does not follow this allocation
and places it in B. peloponne.siu,s. The existence of
and B. meridionumorc differences between B. pc>~~.nyi
lis than are found between B. huusi and B. meridionulis induces us to give it a specific rank.
- B. m~gniutlunti~s:it has been included as a
synonym of B. nu.su.s (Esteve, 1947; Lévéque and
Daget, 1984). Howcvcr, other authors consider it a
well differentiated species (Almaça, 1970a, h): The
existence of numerous diffcrcnces between both species and thcir coexistence in the same river of the
Oum-er-rbia basin suggest B. mugniut1unti.s can be
considered a wcll differentiated species. Curiously,
B. mugniutluntis as well as B. nusus, and especially
the latter species, are spccics of fast mountain rivcrs
with current. This habitat is not frequented by the
other species of the subgenus Lucioburbu.r which are
prescnt in Morocco and it is similar to the habitat of
European species.
- B. guiraonis: it was described by Steindachner
(1886u), but later included as a synonym of
B. gruellsii (Steindachner, 1866b). Almaça (1983 rr)
examined the types from the Natural History
Museum of Vienna and concluded that B. guiraonis
is a synonym of B. graellsii, though he included the
smallest specimen of the type series in B. hocagei. The
absence of denticulations in the last single ray of the
dorsal fin is the main argument of these authors to
consider B. guiraonis a synonym of B. graellsii and,
in our opinion, to also explain the inclusion of the
smallest specimen of the type series in B. hocagei,
sincc it has quite marked denticulations.
Aquat. Living Kesour.
Phylogenetic relationships of species of the genus Barbus
271
FOSSIL RECORD
Figure 4.- Last single rays of the dorsal fin in B. ~uiraonissyntypes. Denticulations can be observed in al1 spccimcns being more
obvious in juveniles.
Our examination of the same type series indicates
that most of the specimens have weak denticulation
Cfig. 4). In small specimens the denticulations are
strong and occupy the whole ray, as it occurs in
B. bocagei, but in the adults they are weaker and
reduced, even disappearing in some cases. In
B. graellsii exceptionally small denticulations appear
in the smallest specimens. Among other characters,
this one distinguishes the two taxa (B. graellsii and
B. guiraonis) which, at the moment, we consider as
being well differentiated. B. guiraonis is distributed
al1 over the Spanish Mediterranean basins, from the
Mijares to the Serpis River.
- B. sclateri: it is a polytypical species withh three
clearly different populations, each one distributed in
a different region: Guadalquivir and Southern Spain,
Segura basin and Guadania basin. A detailed study
of this species and a complete characterization of its
populations will be published separately.
- B. ca1lensi.s: two different populations have been
detected in Morocco, one occupies the Mediterranean
basins and the other the Atlantic ones. However,
extensive study of the species including also Algeria
and Tunisia populations is needed.
- B. moulouyensis and B. pallaryi: they were
included in B. callensis by Lévêque and Daget (1984).
In Our opinion the differences between them are
greater than those between other species within the
genus and for this reason we consider them clearly
distinct.
- B. brachycephalus: it was considered by Howes
(1987) to be a problematic species because the predorsa1 vertebral number is smaller than that found among
Barbus S. str. According to Our analysis it should be
placed in Barbus S. str., having many characters that
justify its inclusion in this genus (seejïgs. 1, 2 and'3).
For the above mentioned reasons we think that
neither the vertebral count nor the ramified ray number in the dorsal fin are definitive features of Barbus.
The ramified ray number is eight in Barbus S. str.,
seven in B. brachycephalus and ranges from 7 to 8 in
some others such as B. haasi.
Vol. 3, no 4
- 1990
Unfortunately there is a poor fossil record for the
family Cyprinidae, the oldest remains being from the
European lower Eocene (Pattcrson, 1975). The genera
appearing in the European Eocene arc Blicca, Chela
and Rutilus (Novacek and Marshall, 1976). Thc latter
genus has also been found in the Soviet Neogenc
(Lebedov, 1959).
In Africa the oldest known cyprinids come from
the middle-lower Miocene and belong to the genus
Labeo (Baker et al., 1971; Van Couvering, 1977). As
for the genus Barbus , the earliest fossil found in
Africa comes from the Upper Miocene of Tunisia
(Robinson and Black, 1974) and Kenya (Van Couvering, 1977).
Tt is appropriate to point out that the Tunisian
Miocene shows a fauna which resembles that of the
Nile, with genera such as cl aria.^, Polypterus, etc.,
and probably Barbus (Greenwood, 1974). The extinction of this fauna, at present represented only by
Barbus, Pseudophoxinus and some ciclids (Kraiem,
1983), happened after the Miocene. In the Plio-Pleistocene the genus Burbus is frcquently found in Africa
(Greenwood, 1974).
In Europe the oldest putative fossils of Barbus
(B. rudeli Piton, 1936) are found in the middle Oligocene, but further studies have questioned their taxonomic allocation, and they have been included in the
Indo-Malayan genus Puntius (Piton and Theobald,
1939). Thus, the oldest Barbus remains date from the
middle Miocene (Woodward, 1901; Banarescu, 1960)
and are said to be B. steinheimensis (Quenstedt, 1852).
In Spain the earliest record of Barbus (Doadrio
and Casado, 1989) appears in the Upper Miocene.
Based on dental morphology they were assigned to
the subgenus Luciobarbus.
The species B. barbus is present in the European
Pleistocene (Gaudant, 1979). In the Iberian peninsula
B. bocagei is found (Morales, 1980), a species
erroneously cited by this author as B. barbus (see
Doadrio and Casado, 1989).
In Asia, some remains of pharyngeal teeth of Cyprinidae from the Upper Miocene of Japan have been
found. This material has not been referred to any
generic or specific taxon (Yasuno, 1982). In Our
opinion they belong to the subgenus Luciobarbus and
are quite similar to those found in the deposits of
Cullar-Baza Spain (Doadrio and Casado, 1989).
We have discovered some material, from the middle
Miocene of Simancas (Valladolid; Spain), conserved
in the Museo Nacional de Ciencias Naturales
(Madrid) collection and including five pharyngeal
teeth and some vertebral centra, that belong to the
genus Barbus. This deposit has been dated at approximately 16-17 million years old (Magna National Plan
of Geology Carthography), corresponding to the
European Continental biozone 4-5 of Mein of the
1. Doadrio
European continental Neogene. These are the oldest
fossils of the genus Barhu.~known in Europe since
the German deposit of Steinheim has been dated
between the 7-8 biozones of Mein.
The teeth are morphologically similar to those
reported by Quenstedt (1852) from Steinheim and
resemble those of the subgenus Burhu~.They are quite
large, elongated and laterally compressed, with a not
very marked hook at the top and a reduced masticatory surface, with no denticulations, or in some cases
absent. The reduction of the masticatory surface
The known fossil record of Barbus is older than
that of 1,uciohurhu.r and limited to Europe. Lucioharhus has been widely distributed over Asia, northern
Africa and Spain since the upper Miocene.
It is not suggested that the environmental conditions during the upper Miocene possibly favoured
Lucioharhus which seems to tolerate salty fluvial
courses (Kraiem, 1986). However, some places where
this subgenus lives nowadays, such as the Ichkleul
Lake in Tunisia (Kraiem, 1986) must be similar to
the small lakes and fluvial courses present during the
Messinian draining of the Mediterranean.
In summary, the known fossil record seems to agree
with our evolutionary scheme and classification. The
important disagreements bctween the modern theories
on colonization of the genus Burhus and the data
provided by the fossil record will be discussed below.
BIOGEOGRAPHY
Figure 5.- Pharynyeal teeth from the middle Miocene of Simancas
(Valladolid, Spain). Scc tcxt for description.
(Jg. 5) is greater than that found in modern species
of the genus. The teeth of B. meridionalis and B. huu.si
most closely resemble those of the fossils. For the
moment, until an extensive study is made (which is
not the object of this work), they are assigned to
B. steinheimensis.
On the basis of the fossil record it can be deduced
that there were two different evolutionary lines of the
genus barbu.^ in the Miocene. The first one with a
not very differentiated dental morphology is assigned
to the subgenus Barbus. The second with a very derived dental morphology (teeth quite compressed, crecent masticatory surface and a tendency for reduction
of tooth number) is assigned to the subgenus Lucioharbus.
The Ostariophysi consists largely of freshwater species most of which do not show very active dispersal.
Moreover, colonization of new areas seems to bc
restricted by geological and tectonic events. Despite
this fact, most of the current hypothescs on the biogeography of Ostariophysi defend their active dispersal
and share the following points. (1) They agree with
the classic phylogeny of the group (Greenwood et al.,
1966) after which the Characoids are the most primitive (Rosen and Greenwood, 1970); (2) Al1 current
models fix a centre of origin, though this is variable
depending on the authors: Gondwana (Roberts,
1969), Africa (Géry, 1969); South America (Chardon,
1967), Asia (Darlington, 1957; Briggs, 1979). There
is now an alternative hypothesis on the phylogenetic
relationships of the Ostariophysi (Fink and Fink,
198 1). Furthermore, recently the concepts of centre
of origin and dispersal have been heavily critized
(Croizat et al., 1974).
The biogeographical models of cyprinids are al1
very similar, and are also based on the concepts of
centre of origin and dispersion. The genus Burhus is
no exception. It is generally accepted that the group,
sensu stricto, had an Asian origin with a subsequent
dispersal to Europe. In this way Banarescu (1960)
considered West Asia to be the centre of origin for
the Barbinae. The migratory route would have been
from Asia Minor to the Balkans, and from there to
Central Europe, the Iberian Peninsula and northern
Africa. The migration of this genus would have occurred mainly in the middle Miocene, with some more
recent immigrants such as B. meridionalis added later.
Kamaran (1971) indicated that there is no centre of
origin known, but that the genus must have entered
Europe and northern Africa from the Middle East.
Banarescu (1973) suggested a new centre of origin
for Barbus in eastern Asia, from where it would have
Aquat. Living Resour.
Phylogeneîic relationships of species of the genus Barbus
migrated through northern Siberia ultimately to reach
Europe.
Kosswig (1973) felt the genus Barbus had an East
Asian origin and reached Africa between the late
Miocene and the Plcistocene. He is the only author
who does not state that Barbus has colonized the
Iberian Peninsula from Central Europe.
Almaça (1976) indicated that the Iberian peninsula
was colonized by Barbus from Europe in the late
Oligocene, and that some populations moved to northern Africa in the Mio-Pliocene. Almaça (198 1) proposed a similar hypothesis to that of Banarescu
(1973). According to this author the colonization of
Europe would have taken place from Eastern Asia
by two differcnt lines. A line constituted by B. barbus
or its ancestor which would later colonize the lberian
Peninsula and a line that would colonize the meridiona1 peninsulas. Almaça (1988) has implicitly admitted
the existence of two different groups within the genus
Barbus, both with a European dispersal. The Iberian,
northern African and Balkan species would have originated from the first and the remaining species from
the second. The dispersal postulated by al1 authors is
quite extensive. It would have occurred from an Asian
centre, spreading al1 over Europe and arriving in
northern Africa through Spain. A very long geological
time is needed to explain these theories of dispersal,
considering that for fish dispersa1 requires not only a
favourable climatology and availability of resources,
but also important geological changes.
I
1
1 1 1
l
A
t
l
a
s of Morocco
!-Danube,
-5
8
Vistula, Niester
NE.Spain
-
C e n t r a l Europe. $.England
N.Africa
I b e r i a n Penin:uld
Figure 6.- llypothetical rclationships among the dillèrent arcaz.
Principal evcnts known. 1 Uplift of the Urals (35-25 my).
2 Gibraltar Strait? (35-25 my). 3 Paratcthys (20-17 my). 4 Balkans
mountains? (23-12 my). 5 Uphcaval of the Pyrknees (23-12 my).
7 Paratethys (3.5-2.5 my). 9 Red Sea (15-1 7 my). 10 Gibraltar
Strait (6-5 my).
The absence from Europe and Africa of fossil
Barbus until the upper Miocene is the only fact that
seems to confirm the theory of an Asian origin. However, in Asia no earlier record has becn found either.
The European fossils are the oldest known and belong
to the subgenus Burhus. The fossil record indicates
on origin for the genus Barbus at least in the lower
Figure 7.- Present distribution of the species of the genus Barbu.r studied in this work. Vertical= Distribution of the species of the subgenus
Barbus. Horizontal = Distribution of the species of subgcnus Luciobarbus. Solid = Sympatric arca.
Vol. 3, no 4 - 1990
Phylogenetic relationships of species of the genus Barbus
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fishes in Africa. Copeia, 163-166.
Vandervalle P., 1977. Particularités de la tête de deux Poissons Cyprinidés, Barbus barbus (L.) et Leuciscus leuciscus
(L.). Bull. Acarl. r. Bel,. Cl. Sci., 63, 469-479.
Woodvard A. S., 1889-1901. Catalogue of the fossils fishes
in the British Museum, London, 2493 p.
Yasuno T., 1982. Fossil pharyngeal teeth of subfamily
Cyprinidae Fishes Collected from the Miocene Mizunami
Group in Kami Basin, Gifu Prefecture, Japan. Bull.
Mizunami fos.ril Mus., 9, 15-23.
APPENDIX 1
Material List
The material examined is kept in the collections of the
Museo Nacional de Ciencias Naturales of Madrid.
Aquat. Living Resour.
A. Ethmoid morphotypes. 1. Wide (B. callensis). 2. Narrow (B. nusus). 3. Thc ethmoid is wide in its posterior part (B. comizu). 4. Markedly
concave (B. magniatluniis). C. Prevomer morphotypes. 1. Posterior part of the prevomer long and narrow ( B . haasi 1). 2. Posterior part of
the prevomer small and wide (B. barbus). C. Parasphenoid morphotypes. 1. The parasphenoid is widened in its anterior part ( B . bocagei).
2. The parasphenoid is slightly widened in its anterior part (B. hurbu.~).3. The parasphenoid is wide in its antcrior half and narrow in the
posterior one (B. magniutlantis). 4. The parasphenoid media1 part is quite extended (B. haasi 1). D. Sphenotic morphotypcs. 1. Thc process
is wide (B. graellsiz). 2. The process is very sharp (B. magnia~lanfis).E. Casioccipital morphotypes. 1. The cartilage masticatory plate is large
(B. graellsii). 2. The cartilage masticatory plate is small (B. petenyi). 3. The pharyngeal process of the basioccipital is wide (B. pulluryi).
F. Frontal morphotypes. 1. Convex anterior border ( B . guiraoni.~).2. Concave anterior border ( B . cyclolrpi.~).3. More than 10 pores
(B. hrachycephalus). G. Opercular morphotypes. 1. Posterior border slopes weakly ( B . guiraonis). 2. Posterior border slopes slightly
(C. haasi 1).
Aquai. Living Re5our.
. Urohyal
morphotypes. 1. Large basal plate with a rigkt posterior border (0.sclaieri 1). 2. High medial process (B. mugniat1anti.r).
fdarrow basal plate (0.hmsi 1). 1. Cleithrum morphotypes. 1. Long upper branch (0.callensis 1). 2. Short upper branch (B. barbus).
Clritlirum morphotypes. 1. The anterior border is perpendicular (0.cumiza). 2. The anterior border is not perpendicular (B. nasus).
1.: Orbitosphenoid norphotypes. 1. Small (0.guiruonis). 2. Large (0.fritschi). L. Prootic morphotypes. 1. The process is small
(0.mowlouyensis). 2. The process is large (B. capiio). M. Fterotic morphotypes. 1. Narrow ( B . longiceps). 2. VJide (B. rneridionalis).
N. Apophysis of the 4th vertebra morpkotypes. 1. Narrow (0.fritschi). 2. Wide (0.graellsiz].
Vol. 3, no 4
- 1S;3
1. Doadrio
13. Size of the mu.rticatory plate of the busioccipitul.
Description. State 1. The masticatory plate is smaller than
the vertebral centrum. Staie 2. The masticatory platc has
the same sizc than the vertebral centrum. Polurity. Outgroup 1 2.
14. Shape of the unterior border of the frontal (plate 1,
fig. F). Description. State 1. Convex anterior bordcr.
State 2. Concave and sharply pointed antcrior bordcr.
Polurity. Outgroup 1 -+ 2.
15. Pore numbcr of the frontal (platc 1, fïg. F). Bescription. State 1. Less than 10 pores. State 2. More than
10 pores. Polarity. Outgroup 1 -+ 2.
16. Frontal width. Description. The length is mcasured
as thc maximum parallel to thc sagittal axis and the
width as the maximum mcasurcd perpendicular to the
former one. State 1. Widc frontal. Widthilength x 100 > 28.
State 2. Medium-sized frontal. Widthllength x 100> 18<25.
State 3. Narrow frontal. Widthilength x 100< 15. Polurity.
Outgroup 1 -+ 2 -* 3.
17. Operculur posterior border (plate 1, fiK. G). Ijescription. State 1. The opcrcular posterior border slopcs wcakly
in relation to its inncr edge the angle in bctwccn being
greater than 88". State 2. The postcrior border slopes
slightly forming an angle of less than 77". Pohrity. Outgroup 1 -+ 2.
18. Length of the basal plate of the urohyal (platc 2,
fig. H). Description. Statc 1 . The basal plate is small with
respect its media1 process and narrows towards the posterior
part. Statc 2. Large basal plate with a straight posterior
bordcr. Polarity. Outgroup 1 -+ 2.
19. Mediul process of the urohyul (plate 2, fig. H).
Description. State 1. High medial process. Statc 2. Low
medial process. Polarity. Outgroup 1 -+ 2.
20. Width of the basal plate of the urohyul (plate 2,
fig.H).
Description.
State1.Broad
basal
platc.
State 2. Narrow basal plate. Polarity. Outgroup 1 -+ 2.
21. Lcngth of the upper branch of the cleithrum (plate 2,
fig. 1). Description. The branch is measured as the maximum
paralle1 to the inferior border of the cleithrum. The maximum height is perpendicular to the former onc.
State 1. Long upper branch. Lengthiheight x 100 >44.
State 2. Short upper branch. Lengthlheight x 100< 39.
Pohrity. Outgroup 1 -+ 2.
22. Anterior border of the cleithrum (plate 2, fig. J).
Description. State 1. The anterior border is perpendicular to
the external face of the lower branch. State 2. The antcrior
border is at an angle greater than 90" to the external face
of the lower branch. Polarity. Outgroup 1 -* 2.
23. Orhitosphenoid size (plate 2, fig. K). Description.
State I. Small. State 2. Large. Polarity. Outgroup 1 -.2.
24. Size of the sheet of the prootic (plate 2, fig. L).
Description. State 1. Small. State 2. Large. Polarity. Outgroup 1 -* 2.
25. Exoccipital apophysis of the pterotic (plate 2,
fig. M). Description. State 1. Wide and maze-shaped.
State 2. Mcdium-sized. State 3. Narrow and sharply edged.
Polarity. Outgroup 1 -+ 2 -+ 3.
-
O. Pharyngeal tccth morphotypes. 1. Crescent shapc (B. hocugei).
2. Triangular (B.friischi). 3. Right (B. huusi 1). P. Pharyngcal
bonc morphotypes. 1. Lower extrcmity çonvex (B. petenyi).
2. Wide lowcr cxtremity (B. guiruunis). 3. Upper extremity long
(B. comizu). Q. Fourth tooth morphotype. 1. Strong. 2. Quite
strong. 3. Weak.
26. Apoph.ysis width of the 4th certebra (plate 2, fig. N).
Description. State 1. Narrow. State 2. Wide. Polurity. Outgroup 1 2.
-
Aquat. Living Re5our.
2.Maxi:!ar morpkotypes. 1. Palatice process short (0.ciscauca.ricus). 2. Anterior process low (B.selareri 1). 3. Palatine process large and
roiladrd (B.mc~nicitlcintis).4. Palatine process directed backwards (B.comiza). 5. Palatine process directed fonvard (B.frit.~chi).6. High
fritsehi). 2. Quadratojugal process
rnaxilla (C.microcepha~;ls).S . Quadratr morphotypes. 1. Quadratojugal process wide and rounded top (B.
qritr widr (C.guiraonis). 3. Qüadratojugal process narrow ( B . comiza). T. Last single dorsal fin ray morphotypes. 1. No denticulations are
presrnt (B.fritsehi). 2. Drnticulations are only present at the ray base (C.hocagei). 3. Denticulations are strong (B. ciscaucasicus).
4. Denticillations are directed downwards (0.comiza). U. Lachrymal shape morphotypes. 1. Lachrymal long (B.cyclolepis). 2. Lachrymal
short (C.bocqei]. 3. Lachrymal without posterior notch (B.comiza). 4. Posterior part clearly longer than high (B. nasus).
Vol. 3, no 4
- 3433
1. Doadrio
27. Masticatory surfuce shupe of the phuryngeul teeth
(platc 3, jïg. O). Description. State 1. Straight, vcrtical
and weakly devclopcd. State 2. - Crcscent shaped.
State 3. Triangular. Polurity. Outgroup 1 -, 2 -+ 3. Ontogcnetic 1 -+ 2.
28. Phoryngeul teeth numher. Description. Statc 1. Five in
the cxternal row. Statc 2. Four in the extcrnal row. Pohrity.
Outgroup 1 -+ 2. Ontogenetic 1 -+ 2.
Frequently, in the descriptions of some species such as
Barbus hocugei the authors assign them four or fivc tceth
without taking into account that thc fifth tooth disappears
in old agc. This fact has bccn mcntioned alrcady for othcr
African barbels (Matthcs, 1963).
29. Phuryngeul teeth strength. Be.scription. Statc 1. Not
vcry strong. Statc 2. Quite strong. Polurity. Outgroup
1 -+2.
30. Lower brunch shupe of the phuryngeul bone (plate 3,
fïg. P). Description. State 1. In a latcral view, the lowcr
branch is straight. State 2. In a lateral view, thc lower
branch is convex. Polarity. Outgroup 1 + 2.
31. Width of the lower brunch of the phuryngeul hone
(plate 3, fig. P). Description. T o mcasure the pharyngcal
arch we considcr a straight linc cutting the inferior cxtrcme
of the process and the highest point of thc basal plate. The
maximum length is measured parallel to this axis and the
maximum width perpendicular to it. State 1. Wide lower
branch. Widthllcngth x 100> 30. Statc 2. Narrow lowcr
branch. Widthllcngth < 25. Polariiy. Outgroup 1 -t 2.
32. Shupe qf the upper branch of the pharyngcal bone.
Description.
Statc 1. Rounded
antcrior
bordcr.
Statc 2. Sharply pointcd anterior border. Polurity. Outgroup 1 2.
33. Upper brunch length of the phuryngeul honc
(platc 3, fig. P). Description. State 1. Long and straight.
State 2. Short and curved. Polurity. Outgroup 1 -,2.
34. Articulur process of the phuryngeul bone. Description.
State 1. Short and rounded. State 2. Long and sharply
edgcd. Polurity. Outgroup 1 2.
35. Fourth 100th strength ($ the external row (plate 3,
fig. Q). State 1. Quitc narrow. State 2. Narrow.
State 3. Strong. Statc 4. Quite strong. Polurity. Outgroup
1-+2+3+4.
36. Size ($the basal plute ($the phuryngeul bonp. Description. Statc 1. The basal platc is reduced and docs not extend
over the wholc lower branch. Statc 2. The basal platc is
extcnded over the wholc lower branch. Polurity. Outgroup
1 -+ 2. Ontogcnetic 1 -,2.
37. Basal plute shupe of the pharyngeul hone. Description.
State 1 . The basal plate shows no concavity. State 2. The
basal platc is quite concave. Polurity. Outgroup 1 2.
38. Width of the upper brunch of the phuryngeul bone.
Description. State 1. The upper branch narrows to its top.
Statc 2. The upper branch is wider to thc cnd. Polarity.
Outgroup 1 -+ 2.
39. Shape of the plzaryngeal bone. Description.
Statc 1. Not vcry strong. State 2. Quitc strong. Polarit)?.
Outgroup 1 -+ 2.
40. Length of tlre pulatiize process of the maxilla (plate 4,
fi,.. R.). Description. State 1. Short. Statc 2. High. Polurity.
Outgroup 1 4 2.
-+
-+
-+
41. Height of the unterior process oj' the muxillu (platc 4,
Jg. R). Description. Statc 1. Low. State 2. High. Polr~rity.
Outgroup 1 2.
42. Shupe of the pulutine process of the muxillu (platc 4,
Jg. R). Description. State 1. Narrow and not roundcd.
State 2. Large and rounded. Polurity. Outgroup 1 + 2.
43. Palatine process direction (platc 4, fig. R). Description. Statc 1. Directed forwards or straight.
State 2. Dirccted backwards. Polurity. Outgroup 1 + 2.
44. Muxiliur height (plate 4, Jg. R). Description. The
maxilla hcight is measured as the maximum height
pcrpcndicular to the length of the bonc without
taking into account the palatine proccss. State 1. Low
maxilla Hcightllength x 100<24. State 2. High maxilla.
Heigthllcngth x 100 > 30. Polurity. Outgroup 1 -+ 2.
45. Shupe of the unterior proccss qf the muxillu. Description. Statc 1. Straight antcrior border. Statc 2.- Sharply
pointcd antcrjor border elongating ahcad of thc interna1
process. Polurity. Outgroup 1 + 2. Ontogcnetic 1 -+ 2.
46. Quudrudojugul process width (plate 4, fig. S). Description. State 1.- Narrow and sharply pointcd. Statc 2. Wide
with a rounded top. Statc 3. Quite widc. Polurity. Outgroup
1 -+ 2 -+ 3. Ontogcnctic 1 -+ 2.
47.Quudrute height. Description. Statc 1 . High.
Statc 2. Low. Polurity. Outgroup 1 2.
48. Q d r d j u g u l proce.7.r length. Desc.ription. Statc 1. Long.
Statc 2. Short. Pokurity. Outgroup 1 -, 2.
49. I)enticulation of the lust single dordulfin ruy (platc 4,
fig. T). Description. State 1. N o denticulations arc prcsent.
Statc 2. Only somc specimens havc denticulations.
State 3. Thc dcnticulations are only prcsent at the ray base.
Statc 4. The whole ray bears strong denticulations. Pobrity.
Outgroup I 2 -+ 3 -+ 4.
50. Ruy denticulations orientation (plate 4, fig. T).
Description. Statc 1. The denticulations are directed downwards. Statc 2. The denticulations are directed upwards.
Polurity. Ontogcnctic 1 -+ 2.
51. Luchrymul size (plate 4, Jg. U). Description.
State 1. Short. Virtually as short as high. State 2. Short.
Slightly Longer than highcr. State 3. Long. Virtually twice
as long as higher. Polurity. Outgroup 1 -+ 2 -, 3. Ontogcnctic 1 -+ 2 -+ 3.
52. Length of the po.rterior purt qf the hchrymal (platc 4,
jig. U). Description. State 1. Virtually as long as high.
State 2. Clcarly longer than high. Polurity. Outgroup 1 -t 2.
Ontogenetic 1 -+ 2.
53. Luchrymul shupe (platc 4, jig. U). Description
State 1 . Without postcrior notch. State 2. With a marked
posterior notch. Polurity. Outgroup 1 -t 2.
54. Width of the,f&urthundjiyth injruorbituls. Description.
State 1. As wide as the 3rd. State 2. Wider than the 3rd.
Polurity. Outgroup 2 -t 1. Ontogenetic 1 2.
55. Scale number of the lutrrul l i e . Description.
State 1. Less than 53. State 2. Equal to or grcatcr 54. Polurity. Outgroup 1 4 2.
56. Numher of ramified ruys on the dorsal fin. Description.
State 1.- Scven rays. Statc 2. Scvcn or eight rays.
Statc 3. Eight rays. Polurity. Outgroup 1 -+ 2 3.
-+
-+
-+
-+
-+
Aquÿt. Living Resour.