Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana

Abstract

Laimosemion paryagi, new species, is described from the
upper Mazaruni river system, a tributary of the Essequibo
River. It is a member of the Laimosemion breviceps group
former known as the Rivulus breviceps group, and shares a
robust body and deep caudal peduncle with Laimosemion
breviceps (Eigenmann, 1909) and Laimosemion lyricauda
(Thomerson et al., 1991) and to a lesser degree with L.
gransabanae (Lasso et al., 1992) and Laimosemion torrenticola
(Vermeulen & Isbrücker, 2000). These species are all
endemic to the Guiana Highlands in western Guyana and
the neighbouring Gran Sabana in Eastern Venezuela. It is
distinguished from other species in the L. breviceps group
by morphology and its remarkable male color pattern of
red blotches on a turquoise ground color on the flanks and
in the unpaired fins and having a rounded caudal fin as
opposed to one having extension or being spade shaped.
Analysis of mitochondrial DNA sequence reveals that it is
genetically distinct from all other members of this group
and that inhabitants of the Guyana highlands diverged
from each other early in the history of the genus, commensurate
with the geological age of the Guiana Shield.

Full text

Abstract
Laimosemion paryagi, new species, is described from the
upper Mazaruni river system, a tributary of the Essequibo
River. It is a member of the Laimosemion breviceps group
former known as the Rivulus breviceps group, and shares a
robust body and deep caudal peduncle with Laimosemion
breviceps (Eigenmann, 1909) and Laimosemion lyricauda
(Thomerson et al., 1991) and to a lesser degree with L.
gransabanae (Lasso et al., 1992) and Laimosemion torrenti-
cola (Vermeulen & Isbrücker, 2000). These species are all
endemic to the Guiana Highlands in western Guyana and
the neighbouring Gran Sabana in Eastern Venezuela. It is
distinguished from other species in the L. breviceps group
by morphology and its remarkable male color pattern of
red blotches on a turquoise ground color on the flanks and
in the unpaired fins and having a rounded caudal fin as
opposed to one having extension or being spade shaped.
Analysis of mitochondrial DNA sequence reveals that it is
genetically distinct from all other members of this group
and that inhabitants of the Guyana highlands diverged
from each other early in the history of the genus, com-
mensurate with the geological age of the Guiana Shield.
Zusammenfassung
Laimosemion paryagi nov. spec. wird vom Flusssystem des
oberen Mazaruni beschrieben, einem Zuflusssystem des
Essequibo. Diese neue Art gehört zur Laimosemion-brevi-
ceps-Gruppe, die früher Rivulus-breviceps-Gruppe genannt
wurde, und hat einen robusten Körper und einen tiefen
Schwanzstiel mit Laimosemion breviceps (Eigenmann, 1909)
und Laimosemion lyricauda (Thomerson et al., 1991)
gemeinsam, in geringerem Grade auch mit L. gransabanae
(Lasso et al., 1992) and Laimosemion torrenticola (Vermeulen
& Isbrücker, 2000). All diese Arten sind endemisch im
Guyana-Hochland in West-Guyana sowie im benachbarten
Gran Sabana in Ost-Venezuela. Die neue Art unterscheidet
sich von den anderen Arten der L.-breviceps-Gruppe durch
die Körpergestalt und das auffällige Farbmuster der Män-
nchen: rote Flecken auf türkisfarbenem Grund an den
Flanken und den unpaaren Flossen sowie eine abgerundete
Schwanzflosse, während die anderen Arten mit Fortsatz oder
spatenförmigen Schwanzflossen ausgestattet sind. Eine
Analyse der mitochondrialen DNA-Sequenz zeigt, dass die
neue Art sich genetisch von allen anderen Angehörigen der
Gruppe unterscheidet und dass die Bewohner des Guyana-
Hochlandes als Divergenz zu allen anderen Arten in der
Frühzeit der Gattungsgeschichte entstanden sind, gemessen
am geologischen Alter des Guyana-Schildes.
Résumé
Laimosemion paryagi, nouvelle espèce, est décrit comme
originaire du système du haut Mazaruni, un tributaire de
l’Essequibo River. C’est un membre du groupe Laimo -
semion breviceps, jadis groupe Rivulus breviceps, et partage
un corps robuste et un large pédoncule caudal avec
Laimosemion breviceps (Eigenmann, 1909) et Laimosemion
lyricauda (Thomerson et al., 1991) et, à un moindre degré,
avec L. gransabanae (Lasso et al., 1992) et Laimosemion tor-
renticola (Vermeulen & Isbrücker, 2000). Ces espèces sont
toutes endémiques des hauteurs de l’ouest de la Guyana et
du Gran Sabana voisin, à l’est du Venezuela. Il se distingue
des autres espèces du groupe L. breviceps par la morpholo-
gie et par le remarquable patron de coloration du mâle fait
de taches rouges sur fond turquoise sur les flancs et les
nageoires impaires et par une caudale arrondie au contraire
de celle de forme étendue ou en pelle. L’analyse de séquence
mitochondriale d’ADN révèle que l’espèce se distingue
génétiquement de tous les autres membres de ce groupe et
que les espèces des hauteurs de Guyana diffèrent l’une de
l’autre tôt dans l’histoire du genre, proportionnellement à
l’âge géologique du Bouclier guyanais.
Sommario
Laimosemion paryagi, nuova specie, è descritto dal sistema
superiore del fiume Mazaruni, un affluente del fiume Esse-
quibo. Si tratta di un membro del gruppo Laimosemion
breviceps un tempo noto come gruppo Rivulus breviceps, e
condivide con L. breviceps (Eigenmann, 1909), L. lyricau -
da (Thomerson et al., 1991) e, in misura minore, con
L. gran sabanae (Lasso et al, 1992) e L. torrenticola (Ver-
aqua vol. 18 no. 4 - 15 October 2012
181
aqua, International Journal of Ichthyology
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae),
a new species from the upper Mazaruni river drainage of Western Guyana
F. B. M. Vermeulen¹, W. H. Suijker² & G. E. Collier³
1) Tanki Leendert 194c, Aruba. E-mail: vermeulen@setarnet.aw
2) Straatweg 50 Rotterdam, The Netherlands. E-mail: w.suijker@upcmail.nl
3) Department of Biological Science, The University of Tulsa, S. Tucker Dr. Tulsa, OK 74104.
E-mail: glen-collier@utulsa.edu
Received: 16 March 2012 – Accepted: 27 September 2012

meulen & Isbrücker, 2000), un corpo robusto e un altret-
tanto ro bu sto peduncolo caudale. Queste specie sono tutte
en de miche del le Guiana Highlands nella parte occidentale
della Guyana e della vicina Gran Sabana nel Venezuela ori-
entale. Si distingue dalle altre specie del gruppo L. breviceps
per la notevole colorazione del maschio, fatta di chiazze
rosse su un fondo di colore turchese sui fianchi e sulle
pinne impa ri, e per la morfologia tra cui una pinna caudale
arrotonda ta rispetto ad una avente estensione o a forma di
vanga. L'analisi della sequenza di DNA mitocondriale riv-
ela che è ge neticamente distinta da tutti gli altri membri di
questo grup po e che le specie che abitano gli altopiani della
Guya na si allontanarono uno dall'altro piuttosto presto
nella sto ria del genere, commisurato con l'età geologica
dello scu do della Guiana.
INTRODUCTION
The recently erected genus Laimosemion (Cyprin-
odontiformes: Aplocheiloidei: Rivulidae) currently
includes twenty four species (Costa, 2011). It rep-
resents a molecularly defined clade (Murphy et al.,
1999; Hrbek and Larson, 1999; Hrbek et al.,
2004; Suijker and Collier, 2006) within a large,
diverse assemblage of species (formerly assigned to
the genus Rivulus) that are distributed throughout
Northwestern South America; central and eastern
Brazilian Amazon; river basins of Guianas; upper
Orinoco River basin in Venezuela and Colombia;
Rio Negro basin in Brazil; and lower Peruvian
Amazon. They are found in coastal lowland areas as
well as mountainous terrain occasionally up to alti-
tudes of 1300 meters above sea level. Within these
areas they are found in small streams, creeks,
swampy or wet places adjacent to creeks and rain
pools.
Past authors have attempted to arrange related
forms of this larger group into a number of com-
plexes or species groups or subgenera (Hoedeman,
1958 and 1961; Huber, 1992 and 1999; Costa,
2006). Hoedeman (1958) coined the term “brevi-
ceps complex” for “small forms, coarsely scaled”
which included breviceps, frenatus, agilae and geayi.
Huber (1999) grouped these species into the sub-
genus Laimosemion based on small size, distinctive
male patterns, a red opercular shield and that two
had been reported to have bifid epipleural ribs, a
unique character among the Rivulidae. Costa
(2006) refined the morphological and osteological
definition of this subgenus and erected a new sub-
genus Owiyeye for a distinctive set of related species
from west of the Guiana Shield. Subsequently,
Costa (2011) elevated the subgenus Laimosemion
to genus and redefined it to also include those
species he formerly included in Owiyeye primarily
on the basis of previously published molecular
phylogenies (Hrbek and Larson, 1999; Murphy et
al. 1999: Hrbek et al., 2004; and Suijker and Col-
lier, 2006).
The first and second authors have visited Guyana
several times since 1989. These series of expedi-
tions were the first serious attempts to relocate all
species of the former genus Rivulus described by
Eigenmann (1909 and 1912) based upon material
collected during his expedition of 1908. During
these and other expeditions to the neighboring
areas of Venezuela, five additional species have
been described that can be included in this group
of species (Thomerson at al., 1991; Lasso et al.,
1992; Vermeulen and Isbrücker, 2000; Lasso-
Alcala et al., 2006: Suijker and Collier, 2006). The
new species presented in this paper was first col-
lected during a survey of the Guyana plateau by W.
H. Suijker and Y. Suijker, accompanied by S. C.
Paryag on April 4, 1995.
MATERIAL AND METHODS
Specimens analysed: Wild-caught specimens
were used as holotype and paratype while first gen-
eration aquarium bred descendents of wild-caught
fish were used for other paratypes, preparations of
karyotypes and DNA extractions. Specimens of
Laimosemion gransabanae from San Rafael de
Kamoirán, Venezuela, Laimosemion breviceps from
Kaieteur Falls, Guyana and Laimosemion torrenti-
cola from Kamarang, Guyana, were also used for
DNA extractions.
Molecular analyses: DNA was extracted from
ethanol preserved specimens using a DNAeasy kit
(Qiagen). Amplification of portions of the mito-
chondrial genes for 12S rRNA, 16S rRNA, and
cytochrome B was done as previously described
(Murphy and Collier, 1996, and 1997. GenBank
accession numbers for the taxa used are: L. geayi
(AF002433, AF002537, AF002483), L. strigatus
(AF002434, AF002538, AF002484), L. agilae
(AF002432, AF002536, AF002482), L. frenatus
(AF002435, AF002539, AF002485), L. xiphidius
(AF002436, AF002540, AF002486), L. lyricauda
(AF002439, AF002543, AF2489), L. rectocauda-
tus(AF002440, AF002544, AF002490), L. sp.
Tobogan (AF002437, AF002541, AF002487), L.
sp. DCT89-132 (AF002438, AF002542,
AF002488), L. mahdianensis (DQ501248,
DQ501249, DQ501250), R. cylindraceus
(U41799, AF002533, U41781) and R. roloffi
aqua vol. 18 no. 4 - 15 October 2012 182
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana

(U41798, AF002434, U41780). The GenBank
accession numbers for L. breviceps and L. paryagi
are JX885658 through JX885663.
Sequences were aligned by Clustal W and ana-
lyzed using MEGA 4.1 (Tamura et al., 2007).
Karyotype: Metaphase preparations of chromo-
somes were prepared from gill epithelium of a sin-
gle wild-caught male as described by Kligerman
and Bloom (1977).
Morphometrics: Measurements and counts fol-
low Huber (1992). All visible and minute rays
from the anal and dorsal fins were counted. Mea-
surements are taken with a Mitutoyo Dial Calliper
to the nearest tenth of a millimetre and are pre-
sented in percentages of the standard length.
Clearing and staining of a specimen was per-
formed as described by Taylor and Van Dyke
(1985).
Laimosemion paryagi, n. sp.
Figs 1-2, Table I
Holotype: ZMA (Zoölogisch Museum Amster-
dam) 124.896, male, 39,2 mm SL: Guyana, Upper
Mazaruni District, upper Mazaruni river about 7
aqua vol. 18 no. 4 - 15 October 2012
183
F. B. M. Vermeulen, W. H. Suijker and G. E. Collier
Fig. 1. Laimosemion paryagi n. sp. (not preserved). Male from the terra typica. Photo by F. Vermeulen.
Fig. 2. Laimosemion paryagi n. sp. (not preserved). Female from the terra typica. Photo by F. Vermeulen.

miles downstream from Kamarang, a little creek
about 15 minutes walk on a bush trail in eastern
direction starting at the right bank near a rapid
locally called “Sand Landing” (Figures 3 and 4).
Coordinates: 05°43’50”N – 60°30’55” W, altitude
approx. 495 meters above sea level. April 4, 1995.
Paratypes: ZMA 124.897, one female, 36,9 mm
SL: Collected same locality and date as holotype
and 6 males F1 generation, original from same
location.
Diagnosis: Laimosemion paryagi is a member of
the L. breviceps species group as indicated by its rel-
ative small size, short snout length, bright col-
oration and relatively low number of scales
(LL+C). It is distinguished from lowland close rel-
atives (L. agilae, L. xiphidius,L. cladophorus,L.
geayi and L. mahdiaensis) by having a deeper cau-
dal peduncle (14.1-17.3 % of SL, vs. 11.5-13.6 %
of SL). It is distinguished from the remaining
species in the group by having a rounded caudal fin
(versus a spade-shaped caudal fin in L. breviceps,L.
gransabanae and L. torrenticola, and a lyre-shaped
caudal fin in L. lyricauda). Its remarkable col-
oration of red blotches on a turquoise background
on the flanks and in all unpaired fins in males (ver-
sus absence of turquoise color and red blotches) is
unique within the group.
Description: Morphometric data are presented in
Table I. Largest specimen 39.2 mm SL, male. The
dorsal outline of the body has a weak convex pro-
file, straight from anterior of the dorsal fin along
the caudal peduncle. Ventral profile convex, to the
caudal peduncle more straight. Body cylindrical.
Greatest body depth at level of pelvic fin insertion.
Dorsal fin short triangular shaped, no filaments in
either males or females, somewhat larger in males.
Dorsal fin origin directly above 6th or 7th anal fin-
ray. Anal fin rectangular in males, more rounded or
slightly rectangular in females, no fin rays extended
as filaments. Caudal fin rounded in both sexes.
Pelvic fins rounded, in males reaching the second
ray of anal fin, in females smaller not reaching anal
fin. Pectoral fins in males somewhat larger than in
females but both not reaching base of pelvic fin.
Dorsal fin-rays 9; anal fin-rays 11-12; pelvic fin-
rays 6.
Scales large and cycloid, head scaled except the
ventral part of cheeks. Longitudinal series of scales
31-32 upon the end of lateral plate, some scales (3-
5) on the caudal fin base, no scales on base of other
fins. Frontal scales are circularly arranged around
the central A-scale. The squamation pattern is of
the E type in most samples and of the E-D type in
2 of the 8 samples. Scales along the mid- lateral
line have single minute sensory organs.
Epipleural ribs are not bifid (Fig. 8).
Coloration in life: Male: Dorsum brown
to dark brown, laterally the body is light brown
with aubergine red spots arranged to form weak
and broken lines, more intense on the anterior por-
tion. These clear, visible red spots are shown only
on the upper half of the sides and are only weakly
present on the lower half. The lower half of the
sides is iridescent turquoise from the lower edge of
the eye, over the gill-cover, to the end of the caudal
peduncle. Venter and cheeks are whitish to grey.
Iris of the eye is golden. No opercular or humeral
markings are present. The side of the head is
orange to yellow. The dorsal fin is turquoise with
large aubergine- red blotches that tend to form
aqua vol. 18 no. 4 - 15 October 2012 184
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana
Fig. 3. The habitat with the tea colored water lighting up in
the sun. L. paryagi was not seen in the creek itself but in
swampy places beside the creek. Photo by S. Sladkowski.

aqua vol. 18 no. 4 - 15 October 2012
185
F. B. M. Vermeulen, W. H. Suijker and G. E. Collier
lines along the rays, while the distal portion of the
dorsal fin fades to more red than turquoise to form
a marginal aubergine- red zone. Anal fin is bluish
to turquoise near the base with some weak red
blotches. The distal two thirds of the anal fin
becomes more red than turquoise to form a mar-
ginal aubergine- red zone. Caudal fin basis is
turquoise, with the center more iridescent and the
edges of which fade to form an aubergine-red mar-
gin around the outer edge. The central part of the
caudal has rows of numerous red dots that form 3-
4 vertical bars. The pelvic fin is almost completely
aubergine-red with a weak bluish at the base. Pec-
toral fin is hyaline orange.
Female: Dorsum light brown, pale brown on the
sides and yellow brown to the vent. The anterior
part of the flanks show hyaline red markings. Dor-
sal fin is hyaline orange with weak darker dots
forming horizontal lines. Anal fin is hyaline orange
with more orange toward the distal end. Caudal fin
is hyaline orange without any markings or mar-
ginal band. Pelvic fins hyaline. Pectoral fins hyaline
orange. Eye with golden iris.
No “Rivulus spot” or supracaudal ocellus is present
in either sex or at any stage of life. There is a dark
(black) stripe that appears below the midline from
behind the pectoral fin to nearly the base of the cau-
dal. This is seen in both sexes when they are stressed.
Table I. Morphometric data of the holotype and paratypes of Laimosemion paryagi. Standard length is given in mm, other
morphometric data are presented as percentages of standard length. The abbreviation “dev” = deviation.
Holotype Paratypes 8 samples
male female male male male male male male lowest highest average dev
Standard length in mm 39.2 36.9 28.1 26.0 23.0 26.7 35.2 36.9 23.0 39.2 31.5 5.8
In % of SL
Total length 124.0 110.8 119.8 120.9 124.9 119.6 114.7 120.7 110.8 124.9 119.4 4.4
Pre-dorsal length 69.4 69.4 71.7 69.0 73.6 71.3 70.2 69.4 69.0 73.6 70.5 1.5
Pre-anal length 62.0 59.2 58.4 62.6 59.8 60.0 59.7 61.4 58.4 62.6 60.4 1.4
Body depth at anal level 23.0 20.8 22.1 24.7 22.3 23.9 19.8 23.9 19.8 24.7 22.6 1.5
Head length 25.7 21.9 26.2 26.5 27.9 25.8 20.9 24.9 20.9 27.9 25.0 2.2
Snout length 4.1 3.0 5.7 6.5 4.5 4.2 5.2 5.1 3.0 6.5 4.8 1.0
Interorbital space 11.5 12.0 12.7 10.0 12.5 11.7 11.9 13.4 10.0 13.4 12.0 0.9
Eye diameter 7.9 7.1 9.4 7.4 10.5 8.5 7.9 8.5 7.1 10.5 8.4 1.0
Depth of caudal peduncle 15.3 14.7 16.0 17.4 15.3 14.9 14.1 16.5 14.1 17.4 15.5 1.0
Snout to dorsal 69.4 69.4 71.7 69.0 73.6 71.3 70.2 69.4 69.0 73.6 70.5 1.5
Snout to anal 62.0 59.2 58.4 62.6 59.8 60.0 59.7 61.4 58.4 62.6 60.4 1.4
2. Meristic data
Number of dorsal fin rays 9.0 9.0 9.0 9.0 9.0 9.0 9.0 10.0 9.0 10.0 9.1 0.3
Number of anal fin rays 13.0 12.0 12.0 12.0 13.0 13.0 13.0 13.0 12.0 13.0 12.6 0.5
Dorsal insertion to 7.0 6.5 6.5 6.0 6.5 6.5 7.0 7.0 6.0 7.0 6.6 0.3
anal insertion
Longest dorsal fin ray 6.0 4.4 4.2 3.7 3.5 3.7 4.2 6.3 3.5 6.3 4.5 1.0
Longest anal fin ray 7.1 3.1 4.6 4.0 3.6 4.0 4.6 6.0 3.1 7.1 4.6 1.2
3. Scalation
Number scales LL + C 32.0 32.0 32.0 33.0 32.0 34.0 32.0 32.0 32.0 34.0 32.4 0.7
Number scales to 22.0 22.0 22.0 22.0 23.0 22.0 22.0 22.0 22.0 23.0 22.1 0.3
D insertion
Head scalation type E E E E E damaged D-E E-D

aqua vol. 18 no. 4 - 15 October 2012 186
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana
Behaviour: In general, species that were formerly
placed in the genus Rivulus (now divided into
Anablepsoides, Atlantirivulus, Cynodonichthys, Kryp-
tolebias, Laimosemion, Melanorivulus and Rivulus
by the classification as proposed by Costa, 2011),
are non-schooling, solitary fish that are found in
very small streams and swamps. In our collecting
experience, they are found in the dense vegetation
of swamps, under rocks or between logs and leaf
litter in cascading mountain creeks. Often they can
be found in the wet leaf litter adjacent to small
streams. As a result they are often missed in general
Fig. 4. Main river systems of Guyana.

ichthyological surveys of larger fishes or larger bod-
ies of water. They are most often discovered by
coincidence during expeditions with other objec-
tives. Collectors of Laimosemion almost never see
the species that they are looking for until they see
the fish in their nets after “blind” collecting.
Members of the L. breviceps species group display
behaviour that is an exception to the general pat-
tern described above for the larger group of related
genera. Although they do not school, they are
clearly seen not hiding and in open water. This
behaviour, witnessed in the field by authors in
many occasions, is also shared by Anablepsoides
waimacui and Anablepsoides amphoreus. All mem-
bers of the Laimosemion breviceps group, as well as
the two species mentioned above, do not share
their habitat with any other fish species. The lack
of fish predators may be responsible for this
remarkable behaviour.
Molecular phylogeny: Sequences of portions of
the mitochondrial genes for 12S and 16S RNA and
cytochrome b for L. paryagi were added to those
previously determined (Murphy et al., 1999;
Suijker and Collier, 2006). The resulting phy-
logeny places the new species as a member of this
group with the closest relationship being to L. bre-
viceps. Partial DNA sequences for L. gransabanae
and L. torrenticola place them as close relatives of L.
breviceps as well.
Karyotype: Metaphase spreads (Figure 7) reveal
2n=26. The haploid complement consists of one
large metacentric and one medium size metacen-
tric element and a graded series of smaller meta-
centric and submetacentric elements. This is the
lowest number recorded to date for a species of the
genus Laimosemion. Low diploid numbers also
characterize other members of the group shown in
Figure 6 (Suijker and Collier, 2006).
Distribution: Known only from the type locality,
a small black- water creek, about 15 minutes walk
on a hunting trail starting at the right bank of the
upper Mazaruni river about 10 km downstream
from Kamerang near rapids locally called “Sand
Landing”.
Etymology: The name paryagi is chosen to hon-
our Mr. Subhas Chand Paryag, from Georgetown,
Guyana, co-collector of the new species and local
helper during most of the expeditions in Guyana
made by first and second author.
Discussion: Laimosemion paryagi is a member of
the L. breviceps species group as indicated by its
relative small size, short snout length and bright
aqua vol. 18 no. 4 - 15 October 2012
187
F. B. M. Vermeulen, W. H. Suijker and G. E. Collier
Fig. 5. Upper Mazaruni River at Kamarang.

coloration. It is distinguished from other members
of the group principally by the shape of caudal fin
and a unique color pattern. This placement is also
supported by the molecular and cytogenetic data.
The dark line displayed below the midline when
stressed is a character that is shared with L. brevi-
ceps,L. agilae and L. lyricauda. This unique, shared
color pattern may be another character that unites
the group.
This group has also been assigned to the subgenus
Laimosemion (Huber, 1999; Costa, 2006). However,
the primary osteological diagnostic characteristic of
this subgenus was given as “bifid epipleural ribs”
(Costa, 2006). The subgenus Owiyeye was erected
(Costa, 2006) to include species from west of the
Guiana Shield. In Figure 6 this subgenus is repre-
sented by L. rectocaudatus, L. sp. Tobogán and L. sp.
DCT 89-132 (Maroa). This subgenus is character-
ized by, among other characters, “frontal scales
transversely arranged” and “squamation S-pat-
terned”. None of these characters are found in L. pa -
ryagi. Thus, L. paryagi cannot be assigned to either
of these subgenera as defined by Costa (2006).
aqua vol. 18 no. 4 - 15 October 2012 188
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana
Fig. 6. Molecular phylogeny for L. paryagi and selected members of the genus Laimosemion. Neighbor-joining tree based
upon 1048 base pairs of portions of the cytochrome b, 12S, and 16S mitochondrial genes. The values below the nodes are
bootstrap values. Rivulus cylindraceus and Rivulus roloffi were used as outgroups to root the tree. Previous subgeneric group-
ings (Costa, 2006) and current generic assignment (Costa, 2011) are indicated by brackets.
Fig. 7. Metaphase chromosomes from gill epithelia of Lai -
mo semion paryagi.
L. gaeyi
L. strigatus
L. agilae
L. frenatus
L. xiphidius
L. lyricauda
L. mahdiaensis
L. paryagi
L. breviceps
L. rectocaudatus
L.
sp. Tobogan
L.
sp. DCT89-132
R. cylindraceus
R. roloffi

Costa (2011) conceded that the morphological
characters had not been fully examined in all
species of the subgenus Laimosemion and recog-
nized the monophyly of the set of species included
in the subgenera Laimosemion and Owiyeye
revealed by previous published molecular phyloge-
nies. Accordingly Laimosemion was raised to
generic rank to include all species of the two for-
mer subgenera.
However, the species of Owiyeye differ from the
other species in the new genus in their size, behav-
iour and, most importantly, geographical distribu-
tion. Unfortunately, most species formerly assigned
to Owiyeye have not been made available for mole-
cular analysis. Until there can be a comprehensive
analysis of both morphological and molecular
characters, the taxonomic status of these species
will remain ambiguous. For these reasons, we pre-
fer to informally use the term breviceps group to
refer to those small species found along the eastern
edges of the Guiana Shield and adjacent lowlands
where they are found in creeks and wetland areas
with stony or sandy bottoms.
The Guiana Shield is an ancient geological for-
mation that forms one of the three cratons of the
South American plate. The Guiana highlands are
the higher elevations of this formation found in
west central Guyana. Some of the most spectacu-
lar waterfalls in world flow off these highlands to
the surrounding lowlands. These highlands were
presumably never inundated by epicontinental
seas. Hence species of freshwater fishes endemic to
this region likely diversified over a longer period of
time than species from the surrounding lowlands.
The molecular phylogeny is consistent with this
notion in that all members from the interior and
highland areas of the Guiana Shield are deeply
diverged from one another. Even though these
species are each other’s closest relatives, they repre-
sent a diversity of morphology and coloration
unparalleled in other parts of the former genus
Rivulus. Whether this is simply a function of the
age of the Guiana Shield or other factors remain to
be seen.
ACKNOWLEDGEMENTS
Authors are grateful to the Ministry of Fisheries
and Wildlife of the Co-operative Republic of
Guyana providing permits, to Subhas Chand
Paryag and his wife Jenny for their hospitality and
help during most expeditions of authors and to
Mr. Vishnu Mesir, at that time head of the Upper
Mazaruni district, for his help and hospitality in
Kamarang. They also thank Mr. Siegmund Slad-
kowski, Germany for the use of his image from the
Terra Typica and his assistance in the field.
aqua vol. 18 no. 4 - 15 October 2012
189
F. B. M. Vermeulen, W. H. Suijker and G. E. Collier
Fig. 8. Fifth epipleural rib dissected from the right side of a cleared and stained specimen of L. paryagi.

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aqua vol. 18 no. 4 - 15 October 2012 190
Laimosemion paryagi (Cyprinodontiformes: Aplocheiloidei: Rivulidae), a new species from the upper Mazaruni river drainage of Western Guyana