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Contributions to Zoology, 74 (1/2) 97-115 (2005)
Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paría
Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907
and other subgenera of Poecilia Bloch and Schneider, 1801
(Teleostei, Cyprinodontiformes, Poeciliidae)
Fred. N. Poeser1, Michael Kempkes2, Isaäc J. H. Isbrücker1
1 Zoological Museum Amsterdam, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, The Nether-
lands, e-mail: poeser@science.uva.nl; 2 Am Mühlenberg 25, D-46419 Isselburg – Anholt, Germany
Keywords: Guppies, Poecilia reticulata, P. wingei, melanophore patterns, character displacement, Paría
Peninsula, despeciation
Abstract
The taxonomy of the common guppy, Poecilia reticulata Peters,
1859, is reviewed and the closely related Campoma guppy, P.
wingei n. sp., is described. Formerly, the common guppy was not
judged to be closely related to any other species of Poecilia, but
the new species is the second species to be allocated in the sub-
genus Acanthophacelus Eigenmann, 1907. The recognition of P.
wingei results from observed character displacement, i.e., on the
interaction between two closely related species in a shared envi-
ronment. In addition to differences in coloration, behaviour also
indicates specic differences. The area in which P. wingei occurs,
the Campoma region at the base of the Paría Peninsula in Vene-
zuela, hints to an origin of the subgenus Acanthophacelus prior
to the uplift of the Cordilleras, i.e., the eastern orogenesis of the
Andes. Moreover, an explanation is offered for aberrant molecu-
lar data in Trinidadian guppies.
Contents
Introduction ..................................................................................... 97
Methods .......................................................................................... 99
Material .......................................................................................... 99
Collection data of the 2002 expedition ............................. 102
Systematic section ....................................................................... 103
Poecilia (Acanthophacelus) wingei n. sp. ........................ 103
Characterization of the genus Poecilia ............................. 105
Characterization of the subgenus Acanthophacelus ....... 105
Comparison of the colour patterns between the
subgenera Acantophacelus and Micropoecilia .......... 106
Spatial distribution of the phenotypes of the
freshly collected guppies ................................................ 108
Evidence of character displacement .................................. 108
Behavioural differences between Poecilia reticulata and
P. wingei ................................................................................... 109
Behaviour of the common guppy ....................................... 109
Behaviour of the Campoma guppy .................................... 109
Remarks on population differentiation in Trinidadian
guppies ..................................................................................... 111
Remarks on the 'Endler’s live-bearer' ...................................... 113
Acknowledgements ..................................................................... 114
References ..................................................................................... 114
Introduction
The common guppy, Poecilia reticulata Peters, 1859,
has a long history as a monotypic taxon. Poeser
(2003, chapter 11) considered Poecilia Bloch and
Schneider, 1801 as a monophyletic assemblage of
nine morphological distinct species groups, all with
their own evolutionary origin but with no species
group more related to one group or the other. Because
the morphological distinctions are sometimes quite
remarkable, a further in-group analysis was per-
formed (Poeser, unpubl.) and the apparent autapo-
morphies for P. reticulata (cf. Poeser, 2003, chapter
9) proved to be synapomorphies for P. reticulata and
the species described as new herein. The subgenus
Acanthophacelus Eigenmann, 1907 is again recog-
nised based on these synapomorphies (cf. Poeser and
Isbrücker, 2002; Poeser and Isbrücker, in Poeser,
2003), prompting a re-examination of all species
groups mentioned in Poeser (2003, chapter 9).
Poecilia (Acanthophacelus) reticulata is a well-
known little sh, whereas its taxonomic allocation
was fuzzy. Based on a single female, Peters (1859)
allocated a new species from Venezuela to the genus
Poecilia, viz., P. reticulata. Since Peters (1859) did
not mention more specimens, the taxonomic status
of this species is based on this single specimen (Poe-
ser and Isbrücker, 2002; Poeser and Isbrücker, in
Poeser, 2003, chapter 6). In addition, conspecic
specimens from the same lots as the holotype (topo-
98 F.N. Poeser et al. – A new guppy from Venezuela
P. reticulata. He established his Mollienesia on the
basis of the terminal hooks on the gonopodium and
Poecilia by the bumps on ray 4a. Poecilia reticulata
possesses both a hook on the gonopodium and has a
‘bumpy’ gonopodial ray 4a, i.e., it has both of the
above features. Rodriguez (1997: 673) thus was con-
tradicting his own character assessment. Also phylo-
genetic analyses based on molecular data (Ptacek and
Breden, 1998; Breden et al., 1999) did not render a
resolved taxonomy. Moreover, the latter phylogeny
clustered P. reticulata with P. parae and P. picta,
separating these species from the remaining species
of Poecilia. Based on the very distinct morphology
that is presented in this paper, we hold the guppy dis-
tinct from the other species of Poecilia and we split
P. reticulata and the new species off from the species
of Micropoecilia, viz., P. parae Eigenmann, 1894, P.
branneri Eigenmann, 1894, P. bifurca (Eigenmann,
1909), P. picta Regan, 1913 and P. minima Costa and
Sarraf, 1997. Differences between P. reticulata, P.
parae and P. branneri are summarised in Table II.
Another method to unravel taxonomic difculties
is to study behavioural patterns, preferably under
natural conditions (Clark and Aronson, 1951; Liley,
1966). Studies on guppy behaviour yielded a standard
description for mating behaviour in P. reticulata (cf.
Baerends et al., 1955; Table III). Because the initial
attempts to distinguish a new species of guppies
based on preserved material was inconclusive, it was
decided to make eld observations on behavioural
aspects, in order to gain insight on possible specic
differentiations. Collection data of this eld trip are
herein presented separately from the materials exam-
ined prior to this journey.
Detailed information on guppy populations is
fragmented and concerns examinations on a local
scale, e.g., Trinidad. General information on mor-
phometric and pigmentational data comprising the
types) were shipped by Peters from Berlin to London
(Paepke, 1986). They were recorded by Günther
(1866) as Girardinus reticulatus, and were compared
to Girardinus guppii Günther, 1866. Girardinus Poey,
1854 (type species, G. metallicus Poey, 1854), is a
valid genus, distantly related to Poecilia. The type
material of Girardinus guppii was examined (Poeser,
submitted manuscript) and conrmed as identical to
P. reticulata.
Eigenmann (1907) examined specimens collected
from Barbados and Guyana. He allocated the guppy
to a distinct genus, viz., Acanthophacelus Eigenmann,
1907, mainly based on structures of the gonopodial
tip. Eigenmann (1907) designated Poecilia reticu-
lata Peters, 1859 type species for his new genus.
In a breeding report concerning guppies in the
Royal Botanical Gardens in London, Boulenger
(1912) mentioned guppies as the only freshwater sh
on Barbados, which island was the alleged type local-
ity of Lebistes poecilioides De Filippi, 1861, thus
naming this species Girardinus poecilioides. Regan
(1913) changed this by re-naming this species Lebi-
stes reticulatus. Lebistes remained monospecic
until Rosen and Bailey’s (1963) revision of the Poe-
ciliidae, in which Poecilia (Lebistes) reticulata was
joined in one subgenus with four species of Micro-
poecilia Hubbs, 1926 (Table I), and with Cnesterodon
scalpridens Garman, 1895. The latter species was
re-allocated by Costa (1991) as Pamphorichthys
scalpridens, whereas Poecilia (Micropoecilia) was
considered as a distinct genus by Meyer (1993).
Thereafter, the guppy largely remained in the mono-
specic subgenus Lebistes, although not without
debate (cf. Costa and Sarraf, 1997; Poeser and Is-
brücker, 2002; Poeser and Isbrücker, in Poeser,
2003).
The phylogenetic analysis of Rodriguez (1997)
did not adequately resolve the taxonomy concerning
Table I. Allocation of species of the subgenus Lebistes sensu Rosen and Bailey, 1963.
Poecilia reticulata Peters, 1859 Poecilia (Acanthophacelus) reticulata
Poecilia wingei n. sp. Poecilia (Acanthophacelus) wingei
Poecilia parae Eigenmann, 1894 Poecilia (Micropoecilia) parae
Poecilia branneri Eigenmann, 1894 Poecilia (Micropoecilia) branneri
Poecilia bifurca (Eigenmann, 1909) Poecilia (Micropoecilia) bifurca
Poecilia picta Regan, 1913 Poecilia (Micropoecilia) picta
Poecilia minima Costa and Sarraf, 1997 Poecilia (Micropoecilia) minima
Poecilia scalpridens Garman, 1895 Pamphorichthys scalpridens (cf. Costa, 1991).
99
Contributions to Zoology, 74 (1/2) – 2005
total natural range of variation in guppies is not
fully comprehended, although local studies have
provided information on evolutionary mechanisms
shaping guppy characteristics. While it is unknown
what the complete natural range of P. reticulata is,
what the natural variation in colour patterns is, or
what the variation in their anatomy is, no standard is
available to which potential new species can be
compared. In order to provide a more specic descrip-
tion of this otherwise well studied species, this study
rst provides such descriptions for the two species
of Venezuelan guppies.
Methods
To catalogue variation in melanophore patterns, material of the
Zoological Museum in Amsterdam (ZMA) and the University of
Michigan (UMMZ) was examined. Arbitrarily, ten males and ten
females from UMMZ 158720, Ciudad Bolivar, Venezuela, were
selected to set an initial standard. Primarily, only black spots were
noted. Both sides of at least ve males of the additional UMMZ
samples were examined. Horizontal lines were scored on the
anterior part of body (body line) and on the caudal peduncle
(caudal line). Some melanophore patterns are illustrated (Figs.
1-5), and the gonopodia of these specimens (Fig. 6). Morphomet-
ric features scored were standard length (SL) and the least depth
of the caudal peduncle. The average standard lengths of the
populations and the proportional size of the caudal peduncle (as
percents of SL) are recorded as follows:
(number of individuals measured; range of measurements):
average.
Extralimital material of the Zoological Museum of Amster-
dam (ZMA) was examined to demonstrate variation in pigmen-
tation. In July 2002, Poeser and Kempkes made a eld trip to
Venezuela. On the west side of the Paría Peninsula, they col-
lected the species that is described as new in the present paper.
In the diagnosis, the following abbreviations are used. A. =
number of anal n rays; D. = number of dorsal n rays; C. =
number of caudal n rays; LLS. = number of scales in a lateral
series; CPS. = number of scales around the caudal peduncle.
Material
Standard population for Poecilia reticulata, UMMZ 158720 (44
specimens), Venezuela, Ciudad Bolivar, Río Tanguati, tributary
to Río Caroni to Orinoco, 11-03-1939, coll. F. F. Bond. Average
length females (10; 18.4-23.8): 20.4 mm SL, males (10; 12.9-
15.0): 14.2 mm SL. Average least depth caudal peduncle females
(10; 9.0-11.9): 11.0, males (10; 10.1-14.0): 11.9. Some females
may have weak blotches. Males have caudal lines, four (of ten)
have a body line as well. All males have a spot on the anterior
part of body: six have this spot dorsally and four ventrally, one
male only has a spot on the left side of his body. Five males (of
ten) have a black spot halfway on the body: two dorsally and three
ventrally. Two of the latter males only on the left side of the body,
one male only on the right side. Six males have a spot on the
caudal peduncle, just in front of, or just on the caudal n, two
males with a caudal spot only at the right side of the body. One
male has the left spot just in front of the caudal n, the right spot
just on the caudal n.
Table II. Anatomical differences between Poecilia reticulata, P. parae and P. branneri.
Poecilia reticulata Poecilia parae Poecilia branneri
Polychrome body Males are all different, Males exhibit three or Males exhibit a single
no xed patterns four xed colour patterns xed colour pattern
Female body pigmentation Females without Females frequently with a Females with a humeral blotch
humeral blotch humeral blotch and a blotch at the caudal base
Gonopodial structures Palp on ray 3 extending Palp on ray 3 reaching tip; Palp on ray 3 reaching tip;
clearly beyond the tip; no hooks or spine no hooks or spine
hook on ray 5p
Meristic data Nine anal n rays; six or Eight anal n rays; eight Eight anal n rays; eight dorsal
seven dorsal n rays (modally) dorsal n rays (modally) n rays (modally)
Table III. Standard of behavioural sequences in Poecilia reticulata (After Baerends et al., 1955)
1a. Male approaches female. If the female disengages this contact, he will follow her.
1b1. Non-receptive females accelerate the ight, followed by the male. The male will attempt to nip her genital region (gonopore) to
stimulate a sexual response, or he will attempt a sneak-copulation, i.e., to copulate without female co-operation.
1b2. Receptive females slow down to signal the male that she will co-operate.
2a. The male positions in front of the female, slightly swimming backwards and circling in front of the female.
2b. The male displays a sigmoid body posture, showing court-jumps, i.e., sudden jumps centimetres backward.
2c. The male circles to the side of the female and tries to insert his gonopodium.
100 F.N. Poeser et al. – A new guppy from Venezuela
UMMZ 158704 (30 of 1074), Venezuela, 5 km east of San
Mateo, 70 km south of Barcelona, Río Punctual, tributary to Río
Araque, 21-03-1939, coll. F. F. Bond. Average length females (5;
23.1-25.9): 23.9 mm SL, males (5; 13.7-15.6): 14.9 mm SL.
Average least depth caudal peduncle females (5; 11.3-137): 12.8,
males (5; 12.7-14.1): 13.4. This population is seemingly more
coloured than UMMZ 158720. One female has a black spot at
the caudal peduncle. All males have caudal lines, some have a
body line as well. One male exhibits a double body line, as mar-
gins of a brighter band between them. Three males (of ve) have
a spot in the anterior section of their bodies, two other males have
a spot halfway the body. All males have a caudal spot, in some it
is more anterior than in others. Several males have brighter areas
on their anks, suggesting that other colour spots were present.
The unpaired ns are pigmented.
UMMZ 158706 (30 of 656), Venezuela, Isla de Margarita, El
Valle, Río Porlamar, 31-03-1939, coll. F. F. Bond. Average length
females (5; 35.0-40.4): 37.9 mm SL, males (5; 18.5-19.0): 18.8
mm SL. Average least depth caudal peduncle females (5; 15.2-
17.3): 16.2, males (5; 16.8-19.5): 18.0. Two females with a spot
in the caudal n, one up, one down. Males mostly multi-spotted,
varied on left and right side of body (Figs. 1a-d). Body- and
caudal lines frequently thickened into spots. Many males with
caudal n patterns.
The gonopodium of one of the males (Fig. 1c) is also gured
(Fig. 6).
UMMZ 158711 (69), Venezuela, 30 km east of Upata, Río
Charichapo, tributary to Río Yuruori, 12-03-1939, coll. F. F. Bond.
Average length females (5; 18.0-21.2): 19.8 mm SL, males (5;
13.7-15.4): 14.4 mm SL. Average least depth caudal peduncle
females (5; 10.4-13.4): 11.4, males (5; 8.8-11.6): 10.4. One female
has a thin lateral line, best seen on right side of body. Males with
two, sometimes three spots (Fig. 2a-c). Males always with a
caudal line, sometimes with a weak body line. Sometimes the
spots are nothing more than thickened lines. Basal caudal n
pigmentation is present in one examined male, in others similar
markings are found.
UMMZ 158717 (30 of 445), Venezuela, Lagoon 3 km W of
Cumaná, 26-03-1939, coll. F.F. Bond. Average length females (5;
25.9-30.4): 28.4 mm SL, males (5; 17.1-18.0): 17.4 mm SL.
Average least depth caudal peduncle females (5; 11.4-13.7): 13.0,
males (5; 15.0-17.0): 16.1. This population is seemingly more
coloured than UMMZ 158720 or UMMZ 158704. Males have a
spot in the anterior half of the body or more centrally, and have
a spot in C. All males have a varied pattern of lines and bands,
bordering brighter areas. Although most unpaired ns are badly
damaged, some males exhibit caudal n patterns resembling
Ferrugineus (cf. Winge, 1922; here reproduced as Fig. 7), i.e., a
crescent of pigment spots at the caudal base.
UMMZ 158723 (30 of 402), Venezuela, 12 km northwest of
Cumanacoa, 45 km southeast of Cumaná, tributary to Río Man-
zanare, 26-03-1939, coll. F. F. Bond. Average length females
(530.3-40.7): 34.4 mm SL, males (5; 19.0-21.8): 20.7 mm SL.
Average least depth caudal peduncle females (5; 13.3-15.2): 14.2,
males (5; 16.3-18.5): 17.5. The pigmentation in this population is
comparable with UMMZ 158717, but darker. No Ferrugineus
caudal n pattern was observed.
UMMZ 158729 (30 of 822), Venezuela, Caracas, Río Guaire,
Quebrada Cano, la Florida, 10-01-1939, coll. F. F. Bond. Aver-
age length females (5; 26.6-30.7): 29.1 mm SL, males (5; 18.7-
25.1): 21.2 mm SL. Average least depth caudal peduncle females
(5; 12.8-15.0): 14.0, males (5; 13.9-16.1): 15.2. Males have a
varied number of spots (Figs. 5a-e), sometimes differing mark-
edly on left side (Fig. 4a) and right side (Fig. 4b). Pigmentation
is also dominated by horizontal lines and bright areas (Figs.
4c-d), which is reduced in some specimens (Fig. 4e). Most
specimens have a variegated caudal n pattern.
Fig. 1a-c. (Photographs by J. v. Arkel). Pigmentation patterns of
male guppies from Isla de Margarita, Venezuela (UMMZ 158706).
Scale is 1 cm.
Fig. 2a-b (Photographs by J. v. Arkel). Pigmentation patterns of
male guppies from Río Charichapo, Venezuela (UMMZ 158711).
Scale is 1 cm.
a
c
ba
b
101
Contributions to Zoology, 74 (1/2) – 2005
Stock, 1982. One male (20 mm) with a spot on the anterior part
of the body and one at midsection. Caudal peduncle lateral pattern,
caudal n pattern.
ZMA 100.595 (1 ex.), Venezuela, Isla de Margarita, Porlamar,
coll. P. Wagenaar Hummelinck, 17-07-1936. One pale male. Body
spot at dorsal n base, one at caudal n base.
ZMA 123.631 (34), Venezuela, Isla de Margarita, Toma de
agua de Encañado, coll. P. Wagenaar Hummelinck, 13-07-1936.
UMMZ 158730 (75), Venezuela, Río Mariara, 12 km west
of Maracay, 12-01-1939, coll. F. F. Bond. Average length fe-
males (5; 22.5-24.6): 23.5 mm SL, males (5; 13.4-17.1): 14.9
mm SL. Average least depth caudal peduncle females (5; 12.4-
13.8): 13.3, males (5; 11.7-15.2): 12.9. One female with heavy
pigmented fore body. Smaller males all have caudal stripes
and two or three spots, larger specimens have less pigment.
Two larger males (15.4 mm and 16.5 mm) have very little
pigment and immature gonopodia.
UMMZ 158736 (30 of 259), Venezuela, 40 km west of Puerto
Cabello, Río Urama, tributary to Río Yaracuy, 26-01-1939, coll. F.
F. Bond. Average length females (5; 21.6-23.0): 22.3 mm SL, males
(5; 14.1-15.5): 14.5 mm SL. Average least depth caudal peduncle
females (5; 13.0-14.8): 14.0, males (5; 12.8-16.6): 14.8. Most males
have a shoulder spot, with a spot present ranging from mid-body,
to caudal base (Figs. 5a-d). Caudal stripe thin or absent.
ZMA 120.725 (7), Venezuela, Well Cipare, 2.85 km from
Guaibacoa, 3.2 km on side road Sierra de San Luiz, coll. J.H.
Fig. 3a-d (Photographs by J. v. Arkel). Pigmentation patterns of
male guppies from the Paría Peninsula, Venezuela (UMMZ
158715). Scale is 1 cm.
Fig. 4a-e (Photographs by J. v. Arkel). Pigmentation patterns of
male guppies from Río Guaire, Venezuela (UMMZ 158729).
Scale is 1 cm.
a
b
c
d
a
b
c
d
e
102 F.N. Poeser et al. – A new guppy from Venezuela
The males have spots on the anal base, caudal n base with
pigmentation pattern.
ZMA 123.632 (6), Venezuela, Isla de Margarita, Toma de agua
de Tacarigua, coll. P. Wagenaar Hummelinck, 11-08-1936. Three
subadult males, pale. Spots on the anterior part of the body and
midsection.
ZMA 123.634 (46), Venezuela, Isla de Margarita, Asuncion,
Río Asuncion, coll. P. Wagenaar Hummelinck, 11-05-1936. Males
are multi-spotted with a vertical pattern on caudal peduncle.
Caudal n has upper margins, lower margins, midsections or both
margins coloured, or is even completely coloured. Some females
with spots on caudal n base or on caudal n.
ZMA 123.635 (31), Venezuela, Isla de Margarita, Asuncion,
Río Asuncion, coll. P. Wagenaar Hummelinck, 03-07-1936. Dark
specimens, including females.
ZMA 119.916 (20), Guyana, East Coast Demarara, Ogle, coll.
M. Temassar, 27-01-1987. Pale specimens. Males with midsection
spot, one or two spots on caudal n base.
ZMA 119.918 (16), Guyana, East Coast Demarara, Industry,
coll. M. Temassar, 28-01-1987. Multi-spotted males with two to
four spots, vertical stripes on caudal peduncle. Caudal n pattern
present.
ZMA 115.037 (4), Guyana, Canal to Demarara River, near
Georgetown, coll. P. de Rham, 11-10-1976. Anterior spot, caudal
spot, and a small spot in the midsection.
ZMA 109.626 (1 ex. [ex Carnegie museum]), Guyana, Geor-
getown trenches, det. C. H. Eigenmann, no further data. Male with
spots on anterior part of body and caudal peduncle. Vertical pat-
tern caudal peduncle.
ZMA 100.527 (16), Suriname, District Marowijne, coll. P.
Florschutz, December 1952. 16 Multi-spotted males with lateral
patterns and caudal n patterns
ZMA 115.116 (1 ex.), Suriname, Marowijne district, Pankoe-
koe Creek, coll. M.P. Panday, June 1974.
Body with anterior spot and caudal n base spot. Lower
margin caudal n pigmented.
ZMA 119.999 (10), Suriname, Paramaribo, coll. W.C. v. Heurn,
August 1911. Males are multi-spotted, spots on anterior part of
body and midsection, caudal peduncle and caudal n base.
ZMA 120.724 (4), Barbados, Well of Thicket, coll. J.H. Stock,
10-06-1978.
One multi-spotted male, spots on anterior part of body, midsec-
tion, caudal n base. Caudal n pattern present.
ZMA 109.627 (5, ex Carnegie museum), Barbados, det. C.H.
Eigenmann, no further data. Three males, dark. Spots on midsec-
tion and anterior part of body, caudal n pattern present.
ZMA 120.727 (78), Barbuda Island, coll. P. Wagenaar Hum-
melinck, 09-05-1955.
Dark specimens. Males are multi-spotted with a caudal n
pattern. ZMA 120.006 (27), Pacic Ocean, Cook Islands, Vai
Momoiri, Sink hole in Makatea, coll. N.W. Broodbakker, 08-08-
1986. Males have spots in midsection, anal n base and caudal
n base.
Collection data of the 2002 expedition
In the summer of 2002, Poeser and Kempkes stayed in a posada
near Cariaco, on the highway to Casanay. From this base daily
trips were made to localities in the surroundings, guided by a
local taxi driver, Mr. Luis Palacio, who grew up in the region.
Mr. Palacio thus provided much information, e.g., the names of
the waters, villages, etc. At nearly all localities in the proximity
of the posada guppies were abundant, easy to catch with a simple
dipping net. The samples were divided later into 'Campoma' gup-
pies (Poecilia wingei n. sp.) and common guppies (Poecilia re-
ticulata), hereafter indicated as such.
13-07-2002. Station 1: Road from Cariaco to Carúpano, bridge
at the division of Laguna Campoma and Laguna Buena Vista, ca.
3 km north of Cariaco. Male guppies had a melanophore pattern
as in UMMZ 158715. All other colours much enhanced by metal-
lic sheen, they are brilliant. Also females exhibited a more
brightly sheen on their bodies, compared to the guppy females
we were used to. Some other, larger sh, probably cichlids, were
present. A collection of guppies was made, deposited in ZMA
123.636 (Campoma guppy).
13-07-2002. Station 2: Irrigation ditch ca. 50 meters from the
road from Cariaco to Carúpano, ca. 2 km north of Cariaco. Gup-
Fig. 5a-d (Photographs by J. v. Arkel). Pigmentation patterns of
male guppies from Río Urama, Venezuela (UMMZ 158736). Scale
is 1 cm.
a
b
c
d
103
Contributions to Zoology, 74 (1/2) – 2005
pies as in station 1, which were deposited in ZMA 123.637
(Campoma guppy).
13-07-2002. Station 3: Same irrigation ditch as locality 2 at
the outskirts of Cariaco. Same guppies as in stations 1 and 2
(Campoma guppy). No guppies were deposited.
13-07-2002. Station 4: Bridge over wide stream next to road
from Cariaco to Chocopata, ca. 3 km from Cariaco. Same guppies
as in stations 1-3, deposited in ZMA 123.638 (Campoma guppy).
Also sh of the genus Rivulus was collected, deposited in ZMA
123.639.
13-07-2002. Station 5: Small stream at Heullo de Cariaco,
small village at the Golfo de Cariaco. Common guppies, viz.,
melanophores in spots, little trace of metallic sheen, deposited
in ZMA 123.640 (Common guppy). In this small biotope, a
remarkable collection of mollies (Poecilia koperi Poeser, 2002;
ZMA 123.6341), Rivulus sp. (ZMA 123.642), Gobiidae gen. sp.
indet. (ZMA 123.643), Poecilia picta (ZMA 123.644) and a
shrimp was made.
13-07-2002. Station 6: Waterfall El Cordon, ca. 5 km west of
Cariaco. Large common guppies, males ca. 3 cm TL, females
4-5 cm TL. Variegated colour patterns, with large metallic, silvery
blotches. Also a cichlid predator was collected. No specimens
were deposited.
14-07-2002. Station 7: Río de Oro, at the entrance of Aguas
de Moises. Small river with many bends. Guppies as in stations
1-4 (Campoma guppy), together with characid sh, Crenicichla
sp., Cichlasoma sp., a rivulid sh, and crabs. No specimens were
deposited.
14-07-2002. Station 8: Junction of Río Catuaro and Río Santa
Lucia, tributary to Río San Juan, tributary to Río Orinoco, ca. 10
km south of Casanay. Common guppies, chromatophores in spots,
no extensive metallic sheen. No specimens were deposited.
14-07-2002. Station 9: Río Tonoro (= Río Domingo), tributary
to Río San Juan, tributary to Río Orinoco, ca. 20 km south of
Casanay. Guppies like in locality 8 (Common guppy). No speci-
mens were deposited.
14-07-2002. Station 10: Río Grande, at Río Grande, tributary
to Río San Juan, tributary to Río Orinoco, ca. 25 km south of
Casanay. Guppies like in stations 8 and 9 (Common guppy). No
specimens were deposited.
15-07-2002. Station 11: Small ditches, parallel to the road
from Cariaco to Carúpano, ca. 3 km west of Carúpano. Guppies
with a mix of characters, i.e., melanophores concentrated in spots,
with other colours brightly metallic (as in stations 1-4, 7). These
sh were syntopic with a sailn molly, identied as Poecilia
latipinna (LeSueur, 1821). The guppies were deposited in ZMA
123.645 (Campoma guppy), the sailns in ZMA 123.646.
15-07-2002. Station 12: Laguna north of the road from Cariaco
to Carúpano (= seaside), ca. 2.5 km west of Carúpano. Guppies
like in station 11, deposited in ZMA 123.647 (Campoma guppy).
15-07-2002. Station 13: Laguna south of the road from Cari-
aco to Carúpano (= land side), ca. 2 km west of Carúpano. Gup-
pies like in stations 11 and 12 (Campoma guppy), but much less
colourful. No specimens were deposited.
15-07-2002. Station 14: Río El Pilar, bridge on the road from
Carúpano to El Pilar, ca. 15 km southeast from Carúpano. Common
guppies, like in stations 8-10. No specimens were deposited.
15-07-2002. Station 15: Río Casanay, at Guarapiche. Guppies
as in stations 11 and 12 were deposited in ZMA 123.648 (Cam-
poma guppy).
16-07-2002. Behavioural studies at station 7. Nearby, in a
stream close to the road from Cariaco to Casanay, ca. 4 km from
Casanay, we discovered another stream containing guppies like
in stations 1-4, 7 (Campoma guppy).
17-07-2002. Several locations north of the Golfo de Cariaco.
These locations are not noted as stations, they did not yield any
guppies. One molly, viz., Poecilia koperi, was collected in La-
guna de los Patos and deposited in ZMA 123.649.
18-07-2002. Behavioural studies at stations 14. Guppies like
in stations 1-4, 7 (Campoma guppy), near the Posada Elvira, on
the road from Cariaco to Casanay, ca. 2 km from Cariaco.
Systematic section
Poecilia (Acanthophacelus) wingei n. sp.
Type material. Holotype. ZMA 123.704 (male), Venezuela, Esta-
do Sucre, Paría Peninsula, ca. 3 km north of Cariaco, bridge on the
road from Cariaco to Carúpano, division between Laguna Buena
Vista and Laguna Campoma, coll. F.N. Poeser, M. Kempkes, and
L. Palacio, 13-VII-2002. Allotype. ZMA 123.705, (female) same
data as ZMA 123.704. Paratypes. ZMA 123.636 (18), same data
as ZMA 123.704; ZMA 123.637 (67), Venezuela, Paría Peninsula,
irrigation ditch ca. 50 meters from the road from Cariaco to
Carúpano, coll. F.N. Poeser, M. Kempkes, and L. Palacio, 13-VII-
2002; ZMA 123.638 (25), Venezuela, Paría Peninsula, ca. 3 km
from Cariaco, bridge over wide stream beside road from Cariaco
to Chocopata, coll. F.N. Poeser, M. Kempkes, and L. Palacio, 13-
VII-2002; ZMA 123.645 (7), Venezuela, Paría Peninsula, ca. 3 km
west of Carúpano, small ditches, parallel to the road from Cariaco
to Carúpano, coll. F.N. Poeser, M. Kempkes, and L. Palacio, 15-
VII-2002; ZMA 123.647 (38), Venezuela, Paría Peninsula, ca. 2,5
km west of Carúpano, laguna north of the road from Cariaco to
Carúpano (= sea side), coll. F.N. Poeser, M. Kempkes, and L.
Palacio, 15-VII-2002; ZMA 123.648 (10), Venezuela, Paría Pe-
ninsula, Río Casanay, at Guarapiche, coll. F.N. Poeser, M. Kemp-
kes, and L. Palacio, 15-VII-2002; UMMZ 158715 (30 paratypes
of 585), Venezuela, Paría Peninsula, stream 35 km west of Carú-
pano, 28-03-1939, coll. F.F. Bond.
Description of the types. The holotype is a male, 14.3
mm SL, with a black band at the most anterior part of
the caudal peduncle (cf. Fig. 3a, b). On the right side
of the caudal peduncle are two elongate and intercon-
nected metallic spots. Body scales contain many
melanophores, darkening the body and forming a
vague spot between the ventral and pectoral ns. The
pigmentation on the left side is like the right side, but
with the vague spot elongate as a band, bordering a
metallic body colour. This male has a denite caudal
n pattern, with a slightly enlarged and dark upper
margin. The upper and central section shows small
black spots, whereas the lower margin is pigmented
at the base, i.e., a thin black line on the lower margin
104 F.N. Poeser et al. – A new guppy from Venezuela
of the n and a white area around a black spot just
above this margin. The gonopodium contains remark-
ably many melanophores, also extending to the go-
nopodial palp.
All other diagnostic features are as mentioned in
the characterisation of the subgenus.
The allotype is a slender female, 14.1 mm SL. The
unpaired ns and the body with many melanophores,
whereas melanophores are concentrated along the
margins of the scales. No further specic markings
are present. Females of P. wingei are indistinguish-
able from those of P. reticulata, with the possible ex-
ception of a slight metallic sheen in life specimens.
The paratopotypes are 12 females and juveniles
without specic pigmentation, 4 subadult males with
one or two metallic spots on the body and caudal pe-
duncle. These males have a caudal stripe and a basal
spot on the caudal peduncle. The two remaining
males show more developed markings. The smaller
of these two has an anal black spot and black hori-
zontal bands on body and caudal peduncle, border-
ing a lighter, mostly metallic area. The caudal n pig-
mentation is more or less an extension of the body
pigmentation, i.e., a central lighter area bordered by
a thin black line. The larger of the two adult males is
coloured like the holotype, but with a thinner blotch
halfway the body. This band splits the anterior of the
two metallic spots. The anterior part of the body is
very dark.
One female has a thin lateral line. Males do not
have spots, but blotches from the origin of the go-
nopodium to under the dorsal n (Figs. 4a-d). Ante-
rior side of the body sometimes has broken spots, or
little horizontal lines. One or two bands over the
caudal peduncle, caudal peduncle with a brighter area
or a bright spot. The gonopodial palp on ray 3 exceeds
the tip, but is seemingly smaller in this population
than in others.
The gonopodia from two of the males (Fig. 3b-c)
are also gured (Fig. 6).
Diagnosis. Poecilia wingei is a small species, with
males measuring up to about 15 mm SL and females
up to 20 mm SL, and with a distinct sexual dimor-
phism, with polychromatic males and uniform females.
Meristic data. A. 9; D. 6-7; C. 12-14; LLS. 25-27; CPS.
14. The gonopodium has a retrorse hook on ray 5, but
not on ray 3. The eshy palp on ray 3 is extending
beyond the tip. The ventral spines on ray 3 are large,
resulting in a bulb-like appearance of the gonopodial
tip. The dorsal bulbs on ray 4a are much smaller than
the spiny dorsal extensions on ray 4p.
Comparison. Poecilia wingei differs from most spe-
cies of Poecilia by the following characters. The go-
nopodial palp extends clearly beyond the tip of the
gonopodium. The gonopodium also lacks a terminal
hook at ray 3, present in the short nned and sailn
mollies and in Psychropoecilia. With the exception
of Micropoecilia, none of the subgroups of Poecilia,
viz., the P. sphenops species group, the P. latipinna
species group, the P. caucana species group, Cur-
tipenis and Psychropoecilia, have polychromatic
males. It also differs from most species groups in the
number of scales around the caudal peduncle, i.e., 14
vs. 16 or more, sharing this characteristic only with
the P. caucana species group and Micropoecilia. Fe-
males are further distinguished from all other species
groups by the unique combination of nine anal n
rays and less than eight dorsal n rays.
Its closest relative is the common guppy, P. re-
ticulata, sharing identical meristic data, but differing
by its enhanced metallic body pigmentation. This
brightness in body pigmentation is also noticed in
the females of P. wingei. Moreover, in the zone ad-
jacent to the distribution area of the common guppy,
P. wingei males exhibit a unique melanophore pat-
tern, viz., a large band in the midsection of its body.
The importance of this feature, i.e., the spatial distri-
bution of melanophore patterns, is decisive for its
recognition as a valid species (see below).
UMMZ 158715 was examined and compared to
the common guppy (see above). Average length fe-
males (5; 26.9-30.4): 29.0 mm SL, males (5; 18.8-
21.3): 19.6 mm SL. Average least depth caudal
peduncle females (5; 10.4-13.4): 11.5, males (5;
10.6-13.2): 11.6.
Distribution. The Campoma guppy occurs in fresh
waters in the Campoma- and Buena Vista Lagoons,
unto Carúpano.
Etymology. We dedicate this species to Dr Øjvind
Winge, born in May 1886, died in April 1964. He was
head of Department of Physiology, Carlsberg labora-
tory (1933-1956), and was not only the father of
“genetic engineering” (Szybalskia, 2001), but also
described many colour patterns and the genetics of
105
Contributions to Zoology, 74 (1/2) – 2005
sex-determination in the common guppy as a result of
extensive breeding experiments. His work provided
the basis for understanding the colour polymorphism
in guppies in general (answering to the question posed
by Houde [1997: 5]) and the means for recognition of
character displacement between the common guppy
and the Campoma guppy in particular.
Hypothetical origin.The area of distribution of this
new species invites speculation to its origin. It is
separated from the Orinoco River and the common
guppy by the Cordilleras de la Costa, it is, therefore,
obvious to propose that the upraise of these mountains
separated the shared ancestor of both species. The
Campoma guppy is restricted to the coastal side of
the Cordilleras, whereas the common guppy initially
occupied the landside, or Orinoco side, of Venezue-
la. The common guppy somehow re-migrated to the
coastal areas, maybe through areas near present-day
Barcelona, where rivers with a Caribbean drainage
originate next to rivers of the Orinoco drainage. The
Campoma guppy managed to keep its specic integ-
rity in the base of Paría through sexual selection based
on character displacement in male body coloration
(cf. Houde, 1997). Differences in male coloration
might be caused by accumulation (or mutation) of
genes responsible for iridocytes, i.e., the metallic
colours, accentuated by the black midsection band
on the males in the probable zone of sympatry, the
Campoma region. Differentiation in pigmentation
within the type series suggests that the midsection
band develops later in life.
Characterisation of the genus Poecilia
The genus Poecilia was dened by three synapomor-
phies (Poeser, 2003: 119, 137): (i) Serrated serrae
subdistally on gonopodial ray 3; (ii) A protruding
hook, distally on gonopodial ray 5p; (iii) Hook found
distally on gonopodial ray 3. The phylogenetic
analysis on which these conclusions were based,
compared the genera of the Poeciliini (sensu Rosen
and Bailey, 1963) with Brachyrhaphis Regan, 1913,
lumping all subgroups of Poecilia in an extensive
polytomy. This prompted Poeser (2003, chapter 9)
to consider the P. reticulata species group phyloge-
netically indenable as a subgenus. A re-examination
of eleven characters strictly applicable to Poecilia
was made, this time including all species (cf. Poeser,
2003, chapter 11). To root the tree, Alfaro cultratus
(Regan, 1908) was chosen as outgroup.
The analysis rendered a surprising arrangement of
taxa. At the base, the three Hispaniolan species of
Poecilia are the sistergroup of all remaining species.
These species are classied according to the available
subgeneric names, rendering them P. (Curtipenis)
elegans (Trewavas, 1948), P. (Psychropoecilia)
dominicensis (Evermann and Clark, 1906) and P.
(Psychropoecilia) hispaniolana Rivas, 1978. Much
to our surprise, also P. (Acanthophacelus) reticulata
was allocated at the base, separating P. reticulata
from its South American congeners. The species
described herein as new shares these characters.
Therefore, we conclude that the phyletically closest
relatives are not among the geographically closest
relatives. Since both species of guppies are less re-
lated to P. vivipara Bloch and Schneider, 1801 and
to P. parae, we re-instate subgeneric distinctions in
Poecilia to emphasise this distinction. In the clado-
gram (Poeser, unpublished), Acanthophacelus is
closest related to the Central American mollies, as is
Poecilia (Poecilia) vivipara, but they are not each
other closest relatives (conrming the phylogenetic
relationships within Poecilia based on mitochon-
drial DNA [Ptacek and Breden, 1998]). Micropoecil-
ia spp. are closely related to P. vivipara.
Characterisation of the subgenus Acanthophacelus
The allocations in species groups are largely prompt-
ed by the gonopodial features and number of dorsal
n rays (Table IV). These differences are geoclinal
for the species of the subgenus Poecilia (cf. Poeser,
2003, chapter 5), combining the P. vivipara species
group with the P. latipinna and the P. sphenops spe-
cies groups. The remaining species groups are iden-
tiable by these characters, yielding the following
characterisation of Acanthophacelus:
Meristic data. A. 9; D. 6-7; C. 12-14; LLS. 25-27;
CPS. 14.
Both species are small with marked sexual dimor-
phism in size and in pigmentation. Mature males are
about 14 to 21 mm SL, females are about 20 to 38 mm
SL. Females are relatively more slender, least depth
of the caudal peduncle averaging from 10 to 16% SL,
whereas males are slightly more stout, from 10 to 18%
SL. Females are generally without markings, except
for a reticulate pattern of melanophores along the
scales margins. Males always have black markings in
F.N. Poeser et al. – A new guppy from Venezuela
106
their variegated colour pattern. While females have no
diagnostic markings on their silver grey to brownish
bodies, males have patterns that are hard to describe.
Within a population no male is exactly the same as
another. Although this is largely true for male speci-
mens within one population, some generalities are
apparent between populations:
Poecilia reticulata: Sometimes a population consists
of males with only one spot, e.g., UMMZ 158730
and UMMZ 158753. Most populations contain males
that are multi-spotted, e.g., UMMZ 158704, UMMZ
158706, UMMZ 158711, UMMZ 158717, UMMZ
158720, and UMMZ 158723 (Figs. 1, 2, 5). Occasion-
ally, males exhibit a horizontal line on the body,
and/or on their caudal peduncle. A vertical elongated
band may be present on the caudal peduncle, e.g.,
UMMZ 158736 (Fig. 4).
Poecilia wingei: A large vertical band is sometimes
present in the midsection of the body, at the anterior
part of the caudal peduncle, e.g., UMMZ 158715 (Fig.
3). Populations near Carúpano contain males that are
multi-spotted, like in P. reticulata.
These patterns concern preserved melanophores. Life
colours also include differently coloured spots, e.g.,
red, orange, yellow, blue, green and blue, dull or iri-
descent spots and patches, whereas also red horizontal
lines are not uncommon. Some individuals exhibit
only a single colour, e.g., red or green, whereas most
individuals have colour patterns combining spots and
patches of different colours. Also the unpaired ns,
except the gonopodium, can have colours, sometimes
with an elongation of the upper or lower caudal n
lobe. We are not aware of any study that correlates
specic colour patterns to particular areas, although
several studies have shown correlation between colour
patterns and ecological factors, e.g., predation pres-
sures (Reznick, 1982, 1983; Reznick et al., 1996a,
1996b, 1996c; Albers, 2000), and food regimes (Rob-
inson and Wilson, 1995). A recent study (Alexander
and Breden, 2004) that shows this correlation is dis-
cussed below. The genetic basis of these spots has been
investigated since Schmidt (1920) and Winge
(1922).
The occurrence of a wide polymorphy of melano-
phore patterns conicts with the genetic mechanism
proposed by Winge (1922, 1927; Fig. 7) which only
allows for a moderate amount of variation in these
patterns. However, other genetic mechanisms have been
proposed (Turing, 1952; Murray, 1988) explaining the
variation in the observed patterns. One characteristic
of this alternative explanation explains why bodies are
never striped when the caudal is spotted, which is true
for all guppies examined. It also explains the differ-
ences in spot size, e.g., correlated with predation
regiments. The genetics of this mechanism is highly
speculative, and therefore not discussed herein.
Comparison of the colour patterns between the sub-
genera Acanthophacelus and Micropoecilia
Generally, some similarities between melanophore
patterns in the guppy and in the species of Micropo-
ecilia (cf. Meyer, 1993) are observed. A number of
guppy populations have colour patterns like those
found in the species of Micropoecilia. A spot on the
caudal base combined with a pigmented upper margin
of the caudal n, found in many populations of Mi-
cropoecilia, was named P. reticulata 'Lineatus' (cf.
Winge, 1927). The large red and black spots are present
in nearly all guppy populations, as well as in P. parae
'amazonica' and P. picta (Poeser, 2003, chapter 11).
Fig. 6a-c (Photographs by J. van Arkel), detailed photographs of
gonopodia. (a) Poecilia reticulata, gonopodium of specimen in Fig.
1a; (b, c) Poecilia wingei, gonopodia of specimens in Figs. 3b, c.
a
c
b
107
Contributions to Zoology, 74 (1/2) – 2005
There can be a bright area on the caudal peduncle,
margined by heavily pigmented bars (Figs. 3-4), like
in P. parae 'melanzona' (Fig. 8). Two explanations are
possible. If these patterns occur sympatrically, it might
be that these reect better shoaling possibilities or
adaptations to similar environmental demands. If these
patterns do not co-occur, a special form of character
displacement (cf. Poeser, 1995, 1998; present paper)
might be present. Examination of these patterns, both
genetically and bio-geographically, e.g., correlated to
mimicry or character displacement, therefore, is a
promising eld of investigation, possibly explaining
the occurrence of these patterns in wild guppies and
in Micropoecilia.
Gonopodia. The gonopodium of both species of gup-
pies (cf. Meyer, 1993; Fig. 6) has a retrorse hook on
ray 5, but not on ray 3. The eshy palp is much elon-
gated, extending well beyond the tip. The ventral
spines on the 3rd gonopodial ray are large, giving the
gonopodium a bulb-like appearance. Both ray 4a and
ray 4p have dorsal spines, the spines on 4a are much
smaller than on 4p.
Habitat. Guppies are known to inhabit a wide vari-
Fig. 8. Habitus of Poecilia parae 'melanzona'. The pigmented
bars on the caudal peduncle resemble the phenotypes of guppies
shown in Figures 3c and 4c.
Table IV. Characterisation of the subgenera in Poecilia.
Poecilia vivipara represents Poecilia; P. sphenops and P. latipinna represent Mollienesia; P. dominicensis represents Psychtopoecilia;
P. elegans represents Curtipenis; P. parae represents Micropoecilia; P. reticulata represents Acanthophacelus; P. caucana represents
Allopoecilia.
Species group Serrae on ray 3 Hook on ray 3 Hook on ray 5p Number of dorsal n rays
Poecilia sphenops + + + 8-11
Poecilia latipinna + + + 12 or more1
Poecilia dominicensis - + + 8-11
Poecilia elegans + - + 8-11
Poecilia vivipara + -2 -3 7-11
Poecilia parae + -4 -5 8
Poecilia reticulata + - + 6-7
Poecilia caucana + +/-6 + 8
1 Poecilia latipunctata seems a noticable exception to this. Molecular data clearly indicate close relations with the P. latipinna species
group (12-20 dorsal n rays), although it does not have more than 11 dorsal n rays (cf. Poeser, 2003).
2 This character is “reduced to absent” in this species group (cf. Poeser, 2003).
3 This character is “reduced to absent” in this species group (cf. Poeser, 2003).
4 This character is “reduced to absent” in this species group (cf. Poeser, 2003).
5 This character is “reduced to absent” in this species group (cf. Poeser, 2003).
6 Frequently much reduced in this species group (cf. Poeser, 2003)
Fig. 7. Drawings of guppy phenotypes (after Westerhof, 1960) Ferriguneus, Auratus, Armatus and Variatus represent ‘multi-spotted’
males with different caudal n patterns, Lineatus only indicate the caudal n pattern.
108 F.N. Poeser et al. – A new guppy from Venezuela
ety of habitats with temperatures ranging from 18-
30°C. They are not adapted to special biotopes but do
not occur in purely marine environments.
Distribution area. The common guppy is presently
circumtropical. However, it is believed to occur natu-
rally only in the northeastern part of South America
and on the Lesser Antilles (Fig. 9). We consider only
the mainland areas of Venezuela, east of Lago de
Maracaibo, further east to the Guiana’s, the adjacent
part of Brazil, Para district, and upstream the Amazon
river (Rio Solimões) as its natural area of distribution.
Based on data presented in the discussion section, the
occurrence on Trinidad is undisputed. The island of
Barbados (cf. Boulenger, 1912) is not considered part
of its natural range (Poeser and Isbrücker, 2002).
Spatial distribution of the phenotypes of the freshly
collected guppies
Life colours of the Orinoco variety of P. reticulata are
characterised by one or two black spots, often en-
hanced by a metallic patch. This morph is named for
the guppies from stations 8-10, which eventually are
tributaries of the Río Orinoco. Guppies from stations
5 and 14 are similar to these phenotypes and are there-
fore also classied as Orinoco guppies. These speci-
mens are like most guppies (P. reticulata) examined
from the ZMA and UMMZ material.
Guppies caught below the El Cordon waterfall
(El Cordon variety, station 6) differ from the Ori-
noco variety. Not only are they substantially bigger,
body colorations are different too. Two specimens
(of ve collected males) had enormous 'peacock'
caudal ns, i.e., a large black spot margined by a
bright metallic, silvery blotch. One specimen was
without any specic colours, except for exhibiting
a metallic silver body. This latter specimen also had
an enormous gonopodium, extending almost to the
caudal base. The possible origin of the characteris-
tics of this variation is discussed below.
In the areas of stations 11-13, 15 (Carúpano re-
gion), the guppies exhibited similar colour patterns
as the Orinoco variety, with the addition of metallic
polychromatic patterns. Where in the Orinoco gup-
pies the metallic sheen is restricted to areas around
the black spots, in P. wingei all colours are a brilliant
array of metallic colours: red, blue and green to yel-
low (= gold). Dorsal n is usually transparent, how-
ever, sometimes black or white markings are present.
Caudal n variegated, i.e., with red or black upper,
lower or both margins, or a brownish blotch extend-
ing from the caudal base. Females are greyish, with
a bright sheen over their bodies.
In the area around stations 1-4, 7 (Campoma re-
gion), guppies are found with colours like the
Carúpano variety, with addition of a characteristic
black midsection of the body. Individual specimens
exhibit a black anterior part of the gonopodium, or a
ne striped pattern on the anterior part of body, or a
completely green body.
Evidence of character displacement
Character displacement is observed “when the areas
of distribution of two species of animals overlap
geographically, the differences between them are
accentuated in the zone of sympatry and weakened
or lost entirely in the parts of their ranges outside this
zone (Brown and Wilson, 1956)”. Summarising the
spatial distribution of the phenotypes, the least vari-
egated guppies (hereafter named type A) are the El
Cordon variety, followed by the Orinoco variety that
has black spots and some metallic fringes (type B).
The Carúpano morph also has black spots, but is
brilliantly metallic (type C), whereas the most exten-
sive black and metallic colouring (type D) is found
in the Campoma variety.
When plotted on a map (Fig. 10), the distribution
of phenotypes shows coherence. The brilliant guppies
occur exclusively in the Cariaco-Carúpano region, a
valley of marshlands surrounded from south to east
by the Cordilleras de la Costa. The northern parts of
this region are enclosed by the dry Araya Peninsula
and in the west it is bordered by the Golfo de Cariaco.
It is from this latter area that P. reticulata can enter
this territory. In the western part of this region, where
P. wingei and P. reticulata may co-occur, both popu-
lations have their most extreme phenotypes, i.e., the
Campoma variety of the Campoma guppy and the El
Cordon variety of the Orinoco, or common, guppy.
In the areas in which these two kinds of guppies do
not co-occur the phenotypes are similar: the Carúpano
variety of the Campoma guppy seems merely a bril-
liant version of the Orinoco guppy. Both phenotypes
are here characterised by the typical black spots. So,
while conspecic genetic variation supposedly would
follow an A-B-C-D pattern in colour-type distribu-
tion, observed distribution is actually B-A-D-C. This
109
Contributions to Zoology, 74 (1/2) – 2005
represents a clear example of character displacement,
leaving little doubt that the Campoma guppy indeed
represents a new species.
Character displacement as a pre-copulatory isola-
tion mechanism has already been suggested to have
mediated in the speciation of Hispaniolan short n
mollies (Rivas, 1982), short n mollies from El Sal-
vador (Poeser, 1995) and sailn mollies (Gabor and
Ryan, 2001). The example in guppies is new to sci-
ence and its characteristics, i.e., easily accessible
populations of a species already thoroughly studied
and clearly limited geographical dimensions, makes
it an almost perfect subject for further studies. The
occurrence of this type of speciation was already
predicted (Houde, 1997: 55): “… a distinctive gup-
py population from Venezuela that, at least in labo-
ratory stocks, appears to be monomorphic, but is in-
terfertile with Trinidad guppy populations … A spe-
cies-recognition function for colour patterns thus
cannot be ruled out…”
The case study of Alexander and Breden (2004),
examining this phenomenon, is fully discussed be-
low.
While there is little doubt about the distribution
of phenotypes of P. wingei, the origin of the El Cor-
don phenotype (= P. reticulata), based on only a small
collection at one locality, remains the subject of
further study. This phenotype can have arisen either
by character displacement or by hybridisation.
Behavioural differences between Poecilia reticu-
lata and P. wingei
Behaviour of the Common guppy
Guppies differ from all other species of Poecilia in
their mating behaviour (Liley, 1966). Behavioural
patterns herein described are personal observations,
obtained from the eld trip to Venezuela described
herein, as well as from an earlier eld trip (Kempkes,
in 1993) and from aquarium observations. Our stud-
ies agree with the observations of other authors on
P. reticulata (cf. Houde, 1997). Courtship behaviour
in P. reticulata proceeds in strict sequential ordered
elements (Table III). However, some variation in
behaviour was observed when behavioural elements
were occasionally skipped by courting males.
In the El Pilar river (Fig. 10) two isolated popula-
tions of P. reticulata were observed (15-07-2002, from
13.00 to 13.30 hr, and 18-07-2002 from 9.40 to 10.10
hr). Both populations occupied about a half square
meter and the water was 10 - 20 cm deep, and both
microhabitats were located near the riverbanks. No
sh were observed to enter the fast owing waters in
the midsections of the river. Therefore, the population
was found to be relatively constant, with no individu-
als leaving or entering the community. The action
radius of males within the population appeared to be
bigger than that of females. In these groups, being
‘local populations’, some territorial behaviour was
observed in which the largest females maintained a
bigger individual distance to other larger females.
These females reacted to approaches of other females
by biting them and chasing them away. Juvenile gup-
pies predominantly inhabited the banks of the river,
close to the main population. We think that in the shal-
low water, young guppies were relatively save from
predators. Sexually mature males were continuously
near the females of their group. The intra-sexual com-
petition between the males was especially perceptible
during the courtship: males constantly tried to disturb
the courtship of other males, after which they at-
tempted to court the same female. Female choice is
apparent (cf. Houde, 1997), because of energy costs
of internal maturation of eggs and the additional costs
of swimming around highly pregnant. Furthermore,
the supply insemination adds to what the males invest
to the reproduction. In a laboratory test, Nicoletto
(1993) described that female guppies favour large
guppy males. The orange and iridescent colour areas
at the bodies of the males are probably indications
of the tness of males. Female choice, combined
with the establishment of local populations, should
favour genetic drift. When populations remain at
more or less the same geographical position, these
populations will adapt to the specic requirements
of these localities. It has been postulated that sneak
copulation alone counteracts possible genetic drift
(Magurran, 1998).
Behaviour of the Campoma guppy
The behaviour of P. wingei is new to science and is
therefore extensively described. Observations were
made on the 14th, 16th and 18th of July at Las Aguas
de Moises, near the highway between Cariaco and
Casanay. The Campoma guppies were observed in a
110 F.N. Poeser et al. – A new guppy from Venezuela
small stream of clear water. The river was about 150
cm deep with the bottom clearly visible, about 180 to
300 cm wide and no submerged vegetation was ob-
served, the observation area was a stretch of about 40
meter. There were trees and some dense vegetation at
several positions at the bank. The Campoma guppy
was syntopic with the following species: Catoprion
spec., Crenicichla spec., Cichlasoma spec., Rivulus
spec., Ampullaria spec., and some unidentied crus-
taceans. The observed subpopulations, i.e., groups of
about twenty adults of both sexes and about 20 sub-
adult and juvenile shes, occupied positions near the
river banks. At different locations of this stream we
observed other groups with approximately 60 adults,
with about 50 subadults and juvenile shes. The fe-
males were continuously grazing the loamy ground,
swimming in groups of 6 to 10 shes. In this stable
environment, subgroups are formed consisting only
of females. Stable all-female subgroups stay a rela-
tively long time, about 20 to 30 minutes, at a certain
locality within the territory, circa 100 square cm, where
they seem to be constantly foraging. Strange females
rarely enter this sub-community. Intra-sexual aggres-
sion, in which the largest females chase smaller fe-
males away, was also observed. When two females
are equally big, they show a particular posture. These
female opposed each other in a T-shaped position for
half a second, spreading their ns. After this showing
off they feed again. Only at one observation we saw
two opponent females actually ght, one female was
picking at the caudal peduncle of the other female.
After about ten seconds they started feeding again.
This behaviour had no apparent effect on the group
structure. Two explanations for this behaviour are
possible. Firstly, the females have some territorial
behaviour related to foraging. Secondly, we suspect
female-to-female competition related to sexual selec-
tion, i.e., male sexual selection not observed in P. re-
ticulata. This latter conclusion is supported by further
observations (see below).
During the observation period, males were con-
tinuously found in the proximity of the females. With
every all-female subgroup, we observed how two or
three males remained proximate, i.e., the males seemed
less promiscuous compared to P. reticulata. In one
case, such group was accompanied by a single male.
In groups with more shes, i.e., more than 40 adults,
the males court more intensive. In relatively small
groups, e.g., one to three males and one to seven fe-
males, the males show less intensive courtship behav-
iour compared to bigger groups. Males in smaller
groups apparently need not show their tness so often,
because the females probably recognise individual
males. When a male not already belonging to the
subgroup started to show courtship behaviour to one
of the females, the male already present showed the
strange male his sigmoid display, after which the in-
truder stopped his courtship attempt. Males that initi-
ate courtship approach the female from behind, prefer-
ably while she is busy searching food. As noted above,
females are mostly stationary during foraging.
Observations indicated that the introduction phase
in the Campoma guppy is relatively short compared
to common guppies, whereas the last phase of their
courtship behaviour, i.e., high courtship, is longer
and more intensive. Correlated to the behavioural
characteristics of P. reticulata (Table III), we record-
ed the following.
1a. A male attempts to position himself under the
female, while his colour pattern darkens if he
is successful.
2a. The male will attempt to become visible in the
face-to-face position.
2b. He initiates the sigmoid-display. During the
sigmoid-display the male turns both sides to the
female, like the common guppy.
2c. Finally the male circles around the female and
attempts to copulate with his forwardly turned
gonopodium.
The signicant differences between the courtship
behaviour of the common guppy and the Campoma
guppy, documented from about 35 observational mo-
ments, is summarised as follows.
• The Campoma guppy males court more from below
during the rst phase of the courtship behaviour.
The males from the common guppy swim more
from behind or from the side of the females.
• Because the female of the Campoma guppy did
not ee, we did not observe any chasing from the
males. The females oat slowly in front of the
males and show that they are co-operative. Fe-
males of the common guppy make clear, in a
relatively early stage of courtship, whether they
are inclined to copulate or not.
• The characteristic display jump of the common
guppy during the courtship is only rarely observed
in the Campoma guppy. We, therefore, assume that
the display jump is not important for the courtship
behaviour of the Campoma guppy.
111
Contributions to Zoology, 74 (1/2) – 2005
• The females of the Campoma guppy swim away
from the courting males when they attempt to
copulate, i.e., they avoid courting males relatively
late. The females from P. reticulata often ee at
the beginning of the courtship. We never saw any
aggression from the Campoma guppy females
towards courting males. Under natural conditions,
as well as in an aquarium, females of the common
guppy attack obtrusive males. In general, the fe-
males of the Campoma guppy seem longer co-op-
erative and receptive, rejecting males in a much
later stage than do females of the common guppy.
• The courtship of the Campoma guppy male is
relatively docile. Moreover, males attempt to court
the same female longer and, after failure, often
start to court the same female again.
• Although the competition is strong in big popula-
tions, we never saw a Campoma male attempting
to court a female that is already courted by an-
other male. The males in a big group show im-
pressing behaviour only on occasion. We assume
that inter-male aggression is not relevant for court-
ship in a big group.
• We observed sneak-copulation, i.e., copulation
attempts without elaborate courtship, in the Cam-
poma guppy too, but it is another kind of sneak-
copulation, i.e., we never observed sneaky copula-
tions without previous courtship. Common gup-
pies will try to copulate without any prior courtship
(cf. Houde, 1997).
Despite intensive observations no particular preda-
tor was found that directly attacked guppies. For ex-
ample, in all locations, juvenile and subadult Cam-
poma guppies were close to the Rivulus-like sh near
the riverbanks. We did not nd any Rivulus attacking
any young guppies. At Las Aguas des Moises, adult
guppies were found near a pair of Crenicichla with
juveniles. No Crenicichla did pursue guppies, and in
one occasion a subadult Crenicichla was seen to actu-
ally ‘pass through’ a shoal of guppies. No reason for
this ‘non-aggression pact’ is apparent.
Remarks on population differentiation in Trini-
dadian guppies
In a paper on guppy population differentiation, Ma-
gurran (1998) posed an enigmatic dilemma: “Guppy
populations evolve rapidly … The rates of evolution
involved can be up to seven orders of magnitude
greater than those seen in the fossil record … female
choice appear to reinforce the divergence …[howev-
er] perplexingly … there is no reproductive isolation
… between populations …” A substantial counterac-
tion against speciation seemed to be 'sneaky mating',
i.e., the mechanism enabling males to copulate with-
out the consent of the females. Also different forag-
ing niches for males and females is supposed to in-
hibit the development of feeding polymorphism, and
therefore inhibit sympatric speciation (Magurran,
1998). This enigma was generated by the occurrence
of genetic differentiation between guppies from the
Quare-Orupuche drainage versus guppies from the
Paría drainage on Trinidad (Fajen and Breden, 1992;
Taylor and Breden, 2000). The guppies from the
Trinidadian Paría drainage are closely related to Ven-
ezuelan and Guyanan mainland populations, whereas
the Quare-Orupuche guppies form a separate mono-
phyletic clade. Our discovery of the Campoma guppy
gives the derivation of the Quare-Orupuche guppies a
possibly different perspective.
Trinidadian guppies interbreed readily in labora-
tory circumstances with Campoma guppies (Houde,
1997; pers. obs.), although this seemed not to be the
case in the late 1970’s (see below in our discussion
on the Endler’s Live-bearer; Alexander and Breden,
2004). The geological environment, protecting the
Campoma region from invasion of the Orinoco
guppy, allows only a limited inux of foreign gup-
pies into the region. Here, separation of the two
guppy populations is secured by sexual selection,
and subsequent character displacement reinforcing
divergence (cf. Houde, 1997; Magurran, 1998).
However, it is not unlikely that, while there might
have been an initial separation on species level
present on Trinidad, the geology of Trinidad did not
allow for an enduring separation of these two spe-
cies. While the dispersal from river to river in the
Cumaná-Cariaco coastal strip limits the possibilities
for guppies to enter the Campoma region, the Ori-
noco river probably provides a continuous ‘bom-
bardment’ of heterospecic genetic material in the
original Trinidadian guppies. These latter guppies
might have been the same species as the Campoma
guppies, or a closely related population. Ongoing
introgression of Orinoco guppy genes has despeci-
ated the original Trinidadian guppy, causing the
genetic divergence observed today (Fajen and
112 F.N. Poeser et al. – A new guppy from Venezuela
Breden, 1992; Taylor and Breden, 2000). Therefore,
the initial situation postulated by Magurran (1998)
must be redened. While she initially situated a
single guppy species on Trinidad, diverging to the
limit of speciation but, 'perplexingly', not going
beyond that specic boundary, we postulate an-
other scenario. The present day situation results
from the introgression of genetic material of one
species, viz., the Orinoco or common guppy, into a
heterospecic population of ‘original’ Trinidadian
guppies, actually converging genetic differences,
i.e., despeciating the latter species. This despecia-
tion in action was also already recorded, though not
recognised as such. Magurran et. al. (1992) sampled
molecular data from a guppy population that was
supposed to be re-allocated by Haskins in 1957 from
the Caroni drainage, i.e., part of the Paría drainage
guppy populations, viz., Orinoco guppies, and re-
moved them to the Orupuche drainage. There these
Paría guppies replaced the residing guppies almost
totally, a striking example of obvious higher tness
of the common guppy.
Fig. 9. Distribution area of the subgenus Acanthophacelus. The occurrence of Common guppies in the Río Santo Domingo, near Merida
and the Río Apure, Llanos, Venezuela (both western rivers of the Río Orinoco drainage) is conrmed by Kempkes (pers. obs.). Guppies
claimed from the Rio Solimões are kept alive by Poeser. Occurrence of guppies upstream of the Río Orinoco and in the Rio Negro awaits
conrmation. Numbered locations indicate examined museum material (bold numbers are gured): 1. ZMA 120.725; 2. UMMZ 158750;
3. UMMZ 158736; 4. UMMZ 158753; 5. UMMZ 158709; 6. UMMZ 158729; 7. UMMZ 158704; 8. UMMZ 158705; 9. UMMZ 158715;
10. UMMZ 158706; 11. UMMZ 158716; 12. UMMZ 158715; 13. UMMZ 158711; 14. UMMZ 158720.
113
Contributions to Zoology, 74 (1/2) – 2005
Magurran (1998) dated the renewed contact of the
common guppy with the original Trinidadian guppy
very recent, i.e., 1,000-10,000 years ago, i.e., during
the period that Venezuela was linked to Trinidad by
the Cordilleras de la Costa. Molecular data (Fajen
and Breden, 1992) date the break up between these
two guppy populations long before this, i.e., 600,000
years ago. We, therefore, postulate that the Cordill-
eras were formed 600,000 years ago, blocking gene
ow between the several populations of guppies.
The Cordilleras also blocked the Río Orinoco from
its northerly ow, redirecting its course just south of
a rocky strip that is the present day northern range
on Trinidad. Since about 10,000 years ago, this
mountainous strip was severed from the Paría Pe-
ninsula and collected substantial strips of sand de-
posited by the Orinoco river, forming present-day
Trinidad. The common guppy established itself rm-
ly in the southeasterly drainages, whereas the Oru-
puche guppies retained some of their original ge-
netic content. On the Paría Peninsula, the Campoma
guppies have kept their specic identity since, where-
as the Trinidadian guppies have lost this identity by
the constant intrusion of Orinoco inuences.
This hypothesis is testable. Obviously, when the
molecular data of the Campoma guppy are compared
with those of the two clades found on Trinidad
(Taylor and Breden, 2000), we predict a closer re-
lationship of the Campoma guppy with the guppies
from the Orupuche drainage than to those from the
Paría drainage.
Remarks on the 'Endler’s Live-bearer'
The populations of Campoma-like guppies collected
in a coastal area of Venezuela, in Cumaná, Laguna
de los Patos (Endler, pers. comm.), might very well
Fig. 10. Map of the collection sites in and around the Paría Peninsula, Venezuela. Indicated are river systems, the Campoma- and Carúpano
regions, and the 250 m altitudes. Localities 1-4, and 7 rendered Campoma guppies, Campoma variety. Localities 11-13, and 15 rendered
Campoma guppies, Carúpano variety. All other, non-Paría localities, rendered Common guppies.
114 F.N. Poeser et al. – A new guppy from Venezuela
be an established local population of P. wingei. How
guppies were distributed there is unknown; they
might be remains of an earlier, wider range of the
Campoma guppy, but most likely they are released
aquarium specimens originating from the Cariaco-
Carúpano region.
The Endler’s live-bearer is renamed as “Cumaná
guppy” by Alexander and Breden (2004). This ver-
nacular name is misleading because not all guppies
from Cumaná they collected are named as such
(Alexander and Breden, 2004: 3), only those from
the west part of Cumaná. The paper of Alexander
and Breden (2004) adds a lot of quantitative data to
our ndings, conrming P. wingei as a valid species.
They quantied sexual isolation, adding sexual
selection and different male display traits to our
ndings of behavioural differences (Alexander and
Breden, 2004: 5). They also added clear morpho-
logical differences to the description of the two
guppy species (Alexander and Breden, 2004: 4, g.
6), as well as a full examination of the differences
in colour patterns also mentioned in the present
paper (Alexander and Breden, 2004: 3, 7, gs. 3-5).
Finally, they recorded only common guppies in a
radius of 100 km outside Cumaná, conrming the
presence of exclusively P. reticulata populations
between Cumaná and the Paría Peninsula. Alexan-
der and Breden (2004) recorded a total lack of re-
productive (postzygotic) isolation between their
Cumaná guppies and other guppy populations, al-
though they did mention the initial incompatibility
of Endler’s guppies with P. reticulata. This makes
sense in the light of our hypothesis of human intro-
duction: the initial isolation, existing in the late
1970’s, is now broken down, possibly by introduc-
tion of P. reticulata genes into the Cumaná popula-
tion of P. wingei by sneak copulations.
Acknowledgements
Our warm thanks extend to Mr. Luis Palacio, who drove us to
places we would not nd without him. We also thank Mr. Latsy
Nyari for loaning us his notes on the Endler’s Live-bearer. Mr.
Jan van Arkel provided us with the photographs of the specimens
shown in Figs. 1-6, and Fig. 8, for which we are grateful. Mr.
Pieter J. Michels kindly read an earlier version of the manuscript
and offered valuable suggestions.
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Received: 22 December 2004
Accepted: 8 September 2005
