Journal of Biogeography, 29, 16

Biogeography of the Amazon molly, Poecilia

formosa
Ingo Schlupp1,2*, Jakob Parzefall1 and Manfred Schartl3 1Zoologisches Institut und
Zoologisches Museum,Universitat Hamburg, Martin-Luther-King Platz 3, D-20146 Hamburg, Germany, 2Section of Integrative Biology, University of Texas at Austin, Austin, TX
78712, USA, and 3Physiologische Chemie I, Am Hubland, Theodor Boveri Institut,
Universitat Wurzburg, D - 97074 Wurzburg, Germany

Abstract
Aim The unisexual Amazon molly (Poecilia formosa) is a clonal, all-female sh but
depends on sperm of heterospecic males to trigger embryogenesis. Thus, one very
important factor shaping its geographical range is the presence of suitable host males.
Several species of the genus Poecilia from Central America, Mexico and the USA can
provide sperm in the laboratory, but are not utilized as hosts in nature. Consequently the
potential geographic range of the Amazon molly is much larger than the actual range.
This raises the question of what is responsible for the biogeographical range of the
Amazon molly?
Location Southern USA, Mexico and Central America.
Methods We review the current data available for the distribution of the Amazon
molly. We further tested whether salinity might hinder the dispersal of the species.
Results Amazon mollies tolerate marine conditions. We review the available data on
recent and human inuenced introductions of Amazon mollies.
Main conclusions We argue that male preferences are not responsible for the current
range. We propose that prevailing near-shore marine currents act as an effective barrier
against further dispersal. Furthermore, we discuss recent changes in the biogeography of
the Amazon molly.
Keywords
Poeciliidae, marine currents, salinity, dispersal, gynogenesis, unisexuality, Poecilia
latipinna, Poecilia mexicana.

INTRODUCTION
The study of biogeography has contributed greatly to our
understanding of evolution (Cox & Moore, 1993). Most
species distributions are shaped by current and historical
environmental factors, mainly the presence or absence of
crucial resources and limiting effects of physical barriers.
For sperm-dependent unisexual vertebrates (Vrijenhoek
et al., 1989) one especially crucial resource is the presence of
host males. All known unisexual shes and amphibians are
probably hybrids and reproduce by gynogenesis or hybridogenesis. Sperm is required to trigger embryonic development.
*Correspondence: Present address: Dr Ingo Schlupp, Zoologisches Institut,
Universitat Zurich, Winterthurerstrasse 190, CH-8057 Zurich, Switzerland.
E-mail: schlupp@zool.unizh.ch

2002 Blackwell Science Ltd

As these species have no males, this sperm must be obtained

from males of different species, a situation that has been
dubbed `sperm-parasitism' (Hubbs, 1964). In a mating
system like this the biogeographies of two species are tightly
linked. Some unisexuals do cover much of the geographical
range of their respective sperm donor species (e.g. Poeciliopsis: Wetherington et al., 1989; Ambystoma: Conant,
1986; Rana: Gunther, 1990; Kuzmin, 1995). These species
successfully exploit most of the available host populations,
although in no case do the unisexual species seem to exploit
this area completely.
The livebearing Amazon molly [Poecilia formosa (Girard
1859)] (Hubbs & Hubbs, 1932; Balsano et al., 1989;
Schlupp et al., 1998) on the other hand has a very limited
range (Darnell & Abramoff, 1968) (Fig. 1). It uses sperm
from males of one of three species: P. latipinna (Le Sueur

2 I. Schlupp et al.

Figure 1 Biogeographical range of the Amazon molly (Poecilia formosa and parts of its host species (P. mexicana and P. latipinna). The range
of P. formosa is depicted as dark grey. In Texas the natural range is restricted to the Rio Grande valley and the coastal area.
Blackwell Science Ltd 2002, Journal of Biogeography, 29, 16

Biogeography of Poecilia formosa 3

1821), P. mexicana Steindachner 1863 and P. latipunctata

Meek 1904. The two main sperm donor species (or hosts),
the sailn molly (P. latipinna) and the Atlantic molly
(P. mexicana) are widely distributed in the USA and Mexico.
These two species also were involved in the single hybridization event about 40,000100,000 years ago that originally
gave rise to P. formosa (Turner, 1982; Avise et al., 1991,
1992; Schartl et al., 1995; Moller et al., in preparation).
Poecilia latipinna ranges from North Carolina along the
Atlantic coast south to the Rio Tuxpan in northern Mexico
(Miller, 1983; Page & Burr, 1991). Poecilia mexicana occurs
from the Rio San Fernando south at least to Honduras, and
possibly into Costa Rica (Miller, 1983). In other words, the
Amazon molly has an enormous potential range.
Nonetheless P. formosa has a rather limited range from
the Nueces River in Texas southward to the mouth of the
Rio Tuxpan, north of the Sierra del Abra, in Mexico
(Darnell & Abramoff, 1968; Miller, 1983). The southern
limits of P. latipinna and P. formosa are identical and the
northern limits of P. mexicana and P. formosa are only some
drainages apart. Additionally some successful introductions
of the Amazon molly together with P. latipinna have been
reported from Central Texas (Hubbs et al., 1991), for
example into the San Marcos River. This distribution raises
the question of why P. formosa has not colonized more of its
host population ranges?
Several hypotheses may explain the absence of P. formosa
from much of its potential range. (1) The species could still
be expanding after its formation via hybridization, but this is
occurring slowly or it is phylogenetically extremely young.
(2) The species could have originally occurred over a larger
area, but its range was subsequently reduced. (3) The species
range may have been stationary because barriers have
prevented further dispersal. (4) Males from allopatric
populations and also other species might reject matings with
Amazon mollies. This latter hypothesis has been tested in
several studies, however, which consistently show that
despite more or less clear preferences for conspecics,
matings with Amazon mollies will occur (Hubbs, 1964;
Schlupp et al., 1991; and references therein; Schlupp et al.,
1994; Ryan et al., 1996). Gabor & Ryan (2001) showed
character displacement for male mating preferences in sailn
mollies, but even males from allopatric populations mate
with Amazon molly females, making invasions possible.
Amazon mollies generally do not produce male offspring, so
they probably need only a few matings to establish themselves in a population.
Earlier work by Hubbs & Hubbs (1946a,b) and Hubbs
(1955) showed that several species of the family Poeciliidae
can be used to breed Amazon mollies. They found that
among others P. sphenops Valenciennes 1864, P. latipinna,
P. velifera (Regan, 1914) and P. petenensis (Gunther 1866)
can serve as sperm donors, but males of the genus Xiphophorus were unsuitable. The taxonomy of the genus has
since been revised (Miller, 1983), the species which were
actually tested are unclear.
In the present study we used males of several species of the
genus Poecilia from known sympatry or allopatry to test
Blackwell Science Ltd 2002, Journal of Biogeography, 29, 16

whether they might serve as sperm-donors. We selected

species that occur along the Atlantic coast because this
matches the actual distribution of the Amazon molly. We
further examined salinity as a potential limiting factor. We
review the current situation of the species and propose a new
hypothesis explaining the limited distribution of the Amazon
molly.
MATERIALS AND METHODS
We used individuals from randomly outbred laboratory
populations in our laboratories in Hamburg and Wurzburg.
The sh were maintained in 25200 l aquaria under a
12 : 12 h light : dark cycle. They were fed ake food twice a
day, supplemented daily with brine shrimp naupliae. The
temperature was maintained at c. 25 C (for additional
details see Schlupp et al., 1991).
To determine the suitability of males as sperm donors we
tested males of different species under consistent experimental conditions to evaluate their potential as sperm donor. We
used males of all species of the genus Poecilia occurring on
the Atlantic side of Mexico. Based on sympatry with
P. formosa we tested P. mexicana mexicana, P. mexicana
limantouri, P. latipinna and P. latipunctata and Xiphophorus variatus (Meek 1904). Based on allopatry with P. formosa
we tested P. catemaconis Miller 1975, P. sphenops, P. gilli
(Kner & Steindachner 1864), P. petenensis, P. sulphuraria
(Alvarez 1948), P. velifera and P. reticulata Peters 1859 and
X. helleri Heckel 1848. In each experiment one male of the
species in question and one to ve, virgin P. formosa were
placed in a 25-l aquarium. Virgin females were obtained by
isolating young directly after birth and raising them to
sexual maturity without contact with males. The test aquaria
were observed daily for offspring. The experiment lasted
6 months or until offspring were obtained.
We tested salinity tolerance in nine Amazon mollies by
gradually increasing salinity by adding sea-salt from freshwater to 35& over 13 days. Five of the sh were adult, the
rest juvenile. The sh were kept in marine conditions for
1 week and were then gradually returned to freshwater
conditions. In another experimental series nine groups of 10
P. formosa each were transferred from freshwater conditions
to 35& salinity and returned to freshwater conditions after
48 h. In this series we recorded mortality after 48 h in
marine conditions and again 48 h after returning the sh to
freshwater. In this experiment we used two different strains
of P. formosa: diploid P. formosa I from the Brownsville
area (ve replicates) and triploid P. formosa VI/16 from the
Rio Puricacion (four replicates). We maintained one group
of each strain in 35& conditions for an additional week.
RESULTS AND DISCUSSION
Establishing potential hosts
Males of all tested species of the genus Poecilia proved to
be potential sperm donor species. All of them had
offspring with P. formosa females. The two experiments

4 I. Schlupp et al.

with members of the genus Xiphophorus, including a

syntopic X. variatus, did not result in offspring. This
corroborates earlier ndings by Hubbs & Hubbs (1946a,b)
and Hubbs (1955). Although our experiments were not
conducted in the presence of competing conspecic females,
they indicate that males of a number of poeciliids will
readily mate with P. formosa. Other experiments have
shown that males prefer conspecics in a variety of choice
experiments, but will still mate with P. formosa under
certain conditions (Hubbs, 1964; Schlupp et al., 1991 and
references therein; Ryan et al., 1996; Gabor & Ryan 2001).
Thus male mate discrimination is unlikely as an explanation
for the absence of P. formosa over large areas of the ranges
of potential hosts. Assuming these species could indeed
serve as sperm donors under natural conditions, this
essentially extends the potential range of P. formosa to
most of Central America.
Salinity tolerance
Poecilia formosa is a euryhaline secondary division freshwater species (Myers, 1938). During the salinity treatment
with gradually increasing salinity the sh showed no signs of
stress and no mortality was observed. More than fty fry were
hatched during the experiment. In the second series of
experiments even when transferred directly from 0 to 35&
salinity and back to 0& the sh showed no signs of stress
and mortality was three of fty for the P. formosa I strain
and no mortality from the forty in the triploid VI/17 strain.
This indicates that P. formosa has a high salinity tolerance
and can survive marine conditions. Furthermore it indicates
that there are no detectable differences in salinity tolerance
among P. formosa differing in ploidy.
Experiments with guppies (P. reticulata) and sailn
mollies (P. latipinna) (Gargiulo et al., 1992; Nordlie et al.,
1992) revealed a similar salinity tolerance. Poecilia formosa
and P. latipinna are often encountered in brackish coastal
lagoons (e.g. Renfro, 1960; personal observation). Furthermore, P. latipinna has been reported from semi-permanent
brackish pools on the barrier islands of the Gulf coast (Ross
& Doherty, 1994).
Changes in the distribution
Darnell & Abramoff (1968) compiled the last account of the
biogeography of the Amazon molly. Since then many
localities have changed dramatically, mainly because of
increased human activities. In the Rio Mante area, for
example, the Amazon molly has increased in abundance,
probably because of increased sugar cane production and
currently can be found together with P. mexicana and
P. latipunctata in numerous drainage ditches. In contrast to
Darnell (1962), who reported that P. latipunctata does not
serve as sperm donor species, we observed matings in nature
between P. latipunctata males and P. formosa. This nding
was conrmed in a laboratory study (Niemeitz et al.,
unpublished) and might represent a case of rapid evolution
in association with the Amazon molly.

Another range expansion occurred in central Texas. Here

P. latipinna was introduced in the 1930s and established a
thriving population (Brown, 1953). In the 1950s, a few
P. formosa from the Brownsville area were introduced into
the San Marcos River and were able to use the P. latipinna
already present to establish a new mating complex (Hubbs
et al., 1953, 1991). The origin of P. latipinna was from
Louisiana (Moller et al., in preparation), which is clearly
from outside the natural range of P. formosa and those
males must be considered as naive in respect to mating
with P. formosa females. Males from the San Marcos River
have a mating preference for conspecic females (Ryan
et al., 1996; Schlupp & Ryan, 1997; Gabor & Ryan,
2001). Nonetheless P. formosa successfully established
themselves there. Very recently, since 1999, P. formosa
was found in the spring of the San Marcos River (Hubbs,
personal communication, personal observation), an area
that was previously free of Amazon mollies, because it was
separated from the downstream region by a dam. A similar
tendency of very recent upstream expansions has been
reported by Edwards (2001) for a population in the San
Antonio River.
We revisited some localities described for P. formosa
south of Tampico in the northern part of the Laguna
Tamiahua (Darnell & Abramoff, 1968) in February 1994
and found Amazon mollies to be absent. Apparently the
habitats had been degraded signicantly and molly populations lost.
Another important change occurred in the area south of
Tampico. Here we found only one population that comprised of P. formosa and both P. latipinna and P. mexicana.
Darnell & Abramoff (1968) described this situation as a
common occurrence. This population is one of very few
where Amazon mollies interact with two sperm donor
species (Korner et al., 1999). The only other area where this
is the case is the Rio Mante system, where P. formosa utilizes
both P. mexicana and P. latipunctata as hosts.
The complex pattern of the Amazon molly biogeography
warrants a careful consideration of the potential evolutionary consequences. Currently two mating complexes are
recognized: P. formosa may occur together with P. mexicana
or P. latipinna. This, however, only roughly describes the
true situation. Two subspecies of P. mexicana have been
described (Menzel & Darnell, 1973), P. m. mexicana and
P. m. limantouri. Both are sperm donors for Amazon mollies
(Turner, 1982). Furthermore, in the case of the introduction
of P. formosa into the San Marcos River a new mating
complex with young sympatry with P. latipinna arose. In the
drainage of the Rio's Guayalejo and Mante we recognize
associations with P. mexicana and P. latipunctata. Theoretically an association with P. latipunctata alone may be
expected, potentially on a microhabitat scale. In the coastal
area south of Tampico, P. formosa may occur with either
P. latipinna or P. mexicana or both of them.
Thus, we recognize six rather than two mating complexes
of the Amazon molly, each of which provides a unique
background for evolutionary and ecological interaction of
two or three species.
Blackwell Science Ltd 2002, Journal of Biogeography, 29, 16

Biogeography of Poecilia formosa 5

The current hypothesis

We propose that near-coast marine currents constitute a
barrier to the Amazon molly's dispersal. In the Gulf of
Mexico two marine currents dominate the near-shore
sediment transport (Britton & Morton, 1989). One is
coming from the north owing in a counter-clockwise
direction south to approximately the mouth of the Nueces
River, TX, where it meets another clockwise-current coming
from the south. Where the two currents meet, a strong
upwelling occurs. This coincides with the natural northern
limit of the Amazon molly's geographical distribution
(Darnell & Abramoff, 1968).
Amazon mollies had been reported from the Nueces River
by Darnell & Abramoff (1968) but were apparently not
found there after 1977 (Turner, 1982). We were also unable
to collect them there. We found Amazon mollies in the Los
Olmos Creek (TX), a small creek south of the Nueces River
together with P. latipinna in 1995, but not in 1994. We also
collected Amazon mollies from a small drainage ditch
approximately 15 km north of the mouth of the Nueces
River in 1993. This may indicate a uctuating northern
border of the Amazon molly's distribution. However,
inadvertent introductions of mollies used as bait cannot be
ruled out.
The Rio Grande drainage is a major divide for the North
American sh fauna and is rich in endemic species. Several
species (e.g. Astyanax fasciatus, Cichlasoma) have their
northern limit here (Banarescu, 1992). The southern limit of
the Amazon molly is the area of the Laguna Tamiahua and
the Rio Tuxpan (Darnell & Abramoff, 1968; Miller, 1983;
Obregon-Banarescu, 1992; Espinosa et al., 1993). About
140 km south, a mountain range, the Sierra del Abra,
extends from the Sierra Madre Oriental to reach the coast.
These mountains constitute an important biogeographical
barrier for inland dispersal of the ichthyofauna (ObregonBanarescu, 1992).
Together with the prevailing near-shore northern currents,
the Sierra del Abra prevents southward range extensions. We
hypothesize that Amazon mollies dispersed from a single
point of origin upstream and from river-to-river system
through estuaries, occasionally passing through coastal
waters where abiotic factors such as salinity or temperature
probably do not constitute effective barriers. Our model
would place the origin of the Amazon molly in the southern
part of its present range (see also Turner, 1982), with
subsequent dispersal to the north.
ACKNOWLEDGMENTS
We thank H. Espinosa-Perez, L. Dries, M. Dobler, K. Korner,
M. J. Ryan and R. Wanker for critical comments. D. Moller
provided the map. This work was supported by the DFG
(PA 14810/12; SCHL 3445/1) and a DAAD HSP II
fellowship to I.S. I. Dibble, D. Franck, and H. Wilkens
kindly supplied sh specimens. We thank the Mexican
Government for issuing permit no. 210696-213-03 and
DOO 7501576 to collect shes and the numerous individ Blackwell Science Ltd 2002, Journal of Biogeography, 29, 16

uals who helped to collect and granted access to their

premises. A special thanks goes to M. J. Ryan for his
continued generous support.
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BIOSKETCHES
Ingo Schlupp is a Visiting Research Faculty Member at
the University of Texas at Austin and the University of
Zurich. He is currently a Heisenberg Fellow of the DFG.
He is interested in the evolution and maintenance of
recombination and sexual selection. He has been studying
the ecology and evolution of the unisexual Amazon molly
for over a decade.
Jakob Parzefall is Professor of Zoology at the University
of Hamburg. His main interest is the evolution and
behaviour of cave dwelling organisms, particularly of
cave shes and crustaceans.
Manfred Schartl is Professor of Biochemistry at the
Biocentre of the University of Wurzburg and Head of the
Department of Physiological Chemistry of the Medical
School. His research focus is on the molecular genetics
and biochemistry of tumour formation using melanoma
in sh of the genus Xiphosphorus as a model. He also
uses molecular methods to study sex determination and
the effect of gene and genome duplication on species
evolution.

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