Phylogeny and biogeography of the Malagasy and Australasian
rainbowfishes (Teleostei: Melanotaenioidei): Gondwanan
vicariance and evolution in freshwater
John S. Sparksa,*, W. Leo Smitha,b,*
aDivision of Vertebrate Zoology, Department of Ichthyology, American Museum of Natural History,
Central Park West at 79th Street, New York, NY 10024, USA
bCenter for Environmental Research and Conservation, Department of Ecology, Evolution, and Environmental Biology,
Columbia University, New York, NY 10027, USA
Received 16 March 2004; revised 21 June 2004
Available online 25 August 2004
Phylogenetic relationships of the Malagasy and Australasian rainbowfishes are investigated using 4394 characters derived from
five mitochondrial genes (12S, 16S, tRNA-Valine, ND5, and COI), three nuclear genes (28S, histone H3, and TMO-4c4), and 102
morphological transformations. This study represents the first phylogenetic analysis of the endemic Malagasy family Bedotiidae and
includes a nearly complete taxonomic review of all nominal species, as well as numerous undescribed species. Simultaneous analysis
of the molecular and morphological datasets results in two equally most parsimonious trees. Results indicate that Bedotiidae (Bedo-
tia + Rheocles) and Bedotia are monophyletic, whereas Rheocles is paraphyletic with the inclusion of two recently described species
from northeastern Madagascar, R. vatosoa, and R. derhami. Rheocles vatosoa and R. derhami are sister taxa, and this clade is recov-
ered as the sister group to Bedotia. The remaining species of Rheocles are not sexually dimorphic and comprise a clade that is recov-
ered as the sister group to Bedotia + (R. derhami + R. vatosoa), all of which are sexually dichromatic, and sexually dimorphic for
pigmentation and fin development. Three geographically distinct clades are recovered within Bedotia, one comprising species with
distributions ranging from mid- to southeastern Madagascar, another including species restricted to eastern drainages north of the
Masoala Peninsula, and a third comprising taxa with distributions extending from the Masoala Peninsula south to the Ivoloina
River. The Australian/New Guinean melanotaeniids are monophyletic and are recovered as the sister group to Bedotiidae. The Aus-
tralasian Telmatherinidae and Pseudomugilidae comprise a clade that is recovered as the sister group to the Melanotaeniidae-Bed-
otiidae clade. This sister-group relationship between Malagasy bedotiids and a clade restricted to Australia–New Guinea, and the
absence of a close relationship between bedotiids and African or Mascarene atheriniforms, is congruent with the break-up of
Gondwana, not a scenario reliant on Cenozoic trans-oceanic dispersal. Finally, results of the phylogenetic analysis indicate that
Atheriniformes is polyphyletic and further corroborate recent morphological hypotheses, which have recovered Bedotiidae in a
derived position within Atherinoidei.
? 2004 Elsevier Inc. All rights reserved.
Keywords: Bedotia; Rheocles; Bedotiidae; Melanotaeniidae; Systematics; Madagascar; Gondwana
The atherinoid family Bedotiidae is endemic to Mad-
agascar and comprises two genera, Bedotia and Rheocles.
These small (most species <100.0mm SL), laterally
compressed, and generally colorful fishes are commonly
1055-7903/$ - see front matter ? 2004 Elsevier Inc. All rights reserved.
*Corresponding authors. Fax: 1 212 769 5642.
E-mail addresses: firstname.lastname@example.org (J.S. Sparks), leosmith@
amnh.org (W.L. Smith).
Molecular Phylogenetics and Evolution 33 (2004) 719–734
referred to as Malagasy rainbows or rainbowfishes. Most
bedotiids exhibit varying degrees of sexual dimorphism,
which ranges from subtle to striking. Bedotia and Rheo-
cles occur exclusively in freshwater environments and are
distributed in small to medium-sized forested rivers and
streams, occasionally in swamps and marshes, spanning
nearly the entire eastern slope of Madagascar (a single
species of Rheocles is recorded from a westward draining
basin in the northeast of the island). Bedotia (Regan,
1903) comprises six nominal species [B. madagascariensis
(Regan, 1903), B. geayi (Pellegrin, 1907), B. longianalis
(Pellegrin, 1914b), B. tricolor (Pellegrin, 1932), B. maro-
jejy (Stiassny and Harrison, 2000), and B. masoala
(Sparks, 2001)]. Relationships among constituent taxa
remain, for the most part, unresolved. Except for color-
ation and pigmentation, Bedotia are morphologically
conservative fishes. In a recent checklist of the freshwater
fishes of Africa, Mauge ´ (1986) synonymized B. longia-
nalis and B. tricolor with B. geayi, although no justifica-
tion for this taxonomic rearrangement was given
(Sparks, 2001). Based on examination of the primary
types for all species of Bedotia and comparison of collec-
tions made throughout eastern Madagascar, it is appar-
ent that several distinct species of Bedotia inhabit the
island, many of which are undescribed (Sparks and
Rheocles (Jordan and Hubbs, 1919) comprises seven
valid species [viz., R. sikorae (Sauvage, 1891), R. alaotr-
ensis (Pellegrin, 1914a), R. pellegrini (Nichols and La-
Monte, 1931), R. wrightae (Stiassny, 1990), R. lateralis
(Stiassny and Reinthal, 1992), R. derhami (Stiassny
and Rodriguez, 2001), and R. vatosoa (Stiassny et al.,
2002)]. Rheocles was recently revised by Stiassny
(1990), in which a third Malagasy bedotiid genus, Rheo-
cloides (Nichols and LaMonte, 1931), was formally syn-
onymized with Rheocles. The number of described
species has nearly doubled since Stiassny?s (1990) revi-
sion, and placement of the two most recently described
species within the genus is problematic.
Monophyly of Bedotia, Rheocles, and a clade com-
prising Bedotia and Rheocles (= Bedotiidae) has been
addressed in several recent studies (Aarn and Ivantsoff,
1997; Dyer and Chernoff, 1996; Stiassny, 1990). This
monophyletic assemblage has been given either familial,
Bedotiidae (Rosen and Parenti, 1981; Saeed et al., 1994;
Stiassny, 1990), or subfamilial rank, Bedotiinae, within
Melanotaeniidae (Dyer and Chernoff, 1996). Regardless
of rank, neither monophyly nor the composition of Bed-
otiidae is disputed, yet their placement within the so-
called ‘‘atherinoid’’ group of Atherinomorpha has been
the subject of considerable debate (Aarn and Ivantsoff,
1997; Dyer and Chernoff, 1996; Stiassny, 1990) (Fig.
1). Rosen and Parenti (1981) concluded that evidence
was lacking to support monophyly of atherinoids, an
assemblage of six families (Atherinidae, Bedotiidae,
Isonidae, Melanotaeniidae, Phallostethidae, and Tel-
matherinidae), which they referred to as Division I of
Series Atherinomorpha. Stiassny (1990) and Saeed
et al. (1994) failed to provide evidence in support of
monophyly of Atherinoidei. A monophyletic Atherinoi-
dei, comparable to that delimited by Rosen and Parenti
(1981), however, was recovered by Dyer and Chernoff
(1996). Although assigned different ranks by various
authors, for clarity and consistency familial-level rank
is retained throughout for bedotiids, melanotaeniids,
and pseudomugilids (herein including telmatherinids),
which we refer to collectively as rainbowfishes.
The similarity between the Malagasy bedotiids and
Australian/New Guinean melanotaeniids is striking,
and a close relationship has been intimated by numerous
authors dating back to the early 20th century (Jordan
and Hubbs, 1919). In recent cladistic analyses, bedotiids
and melanotaeniids are either recovered as sister taxa
(Saeed et al., 1994), bedotiids are the sister taxon to a
clade comprising melanotaeniids, pseudomugilids, and
telmatherinids (Dyer and Chernoff, 1996), or bedotiids
are nested within a paraphyletic melanotaeniid assem-
blage (Aarn and Ivantsoff, 1997) (Fig. 1). The placement
and interrelationships of telmatherinids and pseudo-
mugilids, likewise, varies considerably among analyses.
Fig. 1. Hypotheses of inter- and intrafamilial relationships of Bed-
otiidae and Melanotaenioidei based on morphological evidence
according to (A) Stiassny (1990), (B) Saeed et al. (1994), (C) Dyer
and Chernoff (1996), and (D) Aarn and Ivantsoff (1997).
J.S. Sparks, W.L. Smith / Molecular Phylogenetics and Evolution 33 (2004) 719–734
According to Dyer and Chernoff (1996), pseudomugilids
and telmatherinids comprise the sister taxon to melano-
taeniids, and this clade is in turn the sister taxon to bed-
otiids, whereas the hypotheses presented by Saeed et al.
(1994) and Aarn and Ivantsoff (1997) do not unite
telmatherinids or pseudomugilids in a clade with bedoti-
ids and melanotaeniids (Fig. 1).
Genotypic and phenotypic characters will be used to
test these hypotheses of relationships and to examine the
biogeography of Malagasy and Australasian rainbowf-
ishes within a phylogenetic context. Results of recent
molecular studies suggest that many of Madagascar?s
terrestrial vertebrate lineages owe their origin to random
dispersal from Africa or Asia, well after the Mesozoic
break-up of the Gondwanan supercontinent (Caccone
et al., 1999; Jansa et al., 1999; Mausfeld et al., 2000;
Nagy et al., 2003; Raxworthy et al., 2002; Yoder and
Yang, 2004; Yoder et al., 1996, 2003). The conclusions
of these studies reiterate the prevailing opinion offered
by paleontologists and many ichthyologists (e.g., Aarn
and Ivantsoff, 1997) that Madagascar?s extant verte-
brates, including freshwater fishes, arrived via oceanic
dispersal (Gottfried and Krause, 1998; Krause et al.,
1997; Lundberg, 1993; Murray, 2001). Paleontologists
have defended dispersal hypotheses by citing a lack of
Cretaceous-age fossils for Madagascar?s extant assem-
blages of freshwater fishes (Gottfried and Krause,
1998; Krause et al., 1997; Lundberg, 1993; Murray,
2001) and by pointing to the alleged salinity tolerance
of some Malagasy freshwater fishes (Lundberg, 1993;
Murray, 2001). Our study will test the sister group rela-
tionship of Malagasy and Australasian rainbowfishes
recovered by Dyer and Chernoff (1996), a topology con-
sistent with the fragmentation of Gondwana. We will
examine the distribution patterns of rainbowfishes
throughout their range for congruence with regional
vicariance events. For example, it is generally assumed
that rainbowfishes have dispersed via the ocean in the
Australasian region; we will examine these conjectures
in light of phylogenetic and paleogeographic evidence.
We present a phylogenetic hypothesis for rainbowf-
ishes based upon the simultaneous analysis of 4292
molecular characters and the 102 morphological trans-
formations described by Dyer and Chernoff (1996). This
analysis includes all nominal bedotiid species, less two
members of Rheocles (viz., R. sikorae and R. pellegrini)
for which tissue samples could not be obtained. Rheocles
sikorae, a species from eastern forests that has not been
collected in recent years, is presumed to be extinct, and
R. pellegrini, from northern Madagascar, is known only
from the holotype and two (out of three original)
remaining paratypes. Representatives from several pop-
ulations of Bedotia hypothesized to represent unde-
comparisons (Loiselle and Stiassny, 2003; Sparks and
Stiassny, 2003; Stiassny, pers. comm.) are also included.
The objectives of this study are: (1) to test monophyly of
both Bedotiidae and an assemblage comprising the three
Australasian rainbowfish families; (2) to determine if a
sister-group relationship between the Malagasy bedoti-
ids and Australasian rainbowfishes is supported with
the addition of nucleotide characters; (3) to determine
if the addition of molecular characters corroborates a
derived placement for bedotiids and melanotaeniids
within Atheriniformes; (4) to test monophyly of both
Rheocles and Bedotia and resolve intrageneric relation-
ships within these genera; and (5) to determine if the
recovered phylogenetic pattern for the Malagasy and
Australasian rainbowfishes is congruent with Earth
2. Materials and methods
2.1. Taxon sampling
The resulting hypotheses were rooted using the basal
acanthomorph Percopsis. The putative atherinomorph
sister-group, Mugilidae (following Stiassny, 1990), and
three non-atheriniform atherinomorphs (Cololabis, Fun-
dulus, and Oryzias) were included to rigorously test the
monophyly of the Atheriniformes, rainbowfishes (Mela-
notaenioidei), and Bedotiidae. Fifty-nine atheriniform
taxa were analyzed, including representatives from all
atheriniform families. Ten Australasian rainbowfishes,
all nominal bedotiid species (except Rheocles pellegrini
and R. sikorae), and 11 undescribed Bedotia species were
included in the analysis. Finally, both Malagasy non-
bedotiid atheriniform genera with members that occur
in freshwater (Atherinomorus and Teramulus) were
2.2. Acquisition of nucleotide sequences
Fish tissues were preserved in 70–95% ethanol prior
to DNA extraction. Total genomic DNA was extracted
from muscle, liver, or fin clips via use of a Qiagen Tissue
Extraction Kit (QIAamp or DNeasy Tissue Extraction
Kit) following the manufacturer?s protocol. PCR was
used to amplify fragments from five mitochondrial genes
(12S, 16S, tRNA-Val, ND5, and COI) and three nuclear
genes (28S, histone H3, and TMO-4c4). Double-
stranded amplifications were performed in either 25 or
50lL volumes containing 1· PCR buffer (Qiagen),
2mM MgCl2, 0.2mM of each dNTP, 0.2–0.5lL of each
primer (10lM), 10–1000ng of genomic DNA (1–2lL),
and 1lL of Taq polymerase (Qiagen), or in a 25lL vol-
ume containing one Ready-To-Go PCR bead (Amer-
sham Biosciences), 1.25lL of each primer (10lM) and
2–5lL of genomic DNA. PCR primers and profiles
for all genes can be found in Smith and Wheeler (2004)
or Sparks (2004), except for NADH dehydrogenase
J.S. Sparks, W.L. Smith / Molecular Phylogenetics and Evolution 33 (2004) 719–734
Gottfried, M.D., Krause, D.W., 1998. First record of gars (Lepiso-
steidae, Actinopterygii) on Madagascar: Late Cretaceous remains
from the Mahajunga basin. J. Vert. Paleont. 18, 275–279.
Grande, L., Eastman, J.T., 1986. A review of Antarctic ichthyofaunas
in the light of new fossil discoveries. Palaeontology 29, 133–137.
Hall, R., 1998. The plate tectonics of Cenozoic SE Asia and the
distribution of land and sea. In: Hall, R., Holloway, J.D. (Eds.),
Biogeography and Geological Evolution of SE Asia. Backhuys
Publishers, Leiden, The Netherlands, pp. 99–131.
Hall, R., 2001. Cenozoic reconstructions of SE Asia and the SW
Pacific: changing patterns of land and sea. In: Metcalfe, I., Smith,
J.M.B., Morwood, M., Davidson, I.D. (Eds.), Faunal and Floral
Migrations and Evolution in SE Asia-Australasia. Swets and
Zeitlinger Publishers, Lisse, pp. 35–56.
Hay, W.W., De Conto, R.M., Wold, C.N., Wilson, K.M., Voight, S.,
Shulz, M., Wold, A.R., Dullo, W.-C., Ronov, A.B., Balukhovsky,
A.N., So ¨ding, E., 1999. Alternative global Cretaceous paleogeog-
raphy. In: Barrera, E., Johnson, C.C. (Eds.), Evolution of the
Cretaceous Ocean-Climate System. Geol. Soc. America, Special
Paper 332, pp. 1–47.
Jansa, S.A., Goodman, S.M., Tucker, P.K., 1999. Molecular phylog-
eny and biogeography of the native rodents of Madagascar
(Muridae: Nesomyinae): a test of the single-origin hypothesis.
Cladistics 15, 253–270.
Jordan D.S., Hubbs, C.L., 1919. Studies in ichthyology. A mono-
graphic review of the family of Atherinidae or silversides. Leland
Stanford Jr. Univ. Publ., Univ. Ser. 919 pp. 1–87.
Kluge, A.G., Farris, J.S., 1969. Quantitative phyletics and the
evolution of anurans. Syst. Zool. 18, 1–32.
Kottelat, M., 1991. Sailfin silversides (Pisces: Telmatherinidae) of Lake
Matano, Sulawesi, Indonesia, with descriptions of six new species.
Ichthyol. Explor. Freshwaters 1, 321–344.
Kottelat, M., Whitten, A.J., 1993. Freshwater Fishes of Western
Indonesia and Sulawesi. Periplus Editions (HK), Indonesia.
Krause, D.W., Hartman, J.H., Wells, N.A., 1997. Late Cretaceous
vertebrates from Madagascar: implications for biotic change in
deep time. In: Goodman, S.M., Patterson, B.D. (Eds.), Natural
Change and Human Impact in Madagascar. Smithsonian Institu-
tion Press, Washington, DC, pp. 3–43.
Li, Z.X., Powell, C.McA., 2001. An outline of palaeogeographic
evolution of the Australasian region since the beginning of the
Neoproterozoic. Earth Sci. Rev. 53, 237–277.
Loiselle, P.V., Stiassny, M.L.J., 2003. Bedotia. In: Goodman, S.M.,
Benstead, J.P. (Eds.), The Natural History of Madagascar.
University Chicago Press, Chicago, pp. 867–868.
Lundberg, J.G., 1993. African-South American freshwater fish clades
and continental drift: problems with a paradigm. In: Goldblatt, P.
(Ed.), Biological Relationships between Africa and South America.
Yale University Press, New Haven, pp. 156–199.
Mauge ´, A.L., 1986. Atherinidae. In: Daget, J., Gosse, J.-P., Thys van
den Audenaerde, D.F.E. (Eds.), Checklist of the Freshwater Fishes
of Africa, vol. II. Institut Royal des Sciences Naturelles de
Belgique, Bruxelles, Muse ´e Royal de l?Afrique Centrale, Tervuren,
Office de la Recherche Scientifique et Technique Outre-Mer, Paris,
Mausfeld, P., Vences, M., Schmitz, A., Veith, M., 2000. First data on
the molecular phylogeography of scinid lizards of the genus
Mabuya. Mol. Phylogenet. Evol. 17, 11–14.
McGuigan, K., Zhu, D., Allen, G.R., Moritz, C., 2000. Phylogenetic
relationships and historical biogeography of melanotaeniid fishes in
Australia and New Guinea. Mar. Fresh. Res. 51, 713–723.
Miya, M., Takeshima, H., Endo, H., Ishiguro, N., Inoue, J.G., Mukai,
T., Satoh, T.P., Yamaguchi, M., Kawaguchi, A., Mabuchi, K.,
Shirai, S.M., Nishida, M., 2003. Major patterns of higher teleos-
tean phylogenies: a new perspective based on 100 complete
mitochondrial DNA sequences. Mol. Phylogenet. Evol. 26, 121–
Moss, S.J., Wilson, M.E.J., 1998. Biogeographic implications of the
Tertiary paleogeographic evolution of Sulawesi and Borneo. In:
Hall, R., Holloway, J.D. (Eds.), Biogeography and Geological
Evolution of SE Asia. Backhuys Publishers, Leiden, The Nether-
lands, pp. 133–163.
Morrone, J.J., Carpenter, J.M., 1994. In search of a method for
cladistic biogeography: an empirical comparison of component
analysis, Brooks parsimony analysis, and three-area statements.
Cladistics 10, 99–153.
Murphy, W.J., Collier, G.E., 1997. A molecular phylogeny for
aplocheiloid fishes (Atherinomorpha, Cyprinodontiformes): the
role of vicariance and the origins of annualism. Mol. Biol. Evol. 14,
Murray, A.M., 2001. The fossil record and biogeography of the
Cichlidae (Actinopterygii: Labroidei). Biol. J. Linn. Soc. 78, 517–
Nagy, Z.T., Joger, U., Wink, M., Glaw, F., Vences, M., 2003. Multiple
colonization of Madagascar and Socotra by colubrid snakes:
evidence from nuclear and mitochondrial gene phylogenies. Proc.
R. Soc. Lond. B 270, 2613–2621.
Nelson, J.S., 1994. Fishes of the World, third ed. Wiley, New York.
Nelson, G.J., Platnick, N.I., 1981. Systematics and Biogeography:
Cladistics and Vicariance. Columbia University Press, New York.
Nichols, J.T., LaMonte, F.R., 1931. Rheocloides, a new atherinid fish
from Madagascar. Am. Mus. Novit. 508, 1–2.
Nixon, K.C., 1999. The parsimony ratchet, a new method for rapid
parsimony analysis. Cladistics 15, 407–414.
Page, R.D.M., 1988. Quantitative cladistic biogeography: constructing
and comparing area cladograms. Syst. Zool. 37, 254–270.
Patterson, C., 1993a. An overview of the early fossil record of
acanthomorphs. Bull. Mar. Sci. 52, 29–59.
Patterson, C., 1993b. Osteichthys: Teleostei. In: Benton, M.J.
(Ed.), The Fossil Record 2. Chapman and Hall, London, pp.
Pellegrin, J., 1907. Liste des poissons recueillis a ` Madagascar par M. F.
Geay. Description d?une espe ´ce nouvelle. Bull. Mus. Natl. Hist.
Nat. (Se ´r. 1) 13, 201–206.
Pellegrin, J., 1914a. Sur une Athe ´rine nouvelle des eaux douces de
Madagascar. Bull. Soc. Zool. Fr. 39, 46–47.
Pellegrin, J., 1914b. Sur un Athe ´rinide ´ nouveau de Madagascar
appartenant au genre Bedotia. Bull. Soc. Zool. Fr. 39, 178–180.
Pellegrin, J., 1932. Athe ´rinide ´ nouveau de Madagascar appartenant au
genre Bedotia. Bull. Soc. Zool. Fr. 57, 84–86.
Rabinowitz, P.D., Coffin, M.F., Falvey, D., 1983. The separation of
Madagascar and Africa. Science 220, 67–69.
Raxworthy, C.J., Forstner, M.R.J., Nussbaum, R.A., 2002. Chame-
leon radiation by oceanic dispersal. Nature 415, 784–787.
Reeves, C., de Wit, M., 2000. Making ends meet in Gondwana:
retracing the transforms of the Indian Ocean and reconstructing
continental shear zones. Terra Nova 12, 272–280.
Regan, C.T., 1903. Descriptions de poissons nouveaux faisant partie
de la collection du Muse ´e d?Histoire Naturelle de Gene `ve. Rev.
Suisse Zool. 11, 413–418.
Riseng, K.J., 1997. The distribution of fishes and the conservation of
aquatic resources in Madagascar. Unpublished M.Sc. thesis.
University of Michigan, Ann Arbor.
Rosen, D.E., 1964. The relationships and taxonomic position of the
halfbeaks, killifishes, silversides, and their relatives. Bull. Am. Mus.
Nat. Hist. 127, 217–268.
Rosen, D.E., 1978. Vicariant patterns and historical explanation in
biogeography. Syst. Zool. 27, 159–188.
Rosen, D.E., Parenti, L.R., 1981. Relationships of Oryzias, and the
groups of Atherinomorph fishes. Am. Mus. Novitates 2719,
Saeed, B., Ivantsoff, W., 1991. Kalyptatherina, the first telmatherinid
genus known outside of Sulawesi. Ichthyol. Explor. Freshwaters 2,
J.S. Sparks, W.L. Smith / Molecular Phylogenetics and Evolution 33 (2004) 719–734
Saeed, B., Ivantsoff, W., Allen, G.R., 1989. Taxonomic revision of the
family Pseudomugilidae (Order Atheriniformes). Aust. J. Mar.
Freshw. Res. 40, 719–787.
Saeed, B., Ivantsoff, W., Crowley, L.E.L.M., 1994. Systematic
relationships of atheriniform families within Division I of the
series Atherinomorpha (Acanthopterygii) with relevant historical
perspectives. Voprosi Ikhtiologii 34, 1–32.
Sauvage, H.E., 1891. Histoire naturelle des poissons. In: Grandidier,
A. (Ed.), Histoire physique, naturelle et politique de Madagascar.
Vol. 16. Imprimerie Nationale, Paris, pp. 1–543.
Scotese, C.R., Boucot, A.J., McKerrow, W.S., 1999. Gondwanan
palaeogeography and palaeoclimatology. J. African Earth Sci. 28,
Smith, A.G., Smith, D.G., Funnell, B.M., 1994. Atlas of Mesozoic and
Cenozoic Coastlines. Cambridge University Press, Cambridge.
Smith, W.L., Wheeler, W.C., 2004. Polyphyly of the mail-cheeked
fishes (Teleostei: Scorpaeniformes): evidence from mitochondrial
and nuclear sequence data. Mol. Phylogenet. Evol. 32, 627–646.
Sparks, J.S., 2001. Bedotia masoala, a new species of atherinoid
rainbowfish (Teleostei: Bedotiidae) from the Masoala Peninsula,
northeastern Madagascar. Copeia 2001, 482–489.
Sparks, J.S., 2004. Molecular phylogeny and biogeography of the
Malagasy and South Asian cichlids (Teleostei: Perciformes:
Cichlidae). Mol. Phylogenet. Evol. 30, 599–614.
Sparks, J.S., Smith, W.L., in press. Freshwater fishes, dispersal ability,
and non-evidence: ‘‘Gondwana life rafts’’ to the rescue. Syst. Biol.
Sparks, J.S., Stiassny, M.L.J., 2003. Introduction to the freshwater
fishes. In: Goodman, S.M., Benstead, J.P. (Eds.), The Natural
History of Madagascar. University Chicago Press, Chicago, pp.
Springer, V.G., 1982. Pacific plate biogeography, with special reference
to shorefishes. Smithsonian Cont. Zool. 367, 1–182.
Stiassny, M.L.J., 1990. Notes on the anatomy and relationships of the
bedotiid fishes of Madagascar, with a taxonomic review of the
genus Rheocles. Am. Mus. Novitates 2979, 1–33.
Stiassny, M.L.J., Harrison, I.J., 2000. Notes on a small collection of
fishes from the Parc National de Marojejy, northeastern Mada-
gascar, with a description of a new species of the endemic genus
Bedotia (Atherinomorpha: Bedotiidae). In: Goodman, S.M. (Ed.),
A Floral and Faunal Inventory of the Parc National de Marojejy,
Madagascar: With Reference to Elevational Variation. Fieldiana
Zool., new series, 97, 1–286, pp. 143–156.
Stiassny, M.L.J., Reinthal, P.N., 1992. Description of a new species of
Rheocles (Atherinomorpha, Bedotiidae) from the Nosivolo Tribu-
tary, Mangoro River, Eastern Malagasy Republic. Am. Mus.
Novitates 3031, 1–8.
Stiassny, M.L.J., Rodriguez, D.M., 2001. Rheocles derhami, a new
species of freshwater rainbowfish (Atherinomorpha: Bedotiidae)
from the Ambomboa River in northeastern Madagascar. Ichthyol.
Explor. Freshwaters 12, 97–104.
Stiassny, M.L.J., Rodriguez, D.M., Loiselle, P.V., 2002. Rheocles
vatosoa, a new species of freshwater rainbowfish (Atherinomorpha:
Bedotiidae) from the Lokoho River basin in northeastern Mada-
gascar. Cybium 26, 71–77.
Storey, B.C., 1995. The role of mantle plumes in continental breakup:
case histories from Gondwanaland. Nature 377, 301–308.
Storey, B.C., Mahoney, J.J., Saunders, A.D., Duncan, R.A.,
Kelley, S.P., Coffin, M.F., 1995. Timing of hot spot-related
volcanism and the breakup of Madagascar and India. Science
Torsvik, T.H., Tucker, R.D., Ashwal, L.D., Carter, L.M., Jamtveit, B.,
Vidyadharan, K.T., Venkataramana, P., 2000. Late Cretaceous
India-Madagascar fit and timing of break-up related magmatism.
Terra Nova 12, 220–224.
Vari, R.P., 1992. Redescription of Mesopristes elongatus (Guichenot,
1866), an endemic Malagasy fish species (Pisces, Terapontidae).
Am. Mus. Novitates 3039, 1–7.
Voris, H.K., 2002. Maps of Pleistocene sea levels in Southeast Asia:
shorelines, river systems and time durations. J. Biogeogr. 27, 1153–
Wheeler, W.C., 1996. Optimization alignment: the end of multiple
sequence alignment in phylogenetics. Cladistics 12, 1–9.
Wheeler, W.C., 2003a. Iterative pass optimization of sequence data.
Cladistics 19, 254–260.
Wheeler, W.C., 2003b. Implied alignment: a synapomorphy-based
multiple sequence alignment method. Cladistics 19, 261–268.
Wilson, M.E.J., Moss, S.J., 1999. Cenozoic paleogeographic evolution
of Sulawesi and Borneo. Palaeogeogr. Palaeoclimatol. Palaeoecol.
Yoder, A., Yang, Z., 2004. Divergence dates for Malagasy lemurs
estimated from multiple gene loci: geological and evolutionary
context. Mol. Ecol. 13, 757–773.
Yoder, A.D., Cartmill, M., Ruvolo, M., Smith, K., Vilgalys, R., 1996.
Ancient single origin for Malagasy primates. Proc. Natl. Acad. Sci.
USA 93, 5122–5126.
Yoder, A.D., Burns, M., Zehr, S., Delefosse, T., Veron, G., Goodman,
S.M., Flynn, J.J., 2003. Single origin of Malagasy Carnivora from
an African ancestor. Nature 421, 734–737.
J.S. Sparks, W.L. Smith / Molecular Phylogenetics and Evolution 33 (2004) 719–734