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versión impresa ISSN 0327-9545

Biocell v.30 n.1 Mendoza ene./abr. 2006


Systematic revision of the South American silversides (Teleostei, Atheriniformes)

Brian S. Dyer H.

Escuela de Recursos Naturales, Universidad del Mar

Address correspondence to: Brian S. Dyer. Escuela de Recursos Naturales, Universidad del Mar, Amunátegui 1838, Recreo, Viña del Mar, CHILE. E-mail:

Key words: Atherinopsinae; Odontesthes; Phylogeny; Taxonomy; Morphology

The taxonomic and systematic history of the South American silversides has changed significantly since Campos (1984) review. As a result of phylogenetic studies of the subfamilies Menidiinae (Chernoff, 1986b), Atherinopsinae (White, 1985; Crabtree, 1987; Dyer 1997, 1998) and of the order Atheriniformes (Saeed et al., 1994; Dyer and Chernoff, 1996) the taxonomy and classification of silversides has changed notoriously (Table I). Two systematic studies with different methodologies and divergent results in many ways coincided in that the subfamilies Menidiinae and Atherinopsinae, traditionally considered as atherinids, were to form part of the family Atherinopsidae (Saeed et al., 1994; Dyer and Chernoff, 1996).
The purpose of this paper is to review the present state of the systematics (taxonomy, relationships and distribution) of the South American silversides in the context of phylogenetic revisions of Atheriniformes and Atherinopsinae. All systematic results presented herein are based on published material (Dyer and Chernoff, 1996; Dyer, 1993, 1997, 1998, 2000, 2003a,b; Dyer and Gosztonyi, 1999; Malabarba and Dyer, 2002). In addition to some diagnostic features of the taxa involved (genera, tribes, subfamilies, and families), an identification key and some identification and diagnostic problems, are presented.


Atheriniformes is phylogenetically diagnosed by ten characters (Dyer and Chernoff, 1996) and is sister to the superorder Cyprinodontea (Fig. 1), formed by the orders Beloniformes and Cyprinodontiformes.1 These three orders form part of the series Atherinomorpha (Fig. 1). Six families and 49 genera are recognized in Atheriniformes (Dyer and Chernoff, 1996; Table 1). More recent studies of relationships among families of Atheriniformes involve only the melanotaeniids, pseudomugilids, and telmatherinids (Aarn and Ivantsoff, 1997; Aarn et al., 1998). One of the main conclusions of Saeed et al. (1994) and Dyer and Chernoff (1996) is that subfamilies and genera traditionally considered as a part of Atherinidae, now form part of other families such as: Atherinopsidae (Menidiinae and Atherinopsinae), Atherionidae (Atherion), and Phallostethidae (Dentatherina). Atherinidae now includes only the subfamilies Atherininae, Atherinomorinae, and Craterocephalinae (Fig. 1), and is represented in America by three atherinomorine species found in the Caribbean (Dyer, 2003a): Hypoatherina harringtonensis, Atherinomorus stipes, and Alepidomus evermanni. According to the phylogenetic hypothesis of Dyer and Chernoff (1996), Atherinopsidae is the sister group of the remaining atheriniform families (Fig. 1). The other atheriniform family present in American waters of the Southern Hemisphere is Notocheiridae, and is the sequential sister group of the remaining atheriniforms (Fig. 1).

FIGURE 1. Phylogenetic relationships among families of Atheriniformes (Dyer and Chernoff, 1996), modified in Melanotaeniidae reflecting alternative hypotheses proposed by Aarn and Ivantsoff (1997) and Aarn et al. (1998).

Genera and species are organized by families, Notocheiridae and Atherinopsidae, and ordered hierarchically within each family. The systematic history is discussed briefly for each taxonomic group.

Notocheiridae Schultz

Schultz (1948) proposed Notocheirus and Iso as atherinid sister groups within Tropidostethinae. Because Tropidostethinae was preocuppied, Schultz (1950) corrected the name to Notocheirinae and designated Notocheirus hubbsi Clark (1937), as type species. This correction was followed by Fowler (1951) and is why Notocheiridae has priority over Isonidae of Rosen (1964). Isonidae has been the name most widely used for this group (Greenwood et al., 1966; Rosen and Parenti, 1981; Nelson, 1984). Saeed et al. (1994) proposed Notocheiridae and Isonidae for each of the genera, an unnecessary and unjustified position according to Dyer and Chernoff (1996). Notocheiridae, with the single species Notocheirus hubbsi is distributed from Valparaiso in the Pacific and Puerto Deseado in the Atlantic, to the southern tip of Tierra del Fuego. Notocheirus hubbsi is a rare species and scarce in collections because it inhabits the surf and has been collected only accidentally in intertidal pools in the South of Chile. Gosztonyi (1972) is the only person that has had the privilege to study a collection of 22 female specimens from a single locality. The presence in some specimens of odontodes (teeth on head bones related to laterosensory canals) is a signal of a potential new species (Dyer, 2000), for which more specimens need to be examined.

Atherinopsidae Fowler

Atherinopsidae is diagnosed by 20 characters that identify it as a monophyletic group and sister to the remaining families of Atheriniformes (Atherinoidei: Dyer and Chernoff, 1996). Composed of 13 genera and 104 species (Dyer, 2003b) grouped in the subfamilies Menidiinae y Atherinopsinae (Fig. 2), endemic to the continental and coastal marine waters of the Americas (Dyer, 1997). The sister group relationship between both subfamilies was proposed by White (1985) and Chernoff (1986b) in their systematic revisions of Atherinopsinae and Menidiinae, respectively, and corroborated by Dyer and Chernoff (1996).

FIGURE 2. Phylogenetic relationships among genera of Atherinopsidae (Chernoff, 1986b; Dyer, 1997, 1998).

Menidiinae Schultz

Menidiinae was proposed by Schultz (1948) to lodge all American atherinids that lack a haemal funnel, however its monophyly was confirmed much later by Chernoff (1986b). Menidiinae is significantly larger than Atherinopsinae, with 74 species distributed mostly in continental waters of North and Central America. Menidiinae is diagnosed by 13 characteres (Chernoff, 1986b; Dyer, 1997) and is composed of two tribes and five genera (Fig. 2): Menidiini (Menidia [inc. Chirostoma], Labidesthes) and Membradini (Membras, Atherinella, Melanorhinus), of which only Atherinella has representatives in the Southern Hemisphere (Chernoff, 1986b).
Membradini is diagnosed by 6 characters (Chernoff, 1986b; Dyer, 1997, 1998) and is composed of 43 species, 35 of which are grouped into four subgenera of Atherinella. The subgenus Eurystole is composed of 17 freshwater and marine species, the latter of which are found in the Atlantic (from Costa Rica to Central Brazil) and Pacific (from Gulf of California to northern Chile, including the Galapagos Islands) (Chernoff, 1986b). Atherinella (Eurystole) nocturna (Myers and Wade, 1942) is distributed along the coasts of Ecuador, Peru and northern Chile (Chernoff, 1986b; Kong and Bolados, 1987). The subgenus Xenomelaniris is composed of only three marine and estuarine species: A. robbersi en Lake Totumo, Colombia; A. venezuelae in Trinidad & Tobago and Venezuela; and A. brasiliensis in the Atlantic from Venezuela to Uruguay (Dyer, 2003b).

Atherinopsinae Fowler

Atherinopsinae was created by Fowler (1903) for all atherinids without premaxillary protrusion. Schultz (1948) redefined Atherinopsinae to include those species with distally dilated premaxillaries and a haemal funnel. White (1985) revised the group phylogenetically and determined it as a monophyletic group with eight diagnostic characters. Atherinopsinae is diagnosed by a different set of eight characters (Dyer 1997, 1998)2 and is composed of six genera in two antitropical tribes: Atherinopsini in North America (Atherinops, Atherinopsis, Colpichthys, Leuresthes) and Sorgentinini in South America (Basilichthys, Odontesthes). Crabtree (1987) tested White's (1985) hypothesis using electrophoretic evidence, but had ambiguous results depending on the method of coding the data. Dyer (1997) analyzed the accumulated morphological evidence together with the enzymatic evidence and corroborated the monophyly of the tribes as did the monophyly of Basilichthys and Odontesthes (Figs. 2, 3).

FIGURE 3. Phylogenetic relationships among species of Sorgentinini (sp-gr= species group), showing subgenera Odontesthes, Cauque y Austromenidia.

Sorgentinini is diagnosed by eight characters and composed of two genera and at least 19 species (Dyer, 1997, 1998). White (1985) originally called this tribe Basilichthyini, not knowing the previous availability of Sorgentinini (see White, 1989).

Basilichthys Girard

Basilichthys is diagnosed by 22 characters and is composed of five species in two species groups (Dyer, 1997): the microlepidotus species group (B. australis and B. microlepidotus) and the semotilus species group (B. archaeus, B. semotilus and B. sp.). Basilichthys is found only in lakes and rivers of the western versant of the Andes from Reque River in Lambayeque (7° S), Peru, to rivers of Chiloé Island (43° S), Chile. Basilichthys microlepidotus records in Argentina (Evermann and Kendall, 1906; Aramburu and Ringuelet, 1965) are nomenclatorial mistakes attributed to the Patagonian silverside Odontesthes hatcheri.
The microlepidotus species group, though without a phylogenetically diagnostic character, has descriptive features that clearly identify it as a group (see key, Gajardo, 1987). It is distributed in Central Chile from Huasco River, III Región (28° - 29°S) to Chiloé Island, X Región (42° - 43°S) (Fig. 4).

FIGURE 4. Distribution map of marine and freshwater species of sorgentinins and superimposed cladogram of species relationships. Root of tree and node 04 represented by blackfilled circle. Subgenus Cauque is not included in this diagram of relationships.

The semotilus species group is diagnosed by four characters and is distributed from Reque River, Lambayeque (7°S), Perú, to the Loa River, Iquique (22°S), I Región, Chile (Fig. 4).

Odontesthes Evermann & Kendall

Odontesthes is diagnosed by seven characters of which three are unique to the family. It is the genus with most species (19 recognized species) and most ample distribution, in marine coastal waters and temperate freshwater drainages of South America (Fig. 4). Marine forms are distributed from the austral tip of Tierra del Fuego to Piura, Peru, including the Juan Fernandez Archipelago in the southeastern Pacific, and to Santos, Brazil, including the Falkland or Malvinas Islands in the southwestern Atlantic. Freshwater forms are extended from Patagonia to La Serena, IV Region, Chile, and to Rio Grande do Sul, southern Brazil.
Of the eleven available generic names that are junior synonyms of Odontesthes, only Austromenidia (Hubbs, 1918) and Cauque (Eigenmann, 1928) are recognized at present because, based on the hypothesis of relationships, they are the only clades with two or more species. Other generic names available such as Austroatherina (Marrero, 1950), Kronia (Miranda- Ribeiro, 1915), Patagonina (Eigenmann, 1928), Tupa and Yaci (de Buen, 1953) have a single species (monotypic) and there is no sense in recognizing them as such. Furthermore, if they were to be recognized as such, at least three new genera would have to be described for species such as O. humensis, O. bonariensis and O. argentinensis. This situation could change for Kronia if the morphological, ecological and molecular information (Brugger et al., 1990; Phonlor and Cousin, 1997; Bemvenuti, 1993; Beheregaray and Levy, 2000; Beheregaray and Sunnucks, 2001) were to form a pattern sufficient to propose an estuarine species and a marine species for what is now recognized as divergent populations of O. argentinensis.

Subgenus Odontesthes Evermann & Kendall

The subgenus Odontesthes includes the type species O. perugiae and O. orientalis, both inhabitants of the Uruguay and La Plata rivers and their tributaries, and another four species in the Mirim and dos Patos lagoons (O. mirinensis; Bemvenuti, 1995), and Tramanda’ River system (O. bicudo, O. ledae, O. piquava; Malabarba and Dyer, 2002). A molecular phylogeny based on microsatellite markers and mitochondrial DNA proposes that the species of the Mirim-Patos- Tramandaí system conform a monophyletic group, however not the three endemic species from the Tramandaí River system (Beheregaray et al., 2002). Three new species are in the process of being described from the northern part of the dos Patos Lagoon, RS, Brazil (Malabarba, pers. com.) and a phylogenetic analysis of the subgenus is required.

Subgenus Austromenidia Hubbs

Austromenidia is diagnosed by five characters and is composed of three marine species: in the Pacific from Piura, Peru, to Aysen, XI Region (O. regia), and the Juan Fernandez Archipelago (O. gracilis), and in the Atlantic from Mar del Plata, Argentina, to the Straits of Magellan reaching up to Puerto Natales, including the Falkland or Malvinas Islands (O. smitti). Austromenidia was described by Hubbs (1918) and characterized by species that had a combination of upper jaw protrusion, small scales, and an anterior position of the first dorsal fin. It was distinguished in this way from Basilichthys that has no upper jaw protrusion, and from Odontesthes and Kronia that have large scales and a posterior position of the first dorsal fin. Odontesthes hatcheri, O. nigricans, and the subgenus Cauque also fit into this definition of Hubbs, and were included in Austromenidia by Schultz (1948). However, based on the hypothesis of relationships presented in Figure 3, O. hatcheri could not be included in the subgenus Austromenidia, and the species of the subgenus Cauque and O. nigricans are potential candidates. The characters supporting the relationships among the species of Cauque, O. nigricans and O. incisa, O. platensis and Austromenidia are in conflict among themselves and more evidence is required to confirm or propose a new set of relationships (Dyer and Gosztonyi, 1999). It is for this reason that the species now included in Austromenidia are only those for which a substantial amount of evidence is available to justify the monophyly of Austromenidia, that is O. regia, O. gracilis and O. smitti (Dyer and Gosztonyi, 1999). This does not preclude the possibility that other species be included as long as a solid base of characters supports the monophyly of the group.

Subgenus Cauque Eigenmann

Cauque is diagnosed by seven characters, is endemic of South-Central Chile, and is present in rivers and estuaries from La Serena to Chiloé (O. brevianalis), and in lakes and deep rivers from Maule River to Llanquihue Lake (O. mauleanum). The identity and relationships of the species require an exhaustive revision. Tentatively only two species are recognized: O. brevianalis, a predominantly estuarine species and in the lower courses of the rivers; and O. mauleanum, a predominantly lacustrine species and in deep rivers. Odontesthes wiebrichi is considered as a possible hybrid between O. brevianalis and O. regia because in addition to having intermediate meristic counts between both species, it is collected only in the Valdivia River mouth (Corral), together with the above mentioned species. The type specimens of Odontesthes itatanum have meristic counts of scales and vertebrae much higher than the other species, for which there is some evidence that it is a valid species. More specimens from the Itata River are required to verify and compare them to the types.

Other species

Odontesthes hatcheri is present in lakes and rivers of Patagonia, including the Argentine precordilleran drainages of San Juan and Mendoza, as far south as Santa Cruz River. It is found in rivers of southern Chile that extend into Argentina, from Puelo River to Baker River. This species is characterized by having a ventral mouth, molariform pharyngeal teeth, small scales with a black posterior margin, a haemal funnel without expansions, and 21-27 gill rakers on the lower limb of the first arch. This species hybridizes with O. bonariensis (see Odontesthes bonariensis).
Odontesthes humensis is present in lakes and large bodies of water of the drainages of the La Plata and Uruguay rivers, dos Patos and Mirim lagoons. The main difference between O. humensis and O. guazu is in the pectoral fin extending or not to the base of the pelvic fin (de Buen, 1953), a condition quite variable in other species of silversides. This species is characterized by having a ventral mouth, 20 to 24 short and thick gill rakers on the lower limb of the first arch, and molariform pharyngeal teeth that provide it with a benthic diet, mainly mollusks (Bemvenuti, 2004). This species hybridizes with O. bonariensis (see Odontesthes bonariensis).
Odontesthes retropinnis is present in lakes and slow moving rivers of the La Plata and Uruguay rivers, and Mirim lagoon drainages. This species is characterized by having small mandibular teeth and 45 to 60 gill rakers of the lower limb, the highest number in all of South American silversides. The absence of expansions of the haemal funnel (also in O. hatcheri) is the feature de Buen (1953) used to create the genus Yaci.
Odontesthes bonariensis has its origins in lakes and lagoons of the Province of Buenos Aires, Argentina, and Rio Grande do Sul, Brazil. Despite there are no records of O. bonariensis being native to Uruguay, the populations of this species in dos Quadros and dos Patos lagoons in Brazil are presumed to be native because there are no records of introductions. This is a euryhaline species, with 30 to 40 gill rakers on the lower branch. Specimens of this species have the largest registered size (52 cm LS) than any other atherinopsid or atheriniform. The aquaculture of this species begun in Chascomús Lake, Argentina, in 1904 (Evermann and Kendall, 1906; Valette, 1939; Berasain et al. 2004), in dos Quadros Lagoon, RS, Brazil since 1943 (Kleerekoper, 1945), in Japan since 1966 (Ohashi, 2004), and in Italy since 1974 (Tortonese, 1985). This species was introduced during the 1940's into Chile (Riegel, 1960) and into Bolivia in the late 1940's entering Lake Titicaca in 1955 or 1956 (Calsina-Cota,
Studies on the reproductive biology of this species have determined not only the ages of sexual differentiation but also evidence of temperature dependent sex determination (Strüssmann and Patiño, 1995; Strüssmann et al., 1996a, b, c, 1997b ). Also, an efficient way of inducing triploidy has been developed, but its use for aquaculture purposes is still to be determined (Strüssmann et al., 1993). Hybrids of this species with O. hatcheri are found in some lakes of the Argentine Patagonia (ej., Pellegrini Lagoon), partly due to the artificial insemination practices carried out by aquaculturists for stocking purposes (M. Amalfi, 1990, C. Strüssmann, 2004 com. pers) 3. The same would be happening with O. humensis in the southern part of the dos Patos Lagoon, Brazil (Bemvenuti, 2004) , possibly related to the aquaculture station at Pelotas. In Japan, where the aquaculture of O. bonariensis and O. hatcheri is well developed, both species have hybridized spontaneously and genetic markers have been developed for their identification and differentiation (Yoshizaki et al., 1997; Strüssmann et al., 1997a).
Odontesthes argentinensis is considered at present as a widely distributed western Atlantic coastal species, in marine and estuarine environments from the Province of Sao Paulo, Brazil, to south of Chubut, Argentina. It is distinguished by having crenulate predorsal scales and 26 to 30 gill rakers on the lower branch (see key here and in Malabarba and Dyer, 2002). Numerous studies of this species in southern Brazil (Bemvenuti, 2004) have compared the estuarine and the marine groups in terms of egg morphology (Phonlor and Cousin, 1997), morphometrics (Bemvenuti, 1993; Cuello and Garc’a, 2004), electrophoresis (Beheregaray and Levy, 2000) and DNA (Beheregaray and Sunnucks, 2001). Nevertheless, the differences found are not conclusive to propose two species as did de Buen (1953) with Kronia rex and K. alba.
Odontesthes incisa is a western Atlantic coastal marine species, distributed from Rio Grande do Sul, Brazil, to Santa Cruz, Argentina. It is characteristically a small species (known as "cornalito") with large crenate4 scales on all of body and canine-type teeth on both mandibular jaws. The osteology of this species was described in detail by Pianta de Risso and Risso (1953) in a journal of limited distribution and placed in a new genus and subfamily.
Odontesthes nigricans is a southwestern Atlantic species, with estuarial incursions for reproduction, distributed from the Province of Buenos Aires to Cape Horn and Falklands or Malvinas Islands. It is distinguished for having small and crenate scales, less than 20 gill rakers on the lower branch, and the first dorsal fin placed anteriorly over the base of the pelvic fin. In two studies of the axial skeleton of marine silversides (Piacentino and Torno, 1987; Piacentino, 1990), this species was described as having a haemal funnel of the type that is diagnostic of the subgenus Austromenidia. Radiographs of the type specimens of O. nigricans and O. alburnus clearly show an absence of a haemal funnel for this species.
Odontesthes platensis is a western Atlantic coastal species, distributed from La Plata River to the south of Chubut, Argentina. Known as "panzón" or big belly, this is a species that is not frequent in collections, though apparently very abundant towards the beginning of the 20th century (Garc’a, 1998).


Placing the hypothesis of evolutionary relationships of the South American atherinopsines over the distribution map of the species (Fig. 4), two biogeographic hypotheses can be proposed: a) the elevation of the Andes during the middle to late Miocene was the vicariant process
that separated Basilichthys (to the West) and Odontesthes (to the East), this coinciding with the earliest fossil remains of Basilichthys found in Chile (Rubilar, 1994) and Odontesthes (Cione, ; and b) Sorgentinini is primitively a freshwater group, because the basal species of both Basilichthys and Odontesthes are primarily in freshwater and, consequently the marine silversides of Odontesthes are derived, not primitive as was the traditional assumption (Lahille, 1929a,b; White, 1986, among others).
Contrary to the assumptions of White (1985, 1986) the biogeographic analysis of Sorgentinini indicates that this group is primitively freshwater. Basilichthys and the basal species of Odontesthes are freshwater (Dyer, 1998; Fig. 4), being marine a derived condition within the group (Dyer, 1998). The use of freshwater as a character is ambiguous because the sister group of Sorgentinini (Atherinopsini) is marine and the primitive condition of Menidiinae is ambiguous.

Artificial identification key for South American silverside species of the southern cone (based only characters present in adult specimens)

FIGURE 5. Dorsal view of the head of Basilichthys; A, B. micolepidotus ; B, B. semotilus.

FIGURE 6. Lateral view of the snout region; A, Odontesthes perugiae; B, Basilichthys semotilus.

FIGURE 7. Lateral view of median fins and axial skeleton; A, Odontesthesretropinnis; B, Odontesthes bonariensis; C, Odontesthes regia.

FIGURE 8. Scales posterior border up; A, Odontesthes humensis, smooth; B, O. argentinensis, crenulate; C, O. incisa, crenate. Figures modified from de Buen (1953).

FIGURE 9. Ventral and lateral views of the urohyal; A, O. regia; B, O. smitti.


List of atherinopsine species present in South America.
Museum acronyms follow Leviton et al. (1985). Family Notocheiridae Schultz, 1948
Genus Notocheirus Clark, 1937
            Type species: Notocheirus hubbsi Clark, 1937 {CAS-SU 5525}
                        Type locality: Valparaiso Bay, Chile.
Family Atherinopsidae Fowler, 1903
Subfamily Menidiinae Schultz, 1948
Tribe Membradini Chernoff, 1986
Genus Atherinella Steindachner, 1875
            Type species: Atherinella panamensis Steindachner, 1875:477 {NMW 76439}
            =Eurystole Jordan & Evermann en Jordan, 1895: 418
            Type species: Atherinella eriarcha Jordan & Gilbert, 1882: 348
            =Thyrina Jordan & Culver en Jordan, 1895: 419 (preoccupied)
            Type species: Thryrina evermanni Jordan & Culver en Jordan, 1895:419
            =Melaniris Meek, 1902: 117
            Type species: Melaniris balsana Meek, 1902:117
            =Xenatherina Regan, 1907: 64
            Type species: Menidia lisa Meek, 1904:182
            =Thyrinops Hubbs, 1918: 306
            Type species: Atherinichthys pachylepis Günther, 1864:25
            =Archomenidia Jordan & Hubbs, 1919: 54
            Type species: Atherinichthys sallei Regan, 1903: 60
            =Nectarges Myers & Wade, 1942: 126
            Type species: Nectarges nepenthe Myers & Wade, 1942: 130
            =Euryarges Myers & Wade, 1942: 128
            Type species: Nectarges nesiotes Myers & Wade, 1942: 128
            =Coleotropis Myers & Wade, 1942: 136
            Type species: Menidia starksi Meek & Hildebrand, 1923: 267
            =Xenomelaniris Schultz, 1948: 13, 33
            Type species: Atherina brasiliensis Quoy & Gaimard, 1825: 332
            =Atherthyrina Fowler, 1958: 16
            Type species: Thyrina evermanni Jordan & Culver en Jordan, 1895: 419
            =Allomastax Chernoff, 1986a: 243
            Type species: Melaniris sardina Meek, 1907: 114Atherinella (Eurystole) nocturna (Myers & Wade, 1942)
            Eurystole nocturna Myers & Wade, 1942 {USNM 88712}
            Type locality: Guayaquil, Ecuador, probably opposite Estero Guayas.
Atherinella (Xenomelaniris) brasiliensis (Quoy & Gaimard, 1825)
            Atherina brasiliensis Quoy & Gaimard, 1825: 332 {MNHN A.4374}
            Type locality: Rio de Janeiro Bay, Brazil. Subfamily Atherinopsinae Fowler, 1903
Tribe Sorgentinini Pianta de Risso & Risso, 1953
Genus Basilichthys Girard, 1855
            Type species: Atherina microlepidota Jenyns, 1841
            =Protistius Cope, 1874: 66
            Type species: Protistius semotilus Cope, 1874
            =Gastropterus Cope, 1878: 700
            Type species: Gastropterus archaeus Cope, 1878
            =Pisciregia Abbott, 1899: 342
            Type species: Pisciregia beardsleei Abbott, 1899 Species group microlepidotus de especies microlepidotus
Basilichthys microlepidotus (Jenyns, 1841)
            Atherina microlepidota Jenyns, 1841: 78 {BMIII 403}
                        Type locality: Valparaiso, Chile.
Basilichthys australis Eigenmann, 1928
            Basilichthys australis Eigenmann, 1928: 59
            {CAS-SU 11678, 44699, 44703, 45179, 45180, 45182-45184, 45188-45190, 45192}
                        Type locality: from Maipo River to Rahue River, Chile. Species group semotilus
Basilichthys semotilus (Cope, 1874) de especies semotilus
            Basilichthys semotilus (Cope, 1874): 66 {ANSP 14404}
                        Type locality: Peruvian Andes, at 4.000 m altitude.
            =B. beardsleei (Abott, 1899): 342 {CAS-SU 11961}
                        Type locality: Callao, Lima, Peru.
Basilichthys archaeus (Cope, 1878)
            Basilichthys archaeus (Cope, 1878):700 {ANSP 22002, 22003}
                        Type locality: Arequipa, Peru, at 2.500 m altitude.
Basilichthys sp.
                        Locality: Loa and Codpa rivers, Chile. Genus Odontesthes Evermann & Kendall, 1906
            Type species: Odontesthes perugiae Evermann & Kendall, 1906: 94
            =Kronia Miranda-Ribeiro, 1915: 9
            Type species: Kronia iguapensis Miranda-Ribeiro, 1915: 10
            =Pseudothyrina Miranda-Ribeiro, 1915: 11
            Type species: Pseudothyrina jheringi Miranda-Ribeiro, 1915: 11
            =Austromenidia Hubbs, 1918: 307
            Type species: Basilichthys regillus Abbott, 1899: 339
            =Cauque Eigenmann, 1928: 56
            Type species: Chirostoma mauleanum Steindachner, 1896: 231
            =Patagonia Eigenmann, 1928: 56 (typographic error of Patagonina)
            =Patagonina Eigenmann, 1928: 56
            Type species: Menidia hatcheri Eigenmann, 1909
            =Austroatherina Marrero, 1950: 113
            Type species: Atherina incisa Jenyns, 1841 (available by subsequent designation)
            =Bachmannia Nani in Szidat & Nani, 1951: 336 (preoccupied in Siluriformes)
            Type species: Basilichthys smitti Lahille, 1929a
            =Sorgentinia Pianta de Risso & Risso, 1953: 13
            Type species: Atherina incisa Jenyns, 1841
            =Tupa De Buen, 1953: 48
            Type species: Atherinichthys platensis Berg, 1895
            =Yaci De Buen, 1953: 51
            Type species: Yaci retropinnis De Buen, 1953Odontesthes hatcheri (Eigenmann, 1909)
            Menidia hatcheri Eigenmann, 1909: 281 {types lost}
                        Type locality: Pueyrredón Lake, Santa Cruz, Argentina.
            =?Basilichthys cuyanus Burmeister, 1861: 534 {types lost}
                        Type locality: Guanacache Lagoon, San Juan, Argentina.
            =Basilichthys andinus Lahille, 1929b: 324 {types lost}
                        Type locality: Traful Lake, Rio Negro, Argentina.
            =Basilichthys patagonicus Marrero, 1950: 67 {types lost}
                        Type locality: Limay River, Rio Negro, Argentina. Odontesthes humensis de Buen, 1953
            Odontesthes humensis de Buen, 1953:34 {MNHN-M 1807}
                        Type locality: Negro River, Uruguay.
            =Odontesthes guazu de Buen, 1953: 40 {MNHN-M 1805}
                        Type locality: Uruguay River, Uruguay. Odontesthes retropinnis (de Buen, 1953)
            Yaci retropinnis de Buen 1953:52 {MNHN-M 1809}
                        Type locality: Negro River, Uruguay. Odontesthes bonariensis (Valenciennes, 1835)
            Atherina bonariensis Valenciennes in Cuvier & Valenciennes, 1835:469 {MNHN-P A.44 07}
                         Type locality: Buenos Aires, Argentina.
            =Atherina lichtensteinii Valenciennes in Cuvier & Valenciennes, 1835:476 {ZMB 1883}
                        Type locality: Montevideo, Uruguay.
            =Basilichthys chascomunensis Lahille, 1929b:305 {types lost}
                        Type locality: Buenos Aires, Argentina.
            =Basilichthys puntanus Lahille, 1929b:305 {types lost}
                         Type locality: San Luis, Argentina. Odontesthes argentinensis (Valenciennes, 1835)
            Atherina argentinensis Valenciennes in Cuvier & Valenciennes, 1835:472 {MNHN-P A.4362, A.4363}
                        Type locality: Montevideo, Uruguay.
            =?Atherina lessoni Valenciennes in Cuvier & Valenciennes, 1835:471 {types lost}
                        Type locality: Santa Catarina, Brazil.
            =Kronia iguapensis Miranda Ribeiro, 1915:10 {MNRJ 1351}
                        Type locality: Iguape, São Paulo, Brazil.
            =Pseudothyrina iheringi Miranda Ribeiro, 1915:10 {MNRJ 2369}
                        Type locality: Rio Grande do Sul, Brazil.
            =Basilichthys bonariensis charruanus Lahille, 1929b:319 {MACN 5178?}
                        Type locality: Patagonian coast, Mar del Plata, Buenos Aires, Argentina.
            =Basilichthys bonariensis propinquus Lahille, 1929b:320 {types lost}
                        Type locality: Buenos Aires, Argentina.
            =Menidia thomasii Meinken, 1931:377 {?KHMM}
                        Type locality: Uruguay.
            =Kronia rex de Buen, 1953:64 {MHNM 1803}
                        Type locality: La Paloma, Rocha, Uruguay.
            =Kronia alba de Buen, 1953:59 {MHNM 1801}
                        Type locality: Rocha Lagoon, Rocha, Uruguay. Subgenus Odontesthes Evermann & Kendall, 1906
Odontesthes perugiae Evermann & Kendall, 1906
            Odontesthes perugiae Evermann & Kendall, 1906:94 {USNM 55572}
                        Type locality: Argentina.
            =Basilichthys microather Marrero, 1950:75 {types lost}
                        Type locality: La Plata River, Argentina. Odontesthes orientalis de Buen, 1950
            Odontesthes orientalis de Buen, 1950:149 {MNHN-M 1808}
                        Type locality: Negro River, Uruguay.
Odontesthes mirinensis Bemvenuti, 1995
            Odontesthes mirinensis Bemvenuti, 1995:885 {MCP 17696}
            Type locality: Mirim Lagoon, RS, Brazil.

Odontesthes piquava Malabarba & Dyer, 2002
            Odontesthes piquava Malabarba & Dyer, 2002:261 {MCP 26152}
                        Type locality: da Pinguela Lagoon, Tramanda’ (29º49'S; 50º10'14"W), RS, Brazil.
Odontesthes bicudo Malabarba & Dyer, 2002
            Odontesthes bicudo Malabarba & Dyer, 2002: 264 {MCP 26153}
                        Type locality: Emboaba Lagoon, Osório (29º57'57"S; 50º13'45"W), RS, Brazil.
Odontesthes ledae Malabarba & Dyer, 2002
            Odontesthes ledae Malabarba & Dyer, 2002: 266{MCP 26151}
                        Type locality: Fortaleza Lagoon, Cidreira (30º09'33" S; 50º13'44" W), RS, Brazil.
Subgenus Cauque Eigenmann, 1928
Odontesthes mauleanum (Steindachner, 1896)
            Atherinichthys mauleanum Steindachner, 1896:231 {NMW 62506, 62507, 16979}
                        Type locality: Maule River, VII Region, Chile.
            =Atherinichthys itatanum Steindachner, 1896:232 {NMW 62608}
                        Type locality: Itata River, VIII Region, Chile.
            =Cauque molinae Fowler, 1940:183 {ANSP 69147}
                        Type locality: Malleco River at Angol, IX Region, Chile.
Odontesthes brevianalis (Günther, 1880)
            Atherinichthys brevianalis Günther, 1880:25 {MNHN 1890-119}
                        Type locality: Valparaiso, V Region, Chile.
            =Cauque wiebrichi Eigenmann, 1928:58 {CAS 49902}
                        Type locality: Valdivia, X Region, Chile.
            =Odontesthes (Cauque) debueni Fischer, 1962 {ZMH 1698}
                        Type locality: Estero Lenga, Concepcion, VIII Region, Chile.

Odontesthes nigricans (Richardson, 1848)
            Atherina nigricans Richardson, 1848:77 {BMNH 1848.3.10:29}
                        Type locality: Malvinas or Falkland Islands.
            =Atherinichthys alburnus Günther, 1861:404 {BMNH 1859.10.12:30.33}
                        Type locality: Straits of Magellan, Chile.
            =?Menidia patagoniensis Eigenmann, 1909:280 {types lost}
                        Type locality: ?Punta Arenas, Straits of Magellan, Chile.
            =?Basilichthys nigricans macropterus Lahille, 1929b:332 {types lost}
                        Type locality: Malvinas Islands and R’o Gallegos, Straits of Magellan, Argentina.
            =Basilichthys malvinensis Marrero, 1950:121 {types lost}
            Type locality: Southwest Atlantic, in Argentina from Pen’nsula Valdés to Los Estados Island and Malvinas Islands.

Odontesthes incisa (Jenyns, 1841)
            Atherina incisa Jenyns, 1841:79 {BMIII 405}
                        Type locality: 39º S; 61º W, several miles from continent; opposite Punta Sauce, Buenos Aires, Argentina.
            =Menidia uruguayensis Devincenzi, 1924:205 {MNHN-M 1804}
                        Type locality: La Plata River, Montevideo, Uruguay.

Odontesthes platensis (Berg, 1895)
            Atherinichthys platensis Berg, 1895:27 {MACN 5162, 5195}
                        Type locality: Mar del Plata, Argentina.

Subgenus Austromenidia Hubbs, 1918
Odontesthes regia (Humboldt, 1821)
            Atherina regia Humboldt in Humboldt & Valenciennes, 1821: 187 {types lost}
                        Type locality: Callao, Peru.
            =Atherina laticlavia Valenciennes in Cuvier & Valenciennes, 1835: 473 {MNHN-P 2980}
                        Type locality: Valparaiso, Chile.

=Chirostoma affine Steindachner, 1898: 313 {ZMB 15674}
                        Type locality: Iquique, Chile.
            =Basilichthys jordani Abbott, 1899: 341 {CAS-SU 6070}
                        Type locality: Callao, Peru.
            =Basilichthys octavius Abbott, 1899: 340 {CAS-SU 6069}
                        Type locality: Callao, Peru.
            =Basilichthys regillus Abbott, 1899: 339 {CAS-SU 6071}
                        Type locality: Callao, Peru.

Odontesthes gracilis (Steindachner, 1898)
            Chirostoma gracile Steindachner, 1898: 314 {ZMB 15675}
                        Type locality: Cumberland Bay, Robinson Crusoe Island, Juan Fernandez Archipelago, Chile.
Odontesthes smitti (Lahille, 1929a)
            Basilichthys smitti Lahille, 1929a: 84 {types lost}
                        Type locality: "Fin de Barrancas", Gulf of San Mat’as, SE Atlantic, Argentina.
            =Atherina jacksoniana Quoy & Gaimard, 1825: 333 {MNHN 3096, A.2895}
                        Type locality: original card: Port Jackson, NSW, Australia)
            ?=Menidia patagoniensis Eigenmann, 1909: 280 {types lost}
                        Type locality: Straits of Magellan (?Punta Arenas), Chile.
            =Basilichthys smitti var. australis Lahille, 1929a: 84 {types lost}
                        Type locality: R’o Gallegos, Argentina, and Puerto Natales, Chile.
            =Basilichthys madrynensis Lahille, 1929b: 326 {types lost}
                        Type locality: Port Madryn, Golfo Nuevo, Chubut, Argentina.


1. Phylogenetic relationships among families are based on a data matrix of 83 morphological characters and 31 taxa (Dyer and Chernoff, 1996).
2. The phylogenetic relationships among genera and species of Atherinopsinae are based on a data matrix of 25 taxa and 123 characters, analyzed using a cladistic methodology based on parsimony in PAUP (Swofford, 1993) and MacClade (Maddison and Maddison, 1992). Characters were polarized by multiple-outgroup comparison (Farris, 1982; Maddison et al., 1984; Clark and Curran, 1986).
3. Lic. Marina Amalfi, Cinco Saltos, Prov. R’o Negro, Argentina.
4. Scale nomenclature follows that of Roberts (1993).
5. Alberto Cione, Div. Paleontologia, Museo La Plata, Argentina.
6. The dorsal scales were counted at the level of the pectoral fin's distal half, from side to side over the dorsum, beginning and ending on the row of scales that form half-part of the lateral band. This meristic is considered to represent scale size.


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Received on April 21, 2005.
Accepted on May 11, 2005.

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