Two new Neotropical species of Drosophila peruensis species group (Diptera, Drosophilidae)

Duas novas espécies neotropicais do grupo de espécies Drosophila peruensis (Diptera, Drosophilidae)

Jonas S. Döge Marco S. Gottschalk Vera Lúcia S. Valente About the authors

Abstracts

The Drosophila peruensis species group was recently proposed and includes four taxa: D. atalaia Vilela & Sene, 1982, D. boraceia Vilela & Val, 2004, D. pauliceia Ratcov & Vilela, 2007, and D. peruensis Wheeler, 1959. All these species have most of setae or setulae of mesonotum arinsing from dark spots, wings with crossveins darker (except in D. atalaia) and hypandrium squared-shaped mostly fused to gonopods. Here, we describe two new species, Drosophila itacorubi sp. nov. and Drosophila paraitacorubi sp. nov., belonging to this species group. The male genitalia of these species are figured. An identification key to the D. peruensis species group is provided.

Atlantic Rainforest; male genitalia; mangrove; southern Brazil; spotted thorax


O grupo de espécies Drosophila peruensis foi proposto recentemente e inclui quatro táxons: D. atalaia Vilela & Sene, 1982, D. boraceia Vilela & Val, 2004, D. pauliceia Ratcov & Vilela, 2007, D. peruensis Wheeler, 1959. Todas possuem a maioria das cerdas do mesonoto emergindo de manchas escuras, veias transversais esfumaçadas (com exceção de D. atalaia) e hipândrio com formato quadrado e fortemente fusionado aos gonópodes. Neste artigo são descritas duas novas espécies, Drosophila itacorubi sp. nov. e D. paraitacorubi sp. nov., pertencentes a este grupo de espécies. A genitália masculina destas espécies são ilustradas. Uma chave de identificação para as espécies do grupo D. peruensis é fornecida.

Genitália masculina; manguezal; Mata Atlântica; Sul do Brasil; tórax manchado


Two new Neotropical species of Drosophila peruensis species group (Diptera, Drosophilidae)

Duas novas espécies neotropicais do grupo de espécies Drosophila peruensis (Diptera, Drosophilidae)

Jonas S. DögeI; Marco S. GottschalkII; Vera Lúcia S. ValenteIII

IPrograma de Pós-Graduação em Ecologia, Universidade de Brasília (UnB), Campus Darcy Ribeiro, IB, GEM, Laboratório de Biologia Evolutiva, 70910-900, Brasília, DF, Brazil. (jsdoge@gmail.com)

IIInstituto de Ciências Biológicas, Universidade Federal do Rio Grande, Avenida Itália, Km 08, Campus Carreiros, 96201-900, Rio Grande, RS, Brazil. (marcogottschalk@furg.br)

IIIDepartamento de Genética, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Prédio 43323, Sala 210, Agronomia, Caixa Postal 15053, 91501-970, Porto Alegre, RS, Brazil. (vera.gaiesky@ufrgs.br)

ABSTRACT

The Drosophila peruensis species group was recently proposed and includes four taxa: D. atalaia Vilela & Sene, 1982, D. boraceia Vilela & Val, 2004, D. pauliceia Ratcov & Vilela, 2007, and D. peruensis Wheeler, 1959. All these species have most of setae or setulae of mesonotum arinsing from dark spots, wings with crossveins darker (except in D. atalaia) and hypandrium squared-shaped mostly fused to gonopods. Here, we describe two new species, Drosophila itacorubi sp. nov. and Drosophila paraitacorubi sp. nov., belonging to this species group. The male genitalia of these species are figured. An identification key to the D. peruensis species group is provided.

Keywords: Atlantic Rainforest, male genitalia, mangrove, southern Brazil, spotted thorax.

RESUMO

O grupo de espécies Drosophila peruensis foi proposto recentemente e inclui quatro táxons: D. atalaia Vilela & Sene, 1982, D. boraceia Vilela & Val, 2004, D. pauliceia Ratcov & Vilela, 2007, D. peruensis Wheeler, 1959. Todas possuem a maioria das cerdas do mesonoto emergindo de manchas escuras, veias transversais esfumaçadas (com exceção de D. atalaia) e hipândrio com formato quadrado e fortemente fusionado aos gonópodes. Neste artigo são descritas duas novas espécies, Drosophila itacorubi sp. nov. e D. paraitacorubi sp. nov., pertencentes a este grupo de espécies. A genitália masculina destas espécies são ilustradas. Uma chave de identificação para as espécies do grupo D. peruensis é fornecida.

Palavras-chave: Genitália masculina, manguezal, Mata Atlântica, Sul do Brasil, tórax manchado.

Since the beginning of the twentieth century the male genitalia of drosophilids was considered important to separate sibling species (Sturtevant, 1921) and several authors described this apparatus (Hsu, 1949). In the middle of that century, Hsu (1949) compared the external genitalia of about 170 species and concluded that close related species shows a striking similarity and this trait should be considered important to systematics. However, this author noticed that this apparatus was insufficient to distinguish some couples of species and, on that basis, the internal structures of the male genitalia became drawn (Frota-Pessoa, 1954).

Nowadays, descriptions in the field of taxonomy have been focused on both internal and external morphology of the male genitalia (Vilela, 1983). It was observed that, on the morphological level, many of the species of Drosophila could be distinguished only by the analysis of such structure and mainly by the internal genitalia (aedeagus) (Vilela, 1992). For this reason, the male genitalia, and specially the aedeagus, became the most important character used by taxonomists to recognize and describe species (Vilela & Bächli, 1990). The use of this structure as a morphological character to delimit species is considered effective because, in a large number of animal groups, are under strong selection and evolve rapidly (Eberhard, 1985).

The Drosophila peruensis species group was recently proposed by Ratcov & Vilela, (2007) to include four species: D. atalaia Vilela & Sene, 1982, D. boraceia Vilela & Val, 2004, D. pauliceia Ratcov & Vilela, 2007 and D. peruensis Wheeler, 1959. The first described species, D. peruensis, was formerly described as D. maculipennis by Duda (1927) based on a single female and ascribed to D. repleta species group by Sturtevant (1942) probably due to it spotted thorax. Later, the absence of known males and the misidentification of specimens of D. tucumana Vilela & Pereira, 1985 and D. urubamba Vilela & Pereira, 1993 (both belonging to the D. guarani species group) as D. peruensis has caused it to be included in the D. guarani species group by Vilela (1983). In more recent analysis, Ratcov & Vilela (2007) noted that D. peruensis is closed related to D. atalaia, D. boraceia and to an undescribed species (actually described as D. pauliceia), and assigned them to a new group, the D. peruensis.

Up to now, the D. peruensis group was recorded only in South America. Drosophila peruensis were recorded in Peru and Colombia (Duda, 1927; Vilela & Bächli, 1990), D. boraceia and D. pauliceia in Brazil (states of São Paulo and Santa Catarina) (Vilela & Val, 2004; Gottschalk et al., 2007; Ratcov & Vilela, 2007; Döge et al., 2008), and D. atalaia in Brazil (states of Espírito Santo, Rio de Janeiro and Mato Grosso do Sul), Argentina and El Salvador (Vilela & Sene, 1982).

This paper presents the description of two new species from the D. peruensis species group, focusing on external morphology and traits of the internal and external genitalia, and proposes an identification key to the D. peruensis species group.

MATERIAL AND METHODS

The postabdomen of specimens were removed and dissected and male and female genitalia are stored in microtubes with glycerol and attached to their respective specimens. The material studied is deposited in the Museu de Ciências Naturais of the Fundação Zoobotânica do Rio Grande do Sul (MCNZ, Porto Alegre, Brazil). Pigmentation of specimens was observed on dried individuals. The terminology was based on Vilela & Bächli (2000) and indexes on Vilela & Bächli (1990) and Vilela (1983). Averages are followed by ranges (in parenthesis) when two or more specimens were analyzed. Illustrations were drawn using a microscope with a 20x magnification object lens and a camera lucida.

Drosophila itacorubi sp. nov.

(Figs. 1, 3, 5, 7-15 )




  •  


  • Drosophila sp. 1 Schmitz et al., 2007:57 (tab. 1), 61.

    Type material. Holotype (postabdomen dissected), labelled "Brasil. SC. Florianópolis. Manguezal do Itacorubi, 27º34'14"S 48º30'33"W, H. J. Schmitz col. 22.IV.2005 / Drosophila itacorubi Döge, Gottschalk & Valente Holótipo". Paratypes: 2 labeled "Brasil. SC. Florianópolis. Manguezal do Itacorubi, 27º34'14"S 48º30'33"W, H. J. Schmitz col. 22.IV.2005 / Drosophila itacorubi Döge, Gottschalk & Valente Parátipo" and 2 labeled "Brasil. SC. Florianópolis. Manguezal do Rio Tavares, 27º39'13"S 48º32'12"W, H. J. Schmitz col. 21.IV.2005 / Drosophila itacorubi Döge, Gottschalk & Valente Parátipo", one of them with the postabdomen dissected. Type locality: Manguezal do Itacorubi, Florianópolis, state of Santa Catarina, Brazil.

    Diagnosis. Main color yellowish brown. Frons yellowish brown; arista with 5-6 dorsal and 2-3 ventral branches, plus terminal fork. Most setae and setulae of the thorax and head arising from dark brown spots; two large prescutellars. Wings with both main crossveins darker than longitudinal ones, with lappet (not so evident). Surstylus mostly microtrichose; hypandrium somewhat square-shaped; aedeagus stout.

    Description. . Head (Fig. 1). Main color yellowish brown. Frons yellowish brown. Frontal length = 0.50 mm. Frontorbital plates yellowish brown, with dark brown spots at base of orbital setae. Anterior reclinate orbital setae behind proclinate one. Length ratio of mid to anterior orbital = 0.76. Length ratio of mid to posterior orbital = 0.76. Vt index = 1.02. Frontal triangle dark brown, medially interrupted by a yellowish brown strip. Ocellar triangle yellowish brown with dark brown spots around the ocelli, about 53.1% of frontal length. Face brown. Facial carina prominent, sulcate, and slightly darker than face. Antennae, proboscis, palps and gena yellowish brown. Palps clef-shaped with two prominent setae, one terminal and another subterminal. Red eyes, with short black pile. Cheek index = 9.86. Eye index = 1.44. Aristae with 5-6 dorsal and 2-3 ventral branches, plus terminal fork.

    Thorax (Fig. 3). Ground color yellowish brown, with dark brown spots at the base of each setae. Length = 1.57 mm. Width = 1.04 mm. Eight irregular rows of acrostichals. Two large prescutellars. Dc index = 0.58. Scutellum dark brown with one central spot at the base, and five little spots in the marginal area. Basal scutellar setae convergent. Scut position index = 0.79. Scut index = 0.76. Two prominent katepisternal with a row of setulae between both. Anterior katepisternal about 84.8% of the posterior one. Pleura brown with yellowish spots, somewhat fused. Halteres light brown. Trochanter, coxae and femur brown, tibia yellowish brown and tarsi yellow.

    Wing (Fig. 5). Main color light brown. Crossveins slightly darker than longitudinal veins. Lappet present but not so evident. Length = 2.39 mm. Length to width ratio = 1.83. Indices: C = 3.15; 4v = 1.29; 5x = 1.05; 4c = 0.67; M = 0.41; hb = 0.39; prox. x = 0.70.

    Abdomen. Ground color yellowish brown. Tergites with broad posterior dark brown bands enlarged paramedially (but medially interrupted in tergite 2), which bends towards and reaches anterior margin at angle of tergites, leaving a yellowish area laterally. Body length = 3.15 mm.

    Terminalia (Figs 7-14). Epandrium mostly microtrichose. Ventral lobe not covering surstylus. Cerci mostly microtrichose, anteriorly fused with epandrium at lower half (Fig. 7). Tip of cerci with about seven small setae. Surstylus mostly microtrichose with 17 prensisetae, about nine long outer and ten long inner setae (Fig. 8). Hypandrium square-shaped, shorter than epandrium (Fig. 9). Dorsal arch absent. Gonopod mostly fused with hypandrium. Aedeagus expanded at extremity, rounded at tip and with a little invagination, and slightly curved in profile (Figs 10-14). Dorsal cleft partially covered with a membranous tissue and showing a reduced anterior opening, close to the fusion of aedeagus and aedeagal apodeme. Aedeagal apodeme shorter than aedeagus, flattened laterally and curved in profile. Ventral rod robust and fused with aedeagal apodeme. Paraphysis slightly pointed at distal portion and proximally rounded, with one large and curved subapical setae.

    . Differences from male: The two prominent setae of palps smaller. Posterior band in Tergite 1 medially interrupted in one female. Measurements: Frontal length = 0.61 (0.59 - 0.61) mm. Length ratio of mid to anterior orbital = 0.69 (0.56 - 0.88). Length ratio of mid to posterior orbital = 0.65 (0.59 - 0.76). Vt index = 1.04 (0.93 - 1.12). Ocellar triangle about 57.4% (54.5 - 62.0%) of frontal length. Cheek index = 9.90 (9.07 - 11.06). Eye index = 1.30 (1.27 - 1.32). Thorax length = 1.74 mm (1.55 - 1.89). Thorax width = 1.39 mm (1.26 - 1.63). Dc index = 0.70 (0.64 - 0.73). Scut position index = 0.69 (0.59 - 0.80). Scut index = 0.92 (0.88 - 0.99). Anterior katepisternal about 76.1% (68.8 - 87.7%) of the posterior one. Wing length = 3.05 mm (2.92 - 3.14). Length to width ratio = 2.05 (2.00 - 2.08). Indices: C = 3.40 (3.20 - 3.76); 4v = 1.44 (1.32 - 1.55); 5x = 0.87 (0.85 - 0.89); 4c = 0.72 (0.60 - 0.79); M = 0.39 (0.33 - 0.45); hb = 0.50 (0.43 - 0.61); prox. x = 0.58 (0.56 - 0.61). Body length = 3.87 mm (3.66 - 4.05).

    Terminalia (Fig. 15). Valves of oviscapt apically rounded with about 17 marginal and seven discal heavily sclerotized, stout and peg-shaped ovisensilla (the three discal ovisensillas closer to the valves apex are setaeshaped but not so large). One large and two tiny setae at apex of valves.


    Etymology. The species name refers to the type locality Manguezal do Itacorubi.

    Drosophila paraitacorubi sp. nov.

    (Figs. 2, 4, 6, 16-21 )




  •  

    Drosophila sp.bo3 Döge et al., 2008:620 (tab. II).

    Type material. Holotype (postabdomen dissected), labelled "Brasil. SC. Joinville. Piraí. 26º17'15"S 49º01'00"W, J. S. Döge col. 27.II.2005 / Drosophila paraitacorubi Döge, Gottschalk & Valente Holótipo". Type locality: Piraí, Joinville, state of Santa Catarina, Brazil.

    Diagnosis. Main color yellowish brown. Frons yellowish brown, arista with 6 dorsal and 2 ventral branches, plus terminal fork. Most setae and setulae of the thorax and head arising from dark brown spots; basal scutellar setae convergent; two large prescutellars. Wings with both main crossveins darker than longitudinal ones, with lappet (not so evident). Surstylus mostly microtrichose; hypandrium somewhat square-shaped; aedeagus stout. Female unknown.

    Description. . Head (Fig. 2). Main color yellowish brown. Frons yellowish brown. Frontal length = 0.54 mm. Frontorbital plates yellowish brown, with dark brown spots at base of orbital setae. Anterior reclinate orbital setae behind proclinate one. Ratio of mid to anterior orbital = 0.93. Ratio of mid to posterior orbital = 0.72. Vt index = 1.11. Frontal triangle dark brown, medially interrupted by a yellowish brown strip. Ocellar triangle yellowish brown with dark brown spots around the ocelli, about 72.45% of frontal length. Face brown. Facial carina appears prominent, sulcate, and slightly darker than face. Antennae, proboscis, palps and gena yellowish brown. Palps clef-shaped with two prominent setae, one terminal and another subterminal. Red eyes, with short black pile. First genal setae about half length of the vibrissae, and stout. Cheek index = 7.89. Eye index = 1.43. Aristae with 6 dorsal and 2 ventral branches, plus terminal fork.

    Thorax (Fig. 4). Ground color yellowish brown, with dark brown spots at the base of each setae. Length = 1.65 mm. Width = 1.18 mm. Eight irregular rows of acrostichals. Two large prescutellars. Dc index = 0.61. Scutellum dark brown with one central spot at the base, and about five little spots in the marginal area. Basal scutellar setae convergent. Scut position index = 0.68. Two prominent katepisternal with a row of setulae between them. Anterior katepisternal about 75.16% of the posterior one. Pleura brown with yellowish spots, somewhat fused. Halteres light brown. Trochanter, coxae and femur brown, tibia yellowish brown and tarsi yellow.

    Wing (Fig. 6). Main color light brown. Crossveins slightly darker than longitudinal veins. Lappet present but not so evident. Length = 2.86 mm. Length to width ratio = 2.22. Indices: C = 3.37; 4v = 1.38; 5x = 0.79; 4c = 0.67; M = 0.34; hb = 0.48; prox. x = 0.64.

    Abdomen. Ground color yellowish brown. Tergites with broad posterior dark brown bands paramedially enlarged (but medially interrupted in tergites 2 and 3), which bends towards and reaches anterior margin at angle of tergites, leaving a laterally yellowish area. Body length = 3.27 mm.

    Terminalia (Figs 16-21 ). Epandrium mostly microtrichose, no upper setae, 11 lower setae (Fig. 16 ). Ventral lobe not covering surstylus. Cerci mostly microtrichose, anteriorly fused with epandrium at lower half. Tip of cerci with seven small setae. Surstylus mostly microtrichose with 18 prensisetae, about eight long outer and seven long inner setae. Hypandrium square-shape, shorter than epandrium (Fig. 17 ). Dorsal arch absent. Gonopods mostly fused with hypandrium. Aedeagus expanded at extremity, slightly rounded at tip and with a little invagination, and curved in profile (Figs. 18-21 ). Dorsal cleft partially covered with a membranous tissue and showing a reduced anterior opening, close to the fusion of aedeagus and aedeagal apodeme. Aedeagal apodeme shorter than aedeagus, flattened laterally and curved in profile. Ventral rod robust and fused with aedeagal apodeme. Paraphysis slightly pointed at distal portion and proximally rounded, with one large and curved subapical setae.

    . Unknown.

    Etymology. The species name refers to its sibling species D. itacorubi sp. nov.

    Notes. As the other species from the D. peruensis group, the species described here have most of the setae and setulae of the thorax arising from dark spots; wings with crossveins clouded; surstylus mostly microtrichose; hypandrium square-shaped, mostly fused with the gonopods, without a dorsal arch. The aedeagus is stout, but devoid the pair of ventral, subapical spurs. This last character is inconstant among the species of the D. peruensis group, once D. boraceia have tiny spurs, while D. atalaia and D. pauliceia have large spurs. Drosophila itacorubi sp. nov. and D. paraitacorubi sp. nov. are very similar in morphology (and probably close related), but their affinity with the other members from D. peruensis species group is uncertain.

    The distinction between D. itacorubi sp. nov. from the other species from the D. peruensis species group can be based on some external morphological traits and by the male and female terminalia morphology. Among these species, D. itacorubi sp. nov. differs from D. atalaia and D. peruensis by the prescutellars setae, which is absent in the last species; from D. boraceia and D. paraitacorubi sp. nov. by the presence of first genal setae about half length of the vibrissae, and stout; from D. boraceia by the absence of two lines between dorsocentral setae and two interrupted ones outside dorsocentral setae formed by dark brown spots at the base of each setae on scutum; and from D. pauliceia by the yellowish brown antennae, instead of dark brown. Wings with both main crossveins darker than longitudinal ones, but not in D. atalaia. Regarding the male terminalia traits, D. itacorubi sp. nov. differs from D. paraitacorubi sp. nov. by the presence of 17 prensisetae, about nine long outer and 10 long inner setae, instead of 18 prensisetae, about eight long outer and seven long inner setae, as in the latter; the aedeagus of D. itacorubi sp. nov. is more rounded at the tip than that from the D. paraitacorubi sp. nov., and it exhibits a more curved profile in D. paraitacorubi sp. nov. than in D. itacorubi sp. nov.; differently of D. atalaia, D. pauliceia and D. boraceia, D. itacorubi sp. nov. does not possesses subapical spurs in the aedeagus; D. atalaia and D. pauliceia possesses a series of small lateral spur-like ornamentation (= scales, Ratcov & Vilela, 2007), which are absent in D. itacorubi sp. nov. The aedeagus shape of D. itacorubi sp. nov. is significantly similar to that of D. paraitacorubi sp. nov., and these species do not possess setae in the gonopods. The female terminalia of D. itacorubi sp. nov. differs from D. peruensis, D. pauliceia and D. atalaia in the number of marginal and discal ovisensilla of oviscapt. The former species has 17 marginal and seven discal ovisensilla, while D. peruensis has 14-15 marginal and three discal, D. pauliceia has 12 marginal and four discal, and D. atalaia has 10 marginal and three discal ovisensilla, respectively.

    As for D. itacorubi sp. nov., the distinction between D. paraitacorubi sp. nov. from the other species from the D. peruensis group can be based on external morphological and in the male terminalia traits. Drosophila paraitacorubi sp. nov. differs from D. atalaia and D. peruensis by the prescutellars setae, present in the new species; from D. atalaia, D. pauliceia and D. itacorubi sp. nov. by the presence of a small first genal setae, instead of a long one as in the last species; from D. boraceia by the absence of two lines between dorsocentral setae and two interrupted ones outside dorsocentral setae formed by dark brown spots at the base of each setae on scutum; from D. pauliceia by the yellowish brown antennae, instead of dark brown. See the D. itacorubi sp. nov. diagnosis on the male terminalia traits for differences between D. itacorubi sp. nov. and D. paraitacorubi sp. nov.

    Probably the species mentioned by Vilela & Cunha (2006) as undescribed Drosophila sp.1, illustrated by Marta E. Breuer and collected in Angra dos Reis (state of Rio de Janeiro), belongs to D. itacorubi sp. nov. or D. paraitacorubi sp. nov. The male genitalia of the Marta Breuer's specimen (shown in fig. 9d, Vilela & Cunha, 2006) and the few traits (characterized in one of her sketchbooks), though, are insufficient to certificate this statement. Such traits are "anterior scutellars divergent; main wing crossveins clouded; setae (and setulae?) arising from dark spots" (Vilela & Cunha, 2006). The first of these traits is inconsistent with our observations but there is a high similarity between the male genitalia drawn of "Undetermined Drosophila sp. 1" and the male genitalia of both new species described here. Based on Breuer's sketchbooks, these authors also point out that the male specimen (probably only one) was collected at Angra dos Reis, state of Rio de Janeiro, Brazil, on an unspecified date ("before March 2nd, 1959 [date of the terminalia drawing]"). Additionally, Ratcov & Vilela (2007) suggest that the species belongs to D. peruensis species group.

    Key to species of the Drosophila peruensis species group

    1. Prescutellars absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1'. Prescutellars present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2. Transverse veins of wing not clouded . . . . . . . . . . . . . . . . . . . . . . . D. atalaia 2'. Transverse veins of wing clouded . . . . . . . . . . . . . . . . . . . . . . . . D. peruensis 3. First genal setae less than half of the length of vibrissae. Tergites with a broad marginal brown band, medially interrupted . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3'. First genal setae as long as vibrissae. Tergites with a broad or narrow marginal brown band, medially interrupted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. The spots of the anterior area of the thorax are fused, what gives rise to four narrow lines. Surstylus with about 11 prensisetae. Aedeagus with a pair of short subapical spurs . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . .D. boraceia 4'.

    The spots of the thorax are not fused. Surstylus with about 17 prensisetae. Aedeagus without subapical spurs. . . . . . . . . . . . . . . . . . . D. itacorubi sp. nov.

    5. Pleura with spots. Tergites with a broad marginal brown band, medially interrupted. Surstylus with about 18 prensisetae. Aedeagus without subapical spurs. . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. paraitacorubi sp. nov. 5'. Pleura without spots. Tergites with a narrow marginal brown band. Surstylus with about 11 prensisetae. Aedeagus with a pair of subapical spurs. . . . . . D. pauliceia

    Recebido em 3 de março de 2011.

    Aceito em 22 de novembro de 2011.

    • Döge, J. S.; Valente, V. L. S. & Hofmann, P. R. P. 2008. Drosophilids (Diptera) from an Atlantic Forest area in Santa Catarina, Southern Brazil. Revista Brasileira de Entomologia 52:615-624.
    • Duda, O. 1927. Die sudamerikanischen Drosophiliden (Dipteren) unter Berucksichtigung auch der anderen neotropischen sowie der nearktischen Arten. Archiv für Naturgeschichte 91(A):1-228.
    • Eberhard, W. G. 1985. Sexual selection and animal genitalia Cambridge, Harvard University Press. 244p.
    • Frota-Pessoa, O. 1954. Revision of the tripunctata group of Drosophila with description of fifteen new species (Drosophilidae, Diptera). Arquivos do Museu Paranaense 10:253-304.
    • Gottschalk, M. S.; De Toni, D. C.; Valente, V. L. S. & Hofmann, P. R. P. 2007. Changes in Brazilian Drosophilidae (Diptera) assemblages across an urbanization gradient. Neotropical Entomology 36:848-862.
    • Hsu, T. C. 1949. The external genital apparatus of male Drosophilidae in relation to systematics. University of Texas Publications 4920:80-142.
    • Ratcov, R. & Vilela, C. R. 2007. A new Neotropical species of spot-thoraxed Drosophila (Diptera, Drosophilidae). Revista Brasileira de Entomologia 51:305-311.
    • Schmitz, H. J.; Valente, V. L. S. & Hofmann, P. R. P. 2007. Taxonomic survey of Drosophilidae (Diptera) from mangrove forests of Santa Catarina Island, Southern Brazil. Neotropical Entomology 36:53-64.
    • Sturtevant, A. H. 1921. The North American species of Drosophila Carnegie Institute of Washington Publication 301:1-150.
    • ______. 1942. The classification of the genus Drosophila, with description of nine new species. University of Texas Publications 4213:5-51.
    • Vilela, C. R. 1983. A revision of the Drosophila repleta species group (Diptera, Drosophilidae). Revista Brasileira de Entomologia 27:1-114.
    • ______. 1992. On the Drosophila tripunctata species group (Diptera, Drosophilidae). Revista Brasileira de Entomologia 36:197-221.
    • Vilela, C. R. & Bächli, G. 1990. Taxonomic studies on neotropical species of seven genera of Drosophilidae (Diptera). Mitteilungen der Schweizerische Entomologischen Gesellschaft 63:1-332.
    • ______. 2000. Morphological and ecological notes on the two species of Drosophila belonging to the subgenus Siphlodora Patterson & Mainland, 1944 (Diptera, Drosophilidae). Mitteilungen der Schweizerische Entomologischen Gesellschaft 73:23-47.
    • Vilela, C. R. & Cunha, A. B. 2006. On Marta Breuer and some of her unpublished drawings of Drosophila spp. male terminalia (Diptera, Drosophilidae). Genetics and Molecular Biology 29:580-587.
    • Vilela, C. R & Sene, F. M. 1982. A new spotted thorax species of the genus Drosophila (Diptera, Drosophilidae). Revista Brasileira de Entomologia 26:343-347.
    • Vilela, C. R. & Val, F. C. 2004. A new spot-thoraxed of Drosophila from the Atlantic Forest of southeastern Brazil (Diptera, Drosophilidae). Revista Brasileira de Entomologia 48:45-48.

    Publication Dates

    • Publication in this collection
      08 Mar 2012
    • Date of issue
      Dec 2011

    History

    • Received
      03 Mar 2011
    • Accepted
      22 Nov 2011
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