SciELO - Scientific Electronic Library Online

vol.48 issue2Cerambycidae (Coleoptera) coletados na Venezuela na copa de Matayba (Sapindaceae) e Vochysia (Vochysiaceae)Nothoprodontia, um novo gênero de Trachyderini (Coleoptera, Cerambycidae, Cerambycinae) author indexsubject indexarticles search
Home Pagealphabetic serial listing  

Services on Demand




Related links


Revista Brasileira de Entomologia

On-line version ISSN 1806-9665

Rev. Bras. entomol. vol.48 no.2 São Paulo June 2004 



Drosophila angustibucca Duda sensu Frota-Pessoa is an undescribed species (Diptera, Drosophilidae)



Carlos R. VilelaI; Vera L. S. ValenteII; Luciano Basso-da-SilvaIII

IDepartamento de Biologia, Instituto de Biociências, Universidade de São Paulo. Caixa Postal 11461, 05422-970 São Paulo-SP, Brazil. E-mail:
IIDepartamento de Genética, Instituto de Biociências, Universidade Federal do Rio Grande do Sul. Caixa Postal 15053, 91501-970, Porto Alegre-RS, Brazil. E-mail:
IIIInstituto de Ciências da Saúde, Centro Universitário FEEVALE. Rua Emílio Hauschild 70, 93525-180 Novo Hamburgo-RS, Brazil. E-mail:




Drosophila nappae sp. nov. , belonging to the subgroup I of the Drosophila tripunctata species group of the subgenus Drosophila, is described from flies of one strain established from several females collected from July 1994 through April 1995 at Morro Santana, Porto Alegre, state of Rio Grande do Sul, Brazil. This species has been misidentified during the past fifty years as Drosophila angustibucca (sensu Frota-Pessoa, 1954; non Duda, 1925, described from Costa Rica). Illustrations of male and female terminalia are also provided.

Keywords: Brazil; Neotropical region; new species; Paraguay; tripunctata group.


Drosophila angustibucca Duda sensu Frota-Pessoa é uma espécie ainda não descrita (Diptera, Drosophilidae). Uma nova espécie sul-americana de Drosophila, incluída no subgrupo I do grupo tripunctata e identificada incorretamente nos últimos 50 anos como sendo a espécie costarriquenha Drosophila angustibucca (sensu Frota-Pessoa, 1954; non Duda, 1925), é descrita no presente artigo sob o binômio Drosophila nappae sp. nov. Os espécimens-tipo foram retirados de uma linhagem obtida a partir de várias fêmeas coletadas entre julho de 1994 e abril de 1995 no Morro Santana, Porto Alegre (RS), Brasil. A descrição é acompanhada de ilustrações detalhadas das terminálias masculina e feminina.

Palavras-chave: Brasil; espécie nova; grupo tripunctata; Paraguai; região Neotropical.



Duda (1925) based his description of Drosophila angustibucca on 15 specimens (1 male, 14 females) collected in 1921, at Suiza de Turrialba, Costa Rica and deposited at the Magyar Természettudomanyi Múzeum [Hungarian National History Museum](HNHM) in Budapest, Hungary. As no holotype was designated in the original description they automatically acquired the status of syntypes. Thirty-eight years later, WHEELER (1963) analyzed two out of the 14 female syntypes, designated one of them as lectotype and, following a personal communication from the former curator (Dr. F. Mihályi) of the above-mentioned Museum, assumed the remaining 13 specimens (1 male, 12 females) of the original type series had been lost. Later on, however, BÄCHLI (1984:28) did realize that 12 (1 male, 11 females) of the remaining paralectotypes of Drosophila angustibucca had been overlooked by the former curator as they were not lost but preserved together with the lectotype and one paralectotype (both females) in the collection of the HNHM in Budapest and only one out of the 14 original females was in fact lost [just one laBELLed empty pin remains].

WHEELER (1963), while selecting the female lectotype of Drosophila angustibucca, was the first to raise a suspicion that the Brazilian specimens used by Frota-Pessoa (1954) to redescribe Drosophila angustibucca Duda, 1925 belong to an undescribed species. This suspicion was later confirmed by VILELA & BÄCHLI (1990), who redescribed the male paralectotype (the only male among the 15 syntypes of the original type series) of Drosophila angustibucca and realized that the males belonging to this species have a terminalia undoubtedly different from those of the Brazilian specimens mistakenly identified and illustrated by FROTA-PESSOA (1954), and cited by most of the subsequent Brazilian authors, as Drosophila angustibucca.

On the other hand, BÄCHLI et al. (2000) had the opportunity to dissect the sole female paralectotype of Drosophila paraguayensis Duda, 1927 and, mostly through the analysis of the spermathecae and oviscapt valves, to realize that it does not belong to the same species as the male lectotype but it rather belongs to one undescribed species most probably known in the literature as Drosophila angustibucca sensu Frota-Pessoa 1954 (non Duda, 1925).

The purpose of the present paper is to formally describe this relatively well-known but still undescribed species of Neotropical Drosophila belonging to the tripunctata group.

Refer to KANESHIRO (1969) and VILELA & BÄCHLI (1990, 2000) for details respectively on the methods and terminology used.

Drosophila nappae sp. nov.
(Figs. 1-10)





Drosophila paraguayensis Duda, 1927 (partim, paralectotype female [deposited in the Staatliches Museum für Tierkunde, Dresden, Germany]):186 (fig.59, female terminalia); Frota-Pessoa, 1954:265 but not 266 (key); indirectly designated by Vilela & Bächli, 1990:102, formally laBELLed as such by Bächli et al., 2000:76, and cited by them as "undescribed Drosophila species", 90 (figs. 14A, left wing; B, left oviscapt valve; C, right oviscapt valve; D, inner spermathecal capsules).
Drosophila angustibucca (sensu Frota-Pessoa, 1954; non Duda, 1925); Frota-Pessoa, 1954 (misidentification):266 (key), 278 (as a redescription), plate XIV (figs. 4, male terminalia, ventral view; 5, aedeagus, aedeagal apodeme, left gonopod + paraphysis, dorsal view), plate XVII (fig. 38, right oviscapt valve), plate XVIII (fig. 48, inner spermathecal capsule; fig. 57, egg), plate XIX (fig. 67, male terminalia, right lateral view), plate XXI (fig. 85, posterior leg), plate XXII (fig. 94, right wing); Araújo & Valente, 1981:1488 (table I, seasonal fluctuation, abundance), 1489 (table III, feeding site); Franck et al., 1984:244 (fig. 1, metaphase plates), 245 (fig. 2, photomap of polytene chromosomes), 248 (fig. 3, chromosome inversions); Franck & Valente, 1985:135-138 (tables II-IV, figs. 1-4, seasonal fluctuation); Valente & Araújo, 1991:241, 242 (tables I, II, seasonal fluctuation, abundance) 244-247 (tables III, IV, feeding sites); Franck & Napp, 1992:125, 126 (figs. 1c, 2c, electrophoretic patterns), 127, 128 (tables 1, 2, genetic similarity) 129 (fig. 3, dendrograms); De Toni & Hofmann, 1995:349 (table I, seasonal fluctuation, abundance); Saavedra et al., 1995:63 (table 1, distribution, abundance), 65 (fig. 3 , seasonal fluctuation), 67, 68 (table 3, feeding and breeding sites), 69, 70 (tables 4, 5, niche-breadth, niche overlapping), 71 (fig. 4, dendrogram); Loreto et al., 1998: 154-155 (table 1, transposable elements); Yotoko et al., 2003:615 (footnote of table 1, COXII gene).
Drosophila species R3 of the tripunctata group; Tidon-Sklorz et al., 1994:631 (table II, geographical distribution, abundance), 634 (subgroup affiliation).
Drosophila species 1 of the tripunctata group; Yotoko et al., 2003:615 (table 1, COXII gene), 617 (fig. 1, phylogenetic tree).

Holotype male (postabdomen dissected), laBELLed "Brasil - RS [Rio Grande do Sul], Porto Alegre, VII.1994 - IV.1995, L. Basso-da-Silva col. / Drosophila nappae Vilela, Valente & Basso-da-Silva / HOLOTIPO [red label]". Paratypes: 4 males (2 dissected) and 8 females (3 dissected): same data as holotype. The disarticulated terminalia are kept in a microvial filled with glycerin and attached by the stopper to the pinned specimen. The type specimens were obtained in late May 1995 from a strain established by a mixture of several males and females collected by net sweeping over banana-baited traps between July 1994 and April 1995 at Morro Santana, Porto Alegre. They are all deposited in the Museu de Zoologia, Universidade de São Paulo, São Paulo, SP, Brazil (MZSP). Type locality: Morro Santana, Porto Alegre, state of Rio Grande do Sul, Brazil.

Additional specimen examined (not used in the description, therefore not considered paratype). BRASIL. Minas Gerais: 1 male (previously dissected) labelled : "Serra do Cipó. MG. BR. 1130m. 24-III-77, Sene e Carsons" [terminalia in an attached microvial, collection code A40 = gallery forest located at 27 km from the bridge over Cipó river, at an elevation of 1,130 m above sea level, municipality of Santana do Riacho, highway MG-010 in the direction of Conceição do Mato Dentro], deposited in the MZSP.

Diagnosis. Scutum brown, anterior half with two lighter stripes just within the dorsocentral rows; facial carina large and broad, not sulcate; wing brownish, crossveins clouded, C index = 3.7-4.4; three strong black setae at base of posterior tarsomere I; abdomen shining yellow, tergites 2-4 with posterior dark brown bands medially broadened; aedeagus anterodorsally bearing a distinctive pair of slightly membranous, finger-shaped processes, which are backwards directed and covered with tiny spines.

Description. Head mainly yellowish brown. Frontal length = 0.32-0.40 mm; top to bottom width ratio = 1.4-1.7. Frontal triangle indistinct; ocellar triangle dark brown, about 31-38 % of frontal length. Fronto-orbital plates indistinct. Mid orbital as tiny as fronto-orbital setulae, distance of posterior orbital to anterior one ca. 1/2 of that to inner vertical. Length ratio of anterior to posterior orbital = 0.6-0.7, of mid to anterior orbital = 0.2-0.4; poc = 62-80%, oc = 79-93% of frontal length; vt index = 1.2; vibrissal index = 0.3-0.5. Facial carina yellow, broad, not sulcate. Cheek index = 5.4-7.0. Eye index = 1.2-1.3. First flagellomere short-haired; length to width ratio = 1.8-2.0. Arista with 6-8 upper and 3-4 lower branches plus terminal fork, 7-9 relatively long inner branches. Proboscis and palpi light brown. Face light yellow.

Thorax dull, brown; anterior half with two lighter stripes just within the dorsocentral rows; length = 1.4-1.6 mm; h index = 0.6-0.9; 8 irregular rows of acrostichals. Transverse distance of dorsocentrals 1.9-2.5x longitudinal distance; dc index = 0.6-0.8. Scutellum brown; scut position index = 0.7-0.9; basal scut divergent; scut index = 1.0-1.1; prescutellars absent; sterno index = 0.5-0.6; mid katepisternal = 58-100% of the anterior one. Halteres light brown, dorsally darker. Legs uniformly light brown. Three strong black setae at base of posterior tarsomere I, which is slightly wider than posterior tarsomere II, but not twice as wide, as it remarkably occurs in Drosophila platitarsus Frota-Pessoa, 1954. Apical setae on first and second tibiae, the latter spur-shaped, preapicalls on all three.

Wing brownish, slightly pointed at tip of R4+5, crossveins clouded; length = 2.8-3.2 mm; length to width ratio = 2.1-2.3. Indices: C = 3.7-4.4, ac = 1.6-2.0, hb = 0.4-0.5, 4c = 0.6-0.7, 4v = 1.4-1.6, 5x = 0.7-1.0, M = 0.3-0.4, prox. x = 0.4-0.5.

Abdomen shining brownish yellow, tergite 2 with a posterior, usually medially interrupted dark brown band, tergites 3-5 with a posterior dark brown band, usually not interrupted but enlarged and somewhat triangle-shaped in the mid line, mostly not reaching anterior margin, tergite 6 slighty darker in the mid line.

Male terminalia (Figs. 1-8). Epandrium slightly microtrichose on posterior dorsal area with 3-6 upper and no lower setae; ventral lobe slightly covering surstylus. Cerci slightly microtrichose on dorsal area, linked to epandrium by membranous tissue. Surstylus not microtrichose, with 7-8 cone-shaped prensisetae, 3-4 long, strong outer setae and 7-9 long, thin inner setae. Decasternum as in Fig. 2. Hypandrium longer than epandrium; dorsal arch present, strongly sclerotized; gonopod fused to paraphysis, bearing one long seta on anterior inner margin. Aedeagus lateroventrally strongly sclerotized, anterodorsally bearing a distinctive pair of slightly membranous, finger-shaped, and backwards directed processes, covered with tiny spines; dorsally bearing a small, slightly sclerotized, crescent-shaped plate at subdistal area; distal end medially membranous in dorsal and ventral views, laterally expanded and ventrally covered with tiny spines. Aedeagal apodeme rod-shaped, slightly longer than aedeagus. Ventral rod as in Figs. 5-8.

Female terminalia (Figs. 9, 10). Valves of oviscapt (Fig. 9) with 16-19 marginal and 5-7 discal peg-like ovisensilla. Spermathecal capsule (Fig. 10) somewhat elliptical, with some furrows at base; spermathecal duct with a spherical dilatation, becoming gradually wider apically, sclerotized. FROTA-PESSOA (1954:279) states the dilatation is positioned at the base of the capsule, as it is shown in his figure 48 (plate XVIII). This feature is also seen in the photomicrograph of the spermathecal capsules taken by BÄCHLI et al. 2000:90 (fig. 14D). However, the dilatation is more apically positioned in our Fig. 10 probably because the spermathecal duct has moved to an inner position during the chemical treatments.

Etymology. The epithet is a genitive patronym to honor our good friend the late Marly Napp from the Departamento de Genética, Universidade Federal do Rio Grande do Sul.

Geographical distribution. Based on its type locality (Porto Alegre) and assuming that all the material examined (unknown depository) and misidentified by FROTA-PESSOA (1954: 279) as Drosophila angustibucca (not Duda, 1925) belongs in fact to Drosophila nappae sp. nov., its known distribution includes southeastern and southern Brazil, where it has been collected in the states of Rio de Janeiro (Rio de Janeiro [then Distrito Federal]), São Paulo (Campos do Jordão, Mogi das Cruzes, Mongaguá [cited as Vila Atlântica], Pirassununga) and Rio Grande do Sul (Feliz and Porto Alegre). Making the same above-cited assumption for the specimens identified (apparently using the key by FROTA-PESSOA 1954) and cited in most subsequent papers (i.e. ARAÚJO & VALENTE 1981; FRANCK et al. 1984; FRANCK & VALENTE 1985; VAL & KANESHIRO 1988; VALENTE & ARAÚJO 1991; DE TONI & HOFMANN 1995; SAAVEDRA et al. 1995), the following additional localities of the state of São Paulo: Salesópolis [Estação Biológica de Boracéia], state of Rio Grande do Sul: Bento Gonçalves, Esmeralda [Estação Ecológica de Aracuri], Guaíba [Estação experimental agronômica], Viamão [Parque de Itapuã], Tenente Portela [Parque Florestal Estadual do Turvo] and state of Santa Catarina: Santa Catarina Island [Morro da Lagoa da Conceição] must be included. Moreover, assuming that the female paralectotype (but not the male lectotype) of Drosophila paraguayensis Duda, 1927 also belongs, according to Bächli et al. (2000:79,80,90), to Drosophila nappae sp. nov. (cited as undescribed Drosophila species), the distribution of the latter species must includes Paraguay (Hohenau [Department of Itapúa]) as well. However, it should be pointed out that, for the time being, we have no way to know whether or not the male specimen collected by C.A.W. Schnuse and O. Garlepp in Peru [Department of Ucayali], Mishagua [river, mispelled as Meshagua on the label] (by the Urubamba river), on 03.X.1903, and identified by DUDA (1927:184) as Drosophila angustibucca belongs to Drosophila nappae sp. nov. If extant, it should be checked regarding to its identification. Additionally, we have no way to verify whether or not the specimens cited as occurring in Minca (HOENIGSBERG & LIN 1977:58) and Santa Marta's highlands (HOENIGSBERG 1995a:87, b:177), Colombia, belongs to Drosophila angustibucca Duda, 1927 or Drosophila nappae sp. nov. as there are no statements regarding their depositories or about the key used to identify them. So, the known (confirmed) northernmost record of Drosophila nappae sp. nov. is represented by just one analyzed male specimen (cited above) collected in the Serra do Cipó (municipality of Santana do Riacho), state of Minas Gerais, Brazil.

Larval breeding sites (n = emerged imagines). Assuming that all the identifications of Drosophila angustibucca cited in the literature (except those by Duda) were based on FROTA-PESSOA (1954:266) only 11 specimens of Drosophila nappae sp. nov. have been so far recorded as being emerged from fruits, which belong to two plant families. Thus, it seems that they do not represent the main substrate where the flies develop. They are as follows: fruits of Syagrus romanzoffiana (Cham.) Glassman (Arecaceae) collected in the Parque Florestal Estadual do Turvo (Spring 1982, n = 9) and at Guaíba (Winter 1982, n = 1) (SAAVEDRA et al. 1995:67, 68; cited as Arecastrum romanzoffianum), fruit of Randia armata (Sw.) DC (Rubiaceae) collected in the Parque Florestal Estadual do Turvo (Spring 1982, n = 1, according to SAAVEDRA et al. 1995:68).

Adult feeding sites (n = collected imagines). Assuming that all the identifications of Drosophila angustibucca in the literature (except those by Duda) were based on FROTA-PESSOA (1954), a total of 232 specimens (listed below) have been recorded as being collected as adults over fruits belonging to eight plant families, of which just one (Arecaceae) has also been recorded as a breeding site for Drosophila nappae sp. nov. Fruits of Ficus organensis (Miq.) Miq. (Moraceae) in the Parque de Itapuã (III.1980, n = 1 and VII.1981, n = 14 in VALENTE & ARAÚJO 1991:244). Fruits of Diospyros inconstans Jacquin (Ebenaceae) in the Parque de Itapuã (VII.1981, n = 8 in VALENTE & ARAÚJO 1991:244; cited as Maba inconstans). Fruits of Passiflora elegans Mart.(Passifloraceae) in the Parque de Itapuã (I.1981, n = 120 in VALENTE & ARAÚJO 1991:244). Fruits of Maclura tinctoria (L.) D. Don ex Steud (Moraceae) in the Parque Florestal Estadual do Turvo (I.1982, n = 6 in VALENTE & ARAÚJO 1991:246 and SAAVEDRA et al. 1995:67; cited as Chlorophora tinctoria). Fruits of Syagrus romanzoffiana (Cham.) Glassman (Arecaceae) in the Parque Florestal Estadual do Turvo (X.1980, n = 2 in Valente & Araújo 1991:245), (Spring 1982, n = 28) and at Guaíba (Winter 1982, n = 4) (SAAVEDRA et al. 1995:67, 68; cited as Arecastrum romanzoffianum). Fruits of Campomanesia guazumifolia (Cambessedes) (Myrtaceae) in the Parque Florestal Estadual do Turvo (II.1981, n = 5 in VALENTE & ARAÚJO 1991:247; cited as Brittoa sellowiana). Fruits of Bromelia antiacantha Bertol (Bromeliaceae) in the Parque Florestal Estadual do Turvo (X.1980, n = 3 in VALENTE & ARAÚJO 1991:246). Fruits of Cabralea glaberrima in the Parque Florestal Estadual do Turvo (X.1980, n = 20 in VALENTE & ARAÚJO 1991:246). Fruits of Didymopanax morototonii (Aubl.) Dcne et Planch. (Araliaceae) in the Parque Florestal Estadual do Turvo (VII.1980, n = 20 in Valente & Araújo 1991:246). Fruit of Chrysophyllum gonocarpum (Martius & Eichler) Engler (Sapotaceae) in the Parque Florestal Estadual do Turvo (Spring 1982, n = 1 in SAAVEDRA et al. 1995:68).

Relationships. It belongs to the subgroup I (cf. VILELA 1992:198) of the Drosophila tripunctata species group of the subgenus Drosophila. It shares with D. angustibucca Duda, 1925, D. neoguaramunu FrydenBERG, 1956, D. platitarsus Frota-Pessoa, 1954, D. rostrata Duda, 1925, and D. setula Heed & Wheeler, 1957, the following remarkable features: aedeagus dorsally with a small, sclerotized, crescent-shaped plate at subdistal area, and anterodorsally bearing a pair of finger-shaped and backwards directed processes. The processes are smaller and mostly membranous in those five species but only slightly membranous and proportionally longer in Drosophila nappae sp. nov. Additionally, all of them but the latter, have in common a remarkable, sclerotized, T-shaped area partially surrounded by the pair of processes in the anterodorsal, mostly membranous surface of aedeagus.


Acknowledgments. We are indebted to CNPq, FAPERGS, FINEP and PROPESQ-UFRGS for financial support.



ARAÚJO, A. M. & V. L. S. VALENTE. 1981. Observações sobre alguns lepidópteros e drosofilídeos do Parque do Turvo, RS. Ciência e Cultura 33(11): 1485-1490.         [ Links ]

BÄCHLI, G. 1984. Catalog of the types of Drosophilidae in the Hungarian Natural History Museum, Budapest (Diptera). Folia Entomologica Hungarica 45(2): 27-41.         [ Links ]

BÄCHLI, G.; C. R. Vilela & V. Ratcov. 2000. Morphological differences among Drosophila paraguayensis Duda, 1927 and its close relatives (Diptera, Drosophilidae). Mitteilungen der schweizerischen Entomologischen Gesellschaft 73(1-2): 67-92.         [ Links ]

DE TONI, D. C. & P. R. P. HOFMANN. 1995. Preliminary taxonomic survey of the genus Drosophila (Diptera, Drosophilidae) at Morro da Lagoa da Conceição; Santa Catarina Island; Brazil. Revista Brasileira de Biologia 55(3): 347-350.         [ Links ]

DUDA, O. 1925. Die costaricanischen Drosophiliden des Ungarischen National-Museums zu Budapest. Annales Historico-Naturales Musei Nationalis Hungarici 22: 149-229.         [ Links ]

DUDA, O. 1927. Die südamerikanischen Drosophiliden (Dipteren) unter Berücksichtigung auch der anderen neotropischen sowie der nearktischen Arten. Archiv für Naturgeschichte 91 (A)11/12(1925): 1-228.         [ Links ]

FRANCK, G. & M. NAPP. 1992. Electrophoretic analysis of interspecific relationship in six species of the Drosophila tripunctata group from south of Brazil. Evolución Biológica 6: 121-133.         [ Links ]

FRANCK, G. & V. L. S. VALENTE. 1985. Study on the fluctuation in Drosophila populations of Bento Gonçalves, RS, Brazil. Revista Brasileira de Biologia 45(1/2): 133-141.         [ Links ]

FRANCK, G.; V. L. S. VALENTE & N. B. MORALES. 1984. Chromosome characteristics of Drosophila angustibucca Duda. Revista Brasileira de Genética 7(2): 241-253.         [ Links ]

FROTA-PESSOA, O. 1954. Revision of the tripunctata group of Drosophila with description of fifteen new species (Drosophilidae Diptera). Arquivos do Museu Paranaense 10(6): 253-304 + 13 plates.         [ Links ]

HOENIGSBERG, H. F. 1995a. Collecting Drosophila species in natural surroundings. Drosophila Information Service 76: 86-87.         [ Links ]

HOENIGSBERG, H. F. 1995b. Drosophila baits in northern lowlands of Colombia and high altitude mountainous Andes can change according to natural fruits in the area. Drosophila Information Service 76: 176-177.         [ Links ]

HOENIGSBERG, H. F. & F. J. Lin. 1977. Population genetics in the American Tropics. XIV. Drosophila melanogaster Meigen in Colombian and Chinese natural habitats. Mitteilungen der schweizerischen Entomologischen Gesellschaft 50: 57-58.         [ Links ]

KANESHIRO, K. Y. 1969. A study of the relationships of Hawaiian Drosophila species based on external male genitalia. University of Texas Publications 6918: 55-70.         [ Links ]

LORETO, E. L. S.; L. BASSO DA SILVA; A. ZAHA & V. L. S. VALENTE. 1998. Distribution of transposable elements in neotropical species of Drosophila. Genetica 101(2): 153-165.         [ Links ]

SAAVEDRA, C. C. R.; S. M. CALLEGARI-JACQUES; M. NAPP & V. L. S. VALENTE. 1995. A descriptive and analytical study of four neotropical drosophilid communities. Journal of Zoological Systematics and Evolutionary Research 33(2): 62-74.         [ Links ]

TIDON-SKLORZ, R.; C. R. VILELA; F. M. SENE & M. A. Q. R. PEREIRA. 1994. The genus Drosophila (Diptera, Drosophilidae) in the Serra do Cipó, state of Minas Gerais, Brazil. Revista Brasileira de Entomologia 38(3/4): 627-637.         [ Links ]

VAL, F. C. & K. Y. KANESHIRO. 1988. Drosophilidae (Diptera) from the Estação Biológica de Boracéia, on the coastal range of the state of São Paulo, Brazil : geographical distribution, p. 189-203. In: P. E. VANZOLINI & W. R. HEYER (eds.). Proceedings of a workshop on Neotropical distribution patterns. Rio de Janeiro, Academia Brasileira de Ciências, 488 p.         [ Links ]

VALENTE, V. L. &. A. M. ARAÚJO.1991. Ecological aspects of Drosophila species in two contrasting environments in southern Brazil (Diptera, Drosophilidae). Revista Brasileira de Entomologia 35(2): 237-253.         [ Links ]

VILELA, C. R. & G. BÄCHLI. 1990. Taxonomic studies on neotropical species of seven genera of Drosophilidae (Diptera). Mitteilungen der schweizerischen Entomologischen Gesellschaft 63(suppl.): 1-332.         [ Links ]

VILELA, C. R. & G. BÄCHLI. 2000. Morphological and ecological notes on the two species of Drosophila belonging to the subgenus Siphlodora Patterson & Mainland, 1944 (Diptera, Drosophilidae). Mitteilungen der schweizerischen Entomologischen Gesellschaft 73(1-2): 23-47.         [ Links ]

WHEELER, M. R. 1963. Notes on the extant types of Dr. O. Duda's Costa Rican Drosophilidae (Diptera). Bulletin of the Brooklin Entomological Society 58(2/3): 51-61.         [ Links ]

YOTOKO, K. S. C.; H. F. MEDEIROS; V. N. SOLFERINI & L. B. KLACZKO. 2003. A molecular study of the systematics of the Drosophila tripunctata group and the tripunctata radiation. Molecular Phylogenetics and Evolution 28(3): 614-619.        [ Links ]



Received 30.IX.2003; accepted 30.I.2004

Creative Commons License All the contents of this journal, except where otherwise noted, is licensed under a Creative Commons Attribution License