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Diptera: Drosophilidae) in the Biomes Caatinga and Atlantic Forest, Northeastern Brazil

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Diptera: Drosophilidae) in the Biomes Caatinga and Atlantic Forest, Northeastern Brazil ARTHROPOD BIOLOGY Abundance and Richness of Cryptic Species of the willistoni Group of Drosophila (Diptera: Drosophilidae) in the Biomes Caatinga and Atlantic Forest, Northeastern Brazil ANA CRISTINA LAUER GARCIA,1,2 DIVA MARIA IZABEL DE OLIVEIRA SILVA,3 AMANDA GABRIELA FELIX MONTEIRO,1 GEO´ RGIA FERNANDA OLIVEIRA,1 4 1 MARTI´N ALEJANDRO MONTES, AND CLA´ UDIA ROHDE Downloaded from https://academic.oup.com/aesa/article/107/5/975/64874 by guest on 27 September 2021 Ann. Entomol. Soc. Am. 107(5): 975Ð982 (2014); DOI: http://dx.doi.org/10.1603/AN14052 ABSTRACT Cryptic species meet the biological deÞnition of species, but are morphologically identical or quite similar. Several ecological studies underestimate richness and neglect important information on cryptic species, as they are rather difÞcult to identify. Among insects, drosophilids of the willistoni subgroup, which includes Drosophila willistoni Sturtevant, Drosophila paulistorum Dobzhansky & Pavan, Drosophila equinoxialis Dobzhansky, Drosophila tropicalis Burla & da Cunha, Drosophila insularis Dobzhansky, and Drosophila pavlovskiana Kastritsis & Dobzhansky, are good examples of cryptic species. Although several studies have shown that this subgroup is one of the most abundant in the Neotropical region, no identiÞcation to species level has been reported for areas where these individuals live in sympatry. This study evaluates the seasonal oscillations in abundance of this subgroup in biomes with contrasting vegetation and rainfall regimes: the Caatinga and the Atlantic Forest, in northeastern Brazil. Approximately 39,000 drosophilids were captured in 24 collections, of which 18,000 belonged to the willistoni subgroup. The most abundant were D. willistoni, D. pauli- storum, and D. equinoxialis, in this order. D. equinoxialis was recorded in only one of the environments surveyed, represented by few individuals. In all environments, individuals of the willistoni subgroup were more abundant in the rainy season, when richness often was higher. The results underline the importance of humidity and of temperature for the subgroup willistoni and indicate the ecological versatility of some of its species. KEY WORDS ecology, neotropical region, pernambuco, temperature, humidity Cryptic species establish morphologically identical or by cryptic species (Salles 1947; Burla et al. 1949; quite similar natural populations that nevertheless re- Breuer and Pavan 1950; Magalha˜es and Bjo¨rnberg main isolated from the reproductive standpoint (Mayr 1957; Vilela 1983, 1992; Vilela and Ba¨chli 1990; Vilela 1963). The notion of cryptic species is one of the most et al. 2002). Examples include the willistoni group of stringent conditions that defy the morphological con- Drosophila, which is composed of three subgroups. cept of species and poses a signiÞcant problem to One of these is the willistoni subgroup, represented by taxonomists. Dobzhansky (1970) underscores the im- six cryptic species. Of these, Drosophila insularis Dob- portance of the identiÞcation of these individuals in zhansky and Drosophila pavlovskiana Kastritsis & evolutionary and ecological studies, and emphasizes Dobzhansky are endemic to the Lesser Antilles and that species are natural entities that exist irrespec- Guiana, respectively. The other four species are tively of our competence to distinguish one from the widely distributed and are often sympatric. Drosophila other. willistoni Sturtevant occurs from the south of the A considerable share of the diversity of small ßies United States to northern Argentina and Uruguay, that comprise the Drosophilidae family is represented Drosophila paulistorum Dobzhansky & Pavan spreads from northern Guatemala to southern Brazil, Drosoph- 1 Universidade Federal de Pernambuco, Centro Acadeˆmico de ila equinoxialis Dobzhansky distributes from southern Vito´ria, Programa de Po´s-Graduac¸a˜oemSau´ de Humana e Meio Am- Mexico and practically reaches central Brazil and biente, Rua do Alto do Reservato´rio, s/n, Bela Vista, 55608-680 Vito´ria Peru, while Drosophila tropicalis Burla & da Cunha is de Santo Anta˜o, PE, Brazil. 2 Corresponding author, e-mail: [email protected]. recorded from northern Guatemala down to central 3 Universidade de Pernambuco, Programa de Po´s-Graduac¸a˜oem Brazil and central Bolivia (Spassky et al. 1971, Dob- Biologia Celular e Molecular Aplicada, Rua Arno´bio Marques, 310, zhansky and Powell 1975, Ehrman and Powell 1982, Bairro Santo Amaro, 50100-130 Recife, PE, Brazil. Santos and Valente 1990). 4 Universidade Federal Rural de Pernambuco, Departamento de Biologia, Campus Dois Irma˜os. Rua Dom Manoel de Medeiros, s/n. In the Neotropical region, the willistoni subgroup is CEP 52171-900 Recife, PE, Brazil. one of the most abundant in numerous environments, 0013-8746/14/0975Ð0982$04.00/0 ᭧ 2014 Entomological Society of America 976 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 5 Downloaded from https://academic.oup.com/aesa/article/107/5/975/64874 by guest on 27 September 2021 Fig. 1. Collection sites of drosophilids in the state of Pernambuco, northeastern Brazil, in the Caatinga biome (black squares) and Atlantic Forest biome, biogeographical subregions Brejos de Altitude (triangles) and Pernambuco (circles). Inset shows the map of South America, Brazil (light gray), and the northeastern region of the country (dark gray). Site codes: SET-1 (Serra Talhada, 07Њ 57Ј S, 38Њ 18Ј W); SET-2 (Serra Talhada, 07Њ 56Ј S, 38Њ 17Ј W); CAT-1 (Buõ´que, 08Њ 24Ј S, 37Њ 09Ј W); CAT-2 (Buõ´que, 08Њ 36Ј S, 37Њ 14Ј W); TRI (Triunfo, 07Њ 50Ј S, 38Њ 06Ј W); MAD (Brejo da Madre de Deus, 08Њ 12Ј S, 36Њ 24Ј W); BEZ (Bezerros, 08Њ 53ЈS, 36Њ 29Ј W); BON (Bonito, 08Њ 30Ј S, 35Њ 42Ј W); TAP (Sa˜o Lourenc¸o da Mata, 08Њ 07Ј S, 35Њ 17Ј W); POM (Pombos, 08Њ 08Ј S, 35Њ 23Ј W); ZOO (Recife, 08Њ 03Ј S, 34Њ 52Ј W); SAL (Tamandare´ ,08Њ 45Ј S, 35Њ 06Ј W). and sometimes accounts for the largest proportion of gimes: the Caatinga and the Atlantic Forest in north- individuals collected (Dobzhansky and Pavan 1950, eastern Brazil. In the latter, the biogeographic subre- Martins 2001, Medeiros and Klaczko 2004, Tidon 2006, gions Brejos de Altitude and Pernambuco were Do¨ge et al. 2008, Acurio and Rafael 2009, Rohde et al. surveyed. 2010). Nevertheless, in several studies carried out in areas where more than one species of this group has Materials and Methods been recorded, its representatives are not identiÞed to species level. This situation is observed in Brazilian Study Sites. Adult drosophilids were collected in biomes like the Atlantic Forest (Pavan 1950, Sene et 12 sites in the state of Pernambuco, northeastern Bra- al. 1980, Tidon-Sklorz and Sene 1992, De Toni et al. zil. Of these, four are located in the Caatinga biome 2001, Silva et al. 2005, Mateus et al. 2006, Torres and and eight are in the Atlantic Forest biome. In the Madi-Ravazzi 2006, De Toni et al. 2007, Cavasini et al. latter, four are in the Brejos de Altitude subregion and 2008, Do¨ge et al. 2008, Gottschalk et al. 2009, Bizzo et four are in the Pernambuco subregion (Fig. 1). In each al. 2010), the Cerrado (Sene et al. 1980, Torres and site, two collections were conducted, totaling 24 sam- Madi-Ravazzi 2006, Tidon 2006, Blauth and Gottschalk plings, at periods characterized by contrasting rainfall 2007, Mata et al. 2008), the Amazon Forest (Martins levels (the dry and the rainy season). 1989, 2001), and the Caatinga (Sene et al. 1980). The vegetal formations in Caatinga are character- In light of the importance and of the paucity of ized by xerophilous steppic savanna, and the dry sea- information about species of the willistoni subgroup, son is long (Eiten 1982). Mean annual rainfall varies the current study assessed the seasonal oscillations in between 400 and 600 mm, most of which in three willistoni representativesÕ richness and abundance in consecutive months between November and June biomes with contrasting vegetation and rainfall re- (AbÕSaber 1977). The Caatinga is the only exclusively September 2014 GARCIA ET AL.: CRYPTIC SPECIES OF THE willistoni GROUP OF Drosophila 977 Table 1. Absolute abundance of species of the willistoni subgroup of Drosophila and of the other drosophilid species captured in different collection efforts in each of the three environments surveyed in the dry and rainy seasons Locality willistoni Subgroup Other Biome Environment Date Season Total code D. willistoni D. paulistorum D. equinoxialis Total species Caatinga SET-1 15 Feb. 2009 Dry 3 Ð Ð 3 275 278 23 June 2009 Rainy 1,611 133 Ð 1,744 771 2,515 SET-2 15 Feb. 2009 Dry 1 Ð Ð 1 783 784 23 June 2009 Rainy 1,975 154 Ð 2,129 1,069 3,198 CAT-1 13 Mar. 2009 Dry Ð Ð Ð Ð 273 273 25 June 2008 Rainy 312 Ð Ð 312 630 942 CAT-2 13 Mar. 2009 Dry Ð Ð Ð Ð 350 350 25 June 2008 Rainy 149 55 Ð 204 1,011 1,215 Total 4,051 342 Ð 4,393 5,162 9,555 Atlantic Forest Brejos de altitude TRI 30 Jan. 2009 Dry 8 7 Ð 15 284 299 Downloaded from https://academic.oup.com/aesa/article/107/5/975/64874 by guest on 27 September 2021 23 June 2009 Rainy 774 298 Ð 1,072 602 1,674 MAD 16 Dec. 2010 Dry 31 4 Ð 35 35 70 10 Nov. 2010 Rainy 274 5 3 282 315 597 BEZ 8 Dec. 2009 Dry 44 Ð Ð 44 395 439 12 July 2008 Rainy 117 11 Ð 128 4,064 4,192 BON 6 Feb. 2010 Dry 994 11 Ð 1,005 862 1,867 17 May 2010 Rainy 3,028 262 Ð 3,290 894 4,184 Total 5,270 597 3 5,871 7,451 13,322 Pernambuco TAP 5 Mar. 2011 Dry 682 42 Ð 724 2,960 3,684 14 Oct. 2010 Rainy 746 73 Ð 819 1,152 1,971 POM 5 Jan.
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