Light-attracted hawkmoths (Lepidoptera: Sphingidae) of Boracéia, municipality of Salesópolis, state of São Paulo, Brazil

The light-attracted hawkmoths (Lepidoptera: Sphingidae) of the Estação Biológica de Boracéia, municipality of Salesópolis, state of São Paulo, Brazil were sampled during a period of 64 years (1940-2004). A total of 2,064 individuals belonging to 3 subfamilies, 6 tribes, 23 genera and 75 species were identified. Macroglossinae was the most abundant and richest subfamily in the study area, being followed by Sphinginae and Smerinthinae. About 66 % of the sampled individuals were assorted to the macroglossine tribes Dilophonotini and Macroglossini. Dilophonotini (Macroglossinae) was the richest tribe with 26 species, followed by Sphingini (Sphinginae) with 18 species, Macroglossini (Macroglossinae) with 16 species, Ambulycini (Smerinthinae) and Philampelini (Macroglossinae) with seven species each one, and Acherontiini (Sphinginae) with only one species. Manduca Hübner (Sphinginae) and Xylophanes Hübner (Macroglossinae) were the dominant genera in number of species. Only Xylophanes thyelia thyelia (Linnaeus) and Adhemarius eurysthenes (R. Felder) were recorded year round Introduction Hawkmoths (Lepidoptera: Sphingidae) comprise about 200 genera and 1300 species (Kitching and Cadiou 2000). They are distributed throughout the world, except Antarctica and Greenland (Moré et al. 2005). Approximately one third of these moths belong to the Neotropical fauna (Heppner 1991; 1998). In Brazil, they are represented by 29 genera and 210 species (compilation of Rothschild and Jordan 1910; Moss 1920; Hambleton and Forbes 1935; Oiticica 1939; 1942; Zikán and Zikán 1968; Laroca and Mielke 1975; Schreiber 1978; Biezanko 1981; Ferreira et al. 1986; Laroca et al. 1989; Motta et al. 1991; 1998; Carcasson and Heppner 1996; Motta and Soares 1997; Marinoni et al. 1999; Oliveira et al. 1999; Brown and Freitas 2000; Corseuil et al. 2001; Duarte Jr. et al. 2001; Motta and Andreazze 2001; 2002; Becker 2001; Soares and Motta 2002; Darrault and Schlindwein 2002; Gusmão et al. 2003; Gusmão and Creão-Duarte 2004a; b; Duarte Jr. and Schlindwein 2005; Motta and Xavier-Filho 2005). Hawkmoths have long been recognized as major pollinators of flowering plants (Baker 1961; Gregory 1963; Silberbauer-Gottsberger and Gottsberger 1975; Janzen 1984; Haber and Frankie 1989; Darrault and Schlindwein 2002; Kitching 2002). Because of their capability to fly far away, these moths are potential long distance pollen dispersers (Linhart and Mendenhall 1977; Nilsson et al. 1992; Chase et al. 1996). They may be diurnal, crepuscular or nocturnal. In South America, however, most species are more active at night, when they may be easily observed visiting flowers with white or pale corollas, very long tubes or spurs and copious amounts of nectar (Silberbauer-Gottsberger and Gottsberger 1975; Baker and Baker 1983; Haber and Frankie 1989). According to some authors (e.g. Minet 1994; Carcasson and Heppner 1996; Lemaire and Minet 1998; Kitching and Cadiou 2000), the hawkmoths are classified in three subfamilies each with at least one genus occurring in Brazil. Smerinthinae has three tribes (sensu Kitching and Cadiou 2000), but only Ambulycini is represented in Brazil, comprising the genera Adhemarius Oiticica, 1939, Orecta Rothschild & Jordan, 1903 and Protambulyx Rothschild & Jordan, 1903. Sphinginae includes the tribes Acherontiini and Sphingini. Agrius cingulata (Fabricius, 1775), a very common and recognizable species, is the only acherontine hawkmoth recorded to the New World (Kitching 2002). In the same region, the Check List 4(2): 123–136, 2008.

Hawkmoths have long been recognized as major pollinators of flowering plants (Baker 1961;Gregory 1963;Silberbauer-Gottsberger and Gottsberger 1975;Janzen 1984;Haber and Frankie 1989;Darrault and Schlindwein 2002;Kitching 2002).Because of their capability to fly far away, these moths are potential long distance pollen dispersers (Linhart and Mendenhall 1977;Nilsson et al. 1992;Chase et al. 1996).They may be diurnal, crepuscular or nocturnal.In South America, however, most species are more active at night, when they may be easily observed visiting flowers with white or pale corollas, very long tubes or spurs and copious amounts of nectar (Silberbauer-Gottsberger and Gottsberger 1975;Baker and Baker 1983;Haber and Frankie 1989).
Since sphingids can easily be recorded by lighttrapping and are taxonomically well known (Kitching and Cadiou 2000), they have served as model organisms in a number of diversity studies (e.g.León-Cortés et al. 1998).Considering that the sphingids of the state of São Paulo are still poorly known, the purpose of this work is to document the composition, relative abundance, and phenology of the sphingid fauna of a natural reserve at Boracéia, municipality of Salesópolis, which is well preserved and located approximately 80 km east of the city of São Paulo.A long-term survey with a smaller and better known group of macrolepidopterans, viz.Saturniinae (Saturniidae), has revealed how important this reserve may be to the maintenance of the lepidopterofauna of São Paulo (as well as for other organisms, see Heyer et al. 1990).Of the 11 saturniine species hitherto recorded in the state of São Paulo (data gathered from Lemaire 1978), 10 have been sampled in Boracéia (M.Duarte and collaborators, in progress).

Study Site
The survey of the hawkmoths (Sphingidae) of Boracéia was conducted at the Estação Biológica de Boracéia (EBB), municipality of Salesópolis, state of São Paulo, Brazil (23 o 38'S, 45 o 52'W; 900 m a.s.l.; see map in Heyer et al. 1990).This station has been supported and administrated by the Museu de Zoologia, Universidade de São Paulo (details on the site http://www.mz.usp.br).
The EBB is in a 16,450 ha reserve of the Companhia de Saneamento Básico do Estado de São Paulo.Even before its establishment in March of 1954, the reserve has been a reference site for anyone interested in moths and butterflies of Atlantic Rainforest (Travassos and Camargo 1958).According to Setzer (1946), Boracéia is among the wettest areas in Brazil; average annual rainfall is usually above 1,500 mm (Leemans and Cramer 1991).The vegetation is relatively continuous except for the gaps formed by the narrow dirt access road, the aqueduct line, several small rivers and the small man-made clearings around the station itself (Heyer et al. 1990).The forest has a low canopy, averaging ca.5-10m.Palm trees (especially Euterpe edulis), tree ferns and giant bamboos (Merostachys) are common.The understory is relatively open in most parts of the forest, being more dense along streams, where the presence of the plant genus Heliconia is characteristic (Bertoluci and Rodrigues 2002).There are 240 species of trees, 130 shrubs, 115 epiphytes, 90 lianas, and 89 herbs in the area (Wilms 1995 apud Bertoluci and Rodrigues 2002).

Data Collection
The first and most extensive collections of Lepidoptera from Boracéia were made by the entomologists Romualdo Ferreira D'Almeida and Lauro Travassos Filho in the 1940's and 1950's (Travassos and Camargo 1958).However, only in 1948 and 1949, the hawkmoths were monthly sampled, with 78 and 85 days of field work, respectively (Travassos and Camargo 1958;M. Duarte and collaborators, in progress).Moth sampling extended through the following decades until 2004 (Table 1).The hawkmoths were attracted with mixed mercury vapor bulbs and manually collected on the walls of the scientist's residence (one of the EBB's buildings; for details see Travassos and Camargo 1958).They were killed by direct injection of aqueous ammonia solution in the thorax (specimens collected before 2004 may have been killed with different techniques -see Winter Jr. 2000).All specimens sampled from September of 1940 to April of 2004 were sorted and identified, and their records were included in a digitized database, which is intended to be of free public access through the World Wide Web.

Results and discussion
A total of 2,064 individuals (473 females and 1591 males) belonging to 3 subfamilies, 6 tribes, 23 genera, and 75 species were sampled in the EBB, from 1940 to 2004 (Tables 2 and 3).Macroglossinae is the most abundant subfamily with 66 % of the collected material, being followed by Sphinginae (23 %), and Smerinthinae (11 %).
The species frequency distributions are shown in Figure 2 (see also Table 4).As is noted, the number of rare species is much higher than that of common species.This is typical of organism communities in a complex and relatively stable ecosystem (Preston 1948;1960).--------------------Xylophanes t. thyelia (Linnaeus, 1758) (Macroglossinae: Macroglossini) was the most collected hawkmoth in the EBB (Figure 3).This species occurs year-round with Adhemarius eurysthenes (R. Felder, [1874]) (Smerinthinae: Ambulycini) (Table 5).Six other species appear to be constant in the EBB, although they have not been recorded in only one month during the 64 years of sampling (Table 3)

Figure 1 .
Figure 1.Relative proportions in the number of individuals and species collected per subfamilies and tribes at the Estação Biológica de Boracéia, Salesópolis, São Paulo, Brazil, from 1940 to 2004.A, relative abundance; B, species richness.

Figure 2 .
Figure 2. Species frequency distributions (observed and calculated by means of Fisher's log series) of hawkmoths (Lepidoptera: Sphingidae) with different number of individuals collected at the Estação Biológica de Boracéia, Salesópolis, São Paulo, Brazil.