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Structure and natural history of an assemblage of bats from a xerophytic area in the Caatinga of northeastern Brazil a

b

c

Roberto Leonan Morim Novaes , Rafael De Souza Laurindo & Renan De França Souza a

Campus Fiocruz da Mata Atlântica, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil

b

Departamento de Biologia, Universidade Federal de Lavras, Minas Gerais, Brazil

c

Programa de Pós-Graduação em Ecologia e Evolução, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil Published online: 20 Feb 2015.

Click for updates To cite this article: Roberto Leonan Morim Novaes, Rafael De Souza Laurindo & Renan De França Souza (2015): Structure and natural history of an assemblage of bats from a xerophytic area in the Caatinga of northeastern Brazil, Studies on Neotropical Fauna and Environment, DOI: 10.1080/01650521.2015.1006478 To link to this article: http://dx.doi.org/10.1080/01650521.2015.1006478

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Studies on Neotropical Fauna and Environment, 2015 http://dx.doi.org/10.1080/01650521.2015.1006478

Structure and natural history of an assemblage of bats from a xerophytic area in the Caatinga of northeastern Brazil Roberto Leonan Morim Novaesa*, Rafael De Souza Laurindob & Renan De França Souzac a

Campus Fiocruz da Mata Atlântica, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil; bDepartamento de Biologia, Universidade Federal de Lavras, Minas Gerais, Brazil; cPrograma de Pós-Graduação em Ecologia e Evolução, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil

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(Received 2 July 2014; accepted 8 January 2015) The Caatinga biome is restricted to Brazil, and its bat fauna is among the least studied in South America, with scarce information on species occurrence, distributions, and structure of assemblages. Moreover, most of the information available on bats from this biome comes from relicts of other ecosystem formations. From 2010 to 2012 we conducted bat surveys in different sites along the Serra da Jitirana, a xerophytic locality in the Caatinga of Piauí state, northeastern Brazil. We recorded 20 species in six families. Representatives of animalivorous guilds predominated in both the number of individuals and species. We speculate that the low numbers of frugivores is a response to the environmental constraints imposed by the drought. Along with an analysis of this assemblage, we also report here new information on roosts, behavior, and feeding items for several species about which little is yet known. Keywords: Chiroptera; feeding habits; roost; seasonality; species richness

Introduction The Caatinga biome covers c.740,000 km2 in northeastern Brazil, and is mostly characterized by its semiarid climate. The region is undergoing exceptional habitat loss. Around 30% of its coverage has been drastically altered by human activities, turning the area into the third most altered Brazilian biome (Myers et al. 2000; Leal et al. 2005). The Caatinga is among the least studied South American biomes (Santos et al. 2011), and for a long time its biota was considered a simplified subset of that from Cerrado (Mares et al. 1981, 1985; Willig and Mares 1989), receiving fewer conservation resources than other biomes (Tabarelli and Silva 2003; Santos et al. 2011). However, after more careful analyses, previous hypotheses of an impoverished fauna with low endemism have been refuted, and a new scenario has emerged, with a rich and peculiar assemblage of mammals associated to open habitats (see Oliveira et al. 2003; Carmignotto et al. 2012). As this rich biodiversity been revealed, the evolutionary history and zoogeography of this fauna has become the focus of several recent studies (e.g. Carmignotto et al. 2012; Nascimento et al. 2013). Checklists of mammals report 153 species in 10 orders, with 6.5% of endemism (Paglia et al. 2012). With the intensification of research, these numbers have grown for bats and other taxa, with new occurrence records and undescribed species emerging *Corresponding author. Email: [email protected] © 2015 Taylor & Francis

(Gregorin and Ditchfield 2005; Gregorin et al. 2006; Taddei and Lim 2010; Moratelli et al. 2011; Novaes et al. 2013a). Although several studies have focused on Caatinga bats (e.g. Mares et al. 1981; Willig 1983, 1985a; Gregorin and Mendes 1999; Guedes et al. 2000; Silva et al. 2004; Gregorin et al. 2008; Sá-Neto and Marinho-Filho 2013), capture efforts were much lower than for other Brazilian biomes (Bernard et al. 2011a). Also, most surveys are based on low sampling efforts, resulting in preliminary species lists (e.g. Silva et al. 2001, 2004; Gregorin et al. 2008). We claim that additional research is necessary to better understand the structure of assemblages and species distributions, both essential to define conservation strategies for this region. In this context, we report here the results of a two-year field study in the Serra da Jitirana, Piauí state. Along with an analysis of the bat assemblage, we present insights for the conservation of Caatinga bats, and new information on natural history for several species whose biology is still very poorly known. Materials and methods Study area Fieldwork was conducted in the Serra da Jitirana, a xerophytic area in São João do Piauí, Piauí state,

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northeastern Brazil. Köppen’s climate classification is semiarid BSh (Peel et al. 2007); and the vegetation is classified into two prevailing physiognomies, Savana Estépica Arborizada (arboreal vegetation) and Savana Estépica Parque (shrub vegetation), both peculiar of the Caatinga sensu stricto (IBGE 2012) (Figure 1). Four localities were selected for sampling (Table 1). They are adjacent to the Piauí River, and located c.30 km away from the Serra da Capivara National Park. This region is currently classified as a priority area for conservation in the Caatinga, and as of extreme biological importance (Tabarelli and Silva 2003). The region contains countless natural cavities formed by sandstone massifs. Sampling sites covered large areas of preserved Caatinga, forming a mosaic of different fragments, which included continuous areas (>1000 ha), and agricultural properties converted into pastures. Some areas had small artificial ponds to store water during the dry season.

Sampling and analysis Fieldwork protocols followed the guidelines of the American Society of Mammalogists (Animal Care and Use Committee of the American Society of Mammalogists 2011). Samplings were split into five six-day expeditions (totaling 30 sampling nights), covering the rainy and dry seasons, and different moon phases. Twelve sampling nights were in the rainy season (February and March of 2010 and 2011) and

Table 1. Sampling sites in the Serra da Jitirana, São João do Piauí municipality, Piauí state, northeastern Brazil. Locality Açude dos Jacarés Sítio União Vila São Domingos Açude Paraíso

Coordinates 08°19ʹ43″ 08°15ʹ37″ 08°18ʹ28″ 08°19ʹ03″

S, S, S, S,

42°21ʹ17″ 42°05ʹ35″ 42°01ʹ19″ 42°19ʹ24″

Altitude (m) W W W W

270 379 388 231

Figure 1. Sampling localities in the Serra da Jitirana, São João do Piauí municipality, Piauí state, northeastern Brazil: (A) Açude dos Jacarés, shrubby vegetation with artificial pond; (B) Vila São Domingos, sandstone plateau with shrubby vegetation; (C) Sítio União, arboreal vegetation; (D) Açude Paraíso, open area with prominent presence of cacti. Photos: Roberto L.M. Novaes.

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Studies on Neotropical Fauna and Environment 18 sampling nights were in the dry season (October) of 2010–2012. In each sampling site, 10 ground level mist nets (9 m × 3 m, 20 mm mesh) were set up on trails and clearings in the vegetation and along the margins of artificial ponds (following Kunz and Kurta 1988). Mist nets were opened at dusk and closed six hours later, covering the period of highest activity of most species (Aguiar and Marinho-Filho 2004). Bats were identified based on criteria of Vizotto and Taddei (1973), Simmons and Voss (1998), Gregorin and Taddei (2002), Reis et al. (2007), Peracchi et al. (2010), and Moratelli et al. (2011). Most individuals were banded and released at the place of capture. Specimens of all species were collected and deposited at the Museu Nacional – Universidade Federal do Rio de Janeiro (IBAMA permit 02001.001113/2008–11, authorization no. 192/ 2010; Appendix). The sampling effort was calculated following Straube and Bianconi (2002), and capture efficiency was estimated dividing the total number of captures by the sampling effort. The species accumulation curve with confidence interval of 95% was constructed using the software EstimateS 9.0 (Colwell 2013). The maximum species richness was estimated through the Jackknife-1 method (Shao and Tu 1995), built in software SigmaPlot 12.0. Bats were classified into trophic guilds following Kalko et al. (1996). Roosting, feeding, and other behavioral observations were based on active searches for shelters that were conducted in the morning and in the late afternoon, totaling 96 h of active search. The search was performed along the arboreal Caatinga and near the slope of the plateaus in areas with the presence of fractures in the sandstone rocks (Trajano 1995). When possible, bats were captured in their shelters using nets or tweezers. Individuals found in shelters (captured or just observed) were not considered in the sampling effort and other quantitative analyses. Results Mist net sampling effort comprised 48,600 m2 h, resulting in the collection of 126 individuals in 20 species, 16 genera and six families (Table 2). The capture efficiency was 0.002 bats/m2 h, and there were no recaptures. The species accumulation curve did not reach an asymptote (Figure 2), indicating that we sampled just part of the assemblage, and other species may be added in a more extensive sampling. The richness estimator (Jackknife-1) predicted a maximum richness around 25 species (Figure 3), indicating that our sample is 80% complete. Phyllostomidae accounted for the largest number of species (8 spp.) and individuals (34%). Molossidae

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was represented by four species (14% of the captures), Vespertilionidae by three species (32% of the captures), Mormoopidae and Noctilionidae by two species each (5% and 19% of the captures, respectively), and Emballonuridae by one species (8% of the captures). Four species represented about 49% of the sampling (Peropteryx macrotis, Artibeus planirostris, Noctilio leporinus, Myotis lavali), with dominance of M. lavali (17%) and N. leporinus (16%). The remaining species were represented by a few individuals. Background aerial insectivores (26% of the captures; 6 spp.) and uncluttered aerial insectivores (22%; 5 spp.) were the most abundant and species rich guilds. Gleaning piscivores represented 16% of the sample in one species; gleaning insectivores and frugivores corresponded to 15% of the captures with three species each, and sanguivores and nectarivores represented 3% and 2%, respectively, guilds with only one species each. Phytophagous bats (frugivorous and nectarivorous; 16%) were less abundant than the insectivores (63%). Mist nets over water bodies proved to be essential for sampling, accounting for 56% of all captures. Sampling at the edge of ponds resulted in 90% of the captures of piscivorous bats, and 84% of the captures of insectivorous bats. Seven out of the 13 insectivorous species recorded (Pteronotus gymnonotus, Noctilio albiventris, Eumops perotis, Cynomops planirostris, Molossus molossus, Lasiurus ega, and Myotis riparius) were captured exclusively foraging over ponds, and three species (Pteronotus parnellii, Neoplatymops mattogrossensis, and Myotis lavali) had more than 75% of the captures in mist nets over water. Seasonal differences in species richness and abundance of captured bats were observed, with the rainy season yielding twice as many species and captures than the dry season (Figure 4). There was a seasonal variation in the abundance of different trophic guilds (Figure 5), with a lower abundance of captured bats, mostly insectivorous species, in the dry season. Observations on roosts and diet A colony composed of at least six Neoplatymops mattogrossensis was found roosting under a flat stone on the slopes of a sandstone plateau with scrub vegetation and abundant cacti. A 1.5 cm wide horizontal slit between the stone and the ground formed the entrance (Figure 6). Fragments of Diptera, Hymenoptera, Isoptera, and other unidentified insects were found in feces collected in the roost. A colony of Molossus molossus was found in a 3 cm wide slit in a sandstone rock 2 m above the

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R. L. M. Novaes et al. Table 2. Bats from Serra da Jitirana, São João do Piauí municipality, Piauí state, northeastern Brazil. Species were assigned to trophic guilds according to Kalko et al. (1996): UAI: uncluttered space/aerial insectivore, SAN: highly cluttered space/gleaning sanguivore, NEC: highly cluttered space/gleaning nectarivore, GIN: highly cluttered space/gleaning insectivore, FRU: highly cluttered space/gleaning frugivore, BAI: background cluttered space/aerial insectivore, PIS: highly cluttered space/gleaning piscivore. Captures

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Taxon Emballonuridae Peropteryx macrotis (Wagner, 1843) Phyllostomidae Desmodus rotundus (É. Geoffroy, 1810) Glossophaga soricina (Pallas, 1766) Micronycteris sanborni Simmons, 1996 Mimon crenulatum (É. Geoffroy, 1810) Trachops cirrhosus (Spix, 1823) Carollia perspicillata (Linnaeus, 1758) Artibeus lituratus (Olfers, 1818) Artibeus planirostris Spix, 1823 Mormoopidae Pteronotus gymnonotus Natterer, 1843 Pteronotus parnellii (Gray, 1843) Noctilionidae Noctilio albiventris Desmarest, 1818 Noctilio leporinus (Linnaeus, 1758) Molossidae Cynomops planirostris (Peters, 1865) Eumops perotis (Schinz, 1821) Molossus molossus (Pallas, 1766) Neoplatymops mattogrossensis (Vieira, 1942) Vespertilionidae Lasiurus ega (Gervais, 1856) Myotis lavali Moratelli et al. (2011) Myotis riparius Handley, 1960 Total

ground. The slit was about 6 m above the ground, and during the twilight period some individuals left the roost from the topmost part of the slit and raised flight. Three individuals of Micronycteris sanborni (1 ♂, 2 ♀♀) were captured in a cut-and-burned hollow of a mango tree (Mangifera indica L., Anacardiaceae). Inside the roost, we found fragments of partially consumed Lepidoptera, Ephemeroptera, Orthoptera, Blattodea, and Coleoptera. Analyses of feces of three males of Artibeus planirostris revealed one with mandacaru seeds (Cereus jamacaru Dc., Cactaceae), and an undetermined yellow fruit pulp; another with the same yellow fruit pulp and undetermined fragments of insects; and a third with seeds of Vismia sp. (Clusiaceae). A female of Mimon crenulatum captured in a mist net had a partially consumed cockroach (Blattodea) in its mouth.

Trophic guild

Dry

Rainy

Total

UAI

8

2

10

SAN NEC GIN GIN GIN FRU FRU FRU

1 1 3 1 — 1 2 5

3 1 6 8 1 5 6

4 2 9 9 1 6 2 11

BAI BAI

— 1

2 4

2 5

BAI PIS

1 12

3 8

4 20

UAI UAI UAI UAI

— 1 3 3

1 5 5

1 1 8 8

BAI BAI BAI

— 3 — 46

1 18 1 80

1 21 1 126

—

—

One individual of Glossophaga soricina mist netted in front of a flowering mandacaru had pollen adhering to its fur.

Discussion Assemblage structure Although our capture efficiency was low, the species richness was within that commonly achieved in other short-term surveys in the Caatinga (see Silva et al. 2004; Gregorin et al. 2008). In the Caatinga lower numbers of captured bats than in Neotropical forests and savannas have been reported (e.g. Silva et al. 2001, 2004; Gregorin et al. 2008; Zortéa and Alho 2008; Esbérard et al. 2010). We believe that these findings reflect a low abundance of bats in the Caatinga which may be related to the scarcity of resources, especially food and water, during the dry

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Figure 2. Species accumulation curve of the bat community assessed by mist netting in the Caatinga area of Serra da Jitirana, Piauí, northeastern Brazil. Note: Sobs is a statistical term for the observed curve.

season (Prado 2003), hindering the maintenance of high population densities (Sá-Neto and MarinhoFilho 2013). The higher abundance of phyllostomids was expected considering that this family is the most diverse among Neotropical bats, representing c.50% of the 178 Brazilian species (Nogueira et al. 2014), with frugivorous bats more easily captured in mist nets (Portfors et al. 2000). In addition, some species of phyllostomids (such as A. lituratus, A. planirostris, and C. perspicillata) are among the most frequently captured in inventories across different biomes, including the Caatinga (see Bernard and Fenton 2002; Esbérard 2003; Bordignon 2006; Faria 2006; Gregorin et al. 2008; Zortéa and Alho 2008; Camargo et al. 2009; Oprea et al. 2009). However, it is important to note that our study reported lower abundance of phyllostomids compared with other studies in Caatinga (Mikalauskas 2005; Gregorin et al. 2008; Rocha 2010; Sá-Neto and Marinho-

Filho 2013). Phytophagous bats (frugivorous and nectarivorous) were less abundant than the insectivores, and this pattern contrasts with other studies that have found phytophagous bats the most abundant (Gregorin et al. 2008; Sá-Neto and Marinho-Filho 2013). As a possible explanation, this sampling was performed during one of the longest droughts in the Caatinga’s history (Novaes et al. 2013b), which may have been biasing results, including low capture rates. Prado (2003) indicates that the Caatinga has low primary productivity, which may restrict the supply of food for phytophagous animals, especially during periods of prolonged drought, affecting population densities of several species (Brown et al. 2004). The study conducted by Gregorin et al. (2008), in a large area of primary Caatinga of Piauí, found a greater abundance of fruit-bats, differing from our results, where the most abundant species were insectivorous bats. This may be associated with the degree of conservation of the sampled area, since the fruit

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Figure 3. Species accumulation curve with richness estimator index (Jackknife-1) of the bat community assessed by mist netting in the Caatinga area of Serra da Jitirana, Piauí, northeastern Brazil.

trees in the Caatinga are the first to be removed during environmental exploitation (Prado 2003; Leal et al. 2005), reducing the availability of food for phytophagous species. This pattern is in stark contrast to tropical forests, where moderate disturbance of the forest may favor frugivorous generalist species due to the rapid process of ecological succession (Bierregaard Jr. et al. 1992; Herrera et al. 1994; Brosset et al. 1996; Schulze et al. 2000). Thus, we suppose that intense environmental disruption in the Caatinga areas can cause a drastic reduction in the richness and abundance of fruit-bats. The conspicuous seasonal variation in the bat assemblage composition is expected since semiarid regions usually present climatic seasonality (Lumsden and Bennet 1995; Prado 2003). The highest abundance in the rainy season is due to the concentration of zoochorous fruit production in that period, which implies greater availability of resources for the phytophagous bat fauna (Prado 2003).

The dominance of M. lavali was not expected considering that insectivorous species are generally reported in low numbers in biological inventories, which is probably a bias of the use of ground level mist nets as the predominant method in species surveys (Portfors et al. 2000). Yet, several studies indicate higher capture rates of insectivorous bats, especially molossids and vespertilionids, on water body margins (Myers and Wetzel 1983; Esbérard 2003; Silva 2007; Lourenço et al. 2010; Costa et al. 2012). Thus, we attribute our successful capture rates of M. lavali (and N. leporinus) to the placing of mist nets in in a V-shape, on the edge of water bodies where these bats forage. The high species richness achieved in this study seems to be associated with the many natural cavities in sandstone massifs, most of them potential roosts for several species. The availability of roosts is an important factor that influences the composition of bat communities in a region, and the presence of

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Figure 4. Comparison of species richness (line) and abundance (bars) of bats recorded with mist nets during six sampling days in the Caatinga area of Serra da Jitirana, Piauí, northeastern Brazil.

natural cavities allows the coexistence of dozens of species (Bredt et al. 1999). Additionally, sandstone caves, such as those found in Serra da Jitirana, may contribute to the high abundance of several bat species, increasing the local diversity (Trajano 1995). The non-stabilization of the species accumulation curve suggests that continuous sampling will increase the number of species. It indicates that additional effort is necessary to adequately sample this locality. Although we have recorded only part of the assemblage (estimated at about 80%), this first assessment is important to further understand the structure of local assemblages and species occurrence and abundance in the Caatinga. Natural history notes Neoplatymops mattogrossensis had already been found roosting in horizontal and vertical slits in rocks in the Caatinga (Willig 1985b; Novaes et al. 2013a) and in the Atlantic Forest (Avilla et al. 2001). These authors found that roosts ranged from 0.5 to 5 m above ground level. However, this is the

first record of this species roosting in a slit at ground level. This is unexpected, because most molossids (except for Tadarida brasiliensis [I. Geoffroy, 1824]; De Knegt and Silva 2007) are unable to take flight from the ground (Dietz 1973). Therefore, it is possible that N. mattogrossensis also has this ability. The little information available about the diet of N. mattogrossensis indicates an insectivorous diet (Willig and Jones 1985). Although Willig (1985b) reported Coleoptera as a common item in the diet of N. mattogrossensis, we did not find evidence of this group in the feces of N. mattogrossensis in the Serra da Jitirana, indicating that the diet of this species has local variations. Molossus molossus is known to occupy various types of roosts, from hollow trees to the roof lining of residences and other human constructions (Fabian and Gregorin 2007), but this is the first record of M. molossus using sandstone rocks as roosts. The behavior of climbing to the highest point of the shelter for takeoff is common in most molossid bats since they are unable to take flight from the ground (Dietz 1973).

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Figure 5. Comparison of the total number of bat captures among trophic guilds and seasons in Caatinga area of Serra da Jitirana, Piauí, northeastern Brazil.

Micronycteris sanborni is known from few individuals, and its ecology and behavior is poorly known (Nogueira, Peracchi, et al. 2007). Due to its occurrence in areas with rocky outcrops, caves were expected to be the roost for this species (Nogueira, Peracchi, et al. 2007). The use of hollow trees by M. sanborni differs from that expected by Simmons (1996) and Nogueira, Peracchi, et al. (2007). Based on the habits of its congeners, an insectivorous diet was expected for the species, but this is the first record of items consumed. This record is also the first one for the species in an area of xeric shrub in the Caatinga (similar to that shown in Figure 1D), and the first description of roost use for this species. According to Simmons (1996), although this species is endemic to the open biomes such as Caatinga and Cerrado, its occurrence seems to be associated with mesic environments, which was corroborated by Nogueira, Peracchi, et al. (2007). Recently, Bolivian savanna populations of this species were assigned to a new species, Micronycteris yatesi Siles et al., 2013 (see Brooks et al. 2002; Siles et al. 2013). Based on the occurrence in the same biogeographic province and

similar vegetation structure, we recommend a reassessment of material assigned to M. sanborni from the Brazilian savanna of Mato Grosso do Sul (see Santos et al. 2010), which may correspond to M. yatesi. Artibeus planirostris is considered a predominantly frugivorous species that complements its diet with other plant items (nectar and leaves) and insects (Zortéa 2007). In an ecotone of Cerrado and Caatinga, A. planirostris maintained its diet based almost exclusively on the consumption of Vismia fruits, although they occasionally also consume insects (Willig et al. 1993). The record we made about the consumption of C. jamacaru fruits is new for this species. Artibeus planirostris has a great potential to act as seed disperser in semiarid areas in Venezuela (August 1981) and other localities in South America (Hollis 2005; Oliveira and Lemes 2010). Fruit bats have an important role in the seed dispersal of cacti in semiarid areas in northern South America, and bats from the Caatinga may also have an important role in the seed dispersion (Ruiz and Soriano 2000).

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Figure 6. Roost of Neoplatymops mattogrossensis under a stone on the ground (arrow). Photo: Roberto L.M. Novaes.

The diet of M. crenulatum consists mainly of insects, although it can also feed on other arthropods, small vertebrates and plant items (Pedro et al. 1994; Emmons and Feer 1997; Nogueira, Peracchi, et al. 2007). Among insects, the consumption of Coleoptera, Lepidoptera, Diptera, Hemiptera, and Orthoptera has been reported (Whitaker and Findley 1980; Humphrey et al. 1983; Rivas-Pava et al. 1996), but this is the first record of Blattodea in the diet of M. crenulatum. Glossophaga soricina is a nectarivorous species that complements its diet with fruits and insects (Gardner 1977; Willig et al. 1993). This species pollinates hundreds of plant species, especially in Bignoniaceae, Bombacaceae, Bromeliaceae, Gentianaceae, Gesneriaceae, Leguminosae, Lythraceae, Myrtaceae, Passifloraceae, and Tiliaceae (Nogueira, Dias, et al. 2007). Consumption of cactus nectar is common among nectarivorous bats, which have a prominent position in the reproduction of these plants in many arid and semiarid regions of South America (Nassar et al. 1997, 2003; Ruiz and Soriano 2000). Understanding of the role of bats in the pollination of plants from the Caatinga is still incipient, and based on

the record of G. soricina with pollen of C. jamacaru stuck in its pelage, we hypothesize that this bat has a role in the reproduction of this cactus. This is the first record of G. soricina visiting (and possibly pollinating) mandacaru.

Acknowledgments Ana Cláudia Delciellos, Carlos Cândido, Igor Catharino de Souza, Eduardo Felberg, William de Paula, Thiago Marques, Mara Silva, and Piktor Benmaman provided assistance during the fieldwork. Ruby Malzoni reviewed the English usage. Dossel Ambiental Consultoria e Projetos Ltda provided support for the fieldwork. Ricardo Moratelli revised the original manuscript, contributing to improving the quality of this article.

References Aguiar LMS, Marinho-Filho J. 2004. Activity patterns of nine phyllostomid bat species in a fragment of the Atlantic Forest in southeastern Brazil. Rev Bras Zool. 21:385–390. doi:10.1590/ S0101-81752004000200037 August PV 1981. Fig-fruit consumption and seed dispersal by Artibeus jamaicensis in the Llanos of Venezuela. Biotropica. 13:70–76. doi:10.2307/2388073 Avilla LS, Rozenzstranch AMS, Abrantes EAL. 2001. First record of the South American flat-headed bat, Neoplatymops

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Appendix Bats from Serra da Jitirana, São João do Piauí municipality, Piauí state, deposited in the Museu Nacional (MN), Rio de Janeiro. Emballonuridae: Peropteryx macrotis (MN75195, MN75196). Phyllostomidae: Artibeus lituratus (MN75198), Artibeus planirostris (MN75192), Carollia perspicillata (MN75193), Desmodus rotundus (MN78407), Glossophaga soricina (MN79933), Micronycteris sanborni (MN75194, MN75211, MN78405), Mimon crenulatum (MN75189), Trachops cirrhosus (MN78408). Mormoopidae: Pteronotus gymnonotus (MN75207, MN78406), Pteronotus parnellii (MN75190, MN78410). Noctilionidae: Noctilio albiventris (MN78411, MN78409, MN79943); Noctilio leporinus (MN75204). Molossidae: Eumops perotis (MN75199), Cynomops planirostris (MN75209, MN75215, MN75216, MN75218), Molossus molossus (MN75197, MN75210), Neoplatymops mattogrossensis (MN75200, MN75212, MN75213, MN75214, MN78412). Vespertilionidae: Lasiurus ega (MN75206), Myotis lavali (MN75191, MN75205, MN75208, MN75217, MN79926, MN79929, MN79932, MN79935, MN79936, MN79938, MN79950), Myotis riparius (MN79930).