Check List 10(1): 10–17, 2014 © 2014 Check List and Authors Chec List ISSN 1809-127X (available at www.checklist.org.br) Journal of species lists and distribution pecies S

of The Social Community (, ) in Olga Coutinho Togni 1* 1 1 ists 2 L anSilveira Area of Atlantic Forest, Ubatuba, , Gabriela de Almeida Locher , Edilberto Giannotti and Orlando Tobias

1 Universidade Estadual Paulista,[email protected] Instituto de Biociências, Departamento de Zoologia, CEP 13506-900, Rio Claro, SP, Brazil. 2 Museu Paraense Emílio Goeldi, Campus de Pesquisas, Departamento de Zoologia, CEP 66040-170, Belém, PA, Brazil. * Corresponding author. E-mail: Abstract:

The Brazilian Atlantic Forest is of great relevance to biological conservation, and is among the areas in with the highest levels of diversity and endemism. The aim of this study was to survey the social wasp species in the subfamily in Ubatuba, São Paulo state, in southwestern Brazil. Collecting work was conducted from May 2007 to May 2008 using attractive PET bottle traps and active searching. Twenty-one species belonging to eight genera were found, among which some may be considered rare in southeastern Brazil such as parallelogrammus and catillifex. The most abundant species were angulata (64.31%), Agelaia nr. centralis (10.08%) and (8.49%). A correlation between species richness and relative humidity (r = 0.6435; p = 0.0176) was observed. Values of species richness were a little higher in the super humid (Sm = 11) than in the less humid (Sm = 9) season. This suggests that this season may have more favorable environmental conditions for a greater richness of species to found colonies. Despite not having a very high species richness compared with other surveys, the collected species in this study can be considered rare in southeastern Brazil, emphasizing the complexity of the Atlantic Forest biome and its relation to the diversity of .

Introduction 2005; Hermes and Köhler 2006; Santos et al. 2007; Souza et al. 2008; Clemente 2009; Gomes and Noll 2009; Lima et al. 2010; Prezoto and Clemente 2010; Tanaka Junior and Social wasps (Vespidae, Polistinae) are remarkably Noll 2011), and the Amazon Forest (Raw 1998; Silveira abundant in Brazil (Raveret-Richter 2000), acting as 2002; Silveira et al. 2005; Morato et al. 2008; Silveira et al. 1975;predators, Gadagkar 1991; collectors Raw 1997) (Suzuki in the1978; food Carpenter chain. and 2008; Silva and Silveira 2009). Marques 2001; Aguiar and Santos 2007), and prey (Jeanne conservation and rational use (Melo et al. 2005). By Due to the importance and abundance of the taxon, Inventorying an area is the first step towards its studies on the diversity and abundance of social wasps have been conducted across different regions and suchundertaking as colony a survey seasonality, of species foragingof social wasps, activity, several and environments, such as etcultivated al. 1993; fields Santos and 1996; human- Lima aspects of their biology and behavior must be considered, modifiedet al. environments et (Rodrigues al. and Machado 1982; SantosMarques et 1989;al. 2009a; Marques Santos et al. et al. nesting habits. Brazilian rainforests represent a peak of 2010; 2000;Auad et Marques al. 2010; Santos 2005; and RibeiroPresley Junior2010; Souza 2008; biodiversity, and inet thisal. 1998), sense, including certain localities in their possessdiversity the of et al. 2011; Tanaka Junior and Noll 2009b; 2011; Alvarenga Locher 2012), greatest abundance and richness of known species on the planet (Machado Brazilian savanna) (Richards 1978; Diniz and Kitayama social wasps (Richards 1978). in areas of “cerrado” (semi-deciduous et al. tropical2005; Souza forest and or the The genera social Mischocyttarus wasps in Brazil, have and the Polybia asynchronous et al. 2007; Souza et al. colony cycle. This pattern has been well-described for 2008;1994; SantosMechi 1996; et al. 2005; Melo et al. Simões et al. (Gobbi et al. SimõesPrezoto et 2006; al. Elpino-Campos and Zucchi 1980; Simões and Mecchi 1983; Gobbi et al. 1984;1995; 2009b; Lima 2010; Henrique- Giannotti 1998). 1985; On Gobbi the other and Simões hand, the1988; foraging Marques activity 2011; Tanaka Junior and Noll 2011), “campos 1992; Giannotti and Machado 1994; Giannotti rupestres” montane savannas (characteristic vegetation of areas above 800-900 m of altitude that presents open of Polistinae may be primarily limited by physical factors Clementevegetation 2009; with aPrezoto stratum and of Clementemonocot herbs,2010), sub-shrubs “caatinga”, (Spradbery 1973), such as light intensity, temperature, air and shrubs of dicots) (Silva-Pereira and Santos 2006; et al. warmesthumidity, andand leastwind humidspeed, hoursand some of the studies day (Giannottihave shown et 2005; Santos et al. 2006; Santos et al. al.that 1995; individual Andrade wasps and Prezotoforage more 2001; intensively Resende et during al. 2001). the (Tropicalforest (Mechi deciduous 1996; Silveira xerophytic et al. woodland)2008; Souza (Melo et al. 2008; Clemente 2009; Souza et al. 2009a), riparian et al. in theThe northern present article coastal presents region theof São results Paulo of state,the first in different environments of the 2010; Atlantic Henrique-Simões Forest (Melo et al. inventory of social wasps in an area of the Atlantic Forest 2011; Pereira and Antonialli-Junior 2011; Locher 2012), southwestern Brazil, emphasizing the importance of this 10 Togni et al. | Social Wasps in Ubatuba, Brazil

Collecting procedures the region. In addition, we sought to analyze and verify group for further studies on the Conservation Biology of Twenty-five points were marked along the studied the existence of relationships between the temporal trail, with a distance of at least 100 meters between each distribution of humidity and temperature and the richness in order to avoid the occurrence of pseudo-replicates; Materialsand abundance and of Methods species identified. that is, individuals from the same wasp colony collected Study area at separate sampling sites. Thirteen monthly collecting The study site was in the Angelim Rainforest, a nature rounds were performed from May 2007 to May 2008, with each consisting of two days of fieldwork at the et al. study site, with a one-week interval between. The reserve in ombrophilous dense forest, located in the Atlantic relative humidity and environmental temperature at Forest biome (Veloso 1991), situated in Ubatuba city, each sampling point were measured by digital thermo- (23°23′ S, 45°03′ W) São Paulo state, southwestern Brazil hygrometer, making it possible to calculate a monthly (Figure 1). The area is privately owned by Paul and Edna average of these climatic variables and correlate them to StateThomsen Park and of contains Serra do a Mar.total Theof 760 farm ha hascovered several by native trails Attractivelocal wasp traps diversity. vegetation. Eighty percent of the area occurs within the (Melo et al. 2001; Souza and Prezoto 2006) were installed that provide access to sites in dense forest, some of which Attractive traps made from two-liter plastic PET bottles accompany the course of the Angelim River that originates in the nearby hills within the property. at each marked point. Four circular holes were made in The climate in the region, according to Köppen’s each bottle and 200 mL of attractive liquid were placed andclassification constant (Köppenhumidity 1948),with an is annual tropical average rainforest humidity with inside. Two types of attractants were used: 1) guava juice, an average annual temperature of approximately 26.72°C, or 2) a blend containing 84 grams of sardines (equivalent to a tin) per liter of water. The traps were placed monthly of 73.45%. The region has two seasons: (1) super humid at each of the 25 sampling points, and removed after one with frequent rains from October to April, and (2) a less week. Individual wasps found in the bottles were collected Morellatohumid season 2002). from May to September with less constant with a sieve and tweezers and placed in vials to be fixed in rainfall, but without any water deficit (Bencke and 70% ethanol.

Figure 1. Location of the study area.

11 Togni et al. | Social Wasps in Ubatuba, Brazil

Active searching Active searching relied on the use of entomological of species in relation et toal. monthly samplings, we used the formula C = P x 100 / N, proposed by Bodenheimer (1955 nets to search for individuals on pathways and trails apud Silveira-Netto 1976), where: in the area, and checking flowers, et al. tree 2007). cavities, Individuals broad- P = number of samples containing a certain species; leafed plants and buildings for the presence of wasps in N = total number of samples; inthe which study the area area (Elpino-Campos was visited for a total of 10 hours of Data were grouped into the following categories: (1) were actively sampled during the two days every month constant species, present in over 50% of the samples; (2) accessory species, present in 25% to 50% of samples (3) Identificationmonthly sampling of collected effort. material accidental species, present in less than 25 % of samples. To verify the existence of significant differences in the richness and abundance of species of wasps collected et al. All specimens were collected in accordance with the during different seasons, the Mann-Whitney U test was permit number 11413-1 of the Brazilian Institute of performed, using the software BioEstat 5.0 (Ayres Environment and Renewable Natural Resources, IBAMA, 2007). In addition, we applied the Spearman correlation and were deposited in the Entomological Collection at the test (BioEstat 5.0) to verify a possible relationship SP,Department Brazil and of at Zoology, the Entomological Universidade Collection Estadual at Paulista Museu monthly.between the diversity of social wasps and environment “Júlio de Mesquita Filho” (UNESP), campus of Rio Claro, temperature and relative humidity, which were measured Results Paraense Emílio Goeldi, Bélem, PA, Brazil. Species were 2007;identified Silveira through et al. the use of dichotomous keys (Richards 1978; Carpenter and Marques 2001; Pickett and Wenzel A total of 2104 social wasp individuals (Vespidae, 2008; Cooper 1997) and by comparison Polistinae) were collected, representing 21 species in eight with specimens in the aforementioned collections. genera (Table 1). orVoucher on the specimens Entomological were deposited Collection on at the Museu Entomological Paraense Table 2 presents the total abundance (A), relative Collection at the Department of Zoology UNESP Rio Claro abundance (RA%), the constancy (C), and richness (S) Agelaiafor species angulata and samples found in the Angelim Rainforest, DataEmílio analysis Goeldi (Table 2). and how they were distributed throughout the year. had the highest abundance (A = 1353), representing more than half of the total number of The relative abundance of each species was calculated individuals collected, with a relative abundance of 64.31%, by dividing the abundance of the species by the total and thus was considered the dominant species. Relative number of individuals collected. To assess the constancy abundances were higher than 1% for the following species: Table 1.

Relative abundance of each species and species richness for each season, less humidRELATIVE (May ABUNDANCE to September) (%) and super humid (October to April). TAXON LESS HUMID SEASON SUPER HUMID SEASON Agelaia angulata 82.60 Agelaia . centralis (Cameron, 1907) 12.17 (Fabricius, 1804) 49.61 (Haliday, 1836) 0.75 0.60 sp. nr 7.47 Angiopolybia pallens (de Saussure, 1854) 0.64 2.49 pallens 0.32 0.17 (Lepeletier, 1836) 2.45 14.14 Polybia bifasciata 0.00 (Fabricius, 1804) Polybia cattilifex 1.28 5.66 de Saussure, 1854 0.43 Polybia fastidiosuscula 0.11 0.60 Moebius, 1856 Polybia ignobilis (Haliday, 1836) 0.32 0.17 de Saussure, 1854 Polybia jurinei 0.53 (Olivier, 1791) 1.07 6.60 de Saussure, 1854 0.34 0.11 0.00 cyanea 0.32 0.09 (de Saussure, 1854) Mischocyttarini (Fabricius, 1775) Mischocyttarus cassununga (von Ihering, 1903) 0.00 0.77 Mischocyttarus parallelogrammus Zikán, 1935 0.69 Mischocyttarus rotundicolis (Cameron, 1912) 0.11 0.43 Mischocyttarus socialis 1.17 0.34 Mischocyttarus wagneri (Buysson, 1908) 0.11 0.00 (de Saussure, 1854) 2.40 Polistini 0.00 0.17 Polistes versicolor (Olivier, 1791) 0.21 2.57 (Fabricius, 1775) Species richness 18 19

12 Togni et al. | Social Wasps in Ubatuba, Brazil

A. nr. centralis Angiopolybia pallens Mischocyttarus cassununga, Polybia occidentalis Polybia catillifex Polybia carnifex and Polistes bifasciata Mischocyttarus socialis (10.08%), Agelaia vicina (8.94%), species were collected and and Polistes versicolor (4.13%), (3.71%), Mischocyttarus wagneri and wereProtopolybia unique Agelaia (1.85%), vicina (1.66%) forexigua this period. In the less humid season, we found 18 (1.52%) species, with The species , in spite of being among beingAgelaia unique angulata for this period. With respect to the the most abundant species, was considered an incidental species that showed relative abundances greater than A.species, multipicta found andin only M. parallelogrammussix samples along the entire period 1.0%, only was moreA. angulata abundant in the studied. Contrary to this situation, the populations of less humid season. Being very abundant and with very presented small populous colonies (Richards 1978), presents relative abundances but were constant species, appearing adaptations to maintain its population during the months in eight and seven of the 13 months Agelaiasampled, angulatarespectively, A. in which food resources are less available, such as nectar (Tablenr. centralis 2). , A. multipicta, Angiopolybia pallens, Polybia and . et al. Furthermore, the attractive liquids: A. nr. centralis, used in catillifexAmong, P. occidentalis the constant, Mischocyttarus species: parallelogrammus, trapsA. vicina, may have Angiopolybia represented pallens an extra Mischocyttarus food source (Elpino- socialis, M. socialis and Polistes versicolor, only Agelaia angulata CamposPolistes versicolor 2007)., PolybiaThe remaining catillifex species and P. occidentalis and A. nr. centralis Angiopolybia pallens M. socialis showed a constancy of 100%, while were collectedMischocyttarus during the super wagneri humid which season. was For collected species was not sampled in June 2007Polybia and with relative abundances less than 1%, itM. is cassununga, noted that, bifasciata was, P.not fastidiosuscula sampled in October, P. ignobilis 2007 (Table, Protopolybia 2). M.excluding parallelogrammus, M. rotundicolis, M. socialis, Polistes exígua,The speciesSynoeca considered cyanea, Mischocyttarus as accidental wagneriwere and carnifexonly once, and all otherP. versicolor independent-founding Polistes carnifex. We collected only one individual of Agelaia Protopolybia exigua, in May 2007, and of were more abundant at the Mischocyttarus wagneri super humid season. As the large populations of the species smallerare better colonies, prepared have toa greater face shortages sensitivity of to resources,decreases in it in August 2007 (Table 2). may be that the tribes Mischocyttarini and Polistini, with In Table 3, the relative abundance of each species and food sources. species richness for each season (less humid and super temperature, humidity and consequently the shortage of Tablehumid) 2. are depicted. In the super humid season, 19 ● □ ■ Total abundance (A), relative abundance (RA%), constancy (C) and richness (S) from May 2007 to May 2008 and species deposition catalogue number. = constant species; = accessory species; X = accidental species; = super humid season; DZRC = Entomological Collection at the Department CATALOGUE ofSPECIES Zoology UNESP Rio Claro;A MPEGRA% = EntomologicalC Collection at Museu ParaenseMONTHS Emílio Goeldi. NUMBER m j j a s o n d j f m a m Agelaia angulata 1353 ● # # # # # # # # # # # # # nr 212 10.08 # # # # # # # # # # # # # A. . centralis 64.31 ● DZRC 1205 - 1219 A. multipicta 0.67 ● # # # # # # # # DZRC 1220 - 1236 11.101.725 MPEG 11.101.714 - A. vicina 1435 1.66 □ # # # # # # 188 # # # # # # # # # # # # Angiopolybia pallens ● DZRC 1237 - 1251 5 # # # # # DZRC 1252 8.94 □ DZRC 1254 Polybia bifasciata 5 0.24 X # # # 11.101.725 MPEG 11.101.714 - P. catillifex 78 0.243.71 ● # # # # # # # # # # 8 0.38 # # P. fastidiosuscula X DZRC 1261 - 1269 5 # # # # P. ignobilis X DZRC 1259 - 1260 9 # # # # P. jurinei 0.24 □ DZRC 1287 - 1287 87 # # # # # # # # P. occidentalis 0.43 ● DZRC 1288 - 1290 Protopolybia exigua 1 4.130.05 X # 1270 - 1286 11.101.725 MPEG 11.101.714 - Synoeca cyanea 0.19 X # # DZRC 1253 Mischocyttarus 49 □ # # # # # # cassununga M. parallelogrammus 12 0.430.57 ● # # # # # # # DZRC 1322 - 1326

M. rotundicolis 5 □ # # # # DZRC 1316 - 1321 11.101.725 MPEG 11.101.714 - M. socialis 39 0.241.85 ● # # # # # # # # # # # # 1 0.05 # MPEG 11.101.713 M. wagneri X DZRC 1302 - 1315 Polistes carnifex 2 0.10 X # # DZRC 1291 P. versicolor 32 1.52 ● # # # # # # # # Number of 186 78 106 99 152 83 100 129 137 162 68 DZRC 1292 - 1301 individuals Richness (S) 212104 13 6 5 40012 10 40410 12 11 11 10 13 10 8

13 Togni et al. | Social Wasps in Ubatuba, Brazil

Although there is no significant difference in species Regarding the number of individuals collected, a slight richness between the two seasons (z = 1.0714, p = 0.2840), seasonal difference (not significant) was observed in one can observe (Figure 2A) that in the super humid mean abundance, with a value of 166.71 individuals at the season (Sm = 11) the average richness was greater than in super humid season and 156.17 in the less humid season. the less humid season (Sm = 9). In the super humid season There was no significant difference in abundance between inthe March. lowest In number the less of humid species season, collected there was was 10, a indecreased October, seasons (z = 0.8571, p = 0.3914), with observed peaks in February and April, and greatest richness was 13 species August and October 2007 (Figure 2B). Also in Figure 2, it can be noted that the environmental richness mainly in the months of June and July, with six and temperature varied little throughout the year, with the five species, respectively. Despite the observed decrease in lowest temperature 23.87°C occurring in June 2007, these months, in May 2007 the richness was equal to the and the highest (29.40°C) occurring in February 2008. highest found in the super humid season, or 13 species. Although lower values of richness, (especially in June and This number is likely due to the fact that the month of July) and lower rates of abundance (such as in May 2008), May represents the beginning of the drier period, and could be noted, probably as a result of the decrease in therefore, wasp colonies have just recently been exposed temperature, an increase in the diversity of social wasps to less favorable conditions. was not observed in the warmer months. By using the Spearman test, it was not possible to establish a significant correlation between temperature and species richness (r = 0.2770, p = 0.3596) or abundance (r = 0.0604, p = 0.8445). However, by examining the values of relative humidity greater than 80% recorded in May and December 2007 and January 2008, greater richness values can be noted (13, 11 coincidentand 11 respectively), with lower while relative in June humidity and July (Figure 2007 2A). and ThisMay 2008, a smaller number of species were collected and are

trend is confirmed by the Spearman test that showed a significant positive correlation between relative humidity and species richness of social wasps in Angelim Rainforest (r = 0.6435, p = 0.0176). Contrary to expectations, in May 2008 the lowest abundance and the lowest humidity were observed. There was no significant correlation between andthese relative variables humidity (r = 0.0604, in this p study, = 0.8445), reflecting the lack of pattern in the relationship between species abundance Discussion

at leastSurvey two studies annual on surveysthis group related of social to thewasps diversity in Brazil of have become relatively frequent (Table 3). Since 2005 the richness of these communities in the Atlantic Forest remainssocial wasps scarce. were published, however, knowledge on Figure 2. The studies of Melo et al. (2005), Hermes and Köhler (2006), Santos et al. (2007), Souza et al. (2008), Gomes Diversity of social wasps related to temperature and relative humidity during the months of the year in Angelim Rainforest, Ubatuba, and Noll (2009), Prezoto and Clemente (2010), Lima et SP: (a) Richness of species (Sm = average richness of each season) and (b) al. (2010), and Tanaka Junior and Noll (2011) addressed Abundance of species (Am = mean abundance of each). Table 3. regions of the Atlantic Forest, reporting quite different Survey studies onN. socialOF waspsN. OFin São Paulo state, Brazil. LOCATIONS AUTHORS SPECIES GENERA richness estimates from the present study. A lack of Rodrigues and Machado studies in São Paulo state is apparent, as evidenced in Rio Claro 33 10 (1982) Table 3. Richards (1978) found 105 species (111 including Santa Rita do Passa 26 8 Mechi (2005) )Apoica in pallens museums around the world originating Quatro from municipalities in the state of São Paulo,Apoica with only pallens the Paulo de Faria 7 species having not been previously listed Pindorama 6 Gomes and Noll (2009) 4 (Rodriguesby him. However, and Machado it should 1982; be Mechi noted 1996; that Locher 2012). Neves Paulista 12 7 4 hadConsidering been listed by the other results authors of inventories for the state carried of São outPaulo in Patrocínio Paulista 30 10 Lima (2010) et al. southeastern Brazil (Rodrigues and Machado 1982; Mechi Magda 20 8 et 13 7 al. et al. 2008; Clemente Matão 13 6 (2011) Bebedouro Tanaka-Junior and Noll 2009;1996; Gomes 2005; Souzaand Noll and 2009; Prezoto Alvarenga 2006; Elpino-Camposet al. 2010; Auad Barretos 19 8 et al.2007; 2010; Ribeiro Lima Junioret al. 2010; 2008; Prezoto Souza and Clemente 2010;

14 Togni et al. | Social Wasps in Ubatuba, Brazil

Souza et al. et al. 2011; Souza et al. 2011; Tanaka Junior and Noll 2011; Locher 2012), it 2010; Henrique-Simões these conditions, but may mean that this period has Agelaia angulata, A. nr. centralis and Angiopolybia pallens “Matabetter doenvironmental Baú” in the conditions,city of Barroso, contributing Minas Gerais to a higher state, wereis observed not collected that the in mostother abundantstudies, with species the latter in Ubatuba, having richness of species to found their colonies. At the localities:

(Santos et al. “Patrocínio Paulista”, SP and “Ibitipoca State Park” in the inbeen the observed inventories in another of the North area ofand Atlantic Northeast Forest regions in Bahia of city of Lima Duarte, MG, the highest species richness values 2007). These three species are more common were observed in the warmer and humid months of the rainforests (Silveira 2002; Melo et al. 2005; Silveira et al. year (Souza and Prezoto 2006; Lima 2008; Ribeiro Junior Brazil,2005; Santos generally et al. covered 2007; Morato by “cerrado” et al. 2008; and Silveira Amazonian et al. 2008; Clemente 2009). Thus, it can be seen in Figure 2B, 2008). that in August and October 2007, which were the months Polybia catillifex and Mischocyttarus with highest abundances, high relative humidity was not parallelogrammus, observed. In the wettest month, January 2008, there was the The Angelim species Rainforest, were not listed in any other a relatively low number of individuals, corroborating which had high abundance M. in the data that humidity and foraging would be inversely parallelogrammus only in the states of Minas Gerais, Rio activityproportional. (Giannotti Moreover, et al. 1995;the high Andrade relative and humidity Prezoto during 2001; desurvey. Janeiro Richards and São (1978) Paulo. reportedThese data the reinforce occurrence the ofneed Resendethis period et al.may 2001). be due to high rainfall that reduces flight for more studies on the northern coast of São Paulo. This

Overall, this study, despite not having observed a very result may be related to the lack of studies in the Atlantic high species richness compared with other surveys, is Forest, but can also be explained by the complexity of lowof great occurrence importance records with inregards southeastern to the need Brazil for suchfurther as this biome. In a study on the community of social wasps Mischocyttarusstudies in the region parallelogrammus and the collection and Polybia of species catillifex with. of three ecosystems of Itaparica Island, in Bahia state (mangrove, ombrophilous dense forest andet al restinga),. 2007). there was a significant correlation between plant diversity Furthermore, this study emphasizes the complexity of the and the diversity of species of wasps (Santos Atlantic Forest biome andAgelaia its relation angulata, to the A. diversity sp. prox. of It was observed that the ombrophilous dense forest has centraliswasps, with and theAngiopolybia frequent collectionpallens of species common a larger and more complex variety of plants than other in the Amazon Forest: biomes studied, and consequently a greater richness of the . A correlation between (Santosfauna, which et al. 2007).can be explained by the heterogeneity of the species richness and relative humidity was also noted, environment and the existence of a wide variety of niches demonstrating the influence abiotic factors can have on those of the Agelaia, showed the greatest populations. In general, species in the Epiponini, especially Acknowledgments:

We thank the Coordenação de Aperfeiçoamento de largeabundances. colonies This that result may have may millions be due to of theindividuals population (Zucchi size LiteraturePessoal de Nível Cited Superior (CAPES) for financial support. etof al. this group of swarm-founding wasps with medium to

Aguiar, C.M.L. and G.M.D.M. Santos. 2007. Compartilhamento deNeotropical recursos 1995). Moreover, in swarm-founding wasps, a greater Entomologyflorais por vespas sociais (Hymenoptera: Vespidae) e abelhas tospecialization a more effective occurs defense among of individuals the colony in(Jeanne a colony, 1991). which (Hymenoptera: ) em uma área de caatinga. reduces the chance of of the queen and contributes 36: 836–842. Alvarenga,in Southeastern R.D., M.M. Brazil. De Castro, Sociobiology H.H. Santos-Prezoto and F. Prezoto. 2010. Andrade,Nesting F.R. of and social F. Prezoto. wasps (Hymenoptera,2001. Horários Vespidae)de atividade in forrageadoraurban gardens e The tribes Mischocyttarini and Polistini are Polistes ferreri 55(2): 445–452. independent-founding and have colonies composed Revista materialBrasileira coletado de Zoociências por Saussure, 1853 (Hymenoptera, of only a few dozen wasps (Richards 1978; Gadagkar Auad,Vespidae), A.M., C.A. nasCarvalho, diferentes M.A. Clemente fases de and seu F. cicloPrezoto. biológico. 2010. Diversity study,1991; ReeveM. parallelogrammus 1991) which may and be P. reflected versicolor in had the low 3: 117–128. Sociobiology abundances found for these species. However, in this Ayres,of socialM., M. wasps Ayres (Hymenoptera) Junior, D.L. Ayres in aand silvipastoral A.A. Santos. system. 2007. BioEstat – Agelaia vicina, Apoica pallens, Polybia 55(2):Aplicações 627–636. estatísticas nas áreas das ciências bio-médicas jurinei,abundances, Mischocyttarus but were considered cassununga constant and M. species. rotundicolis The, remaining species . Belém, PA: Ong Mamiraua. 364 pp. Bencke,sudeste S.C.C. do and Brasil. L.P.C. Revista Morellato. Brasileira 2002. deEstudo Botânica comparativo da fenologia were considered acccidental species. Thus, of the twenty- de nove espécies arbóreas em três tiposContribuição de floresta ao atlântica estudo dos no one species collected, nine (42.86%) were classified as vespídeos do Brasil (Insecta, Hymenoptera, Vespoidae) 25: 237–248.. Cruz das Carpenter, J.M. and O.M. Marques. 2001. constant species, five (23.81%) were considered accessory Clemente, M.A. 2009. Vespas sociais (Hymenoptera, Vespidae) do Parque species and seven (33.33%) were accidental species. The Almas:Estadual Universidade do Ibitipoca-MG: Federal estrutura, da Bahia. composição 147 pp. e visitação floral. M. high percentage of constant species may also be explained by the high complexity of the ombrophilous dense forest, et al. Sc. Dissertation. Juiz de Fora:Megacanthopus Universidade DuckeFederal of de Mischocyttarus Juiz de Fora. 2007).which has numerous food resources that can provide 79 pp. greater chances of survival and (Santos Cooper,Entomologist’s M. 1997. The Monthly subgenus Magazine de Saussure (Hym., Vespidae), with a key and three new species. in more humid conditions does not mean that a larger 133: 217-223. The greatest number of species of wasps sampled Diniz, I.R. and K. Kitayama. 1994.Journal Colony of Hymenoptera densities Research and preferences for habitats of some social wasps in Mato Grosso State, Brazil (Hymenoptera, Vespidae). 3: 133– number of individuals perform foraging activity under 15 Togni et al. | Social Wasps in Ubatuba, Brazil

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