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Home , Agelaia, Agelaia pallipes, Agelaia vicina, Angiopolybia pallens, Apoica, Apoica pallens

doi:10.12741/ebrasilis.v8i1.447 e-ISSN 1983-0572 Publicação do Projeto Entomologistas do Brasil www.ebras.bio.br Distribuído através da Creative Commons Licence v3.0 (BY-NC-ND) Copyright © EntomoBrasilis Copyright © do(s) Autor(es)

Social (: : ) of the Jaú National Park, Amazonas, Alexandre Somavilla¹, Sérgio Ricardo Andena², Marcio Luiz de Oliveira¹

1. Instituto Nacional de Pesquisas da Amazônia, Coordenação de Biodiversidade, e-mail: [email protected] (Autor para correspondência), [email protected]. 2. Universidade Estadual de Feira de Santana, e-mail: [email protected]. ______

EntomoBrasilis 8 (1): 45-50 (2015)

Registered in ZooBank: urn:lsid:zoobank.org:pub:46C8FEF7-E97B-4AC4-B778-987ED7F4AA61

Abstract. Social wasps are common elements in Neotropics, although even elementary data about this taxon in the Amazon region is partially unknown. Therefore the purpose of this work was to increase the knowledge of social at the Jaú National Park. A total of 494 specimens of Polistinae was collected with active search, malaise trap and light trap. Forty-nine belonging to fourteen genera were recorded. The richest genera were (14 species), (07), (05), (04), (04) and Protopolybia (04); the remaining genera were represented by less than two species. The Jaccard similarity coefficient showed a higher similarity of the Jaú National Park with Ducke Reserve (Manaus, Amazonas). Four species were collected for the first in Amazonas state Agelaia flavipennis (Ducke), Polybia affinis Du Buysson, Protopolybia nitida (Ducke) and Protopolybia sedula (de Saussure), only in the Jaú National Park, showing the importance of this park for the social wasps preservation in the Amazon region.

Keywords: Amazon region; Inventory; Paper wasps; Upland .

Fauna de Vespas Sociais (Hymenoptera: Vespidae: Polistinae) do Parque Nacional do Jaú, Amazonas, Brasil

Resumo. Vespas sociais são elementos comuns na região Neotropical, embora até mesmo os dados elementares sobre este táxon na região amazônica ainda sejam parcialmente desconhecidos. Por isso objetivamos contribuir para o conhecimento sobre a fauna de vespas sociais que ocorrem no Parque Nacional do Jaú. Um total de 494 indivíduos de Polistinae foi coletado através de busca ativa, armadilha Malaise e armadilha luminosa. Quarenta e nove espécies de 14 gêneros foram registradas. Polybia (14 espécies), Agelaia (07), Mischocyttarus (05), Apoica (04), Brachygastra (04) e Protopolybia (04) são os gêneros com o maior número de espécies; os demais gêneros coletados foram representados por apenas uma ou duas espécies. De acordo com o coeficiente de similaridade de Jaccard, a composição de espécies de Parque Nacional do Jaú é semelhante a Reserva Ducke (Manaus, Amazonas). Agelaia flavipennis (Ducke), Polybia affinis Du Buysson, Protopolybia nitida (Ducke) e Protopolybia sedula (de Saussure) são considerados novos registros de ocorrência para o estado do Amazonas e foram coletados, até o momento, apenas no Parque Nacional do Jaú, demonstrando a importância deste parque para a preservação de vespas sociais na região amazônica.

Palavras-chave: Floresta de terra firme; Inventário; Região Amazônica; Vespas-papel. ______

he biodiversity of in Brazil is one of the highest of Caxiuanã Reserve, Pará State (Si l v e i r a 2002); Serra do Divisor the world, representing around 20% of all known insects National Park, Acre State (Mo r a t o et al. 2008); Região dos Lagos, Entomologia Geral (Ma r i n o n i et al. 2006), distributed in six main Brazilian Amapá State and Mamirauá Reserve and Alvarães (Si l v e i r a et al. biomes (Amazon Rain Forest, Atlantic Forest, Caatinga, Cerrado, 2008) and Ducke Reserve, Amazonas State (So m a v i l l a 2012). Pantanal and Pampas). The Amazon Forest is the biggest bioma within Brazilian territory, covering an area of 4,196.943 Km² Despite the contribution of these works, the richness of social (Mi n i s t é r i o d o Me i o Am b i e n t e 2014). wasps in Amazon Forest is still underestimated (So m a v i l l a 2012). Si l v e i r a (2002) pointed out that 200 species were recorded in The social wasps (Vespidae: Polistinae) comprise 26 genera and the Brazilian Amazon, which represents about two-thirds of all 958 species widely distributed in the Neotropical region (Pi c k e t t Brazilian Polistinae’s fauna, although So m a v i l l a & Ol i v e i r a (2013) & Ca r p e n t e r 2010). These wasps play an important role and highlighted the need of more studies focusing on distribution of are important components of complex interactions with other wasps in the region. organisms in Neotropical (Si l v e i r a et al. 2012) as , predators or acting in biological control (Hu n t 2007). In this work we present a list of species of social wasps (Polistinae) The can be the size of a thimble or more than a meter long, as collected at the Jaú National Park comparing it with other areas durable as hard felt or more fragile than egg shells, more regular sampled in Amazon Forest. and uniform than the much-celebrated honeybee comb or wildly chaotic with an intricate mazelike interior (We n z e l 1998). Material and Methods

The knowledge of social wasps remains from some few studies The Jaú National Park (JNP) was established in 1980 and is carried out in forest fragments. Du c k e (1904, 1907) conducted considered the second largest Brazilian national park, occupying one of the first surveys of wasp fauna in the eastern region of the an area of 22,720 km². The park covers the Unini river and Jaú Brazilian Amazon, mainly in Pará State. Recently, similar works river hydrographic basins, right bank tributaries of the Negro have been carried out in the Brazilian Amazon, as follow: Maracá River Island Ecological Satation, Roraima State (Ra w 1998); Funding Agencies: FAPEAM and CNPq

www.periodico.ebras.bio.br 46 Identification of the specimens followed the keys proposed by R keysproposed the followed specimens the of Identification surroundings ofbase,onlyinsamples2003and2004. and (igarapés) creeks of margins as such area, the of trails along were methods three applied in all samples periods. Also, nests were actively collected these employed; were sampling, one of from a.m. night 5 to p.m. 6 between light), black and white to one attached sheet white (a net. traps light and entomological sampled, days seven with collected actively the during installed model), (Townes traps malaise Additionally, were wasps The 2001; August and June April, November 2003andMay2004. 1993; June and April during - 1 Area – JNP at (02º00’03”S; points 62º43’54”W) and Area sampling 2 (01º53’04”S; different 61º35’04”W) two in performed levels onOctoberandNovember(Borges river lower the and September and July between occurs season of the levels flood largest rivers occur during June and the July. In opposite, the lowest rainfall however May, thepeak to January from reaches The precipitation seasons. waters flood/low Water annual cycle variation ranges from 6 to 10 meters, between Figure 1.Mapof of JaúNationalPark(PNJ)andthe twoplacesofsamples:Area1and Area2.Imageadaptedfrom river ( h seies ee olce i fs ivnois svn days (seven proposed by inventories as - each) fast in collected were specimens The 2004). (H sand forest (campina) and high white-sand forest (campinarana) white- low fechado), (igapó forest flooded closed aberto), (igapó forest flooded open densa), submontana densedownland firme terra de forest (floresta upland densa), open firme terra de region: (floresta the cover physiognomy upland forest (floresta de terra firme aberta), dense upland forest different Seven c i Social Wasps(Hymenoptera:Vespidae:Polistinae)ofJaú… s e u q i r n e h s d r a B s e g r o (1978), & etal. R l e a f a C 2004)(Figure1). r a 1999; p r e t n e S a v l i H M &

(2011). The inventories were inventories The (2011). al. et s e u q i r n e s e u q r a 2004;

et al. 20) and (2001) P 2004). n i h o r i e & C r a B p r e t n e s e g r o

(P Rare 1% < = D e Recessive 2% – 1 = D Subdominant, 5% – 2 = D individuals collected, where D = > 10% total Eudominant, D = 5 – the 10% Dominant, = t and species particular of a number of total individuals = i (%), formula Dominance = the D using where 100, calculated x D=(i/t) was taxa the of dominance The da Amazônia(INPA)inManaus,Brazil. Collection the at Zoological of Invertebrates at deposited the Instituto Nacional de Pesquisas were specimens Voucher (2004). Polybia genera of Polistinae, were collected in the Jaú National Park area. A total of 494 individuals, forty-nine species, belonging to fourteen (07), we excluded the specimens identified as “morphospecies” or “morphospecies” as “varieties”. identified specimens the excluded we analysis, this In Amazon. Brazilian the in out carried inventories al. et (R Development Core Team 2011), version 2.13, Vegan (O Vegan 2.13, version 2011), Team Core Development (R The Jaccard similarity coefficient was undertaken in the R program located also in Amazonas State. Maracá River Island in Roraima in Island River Maracá State. Amazonas in also located (32%) Mamirauá and (33%) Alvarães by followed State, Pará in Ducke with similar more that the (35%) is Caxiuanã by followed SE, Km 200 situated (39%) Reserve JNP of 2) shows and composition (Figure species coefficient similarity Jaccard The species representedmorethan50%ofthetotal(27species). (25), (Fabricius) (32), (Lepeletier) bistriata Polybia were: species and a s s i l a (04) (Table 1). The Eudominant or Dominant or Eudominant The 1). (Table (04) Protopolybia 2008), comparing the species richness of JNP with others with JNP of richness species the comparing 2008), (05), Mischocyttarus etal. was the richest genera (14 species), followed by followed species), (14 genera richest the was 1979). Result andDicusson Dce (9 specimens), (79 (Ducke) fulvofasciata Agelaia Oie) (27), (Oliver) pallida Apoica Fbiis (78), (Fabricius) Lnau) 2) Rare (24). (Linnaeus) canadensis (04), Apoica H s e u q i r n e e-ISSN 1983-0572 niplba pallens Angiopolybia (2004). Brachygastra Somavilla etal. oyi liliacea Polybia Agelaia n e n a s k (04)

Janeiro - Abril 2015 - www.periodico.ebras.bio.br EntomoBrasilis 8 (1)

Table 1. Species of social wasps collected at Jaú National Park, Amazonas, Brazil, as well as the method used to capture the species. Status: Eudominant (Ed) Dominant (D), Subdominant (Sd), Recessive (Rc) and Rare (Rr). Number of Dominance Species Method specimens Status Agelaia angulata (Fabricius) Active search, Malaise trap 06 Rc Agelaia centralis (Cameron) Malaise trap 06 Rc Agelaia flavipennis (Ducke) Malaise trap 02 Rr Agelaia fulvofasciata (DeGeer) Active search, Malaise trap 79 Ed (Olivier) Malaise trap 08 Rc Agelaia ornata (Ducke) Active search 01 Rr Agelaia testacea (Fabricius) Active search, Malaise trap 02 Rr (Lepeletier) Active search, Malaise trap 32 D Angiopolybia paraensis (Spinola) Active search, Malaise trap 11 Sd Apoica arborea de Saussure Light trap 22 Sd (Fabricius) Light trap 15 Sd Apoica pallida (Olivier) Light trap 27 D Apoica thoracica Du Buysson Light trap 06 Rc Brachygastra augusti (de Saussure) Malaise trap 03 Rr Brachygastra bilineolata Spinola Malaise trap 14 Sd (Latreille) Malaise trap 05 Rr Brachygastra scutellaris (Fabricius) Malaise trap 04 Rr Chartergus chartarius (Olivier) Malaise trap 02 Rr Clypearia sulcata (de Saussure) Malaise trap 01 Rr Leipomeles dorsata (Fabricius) Malaise trap 01 Rr Metapolybia nigra Richards Active search, Colony 22 Sd fraternus (Gribodo) Malaise trap 01 Rr Polybia affinis Du Buysson Malaise trap 01 Rr Polybia belemensis Richards Malaise trap 01 Rr Polybia bicyttarella Richards Malaise trap 01 Rr Polybia bistriata (Fabricius) Active search, Colony 78 Sd Polybia depressa (Ducke) Malaise trap 01 Rr Polybia dimorpha Richards Active search, Malaise trap 21 Sd Polybia jurinei de Saussure Active search, Malaise trap 13 Sd Polybia liliacea (Fabricius) Active search, Malaise trap 25 D (Olivier) Active search, Malaise trap 09 Rc Polybia quadricincta de Saussure Malaise trap 01 Rr (Fabricius) Active search, Malaise trap 06 Rc Polybia scrobalis Richards Malaise trap 06 Rc Polybia singularis Ducke Malaise trap 01 Rr Polybia striata (Fabricius) Active search, Malaise trap 02 Rr Protopolybia bituberculata Silveira & Carpenter Malaise trap 02 Rr Protopolybia chartergoides (Gribodo) Malaise trap 07 Rc Protopolybia nitida (Ducke) Malaise trap 02 Rr Protopolybia sedula (de Saussure) Malaise trap 01 Rr surinama (Linnaeus) Active search 01 Rr Synoeca virginea (Fabricius) Active search 02 Rr Mischocyttarini de Saussure Active search 04 Rr Mischocyttarus labiatus (Fabricius) Malaise trap 03 Rr Mischocyttarus metathoracicus (de Saussure) Active search, Colony 02 Rr Mischocyttarus socialis (de Saussure) Malaise trap 01 Rr Mischocyttarus surinamensis (de Saussure) Malaise trap 01 Rr Polistini 47 (Linnaeus) Active search, Colony 24 Sd (Fabricius) Active search, Colony 08 Rc

e-ISSN 1983-0572 48 Ecological Station(RR)andSerra doDivisorNationalPark(AC). Amazon Region: Ducke Reserve (AM), Alvarães (AM), Mamirauá Reserve (AM), Caxiuanã Reserve (PA), Região dos Lagos (AP), Maracá River Island Figure 2. Dendrogram generated by Jaccard similarity coefficient for the species composition of Jaú National Park (AM) with other inventories in the hs ok ny h genera the only work this mainly three species, (Agelaia abundant genera more sample to efficient are dos Lagos, Amapá, with 31 species (S and work this in recorded as also, and csses n wt as a ifrn cletn efforts. collecting different a different also in with employed been and have ecosystems etc.) troublesome. solution, something attractive search, active are traps, (malaise methods wasps different Unfortunately collecting for Methods Amazonas State. Maracá River Island, Roraima, in with found were 36 forest, species Amazon (R the in found the was richness contrast, lowest In (87). Reserve Ducke by Caxiuanã followed in (97), reserve registered was basin Amazon in richness highest species), both in Amazonas (S Amazonas in both species), Mamirauá Reserve (46 species) and the outskirts of Alvarães (42 with grouped was species) (49 JNP richness, of relation in Even similarity, The 20%). below or grouped areaswithcoefficientofsimilarityplantphysiognomy. to near (values composition species the between similarity lowest the with areas those were Acre in Divisor do Serra and Amapá in Lagos dos Região State, collected in the Amazon region, as In your research, do Divisor,Acre,with20species( bnat eea rbby eas te pce o ti genera this of usually species the because probably genera abundant works, both In abundant. considered were 19) ad osqety ae hge poaiiy f being (S captured of probability higher a have consequently and 1995), al. Social Wasps(Hymenoptera:Vespidae:Polistinae)ofJaú… Polybia

form big colonies with millions of individuals (Z individuals of millions with colonies big form a r i e v l i

Tee or pce ae e rcrs for records new are species four These sedula. , S Polybia

a r i e v l i . 2012). However some species are rarely are species some However 2012). al. et

et al. (2012) showed that fast inventories and Agelaia, a r i e v l i . oee, in However, Mischocyttarus). M Polybia affinis, Polybia A. flavipennis ( a r i e v l i o t a r o

s

Polybia et al. et etal.

2008). However, the However, 2008). Agelaia et al. and 2008). 2008) and Serra a w S Polybia nitida Polybia 1998), Região Angiopolybia a r i e v l i was the most the was c c u 2002), h i

et

ih h hget pce rcns o sca wss n rzl We Brazil. in wasps social of richness species highest the with biome the is Amazon the therefore biomes, these than greater is richness species the JNP and Reserve like Ducke Reserve, , Caxiuanã upland Amazonian of areas in However, changes. composition thespecies but areas, Amazon modified are these of thespeciesarehardlyrecoverable( of species is not available, and data about the relative abundance number certain a finding in spent efforts the about information be employed for maximize the effectiveness of the survey (S must methods different that suggest we problems, such Despite las ofot btce aiig rm non-standardized from arising (S methods obstacles collecting confront always collections in information of using local of Comparisons & forage at night (H the Polistini and Mischocyttarini as well as species of way for collecting social wasps may underestimate the richness of only the as traps Malaise of employment the Therefore, (2009). Protopolybia Epiponini of species to the belonging of collected, majority the work this In 2002). forest or crops areas and anthropogenic environments (H environments anthropogenic and areas crops or forest Atlantic formations), outcrop (rock rupestres (Brazilian campos Savanna), cerrado as such biomes Brazilian other to similar is The species number of Maracá River Island and Serra do Divisor thoracica arborea, capturing these wasps during the day. In the present study has also been reported by reported been also has that pattern a method, this using efficient less were Polistini and Mischocyttarini of species collected contrast, In trap. Malaise in K ö h r e l

Apoica pallens Apoica DuBuyssonwerecollectedonlywithlighttraps. 2006; n sm seis of species some and S t n u a v l i

et al. et al. a r i e v l i Brachygastra, (Fabricius), 1995), which reduces the possibility of 2011; S a r i e v l i

02. n eea, appropriate general, In 2002). T a k a n a

(2002), and (2002), Polybia and pallida Apoica S a r i e v l i -J Leipomeles, Clypearia, e-ISSN 1983-0572 r . & ee atrd only captured were 2002). N Somavilla etal. l l o S a v l i

2011;), since Apoica, that & S Apoica Apoica a r i e v l i a r i e v l i r e m s e

Janeiro - Abril 2015 - www.periodico.ebras.bio.br EntomoBrasilis 8 (1) hope that this work stimulate other surveys in Amazon Region. Ecology Package. R package version 1. Available in: . Acknowledgments Palissa, A.E., M. Wiedenroth & K. Klimt, 1979. Anleitung zum ökologischen Geländepraktikum. Wissenschaftliches We thank to Dr. Augusto Loureiro and Dr. José Albertino Rafael Zentrum der Pädagogischen Hochschule Potsdam, 186p. for the wasps collected and information about the Jaú National Pickett, K.M. & J.M. Carpenter, 2010. Simultaneous analysis Park, to Mateus Clemente and Karine Schoeninger for helping in and the origin of in the Vespidae (Insecta: the discussions of the work and to the two anonymous referees Hymenoptera). Systematics and Phylogeny, 68: for their helpful suggestions. We also thank Instituto Nacional de 3-33. Pesquisas da Amazônia, FAPEAM and CNPq for support. Pinheiro, M.R. & S.H. Borges, 2004. Caracterização das janelas para a biodiversidade do Parque Nacional do Jaú, p. 14-40. References In: Borges, S.H., S. Iwanaga, C.C. Durigan & M.R. Pinheiro Borges, S.H., C.C. Durigan, M.R. Pinheiro, J.L.C. Camargo & A. (Eds.). Janelas para a biodiversidade no Parque Nacional Murchie, 2004. Planejando o estudo da biodiversidade na do Jaú. Fundação Vitória Amazônica, Manaus, Amazonas, Amazônia brasileira: uma experiência no Parque Nacional 273p. do Jaú, p. 3-18. In: Borges, S.H., S. Iwanaga, C.C. Durigan Raw, A., 1998. Social Wasps (Hymenoptera, Vespidae) of the Ilha & M.R. Pinheiro (Eds.). Janelas para a biodiversidade no de Maracá, p. 307-321. In: Ratter, J.A. & W. Milliken (Eds.). Parque Nacional do Jaú. Fundação Vitória Amazônica, Maracá. Biodiversity and environment of an Amazonian Manaus, Amazonas, 273p. . Chichester, John & Sons, London, 508p. Carpenter J.M. & O.M. Marques, 2001. Contribuição ao Estudo Richards, O.W., 1978. 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Noll, M.R. Mechi, S. Mateus, consolidação de sistemas integrados de informação sobre M.V. Baio & S. Shima, 1995. , a swarm- biodiversidade. Centro de Gestão e Estudos Estratégicos/ founding polistine with the largest colony size among wasps Ministério da Ciência e Tecnologia. Brasília, Brasil, 34p. and (Hymenoptera: Vespidae). Journal of New York Ministério do Meio Ambiente, 2014. Biomas, Amazônia. Available Entomological Society, 103: 129-137. in: . [Access in: 01.10.2014]. Received in: 06/10/2014 Morato, E.F., S.T. Amarante & O.T. Silveira, 2008. Avaliação Accepted in: 01/29/2015 ecológica rápida da fauna de vespas (Hymenoptera, Aculeata)

********** 49 do Parque Nacional da Serra do Divisor, Acre, Brasil. Acta Amazonica, 38: 789-798. Oksanen, J., R. Kindt, P. Legendre, B. O’hara, G.L. Simpson, M. Henry, H.H. Stevens & H. Wagner, 2008. Vegan: Community e-ISSN 1983-0572 50 oail, . SR Adn, .. lvia 21. oil ap (yeotr: epde Plsia) f a Ntoa Pr, mzns Brazil. Amazonas, Available in: Park, National Jaú of Polistinae) Vespidae: (Hymenoptera: Wasps EntomoBrasilis, 8(1):45-50. Social 2015. Oliveira, M.L. Andena, S.R. A., Somavilla, Suggested citation: Social Wasps(Hymenoptera:Vespidae:Polistinae)ofJaú… doi:10.12741/ebrasilis.v8i1.44 7 e-ISSN 1983-0572 Somavilla etal.