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Fluctuations in Richness and Abundance of Social Wasps During the Dry and Wet Seasons in Three Phyto-Physiognomies at the Tropical Dry Forest of Brazil

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Fluctuations in Richness and Abundance of Social Wasps During the Dry and Wet Seasons in Three Phyto-Physiognomies at the Tropical Dry Forest of Brazil COMMUNITY AND ECOSYSTEM ECOLOGY Fluctuations in Richness and Abundance of Social Wasps During the Dry and Wet Seasons in Three Phyto-Physiognomies at the Tropical Dry Forest of Brazil 1,2 3 1 G. M. DE M. SANTOS, P. C. BISPO, AND C.M.L. AGUIAR Environ. Entomol. 38(6): 1613Ð1617 (2009) ABSTRACT The social wasp nests were quantiÞed in three different plant physiognomies (forested Caatinga, shrubby Caatinga, and agricultural systems) to analyze the effect of environmental sea- sonality and plant physiognomy on the richness, nest abundance, and species composition of social wasps in the region of tropical dry forest of Brazil. The forested Caatinga physiognomy had the greatest richness of species (S ϭ 16), followed by shrubby Caatinga (S ϭ 13) and by agricultural system (S ϭ 12). The Þrst axis of detrended correspondence analysis (DCA) explained 67.8% of the variability and shows a gradient of the fauna from agricultural system and shrubby Caatinga to forested Caatinga. In the Þrst axis, wet season scores were much higher than those for the dry season in forested Caatinga. The second axis explained 18.7% of the variability and shows a separation of samples collected during the wet or the dry periods in shrubby Caatinga. This separation was less evident in the agricultural system. Variations in nest abundance were more intense in arbustive caatinga (45% decrease in number of active nests in the dry period), moderate in forested Caatinga (24% decrease in number of active nests in the dry period), and low in agricultural systems (8% decrease in the dry period). KEY WORDS Caatinga, Polistinae, community ecology, species richness Studies on diversity of social wasps (Hymenoptera: colonies can be considered semisessile organisms Vespidae) in Brazilian ecosystems are still scarce and (Santos and Gobbi 1998). Studies on nest abundance several refer to the classic studies by Ihering (1904), have shown that seasonality occurs in some neotropical Ducke (1906, 1910), Richards and Richards (1951), social wasps species but not in others (Marques et al. and Richards (1978). Recent research have produced 1992, Ramos and Diniz 1993, Diniz and Kitayama 1998, information on the biodiversity of social wasps in the Santos et al. 2006). Furthermore, the same species may cerrado (Henriques et al. 1992, Diniz and Kitayama present seasonal ßuctuation or not, depending on envi- 1998, Mechi 2005, Elpino-Campos et al. 2007, Santos et ronmental conditions (Ramos and Diniz 1993). al. 2009), rupestrian Þelds (da Silva-Pereira and Santos The semiarid region in the Brazilian Northeast has 2006), caatinga (Santos et al. 2006, Aguiar and Santos two well-deÞned seasons: the dry and the wet seasons. 2007), the Atlantic Forest, mangroves and sandbanks For our purposes, the deÞnition of “dry period” in- (Santos et al. 2007), Araucaria Forest (Hermes and volves scarce food resources for the entomofauna. Ko¨hler 2006), and in agriculture lands (Marques and Conversely, the “wet period,” which extends for 2 mo Carvalho 1993, Hermes and Ko¨hler 2006). after the rain stops, presents intense plant blooming and The diversity and seasonality of species in social abundance of food resources for the entomofauna. wasp communities are usually studied from two per- This study aimed to investigate the relationships spectives: abundance of ßower-visiting wasps (Santos between richness and abundance of wasp species and et al. 1998, 2006; Silva-Pereira and Santos 2006; Hermes plant physiognomies with different levels of complex- and Ko¨hler 2006) or nest abundance (Henriques et al. ity (agricultural systems, shrubby Caatinga, and for- 1992, Marques and Carvalho 1993, Diniz and Kitayama ested Caatinga) in a tropical dry forest region of Brazil 1994, de Santos et al. 2007). This paper is based on nest (Caatinga). It also aimed to understand whether the abundance. ßuctuations in richness and abundance of wasp nests The temporal dynamics of tropical social wasps is in these communities are related to the wet and dry very complex. These insects contribute signiÞcantly seasons. for their colonies and do not abandon them easily; the 1 Laborato´rio de Entomologia, DCBIO, Universidade Estadual de Materials and Methods Feira de Santana, 44031-460 Feira de Santana, Bahia, Brazil. 2 The study was conducted in the municipalities of Corresponding author, e-mail: [email protected]. Њ Ј Њ Ј Њ Ј 3 Faculdade de Cieˆncias e Letras de Assis, Universidade Estadual Itatim (12 42 S and 39 41 W), Itaberaba (13 32 S and Paulista Ju´ lio de Mesquita Filho, Sa˜o Paulo, Brazil. 40Њ18Ј W), and Milagres (12Њ52Ј S and 39Њ51Ј W), State 0046-225X/09/1613Ð1617$04.00/0 ᭧ 2009 Entomological Society of America 1614 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 6 Table 1. Number of social wasp nests registered in three Caatingas physiognomies during the wet and the dry seasons in Brazil Agricultural Shrubby Forested Species system Caatinga Caatinga Dry Wet Dry Wet Dry Wet 01 Apoica pallens (Fabricius, 1804) 3 4 1 2 1 1 02 Brachygastra lecheguana (Latreille, 1824) 16 16 8 8 1 1 03 Clypearia angustior Ducke, 1906 ÑÑÑÑÑ 1 04 Mischocyttarus cassununga (Ihering, 1903) Ñ Ñ Ñ 6 1 4 05 Mischocyttarus cerberus Ducke, 1918 ÑÑÑÑÑ 1 06 Mischocyttarus drewseni Saussure, 1857 ÑÑÑÑÑ 3 07 Parachartergus pseudapicalis Willink, 1959 1 2 Ñ 3 3 6 08 Polistes billardieri (Fabricius, 1804) 2 3 Ñ 1 Ñ Ñ 09 Polistes c. canadensis (L., 1758) 35 33 11 22 8 8 10 Polistes v. versicolor (Olivier, 1791) 7 9 1 10 2 2 11 Polybia ignobilis (Haliday, 1836) 22 21 5 9 1 1 12 Polybia o. occidentalis Richards, 1951 18 27 1 5 2 1 13 Polybia paulista von Ihering, 1896 3 6 Ñ 2 1 Ñ 14 Polybia sericea (Olivier, 1791) 19 14 24 27 4 4 15 Protonectarina sylveirae (Saussure, 1854) 2 2 4 4 1 1 16 Protopolybia e. exigua (Saussure, 1854) 1 3 Ñ 1 1 1 17 Synoeca cyanea (Fabricius, 1775) ÑÑÑÑ 2 2 Total (N) 129 140 55 100 28 37 of Bahia, northeastern Brazil. This is a semiarid trop- used to evaluate the relationship between complexity ical region with high risk of drought. The landscape is of the habitat (Euclidian distance among the samples a mosaic of agricultural environments and several based on a scale of habitat complexity) and the fauna physiognomies of the natural caatinga vegetation. (Morisita-Horn similarity index). The scale of habitat Samples were collected at all three locations from complexity varied from 1 to 3. The value 1 was attrib- each of the following physiognomies: FC, forested uted to agricultural systems, 2 to shrubby Caatinga, caatinga, seasonally dry tropical semideciduous forest, and 3 to forested Caatinga. The Mantel test was done open sclerophyllous forest, 6Ð16 m high; SC, shrubby with 5,000 permutations in NTSYS 2.1 (Rohlf 2000). Caatinga, open physiognomy with predominance of medium-sized vegetation (Ͻ2 m high), important Results presence of cactuses and palm trees; AS, agricultural systems, where the natural vegetation was partially Three hundred nineteen nests of 17 species of social replaced by pasture or small plots of crops (cowpea, wasps were observed. Some of these nests were active Vigna unguiculata; beans, Phaseolus vulgaris; corn, Zea in both dry and wet seasons; thus, we had 489 nests mays). registered (Table 1). The forested Caatinga physiog- The number of nests with active colonies in each nomy had the greatest richness of species (S ϭ 16), physiognomy was used as measure of abundance. Two followed by shrubby Caatinga (S ϭ 13) and the agri- data collectors searched for nests of social wasps with cultural system (S ϭ 12) (Fig. 1). The most abundant active colonies in each physiognomy, during the wet species were Polistes canadensis canadensis (L. 1758), period (January, February, and July 1998) and the dry Polybia sericea (Olivier, 1791), Polybia ignobilis (Ha- period (September, October, and November 1998). liday 1836), Polybia occidentalis occidentalis (Rich- Each collector searched for 80 h in each physiognomy, ards, 1951), Brachygastra lecheguana (Latreille, 1824), a total of 160 h of sampling for each physiognomy. Five and Polistes versicolor versicolor (Olivier, 1791); they to 10 specimens from each colony were registered and identiÞed. Individuals of all species were deposited in the Entomological Collection Johann Becker of the Museu de Zoologia of the Universidade Estadual de Feira de Santana (MZUEFS). Because of the important variation in number of nests among plant physiognomies, we used rarefaction curves to compare the richness of species for the same number of nests. K individuals were randomly re- moved from the sample to estimate the taxa number. The procedure was repeated 1,000 times with the program Ecosim 5 (Gotelli and Entsminger 2000). The abundance matrix was logarithmized before the analysis. This matrix was submitted to detrended correspondence analysis (DCA) (Gauch 1995, Leg- Fig. 1. Rarefaction curves of social wasps registered in endre and Legendre 1998) using the program PCORD three plant physiognomies of Brazilian Steppe (Caatinga). (McCune and Mefford 1999). The Mantel test was ConÞdence intervals (95%). December 2009 SANTOS ET AL.: SOCIAL WASP COMMUNITIES IN THE TROPICAL DRY FOREST 1615 Fig. 3. Fauna of social wasps registered in three plant Fig. 2. Richness (standardized for 40 nests) of social physiognomies. Standardized richness in the wet (W) and wasps registered in three plant physiognomies (AS, agricul- dry (D) seasons for each plant physiognomy (AS, agricul- tural system; SC, shrubby Caatinga; FC, forested Caatinga). tural system; SC, shrubby Caatinga; FC, forested Caatinga). Bahia, Brazil. ConÞdence intervals (95%). ConÞdence intervals (95%). corresponded to 82.4% of all colonies of social wasps (Table 1). tal heterogeneity allows for a larger number of niches, The number of active nests observed varied among setting the conditions for a larger number of species to physiognomies: 171 in the agricultural system, 105 in coexist (Levins 1962, Stehli et al.
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