Diversity of Tiphiidae (Insecta: Hymenoptera) in the Fragmented Brazilian Semi-Deciduous Atlantic Forest
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J Insect Conserv (2016) 20:417–431 DOI 10.1007/s10841-016-9875-9 ORIGINAL PAPER Diversity of Tiphiidae (Insecta: Hymenoptera) in the fragmented Brazilian semi-deciduous Atlantic Forest 1 1 Cı´ntia Eleonora Lopes Justino • Eduardo Fernando dos Santos • Fernando Barbosa Noll1 Received: 31 August 2015 / Accepted: 3 May 2016 / Published online: 7 May 2016 Ó Springer International Publishing Switzerland 2016 Abstract The Atlantic Forest is one of the most important variables also explained the variation in Shannon diversity areas of biodiversity in the world, but it has been largely and species evenness. Variations in species richness and replaced with agropastoral areas and at the present only diversity of Tiphiidae are strongly associated with neutral 12.5 % of the original cover remains. Despite the ecolog- processes, but they are also influenced by forest fragmen- ical importance of insects, few studies have been used in tation and intensive agricultural activities. conservation approaches for the Atlantic Forest, mainly due to a great taxonomic impediment. A group quite eco- Keywords Parasitoids wasps Á Tropical forest Á logically important but deeply neglected includes para- Agricultural landscape Á Deforestation Á Neutral model sitoid wasps that control a great number of invertebrates, like tiphiid wasps that are parasitoids of underground coleopteran larvae. The present study aimed to estimate Introduction Tiphiidae species richness and diversity in 15 patches of a highly fragmented Atlantic Forest region, using factors that The Atlantic Forest is the second largest forest biome in drive the diversity pool from a metacommunity, such as South America (Ribeiro et al. 2009, 2011), and it is con- immigration and speciation probabilities. The parameters sidered a hotspot of biodiversity since Meyer (1988), who were estimated using the Neutral Biodiversity Theory, first defined and described ten tropical forests as hotspots. which is based on the total ecological equivalence of spe- Nowadays, 35 biodiversity conservation hotspots are rec- cies at the same trophic level. Diversity values were mol- ognized and the Atlantic forest is one of the most species ded to the area size, the immigration probabilities, and/or rich areas on the planet (Ribeiro et al. 2011). The original the speciation probability. Eight genera and 460 individuals cover of this biome was approximately 150 million hec- of Thynninae, Myzininae and Tiphiinae were collected. tares under highly heterogeneous environmental condi- Variation in species richness, estimated by both rarefaction tions, encompassing different habitats including deciduous and first-order jackknife methods, was explained by patch and rain forests, mangroves, swamps, sand vegetation size and by immigration and speciation probabilities. These (restingas), inselbergs, high-altitude grasslands (campo rupestre and campo de altitude), and mixed Araucaria pine Electronic supplementary material The online version of this forests (Scarano 2002;Caˆmara 2003 in Ribeiro et al. 2011). article (doi:10.1007/s10841-016-9875-9) contains supplementary However, currently, only 12.5 % of the original cover material, which is available to authorized users. remains (Fundac¸a˜o SOS Mata Atlaˆntica and INPE 2002, 2014; Ribeiro et al. 2009). & Cı´ntia Eleonora Lopes Justino [email protected] Massive areas of Atlantic Forest are being reduced to an archipelago of small, widely separated forest fragments 1 Departamento de Zoologia e Botaˆnica, Instituto de (Ranta et al. 1998), with approximately 97 % of Atlantic Biocieˆncias, Letras e Cieˆncias Exatas, Universidade Estadual Forest fragments having areas smaller than 250 ha (Ribeiro Paulista ‘‘Ju´lio de Mesquita Filho’’, Rua Cristo´va˜o Colombo, 2265, Jd. Nazareth, Sa˜o Jose´ do Rio Preto, SP 15054-000, et al. 2009). Despite a dramatic reduction in Atlantic Forest Brazil deforestation since 2008, more than 60,000 ha were lost 123 418 J Insect Conserv (2016) 20:417–431 between 2010 and 2013 (Fundac¸a˜o SOS Mata Atlaˆntica A great number of species remain unknown to science, and INPE 2014). The main cause of Atlantic Forest especially those from hotspot biomes (Joppa et al. 2011) deforestation is agricultural expansion (Dean 1997), as in such as the Atlantic Forest. The insect fauna is a good other tropical forests worldwide (Kaimowitz and Angelsen example of that, despite of recent efforts to characterize it, 1998; Sodhi et al. 2004; Pirard and Treyer 2010; Kissinger our knowledge on its diversity remains incipient (Lewin- et al. 2012; Adu et al. 2012; FAO 2014). The resultant sohn and Prado 2002; Lewinsohn et al. 2005; Rafael et al. agro-mosaics now represent the predominant landscape 2009; Diniz-Filho et al. 2010) and many groups, such as type in formerly forested lands across the world (Sodhi the Tiphiidae wasps (Genise and Kimsey 1991; Kimsey et al. 2004; Tabarelli et al. 2010). 1991, 1992), have been neglected (Marinoni and Peixoto Forest fragmentation may be understood based on the 2010). Tiphiidae is a cosmopolitan and diverse family (Pate basic conceptual model provided by island biogeography 1947) of solitary and fossorial ectoparasitoid wasps of soil- theory although this model has limitations and it cannot dwelling beetle larvae (Brothers and Finnamore 1995; Pate explain the dynamic that is greatly amplified in fragmented 1947; Potter and Rogers 2008). The family includes more habitats (Haila 2002; Laurance 2008; Mendenhall et al. than 1500 described species representing 120 genera 2014). The island biogeography theory explains variation (Brown 2005). In Brazil, 102 species representing 19 in species richness as a balance between immigration and genera have been described (Justino and Santos 2016). extinction rates (MacArthur and Wilson 1963, 1967; Tiphiidae wasps are mentioned as flower visitors and may Schoener 2010), which are also key components of the be involved in pollination (Kevan 1973); furthermore, unified neutral theory of biodiversity and biogeography some species of Thynninae in Australia have a strong proposed by Hubbell (1997, 2001). The neutral theory relationship with orchids as showed by Alcock (2000), treats the organisms of the same trophic level from a Brown (1998), Handel and Peakall (1993), Mant et al. metacommunity1 as essentially identical in their individual (2002, 2005), Peakall and Beattie (1996), Peakall et al. probability of birth and death, immigration and speciation (2010) and Stoutamire (1983). Santos and Branda˜o(2011) (Hubbell 2001; Mangel 2002; Bell 2001; Chave 2004). In and Santos et al. (2014) carried out the first characteriza- addition, the neutral theory assumes that species are under tions of Tiphiidae diversity in the Atlantic Rain Forest, but no differential selection pressures (Hubbell 2001; Hubbell the tiphiid fauna of Atlantic Semi-deciduous Forest is as and Lake 2003), which means that species do not have yet largely unknown. Recent collections in patches of different competitive ability and the diversity of a meta- Semi-deciduous Forest have resulted in new records (Jus- community is given stochastically by immigration, speci- tino et al. 2013). ation, birth and death (Hubbell 2001; Mangel 2002; Chave Regarding all of the aspects presented above, the present 2004; Economo and Keitt 2008; Rosindell et al. 2011, study aimed to answer the following questions: (1) How 2012; Matthews and Whittaker 2014). Furthermore, it can diverse is the Tiphiidae fauna in the Atlantic Semi-decid- robustly encompass the predictions of island biogeography uous Forest? (2) Are the species richness and diversity of theory, like species-area relationships (Rosindell et al. Tiphiidae affected by fragmentation of the Atlantic Forest 2011). fragmentation ? (3) Is the metacommunity structure of The ratio between local births and immigration has an Tiphiidae driven by stochastic processes that underpin the effect (decreasing or increasing) on the number of species unified neutral theory of biodiversity and biogeography, coexisting in disturbed communities, which is important such as immigration and speciation? for ecosystem management and conservation (Dornelas 2010), whereas disturbance can increase the extinction rate and favor the dominance by a single species (Gardner and Materials and methods Engelhardt 2008). In this way, the neutral theory can be extended to explain patterns of species responses to dis- Study area and collection methodology turbance (Kadmon and Benjamini 2006). The application of this theory in conservation approaches has the advan- The present study was conducted in 15 forest fragments of tages of not requiring species-specific details, which are different sizes (Table 1) from northwestern Sa˜o Paulo State often lacking, and the inclusion of more intrinsic biological (Fig. 1). The landscape in this region consists of an agri- information linked to relative abundances rather than the cultural matrix, which includes cane, citrus and coffee extrapolation of species-area relationships (Rosindell et al. crops, and pasture (Table 1). Forest patches in northwest- 2012). ern Sa˜o Paulo have been characterized as Semi-deciduous Seasonal Forest with different stages of ecological suc- 1 Metacommunity consists of all trophically similar individuals and cession (Necchi Jr. et al. 2012). The climate in the region is species in a regional collection of local communities. defined as Temperate, with two distinct seasons (i.e., dry 123 J Insect Conserv (2016) 20:417–431 419 Table 1 Characteristics of the Area Latitude Longitude Size