In a Floodplain Forest in the Brazilian Pantanal
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Vertical and time distribution of Pseudoscorpiones (Arthropoda: Arachnida) in a floodplain forest in the Brazilian Pantanal Leandro Dênis Battirola¹, Germano Henrique Rosado-Neto², Daniel Augusto Batistella¹, Volker Mahnert³, Antonio Domingos Brescovit4 & Marinêz Isaac Marques5 1. Instituto de Ciências Naturais, Humanas e Sociais (Universidade Federal de Mato Grosso), Av. Alexandre Ferronato 1.200, Setor Industrial, 78557-267, Sinop-MT, Brasil; [email protected], [email protected] 2. Departamento de Zoologia (Universidade Federal do Paraná). Centro Politécnico, Caixa Postal 19020, 81531-980 Curitiba-PR, Brasil; [email protected] 3. Muséum d’Histoire Naturelle, Genève, Switzerland; [email protected] 4. Laboratório Especial de Coleções Zoológicas, Instituto Butantan, Av. Vital Brasil 1.500, Butantã, 05503-900, São Paulo-SP, Brasil; Pesquisador bolsista produtividade CNPq; [email protected] 5. Instituto de Biociências, Universidade Federal de Mato Grosso, Av. Fernando Corrêa da Costa 2.367, Boa Esperança, 78060-900 Cuiabá-MT, Brasil; [email protected] Received 03-V-2016. Corrected 12-I-2017. Accepted 31-I-2017. Abstract: Pseudoscorpions embrace a diverse group of arachnids with approximately 3 500 species that occur in various habitats, such as soil, leaf litter, caves and canopies. This study aimed at evaluating the relationship between the pseudoscorpion assemblages in soil, termite nests, tree trunks and canopies, as well as the temporal distribution as to the abundance and richness of species in these habitats. For this purpose different sampling techniques were applied in an integrated manner in distinct habitats of a seasonally flooded monodominant forest in the Northern Pantanal of Mato Grosso, Brazil. Data regarding the abundance and richness of assem- blage species were organized by means of the Non-Metric Multidimensional Scaling (NMDS). A total of 2 068 Pseudoscorpiones distributed in seven families and 18 species were captured. Chernetidae (8 spp.) and Withiidae (5 spp.) predominated, while Atemnidae, Cheiridiidae, Geogarypidae, Lechytiidae and Olpiidae were present with only one species each. Terrestrial fauna was more abundant (1 035 ind.; 50.0 % of the total catch) with three families and 10 species. In the arboreal fauna (712 ind.; 34.4 %), four families and 13 species were found. Termite mounds (321 ind.; 15.6 %) corresponded to the habitat with the least number of individuals, however, five families and nine species were identified, including Parachernes sp. 2, Geogarypus sp. and Olpiolum sp., not found in other analyzed habitats in this forest. The analyses did not show any temporal variation as to abundance or richness of Pseudoscorpiones in each evaluated habitat; however, the analysis showed that the edaphic environment (soil + termite nests), tree trunks and tree canopies are distinct habitats in relation to the composition of the assemblage. The occurrence of typical terricolous (e.g. Parawithius sp., Withiidae gen. sp. and Parachernes sp.) and other arboricolous (e.g. Cheiridium sp., Americhernes sp. and Lustrochernes sp.) spe- cies highlights the vertical stratification of the assemblage present in this floodplain forest in the Pantanal of Mato Grosso. Rev. Biol. Trop. 65 (2): 445-459. Epub 2017 June 01. Key words: Chernetidae, flood areas, richness, vertical distribution, Withiidae. Floodplains, especially the Brazilian Pan- its geological and geomorphological history, tanal, are characterized by the controlling influ- results in a variety of landscapes, vegetation ence of flood pulses and vegetation, which types (including monodominant forests), and have a major impact on the population dynam- habitats with specific characteristics, consti- ics of species, thereby influencing community tuting real vegetation mosaics (Silva, Mauro, structure (Junk, Bayley, & Sparks, 1989; Junk, Mourão, & Coutinho, 2000; Nunes-da-Cunha 1997). In the Pantanal, the association of & Junk, 2011). The annual and multi-annual Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 65 (2): 445-459, June 2017 445 variability of the flood pulse affects the biota at 2002). Other species are strictly inhabitants of different intensities and time scales (Nunes-da- the forest canopy (Adis et al., 1988). Cunha & Junk, 2004), causing terrestrial organ- Considering the diversity of habitats in isms to develop specific strategies to survive inundated forests and the influence of seasonal and adapt to the strong seasonality, characteris- variations on the Pseudoscorpiones assemblage, tic of this floodplain (Adis, Marques, & Want- different sampling techniques were applied zen, 2001; Battirola, Marques, Rosado-Neto, in an integrated way in this study, in order Pinheiro, & Pinho, 2009). to assess (i) the abundance and richness of As in other regions of South America in Pseudoscorpiones in the soil, leaf litter, termite the Brazilian Pantanal (e.g. Mahnert & Adis, nests, tree trunks and canopies in an inundated 2002), the knowledge about ecological and tax- forest of Vochysia divergens (Vochysiaceae) in onomical diversity of pseudoscorpions is under the Northern region of the Brazilian Pantanal; explored, as well as their relationship with the (ii) evaluate the occurrence over the different different habitats. The Pseudoscorpiones are seasonal periods of the region and (iii) verify if arachnids that can occupy a variety of terres- there is habitat specificity among the Pseudo- trial habitats, mainly edaphic and cave environ- scorpiones which use this area. ments (Murienne, Harvey, & Giribet, 2008), and can colonize the forest canopy (Mahnert MATERIAL AND METHODS & Adis, 1985; Adis, Mahnert, De Morais, & Rodrigues, 1988; Battirola, Adis, Marques, & Study area: This study was carried out Silva, 2007; Mahnert, 2009; Marques, Adis, in the Pantanal of Poconé (16°15’ S - 56°36’ Battirola, Santos, & Castilho, 2011), or live W) in the municipality of Nossa Senhora do associated with the nests of social insects, birds Livramento, Mato Grosso, Brazil. Sampling and mammals (Mahnert & Adis, 2002). Esti- was carried out between January 2004 and mates show that the order Pseudoscorpiones March 2005, in an inundated forest with pre- dominance of V. divergens (Vochysiaceae), has 3 533 species worldwide (Harvey, 2013), which represents one of the region’s typical although earlier projections had already esti- phytofisionomies, called “cambarazal” (Nunes- mated that the number was close to 5 000 spe- da-Cunha & Junk, 2014). The sampling with cies (Adis & Harvey, 2000; Adis, 2002). the Winkler extractor was performed between In Brazil, most of the ecological studies 1999 and 2000. on Pseudoscorpiones were carried out in the This region is characterized by four well- Central Amazon (Adis, 1981; Adis & Mahnert, defined seasonal periods (Heckman, 1998). The 1985; 1990, a, b; 1993; Mahnert & Adis, 1985; dry season occurs between July and September. De Morais, Adis, Mahnert, & Berti-Filho, In this period most of the Northern Pantanal is 1997; Aguiar & Bührnheim, 1998, a, b; 2003; completely dry and rainfall is scarce. The start Aguiar, Gualberto, & Franklin, 2006). Some of the rainy season corresponds to the rising populations of Pseudoscorpiones may have water period (between October and December), undergone extensive evolution within inunda- when some areas may flood up temporarily tion forests in this region, due to forest isolation with water from the rains, but become dry again caused by climatic shifts and micro-geographic after periods of sun. The high water period changes (Adis & Schubart, 1984). In these generally occurs between January and March environments the terricolous species use the or April. In this period much of the Northern soil and leaf litter as a breeding place during region of the Pantanal is submerged, fields the non-flooded period and migrate later to tree and forests are flooded due to the overflow of trunks and regions of the forest canopy, where rivers and lakes. The receding water season is they remain during the flood periods (Adis & the period in which the water level in flooded Mahnert, 1990a; Adis, 1997; Mahnert & Adis, areas drains rapidly, accompanied by reducing 446 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 65 (2): 445-459, June 2017 the amount of rainfall in the region, recorded mounds within the forest and in contact between April and June. with water were selected (March 2004) (Fig. 1F). These mounds had the apical Field procedures: Different collection portion, above the water level, cut with an methodologies were employed for the soil and electric saw, packed in plastic bags, and leaf litter, tree trunks and canopies in the dry, transported to the laboratory for manual rising water, high and receding water periods collection of the pseudoscorpions. (e.g. Adis, 1981; 2002; Battirola et al., 2009). d. Winkler extractor: Samples were collected Only the collections from termite mounds in February (high water), May (receding and trunks following the waterline were per- water), October (dry season), and Decem- formed exclusively during the high water ber (rising water) of 2000 (Pinho, 2003). period, aiming to evaluate specifically this A total of 38 m2 of surface soil (0 to 5 period. Details of each methodology applied cm) and leaf litter were sampled, with are presented below: 10 m2 in the high water, dry season, and rising water periods, and 8 m2 during the a. Pitfall traps: These traps consist of a 20 cm receding water period. Details about the polyethylene flask with a