Seasonal Variation of Ground Spiders in a Brazilian Savanna
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ZOOLOGIA 27 (3): 353–362, June, 2010 doi: 10.1590/S1984-46702010000300006 Seasonal variation of ground spiders in a Brazilian Savanna Marina Farcic Mineo1; Kleber Del-Claro1, 3 & Antonio Domingos Brescovit2 1 Instituto de Biologia, Universidade Federal de Uberlândia. Campus Umuarama, Rua Ceará s/n, 38400-902 Uberlândia, Minas Gerais, Brasil. 2 Laboratório de Artrópodes Peçonhentos, Instituto Butantan. Avenida Vital Brasil 1500, 05503-900 São Paulo, São Paulo, Brazil. 3 Corresponding author. E-mail: [email protected] ABSTRACT.The Brazilian Savanna Ecoregion (Cerrado) is one of the richest biomes in the world, with a characteristic highly seasonal climate a dry season between May and September and a rainy season from October through April. Ground-dwelling spiders from three Cerrado phytophysiognomies, “campo cerrado”, “cerrado” and “cerradão”, were sampled using pitfall traps during two years, totaling 111 species and 3,529 individuals. The abundance of individuals and species richness was higher during the wet season. Fifty-eight species were captured exclusively during that period, whereas only nineteen were restricted to the dry season. Only two species were found in all samples. The number of juveniles was higher than the number of adults in all phytophysiognomies and in all species during both seasons. The highest abundance was registered in October and the lowest in April. Overall sex ratio was male-biased in all vegetation types sampled. Distinct climate variables affected the abundance of spiders depending on sex, age and vegetal physiog- nomy where they were sampled. This study involved the longest sampling of spider abundance and diversity on the ground of a Brazilian Savanna. KEY WORDS. Araneae; Brazil; community ecology; seasonality. Seasonality (WOLDA 1988) is an important factor to con- are quite diverse – in species and micro-habitats, see OLIVEIRA & sider in studies that discuss species richness and abundance. MARQUIS (2002) and GOTTSBERGER & SILBEBAUER-GOTTSBERGER (2006) Despite its relevance, however, seasonal variations have received –, occupying 25% of the Brazilian territory, studies on spiders little attention from Tropical environmentalists until recently are rare and restricted to a single phytophysiognomy or guild (e.g. ROSENZWEIG 1995). Although seasonal changes in tempera- (e.g. RINALDI & FORTI 1996). ture tend to be minimal or absent in the Tropics, it is not accu- This study tested the hypothesis that the seasonal varia- rate to say that there are no seasons (WOLDA 1988). In fact, tion in temperature and humidity observed in a Brazilian sa- rainy seasons alternate with one or two dry seasons each year. vanna promotes a strong seasonal pattern in the species com- Seasonal changes in flora and fauna are determined by position and abundance of ground spiders. abiotic conditions including weather variables. Animals, due to behavioral and physiological characteristics, react differently MATERIAL AND METHODS to the influence of abiotic factors. Some studies have demon- strated that seasonality influences the abundance, growth rate, The present study was carried out at the Panga Ecologi- and size of spiders (GASTON et al. 1993, GASNIER et al. 2002). En- cal Station, an area of 404 hectares, located 30 Km south of the vironmental characteristics also have a strong influence on city of Uberlândia (19°11’10”S, 48°23’30”W), state of Minas spider habitat selection (UETZ 1992). Gerais, southeastern Brazil (Fig. 1). The region, located 800 m Spiders play an important role in many terrestrial eco- above sea level, is characterized by a highly seasonal climate, systems due to their predatory nature, abundance, and ubiq- with dry winters (from April through September) and rainy uity. Because they are at the top of the invertebrate trophic summers (from October through March). The average annual chain these arachnids can regulate decomposer populations precipitation is 1500 mm. Local temperature and precipitation (WISE 1993). were recorded during the two years of the study (Fig. 2). Little is known about the influence of seasonality on the Three different Savanna formations (phytophysiogno- araneofauna of Savanna regions, even though a few well-docu- mies), namely campo cerrado, cerrado and cerradão, were mented studies have been conducted in some African coun- sampled. The campo cerrado is characterized by grasslands with tries (e.g. RUSSEL-SMITH 2002, WHITMORE et al. 2002, MODIBA et al. scattered shrubs and small trees. The cerrado vegetation is pre- 2005). Despite the fact that Brazilian Savannas (Cerrado Biome) dominantly arboreal-shrub with an herbaceous stratum, 20- © 2010 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | All rights reserved. 354 M. F. Mineo et al. 48°24’ N Uberlândia WE S 19°10’ Trails Campo Úmido Campo Sujo BRAZIL Campo Cerrado Cerrado Cerradão Forest Antropic area 19°11’ 0 200 400 600 m Campo Florido Figure 1. Location of the Panga Ecological Station, an area of preserved Brazilian Savanna. Adapted from SCHIAVINI & ARAUJO (1992). 500 40 Adults were sexed and identified to the lowest taxonomic 450 35 level possible. All specimens were identified at least to family. 400 30 350 For the analyses, BioEstat 5.0 was used (AYRES et al. 2007). Spider 300 25 abundances were used in stepwise multiple regressions against 250 20 200 15 the following local environmental variables: maximum, mini- 150 10 mum and medium temperature, relative humidity, total precipi- 100 Temperature (°C) Precipitation (mm) 50 5 tation and maximum precipitation during 24 hours. Because we 0 0 wanted to investigate whether climatic variables affect the araneofauna and, if so, to ascertain whether the effect of these Jul/05 Jul/06 Apr/05 Apr/06 Oct/05 Oct/06 Jun/05 Mai/06 Jan/06 Jun/06 Jan/07 Mai/05 Feb/06 Mar/06 Feb/07 Sep/06 Aug/05 Dec/05 Sep/05 Dec/06 Aug/06 Nov/06 Nov/05 variables is immediate or delayed, we analyzed data pertaining Figure 2. Climate data from the Panga Ecological Station, Minas to the sampling month as well as data from one and two months Gerais, Brazil. (᎐) Minimum temperature, (--) maximum tempera- prior to sampling. Voucher specimens were deposited in the ture and (bars) precipitation. Arachnida and Myriapoda Collection of the Instituto Butantan, São Paulo, Brazil (Curator: A.D. Brescovit). 50% tree cover and average tree height of three to six meters. RESULTS The cerradão is an almost closed woodland with a 50% to 90%crown tree cover (trees are often three to eight meters, or The species composition of the ground-dwelling spiders, as taller), casting considerable shade that reduces surface expo- well as their distribution over the two years of the study, varied sure (OLIVEIRA-FILHO & RATTER 2002). according to habitat type and season. A total of 111 species, 31 Ground spiders were collected using pitfall traps. Each families and 3,529 individuals were collected (Appendix 1). Most trap consisted of a plastic jar 7 cm in diameter and 10 cm long. spiders were collected during the wet season (N = 2,578 individu- Traps were inserted into the ground and were half filled with a als and 90 species). In the dry season, 951 individuals and 51 mixture of 70% ethanol and glycerol. In each species were captured. Of the species sampled, 58 were restricted phytophysiognomy, an area of 2,592 m2 was divided into 12 to the wet season, whereas 19 were found exclusively during the quadrants of 12 x 18 m each. One quadrant in each habitat dry season. Several families (N = 11, 35%) and many species (N = type was sampled every two months. Traps were aligned in five 67, 60%) were represented by less than three individuals. Only rows, each containing eight traps, with a distance of two meters two species were sampled throughout the 12 periods of collec- between rows and between traps. Traps were left in the field tion: Apopyllus silvestrii (Simon, 1905) (N = 73), and Hahniidae sp. for five consecutive days. Each quadrant was sampled only once. 1 (N = 76). Twenty-nine percent of the specimens were captured The total trapping effort involved 2,400 traps for each site, in the campo cerrado, 37% in the cerrado and 34% in the cerradão. sampled between April, 2005 and February, 2007. Climate variables affected spider abundance differently, ZOOLOGIA 27 (3): 353–362, June, 2010 Seasonal variation of ground spiders in a Brazilian Savanna 355 depending on the sex and age of the spiders and the months before sampling. Juvenile abundance and richness in phytophysiognomy of the sampling site (Tab. I). Total spider the Campo cerrado, and male abundance in the Cerradão were abundance was affected primarily by relative humidity two not affected by any of the climate variables measured. months before sampling. Additionally, in the Cerradão, climatic The number of juveniles was higher than that of adults in changes in the sampling month also influenced the spiders. all habitat types and seasons, with the exception of the wet sea- Richness and abundance of males were affected by different son in the Cerrado and Cerradão. During the two years of our climate variables depending on habitat type, for example, abun- study, the highest number of males was observed in October (N dance of females from the three habitat types were influenced = 1,131), except for the Campo Cerrado, where the abundance mainly by maximum temperature and maximum precipitation of males during the first year was higher in December (Figs 3-6). in a period of 24 h during the sampling month. Juveniles in Abundance of females did not change significantly dur- the Cerrado and Cerradão were affected by distinct climatic ing the two years. During the first year, the lowest abundance factors: temperature, humidity and precipitation, starting two of juveniles was recorded in October and, over the second year, Table I. Significant relationships between ground spiders of three phytophysiognomies of the Brazilian savanna and climate variables (maximum, medium and minimum temperature, relative humidity, total precipitation and maximum precipitation in a period of 24 hours) collected one and two months before, and during the sampling month, through multiple linear regression stepwise.