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Araneae: Pholcidae) from the Brazilian Dry Forest Vanessa Stefania,B,F*, Kamila Garciaa , Carliane Vecchiaa , Lígia A

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Araneae: Pholcidae) from the Brazilian Dry Forest Vanessa Stefania,B,F*, Kamila Garciaa , Carliane Vecchiaa , Lígia A Journal of Natural History Vol. 46, Nos. 33–34, September 2012, 2117–2129 Mating behaviour, nympho-imaginal development and description of a new Mesabolivar species (Araneae: Pholcidae) from the Brazilian dry forest Vanessa Stefania,b,f*, Kamila Garciaa , Carliane Vecchiaa , Lígia A. Silvaa , Barbara Guimarãesa , Everton Tizo-Pedrosoc , Ewerton O. Machadod,e, Antonio D. Brescovitd and Kleber Del-Clarof aUniversidade Presidente Antônio Carlos, UNIPAC,/Facisa/Campus Araguari, MG, Brazil; bPrograma de Pós-Graduação em Ecologia e Conservação dos Recursos Naturais, Universidade Federal de Uberlândia, 38400-902, Uberlândia, MG, Brazil; cUniversidade Estadual de Goiás, Un.U. de Morrinhos, GO. Brazil; dLaboratório Especial de Coleções Zoológicas, Instituto Butantan, Av. Vital Brazil, 1500, 05503-900 Butantã, São Paulo, SP, Brazil; eDepartamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Brazil; fLaboratório Ecologia Comportamental e de Interações, LECI, Inst. de Biologia, PPGECRN, Universidade Federal de Uberlândia, 38400-902, Uberlândia, MG, Brazil (Received 28 October 2011; final version received 9 May 2012; printed 13 September 2012) A new species of Mesabolivar is described from Brazilian forests: Mesabolivar delclaroi. The mating behaviour, postembryonic development, number of egg sacs and eggs, birth rate, number of instars, developmental time until adulthood, sex ratio and size of cephalothorax (per instar) were recorded. The sexual behaviour was described and categorized into four steps: courtship, pre-copulation, copula- tion and post-copulation. After hatching, individuals presented five instars until maturity. The mean number of eggs (42 ± 16.6) and live births (31.5 ± 3.4) of the first egg sac were significantly greater than that of a second one (23.8 ± 3.8, and 19.25 ± 3.9, respectively). The developmental time from birth to adult- hood (130.8 ± 9.6 days) did not differ significantly between egg sacs produced (128.61 ± 11.1). The size of the cephalothorax did not differ among adults or between sexes. The sex ratio revealed a shift in favour of females (4: 3). Keywords: developmental stages; life history; Pholcidae; sexual repertoire; taxonomy Introduction The current catalogue of spider species includes 42,055 known species in 3821 genera and 110 families (Platnick 2010). The knowledge of the fauna of spiders in the Neotropical region is incipient and to estimate the proportion of species still unknown would be a difficult task to perform (Santos et al. 2007). In addition to the restricted knowledge about the Neotropical spider fauna is the limited knowledge of the natu- ral history, biology and behaviour of its species. Considering the importance of this taxa inside trophic chains and interactive networks, natural history, biology and tax- onomic studies on spiders of endangered tropical systems are urgent (e.g. Del-Claro and Torezan-Silingardi 2009). *Corresponding author. Email: [email protected] ISSN 0022-2933 print/ISSN 1464-5262 online © 2012 Taylor & Francis http://dx.doi.org/10.1080/00222933.2012.707243 http://www.tandfonline.com 2118 V. Stefani et al. In relation to spider biology, it is known that after the hatching of the egg, the nympho-imaginal period (that between first nymphal instar up to adulthood, sensu Foelix 2011) is determined by multiple ecdyses until the final stage at adulthood (Gullan and Cranston 1994). In spiders, similar to what is known in other arthro- pods, the sexual behaviours are ritualized (Eberhard 1994; Roberts and Uetz 2004). Among adults, reproduction is characterized by behavioural sequences that integrate the courtship process, based on movements and signs performed by both sexes (Uetz and Stratton 1982). The courtship performed by the male is composed of signals to stimulate the female to accept copulation (Huber 2005; Peretti et al. 2006; Costa and Quirici 2007). Pholcidae currently has 84 genera and 1111 species distributed across all continents (Platnick 2010). The genus Mesabolivar González-Sponga 1998 currently includes 36 species and along with Metagonia Simon 1893 is one of the genera most abundant and well distributed in South America (Huber 2000; Machado et al. 2007). However, few species have been studied in detail like Mesabolivar eberhardi Huber 2000 has been (Eberhard and Briceño 1983, 1985). Information about the biology, habitat prefer- ence or temporal variation is still very limited for most species (Machado et al. 2007). In the present study we report a new species in the genus Mesabolivar, describing and analysing its courtship and mating behaviours and characterizing and detail- ing the nympho-imaginal stages (Downes 1987; Wheleer et al. 1990). Additionally we quantified the number of egg sacs, eggs and instars, evaluating birth rate, devel- opmental time to adulthood (e.g. Nentwig 1990; Stefani and Del-Claro 2011). The sex ratio and size of the cephalothorax were also assessed along with the sexual behaviour. Materials and methods Twenty immature males and 20 young females of Mesabolivar delclaroi sp.nov.were collected between January and February 2009 in the John Kennedy Park in the city of Araguari, state of Minas Gerais, Brazil. The grove is a semi-deciduous seasonal forest (11.2 hectares) in the urban perimeter, where trees reach 15 m in height; the soil is rich in litter, especially in the dry season between June and October (Souza and Araújo 2005). Of the 40 individuals collected, all were immature (second and third instar and non-reproductive), seven males and eight females were found within a decaying trunk; four males and eight females were in holes in the ground; and nine males and four females were at the bottom of the funnel web of Aglaoctenus lagotis Holmberg 1876 (Araneae: Lycosidae). The spiders were collected manually, individually stored in 1-litre plastic bottles and transported to the Zoobotanical Laboratory at the President Antônio Carlos University in the Araguari Campus. The M. delclaroi sp. nov. individu- als were kept at an average temperature of 25.5◦C and relative humidity of 60 ± 14.8% (mean ± SD). When the spiders reached sexual maturity (adulthood), 15 males and 15 females were randomly selected and paired up for the observation and description of courtship and copulation. Three females and two males were killed and stored as voucher spec- imens and deposited in the collections of the Instituto Butantan, São Paulo (IBSP, D. Battesti) and Museu de Zoologia da Universidade de São Paulo, São Paulo (MZSP, R. Pinto da Rocha). The other individuals were deposited in the scientific collection of Universidade Presidente Antonio Carlos (UNIPAC, Araguari). Journal of Natural History 2119 The behavioural observations of reproductive behaviour were performed on the spider pairs (n = 15) settled in terraria. The mating arena was 15 × 10 cm and 15 cm high. Fifteen independent observations were performed using 15 different pairs. For each observational event, the female was first introduced into the terrarium and the male was only introduced after the construction of the web by the female, which took 36.8 ± 9.2 hours (mean ± SD; n = 15). All pairings (laboratory-simulated sexual ® encounters) were recorded with a high-definition video camera (Moticam System ), linked to a trilocular stereomicroscope. Films were analysed until each sequential act could be clearly identified (according to Altmann 1974).The observations of sexual behaviour were initiated only after the first vibration performed by the male on the web and ended after the separation of the couple. The males were removed from the terrar- ium 15 hours after their introduction. After mating, the individuals were removed; the terrarium was cleaned with 98% alcohol and reused for the next pair. We considered an interval of at least 24 hours between events (before running the next trial) and repeated the previously described methodology. After removing males from the test arena, the females were returned to their original bottles and followed up until the formation of the oothecae and their hatch- ing. The dispersion of the spiderlings occurred shortly after hatching. The already empty cocoons, the ones that the spiderlings had already left, were taken to the laboratory to count the number of hatched eggs and eggs that did not develop. The number of eggs deposited in each cocoon was also counted. The spiderlings born in the laboratory were maintained individually, followed up until adulthood and fed with beetle larvae (Palembus dermestoides; Coleoptera: Tenebrionidae) and termite nymphs (Armithermes sp.). We considered as the first instar the young spi- ders emerging from the egg sac (which may, or may not, have moulted inside the egg sac). The spiders were observed daily and each exuvium from the instar changes was kept dry, labelled and stored in individual envelopes for each individual. Each exuvium was placed between a slide and cover slip for measurement of all its structures in the same plane with an optical microscope using an ocular scale (Marchioro et al. 2005). The time between ecdyses, the number of survivors at each developmental stage and the sex ratio in adulthood were also evaluated. The normal distributions of the following variables were evaluated using the Kolmogorov–Smirnov normality test with the Lilliefors correction: (1) average num- ber of eggs; (2) number of individuals born alive; (3) developmental time in days; and (4) size of the cephalothorax of individuals from the first and second ovipositions The analysed variables showed non-normal distribution, so the Mann–Whitney U-test was chosen as a non-parametric alternative to the analyses of variables 1 to 4, in systat (Systat Software, Inc., Chicago, IL, USA) program. Descriptions, the terms “vertical hairs”, “curved hairs” and ratio L/dofthe tibia I (length/diameter, a measure of the robustness of the legs) follow Huber (2000). Measurements are given in millimetres. The epigynum was dissected and immersed in clove oil for visualization of internal structures following Levi (1965). Spiders were compared with others of the same locality, one male three females, 20 April 2011, V.
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