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, copulation 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 adulthood (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 natural 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]

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, developmental 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. individuals 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 terrarium 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 hatching. 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 spiders 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 number 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. Stefani,(IBSP161670–161673); Goiás, Morrinhos, Parque Natural de Morrinhos (17◦43 S, 49◦06 W), two females, 18 March 2011, V. Stefani col. (IBSP 161668–161669). All illustrations were made with a camera lucida on a Leica MZ12,5 stereomicroscope. 2120 V. Stefani et al.

Results

Mesabolivar delclaroi Machado and Brescovit sp. nov. (Figure 1)

Types Male holotype from Bosque John Kennedy (18◦38 59 S, 48◦1105 W), Araguari, Minas Gerais, Brazil, September 2009, B. Guimarães and V. Stefani col., deposited in IBSP (160720). Paratypes: one male (IBSP 160722) and one female (IBSP 160721) same data as holotype.

Etymology The speciﬁc name is a patronym in honour of the ethologist Kleber Del-Claro.

Diagnosis Very close to Mesabolivar togatus (Keyserling, 1891), sharing the procursus curvature, the presence of a proximal protrusion and a flat prolateral apophysis in the chelicerae; and the shallow epigynum pocket (see Huber, 2000, figs. 851–863 for comparison). The male is distinguished by the discrete size of proximal protrusion on chelicerae; the rounded and not totally flat tip of prolateral cheliceral apophysis (Figure 1A); and the distinct tip on the procursus in the palp (Figure 1D–F) and presence of a barely visible posterior margin of the epigynum pocket in the female epigynum (Figure 1H).

Description Male (Holotype). Total length 3.8, carapace length 1.3, clypeus 0.4, carapace width 1.5; leg I: 41.3 (10.9+0.6+10.6+19.2+missing), tibia II: 7.5, tibia III: 6.7, tibia IV: 4.8, tibia I L/d: 71. Body very similar to M. togatus. Carapace light brown with a longitudi- nal dark stripe; sternum reddish-brown. Distinct thoracic groove. Eight eyes on slightly elevated ocular area; distance PME-ALE about 100% of PME diameter. Chelicerae light brown, basal segment with a medial apophysis on the prolateral edge, ﬂat pointed and a pointed proximal apophysis (Figure 1B). Palps as in Figure 1(C). Coxa with distinct retrolateral apophysis (Figure 1C). Femur with retrolateral proximal apophysis, rectangular shaped (Figure 1C). Procursus light brown, dorsally curved, with dorsal microspinulation (Figure 1C). Distinct tip of procursus, with projections and a distal membranous projection (Figure 1D–F). Bulb simple, without transparent projection (Figure 1G), embolic division long and straight, membranous distally, sclerotized prolateral stripe present (Figure 1G). Legs light brown, femora with subdistal dark rings; without modiﬁed spines, vertical or curved hairs. Retrolateral trichobothrium on tibia I at 1.5%. Tarsi without serrated ventral spines, tarsus II with approximately 19 pseudo-segments. Oval, greenish-blue abdomen with grouped darker spots.

Female (Paratype, IBSP 160721). Total length 3.6, carapace length 0.9, carapace width 1.3; leg I: 17.3 (7.9+0.5+8.1+11.9+1.4), tibia II: 5.6, tibia III: 3.4, tibia IV: Journal of Natural History 2121

A B C

D E

F G J

I H

Figure 1. Mesabolivar delclaroi sp. nov. (A, B) Male chelicerae, (A) frontal, (B) lateral; (C) male left palp, retrolateral; (D–F) procursus tip, (D) prolateral, (E) dorsal, (F) retrolateral; (G) bulb, prolateral; (H–J) epigynum, (H) ventral, (I) lateral, (J) dorsal. Scale bar 0.5 mm.

5.3, tibia I L/d: 54. In general, very similar to male. Epigynum brown, slightly elevated, without projections, with a discrete epigynal pocket, centred (Figure 1H). Internal genitalia with two porous plates approximately parallel, connected by a dorsal sclerotized region (Figure 1I).

Variation One male: carapace width 1,4; tibia I: 10.1. Two females: carapace width 1.1–1.4; tibia I: 6.2–9.7. 2122 V. Stefani et al.

Distribution Known only in the type locality. Other material examined. Brazil, Minas Gerais, Araguari, Bosque John Kennedy (18◦3859 S, 48◦1105 W), two females September 2009, B. Guimarães and V.Stefani col. (IBSP 160723, 160724).

Courtship and copulation behaviour The courtship and copulation were successful for all 15 experimental couples. The quantification and categorization of the behaviours suggest four distinct steps: courtship, pre-copulation, copulation and post-copulation. Courtship – short and interspersed beats of the first pair of legs of the male on the web were observed, with the cephalothorax always directed to the partner (time in minutes, 7.3 ± 5.4, mean ± SD, n = 15, Figure 2: step I). Alternated chelicerae movements were also documented. Pre-copulation – consists of the first physical contact between the couple. The male quickly walks towards the female, directing its cephalothorax to the partner’s abdomen (cephalothorax directed to opposite sides), positioning the pedipalps on the female’s abdomen (time in seconds, 1.5 ± 0.3, mean ± 1SD, n = 15, Figure 2: step II). Copulation – the simultaneous insertion of the genital bulbs in the female genital orifices (time in minutes, 5.1 ± 2.3, mean ± 1SD, n = 15). The bristles located in the genital cimbia rub the female’s abdomen whenever she moves and the male’s chelicerae engage the female genitalia during the copulation (Figure 2: step III). The release of the bulbs marks the end of the copulation and the beginning of the last phase: Post-copulation – in this phase, the male walks away and cleans the entire pedipalp (Figure 2: step IV).

Egg sac production and births After a single mating, females produced the first oviposition in 19.25 ± 4.5 days (mean ± SD) and the second 33.14 ± 4.2 days after the hatching of the first egg sac. The number of eggs in each egg sac of the first oviposition was 42 ± 16.6 (mean ± SD, n = 504 eggs) whereas in the second oviposition a significantly smaller average was obtained, 23.83 ± 3.76 (n = 286 eggs, U = 126.5, P = 0.002, Mann–Whitney U-test, Figure 3). The average number of births for the first egg sac was 31.5 ± 3.4 (mean ± SD; n = 379 immatures), whereas for the second oviposition it was 19.25 ± 3.9 (n = 231 immatures) significantly smaller than in the first round (Mann–Whitney U-test, U = 143.5, P = 0.001, Table 1). Throughout the development a decrease in the number of survivors until adulthood occurred in the two quantified ovipositions. In adulthood, 4.8 ± 1.1 (mean ± 1SD) survivors were obtained in the first oviposition and 3.9 ± 1.5 were obtained in the second; these numbers did not differ significantly (Table 1). The developmental time from birth to adulthood was 130.8 ± 9.6 days (first egg sac, mean ± SD) and 128.61 ± 11.1 days (second egg sac), not differing significantly (Mann–Whitney U-test, U = 6.45, P = 0.142) and each instar possessed its own specific growth interval (time in days, Table 1). The average length of the cephalothorax in the first instar was 0.23 ± 0.05 mm (mean ± 1SD, n = 379) and in the second instar it was 0.752 ± 0.05 mm (n = 201), reaching 1.85 ± 0.14 (first oviposition, mean ± SD, n = 58) and 1.83 ± 0.33 (second oviposition, n = 47) in adulthood. In the last Journal of Natural History 2123

a) b)

2 3

(II) PRE-COPULATION (III) SPERM TRANSFER

(IV) POST- COPULATION

(I) COURTSHIP 4

Figure 2. Diagram of the sexual repertoire displayed by the Mesobolivar delclaroi sp. nov. spider. The behaviours are divided into four steps: Step I, Courtship; Step II, Pre-copulation – note in the arrow the early introduction of the copulatory bulb into the female’s genitalia; Step III, Copulation, (a) lateral view, (b) ventral view – note the arrows indicating the pedipalp bristles rubbing the abdominal region of the female; Step IV, Post-copulation. developmental instar the male cephalothorax size was of 2.1 ± 0.2mminlength (mean ± 1SD, n = 25) and 1.9 ± 0.3 mm (n = 33) in the female for ﬁrst oviposition, not differing signiﬁcantly (Mann–Whitney U-test, U = 21.3, P = 2.52). The sexual proportion indicated a greater number of females when compared with males, 4: 3, so indicating a bias of the sex ratio towards females.

Discussion In many arachnids the courtship begins with an exchange of signals (e.g. vibrations, odours) so that the female recognizes that the spider courting is a co-specific male (e.g. Suter and Renkes 1982). In the courtship of M. delclaroi, movements of beats of the legs on the webs and of the male’s chelicerae when he was introduced into the arena with the female confirmed this. These data can be interpreted as an intra-specific sig- nal during the courtship, important for the male to show his reproductive intentions to the female (Huber and Eberhard 1997; Schäfer and Uhl 2002). According to Uetz and Stratton (1982) the stridulatory organs may be involved in acoustic communication. In the Pholcidae, the genera Crossopriza (Wiehle 1933), Physocyclus (Brignoli 1981), 2124 V. Stefani et al. 70

* 60

Number of eggs 30

10 1°oviposition 2°oviposition

Figure 3. Average number of eggs (median ± SD) in the ﬁrst and second oviposition. ∗Indicates signiﬁcant difference between the averages, Mann–Whitney U-test, U = 126.5, P = 0.002.

Gertschiola (Brignoli 1981), Ninetis (Kraus 1957), Holocnemus, Hoplopholcus and Holocneminus (Huber 1995) have stridulatory organs (movements of the pedipalps against the chelicerae) that are used during courtship, which is an important behaviour for the female to accept copulation. Peretti et al. (2006) suggested that female stridulation in Physocyclus globosus represents an attempt to induce the male to interrupt genital squeezes: female stridulation was more common when the male was squeez- ing her. Females were more likely to stridulate when individual male squeezes were longer, and when the male did not answer to a previous stridulation by loosening a squeeze. However, stridulations may also be related to other functions, for example, in pseudoscorpions it can be involved in mating as in complex social behaviours (see Tizo-Pedroso and Del-Claro 2011). The pholcids are the only non-entelegyne spiders whose reproductive biology has been carefully studied in several species (Kaster and Jacob 1997; Uhl 1993, 1994; Uhl et al. 1995, Yoward 1998; Huber 1999 – for the haplogyne spiders see Burger et al. 2003, 2006). Other advantages of this family considering the study of sexual selection Journal of Natural History 2125 0.33 0.29 0.04 0.33 0.04 ± ± ± ± ± SD) ± 0.34 1.83 0.35 1.06 0.05 0.73 0.3 1.2 0.05 0.22 (mean ± ± ± ± ± Cephalothorax size (mm) 3.6 1.85 2.63 1.1 2.1 0.75 2 1.34 0.7 0.23 ± ± ± ± ± SD) ± -test. Time (days) 331 2.57 31.7 0.81.67 3.8 28.7 1.5 33.41 U (mean ± ± ± ± ± 1.5 32.3 1.88 31.6 3.92.9 3.9 28.5 1 34.5 ± ± ± ± ± SD) 4.66 0.05; Mann–Whitney 19.25 13.41 ± < ∗ ∗ ∗ P 2.9 1.1 3.9 3.7 3 7.08 3.4 (mean ± ± ± ± ± sp. nov. indicating the number and mean of individuals, time in days and size of the Mesobolivar delclaroi 1st ovip. 2nd ovip. 1st ovip. 2nd ovip. 1st ovip. 2nd ovip. 1st ovip. 2nd ovip. Significant difference between the first and second oviposition with Instar IV 89 56 7.4 Instar V (adults) 58 47 4.8 Instar II 201 161 16.75 Instar III 100 85 8.3 Table 1. Postembryonic development of PhasesInstar I Number of individuals∗ Number of individuals 379 231 31.5 cephalothorax in the five instars observed in the first oviposition (1st ovip.) and second oviposition (2nd ovip). 2126 V. Stefani et al. are the numbers of synanthropic species that are available throughout the world, which are easy to keep in laboratories for studies and also share the environment with a rich fauna (mostly tropical) for comparative studies. Two male structures deserve special attention: the chelicera and the clypeus. At least one of these structures engaged the female during copulation in all species studied so far (Huber 1994, 1995, 1998; Huber and Eberhard 1997). Participation of the chelicera along with male palp was observed in the M. delclaroi copulation. A hypothesis proposed by Eberhard (1985) assumes that the reproductive isolation based on the cheliceral morphological differences evolved to avoid hybridization. Hence, the male of each species will have a different morphology fitting only in the genitalia of the female of the corresponding species (“Lock-and-key” mechanism: Crane 1949; Grasshoff 1975; Foelix 2011). In pholcids, this is especially true for chelicera, studied by Huber (1999, 2003) and Huber et al. (2005), where different cheliceral apophyses fit similar epigynal pockets. Another plausible hypothesis is based on con- flict of interest (Alexander et al. 1997) applied to the modification of the chelicerae in Pholcidae that could be substantiated by physical repression of the male during the mating event overcoming the resistance of the female (Huber 1995). Therefore, the chelicerae can serve as courtship and copulation devices of which the elaborate morphology is used to stimulate or adjust the female in a way that increases the chances of the male increasing its fitness. The results of reproductive biology and development of M. delclaroi showed the capacity of production of two cocoons during the reproductive period and that the second egg sac presented a smaller number of eggs and consequently fewer births when compared with the first egg sac. In the natural environment this strategy of production of additional egg sacs may confer a new reproductive attempt if the first has been predated or parasitized, even when the new oviposition has a smaller number of eggs (Brown et al. 2003). This decline in the number of eggs in the egg sac can be attributed to a decrease in the stocked resources, because during the reproductive period all efforts are directed to the protection of the eggs and not on feeding (Nussbaum 1981; Ferguson et al. 1982; Brown et al. 2003; Stefani et al. 2011). The number of recorded instars in M. delclaroi was five and the average developmental time was similar in the spiderlings from the two ovipositions (which received equal amounts of food). Uhl et al. (2004) observed in an experiment conducted with Pholcus phalangioides Fuesslin 1775 spiderlings that they also presented five instars, but the individuals that ate three times more presented a larger body size in adulthood and a shorter developmental time when compared with the individuals that received a controlled diet. The developmental time, the number of instar changes before reaching maturity and the size of the spiders are fundamental aspects to understand the biology of the studied spider and are important in assisting in the elaboration of a protocol for breeding in captivity (Galiano 1967; Bernardo 1996; Marchioro et al. 2005). In this study M. delclaroi presented a biased sex ratio for females once they reached adulthood. Differences in the width of the cephalothorax between males and females of M. declaroi were not significant, corroborating the monomorphic character of Pholcidae, according to Elgar (1992) and Huber (2005). The present study is the first characterizing all stages of life development of a Pholcidae, M. delclaroi. These data can assist in the initial formation/elaboration of reproductive behaviour descriptions, on studies of the post-larval development of M. delclaroi and providing parameters for comparative studies with other species of Journal of Natural History 2127 this family, specially the less studied New Word Pholcids. The descriptions of species accompanied with other information enrich the knowledge of the species and should be encouraged (e.g. Del-Claro et al. 1997).

Acknowledgements We are grateful to the municipality of Araguari for the permission granted to work in the JK Park. The authors are grateful for the ﬁnancial support from UNIPAC/FUNADESP – Universidade Presidente Antônio Carlos/Fundação Nacional do Desenvolvimento do Ensino Superior Particular (V. Stefani, K. Garcia, C. Vecchia), Fapemig (E. Tizo-Pedroso); CNPq/Fapemig (K. Del-Claro) and CNPq (A. D, Brescovit, E.O. Machado). The authors thank Dr Nice Shindo for the English revision of the manuscript.

References Alexander RD, Marshall DC, Cooley JR. 1997. Evolutionary perspectives on insect mating. In: Choe J, Crespi B. editors. The evolution of mating systems in insects and arachnids. Cambridge: Cambridge University Press, p.4–31. Altmann J. 1974. Observational study of behavior: sampling methods. Behavior 69:227–263. Bernardo J. 1996. The particular maternal effect of propagule size, especially egg size: patterns, models, quality of evidence and interpretations. Am Zool. 36:216–236. Brown CA, Sanford BM, Swedon RR. 2003. Clutch size and offspring size in the wolf spider Pirata sedentarius (Araneae, Lycosidae). J Arachnol. 31:285–296. Burger M, Nentwig W, Kropf C. 2003. Complex genital structures indicate cryptic female choice in a haplogyne spider (Arachnida, Araneae, Oonopidae, Gamasomorphinae). J Morphol. 255:80–93. Burger M, Jacob A, Kropf C. 2006. Copulatory behavior and web of Indicoblemma lannaianum from Thailand (Arachnida, Araneae, Tetrablemmidae). J Arachnol. 34:176–185. Costa FG, Quirici V. 2007. Cortejo e isolamento reprodutivo em aranhas. In: Gonzaga MO, Santos AJ, Japyassú HF. editors. Ecologia e Comportamento de Aranhas. Editora Interciência, Rio de Janeiro. Chapter 5 p. 89–114. Crane J. 1949. Comparative biology of salticid spiders at Rancho Grande, Venezuela. IV. An analysisof display. Zoologica 34:159–214. Del-Claro K, Marullo R, Mound LA. 1997. A new Brazilian species of Heterothrips (Insecta: Thysanoptera) co-existing with ants in the ﬂowers of Peixotoa tomentosa (Malpighiaceae). J Nat Hist. 31:1307–1312. Del-Claro K, Torezan-Silingardi HM. 2009. Insect–plant interactions: new pathways to a bet- ter comprehension of ecological communities in Neotropical Savannas. Neotrop Entomol. 38(2):159–164. Downes MF. 1987. Postembryonic development of Latrodectus hasselti Thorell (Araneae, Theridiidae). J Arachnol. 14:293–301. Eberhard WG. 1985. Sexual selection and animal genitalia. Cambridge (MA): Harvard University Press. Eberhard WG. 1994. Evidence for widespread courtship during copulation in 131 species of insects and spiders, and implications for cryptic female choice. Evolution 48:711–733. Eberhard WG, Briceño RD. 1983. Chivalry in pholcid spiders. Behav Ecol Sociobiol. 13:189–195. Eberhard WG, Briceño RD. 1985. Behavior and ecology of four species of Modisimus and Blechroscelis (Araneae, Pholcidae). Rev Arachnol. 6:29–36. Elgar MA. 1992. Sexual cannibalism in spiders and other invertebrates. In: Elgar MA, Crespi BJ. editors. Cannibalism: ecology and evolution among diverse taxa. Oxford: Oxford University Press. p.128–155. 2128 V. Stefani et al.

Ferguson GW, Brown KL, DeMarco VG. 1982. Selective basis for the evolution of variable egg and hatchling size in some iguanid lizards. Herpetologica 38:178–188. Foelix RF. 2011. Biology of spiders. 3rd ed. Oxford: Oxford University Press. 428p. Galiano ME. 1967. Ciclo biologico e desarollo de Loxosceles laeta (Nicolet 1849). Acta Zool Lill. 23:431–464. Grasshoff M. 1975. Reconstruction of an evolutionary transformation – the copulatory organs of Mangora (Arachnida. Araneae, Araneidae). Proc. 6th Int. Arachnol. Congr. 1974, Free University Amsterdam, p.12–16. Gullan PJ, Cranston PS. 1994. The insects. An outline of entomology. London: Chapman & Hall. 491p. Huber BA. 1994. Genital morphology, copulatory mechanism and reproductive biology in Psilochorus simoni (Berland, 1911). Netherl J Zool. 44:85–99. Huber BA. 1995. Copulatory mechanism in Holocnemus pluchei and Pholcus opilionoides, with notes on male cheliceral apophyses and stridulatory organs in Pholcidae (Araneae). Acta Zool 76:291–300. Huber BA. 1998. Genital mechanics in some Neotropical pholcid spiders (Araneae: Pholcidae), with implications for systematics. J Zool. 244:587–599. Huber BA. 1999. Sexual selection in pholcid spiders (Araneae, Pholcidae): artful chelicerae and forceful genitalia. J Arachnol. 27:135–141. Huber BA. 2000. New World pholcid spiders (Araneae, Pholcidae): a revision at generic level. Bull Am Mus Nat Hist. 254:1–348. Huber BA. 2003. Cladistic analysis of Malagasy pholcid spiders reveals generic level endemism: revision of Zatavua n. ge. and Paramicromerys Millot (Pholcidae: Araneae). Zool J Linn Soc. 137:261–318. Huber BA. 2005. Sexual selection research on spiders: progress and biases. Biol Rev. 80: 363–385. Huber BA, Brescovit AD, Rheims CA. 2005. Exaggerated female genitalia in two new spider species (Araneae: Pholcidae), with comments on genital evolution by female choice versus antagonistic coevolution. Insect Syst Evol. 36:285–292. Huber BA, Eberhard WG. 1997. Courtship, copulation, and genital mechanics in Physocyclus globosus (Araneae, Pholcidae). Can J Zool. 74:905–918. Kaster JL, Jakob EM. 1997. Last-male sperm priority in a haplogyne spider (Araneae: Pholcidae): correlations between female morphology and patterns of sperm usage. Ann Entomol Soc Am. 90:254–259. Levi HW. 1965. Techniques for the study of spider genitalia. Psyche 72:152–158. Machado EO, Brescovit AD, Francisco R. 2007. Three new ground dweller Mesabolivar González-Sponga, 1998 (Araneae: Pholcidae) from Blumenau, Santa Catarina, Brazil. Zootaxa 1560:55–61. Marchioro CA, Fisher ML, Silva, EM. 2005. Desenvolvimento pós-embrionário de Loxosceles intermedia Mello-Leitão, 1934, L. laeta (Nicolet, 1849) e L. gaucho Gertsch, 1967 (Araneae; Sicariidae) criadas sob condições de alimentação monoespecífica. Biotemas 18(1):93–112. Nentwig W. 1990. A zygomycetous fungus as a mortality factor in a laboratory stock of spiders. J Arachnol. 18:118–121. Nussbaum RA. 1981. Seasonal shifts in clutch-size and egg-size in the side-blotched lizard, Uta stansburiana Baird and Girard. Oecologia 49:8–13. Peretti A, Eberhard WG, Briceño, RD. 2006. Copulatory dialogue: female spiders sing during copulation to influence male genitalic movements. Anim Behav. 72:413–421. Platnick NI. 2010. The world spider catalog, version 8.0. American Museum of Natural History, [cited 2011 Feb 2]. Available from: http://research.amnh.org/entomology/spiders/catalog/ index. Roberts JA, Uetz GW. 2004. Species-specificity of chemical signals: Silk source affects discrimi- nation in a wolf spider. 2004. J Insects Behav. 17:477–491. Journal of Natural History 2129

Santos AJ, Brescovit AD, Japyassú HF. 2007. Diversidade de aranhas: sistemática, ecologia e inventários de fauna. In: Gonzaga MO, Santos AJ, Japyassú HF. editors. Ecologia e Comportamento de Aranhas. Rio de Janeiro: Editora Interciência. p.1–23. Schäfer MA, Uhl G. 2002. Determinants of paternity success in the spider Pholcus phalangioides (Pholcidae: Araneae): the role of male and female mating behaviour. Behav Ecol and Sociobiol 51:368–377. Souza JP, Araújo GM. 2005. Estrutura arbustivo/arbórea do subosque de clareiras e áreas sob dossel fechado em floresta estacional semidecidual urbana em Araguari – MG. J Biosci. 21:93–102. Stefani V, Del-Claro K. 2011. Oviposition and post-embryonic development of Aglaoctenus lagotis (Araneae: Lycosidae). Zoologia 28(5):565–570. Stefani V, Del-Claro K, Silva LA, Guimarães B, Tizo-Pedroso E. 2011. Mating behaviour and maternal care in the tropical savanna funnel-web spider Aglaoctenus lagotis Holmberg (Araneae: Lycosidae). J Nat Hist. 45:1119–1129. Suter RB, Renkes G. 1982. Linyphiid spider courtship: releaser and attractant functions of a contact sex pheromone. Anim Behav. 30:714–718. Tizo-Pedroso E, Del-Claro K. 2011. Is there division of labor in cooperative pseudoscorpions? an analysis of the behavioral repertoire of a tropical species. Ethology 117:498–507. Uetz GW, Stratton GE. 1982. Acoustic communication and reproductive isolation in spiders. In: Witt PN, Rovner JS. editors. Spider communication: mechanism and ecological significance. Princeton (NJ): Princeton University Press; p. 123–159. Uhl G. 1993. Sperm storage and repeated egg production in female Pholcus phalangioides Fuesslin (Araneae). Bull Soc Neuchatel Sci Nat. 116:245–252. Uhl G. 1994. Genital morphology and sperm storage in Pholcus phalangioides (Fuesslin. 1775) (Pholcidae; Araneae). Acta Zool. 75:1–12. Uhl G, Huber BA, Rose W. 1995. Male pedipalp morphology and copulatory mechanism in Pholcus phalangioides (Fuesslin, 1775) (Araneae, Pholcidae). Bull Br Arachnol Soc. 10:1–9. Uhl G, Schmitt S, Schäfer MA, Blanckenhorn WU. 2004. Food and sex-specific growth strategies in a spider. Evol Ecol Res. 6:523–540. Wheeler GS, McCafrey JP, Johnson JB. 1990. Developmental biology of Dictyna spp. (Araneae: Dictynidae) in the laboratory and field. Amer Mid Natur 123:124–134. Yoward PJ. 1998. Sperm competition in Pholcus phalangioides (Fuesslin, 1775) (Araneae, Pholcidae) — shorter copulations gain higher paternity reward than first copulations. In: Selden PA. editor. Proceedings of the 17th European Colloquium of Arachnology, Edinburgh 1997. Burnham Beeches, Bucks: British Arachnological Society, p.176–170.