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SHORT COMMUNICATION Anelosimus Oritoyacu, a Cloud Forest Social Spider with Only Slightly Female-Biased Primary Sex Ratios

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SHORT COMMUNICATION Anelosimus Oritoyacu, a Cloud Forest Social Spider with Only Slightly Female-Biased Primary Sex Ratios 2011. The Journal of Arachnology 39:178–182 SHORT COMMUNICATION Anelosimus oritoyacu, a cloud forest social spider with only slightly female-biased primary sex ratios Leticia Avile´s and Jessica Purcell: Department of Zoology University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia V6T 1Z4, Canada. E-mail: [email protected] Abstract. We examine the social characteristics and sex ratio of the recently described Anelosimus oritoyacu Agnarsson 2006. We find that this spider, whose nests occur on tree crowns and bushes in open fields near Baeza, Ecuador, lives in colonies that may contain from one to several thousand adult females and their progeny. It differs from most other social congeners in that it occurs at relatively high elevations (1800–1900 m) and its primary sex ratio, 2.5 females per male, is the least biased of any known social species in the genus. The low sex ratio bias may reflect a low colony turnover rather than high gene flow among colonies, as the colonies occurred in complexes that were few and far between, but appeared to be long-lived. The relatively small body size of adult females and a web that appears to allow the capture of insects from all directions, combined with individual and group foraging, may allow the formation of large colonies at an elevation where insects, albeit abundant, are for the most part small. Keywords: Ecuador, Theridiidae, cooperation, life cycle, quasisocial, subsocial The genus Anelosimus Simon 1891 is of particular interest in the Nest and web structure.—A. oritoyacu’s nests differed from those of study of spider sociality because it contains the largest number of most other social species in the genus in lacking a well differentiated non-territorial permanent-social (or quasisocial) species of any spider basal basket and extensive superior prey capture webbing, as genus (Avile´s 1997; Agnarsson 2006; Lubin & Bilde 2007). Among depicted, for instance, for A. eximius by Yip et al. (2008, fig. 1; see these, Anelosimus eximius Keyserling 1884, Anelosimus domingo Levi also photo in Avile´s et al. 2001, fig. 9). Instead, A. oritoyacu’s webs 1963, Anelosimus dubiosus Keyserling 1881, Anelosimus guacamayos consisted of a core area surrounding a piece of vegetation and prey Agnarsson 2006, Anelosimus lorenzo Levi 1979, and Anelosimus capture strands running away from the core, including inferiorly from rupununi Levi 1956 have been the subject of one to several studies it (Fig. 1), much like the nests of A. rupununi (Avile´s & Salazar 1999), (e.g., Fowler & Levi 1979; Rypstra & Tirey 1989; Rypstra 1993; a canopy species. Also as in A. rupununi, A. oritoyacu’s silk was of a Avile´s & Tufin˜o 1998; Marques et al. 1998; Avile´s & Salazar 1999; lighter texture and whiter coloration than in most other congeneric Avile´s et al. 2007; Purcell & Avile´s 2007; Yip et al. 2007). Here we species. This web structure may reflect the position of the webs on tree report on a new non-territorial permanent social Anelosimus, recently crowns, and the need to capture insects flying from the side and below described by Agnarsson (2006) as Anelosimus oritoyacu. We show that the nests. A. oritoyacu’s nests, as well as those of A. rupununi, thus although this species exhibits social organization similar to that of have characteristics that appear a response to the canopy location other social Anelosimus spiders, it presents some interesting differ- preferred by these species. The nests we observed ranged broadly in ences. Along with A. guacamayos, A. oritoyacu occurs at what size. At least two nests, but possibly as many as seven, of the 55 appears to be the elevational range limit for permanent sociality in recorded contained either a single adult female or what appeared to this genus (Avile´s et al. 2007) and its sex ratio is the least biased be the clutch of a single female. The majority of nests, however, were among known social Anelosimus (Avile´s & Maddison 1991; Avile´s et considerably larger. Several nests in the first nest complex seen in al. 2007). Here we present a brief account of the size and structure of January 2002, for instance, measured on the order of 3–4 m in A. oritoyacu’s nests and colonies, informal observations on the diameter and probably contained several thousand individuals. cooperative nature of its societies, and, given the relevance of sex Among 20 nests measured (two in 2002, three in 2004, eight in ratios as indicators of population structure (Williams 1966; Nagel- 2007, and seven in 2008, from one, two, three, and five different kerke & Sabelis 1996; Hardy 2002), estimates of its primary and colony complexes, respectively), the smallest measured 13 3 13 3 tertiary sex ratios. 23 cm and the largest, 205 3 156 3 100 cm. Location of nest complexes seen.—Over a period of six years Colony age structure.—Of the five nests that we dissected (three in (January 2002–June 2008) we located eight areas, all within a 10 km January 2002 from the initial nest complex found, one in December radius of Baeza, Ecuador (0u279S, 77u539W; 1800–1900 m elev.), that 2002 from a complex 300 m away from the former, and one in 2008 contained from 1–12 A. oritoyacu nests (median 3.5) each, for a total from a seven-nest complex found 500 m away from the original found), of 55 nest records (Table 1). When more than one nest was present four contained a mix of juvenile and/or egg sacs, subadult, and adult within these areas, nests were typically clustered within meters of one spiders, suggesting that reproduction is not strongly synchronized another in what we refer to as ‘‘nest complexes.’’ Distances between within nests (Table 2); the remaining nest contained only subadult identified nest complexes ranged from 25 m to 3.5 km. Most nests males and females (Table 2). We surveyed the age structure of seven were located on the crowns of trees or on bushes growing on open additional nests, both in June–July (2004, 2008) and in December hillsides or roadsides. The nest complexes appeared remarkably stable (2002, 2007) and found that adults and juveniles/egg sacs were present over time—at two of the sites initially discovered in 2002, nests were at both times. Taken together, these findings suggest that A. oritoyacyu still present in 2007 and 2008 (Table 1). In contrast, nest complexes in either has a short generation time and/or that its life cycle is largely species such as Anelosimus eximius rarely last more than 2–3 years (L. independent of the mildly seasonal rain patterns of the region (rainiest: Avile´s unpublished data), and those of species such as Theridion May–July; least rainy: December–February; Neill 1999). Adult males nigroannulatum Keyserling 1884 usually last less than a year (Avile´s et were seen overlapping with adult females in at least eight colonies, al. 2006) suggesting that the opportunity for intracolony mating is present. 178 AVILE´ S & PURCELL—SOCIAL SPIDER WITH FEMALE-BIASED SEX RATIO 179 Table 1.—Location of nest complexes seen and the number of seen nests they contained at the date of inspection. Location code: BZ-TN 5 Baeza-Tena Road; BZ-LA 5 Baeza-Lago Agrio Road; BZ, TN and LA 5 towns of Baeza, Tena, and Lago Agrio, respectively. Km from Baeza shown after each location code. Location code Latitude Longitude Elevation (m) Dates seen # Nests in complex BZ-TN 8.1 0.497083 77.873861 1822 Jan-02 4 Dec-02 few Dec-07 several Jun-08 several BZ-TN 8.1 +333 m 0.4955 77.876306 1881 Dec-02 2 BZ-TN 1.0 0.46322 77.87662 1866 Dec-02 12 Dec-07 4 Jun-08 3 BZ town Jun-04 1 BZ-TN 4.5 0.4729 77.86819 1848 Dec-07 1 Jun-08 2 BZ-LA 2.4 0.45157 77.88392 1818 Jul-04 2 BZ-LA 2.6 0.451395 77.88954 1823 Jun-08 1 BZ-LA 3.0 0.45152 77.88399 1842 Dec-07 8 Jun-08 7 BZ-LA 3.0 + 25 m 0.45152 77.88399 1842 Dec-07 4 Jun-08 4 Clutch size and sex ratio.—In 2004, we collected egg sacs from a Gunnarsson & Andersson 1992 for a solitary species with biased sex single large colony and used the method described by Avile´s & ratios) and from the 10:1 sex ratio found in other social Anelosimus Maddison (1991) to sex the embryos they contained. Egg sacs were species (e.g., A. eximius and A. domingo, Avile´s & Maddison 1991; A. off-white in color, averaged 3.9 6 1.2 mm in diameter (n 5 5), and guacamayos, Avile´s et al. 2007). The tertiary sex ratio of adult and contained between 16 and 46 eggs (n 5 11, median 5 37, mean 5 34, subadult spiders from nests collected in 2002 and 2008 similarly SE 5 3.13). In cytological spreads of individual embryos we counted showed a bias of between two and five females to one male (Table 1). the number of chromosomes contained in at least three dividing cells Spider size and instars.—We measured the length of the tibia + to determine whether the individual was male or female (n 5 130, four patella on leg pair 1 (TP1) and leg pair 2 (TP2), as well as the sternum egg sacs, Table 2). As in other species in the genus, males had 22 and length (SL) and weight for a haphazard sample of subadult and adult females, 24 chromosomes (20 autosomes plus two sex chromosomes spiders collected from one nest belonging to the original complex seen for males, and four sex chromosomes for females).
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