Effect of Kleptoparasitic Ants on the Foraging Behavior of a Social Spider (Stegodyphus Sarasinorum Karsch, 1891)
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Zoological Studies 58: 3 (2019) doi:10.6620/ZS.2019.58-03 Open Access Effect of Kleptoparasitic Ants on the Foraging Behavior of a Social Spider (Stegodyphus sarasinorum Karsch, 1891) Ovatt Mohanan Drisya-Mohan*, Pallath Kavyamol, and Ambalaparambil Vasu Sudhikumar Centre for Animal Taxonomy and Ecology, Department of Zoology, Christ College, Irinjalakuda, Kerala, India. *Correspondence E-mail: [email protected] (Drisya-Mohan). E-mail (other authors): [email protected] (Kavyamol); [email protected] (Sudhikumar) Received 23 February 2018 / Accepted 23 December 2018 / Published 25 February 2019 Communicated by John Wang The term kleptoparasitism is used to describe the stealing of nest material or prey of one animal by another. Foraging and food handling behaviors of social spiders increase the vulnerability to klepto- parasitism. Kleptoparasites of the social spider Stegodyphus sarasinorum Karsch 1891 were identified based on the observations done in the field. Four species of spiders and two species of ants were observed as kleptoparasites and collected from the nest and webs of this social spider. The ants were found to be the most dominant among them. The influence of a facultative kleptoparasitic ant, Oecophylla smaragdina on the foraging behavior of S. sarasinorum was studied in laboratory conditions. The experiments suggested that the web building behavior of S. sarasinorum was influenced by the exposure to ants. However, exposure to ants caused no significant effect in the prey capture, handling time of prey and prey ingestion behaviors of the spider. Key words: Social spider, Ecology, Kleptoparasites, Ants, Prey capturing behavior. BACKGROUND 2000). Kleptoparasites depend on high-quality food and it is available to them due to prolonged handling Kleptoparasitism is a type of feeding behavior, (Giraldeau and Caraco 2000). Lastly, the more in which one animal steals the food or prey captured conspicuous the hunting and food handling behaviors, by another animal. Kleptoparasites feed on prey which the easier it is for the potential thieves to identify the could not be obtained by themselves without time and opportunities for exploitation. effort. There are chances for the kleptoparasites to be Among spiders, web-building spiders are frequent injured by the host organism (one being stolen from) targets of kleptoparasites (Vollrath 1987). Web-building when the latter defends its prey. Kleptoparasitic behavior spiders are relatively sedentary, sit-and-wait type of is seen in a diverse array of taxa including marine predators and capture prey that is often larger than invertebrates (Zamora and Gomez 1996; Morrissette themselves and that takes time to consume. They usually and Himmelman 2000), spiders and insects (Higgins store many prey items in the web or nest for future and Buskirk 1988; Field 1992), birds (Brockmann and consumption (Champion de Crespigny et al. 2001). Barnard 1979), large carnivores (Packer and Ruttan Social spiders built large sized web and capture large 1988) and primates (Di Bitetti and Janson 2001). sized prey cooperatively. Due to these behaviors, social Susceptibility to kleptoparasitism is affected by spiders are also an attractive resource for kleptoparasites. various characteristics of the organism’s behaviors A few species of spiders act as kleptoparasites include foraging and food-handling behaviors along with insects like some scorpionflies (Thornhill (Brockman and Barnard 1979; Giraldeau and Caraco 1975) and ants (Henschel and Lubin 1992; Pasquet et Citation: Drisya-Mohan OM, Kavyamol P, Sudhikumar AV. 2019. Effect of kleptoparasitic ants on the foraging behavior of a social spider (Stegodyphus sarasinorum Karsch, 1891). Zool Stud 58:3. doi:10.6620/ZS.2019.58-03. © 2019 Academia Sinica, Taiwan 1 Zoological Studies 58: 3 (2019) page 2 of 11 al. 2007). The spiders which regularly steal food from sp., Tamarindus indica, Artocarpus heterophyllus and other species of spiders, i.e., kleptoparasitic spiders, Nigeria grass (Pennisetum pedicellatum). are known to occur in five families. As ants can rapidly recruit a large number of individuals and by patrolling, Kleptoparasites of S. sarasinorum ants are likely to be the most prominent kleptoparasites (Henschel 1998; Hölldobler and Wilson 1983). The nests of the spiders were examined in Moreover, ants can also become predators, particularly its habitat at different locations of Christ College in spider broods (Schneider and Lubin 1997). If campus; covering both winter and summer seasons. S. kleptoparasitism shown by ants is a sufficiently serious sarasinorum thrives on shrubs and the lower branches risk to spiders, adaptive responses by spiders should be of tall plants. Kleptoparasites were collected by hand as observed to lower the risk. follows. The specimens were collected by leading them Although most of the studies have dealt with the into glass tubes containing alcohol with the help of a diversity, evolution, and behavior of kleptoparasites, brush dipped in alcohol. The collected spiders and non- the knowledge about its effect on the web-building and spider kleptoparasites were preserved in 70% alcohol. foraging behavior of social spiders are insufficient, The collected specimens were then taken to the due to lack of studies except a few (Cangialosi 1990; laboratory and examined under Magnus MSZ TR stereo Leborgne et al. 2011). A pioneering study conducted microscope for taxonomic identification. Kleptoparasitic by Leborgne et al. (2011), showed that in Eresid sub- spiders were identified with the help of available social spider, Stegodyphus lineatus kleptoparasites literature (Sebastian and Peter 2009; WSC 2018) and negatively influenced the web building behavior kleptoparasitic ants were identified using the catalogue and was independent of the success in the spider’s of ants (Bolton et al. 2007) and AntWeb online database previous foraging. According to the accepted theory (2017, http://www.antweb.org). The specimens are in spiders, non-territorial permanent-sociality is housed in the reference collection maintained at the derived evolutionarily from a sub-social state and is Centre for Animal Taxonomy and Ecology, Department evolutionarily restricted by genetic or ecological factors of Zoology, Christ College, Irinjalakuda, Kerala, India (Aviles 1997). Since there is a difference between social (ID – CATE0306). and sub-social spiders a detailed examination was done on the influence of kleptoparasites on web building Effect of exposure of ants on web building as well as foraging behavior of social spiders. So this behavior of S. sarasinorum study was conducted on the social spider, Stegodyphus sarasinorum, Karsh 1891 (Araneae: Eresidae) which S. sarasinorum was collected from Christ College is commonly known as the Indian cooperative spider. campus. Adult and subadult females were used for This spider exhibits communal predation and feeding, this study. Spiders (two groups, 30 spiders each) were where individual spiders live in large cooperatively built introduced into the 45 × 30 × 30 cm enclosures (Fig. 2B) colonies with a nest or retreat made of silk, and woven made up of nylon mesh and wooden reapers (Wooden using twigs, leaves, food carcasses and a sheet web for sticks used for making the frame of the enclosure). prey capture. Twigs and wet cotton were placed in the enclosure The present study identifies the kleptoparasites of which facilitates web building and maintains moisture S. sarasinorum in the natural colonies at Christ College, content in the enclosure respectively. Grasshoppers Irinjalakuda, Kerala, India. The influence of facultative (Chortopaga viridifasciata (0.22 ± 0.008 g), Oxya hyla kleptoparasitic ants on web-building and prey capturing intricata (0.13 ± 0.007 g), and Neorthacris simulans behaviors of S. sarasinorum was also studied. (0.28 ± 0.014 g)) were collected from the field and grown in an artificial terrarium (60 × 30 × 30 cm) and they were used as prey for the spiders. Two similar- MATERIALS AND METHODS sized grasshoppers were given to each group daily in order to homogenize the feeding status (We observed Study Area the influence of feeding status and exposure of ants). After 5 days, the first group received two grasshoppers The study site was in the Christ College campus (“fed”) while the second group received no prey (10°21'N 76°12'E), located in Irinjalakuda town of (“fasted”) over the next 4 days. Thrissur district, Kerala, India. Colonies of Stegodyphus These two groups were again subdivided into sarasinorum are widely distributed in the campus (Fig. two sub-groups (15 spiders each). The facultative 1) and are found on the branches of plants such as kleptoparasitic ants, Oecophylla smaragdina (K) Eugenia uniflora (Fig. 2A), Acacia disparrima, Acacia were collected from Christ College campus and 10 © 2019 Academia Sinica, Taiwan Zoological Studies 58: 3 (2019) page 3 of 11 individuals each were exposed into one of the fasted and area of the three replicates of the experiment were fed sub-groups. Some ants moved on the web and some measured on 4 consecutive days. others moved on the enclosure (Fig. 2C). After 1 hour, ants were removed and all webs both in the K exposed Statistical analyses and K unexposed groups were destroyed manually. The experimental setup with spiders alone was kept Welch Two independent Sample t-test was undisturbed for the next four days except during daily performed to compare the means of the web size built web size measurements. The day 1 measurement was on the