The Effects of Food Availability of Social Behavior in the Spider, Nephila clavipes

Nephila clavipes

Acknowledgements:
Wendy Welshans & the Forman School Rainforest Project

Kimberly Layman
Advisor: Dr. Karen Cangialosi

Introduction
While most spider species are aggressive and territorial towards their conspecifics, a few species are known to exhibit some level of sociality (Rypstra 1985, Buskirk 1975). Studying the situations under which spiders that are normally solitary, but may congregate can help shed light on the evolution of sociality. Nephila clavipes, otherwise known as the Golden Orb Weaver spider, has been known to exist in colonies as well as solitarily. Their group formations could be due to food abundance in a specific location, however this species has also exhibited cannibalistic and territorial behavior in situations where the spiders other food sources are not in abundance (Rypstra 1985). In my previous observations of these spiders, I did not witness aggressive behavior and was intrigued by these claims of cannibalism within groups of Nephila. Therefore, the main objective of this study was to determine whether Nephila clavipes would acquire some benefits from living in social aggregations.

Discussion
The fact that colonial spiders captured more prey could be due to several factors. One possibility was the location of the colony. Because the colony was on the same side of the building as the kitchen, it is possible that the food smell emitted from the kitchen caused a greater abundance of prey such as flies. Several orb-weaving spiders have been found to aggregate around areas of high food concentration (Uetz & Cangialosi 1986). It has been previously observed that Nephila clavipes have also been known to exist in large groups when food sources are adequate (Rypstra 1985). Groups may form because of the abundance of food in an area, but the amount of competition is likely to affect the prey capture rate of individuals (Rypstra 1985). It is also possible that the higher prey capture in the colony was due to the ricochet effect, meaning when prey are captured after bouncing off several webs in succession (Spiller 1992, Uetz 1989). The spiders found in the colony may have been more successful due to that fact that their structure was more dense and prey were less likely to be able to escape because of the higher density of silk strands in the interconnected webs. It is thought that Nephila clavipes are territorial and have cannibalistic tendencies, causing their social groupings to be an ineffective method of living (Lewis 1996). However, my data shows that the colony was more successful in capturing prey than the spiders found in a solitary environment. Solitary spiders captured a larger range of size classes of insect prey than colonial spiders. This was unexpected due to the fact that a larger amount of webbing from the colony should have intercepted more types of prey. However, this could be due to the location of the solitary spiders, which were found near more dense vegetation, which could have provided a more favorable habitat for a larger variety of insects. The colonial spiders were located farther from this type of habitat. In future studies, I would attempt to increase the amount of observation time as much as possible, and also make a greater attempt to identify the prey captured to the species level.

Results
Nephila clavipes found in the colony caught more prey per spider per hour than
the solitary spiders on average (Figure 1). The social spiders were more successful in catching prey in the small size category. Conversely, solitary spiders caught the largest prey, but also had the highest number of very small prey per individual per hour (Figure 2). The prey captured which I was able to identify included gnats (almost the entire extra small classification was gnats) four black flies and three flying red ants (both ranged in size from small-medium). The total amount of prey captured for each size class, except extra large prey, was greater for the colonial spiders (Figure 3). However the rate of capture per individual for extra small prey was greater for solitary spiders (Figure 2). Solitary spiders also exhibited more variation in their prey size than did colonial spiders (Figure 3). From the data collected, there seemed to be no evident correlation between time, temperature, or humidity at the time of observation and the prey captured. What I also observed during this study was that not all prey that encountered the web was consumed by the spider. The surviving insects that I most often observed were flying ants of an unknown species. Such subjects were examined by the spider and upon investigation allowed to either remove themselves from the web or remained untouched. I also observed that spiders found in solitary conditions were more likely to wrap and save excess captured prey, whereas colonial spiders consumed the prey immediately. The overall prey capture rates (insects per spider) of those in the colony were higher than those of solitary spiders.
Figure 1. Prey Capture Rate Of Solitary and Colonial Spiders
0.44

Methods
This study was performed on the Rara Avis and El Plastico reserves, just outside of the city of Las Horquetes, Costa Rica from March 7 to March 15, 2009. The species I studied, Nephila clavipes, otherwise known as the Golden Orb Weaver spider is found from southern Brazil to Florida and typically ranges from 22mm40mm in body length (Lewis 1996). In order to observe the differences in prey capture rates of colonial spiders vs. solitary spiders, I identified and marked colonial and solitary spider webs of Nephila clavipes, or more commonly the Golden Orb Weaver Spider. Colonial spiders are those found in groups, where individuals are in close proximity to one another and have interconnected webs. Solitary spiders are those isolated from other spiders of the same species in individual webs. I marked one site where I could observe one solitary spider on the Rara Avis reserve in Costa Rica, and the second at El Plastico reserve (about three miles from Rara Avis) where I observed a colony. Also at El Plastico, I marked three separate individuals on the frame of a farmhouse for a total of four solitary individuals and one colony of twenty-two individuals. Observations of natural prey capture were conducted over a period of 19 total hours in this seven-day study. For both the colony of spiders as well as the solitary individuals, I observed the webs for 30 minutes to 2 hours. Each time an insect prey item encountered the web naturally, I recorded the time as well as the size and taxonomic identification of the prey (if possible). Prey were assigned a size category of very small (<2 cm), small (2-3 cm), medium (3-4 cm), large (4-5 cm) and extra large (>5 cm) according to their body length I also recorded detailed descriptions of the behavior. The data were analyzed to determine the prey capture rate (number of insects per hour per spider). In this way, the rates between solitary prey capture and the prey capture of a group could be compared with regard to which social condition seemed to be more effective when collecting prey. The date, time, humidity, temperature, web orientation and number of spiders present at the time of observation were also noted.

Prey Capture Rate (prey per individual per hour)

0.42 0.4 0.38 0.36 0.34 0.32 Solitary Colonial Spider Social Classification

Figure 2. Capture Rate by Insect Prey Size Class

Figure 3. Total Number of Prey Captured per Size Class

References
Buskirk, R. E. 1975. Coloniality, activity patterns and feeding in a tropical orb-weaving spider. Ecology, 56:13141328.
Colonial Solitary

Number of Prey Captured (per individual

0.45 0.4 0.35 0.3 0.25 0.2 0.15 0.1 0.05 0 XS Small Medium Size of Prey Large XL Colonial Solitary

Number of Prey Captured

14 12 10 8 6 4 2 0 XS Small Medium Size of Prey Large XL

Rypstra, A. L. 1985. Aggregations of Nephila Clavipes (L.) (Araneae, Araneidae) in relation to prey availability. J. Archnol., 13:71-78. Spiller, David A. 1992. Relationship Between Prey Consumption and Colony Size in an Orb Spider. Oecologia 90:457-66. Uetz, George W. 1989. The Ricochet Effect and Prey Capture in Colonial Spiders. Oecologia 81:154-59. Uetz, G. W. and K. R. Cangialosi. 1986. Genetic differences in social behavior and spacing in populations of Metepeira spinipes, a communal-territorial orb weaver (Araneae, Araneidae). J. Arachnol., 14:159-173).