Untangling ecotone diversity hypotheses: Spider communities within a forest-meadow interface - UC Natural ...
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Untangling ecotone diversity hypotheses: Spider communities within a forest-meadow interface Melissa Grim1, Jagger Joyner1, Christina Nakhla2, Bautista Tobias3 1 University of California, Santa Barbara; 2University of California, Riverside; 3 University of California, San Diego ABSTRACT Edge effects are important and understudied in ecology. Spiders exhibit a diverse array of predatory strategies which depend on habitat structure and abiotic environmental factors, making them an important study taxon in edge habitats. In this study, we evaluated the relationship between spider family richness, abundance, and diversity with a forest-meadow interface in the Coast Range of California. We expected to observe a difference in response to edge effects depending on the functional guild of spiders, active hunters or web-weavers. We found that web-weavers were more common in forest interiors while hunter abundance did not differ by habitat. However, no difference in diversity was found between habitats. Overall, this study shows that spiders do respond to edge effects and suggests that future research into edge effects should look explicitly at the underlying environmental factors contributing to these patterns. Keywords: spiders, ecotones, edge effect, biodiversity INTRODUCTION forest habitats may create intermediate environments or microhabitats which Biodiversity plays a crucial role in increase the biodiversity and abundance of maintaining ecosystem dynamics and has specialist taxa (Meiners et al. 2000, Stašiov been a focus of research in recent years. et al. 2021). Biodiversity can vary greatly between While it has been hypothesized that edges habitats, making the transitional edge maintain higher biodiversity than adjacent between habitat types an area of interest. habitats (Senft 2009), this may not be the This buffer zone, known as the ecotone, is case in forest-grassland interfaces (Turton subject to changes that impact the habitat and Duff 1992, Harper 1995, Baldissera et as a whole. Edge effects describe the shift in al. 2004) This lack of consensus suggests species composition, vegetative structure, that further study of forest edge effects and and ecosystem processes across an underlying environmental factors would be ecotone, and can impact the abundance of value. and distribution of species (Murcia 1995). The ecotone or edge between open and | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 1/6
Spiders in particular present an METHODS interesting study taxon given their roles as invertebrate predators. Spiders vary in their 2. 1 Study Site predatory strategies and preference of habitat structure. They are highly abundant Research was conducted from May 6–9, throughout most ecosystems, making them 2021, at Angelo Coast Range Reserve in ideal for examining the effects of habitat Mendocino County, California. The reserve transitions. Additionally, spiders have is located in the upper basin of the South strong potential as bioindicators (Pearce Fork Eel River. Angelo has a Mediterranean and Venier 2006). climate, with average summer highs of 30°C Spider families can be separated into two and winters below freezing, and average functional guilds based on their predation annual precipitation of 215.6 cm. In this strategy: sit-and-wait (web-weaving) or study we looked at the edges between four active hunting. Web-weaving spiders meadows and adjacent mixed coniferous depend strongly on habitat structure for forests. Meadows primarily consist of scaffolding to build webs (Robinson 1981, European grasses and forbs, with adjacent Uetz 1991, Rodrigues et al. 2014), while forests of Douglas Fir, manzanita, madrone, active hunters do not and instead are and oak species. the four meadows used in limited more by temperature and moisture this study are Lower Walker Meadow, (Downie et al. 1996). These functional Walker Meadow, White House Meadow, guilds depend on different environmental and South Meadow. factors, but both occupy a similar place in 2.2 Data Collection the food web. How the two guilds respond differently to biotic and abiotic factors may Three belt transects (30 m x 2 m) were help to explain why previous studies have placed parallel to the east edge of each reached contradictory conclusions when meadow with a 30-meter buffer between looking at ecotone diversity. transects (Fig. 1). The edge was defined by The goal of this study was to evaluate the furthest tree within the meadow and differences in family richness, diversity, and was used when measuring the 30-meter abundance of spider communities along buffers between the meadow, edge, and four forest-meadow edges in Angelo Coast forest transects. Where terrain was Range Reserve. We hypothesized that web inaccessible, transects were shifted north to building spiders would be most abundant in the nearest accessible location. forest interiors where there would be more vegetative scaffolding, while family diversity would be greatest in the intermediate edge habitat. Conversely, since hunting spiders are less dependent on scaffolding, we hypothesized that both abundance and diversity would be greatest in the edge habitat. | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 2/6
on abundance of live spiders, webs, and individual families at a transect. RESULTS Overall, 15 families were identified between meadow, edge, and forest habitats. There was no difference in family richness between habitats (n = 4, F = 0.512, p = 0.623). The combined abundance of live Figure 1: Transect placement in South Meadow. The spiders and webs was greatest in forests, first tree protruding into the meadow was used to with no difference in abundance between define edge. A 30 m x 2 m belt transect was set up within each habitat type, with a 30 m buffer meadows and edges (n = 4, F = 12.758, p = between transects. 0.007, Fig. 2). Web-weaver abundance was greater in forests than meadows and edges, Living spiders and individual webs were which were not significantly different (n = 4, recorded along each transect, identified to F = 15.445, p = 0.004, Fig. 2). No difference family level, and sorted into web-building or was found in the abundance of hunting hunting guilds. We limited observation time families between habitat types (n = 4, F = to 45 minutes per transect. Additionally, 2.956, p = 0.128, Fig. 2). Scaffolding index, vegetation scaffolding index, which substrate type, and vegetative cover were describes the amount of structural found to have no effect on family richness vegetation available to web-builders, was or abundance. recorded at two random points on each transect based on methods by Miyashita et al. (2004). Substrate and vegetative cover were recorded in a 0.25 m2 quadrat at 0 m, 15 m, and 30 m, alternating sides along the transect. 2.3 Statistical Analysis All data analysis was completed using RStudio version 1.3.1093 and JMP version 15. We used ANOVA tests were performed to assess differences in live spider, spider Figure 2: Abundance of spider guilds. Abundance of web, and total spider abundance between web-weaver and hunter guilds the in meadow, edge, habitats, using site as a random effect. and forest habitats. Web weavers were significantly Shannon diversity index was calculated for more abundant in forests (n = 4, F = 12.758, p = each transect and compared across 0.007), but no difference in hunter abundance was habitats. We also tested the predictive found between habitat types (n = 4, F = 2.956, p = 0.128). power of vegetative index and quadrat data | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 3/6
The Shannon diversity index showed a marginal trend, with meadows more diverse than edge and forest (n = 4, F = 3.958, p = 0.080). The dominant family in meadow and edge habitats was Theridiidae (tangle-web weavers), and the dominant family in forests was Linyphiidae (sheet-web weavers) (Fig. 3, Table 1). Figure 3: Rank abundance of the meadow, edge and forest habitats. Abundance of spider webs and live hunting spiders was averaged for all transects and ranked from most to least abundant. Individual family ranks can be found in Table 1. Table 1: Average abundance of spider families. Ranks indicate most abundant family within meadow, edge, and forest habitats, and correspond to Figure 3. | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 4/6
DISCUSSION interior forest and meadow provide ideal scaffolding for both of these families, but This study investigated the differences in Agelenidae is better able to take advantage spider family composition between of the tall grasses and thatch found in meadow, edge, and forest habitats. We meadows. This complicates the hypothesis hypothesized that hunting spiders would be that ecotones have increased diversity by more abundant and diverse in the edge providing an intermediate habitat (Meiners habitat. We found that there was no et al. 2000). In the case that organisms have difference in hunting spider abundance or distinct structural habitat preferences, the diversity across all three habitats. In other ecotone may not provide the necessary words, edge effect did not impact hunting resources as demonstrated by two different guild spiders. web types. For web-weaving spiders, we expected to Future studies could improve several see the greatest abundance of families aspects of our experimental design and within the forest interior where we methods. We recommend taking more predicted vegetation scaffolding index to be thorough measurements of substrate and greatest. While we did find web-building vegetative ground cover in order to better families to be most abundant in the forest quantify the impacts of these interior, we found no difference in environmental factors on spider vegetation scaffolding availability between communities. Additionally, we would habitats. However, our measurements of recommend taking note of abiotic factors scaffolding did not match our observations, such as temperature and relative humidity, which indicates that our methods may have both of which influence microclimates. been ineffective for the study system. Previous work has found that microclimates Additionally, we expected family diversity and vegetation cover alter the activity of to be the greatest in the edge. Instead, we spiders, including hunting and weaving found that total family diversity was similar (Downie et. al 1996). The observation of across all habitats, but marginally greater in microclimate differences within the three meadows than both edge and forest habitats in our study may have been useful habitats. in understanding the effects of abiotic When analyzing family level abundance factors on spider community composition across habitats we found that Linyphiidae and potential reasons for an increased were the most abundant family within the diversity in the ecotone. Overall, forest interior (Table 1). Agelenidae had a understanding the interaction between similar abundance in both the interior abiotic and biotic factors within forest and meadow. This difference may be microhabitats is important when due to the specific requirements for each considering the impacts of ecotones on taxa family; for example, Linyphiidae require a such as spiders. When analyzing edge clearing to establish a sheet web, whereas effects, future studies should more broadly Agelenidae construct more dense and consider the impacts of microclimates in robust funnel webs closer to the ground and across ecotones. within dense vegetation (Rojas 2011). The | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 5/6
ACKNOWLEDGMENTS Pearce, J. L., and L. A. Venier. 2006. The use of ground beetles (Coleoptera: Carabidae) and We would like to acknowledge our spiders (Araneae) as bioindicators of sustainable forest management: a review. Ecological amazing teaching assistant, Robert Blenk indicators 6:780–793. for all his guidance throughout the course. We would also like to thank Mary Power, Robinson, J. V. 1981. The effect of architectural Blake Suttle, and Kirsten Hill for their advice variation in habitat on a spider community: An and guiding words. This work was experimental field study. Ecology 62:73–80. performed at the University of California’s Rodrigues, E. N. L., M. D. S. Mendonça, and L. E. Angelo Coast Range Reserve, Costa-Schmidt. 2014. Spider diversity responds doi:10.21973/N3R94R. strongly to edge effects but weakly to vegetation structure in riparian forests of Southern Brazil. REFERENCES Arthropod-Plant Interactions 8:123–133. Baldissera, R., G. Ganade, and S. B. Fontoura. 2004. Rojas, A. 2011. Sheet-web construction by Web spider community response along an edge Melpomene sp. (Araneae: Agelenidae). The between pasture and Araucaria forest. Biological Journal of Arachnology 39:189–193. Conservation 118:403–409. Senft, A. R. 2009. Species diversity patterns at Downie, I. S., J. C. Coulson, and J. E. Butterfield. ecotones. Thesis, University of North Carolina at 1996. Distribution and dynamics of surface- Chapel Hill. dwelling spiders across a pasture-plantation ecotone. Ecography 19:29–40. Stašiov, S., V. Kubovčík, M. Čiliak, A. Diviaková, I. Lukáčik, and M. Dovciak. 2021. Harvestmen Greenstone, M. H. 1984. Determinants of web spider (Opiliones) community structure varies across species diversity: Vegetation structural diversity forest-meadow ecotones in a biodiverse karst vs. prey availability. Oecologia 62:299–304. region. Biodiversity and Conservation 30:1101– 1117. Harper, K. A. 1995. Effect of expanding clones of Gaylussacia baccata (black huckleberry) on Suttle, K. B., M. A. Thomsen, and M. E. Power. 2007. species composition in sandplain grassland on Species interactions reverse grassland responses Nantucket Island, Massachusetts. Bulletin of the to changing climate. Science 315:640–642. Torrey Botanical Club 122:124. Turton, S. M., and G. A. Duff. 1992. Light Miyashita, T., M. Takada, and A. Shimazaki. 2004. environments and floristic composition across an Indirect effects of herbivory by deer reduce open forest-rainforest boundary in northeastern abundance and species richness of web spiders. Queensland. Austral Ecology 17:415–423. Ecoscience 11:74–79. Uetz, G. 1991. Habitat structure and spider foraging. Meiners, S. J., S. N. Handel, and S. T. A. Pickett. 2000. In: Bell S.S., McCoy E.D, Mushinsky H.R. (eds.) Tree seedling establishment under insect Habitat structure. Population and Community herbivory: edge effects and inter-annual variation. Biology Series, Vol. 8, pp. 325–348. Springer, New Plant Ecology 151:161–170. York, New York. Murcia, C. 1995. Edge effects in fragmented forests: implications for conservation. Trends in Ecology & Evolution 10:58–62. | CEC Research https://doi.org/10.21973/N3D951 Spring 2021 Vol. 5, Issue 3 6/6
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