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 - UC Natural ...
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

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Untangling ecotone diversity hypotheses: Spider communities within a forest-meadow interface - UC Natural ...
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.

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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

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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.

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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

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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.
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