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Volume 2, Chapter 7-2: Arthropods: Arachinda-Spider Biology

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Volume 2, Chapter 7-2: Arthropods: Arachinda-Spider Biology Glime, J. M. 2017. Arthropods: Arachnida – Spider Biology. Chapt. 7-2. In: Glime, J. M. Bryophyte Ecology. Volume 2. Bryological 7-2-1 Interaction. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 18 July 2020 and available at <http://digitalcommons.mtu.edu/bryophyte-ecology2/>. CHAPTER 7-2 ARTHROPODS: ARACHNIDA – SPIDER BIOLOGY TABLE OF CONTENTS Subphylum Chelicerata ....................................................................................................................................... 7-2-2 Class Arachnida .................................................................................................................................................. 7-2-3 Arachnid Trapping Limitations .................................................................................................................... 7-2-3 Order Araneae – Spiders ..................................................................................................................................... 7-2-4 Spider Biology ............................................................................................................................................. 7-2-4 Growth Forms and Life Forms ..................................................................................................................... 7-2-4 Bryophytes as Cover .................................................................................................................................... 7-2-5 Trampling ..................................................................................................................................................... 7-2-7 Abundance, Richness, and Specificity ......................................................................................................... 7-2-7 Moisture Relationships ................................................................................................................................ 7-2-9 Importance of Temperature ........................................................................................................................ 7-2-12 Food Sources .............................................................................................................................................. 7-2-13 Reproduction .............................................................................................................................................. 7-2-14 Nests and Webs .......................................................................................................................................... 7-2-16 Dormant Stages .......................................................................................................................................... 7-2-19 Overwintering ............................................................................................................................................ 7-2-20 Spider Guilds ............................................................................................................................................. 7-2-20 Adaptations to Bryophytes ......................................................................................................................... 7-2-21 Anapidae ............................................................................................................................................. 7-2-21 Clubionidae (Sac or Tube Spiders) ..................................................................................................... 7-2-22 Gnaphosidae (Ground Spiders) ........................................................................................................... 7-2-22 Linyphiidae (Sheet Spiders) ................................................................................................................ 7-2-22 Lycosidae (Wolf Spiders) ................................................................................................................... 7-2-24 Symphotognathidae and Micropholcommatidae ................................................................................. 7-2-24 Theridiidae (Tangle-web Spiders, Cobweb Spiders, and Comb-footed Spiders) ................................ 7-2-24 Summary ........................................................................................................................................................... 7-2-25 Acknowledgments ............................................................................................................................................. 7-2-25 Literature Cited ................................................................................................................................................. 7-2-25 7-2-2 Chapter 7-2: Arthropods: Arachnida – Spider Biology CHAPTER 7-2 ARTHROPODS: ARACHNIDA – SPIDER BIOLOGY Figure 1. Arctosa cf. alpigena female on moss, showing disruptive coloration that makes it more difficult to see. It has been reported from mosses in more than one study (Harvey et al. 2002; Almquist 2005). Photo by Walter Pfliegler, with permission. SUBPHYLUM CHELICERATA The subphylum Chelicerata includes the spiders and Following the concept of a niche, bryophytes can mites, both having members associated with bryophytes. provide a number of important "resources" for arachnids. Both spiders and mites are in the class Arachnida, along The most obvious of these are shelter and protection. With with scorpions, harvestmen, ticks, and Solifugae. The disruptive coloration on their backs, spiders and other Chelicerata are characterized by four pairs of walking small arachnids can hide among the bryophytes undetected legs, a pair of chelicerae, and a pair of pedipalps. Although by would-be predators such as birds. This shelter may the arachnids are not as small as many of the organisms in provide a safe site when an arachnid is being chased or preceding chapters, many are small enough that the provide a protected niche for an egg case during bryophytes still provide sufficient space for many of these incubation. The protection also extends to anchorage and taxa to navigate easily among the stems and leaves. Hence, shelter from wind, diffusion of raindrops (avoiding the we should expect to find the bryophytes to be a suitable impact of a free-fallen drop), temperature buffering, and habitat for a number of these. retention of humidity. Further possibilities include having Chapter 7-2: Arthropods: Arachnida – Spider Biology 7-2-3 a place to lie in wait for a walking meal to meander by, or is not necessary and may introduce a bias if the study is perhaps even eating the bryophyte itself, a menu item that quantitative. is poorly documented (and unlikely) for spiders. But bryophytes may also extend their benefits to those arachnids not living among the branches. Bryophytes help to keep the soil beneath humid, soft, and pliable for longer periods than that experienced by bare soil. They permit an arachnid to emerge from a burrow and look around while remaining hidden beneath a canopy of loose bryophytes. Even those arachnids traversing the surface of bryophytes may benefit from the disruptive coloring of mosses that make the disruptive colors of arachnid backs less conspicuous. Or they may simply add a place where humidity is greater, helping arachnids to travel greater distances before risk of drying. And who knows if these arachnids might take advantage of the early morning dew captured by bryophyte leaves to gain a drink of water. With all these possibilities, we would expect some arthropods to have distinct adaptations to that bryophytic habitat. Indeed some do, but I feel certain many stories remain to be discovered. Class Arachnida The arachnids include the spiders (order Araneae), mites (subclass Acarina), ticks (subclass Acarina), and harvestman or daddy-long-legs (order Opiliones). These are creatures that somewhat resemble insects, but as adults they have eight legs. They have one or two main body regions, not three as in insects. Among these, the mites are Figure 2. Pitfall trap with cereal bowl holding alcohol. The fairly common residents in moss clones. Although the bait will bring the organisms to the trap, but most will fall into the other arachnids are not very common among bryophytes, alcohol before reaching the wire that gives them access. Drawing there are, nevertheless, some interesting stories about all of from USDA website. these inhabitant groups. Arachnid Trapping Limitations Little quantitative work exists for any arachnids except that for the moss-dwelling mites. One limitation that might suggest that bryophytes are unimportant is the typical sampling method used for forest floor arthropods, including arachnids. Pitfall traps are typically used for those arthropods that are active above the surface during some part of the 24-hour cycle (Curtis 1980). But if arthropods spend most of their time within the bryophyte mat rather than on the surface, they are not likely to fall into such traps. Curtis found that responses of spider species to four pitfall trapping methods differed, causing distortions in the community species frequency curves. Hence, we should expect even greater differences among a wider range of methods. For example, Komposch (2000) studied the spiders in wetlands of Austria using pitfall traps, light traps, soil sifters, and hand collections. As will be seen in studies cited in this chapter, this broader set of methods gets better representation of groups like the Linyphiidae, a very species-rich
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