2 PRAIRIE SPIDERS VOL. 120(2) Passive Re-colonization of the Spider Assemblage on an Ohio Restored Tall Grass Prairie Compared to Nearby Remnant Prairies and Old Fields RICHARD A. BRADLEY 1, Department of Evolution, Ecology, and Organismal Biology, The Ohio State University at Marion, Marion, OH, USA; SHAUNA L. PRICE, Department of Science and Education, Integrative Research Center, The Field Museum of Natural History, Chicago, IL, USA; WILLIAM L. HICKMAN, Cardington, OH, USA; ROBERT A. KLIPS, Department of Evolution, Ecology, and Organismal Biology, The Ohio State University at Marion, Marion, OH, USA. ABSTRACT. The reconstructed prairie on the Marion Campus of The Ohio State University was established in 1977. Since then restoration has focused on plants. Animals on the site have recolonized without active management. Spider assemblages were sampled in 2000 and compared to those sampled at the time on 2 remnant prairies and 2 old fields. Pitfall traps and sweep nets were used for sampling. In 2000, spiders (n = 1,541) representing 94 species were captured; 91% of these were captured with pitfall traps. The restored Marion Campus Prairie was inhabited by an assemblage of spiders resembling those on nearby remnant prairies and old fields. Publication Date: February 2020 https://doi.org/10.18061/ojs.v120i2.6904 OHIO J SCI 120(2):2-13 INTRODUCTION In recent decades there has been a movement to with a focus on insects. Much of this work has been restore prairie habitat in North America (Samson done on the influence of disturbance, especially fire, and Knopf 1996). This effort followed the realization on arthropod communities—primarily focused on that tall grass prairies have been largely replaced insects in grasslands (Rice 1932; Bulan and Barrett by agricultural development in North America. 1971; Riechert and Reeder 1972; Seastedt et al. 1986; Currently, less than 5% of this once widespread biome Weaver 1987; Hansen 2000; Harper et al. 2000; Jonas remains (Whiles and Charlton 2006). Ohio lies at et al. 2002; Panzer 2002). Fire can depress populations the eastern end of an area that was characterized by briefly, but these studies generally conclude that patches of tall grass prairie and oak savannah openings arthropod populations recover relatively quickly. referred to as the Prairie Peninsula by Transeau (1935). In an extensive review of the literature on arthropods Cusick and Troutman (1978) estimated that these in prairies, Whiles and Charlton (2006) point out openings covered approximately 294,000 ha in that there is a need for work that determines what Ohio during pre-settlement times. One of the larger species of other arthropod predators, in particular, patches, referred to as the Sandusky Plains, was in spiders, are present in North American prairies. Crawford, Marion, and Wyandot Counties in Ohio. Very few studies provide detailed information about Researchers estimate that the Sandusky Plains once prairie spiders in the North American context (Lowrie covered approximately 77,700 ha (Clutter 2001). 1948; Muma and Muma 1949; Peck 1966; Riechert What remains of the original Sandusky Plains flora and Reeder 1972; Bruggeman 1981; Bultman et are mostly relatively small remnants comprising about al. 1982; Weaver 1987; Okins and Johnson 2005). 100 ha (Troutman 1981). More study of spiders in grassland habitats has been The arthropod fauna of North American prairie conducted in Europe—see references in Curry (1994) grasslands has been studied by many workers (Rice and Bell et al. (2001). These studies have highlighted 1932; Shackleford 1942; Peck 1966; Lussenhop the importance of spiders as mid-level arthropod 1976; Kirchner 1977; Rottman and Capinera 1983; predators in structuring grassland communities. Siemann et al. 1997; Siemann 1998; Harper et al. Spiders in abandoned or fallow agricultural fields, 2000; Jonas et al. 2002; Panzer 2002; Fay 2003). Most often referred to as “old fields,” have been examined of these studies were inventories of species present, by a few workers in North America (MacMahon and Trigg 1972; Peck and Whitcomb 1978; Haskins and 1 Address correspondence to Richard A. Bradley, Department of Evolution, Ecology, and Organismal Biology, The Ohio © 2020 Bradley et al. This article is published under a State University at Marion, 1465 Mt. Vernon Ave., Marion, Creative Commons Attribution 4.0 International License OH 43302, USA. Email: [email protected] ( https://creativecommons.org/licenses/by/4.0/) OHIO JOURNAL OF SCIENCE R. A. BRADLEY ET AL. 3 Shaddy 1986; Cangialosi 1989; Okins and Johnson distant woodlots. At the time of sampling, it was 2005). Again, more work has been conducted in subject to relatively frequent grazing by sheep (lat Europe (see review by Bell et al. 2001). Because old 40°43'50.9"N, long 83°05'38.9"W). fields are often located in regions where the principal Claridon Railroad Prairie (CRP): a 2.2 ha tall native vegetation with this physiognomy are tall grass grass prairie located along a railroad right-of-way. prairies, it has been assumed that they would be This remnant supports a diverse plant community colonized by species of animals that inhabit prairies. including 5 state-listed plant species (2 threatened There appear to be no published studies that explicitly and 3 potentially threatened) (Klips 2003). This site is compare the spider faunas of tall grass prairies and located in Marion County and surrounded primarily old fields. There is 1 study of a similar design that by row-crop agricultural fields and nearby low-density investigated ground beetles (Coleoptera: Carabidae) industrial development. It is managed primarily by in original and restored prairies in Iowa (Larsen selective removal of both non-native species and woody and Work 2003). These authors found that ground plants (lat 40°37'08.4"N, long 83°01'33.8"W). beetle diversity was highest in tall grass prairie, but The Marion Campus Prairie (MCP): a restored that activity density was low. They speculate that the prairie on The Ohio State University at Marion high stem density impedes ground beetle activity on campus, Marion County, Ohio. Plantings at the MCP the ground surface. were initiated by Larry Yoder and his students on 1.1 Prairie restorations have generally depended ha in 1977 (Klips 2004). The restoration effort has upon passive re-colonization for their arthropod continued, and at the time of this study the Marion faunas (Trager 1990). The exceptions to this passive Campus Prairie occupied approximately 2 ha. The approach are exemplified by work with charismatic restoration was accomplished by planting seeds Lepidoptera (e.g., the Karner blue Butterfly,Plebejus and transplanted clumps of vegetation exclusively melissa samuelis (Nabokov, 1944) (Andow et al. from sites within the historic Sandusky Plains, 1994)). It is unknown whether passive colonization primarily from the Claridon Railroad Prairie. The is effective at restoring native arthropod assemblages Marion Campus Prairie is surrounded by row- to restored prairies. The current study compared 2 crop agriculture, a small creek with riparian edge, small prairie remnants of the Sandusky Plains—the lawns, parking lots, and residential and campus Claridon Railroad Prairie (CRP) and the Daughmer development. Management activities on the site Prairie Savannah State Nature Preserve (DPSSNP)— include periodic controlled burning (at approximately and 1 small reconstructed prairie, the 1977 restored 2-year intervals), selective removal, and herbicide Marion Campus Prairie (MCP). Additionally, treatment of weedy invasive and native woody species contemporaneous sampling of 2 old fields in Big (lat 40°34'32.3"N, long 83°05'18.0"W). Island Wildlife Area (BIWA) and Delaware Wildlife Big Island Wildlife Area (BIWA): a 830 ha Area (DWA) provide a comparison with local, early old field in a 2,036 ha wildlife area surrounded by successional habitats. This work tested the hypothesis wooded areas, wetlands, riparian floodplain, low- that passive re-colonization has established a spider density residential, and row-crop agriculture (lat assemblage similar to that in remnant prairie sites 40°34'37.5"N, long 83°15'22.4"W). of the Sandusky Plains, Ohio. Delaware Wildlife Area (DWA): a 311 ha old field in a 2,625 ha wildlife area surrounded by METHODS AND MATERIALS woodlots, wooded riparian floodplain, reservoir, Sampling Localities state park, and some row-crop agriculture (lat Spiders were sampled at 5 locations: 2 were 40°26'20.1"N, long 83°03'25.1"W). remnant prairies (DPSSNP, CRP), 1 was a The 2 old field sites (BIWA, DWA) are located in reconstructed prairie (MCP), and 2 were old fields state-owned wildlife areas and are managed primarily (BIWA, DWA). by periodic mowing and woody plant removal. Daughmer Prairie Savannah State Nature They are dominated by herbaceous perennial forbs Preserve (DPSSNP): a 13.8 ha tall grass prairie- (Solidago spp., Aster spp.), herbaceous annual and bur oak (Quercus macrocarpa) savannah (Troutman biennial forbs (Brassica spp., Cirsium spp., Daucus 1981). This Crawford County site is surrounded carota, Erigeron spp.), and non-native grasses (Setaria primarily by row-crop agricultural fields and Bromus Festuca spp., spp., spp.). 4 PRAIRIE SPIDERS VOL. 120(2) Collecting Methods—Pitfall Trapping and RESULTS Sweep Net Sampling A total of 1,541 spiders were collected. The Pitfall trapping. Open pitfall traps were placed pitfall traps were more productive than the sweep in 2 lines of 5 traps each. The individual traps were samples: 1,407 (91%) individuals were captured in spaced 5 m apart and the 2 lines were separated by the pits while 134 (9%) were captured in the sweep at least 25 m and more than 10 m from any edge. samples. Of 94 species captured during this study, Each trap was an open 1-liter plastic cup with a top 69 (73%) were captured exclusively in pit samples, opening diameter of 11 cm and a depth of 15 cm.