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Habitat Relationships Among Grasshoppers (Orthoptera: Acrididae)

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Habitat Relationships Among Grasshoppers (Orthoptera: Acrididae) Am. Midl. Nat. 142:314-327 Habitat RelationshipsAmong Grasshoppers (Orthoptera:Acrididae)at the WesternLimit of the Great Plains in Colorado DAVID P. CRAIG, CARL E. BOCK, BARRYC. BENNETT and JANE H. BOCK Departmentof Environmental, Population and OrganismicBiology, Campus Box 334, Universityof Colorado,Boulder 80309 ABSTRACT.-We measured grasshopperdensities on 66 shortgrass,mixed grass,tallgrass and hayfieldplots on Boulder,Colorado, open space in 1995-1996. Grasshoppersas a group, and mostspecies individually,were more abundant on relativelysparse shortand mixed grass plots than on lusherhayfields and tallgrassplots-a resultconsistent with the hypothesisthat most species require warm and sunny open ground for survivaland reproduction.Band- winged grasshoppers(Oedipodinae) were particularlyassociated with open ground in short- grassvegetation, whereas spur-throated grasshoppers (Melanoplinae) were mostabundant in mixed grass plots witha high percentage of forbvs. graminoidcover. Slant-facedgrasshop- pers (Gomphocerinae) were the most uniformlydistributed among habitats,but were gen- erallyassociated with plots includingrelatively high proportionsof grassvs. forbcover. While 20 of the 25 most common grasshopperspecies on Boulder open space occurred in all four grasslandtypes, each habitatwas dominated by a distinctiveset of species. However,numer- ically dominant grasshopperson short, mixed and tallgrassplots only loosely resembled groups of dominantspecies in the same habitatselsewhere on the Great Plains. While most Great Plains grasshoppersare widelydistributed, they are apparentlynumerically responsive to combinationsof environmentalconditions expressed at local scales. Common species of grasshopperson Boulder open space in 1995-1996 were the same as those collected in the region in the 1950s. Our resultssuggest these protected grasslands,although invaded by alien vegetationand fragmentedby suburbanization,are stilleffectively conserving this par- ticularinsect group. INTRODUCTION Insectsin general,and grasshoppersin particular,are dominantabove ground herbivores in most grassland ecosystemsbut much remains to be learned about the structureand functionof their communities (Otte, 1981; Curry,1994; Tscharntkeand Greiler,1995). Most grasshoppersof the NorthAmerican Great Plains are widelydistributed, with ranges thatencompass eastern tallgrass,western shortgrass and northernmixed grassprairies that comprise the naturalvegetation across broad expanses of this region (Daubenmire, 1978; Otte, 1981, 1984; Sims, 1988; Pfadt,1994). Local and regional studies suggestthat many Great Plains grasshoppershave distincthabitat requirements,despite their large ranges (Joern, 1982; Capinera and Thompson, 1987; Evans, 1988; Kemp et al., 1990; Quinn and Walgenbach, 1990; Welch et al., 1991; Kemp, 1992). However,we are not aware of any investigationattempting a quantitativecomparison of grasshopperdensities among short, mixed and tallgrasshabitats. One difficultyis the notoriousand apparentlyclimate-related volatilityof manygrasshopper populations (Capinera and Horton, 1989; Joernand Gaines, 1990; Kemp, 1992). These fluctuationscan make it difficultto distinguishfundamental habitatdifferences from temporal variations in widelyseparated sites sampled in different years (but seeJoern and Pruess, 1986). The arid plains of eastern Colorado are, or were, dominated by relativelyuniform short- grassprairie. However, fine-scale mosaics of short,mixed and tallgrasshabitats occur where 314 1999 CRAIG ET AL.: GRASSHOPPER HABITAT 315 the plains meet the eastern edge of the Colorado RockyMountains, apparently the result of increased precipitation,topographic heterogeneity and differencesin soils and land-use history(Livingston, 1952; Branson et al., 1965; Moir, 1969; Bennett, 1997). This circum- stancepermitted a comparisonof grasshoppersoccupying the threefundamentally different sortsof Great Plains grasslandsat a local scale where theirregional populations would be experiencingsimilar patterns of temperatureand precipitation,and presumablywhere in- dividualscould move among the available habitats. Grasslandsat the base of the RockyMountains in Colorado (the so-called Front Range Corridor) are rapidlybeing fragmentedand lost to development(Mutel and Emerick,1992; Long, 1997). Some of the largestremaining stands are part of municipalopen space owned and managed by the Cityof Boulder, Colorado. In 1995-1996 we quantifieddensities of grasshopperson 66, 200-mdiameter plots on Boulder open space, placed systematicallyto include stands of short,mixed and tallgrassvegetation, as well as irrigatedhayfields which have replaced manynative grasslands within the past 150 y.We then compared grasshopper populations among the habitattypes and correlatedtheir densities with vegetative attributes of the plots. Our objectiveswere to: (1) reveal individualspecies' habitatpreferences, (2) determineif the differentgrassland plant communitiessupported distinctivegrasshopper communitiesand (3) compare our resultswith those fromother studies across the Great Plains. Cityc m elevation, around the northern,eastern and southern citylimits, all against a Rocky Mountain foothillsbackdrop of ponderosa pine (Pinus ponderosa)savanna. Plains habitats include riparian corridorsalong streams,tallgrass vegetation and agriculturalhayfields in adjacent lowland floodplainsand mixed and shortgrassstands on upland slopes and bench- es (Bock et al., 1995). Soils are principallyclay or sandy loams (Moreland and Moreland, 1975). Averageannual precipitation(47 cm) is sufficientto supportmixed grassvegetation on all of the uplands. Shortgrassoccurs only in comparativelyheavily grazed areas. We established66,200-m diameter circular plots on Boulder open space grasslandsin the fall of 1993. Preliminaryfloristic surveys were conducted in 1994. Each plot was categorized in its predominantgrassland type based on extensiveplant frequencydata collected inJuly of 1995 and 1996 (Bennett, 1997). Botanical nomenclaturewas taken from Great Plains Flora Association (1986), whereas nativevs. exotic statusof each plant species was deter- mined fromWeber (1995). Types and numbersof plots,and theirpredominant plant spe- cies, were as follows: Shortgrassvegetation (13 plots).-These upland plots had a recent or currenthistory of grazingby cattle,horses and/or black-tailedprairie dogs (Cynomysludovicianus). The most abundant native grasses on shortgrassplots were Agropyronsmithii, Bouteloua gracilis and BuchloI dactyloides.Common native forbswere Artemisiafrigida, Plantago patagonica and Sphaeralceacoccinea. Two common exotic annual grasseswere Bromustectorum and B. Ia- ponicus,while the mostabundant exoticforbs were Alyssum minus and Convolvulusarvensis. Mixed grass vegetation(21 plots).-These upland plots were ungrazed, or lightly-grazed, and were dominated by varyingmixtures of midheightand short grasses and sedges, de- pending on soils and topography.Common nativegraminoids included Agropyronsmithii, Andropogongerardii, A. scoparius,Aristida purpurea, Bouteloua gracilis, B. curtipendula,Car- ex heliophila,Koelaria pyramidata, Panicum virgatum,Stipa comataand S. neomexicana.The mostabundant nativeforbs were Ambrosiapsilostachya, Artemisia ludoviciana, Asterfalcatus, 316 THE AMERICAN MIDLAND NATURALIST 142(2) Helianthuspumilis and Psoralea tenuiflora.Common exotics on mixed grass plots included Alyssumminus, Bromus japonicus, Poa pratensis,P compressaand Tragapogondubius. Tallgrassvegetation (11 plots).-These plots occurred in lowland floodplains or on low benches with cobbly soils and high water tables. Some were winter-grazedby cattle,but all remained undisturbedthrough the summergrowing season. They were dominated by mid- height and tall grassescharacteristic of the True Prairie of the eastern Great Plains (Risser et al., 1981):' Andropogongerardii, A. scoparius,Panicum virgatum,Sorghastrum nutans and Sporobolusasper. The most abundant native forbs on tallgrassplots were Ambrosiapsilos- tachya,Aster falcatus, Dicantheliumoligosanthes, Grindelia squarosa and Prunella vulgaris. There were many exotic species on tallgrassplots, including the forbs Cichoriumintybus, Plantago lanceolata, Taraxacum officinale,Trifolium pratense and a varietyof cool-season meadow grasses planted for pasture improvement(Agrostis stolonifera, Festuca pratensis, Phleumpratense and Poa pratensis). Hayfields (21 plots).-These lowland plots were flood-irrigatedin early summer and mowed between mid-Julyand earlyAugust. Most were also winter-grazedby cattle. Some included remnants of the tallgrassvegetation they replaced, but all were dominated by exotic species such as alfalfa (Medicago sativa) and a series of cool-season pasture grasses, including Bromusinermis, Festuca pratensis, Dactylis glomerata and Phleumpratense. METHODS Vegetationcover.-Vegetation canopy cover was sampled on each of the 66 plots in June and again in August of 1995 and 1996. During each samplingperiod 100 m transectswere establishedrunning due east and due westfrom each plot center.An optical point projector was set at 4 m intervalsalong each transectand vegetation was sampled at six random points within1 m of each setting.This resultedin 300 points per plot per samplingperiod (n = 1200 total points per plot over the 2 y). Each point was classifiedas unvegetatedor as covered by one or more of the followingfour vegetationcategories: native graminoids, exotic graminoids,native forbs and/or exotic forbs.Results from the foursampling periods were averaged to generate
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