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(Orthoptera: Tettigoniidae) Species Composition and Size Across Latitude in Atlantic Coast Salt Marshes

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(Orthoptera: Tettigoniidae) Species Composition and Size Across Latitude in Atlantic Coast Salt Marshes Estuaries and Coasts: J CERF (2008) 31:335–343 DOI 10.1007/s12237-007-9026-3 Grasshopper (Orthoptera: Tettigoniidae) Species Composition and Size Across Latitude in Atlantic Coast Salt Marshes Elizabeth L. Wason & Steven C. Pennings Received: 14 June 2007 /Revised: 5 November 2007 /Accepted: 16 November 2007 /Published online: 16 January 2008 # Coastal and Estuarine Research Federation 2007 Abstract Although grasshoppers are common salt marsh Geographic variation in species composition and body size herbivores, we know little about geographic variation in of grasshoppers may help explain documented patterns of their species composition. We documented latitudinal geographic variation in plant palatability and plant–herbivore variation in species composition of the tettigoniid grass- interactions in Atlantic Coast salt marshes. Because it can be hopper fauna of Atlantic Coast salt marshes. Tettigoniids difficult to identify tettigoniids to species, we present a guide (N=740 adults) were collected from the Spartina alterni- to aid future workers in identifying the tettigoniid species flora zone of 31 salt marsh sites across a latitudinal range of common in these marshes. 13.19° (Florida to Maine), with an additional 52 individuals collected from the Juncus roemerianus zone of low-latitude Keywords Biogeography. Latitude . marshes for comparative purposes. Eight species were Plant–herbivore interactions . Bergmann’s rule . collected, but some were common only at a few sites or Orthoptera . Spartina alterniflora . Salt marsh rare throughout the entire collection range. The tettigoniid community was dominated by Orchelimum fidicinium at low latitudes and Conocephalus spartinae at high latitudes. Introduction Several factors might explain this shift, including changes in climate, plant phenology, and plant zonation patterns. Most ecological research is conducted at single sites, but O. fidicinium and C. spartinae increased in body size toward ecologists are interested in generalizing their results as low latitudes. In laboratory feeding assays, O. fidicinium broadly as possible. Our ability to make broad general- readily ate S. alterniflora and J. roemerianus leaves, izations is contingent upon an understanding of how abiotic Orchelimum concinnum, which is largely restricted to the and biotic conditions vary geographically and how this J. roemerianus zone, ate only J. roemerianus leaves, and variation affects ecological processes. Conocephalus spp. ate neither, consistent with literature Salt marshes along the Atlantic Coast of the USA are suggestions that they mainly consume seeds and flowers. superficially similar from northern Florida through Maine (Pennings and Bertness 2001); however, there is extensive latitudinal variation in soil composition and salinity (Pennings and Bertness 1999; Craft 2007), plant phenology, morphology E. L. Wason : S. C. Pennings Biology and Biochemistry, University of Houston, and productivity (Turner 1976;Seliskaretal.2002), Houston, TX 77204-5001, USA interactions between plant species (Bertness and Ewanchuk – S. C. Pennings 2002; Pennings et al. 2003), plant herbivore interactions e-mail: [email protected] (Pennings et al. 2001), and herbivore life history (Denno et al. 1996). Thus, extrapolating results from any one site E. L. Wason (*) requires an understanding of how both the abiotic and biotic University of Michigan, conditions at that site compare with those at other sites. 1300 Kraus Natural Science Building, 830 North University, Ann Arbor, MI 48109-1048, USA In this paper, we focus on latitudinal variation in the e-mail: [email protected] composition of an important group of salt marsh herbivores, 336 Estuaries and Coasts: J CERF (2008) 31:335–343 tettigoniid grasshoppers. Herbivore pressure on salt marsh range of 13.19° (Table 1). Our sampling focused on the plants appears to be stronger at lower latitudes (Pennings and S. alterniflora zone because this is the largest vegetation Silliman 2005; Pennings unpublished data). This increased zone in many Atlantic Coast salt marshes and occurs at all herbivore pressure may cause plants at low latitudes to latitudes. We collected an average of 24 (range 1 to 59) develop stronger defenses against consumers (Siska et al. adult tettigoniids from each site, for a total of 740 2002), resulting in decreased palatability of plants with individuals. We visited most sites once or twice in the decreasing latitude (Pennings et al. 2001). It is likely that summer of each of 3 years (2004–2006). In addition, we latitudinal variation in herbivore pressure is driven, at least in made a few collections (N=99 of the 740) in May, part, by geographic variation in grasshopper species compo- September, and October to confirm that species composi- sition (Pennings and Silliman 2005), but rigorous data to tion did not vary seasonally (it did not), and a small number address this hypothesis are lacking. (N=24 of the 740) of specimens were collected in October Few data on grasshopper species composition exist for of 2003. Collecting methods were consistent across sites, Atlantic Coast salt marshes, except for one study in although the time spent collecting was greater at sites where North Carolina (Davis and Gray 1966). Although no grasshopper density was low. Because our interest was in comparative studies have been conducted across broad the relative abundance of different grasshopper species, we geographic regions, ecological studies in New England did not document absolute densities (numbers per square (Bertness et al. 1987; Bertness and Shumway 1992), meter). We collected only adult tettigoniids, avoiding Georgia (Smalley 1960), and Mississippi (Parsons and de juvenile tettigoniids, which were difficult to identify to la Cruz 1980) have focused on different tettigoniid species, as well as individuals from other grasshopper grasshopper species (Conocephalus spartinae in New families (i.e., Acrididae), which were rare in the S. England, Orchelimum fidicinium in Georgia, and O. alterniflora zone. We collected tettigoniids using a combi- concinnum in Mississippi), which presumably were the nation of sweep netting, targeted netting, and hand most abundant grasshoppers at each site. Whether this collection. In addition, we collected tettigoniids (N=52) reflects local site-to-site variation in species composition from the Juncus roemerianus zone of four low-latitude salt or a broader geographic pattern, however, is unclear. marsh sites for comparative purposes (three sites in GA and Nevertheless, based on these studies, the feeding ecology one in SC) on one date per site in either 2005 or 2006. The of different tettigoniid species appears to vary, with C. Juncus zones at these sites were dominated by J. spartinae feeding primarily on flowers and seeds (Bertness roemerianus, with small amounts of Batis maritima, et al. 1987; Bertness and Shumway 1992)andOrchelimum Borrichia frutescens, Distichlis spicata,orS. alterniflora. spp. feeding on leaves (Smalley 1960; Parson and de la Cruz 1980). This suggests that documenting patterns of grasshop- Identification of Tettigoniid Species per species composition across latitude would improve our understanding of biogeographic patterns in plant–insect We classified tettigoniids to species following Rehn and interactions. Hebard (1915a, b, c) and by comparing our specimens to One obstacle to addressing these issues has been the those in the collection of the Entomology and Nematology difficulty in identifying tettigoniid grasshoppers to species. Department, University of Florida, Gainesville. The eight Although tettigoniids have been the subject of excellent tettigoniid species that we found in Atlantic Coast S. taxonomic monographs (Rehn and Hebard 1915a, b, c), the alterniflora salt marshes can be distinguished based on species are morphologically quite similar and difficult for non- body size, wing length, coloration, and morphology of experts to identify to species. Therefore, in addition to genitalia (cerci in males and ovipositors in females). documenting tettigoniid species composition and distribution Tettigoniids were observed under a dissecting microscope in salt marshes along the Atlantic Coast of the USA, we and measured with electronic calipers (Fig. 1). Body size provide a resource to facilitate tettigoniid species identification was measured as the length of the grasshopper, excluding by future workers. the cerci or ovipositor, from the tip of the head to the end of the abdomen. Measurements of body size, however, are affected by the posture of the preserved grasshopper. A Materials and Methods more reliable indicator of size is the length of the hind tibia, which was measured from the joint between the tibia and Study Sites and Collections femur to the joint between the tibia and tarsus. We focused on tibia length for our analyses, but analyses of body length We collected tettigoniid grasshoppers from the Spartina produced similar results. Wing length was measured from alterniflora zone of 31 salt marsh sites along the Atlantic the abuttal of the pronotum and wing to the distal end of the Coast of the USA from Florida to Maine across a latitudinal wing. Each cercus was measured from the abuttal with the Estuaries and Coasts: J CERF (2008) 31:335–343 337 Table 1 Study sites for tettigoniid collections trials lasted 4 days, with each replicate beginning on the Site name State Latitude Number Number day that each individual insect was collected. Individual collected collected tettigoniids were housed in 1,000-ml glass
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