Prey Preference and Diet of Neonate Eastern Massasaugas (Sistrurus c. catenatus)

Am. Midl. Nat. 152:360 368 Prey Preference and Diet of Neonate Eastern Massasaugas (Sistrurus c. catenatus) DONALD B. SHEPARD, 1 CHRISTOPHER A. PHILLIPS, MICHAEL J. DRESLIK AND BENJAMIN C. JELLEN Illinois Natural History Survey, Center for Biodiversity, 607 East Peabody Drive, Champaign 61820 ABSTRACT. Because prey acquisition in young organisms often has profound effects later in life, understanding the foraging ecology of early age classes is important. We examined diet and prey preference of neonate Eastern Massasaugas (Sistrurus c. catenatus) at Carlyle Lake, Clinton County, Illinois. Prey recovered from free-ranging neonates consisted primarily of southern short-tailed shrews (Blarina carolinensis). In feeding trials, neonates demonstrated a preference for snake prey, disinterest in anuran and insect prey and indifference toward mammal prey. Because of gape limitations, neonates may have difficulty ingesting small mammals, but snakes are comparably easier to ingest and are the most common prey item of young S. c. catenatus in other parts of their range. Blarina carolinensis has not been reported previously from the diet of S. c. catenatus as their ranges overlap only in southwestern Illinois. Blarina carolinensis is considerably smaller than most mammals preyed upon by older age classes and would be easier for neonates to ingest. Thus, at Carlyle Lake, snakes may not be as important a prey resource for neonate massasaugas as in other parts of their range due to the availability of B. carolinensis. INTRODUCTION In many organisms, the ecology of younger age classes differs considerably from older classes. Ontogenetic dietary shifts in prey composition and prey size are common in gapelimited predators such as snakes (Mushinsky et al., 1982; Reynolds and Scott, 1982; Mushinsky, 1987; Henderson and Crother, 1989; Arnold, 1993; Martins et al., 2002). However, due to the secretive nature of many snake species, there is limited information on their ecology with most data derived from studies involving adults (Parker and Plummer, 1987). Because prey acquisition at an early age influences growth, age at first reproduction and adult body size (Ford and Seigel, 1994; Madsen and Shine, 2000), it is critical to understand the foraging ecology of young snakes. Dietary information can also be important in formulating management plans for endangered and threatened species (Holycross et al., 2002). The Eastern Massasauga (Sistrurus c. catenatus) occurs in widely disjunct populations from western New York and southern Ontario to eastern Iowa and Missouri (Conant and Collins, 1991; Ernst, 1992). It is afforded legal protection at the state/provincial level throughout its range (Szymanski, 1998) and is currently a candidate for federal listing under the U.S. Endangered Species Act of 1973 (U.S. Fish and Wildlife Service, 1999). The Massasauga (S. catenatus) exhibits substantial variation in natural history across its range (Ernst, 1992), especially in regard to diet (Holycross and Mackessy, 2002). Young age classes have been underrepresented in previous S. c. catenatus dietary studies (reviewed by Holycross and 1 Corresponding author present address: Department of Zoology and Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, 2401 Chautauqua Ave, Norman. 73072; e-mail: dshepard@ou.edu 360

2004 SHEPARD ET AL.: MASSASAUGAS DIET 361 Mackessy, 2002) and little information is available regarding neonate diet and life history in general (Szymanski, 1998). Previous dietary records for neonate S. c. catenatus are few, but include snakes (Keenlyne and Beer, 1973; Hallock, 1991; Mauger and Wilson, 1999), small mammals (Bielema, 1973; Hallock, 1991) and possibly insects (Hallock, 1991) from freeranging neonates and frogs in captive neonates (Schuett et al., 1984). However, considering the variety of prey reported in spite of the small number of records, the variation in natural history exhibited by S. c. catenatus across its range and its imperiled status, a more detailed examination of neonate diet is desirable. The objective of this study was to determine prey preference and diet of neonate S. c. catenatus and examine other general aspects of neonate foraging ecology. MATERIALS AND METHODS During summer 2001, we captured three gravid female Sistrurus c. catenatus from sites around Carlyle Lake, Clinton County, Illinois, and held them in captivity through parturition in early August. We conducted feeding trials using neonates from these three broods (n ¼ 5, 7 and 8) and neonates incidentally captured around the Carlyle Lake region (n ¼ 6). All neonates had previously completed their first shed as evidenced by the presence of a button rattle segment. For all neonates we measured snout-vent length (SVL) to the nearest 1 mm using a flexible measuring tape and mass to the nearest 0.1 g. We placed neonates into individual plastic containers (18 cm diam 3 8 cm height) and allowed them to acclimate to their surroundings for at least 12 h before the initiation of feeding trials. Our choices of prey types for feeding trials were based upon published diet records, what previous authors speculated as prey in other regions and what we considered potential prey of neonates at Carlyle Lake. In Trial 1 we offered neonates (n ¼ 26) one of three prey types: orthopteran (Acrididae, Tettigoniidae; n ¼ 8), anuran (Acris crepitans; n¼ 9) and mammal (neonate Mus musculus; n ¼ 9). We controlled for possible maternal/genetic effects by dividing individuals from the same brood among prey groups. For prey items, we measured body length and maximum width to the nearest 0.1 mm with digital calipers and mass to the nearest 0.1 g. We introduced prey items into the containers, observed neonate behavior continuously for the first hour and then checked neonates periodically (every 1 4 h) for the next 11 h recording whether the prey item had been ingested. To adequately observe the behavior of neonates during the first hour after prey introduction, we offset the starting times of the different prey groups by 1 or 2 h. We ended feeding trials when 12 h had elapsed since introduction of the prey item. In Trial 2 we offered a second prey item to 19 of the 21 neonates that did not ingest a prey item in Trial 1. We randomly divided these neonates into three prey groups: mammal (neonate Mus musculus; n ¼ 13), snake (neonate Thamnophis sirtalis; n ¼ 5) and lizard (Eumeces fasciatus; n ¼ 1). We scored Trial 2 solely as to whether the item was ingested within 12 h. All neonates were released at their site of capture and snakes born in captivity were released with their mother at the mother s site of capture following the study. In addition to lab trials, we made field observations during late summer and fall 2001 on the diet of free-ranging neonate Sistrurus c. catenatus throughout the Carlyle Lake region. Neonates were easily identified by their small size, presence of an umbilical scar and button rattle segment. We measured all neonates captured (SVL and mass), manually palpated them and forced individuals with stomach contents to regurgitate. We identified prey items to species and measured length, maximum head width and mass when possible. At each neonate capture site we measured percent canopy cover using a spherical densiometer and the distance to the nearest forest edge.

362 THE AMERICAN MIDLAND NATURALIST 152(2) TABLE 1. Mean 6 SE sizes of prey items used in feeding trials (n ¼ size of sample from which mean is based and not necessarily number of prey used in feeding trials) Prey Length (mm) Max. width (mm) Mass (g) Anuran (n ¼ 9) 20.8 6 0.2 6.0 6 0.1 0.8 6 0.1 Mammal (n ¼ 9) 42.7 6 0.6 7.9 6 0.2 2.8 6 0.1 Orthopteran (n ¼ 8) 32.2 6 2.6 4.9 6 0.3 0.8 6 0.2 Snake (n ¼ 39) 125.6 4.7 * 1.4 Lizard (n ¼ 1) 41.2 8.5 2.3 Measurement based on whole brood; SE not available * Measurement based on one stillborn from same brood To test if neonates demonstrated preferences for particular prey types during feeding trials, we used a G -test (Sokal and Rohlf, 1995) on the pooled results from Trials 1 and 2. Because this test uses logarithms and some observed values were zero, it was necessary to add one to the value in each cell of the contingency table. We also used a G -test to test for maternal effects on ingestion during our feeding trials. To determine if prey size affected ingestion of mammal prey, we used ANOVA with Tukey HSD post-hoc tests to test for differences in maximum head width between mammal prey that were ingested in Trial 1, mammal prey that were not ingested in Trial 1 and mammal prey recovered from freeranging neonates. To determine if individuals with meals were associated with particular habitat characteristics, we used two-tailed t-tests to test for differences in canopy cover and distance to forest edge among neonates with and without stomach contents. We used Mann- Whitney U-tests when assumptions of parametric tests were not satisfied. Means are reported 61 SE and all alpha levels were set a priori at 0.05. RESULTS Neonates used in feeding trials (n ¼ 26) averaged 209 6 2.5 mm SVL and 10.4 6 0.2 g mass. Mean sizes of prey items used in feeding trials varied (Table 1), but all were initially believed to be within the range of ingestible sizes. In Trial 1, 5 of 26 neonates ingested the prey item presented to them (Table 2). All nine neonates offered mammal prey tongue flicked, oriented towards and struck at the prey item (usually in less than 30 s). Four TABLE 2. Results from neonate Sistrurus c. catenatus feeding trials. Shown as number of individuals that ingested prey over the number presented with that prey item. Individuals that did not eat in Trial 1 were reused in Trial 2 Mother Prey type #132 #174 #192 Unknown Total Orthopteran Trial 1 0/2 0/1 0/3 0/2 0/8 Anuran Trial 1 0/3 0/2 0/2 1/2 1/9 Mammal Trial 1 2/3 1/2 1/2 0/2 4/9 Subtotal 2/8 1/5 1/7 1/6 5/26 Mammal Trial 2 1/4 0/2 2/4 2/3 5/13 Snake Trial 2 2/2 2/2 1/1 0/0 5/5 Lizard Trial 2 0/0 0/0 1/1 0/0 1/1 Subtotal 3/6 2/4 4/6 2/3 * 11/19 Total 5/8 3/5 5/7 3/6

2004 SHEPARD ET AL.: MASSASAUGAS DIET 363 TABLE 3. Results from Trial 2 in which individuals that did not eat in Trial 1 were presented with a second prey item. Numbers shown are the number of individuals that ate over the number presented with that prey item Trial 2 item Trial 1 item Mammal Snake Lizard Mammal 0/3 0/0 0/0 Anuran 4/7 1/1 0/0 Orthopteran 1/3 4/4 1/1 neonates successfully ingested the prey item, three neonates attempted ingestion but failed and two neonates made no attempt at ingestion. Four of the nine neonates offered anuran prey tongue flicked and orientated towards the prey, but only one neonate ingested the prey. This individual did not strike and release the prey, but ingested it alive. All eight neonates offered orthopteran prey showed no response. In Trial 2, 11 of 19 neonates ingested the prey item presented to them (Table 2). Because there appeared to be no effect of reusing individuals that did not eat in Trial 1 (Table 3), we pooled all observations (n ¼ 45) to determine if neonates demonstrated prey preferences. We excluded lizard prey from the analysis because of small sample size (n ¼ 1). Overall, neonates exhibited significant prey preferences (G Williams ¼ 12.79, df ¼ 3, P ¼ 0.005) with relative disinterest toward orthopterans and anurans, a preference for snakes and indifference towards mammal prey. Further, there was no maternal influence on whether or not a snake ate during our experiments (G Williams ¼ 0.62, df ¼ 3, P ¼ 0.89). From August through November 2001 we captured 19 neonate Sistrurus c. catenatus and recovered eight identifiable prey items from seven individuals with stomach contents. We obtained single prey items from one individual on two occasions 11 d apart. In all cases, prey items were small mammals: seven southern short-tailed shrews (Blarina carolinensis) and one juvenile prairie vole (Microtus ochrogaster). One B. carolinensis had small maggots on it, suggesting that it was scavenged. Unidentifiable mammal hair was also recovered from the feces or digestive tract of three neonates. We excluded two individuals from the analysis of habitat variables because they were encountered in developed areas. Neonates with meals were encountered closer to the forest edge than neonates without meals (with mean¼10.4 6 4.2 m, without mean ¼ 30.9 6 7.5 m; t 15 ¼ 2.56, P ¼ 0.02), but canopy cover did not significantly differ (with mean ¼ 31.4 6 12.3%, mean rank ¼ 10.4, n ¼ 10; without mean ¼ 4.9 6 2.3%, mean rank ¼ 7, n ¼ 7; Mann-Whitney U-test, Z ¼ 1.375, P ¼ 0.17). Because snakes are gape-limited predators and three neonates from Trial 1 had difficulty ingesting mammal prey, we examined whether prey size affected ingestion. To test if prey size had an influence, we compared the maximum head width of mammal prey ingested in Trial 1, mammal prey not ingested in Trial 1 and mammal prey recovered from free-ranging neonates. We were able to accurately measure maximum head width of six prey items (all Blarina carolinensis) recovered from free-ranging neonates. Mean maximum head width of those prey was 8.0 6 0.5 mm (range 6.8 10.5 mm). We observed one neonate (225 mm SVL, 7.3 g mass) that ingested a large B. carolinensis (10.5 mm maximum head width, 10.8 g mass) and was so distended when encountered that it was incapacitated and died before any attempt to force regurgitation. When this outlier was removed, mean maximum head width of prey from free-ranging neonates was 7.5 6 0.2 mm. Overall, the size (width) of the mammal prey did affect ingestion (F 2,11 ¼ 7.21, P ¼ 0.01). In Trial 1 the maximum head widths of ingested mammal prey were narrower than those not ingested (ingested mean ¼ 7.5 6 0.1 mm,

364 THE AMERICAN MIDLAND NATURALIST 152(2) FIG. 1. Neonate snout-vent length (SVL) versus mammal prey maximum head width for Trial 1 and field observations. Trial 1 is demarcated as successful ingestion of prey, failed attempt, or no attempt made. (* neonate died from causes related to ingestion of prey item) n ¼ 4; not ingested mean ¼ 8.2 6 0.2 mm, n ¼ 5; P ¼ 0.03). The maximum head widths of prey from free-ranging neonates (excluding one outlier) were not significantly different than those of mammal prey ingested in Trial 1 (P ¼ 0.96), but significantly less than those that were not ingested (P ¼ 0.01). When all data on mammal prey are examined, neonates (180 300 mm SVL) primarily ingested mammal prey 8.0 mm in head width (Fig. 1). DISCUSSION Orthopterans elicited no response from Sistrurus c. catenatus in our feeding trials or in feeding trials with a population from western Illinois (Bielema, 1973). The diet of S. c. edwardsii includes centipedes (Holycross and Mackessy, 2002) and the diet of S. miliarius includes insects, spiders and centipedes (Hamilton and Pollack, 1955; Ernst, 1992), but there has been only one report of insects (Coleoptera) in the diet of S. c. catenatus (Hallock, 1991). Thus, in spite of their abundance, insects are apparently not eaten by S. c. catenatus. Although anurans elicited tongue flicking in some of our neonates, only one was ingested. Several authors have reported that captive Sistrurus c. catenatus readily ingest anurans (Atkinson and Netting, 1927; Curran, 1935; Schuett et al., 1984); however, documented cases in free-ranging S. c. catenatus are few (see Holycross and Mackessy, 2002). Captive neonate S. c. catenatus and S. c. tergeminus caudal lure for anurans (Schuett et al., 1984; Reiserer, 2002), but there appears to be geographic variation in the stimulus control of caudal luring for anurans in S. catenatus (Reiserer, 2002), which may be linked to the considerable variety of habitats the species occupies throughout its range (Ernst, 1992). Our results are consistent with previous studies on S. catenatus in that frogs may be consumed, but probably constitute a small portion of the diet (Holycross and Mackessy, 2002). However, the importance of frogs in the diet may vary seasonally and/or geographically. Although lizards constitute a significant portion of the diet of Sistrurus c. tergeminus and S. c. edwardsii, especially snakes in smaller size classes, lizards have not been reported from the

2004 SHEPARD ET AL.: MASSASAUGAS DIET 365 diet of S. c. catenatus (Greene and Oliver, 1965; Holycross and Mackessy, 2002). In the United States, lizard species richness is highest in the arid southwest (Schall and Pianka, 1978), thus S. c. edwardsii and S. c. tergeminus are exposed to a greater diversity and abundance of lizard prey. However, in the northeastern part of the range of S. c. tergeminus, where habitats are more mesic and lizard species richness is lower, lizards are not part of the diet (Seigel, 1986; Holycross and Mackessy, 2002). In Illinois, as in much of the east-central and northeastern United States, lizard species richness is low and lizards do not commonly occupy the same habitats as S. c. catenatus. At Carlyle Lake, the only lizard we have encountered, albeit infrequently, is Eumeces fasciatus. Although one neonate S. c. catenatus consumed a lizard in our study, the proportion of lizards in the diet is probably small due to low availability. Neonates consumed snake prey in every case it was offered. Ophiophagy is more common in smaller size classes in Sistrurus c. catenatus and S. c. tergeminus and predation on Thamnophis sirtalis, Storeria dekayi and Opheodrys vernalis has been reported (Keenlyne and Beer, 1973; Seigel, 1986; Hallock, 1991; Mauger and Wilson, 1999; Holycross and Mackessy, 2002). In Illinois, T. sirtalis and S. dekayi are born about the same time as S. c. catenatus in late summer and occur in a variety of habitats (Smith, 1961; Phillips et al., 1999) and, thus, are probably frequently encountered by neonate S. c. catenatus. The diet of adult Sistrurus c. catenatus primarily consists of small mammals (reviewed by Holycross and Mackessy, 2002), including voles (Microtus), mice (Peromyscus and Zapus) and shrews (Blarina and Sorex), which are usually abundant in the snake s habitat. Although all neonates presented mammal prey in Trial 1 struck at the prey, only half ingested it. Because most neonates attempted to ingest mammal prey, they apparently have a strong mammaldirected feeding response, but had problems with prey handling and ingestion. Although the mammal prey used in our experiments were typically one-half to one-third the mass of prey recovered from free-ranging neonates, differences in prey width prevented ingestion. The mammal prey we used were evidently near the threshold limit of ingestible prey size for neonates (Fig. 1). In gape-limited predators such as snakes, ontogenetic dietary shifts in prey composition and prey size often occur (Mushinsky et al., 1982; Reynolds and Scott, 1982; Mushinsky, 1987; Henderson and Crother, 1989; Arnold, 1993; Holycross et al., 2002; Martins et al., 2002). Adult and juvenile Sistrurus c. edwardsii differ in diet with juveniles consuming a higher proportion of lizards (Holycross and Mackessy, 2002). Generalization from previous studies on S. c. catenatus reveals an ontogenetic dietary shift from snakes (in young age classes), to small mammals such as Blarina and Zapus, then to larger rodents such as Peromyscus and Microtus in older classes (Bielema, 1973; Keenlyne and Beer, 1973; Hallock, 1991; Mauger and Wilson, 1999). Snake prey are an especially important resource for young S. c. catenatus because a snake s body shape does not require as great a gape as a small mammal of similar mass. Due to their specialized morphology, viperids are capable of ingesting prey of high relative body mass (Greene, 1983; Pough and Groves, 1983; Cundall and Greene, 2000). In many species, relative prey mass is greatest for smaller size classes (Sazima, 1992; Shine et al., 1998; Martins et al., 2002). Extreme MRs (mass ratio) of 1.30 in Bothrops jararaca (Sazima, 1992), 1.56 in Bothrops atrox (Greene, 1983), 1.57 in Bitis caudalis (Branch et al., 2002) and 1.72 in Crotalus cerastes (Mulcahy et al., 2003) have been reported. Our results on neonate Sistrurus c. catenatus further demonstrate the ability of small viperids to ingest large prey. In our study, one individual had a MR of 1.48, whereas the mean MR for the other five measurable records from free-ranging neonates was 0.47 6 0.06. Additionally, after this study in August 2002, we encountered a dead neonate S. c. catenatus (225 mm SVL, 8.0 g mass) that contained a large B. carolinensis (7.5 g mass, MR ¼ 0.94). Ingestion of a large prey

366 THE AMERICAN MIDLAND NATURALIST 152(2) item may result in mortality through decreased locomotor performance and inability to escape predators or unfavorable environmental conditions (Pauly and Benard, 2002); however, mortality might also result simply from the prey being too large (Mulcahy et al., 2003). Because the majority of neonates in our experiments were naive predators, their responses should be indicative of the innate prey preferences of young Sistrurus c. catenatus at Carlyle Lake. Captive neonates exhibited a propensity for both snake and small mammal prey and we have observed a captive individual eat both a small S. dekayi and a small Mus musculus one week apart. However, our observations on the diet of free-ranging neonates showed their diet consisted primarily of small mammals, mainly the southern short-tailed shrew (Blarina carolinensis). The closely related northern short-tailed shrew (B. brevicauda) has been reported from the diet of all size classes (Hallock, 1991; Anton, 1993; Mauger and Wilson, 1999) including neonates (Hallock, 1991). Because Blarina are considerably smaller than Microtus and Peromyscus (Hoffmeister, 1989), smaller snakes may be able to prey on them. The range of B. brevicauda overlaps the range of S. c. catenatus entirely; however, southwestern Illinois is the only region where the ranges of S. c. catentatus and B. carolinensis overlap (Hoffmeister, 1989; Conant and Collins, 1991). Blarina carolinensis (mean mass ¼ 9.6 g) are substantially smaller than B. brevicauda (mean mass ¼ 18.5 g; Hoffmeister, 1989). At Carlyle Lake, neonate S. c. catenatus are able to prey upon small mammals immediately because the smaller B. carolinensis is available. Thus, snakes may not be as important a prey resource for young S. c. catenatus at Carlyle Lake when compared to other parts of their range and the availability of B. carolinensis may be an important factor influencing the viability of the Carlyle Lake population. Neonates with meals were associated with wooded edge habitats. This indicates dispersal from gestation and birthing sites, which are typically in open areas due to the thermal requirements of gestating females (Johnson, 1995; Dreslik, unpubl.). This habitat shift may be due to foraging tactics. In Illinois, Blarina occur in a variety of habitats, but are most numerous in wooded areas with heavy leaf litter (Hoffmeister, 1989). Neonate Sistrurus c. catenatus likely use chemical cues to select foraging sites as has been shown in S. miliarius (Roth et al., 1999). Our observations suggest that neonates move to wooded habitats to forage where encounters with Blarina would be more frequent. Mortality of neonate snakes is typically highest during postnatal dispersal and long dispersal distances increase their risk of mortality (Bonnet et al., 1999). Thus, at Carlyle Lake, wooded habitat in proximity to neonate birthing sites may be an important factor to consider when formulating habitat management plans. Acknowledgments. We thank the Illinois Department of Natural Resources, U.S. Fish and Wildlife Service, Illinois Wildlife Preservation Fund and U.S. Army Corps of Engineers for financial support; J. Hoffman for assistance in identifying stomach contents; J. Smothers, J. Selle, S. Luebbers and V. Lohr for turning in snakes; J. Bunnell, J. Birdsell, J. Smothers, D. Baum, A. Kuhns, P. Jellen, J. Mui, J. Petzing, G. Tatham, T. Anton and D. Mauger for field assistance; A. Holycross for information on Sistrurus c. edwardsii and S. c. tergeminus; R. Reiserer for information on caudal luring in S. catenatus; A. Holycross, A. Kuhns, and J. Mui for suggestions on improving earlier versions of this manuscript. LITERATURE CITED ANTON, T. G. 1993. Massasaugas in Lake County, Illinois: the Ryerson population, p. 71 77. In: B. Johnson and V. Menzies (eds.). International symposium and workshop on the conservation of the Eastern Massasauga Rattlesnake Sistrurus catenatus catenatus. Toronto Zoo, Toronto. ARNOLD, S. J. 1993. Foraging theory and prey-size predator-size relations in snakes, p. 87 115. In: R. A. Seigel and J. T. Collins (eds.). Snakes: ecology and behavior. McGraw-Hill, N.Y.

2004 SHEPARD ET AL.: MASSASAUGAS DIET 367 ATKINSON, D. A. AND M. G. NETTING. 1927. The distribution and habits of the massasauga. Bull. Antivenin Inst. Amer., 1:40 44. BIELEMA, B. J. 1973. The eastern massasauga (Sistrurus catenatus catenatus) in west-central Illinois. M.Sc. Thesis, Western Illinois Univ., Macomb. 80 p. BONNET, X., G. NAULLEAU AND R. SHINE. 1999. The dangers of leaving home: dispersal and mortality in snakes. Biol. Conserv., 89:39 50. BRANCH, W. R., A. M. BAUER AND T. LAMB. 2002. Bitis caudalis. Prey size. Herpetol. Rev., 33:137 138. CONANT, R. AND J. T. COLLINS. 1991. A field guide to reptiles and amphibians: eastern and central North America, 3rd ed. Houghton Mifflin, Boston. 450 p. CUNDALL, D. AND H. W. GREENE. 2000. Feeding in snakes, p. 293 333. In: K. Schwenk (ed.). Feeding: form, function, and evolution in tetrapod vertebrates. Academic Press, San Diego. CURRAN, C. H. 1935. Rattlesnakes. Natural History, 36:331 340. ERNST, C. H. 1992. Venomous reptiles of North America. Smithsonian Institution Press, Washington, D.C. 236 p. FORD, N. B. AND R. A. SEIGEL. 1994. An experimental study of the trade-offs between age and size at maturity: effects of energy availability. Functional Ecol., 8:91 96. GREENE, H. W. 1983. Dietary correlates of the origin and radiation of snakes. Amer. Zool., 23:431 441. AND G. V. OLIVER, JR. 1965. Notes on the natural history of the western massasauga. Herpetologica, 21:225 228. HALLOCK, L. A. 1991. Habitat utilization, diet and behavior of the eastern massasauga (Sistrurus catenatus catenatus) in southern Michigan. M.Sc. Thesis, Michigan State Univ., East Lansing. 40 p. HAMILTON, W. J., JR. AND J. A. POLLACK. 1955. The food of some crotalid snakes from Fort Benning, Georgia. Nat. Hist. Misc., 140:1 4. HENDERSON, R.W.AND B. I. CROTHER. 1989. Biogeographic patterns of predation in West Indian colubrid snakes, p. 479 517. In: C. A. Woods (ed.). Biogeography of the West Indies: past, present, and future. Sandhill Crane Press, Gainesville, Fla. HOFFMEISTER, D. F. 1989. Mammals of Illinois. Univ. of Illinois Press, Urbana, Ill. 348 p. HOLYCROSS, A. T. AND S. P. MACKESSY. 2002. Variation in the diet of Sistrurus catenatus (Massasauga), with emphasis on Sistrurus catenatus edwardsii (Desert Massasauga). J. Herpetol., 36:454 464., C. W. PAINTER, D. G. BARKER AND M. E. DOUGLAS. 2002. Foraging ecology of the threatened New Mexico Ridge-nosed Rattlesnake (Crotalus willardi obscurus), p. 243 251. In: G. W. Schuett, M. Hoggren, M. E. Douglas and H. W. Greene (eds.). Biology of the vipers. Eagle Mountain Publishing, Eagle Mountain, Utah. JOHNSON. G. 1995. Spatial ecology, habitat preference, and habitat management of the eastern massasauga, Sistrurus c. catenatus in a New York weakly-minerotrophic peatland. Ph.D. Dissertation, State University of New York, Syracuse. 222 p. KEENLYNE, K. D. AND J. R. BEER. 1973. Food habits of Sistrurus catenatus catenatus. J. Herpetol., 7:382 384. MADSEN,T.AND R. SHINE. 2000. Silver spoons and snake body sizes: prey availability early in life influences long-term growth rates of free-ranging pythons. J. Anim. Ecol., 69:952 958. MARTINS, M., O. A. V. MARQUES AND I. SAZIMA. 2002. Ecological and phylogenetic correlates of feeding habits in the neotropical pitvipers of the genus Bothrops, p. 307 328. In: G. W. Schuett, M. Hoggren, M. E. Douglas and H. W. Greene (eds.). Biology of the vipers. Eagle Mountain Publishing, Eagle Mountain, Utah. MAUGER, D. AND T. P. WILSON. 1999. Population characteristics and seasonal activity of Sistrurus catenatus catenatus in Will County, Illinois: implications for management and monitoring, p. 110 124. In: B. Johnson and M. Wright (eds.). Second international symposium and workshop on the conservation of the Eastern Massasauga Rattlesnake, Sistrurus catenatus catenatus: population and habitat management issues in urban, bog, prairie, and forested ecosystems. Toronto Zoo, Toronto. MULCAHY, D. G., J. R. MENDELSON, III, K. W. SETSER AND E. HOLLENBECK. 2003. Crotalus cerastes. Prey/ predator weight-ratio. Herpetol. Rev., 34:64. MUSHINSKY, H. M. 1987. Foraging ecology, p. 302 334. In: R. A. Seigel, J. T. Collins and S. S. Novak (eds.). Snakes: ecology and evolutionary biology. McGraw-Hill, N.Y.

368 THE AMERICAN MIDLAND NATURALIST 152(2), J. J. HEBRARD AND D. S. VODOPICH. 1982. Ontogeny of water snake foraging ecology. Ecology, 63:1624 1629. PARKER, W. S. AND M. V. PLUMMER. 1987. Population ecology, p. 253 301. In: R. A. Seigel, J. T. Collins and S. S. Novak (eds.). Snakes: ecology and evolutionary biology. McGraw-Hill, N.Y. PAULY, G. B. AND M. F. BENARD. 2002. Crotalus viridis oreganus. Costs of feeding. Herpetol. Rev., 33:56 57. PHILLIPS, C. A., R. A. BRANDON AND E. O. MOLL. 1999. Field guide to amphibians and reptiles of Illinois. Ill. Nat. Hist. Surv. Manual, 8:1 300. POUGH, F. H. AND J. D. GROVES. 1983. Specializations of the body form and food habits of snakes. Amer. Zool., 23:443 454. REISERER, R. S. 2002. Stimulus control of caudal luring and other feeding responses: a program for research on visual perception in vipers, p. 361 384. In: G. W. Schuett, M. Hoggren, M. E. Douglas and H. W. Greene (eds.). Biology of the vipers. Eagle Mountain Publishing, Eagle Mountain, Utah. REYNOLDS, R. P. AND N. J. SCOTT, JR. 1982. Use of a mammalian resource by a Chihuahuan snake community, p. 99 118. In: N. J. Scott, Jr. (ed.). Herpetological communities. U.S. Department of the Interior, Fish and Wildlife Service, Washington, D.C. ROTH, E. D., P. G. MAY AND T. M. FARRELL. 1999. Pigmy rattlesnakes use frog-derived chemical cues to select foraging sites. Copeia, 1999:772 774. SAZIMA, I. 1992. Natural history of the jararaca pitviper, Bothrops jararaca, in southeastern Brazil, p. 199 216. In: J. A. Campbell and E. D. Brodie, Jr. (eds.). Biology of the pitvipers. Selva, Tyler, Tex. SCHALL, J. J. AND E. R. PIANKA. 1978. Geographical trends in numbers of species. Science, 201:679 686. SCHUETT, G. W., D. L. CLARK AND F. KRAUS. 1984. Feeding mimicry in the rattlesnake Sistrurus catenatus, with comments on the evolution of the rattle. Anim. Behav., 32:625 626. SEIGEL, R. A. 1986. Ecology and conservation of an endangered rattlesnake, Sistrurus catenatus, in Missouri, USA. Biol. Conserv., 35:333 346. SHINE, R., W. R. BRANCH, P. S. HARLOW AND J. K. WEBB. 1998. Reproductive biology and food habits of horned adders, Bitis caudalis (Viperidae), from southern Africa. Copeia, 1998:391 401. SMITH, P. W. 1961. The amphibians and reptiles of Illinois. Ill. Nat. Hist. Surv. Bull., 28:1 298. SOKAL, R.R.AND F. J. ROHLF. 1995. Biometry, 3rd ed. Freeman, N.Y. 887 p. SZYMANSKI, J. 1998. Status assessment for the eastern massasauga (Sistrurus c. catenatus). U.S. Fish and Wildlife Service, Endangered Species Division. Fort Snelling, Minn. 64 p. U.S. FISH AND WILDLIFE SERVICE. 1999. Endangered and threatened wildlife and plants; review of plant and animal taxa that are candidates or proposed for listing as endangered or threatened; annual notice of findings on recycled petitions; and annual description of progress on listing actions. Federal Register, 64:57534 57547. SUBMITTED 18 FEBRUARY 2003 ACCEPTED 20 FEBRUARY 2004