ILLINOIS NATURAL HISTORY SURVEY

Natural History of the Bird-voiced Treefrog {Hyla avivoca) and Green Treefrog {Hyla cinerea) in Southern lUinois

Michael Redmer Lauren E. Brown Ronald A. Brandon

Illinois Natural History Survey Bulletin Volume 36, Article 2 September 1999 Illinois Natural History Survey, David L. Thomas, Chief A Division of the Illinois Department of Natural Resources

A catalog of the publications of the Illinois Natural History Survey is available without charge from the address below. A price list and an order blank are included with the catalog.

Illinois Natural History Survey Distribution Center Natural Resources Building 607 East Peabody Drive Champaign, Illinois 61820

Citation: Michael Redmer, Lauren E. Brown, and Ronald A. Brandon. 1999. Natural History of the Bird-voiced Treefrog {Hyla avivoca) and Green Treefrog {Hyla cinerea) in Southern Illinois. Illinois Natural History Survey Bulletin 36(2):37-66.

Editor: Thomas E. Rice

Michael Redmer is a biologist and freelance photographer. He completed work on this project while a graduate student in the Department of Zoology, Southern Illinois University at Carbondale 62901-6501.

Lauren E. Brown is Professor of Vertebrate Zoology and Curator of Amphibians and Reptiles, Department of Biological Sciences, Illinois State University, Campus Box 4120, Normal 61790-4120. Ronald A. Brandon

is Professor Emeritus and Curator of Herpetology, Department of Zoology, Southern Illinois University at Carbondale 62901-6501. Brandon and Brown are Affiliate Professional Scientists, Center for Biodiversity, Illinois Natural History Survey.

US ISSN 0073-4918

Printed by Authority of the State of Illinois (RR46854-1M-9-99) ILLINOIS NATURAL HISTORY SURVEY

Natural History of the Bird-voiced Treefrog {Hyla avivoca) and Green Treefrog {Hyla cinerea) in Southern Illinois

Michael Redmer Lauren E. Brown Ronald A. Brandon

Illinois Natural History Survey Bulletin Volume 36, Article 2 September 1999 Dedication

This study is dedicated to the memory of our colleague, the late Dr. Michael A. Morris, with whom LEB initiated this project in the late 1980s as Mike completed his graduate work at SIUC. Mike was a foremost expert on the herpetofauna of Illinois and was associated with the Illinois Natural History Survey several times during his short career. Contents

Acknowledgments iv Introduction 37 Materials and Methods 37 Historical Records 37 Current Distribution 37 Habitat 37 Results 40 Hyla avivoca 40 Description 40 Comparison With Gray Treefrogs 40 Distribution 40 Habitat 43 Hyla cinerea 46 Description 46 Distribution 46 Range Expansion 49 Habitat 52 Discussion 53 Biogeography 53 Range Expansion of Hyla cinerea 54 Conservation Biology 56 Summary 57 Literature Cited 58 Appendix 64 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Acknowledgments

For assistance with field work or for alerting us Malnate (Academy of Natural Sciences of to localities, we thank S. Ballard, M. Bavetz, Philadelphia); R. Vasile (Chicago Academy of

M. Blanford, D. Brandon, E. Brown, J. Brown, Sciences); R. Beer (St. Louis Science Center); L. N. Brown, L. Buckley, B. Burke, M. and D. Rossman (Louisiana State University

DeJesus, M. Heafner, D. Jedele, J. Karnes, S. Museum of Natural Science). R. Crombie and

Karsen, J.S. Keogh, M. MacCallum, W. R. Reynolds provided extensive information on McDowell, E. Mockford, H. Moeller, M. E. Cope and specimens in the U.S. National Morris, G. Paukstis, M.A. Phillippi, M. Museum of Natural History collected in St. Peterson, S. Reilly, D. Shepard, M. Smitka, K. Louis. B. Wainscott and P. Foley provided

Tolch, and A. Wilson. R. Heidinger and J. information on toad tadpoles that arrived in a

Stahl provided impoundment dates for some shipment of fish at the Aquaculture Facility, reservoirs. J. Parmelee alerted us to KU Illinois State University. W. Lewis provided specimens from the introduced locality in information on tadpoles in his fish hatchery. L. Kansas, and examined the specimens. P. Reif Massarolo gave information on tadpoles in fish shared his observations of H. cinerea and shipments to a pet store. R. Axtell gave tadpoles at his fish farm. K. Davie (SIUC) and information on his field experiences in Madi-

J. Miller (U.S. Forest Service) helped with the son County, Illinois. T. Schiebel provided help preparation of range maps. Permits to collect with computer software. R. Goellner searched specimens on lands under their control were the archives of the Naturalists Club (St. Louis) issued by the following agencies: Illinois for J. Hurler's field notes. R. Axtell and T. Department of Natural Resources (Divisions of Johnson facilitated the search for the specimens

Wildlife Resources and Natural Heritage, and and field notes of J. Hurler. A. Resetar Illinois Nature Preserves Commission); United provided photocopies of correspondence States Fish and Wildlife Service (Crab Orchard between F. Cagle and K. Schmidt in the and Cypress Creek National Wildlife Refuges); archives of the Field Museum of Natural and the United States Forest Service (Shawnee History. J. Herkert provided information on the National Forest). Permission was granted to approval of the listing of H. avivoca as a LEB to reside in Pine Hills Swamp during threatened species by the Illinois Endangered several periods in the earlier years of this study Species Protection Board. R. Gray, G. Paukstis, by J. Parsons of the Pine Hills Field Station and and D. Shepard critically reviewed the manu- W.C. Ashby. script. H.M. Smith provided encouragement. The following provided loans or informa- Portions of the field work were funded by tion or both on holdings in their museums or Illinois Department of Natural Resources collections: H. Marx, A. Resetar, and H. Voris contracts with RAB, and a Nature Conservancy (Field Museum of Natural History); S. Minton contract with Southern Illinois University at (Minton Herpetological Collection); L. Page Carbondale. A grant to the late W. Klimstra and C. Phillips (Illinois Natural History provided funds for some of the field work by

Survey); J. Purdue (Illinois State Museum); A. LEB. Publication costs were funded by Illinois Kluge and G. Schneider (University of Michi- State University (facilitated by T. Cheung, S. gan Museum of Zoology); W. Duellman Little, and P. Schollaert); the Department of (University of Kansas Museum of Natural Zoology, Southern Illinois University at History); R. McDiarmid, R. Reynolds, and G. Carbondale (facilitated by W. Muhlach); and Zug (United States National Museum of the Illinois Natural History Survey (facilitated Natural History); C. Phillips (University of by C. Phillips). Illinois Museum of Natural History); E. Natural History of the Bird- voiced Treefrog {Hyla avivoca) and Green Treefrog {Hyla cinerea) in Southern IlHnois

The bird-voiced treefrog, Hyla avivoca Viosca (MAM), Minton Herpetological Collection

1928, Figures 1 and 2, and green treefrog, Hyla (SAM), Southern Illinois University at cinerea (Schneider 1799), Figures 3 and 4, are Carbondale Fluid Vertebrate Collection distributed primarily on the Atlantic and Gulf (SIUC), United States National Museum of coastal plains of the southeastern United States Natural History (USNM), University of Illinois (Conant and Collins 1998). The ranges of both Museum of Natural History (UIMNH). and species reach their northern limits in the University of Michigan Museum of Zoology Midwest in southern Illinois where Smith (UMMZ). See the Appendix for a detailed list

( 1961 ) recorded them from Alexander, Jackson, of localities and preserved specimens exam- and Union counties. Because the few docu- ined. mented localities ofthe.se treefrogs in Illinois were, until recently, mainly in or near remnant Current Distribution Austroriparian swamplands (which are disap- Current distributions were determined in the pearing rapidly), concern has been expressed field between 1961 and 1998 by listening for that their continued existence in the state is in the species-specific mating calls and looking jeopardy (Ackerman 1975; Ashton et al. 1976; for individuals in vegetation and on roads in the

Dyrkacz 1974). Since Smith's ( 1961 ) compre- vicinity of historic localities and other pre- hensive study The Amphibians and Reptiles of sumed suitable habitats. Most field work was

Illinois, few additional records for these species carried out between May and August during have been reported. Carton and Brandon and shortly after the chorusing seasons: May to (1975) studied reproductive ecology and habitat mid-July for H. avivoca and mid-May to early of//, cinerea at a southern Illinois swamp, but August for //. cinerea (this study; Garton and there has been no previous in-depth environ- Brandon 1975). Attempts usually were made mental examination of H. avivoca in Illinois. to collect one or two voucher specimens

The objective of this publication is to report the (deposited at SIUC; Appendix). results of our study of the natural history of these two poorly known treefrogs in southern Habitat

Illinois. Wherever we heard //. avivoca calling at night, we visited the localities again by day and Materials and Methods recorded information on dominant tree species, including their condition (alive or dead), and and perch Historical Record.s understory species. In 1996 1997, calling male //. avivoca were recorded Historical distributions were documented from sites of five locations: Heron Pond (NW 1/4 sec. 30, published literature, unpublished reports, and at South (SE 1/4 museum specimens. Specimens were exam- T13S, R3E), Hogan's Bottoms R2E), Hogan's Bottoms North ined from the Field Museum of Natural History sec. 23, T13S, 1/4 sec. 23, T13S, R2E), Bell Pond (NE 1/ (FMNH), Illinois Natural History Survey (NE R4E). and Reevesville Swamp (INHS), Illinois State Museum (ISM), Illinois 4 sec. 14, T13S, l/4sec. 31,T13S, R5E). Type of perch, State University (ISU), University of Kansas (NW depth of water below each Museum of Natural History (KU), Lauren E. taxon if plant, and Descriptions of habitat Brown Frog Collection (LEB), Louisiana State perch were recorded. are based on our field observa- University Museum of Natural Science for H. cinerea (LSUMZ). Michael A. Morris Collection tions and information from the literature. 38 Illinois Natural History Sur\ev Bulletin Vol. 36 Art. 2

Figure 1. Female bird-voiced treefrog (Hyla avivoca) from Hogan"s Bottoms (Cypress Creek drainage). Union County, Illinois. Photo by M. Redmer.

Figure 2. Tadpole of Hyla avivoca from Reevesville Swamp,

Pope County, Illinois. Photo by M. Redmer. September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 39

Figure 3. Calling male green treefrog (Hyla cinerea) from Mermet Lake Conservation Area, Massac County, Illinois. Photo by M. Redmer.

Figure 4. Tadpole of Hyla cinerea from Reevesville Swamp, Pope County, Illinois. Photo by M. Redmer. 40 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Results Le Conte 1 825 in coloration and habitus. However, two features of live H. avivoca

reliably distinguish them from //. chrysoscelis Hyla avivoca and H. versicolor: (1) the mating call of//. avivoca consists of a series of birdlike whistles Description whereas the mating calls of//, chrysoscelis and Hyla avivoca is a moderately slender, small H. versicolor are trills with fast pulse rates; and treefrog. Maximum reported snout-vent length (2) the light green flash coloration on concealed (SVL) is 52 mm (Conant and Collins 1998). surfaces of the thighs of //. avivoca contrasts Males average smaller than females. The head with yellow to orange flash coloration on the is short with a truncate muzzle. The eyes are thighs of //. chrysoscelis and //. versicolor. large and protuberant. Males have a subgular Other characters sometimes are helpful in vocal pouch. The toe pads are conspicuous. distinguishing //. avivoca from //. chrysoscelis The subarticular pustule of the fourth digit on and //. versicolor but do not provide absolute the hand often (but not always) is divided by a identification. For example, //. avivoca has a furrow (Smith 1953). The hind toes are smaller size at maturation and smaller average extensively webbed. There are a tarsal fold and aduh size (Mecham 1960; Smith 1953, 1961). two metatarsal tubercles (one small, one large) Also, //. avivoca has slightly smoother skin, on each hind limb. The hand has one large (at and the subarticular pustule of the fourth digit the base of the thumb) and a number of small of each hand often is divided by a furrow palmar tubercles. A fold of skin extends across (Smith 1953, 1961, 1966). This pustule more the chest between the axillae. Another fold often is undivided in //. chi-ysoscelis and H. occurs across the dorsal aspect of the wrist. versicolor. The tadpole of //. avivoca is, Dorsally the skin is relatively smooth, but however, distinctive (see "Description") and ventrally it is granular. The background color contrasts with tadpoles of //. chrysoscelis and of the dorsum is green, brown, or most fre- //. versicolor, which are olive green dorsally, quently, various shades of gray. The venter is white or cream ventrally, and sometimes have white or cream. Usually there is one large, red or orange tails that lack bars (Altig 1970; irregular-shaped, dark blotch on the dorsum. Wright 1914). One or sometimes two irregular dark spots occur between the eyes. A white spot (edged in Distribution black) is beneath each eye. The fore and hind Hyla avivoca occurs in isolated colonies along legs have dark bars dorsally. A light green the drainages of the Big Muddy River, Missis- flash coloration occurs in the groin and on the sippi River, Cache River, and Bay Creek in posterior surface of the thighs. Alexander, Johnson, Pope, Pulaski, Union, and The distinctive tadpole of//, avivoca (Fig. possibly Jackson counties of extreme southern 2) was characterized by Altig (1970), Hellman Illinois (Fig. 5, Appendix). However, before (1953), Parker (1951), Redmer (1998), and Smith's (1953, 1961) work, there was a Volpe et al. (1961). It has a mostly black body question as to whether this species occurred in and tail with bold red, orange, or gold markings Illinois because of confusing nomenclature and (saddles on the tail, interorbital and orbitonasal strong resemblance of //. avivoca to frogs of stripes). the //. versicolor complex. Thirty-four years The mating call of H. avivoca consists of a after Cope (1889) described Hyla versicolor series of birdlike whistles. It was analyzed in phaeocrypta from Mt. Carmel, Wabash detail by Blair (1958, 1959). County, Illinois, Viosca (1923) described a distinct gray treefrog with a whistling call from Comparison With Gray Treefrogs Louisiana, which he tentatively identified as There is a strong resemblance between //. Hyla phaeociypta. However, he noted that avivoca and the gray treefrogs Hyla information on the mating call of Cope's chrysoscelis Cope 1880 and Hyla versicolor September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 41

(1889) frog was lacking, so direct comparison Parker 1 95 1 ; and others) referred to the bird-

with the frogs from Louisiana was not possible. voiced treefrog as //. phaeocrypta under the Ridgway (1924) reported hearing birdlike assumption that Cope's (1889) specimen from whistles near Mt. Carmek and speculated that Mt. Carmel, Illinois, was conspecific and had this might have been the call of Cope's H. priority. Smith (1953) clarified the identify of phaeocrypta. Viosca (1928) reconsidered his Cope's specimen as a member of the H. earlier report (1923) and named his treefrog as versicolor complex and demonstrated the a new species, Hyla avivoca. He designated proper application of//, avivoca as a distinctive the type locality as Mandeville, Louisiana treefrog, whose range included far southern (holotype USNM 75017), and also identified a Illinois, well south of Mt. Carmel. sample of 15 specimens of//, avivoca collected Cagle (1942) reported a single specimen of

in 1907 from Olive Branch (= Horseshoe //. avivoca (FMNH 25940) from Little Grand Lake), Alexander County, Illinois (FMNH Canyon, Jackson County. This frog was sent to

2201 ). We located and examined three of the Karl P. Schmidt at the Field Museum of specimens and confirm identification as H. Natural History, Chicago, for identification

avivoca. Nevertheless, a number of workers (Cagle 1937). Schmidt ( 1937) identified the (Edgren 1953; Mittleman 1945; Neill 1948; specimen (probably on the basis of its small

10 20 30 Kilometers

Figure 5. Distribution of Hyla avivoca in Illinois. Solid circles represent localities documented by voucher specimens examined or collected by us; triangles represent localities where we heard and/or saw specimens but collected no vouchers. Some plotted localities represent two or more nearby collection sites. 42 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

size) as H. avivoca, but added that it "requires H-4741). In 1996 we found large populations the confirmation of additional specimens, and in two small swamps east of Reevesville, Pope especially the check of hearing its peculiar County (Redmer 1998) ca. 13 km downstream birdlike whistle." We examined FMNH 25940; (ca. 6 km straight-line distance) from Bell it has undivided subarticular pustules on the Pond, and collected vouchers (SIUC H-5251- fourth digits of the hands (as often is character- 5253, H-5383-5385, H-5543, H-5598). istic of the H. versicolor complex), and is a Most of the known records in Illinois are in sexually immature male with a SVL of 31 mm. the drainage of the Cache River, including

Mean SVL for 52 sexually mature preserved three of its tributaries (Cypress Creek, Dutch- male H. avivoca from La Rue-Pine Hills man Creek, Limekiln Slough). Hyla avivoca

SwampAVolf Lake is almost the same at 3 1 .9 was reported earlier in this drainage by mm (range = 28-37). whereas the mean SVL Klimstra and Hutchison (1965), Evers and Page for 41 sexually mature preserved male H. (1977), and Brandon and Morris (1987, 1988). chrysoscelis from various localities in southern Localities have been found from as far up- Illinois (the only species of the H. versicolor stream as Cypress Pond State Natural Area at complex currently known to occur in the the Union/Jackson county line near the town of region; Appendix) is 36.4% greater at 43.5 mm Mt. Pleasant (Appendix), and as far down- (range = 38-49 mm). The two latter cohorts are stream as east and south of Tamms, Alexander obviously different because their ranges do not County (Burbrink et al. 1995, 1998). Along the overlap. If FMNH 25940 were a H. avivoca, it nearly 80 km of stream between these sites, should be sexually mature, which it is not. suitable habitat is fragmented by drainage Thus, on the basis of size, sexual immaturity, projects, channelization, timber clearing, and and undivided subarticular pustules on the natural topography. In Heron Pond-Little fourth digits of the hands, we conclude that Black Slough State Natural Area, H. avivoca FMNH 25940 was misidentified, and actually may be continuous over a broad expanse of is a male of the H. versicolor complex (prob- baldcypress (Taxodium distichum)/tupe\o ably H. chrysoscelis). However, it is quite (Nyssa aquatica) swampland. Because much probable that H. avivoca does occur just west of this natural area is relatively remote and of Little Grand Canyon at Oakwood Bottoms inaccessible, and part of it (Heron Pond) is (see information that follows). protected as an Illinois Nature Preserve,

Between 1961 and 1991, //. avivoca was preserved specimens from this 6,000-hectare reported or documented from Alexander, area are uncommon in museum collections. In Johnson, Pulaski, and Union counties (Brandon addition to four sites represented by voucher and Morris 1987; Brown and Pierce 1965; specimens, we heard choruses at two additional

Klimstra and Hutchison 1965; Smith 1961, sites within this swamp. 1966). Since then, we have found a number of Hyla avivoca has been collected infre- populations in Alexander, Jackson, Johnson, quently at Horseshoe Lake, Alexander County.

Pope, Pulaski, and Union counties (Fig. 5; We located and examined three of the speci- Appendix). Brandon and Ballard (1997) mens of//, avivoca (FMNH 2201) that Viosca inadvertently included Massac and Williamson (1923, 1928) reported from Olive Branch counties within the known range of H. avivoca. (actually collected at Horseshoe Lake [Viosca However, there are no known vouchers, and we 1923]) and agree with his identification of them have been unable to locate choruses. as H. avivoca. An immature specimen (SVL =

In the Bay Creek drainage, H. avivoca is 26 mm; LSUMZ 51046) collected from common at Bell Pond {= Grantsburg Swamp of Horseshoe Lake by D. Rossman in 1957 was Mohlenbrock 1991), although previously identified by him as H. avivoca. No informa- recorded there only by Brandon and Morris tion on flash coloration is available for this

(1987). We heard large choruses from 1993 specimen, nor is it known whether Rossman through 1996 and collected a voucher (SIUC heard breeding choruses. However, the September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 43

subarticular pustules on the fourth digits of the Hills SwampAVolf Lake, was described in hands are undivided (more often characteristic detail by Mohlenbrock (1959). Voigt and of the H. versicolor coxn\i\

Horseshoe Lake over his many years of field and Mohlenbrock" s (1964) map showing

work there dating back to 196 1 . Furthermore, distribution of these deep swamps in Illinois intensive field work in the area from 1986 to and Mohlenbrock and Ladd's (1978) map of 1996 (Brandon 1994; Brandon and Morris the range of M aqiiatica in Illinois both roughly 1987, 1988; Brandon and Wilson 1996; parallel the range of H. avivoca. Other trees

Heidinger et al. 1994, 1995) did not uncover commonly associated with these swamps are the species in what is now greatly disturbed water hickory {Caiya acjuatica), pumpkin ash habitat (discussion follows). Thus, H. avivoca (Fraxiims tomentosa), water locust (Gleditsia may be extirpated at Horseshoe Lake. aquatica), and swamp red maple (Acer In the drainage of the Big Muddy River, H. drummondi). Common understory plants avivoca is common and well documented in the include buttonbush (Cephalanthus

literature at La Rue-Pine Hills SwampAVolf occidentalis), swamp rose {Rosa palustris), and Lake, Union County (Ballard 1994; Brown and swamp loosestrife (Decodon verticilatus). Pierce 1965; Evers and Page 1977; Minton and While baldcypress and tupelo gum commonly Minton 1948; Rossman 1960; and Smith 1953, occur together in these swamps, mature stands 1961). In 1992. one ofus (MR) heard a calling of one species often nearly exclude the other.

male at Worthen Bayou, Oakwood Bottoms- Voigt and Mohlenbrock (1964) noted that Turkey Bayou Waterfowl Management Area, baldcypress-tupelo gum communities or tupelo approximately 1.5 km southwest of Cagle's gum alone dominate in deep water, and that

( 1942) Little Grand Canyon locality. Michael baldcypress-pumpkin ash communities Morris (pers. comm.) also heard H. avivoca at dominate where water is less than 0.3 m deep. Oakwood Bottoms. We have heard none in Little has been reported previously about recent years, but flooding during late May and the habitats of//, avivoca in Illinois. During

early June prevented visits to the site in 1995 field work in 1996 and 1997 (when all known and 1996. localities in Illinois were examined), we found An immature specimen (SVL = 22 mm; large choruses (>10 or more individuals LSUMZ 51047) was collected in 1957 by D. calling) only in pure or mixed stands of Rossman at Union County Conservation Area baldcypress and tupelo gum (Fig. 6A), or in and identified by him as H. avivoca. This frog adjacent reed thickets {Phragmites australis.

is faded and the subarticular pustules on the Fig. 6B). Where the swamp canopy was dense fourth digits of the hands are undivided (more and the understory diverse and well shaded by often characteristic of the H. versicolor day, H. avivoca usually was the dominant complex). No information on flash coloration calling anuran (other species calling included

or mating call is available, so we cannot justify cricket frogs, Acris crepitans; bullfrogs, Rana

identifying it to species. We visited Union catesbeiana; green frogs, Rana clamitans; and County Conservation Area many times during occasional H. cinerea). Few calling H. avivoca 1991-1997, but never heard H. avivoca. (< 5 individuals) were present in more open shrubby wetlands (Fig. 7A), or where Habitat baldcypress and tupelo gum were sparse, We found H. avivoca only in or on the edges of immature, or appeared stressed or dying (Fig. remnant swamps dominated by baldcypress or 7B). Sometimes none were seen or heard in tupelo gum along rivers and streams. An such open areas on nights when large choruses example of this type of habitat. La Rue-Pine were heard at adjacent shaded swamps. 44 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Figure 6. Habitats of Hyla avivoca in Illinois as they appeared in August. A (Top): Mature baldcypress-tupelo swamp 1.5 km NE Reevesville, Pope County. B (Bottom): Reed thicket adjacent to the above swamp (this was the only place where tadpoles of H. avivoca were collected during this study). Photos by M. Redmer. September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 45

\

\

" \

Figure 7. Habitats of Hyla avivoca and Hyla cinerea in Illinois as they appeared in August. A (Top): Open-water buttonbush-cattail swamp adjacent to mature baldcypress-tupelo swamp 1.5 km NE Reevesville. Pope County. B (Bottom): Declining baldcypress-tupelo swamp at Hogan's Bottoms 3.4 km W Cypress, Union County. Photos by M. Redmer. 46 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Usually H. cinerea was dominant in these open Hyla cinerea wetlands where C. occidentalis was common. Fifty-five male H. avivoca called from ten Description identified species of plants, dead logs (not to Hyla cinerea is a moderately large treefrog species), and the metal rail of an interpretive with a slender, elongate body. Maximum trail. Perch species or type (and frequency of reported SVL is 64 mm (Conant and Collins use) were as follows: N. aquatica (23.6%); T. 1998), with males averaging smaller than distichum (21.8%); C. occidentalis {\6A%)\ A. females. The head is flat and the snout is dmmmondi (9.1%); Rosa palustris (7.3%); P. pointed. Males have a subgular vocal pouch. australis (7.3%); metal rail (5.5%); F. The toe pads are conspicuous and the hind tomentosa (3.6%); Virginia willow, Itea limbs are long. Extensive webbing occurs virginica (3.6%); and floating log (1.8%). between the hind toes. There are two metatar-

Fifty-three frogs were 0.1-2.3 m (mean = 1.5 sal tubercles (one large, one small) and a tarsal m) above water. Two perched on vegetation fold on each hind limb. On each hand there are rooted in soil along the swamp margin. One one large (at the base of the thumb) and a frog was observed calling on the trunk of a number of small palmar tubercles. A fold of tupelo 3 m above water. (On other occasions skin occurs across the chest between the we observed H. avivoca calling on perches >3 axillae. Another fold extends dorsally across m above water.) Water depths below 53 the wrist. Dorsally the skin is smooth but perch sites were 0.1-0.7 (mean = emergent m ventrally it is granular. The dorsum usually is 0.3 m). bright green, but sometimes olive or yellow- At Reevesville Swamp, 13 tadpoles of H. green. Small gold flecks that frequently have a avivoca were collected on 7 and 10 July 1997. black border often are scattered on the dorsum. All were netted from shaded water (< 0.3 m A white lateral stripe extends from the snout, deep) below the interior of a thicket of P. beneath the eye, to midbody or nearly the to australis. Other attempts collect tadpoles of groin. This stripe sometimes is broken but unsuccessful, despite H. avivoca were repeated rarely is absent. The venter is white or cream. daytime searching at chorus sites and known Dickerson (1907, p. 127) wrote: "The Green habitats (especially around bases of tupelo and Tree Frog is perhaps the most beautiful tree baldcypress trees). frog in North America." We occasionally found recently metamor- The tadpole of H. cinerea (Fig. 4) has phosed juveniles in late summer or early yellow orbitonasal stripes, a greenish body after the autumn, breeding season. Most were (dorsally and laterally), yellow tail with dark observed perched in low (< 2 m) shrubs in reticulations, and yellow or buff venter (Altig woodlands adjacent to swamps where breeding 1970; Wright 1932). choruses had been observed previously. On 25 The loud and distinctive mating call (Blair April five subadults, 1996, MR observed one 1958, 1959) of H. cinerea is best described as a adult male, and an adult female prior to the " nasal "frank, frank, frank. . . Large choruses breeding season, all perched on low emergent (e.g., at La Rue-Pine Hills SwampAVolf Lake) vegetation and on trunks of baldcypress and produce a tremendous cacophony that can be tupelo gum trees. However, on 30 September heard 1-2 km or more away. 1995 and 20 September 1996, LEB and MR found one adult and several subadults on the Distribution ground and perched in vegetation in the low Hyla cinerea is widely distributed across much forest adjacent to Heron Pond. LEB found of extreme southern Illinois in Alexander, terrestrial subadult H. avivoca adjacent to Jackson, Johnson, Massac, Pope, Pulaski, Reevesville Swamp and La Rue-Pine Hills Randolph, Union, and Williamson counties on Swamp 26 September 1998. Rossman (Fig. 8; Appendix). The first record in southern also (1960) noted the terrestrial tendencies of Illinois (no specific locality) probably was the species in autumn at Pine Hills Swamp. September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 47

provided by Davis and Rice (1883), who (although it was documented there by museum identified them as squirrel treefrogs Hyla specimens collected as early as 1980) and squi reiki (a similar species found in the Randolph County (Fig. 8; Appendix). southeastern U.S.). Carman (1890. 1892) In the late nineteenth century, two localities subsequently reported them from Bluff Lake, were reported for H. cinerea far north in the

Union County. vicinity of St. Louis. Because these northern

Additional valid records (Fig. 8; Appendix) records are of highly questionable validity, it is of H. cinerea in Illinois include La Rue-Pine pertinent to discuss them in detail. Hills SwampAVolf Lake, Union County A single specimen of "Hyla Carolinensis"

(Ballard 1994; Brown and Pierce 1965; Cagle (= H. cinerea) was found at "Cantine [sic]

1942; Carton and Brandon 1975; Minton and Creek, Cantine [sic], Madison County, 111." by Minton 1948; Rossman 1960; Smith 1961). the pioneer Missouri herpetologist Julius Hurter several localities in Johnson County (Brandon (1893), who spent considerable time in the and Morris 1987; Klimstra and Hutchison vicinity of St. Louis. He did not mention this

1965; Phillippi et al. 1986), in Pulaski County specimen in later publications (1897, 1903,

(Brandon and Morris 1987). and in Massac, 191 1) on the herpetofauna of Missouri, but

Pope, and Williamson counties (Redmer and there is no doubt that he was familiar with H.

Ballard 1995). In all of these counties we cinerea, as evidenced by his collection of other found numerous additional localities. We specimens of the species (USNM 57732, report it for the first time in Jackson County 57736) in Alabama and Louisiana in 1892. We

Figure 8. Distribution of Hyla cinerea in Illinois. Solid circles represent localities documented by voucher specimens examined or collected by us; triangles represent localities where we heard and/or saw specimens but collected no vouchers. Some plotted localities represent two or more nearby collection sites. 48 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

contacted T. Johnson (an authority on the Cope (1889) reported two specimens of//. herpetofauna of Missouri), who indicated that carolinensis (both under USNM 1070) col-

most of Hurter's specimens were deposited in lected by George Engelmann from St. Louis.

the U.S. National Museum of Natural History, This collection locality is pertinent to our study

with some going to the Chicago Academy of because it lies just across the Mississippi River Sciences and the Academy of Science of St. from Madison County, Illinois. R. Crombie Louis (antecedent of the St. Louis Science and R. Reynolds of the U.S. National Museum Center). Johnson also indicated that Hurter's of Natural History provided us with the field notes may be located at these institutions following information pertaining to USNM or the Naturalists Club in St. Louis. No 1070 (as well as subsequent specimens dis- specimens of H. cinerea from Illinois collected cussed). Cope had the bad habit of cutting off by Hurter or any of his pertinent field notes are the tags of hundreds of specimens he borrowed located in the Chicago Academy of Sciences, from the U.S. National Museum of Natural St. Louis Science Center, or Naturalists Club History. He later reassociated tags and (R. Vasile, R. Beer, R. Goellner, pers. comm.). specimens, some correctly, many incorrectly. However, the U.S. National Museum of Natural After Cope died, many specimens were History has a specimen (USNM 57827) returned to the U.S. National Museum of

collected by Hurter on 1 May 1 892 in Madison Natural History without tags. To further

County, Illinois. (No additional information is complicate the issue, in the earlier years at the

available on the locality [G. Zug, pers. U.S. National Museum of Natural History it comm.].) We have examined this juvenile male was a standard practice to assign a single (SVL = 25 mm). Although the color pattern is catalogue number to a lot of specimens from a faded, white spots can still be detected under single collector. When a lot was recognized as the eyes, and the subarticular pustules on the containing more than one species, the extra fourth digits of the hands are undivided (as is species was recatalogued. Presently USNM often characteristic of the H. versicolor 1070 is assigned to a water snake {Nerodia) as complex). We conclude it is a member of the are all those numbers surrounding 1070. H. versicolor complex. {Hyla avivoca also has Interestingly, 1070 was received from Engel-

white spots under the eyes, but the northern mann and was from St. Louis. There is nothing collection locality makes it unlikely that to suggest (other than Cope 1889) that 1070 USNM 57827 is of that species.) Young frogs was ever associated with any frogs. No further of the H. versicolor complex frequently are information is available on 1070 or any other light green dorsally (Dickerson 1907), and possibly associated frogs. superficially similar to H. cinerea. Thus, Other frogs of pertinence in the U.S. Hurter (1893) may have misidentified the National Museum of Natural History include young specimen of the H. versicolor complex USNM 3667 (four specimens) and 104553 (one (USNM 57827) as ''Hyla Carolinensis'' {= H. specimen), which were collected by Engelmann cinerea). We contacted R. Axtell, herpetologist from St. Louis and entered into the catalogue in at Southern Illinois University at Edwardsville, 1858. These specimens are all of the H. about the possibility of H. cinerea occurring in versicolor complex. USNM 3667 is listed by Madison County. In his many years of field Cope (1889) under H. versicolor, but as one experience in that area he never collected any, specimen rather than four. USNM 104553 nor is he aware of anyone else who has originally was catalogued as USNM 3453 (also encountered the species in Madison County. In listed by Cope [1889] as H. versicolor). June 1995 and July 1997 we searched for H. USNM 3453 was a lot also containing two cinerea in the vicinity of Canteen in southern wood frogs {Rana sylvatica), which retained Madison County and St. Clair County. None the original number, while the member of the were heard or seen, but a small chorus of H. H. versicolor complex was recatalogued in

versicolor was heard near Canteen Creek. 1938 as USNM 104553. There is no reason to September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 49

assume that USNM 3667 and/or 104553 were Creek and the Big Muddy River. Thus, H. the specimens Cope (1889) refeired to as H. cinerea from Drury Creek may have colonized carolinensis (= H. cinerea) USNM 1070 from Carbondale during the past 20 years. At least St. Louis. Furthermore, Johnson (1987) found one Austroriparian reptile (the cottonmouth, that the farthest north that H. cinerea occurs in Agkistrodon piscivorus) has been collected in

Missouri along the Mississippi River is well to the drainage of Drury Creek. Additional the south in Cape Girardeau County (across the colonies of H. cinerea have been found since river from Union and Alexander counties, 1991 in southeastern Jackson County, mostly in Illinois). retention ponds or lakes not far from Drury

We conclude that there is no compelling Creek. evidence that the nineteenth century records of In 1995 and 1996 large choruses were

H. cinerea from Madison County, Illinois, and observed in ponds at two commercial fish St. Louis, Missouri, are valid. farms (Fig. 9B) on the floodplain of the

Mississippi River in western Jackson County.

Range Expansion Both fish farms are within 1 .5 km of floodplain

It became apparent to us that H. cinerea swamps containing H. cinerea. The species expanded its range northward and eastward in also occurs in weedy bays of Little Grassy Illinois during the past 10-25 years. LEB Lake (Fig. lOA) less than a km from the Little carried out field work in southern Illinois as a Grassy Fish Hatchery (operated by the Illinois graduate student at Southern Illinois University Department of Natural Resources) along the in the early 1960s and never observed H. Williamson and Jackson county line. The fish cinerea in or near Carbondale, Jackson County. hatchery is in an upland setting ca. 4 km from RAB has lived in Carbondale and carried out the nearest natural wetlands with a colony of H. field research in the area for over 30 years and cinerea. never encountered H. cinerea in the vicinity of Choruses have been observed and docu- Carbondale before the 1980s. In 1982, M. mented in most of the medium to large lakes in

Morris (pers. comm.) was the first to find H. far southern Illinois, all of which were im- cinerea in Carbondale at Evergreen Park. He pounded in this century. These include (with continued to encounter the species there impoundment dates) campus lake at Southern through 1987. In the late 1980s, W. McDowell Illinois University at Carbondale (impounded (pers. comm.) collected H. cinerea from a twice, in slightly different basins, most recently sewage lagoon (Fig. 9A) on the south edge of ca. 1952), Carbondale Reservoir (ca. 1925), Carbondale, and a large colony continues there Cedar Lake (ca. 1973), Crab Orchard Lake (ca. now. Additional large colonies were discov- 1940), Dutchman Lake (ca. 1973), lake at ered at other sites on the south edge of Feme Clyffe State Park (ca. 1960), Kinkaid Carbondale and on the campus of Southern Lake (ca. 1971), Lake of Egypt (ca. 1962),

Illinois University at Carbondale during 1991 Little Grassy Lake (ca. 1940), and Lake and 1994-1997. These were discovered in a Murphysboro (ca. 1970). Most of these are generally south to north order over successive deep (over 5 m) upland reservoirs, many have years, possibly indicating a gradual invasion of clear water, and many have limestone or suitable breeding habitat in Carbondale. sandstone ledges along portions of the shore-

During 1995-1997, H. cinerea was heard and line. All have shallow bays with abundant collected in the drainage of Drury Creek and in emergent vegetation (Fig. lOA). Choruses of ponds as well as along the road in Giant City H. cinerea usually occurred in eutrophic heads State Park near Boskydell and Makanda. Drury of shallow bays where cattail (Typha sp.), Creek passes the southeastern edge of willow {Salix sp.), American lotus (Nelumbo Carbondale (within 1.2 km of the sewage lutea), and other emergent plants were abun-

lagoon) in its flow northward to Crab Orchard dant. 50 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Figure 9. Habitats of //v/a c/nerea in Illinois as they appeared in August. A (Top): Sewage lagoon at a mobile home park, Carbondale, Jackson County. B (Bottom): Commercial fish farm ponds 1.2 km E Gorham, Jackson County. Photos by M. Redmer. B September 1999 Bird-voiced Treefroa and Green Treefrog in Southern Illinois 51

Figure 10. Habitats of //Wa c/W/ra in Illinois as they appeared in August. A (Top): Weedy bay of Little Grassy Lake at Giant City State Park, Jackson/Williamson counties. B (Bottom): Morgan Pond, a karst sink pond ca. 3 km E Dongola, Union County. Photos by M. Redmer. 52 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Habitat in a nearby timbered swamp. In eastern Union We found H. cinerea in a wide variety of County, choruses often were found in karst sink wetland habitats. All chorusing sites were ponds (Fig. 1 OB) and farm ponds. Emergent sunlit by day because tree canopy was sparse or vegetation in these habitats sometimes con- absent. Male H. cinerea usually were observed sisted oi N. lutea, Typha sp., C. occidentalis, calling from elevated perches in N. liitea, and 5^///.v sp., but a few such ponds were

Typhci sp.. shrubs (especially C occidentalis), located in pastures and crop fields where or young trees (especially Sali.x sp.). Den.se emergent vegetation consisted of crop plants or mats of duckweed (mostly Lemna sp.) were stubble. Impoundment lakes in southern present at most localities. Many of these Illinois often have rocky, steeply sloped habitats appeared disturbed or successional shorelines (Harris et al. 1977), which do not because mature baldcypress and tupelo gum offer suitable habitat for H. cinerea. However, trees (if present) usually were dead or stressed, these lakes usually are fed by many small and shrubs (especially C. occidentalis) were tributary streams that deposit sediment into abundant (Fig. 7A, B). Furthermore, the bays (Harris et al. 1977). Consequently, the dominance of Typlia sp. at a number of the sites shallow bays become heavily vegetated (Fig. is indicative of habitat disturbance (Anderson lOA) and provide suitable habitat for H. and Brown 1991: Odum 1988). Phelps and cinerea. In addition to the various impound- Lancia (1995) found H. cinerea in South ment reservoirs, we also found choruses of H. Carolina more abundant in clearcut swamp than cinerea in several borrow-pit ponds along roads (Fig. 11).

:."• *

*.i- i

Figure 11. Habitat of Hyla cinerea in Illinois as it appeared in August: Shallow borrow

pit pond along Interstate 24, 1 .4 km SW Big Bay, Massac County. Photo by M. Redmer. September 1999 Bird-voiced Treefrog and Green Treefrog in Southern Illinois 53

The habitat at Otter Pond. La Rue-Pine Hyla avivoca usually is described as an Hills SwampAVolf Lake was described by inhabitant of large forested swamps along Garton and Brandon (1975). where they found rivers (Blair and Lindsay 1961; Conant and males usually called from branches or leaves of Collins 1998; Dundee and Rossman 1989; several emergent plants including Rosa Krupa 1986; Trauth and Robinette 1990; and pahistris, C. occidentalism D. verticilatus, and others). In Illinois, all localities are floodplain yellow pond lily {Nuphar advena), usually 0.3 swamps in drainages of Bay Creek, Big Muddy m or more above water. During peak breeding, River, Cache River (and its tributaries Cypress calling males were also found on perches Creek, Dutchman Creek, and Limekiln nearer water (within 0.2 m) and on mats of Slough), and (formerly) Lake Creek. One of surface vegetation. these (Big Muddy River) is a tributary of the We found tadpoles to be abundant in warm Mississippi River, while the remainder are (sunlit) water at breeding sites. The tadpoles tributaries of the Ohio River. (usually with tadpoles of cricket frogs, A. Hyla avivoca probably originally invaded crepitans) were most often found in shallow southern Illinois by moving northward up the (< 20 cm deep) water and under mats of Mississippi embayment. The populations along duckweed. the Big Muddy River, lower Cache River, and During September-October, we frequently Mississippi River are a direct result of that found recently transformed H. cinerea in colonization. However, the biogeography of//. emergent vegetation and shrubs around the avivoca along the upper Cache River and Bay peripheries of breeding sites, and even in Creek requires a more complex interpretation. grassy fields. Garton and Brandon ( 1975) as The swamps along these rivers are remnants of well as this study found that young-of-the-year the floodplain habitat of the ancient channel of moved into surrounding floodplain and the Ohio River. This channel cut across (from adjacent upland forest by September and east to west) large parts of Pope, Massac, October. Some may have spent the winter Johnson, Pulaski, and Alexander counties under limestone talus or in rock crevices. before the Ohio River divided and shifted to

assume its current more southern channel Discussion during the Pleistocene (Alexander and Prior

1968; Hutchison 1987). This shift is thought to Biogeography have been caused by meltwaters of Because both H. avivoca and H. cinerea are Wisconsinan glaciation 13,000 to 10,000 years Austroriparian species that reach their northern ago (Alexander and Prior 1968; Hutchison limits in Illinois, their biogeographic histories 1987). The current floodplain of the Cache in the state have much in common. Remnants River now occupies the western portion of the of the baldcypress/tupelo gum swamplands that old channel of the Ohio River (flowing into the formerly were extensive across far southern current Ohio River near Cairo, Alexander

Illinois now are concentrated mostly in three County), while the lower drainage of Bay areas: (1) the floodplain of the Mississippi Creek occupies the eastern portion (flowing River and drainage of the lower Big Muddy into the Ohio River near Bay City, Pope River; (2) the drainages of Bay Creek, Cache County).

River, and former channel of the Ohio River in In presettlement times the drainages of Bay which swampland was extensive before being Creek and the Cache River contained large largely drained for agricultural purposes tloodplain swamps known as Big Black Slough (Alexander and Prior 1968; Hutchison 1987); and Little Black Slough (Hutchison 1987) that and (3) the lowlands of southern Massac and probably provided nearly continuous habitat for Pope counties adjacent to the Cretaceous Hills H. avivoca across this part of the state. Big Section, which drain directly into the Ohio Black Slough drained into the Cache River

River (Harris et al. 1977). through a sluggish stream or received water 54 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

from the Cache River depending on flood level. invade new habitat through dispersal of During 1913-1916. Post Creek Cutoff, a offspring, since recently transformed juvenile shortcut to the Ohio River, was excavated by H. cinerea have been found up to 0.5 km from humans to carry water from the upper Cache the nearest breeding habitats (this study). It River more quickly and reduce flooding of also is possible that previous surveys over- agricultural fields downstream. As a conse- looked small, localized populations that have quence. Big Black Slough and much of Little expanded during recent years. However, the Black Slough were drained, timbered, and increasing number of permanent impoundment converted to agriculture leaving scattered lakes, retention ponds, and farm ponds con- populations of H. avivoca in remnants of structed in southern Illinois since the 1950s has original swamp. Thus, although the popula- facilitated expansion of H. cinerea by provid- tions along Bay Creek and the Cache River ing additional suitable habitat. Many of these might superficially appear to have ancient impoundments and ponds are stocked and independent origins, their isolation actually is a managed for sport fishing. Predatory fishes recent result of human environmental alter- may not exclude H. cinerea from these perma-

ation. nent aquatic habitats as they do some other Elsewhere in the valleys of the Mississippi amphibians. Blouin (1990) presented experi- and Ohio rivers, H. avivoca is known from mental evidence that tadpoles of H. cinerea are western Kentucky (Barbour 1971), western less vulnerable than tadpoles of H. gratiosa to Tennessee (Redmond and Scott 1996), Arkan- predatory fishes (either through greater sas (Trauth and Robinette 1990), Mississippi unpalatability or difference in behavior), and (Lohoefener and Altig 1993), and Louisiana this may help explain the continued occurrence (Dundee and Rossman 1989; Fouquette and of H. cinerea in permanent lakes and swamps. Dalahoussaye 1966), but has not been reported There is further evidence that human from Missouri (Johnson 1987) immediately assistance has allowed H. cinerea to expand its west of some localities in Illinois. range in Illinois. It is likely that fish farms on Many of our biogeographic comments on the floodplain of the Mississippi River were the distribution of H. avivoca in Illinois also colonized by H. cinerea from nearby swamps, apply to H. cinerea. However, while H. although it is possible that the fish farms were

cinerea is associated with Austroriparian colonized by adults or tadpoles introduced with

It is pertinent swamps, it is not limited to them. We have fish stock from other fish farms. considered three possible explanations for the also to consider how H. cinerea entered the currently expanding range of H. cinerea in medium to large reservoirs. Most are far from

Illinois: (1) construction of many new aquatic remnant swamps, but all periodically are breeding habitats, such as farm ponds and stocked with fish. It is possible that tadpoles of impoundment reservoirs; (2) the ability of the H. cinerea were introduced along with fish species to colonize new aquatic habitats rapidly raised in hatcheries. P. Reif, owner of a fish through dispersal of subadults; and (3) intro- farm in Jackson County where we found H. duction of tadpoles through fish stocking into cinerea, confirmed (pers. comm.) that tadpoles many of the new aquatic habitats. are included in deliveries of stock fish from his farm, although efforts are made to exclude Range Expansion of Hyla cinerea them. While Reif was familiar with adult H.

The range of H. cinerea is expanding in cinerea, he could not identify their tadpoles.

southern Illinois. The northernmost Introduction of tadpoles through fish stocking localities we found (Rockwood, Randolph probably is widespread. In 1991 and 1993, MR County, and northwestern Kinkaid Lake, observed tadpoles of R. catesbeiana being Jackson County) are both ca. 27 km farther introduced when catfishes {Ictalurus punctatus, north than Smith's (1961) northern range limit Pylodictus olivaris), largemouth bass for the species. This species has the capacity to {Micropterus salmoides), and walleyes )

September 1999 Bird-voiced Treefroa and Green Treefrog in Southern Illinois 55

{Stizostedion vitreum) were stocked into public the toads.) The hatchery personnel made fishing lakes in DuPage County, Illinois. All persistent but unsuccessful attempts to elimi- the fish and tadpoles originated from fish farms nate the toads over many years. Eggs were >70 km away. LEB also observed tadpoles of killed by tossing them onto the banks, and R. catesheiana being introduced with fishes in adults were transported elsewhere or killed.

the 1970s into the human-made impoundment This seemed to have had little effect because Evergreen Lake. McLean County, in central large numbers of toads returned to breed during Illinois. Furthermore, as R. catesheiana has the three-year study. LEB noted a similar been introduced widely throughout much of abundance of tadpoles of American toads. B. western North America (Wright and Wright americanus, in the fish hatchery ponds of

1949: Stebbins 1951), it is logical to conclude Southern Illinois University at Carbondale in that some (perhaps many?) of these introduc- the early 1960s. W. Lewis (pers. comm.) also

tions occurred as tadpoles through the numer- reported problems in the 1960s and 1970s with

ous fish stockings that have taken place. In large numbers of anuran tadpoles in fish

1996 at the Aquaculture Facility, Illinois State hatchery ponds near Gorham, Illinois (which is

University, B. Wainscott and P. Foley (pers. within our study area). Brown ( 1 97 1 ) sug- comm.) found a number of metamorphosing gested that the filling of the ponds may be a American toads {Bufo americanus) in a factor that stimulates anurans to use them for shipment of yellow perch fry (Perca reproduction. More recently there has been flavescens) that originated from Cambridge. more extensive documentation that "tadpole

Wisconsin (ca. 292 km to the north). In 1997. infestation" is a serious problem in fish E. Massarolo (pers. comm.) found significant hatcheries (Carmichael and Tomasso 1983;

numbers of tadpoles of R. catesheiana in Corse and Metter 1980; Gabbadon and shipments of goldfish (Carassius auratus) and Chapman 1996; Kane and Johnson 1989; Kane

guppies (Poecilia reticulata) to a pet store in et al. 1985; Kane et al. 1992). Tadpoles often

Bloomington, Illinois. Platz et al. (1990) constitute 50-75% of the harvest biomass, and presented evidence for the introduction of the can reach over 2,000 kg/ha (Kane and Johnson

Rio Grande leopard frog Rana herlandieri into 1989; Kane et al. 1985). Thus, there are ample

southwestern Arizona from Uvalde, Texas, and indications that tadpoles occur in fish hatcher-

eastern New Mexico. They implicated trans- ies, and that tadpoles are included in outgoing

port of eggs or tadpoles during fish stockings. shipments of fish.

Thurow (1994) intentionally established wood Hyla cinerea also may be expanding its frogs {R. sylvatica) by introducing eggs, range (either naturally or by introduction)

tadpoles, and adults (collected in Brown elsewhere. Conant (1977) suggested that the County, Indiana) into woodland pools in isolated population(s) in the area near

McDonough County. Illinois. Reinert ( 1991 Brownsville, Texas, initially resulted from and Thurow (1994) noted that introductions of escapees from the pet trade. Snyder and Piatt

early life stages were most likely to be success- (1997) reported possible range expansion in ful in establishing new colonies. South Carolina. An introduced population of

There is considerable evidence that H. cinerea was reported at a fish farm in

tadpoles occur in abundance in fish hatchery Camden County, Missouri, by Johnson (1987)

ponds. Brown ( 1 97 1 ) studied reproductive and recent records (Powell et al. 1995, 1996) ecology of Woodhouse's toad, Bufo from southeastern Missouri could indicate woodhousei, and the Gulf Coast toad, Bufo range expansion there. In 1974 a now-extinct

valliceps, at a federal fish hatchery in Austin, colony was established at a fish farm in

Texas. Large numbers of both species of toads Douglas County, Kansas (Collins 1993; J. as well as R. catesheiana laid massive numbers Parmelee, pers. comm.), and an introduced

of eggs in the 19 hatchery ponds. (The R. population is known in Puerto Rico (Schwartz catesheiana were introduced by the hatchery and Henderson 1991). management in hopes that they would prey on 56 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Conservation Biology (Bogner et al. 1985). Sources of water for the

Smith (1961 ) reported H. avivoca from 3 lake originally were its 39.3-km- drainage, localities in Illinois; we report it from 29 flooding from the Mississippi River, and localities (of which 24 are documented by rainfall. The 76-hectare lake (a seasonal slough vouchers). The increase probably is due more before spillway construction) drained into the to additional field work than to range expan- Cache River by way of Lake Creek. Stop log sion. This species occurs today only in and concrete spillways were constructed on remnant natural swamps, many of which Lake Creek in 1927, 1931, and 1939 (Thomp- currently are difficult to access, and certainly son and Bennett 1938), prior to which the were even less accessible when Smith was meander had existed as a shallow baldcypress surveying the state's herpetofauna. Twenty- swamp that dried partially during July and one (72%) of the current localities are in the August. Lake Creek is subject to annual spring drainage of the Cache River, three (10%) are in backwater flooding from the Cache River and the drainage of Bay Creek, and five (17%) are floodwater occasionally enters the lake over the in the bottomland of the Big Muddy and spillway. Until 1969, when the Fayville Levee Mississippi rivers. This species may have been was completed. Horseshoe Lake occasionally extirpated from Olive Branch/Horseshoe Lake, received overbank flooding from the Missis- Alexander County. Horseshoe Lake went dry sippi River through Big Cypress Creek into in the summer of 1930 (Cahn 1937; Thompson Black Creek (Bogner et al. 1985; Lee 1993). and Bennett 1938), and that might have Horseshoe Lake also is threatened by excessive influenced the decline or extinction. Of greater sedimentation (Lee et al. 1986). During the significance may be the intensive long-term flooding of the Mississippi River in 1993 and management of the lake for recreation by the 1994, several months of inundation killed much State of Illinois, which has involved control of of the surrounding forest, including many vegetation along the shore (M. Morris, pers. tupelo trees. Animal and plant life at Horse- comm.). Hyla avivoca was heard only twice at shoe Lake includes many Austroriparian another locality (Oakwood BottomsAVorthen species, making it one of the most distinctive

Bayou, Jackson County), where it may also be natural areas in Illinois (Evers and Page 1977). close to extinction or extirpated. All popula- In addition to H. avivoca and H. cinerea, tions are in or along the margins of remnant Austroriparian amphibians and reptiles known baldcypress/tupelo gum swamps. Choruses to occur there historically include the mole were largest in less disturbed swamps where salamander (Ambystoma talpoideum), Missis- canopy cover was greatest and the understory sippi green water snake (Nerodia cyclopion), well shaded at midday. Choruses were smallest mud snake {Farancia abacura), broad-banded where mature baldcypress and tupelo gum trees water snake {Nerodia fasciata), plainbelly were sparse or dying and the canopy was more water snake {Nerodia erythrogaster), and open. Similar observations were reported for cottonmouth (A. piscivorus). These species choruses of H. avivoca in Oklahoma (Secor thrive in shallow, bottomland baldcypress and

1988). There also is evidence (based on diet) tupelo swamps subjected to cyclic flooding and that H. avivoca forage in arboreal habitat, drying. They may have been negatively rather than on the ground (Jamieson et al. 1993; affected when the seasonal swamp was Redmeretal. 1999). converted into a permanent lake in the late The management history of Horseshoe 1920s and early 1930s. Subsequently, Horse- Lake clearly shows how habitat disturbances shoe Lake has been protected from most may have led to the decline of H. avivoca and overland flooding of the Mississippi River and other Austroriparian amphibians and reptiles. managed for the benefit of sport fishing and Horseshoe Lake is a cut-off meander, oxbow waterfowl. While a few Austroriparian species lake formed in the wide floodplain of the (A. talpoideum, H. cinerea, N. erythrogaster) Mississippi River about 6,000 years ago remain locally common, the others are rare or September 1999 Bird-voiced Treefroa and Green Treefros in Southern Illinois 57

extirpated (Brandon et al. 1994; Heidinger et al. anurans; the bullfrog, R. catesbeiana, in 1994. 1995; Brandon and Wilson 1996). western North America (Wright and Wright Twenty-three of the twenty-nine (79%) 1949; Stebbins 1951), and the giant toad, Bufo current known breeding sites for H. avivoca are marinus, in Australia and elsewhere (Tyler owned in whole or in part by public (Illinois 1989). Both of these colonizations were Department of Natural Resources, United knowingly facilitated by humans (for sport

States Forest Service [Shawnee National purposes and consumption in the case of R. Forest], United States Fish and Wildlife Service catesbeiana; for biological control of sugar

[Cypress Creek National Wildlife Refuge]) or cane pests in the case of B. marinus). The private (The Nature Conservancy) conservation expansion of H. cinerea probably is natural and agencies, so protection of most remaining also unknowingly facilitated by human beings habitat is possible. Nevertheless, because of through inadvertent introductions of tadpoles the frog's dependence on scattered fragments with game fishes and creation of new habitat. of declining habitat, we formally recommended Whether or not the expansion of//, cinerea will

(while this manuscript was in preparation, 2 reach the magnitude of those of R. catesbeiana

December 1997) that the Illinois Department of and B. marinus is unknown, but it presents a Natural Resources and Illinois Endangered tremendous opportunity to study amphibian

Species Protection Board consider listing H. success and contrast it with species headed avivoca as a threatened species. The Board toward extinction. Such a comparison should voted positively in August 1998 on this increase our understanding of the decline of the recommendation and the species is now listed Amphibia, and perhaps supplement our meager as threatened in Illinois (Illinois Endangered knowledge of how these important animals can Species Protection Board 1998). Additional be conserved. studies will be necessary to determine micro- habitat requirements so that remaining habitats Summary can be managed in a way that is beneficial to the frogs. Management may need to include The bird- voiced treefrog {Hyla avivoca) and cyclic manipulation of water levels to ensure green treefrog {Hyla cinerea) are Austro- that healthy stands of mature tupelo and riparian species that reach the northern limits of baldcypress (as well as diverse understory) their ranges in the Midwest in far southern persist. Illinois. Hyla avivoca is a habitat specialist that

Smith ( 1961 ) reported H. cinerea from 4 persists only in remnant baldcypress/tupelo localities in Illinois, whereas it is now known swamps, which were more widespread in from 127. Although it occurs around the open former times. Its survival in Illinois depends edges of baldcypress/tupelo swamps, H. upon the survival of this swamp habitat. Hyla cinerea is not limited to them. It is less of a avivoca is restricted to isolated colonies (often habitat specialist than H. avivoca, and often is with low numbers of individuals) in drainages abundant in a variety of disturbed, succes- of the Big Muddy River, Mississippi River, sional, or open swamp habitats, as well as sink Cache River, and Bay Creek in Alexander, ponds, farm ponds, borrow pits, and shallow Jackson, Johnson, Pope, Pulaski, and Union bays of many constructed lakes. There is no counties. Males call from a variety of emergent evidence that H. cinerea is in danger of plants, commonly the trunks and branches of extirpation in Illinois. baldcypress and tupelo trees, nearly always With all the recent media publicity con- over water. Tadpoles develop in cool, shaded cerning the "decline of the Amphibia," it could water. Recently transformed juveniles and be regarded as unusual that H. cinerea is subadults often perch in low shrubs in nearby significantly expanding its geographical range woods, on emergent shrubs, and on baldcypress and increasing markedly in numbers. How- and tupelo trees. In the fall they can be ever, within historical times there have been terrestrial. major range expansions by at least two other 58 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Hyla cinerea occurs in a wide variety of Literature Cited disturbed, successional, and open swamp habitats, and during the past 30 years has been Ackerman, K. 1975. Rare and endangered undergoing considerable expansion of its range vertebrates of Illinois. Illinois Department of into farm ponds, borrow pits, and shallow bays Transportation, Bureau of Environmental reservoirs constructed since the 1950s. It is of Science, Springfield. 50 pp. widely distributed across much of Alexander,

Jackson. Johnson, Massac, Pope, Pulaski, and Alexander, C.S., and J.C. Prior. 1968. The Union counties, as well as in parts of Randolph origin and function of the Cache Valley, Williamson counties. Males call from and southern Illinois. Pages 1 1-18 in R.E. shrubs, small trees, and other emergent Bergstrom, ed. The Quaternary of Illinois. vegetation in a variety of wetland habitats University of Illinois College of Agriculture sunlit by day in the open or under sparse tree Special Publication 14, Urbana. canopy. Tadpoles are abundant in warm sunlit waters covered by mats of duckweed. Recently Altig, R. 1970. A key to the tadpoles of the transformed juveniles are found in emergent continental United States and Canada. vegetation, shoreline shrubs, and even nearby Herpetologica 26:180-207. grassy fields.

Both species probably originally invaded Anderson, R.C., and L.E. Brown. 1991. southern Illinois by moving northward up the Establishment of a wetland plant community in Mississippi embayment with subsequent a limestone quarry. Castanea 56: 168- 175. colonization of the drainages of the Cache, Big

Muddy, and Ohio rivers. Although the Ashton, R.E., Jr. (comp.), S.R. Edwards, and populations of H. avivoca along Bay Creek and G.R. Pisani (eds). 1976. Endangered and the Cache River might superficially appear to threatened amphibians and reptiles in the origins, their have independent separation United States. Herpetological Circular 5, actually is a recent result of intervening human Society for the Study of Amphibians and environmental alteration. The currently Reptiles. 65 pp. expanding range of H. cinerea in southern

Illinois probably is a result of (1) construction Ballard, S.R. 1994. Status of the herpetofauna of many new breeding habitats by humans in the La Rue-Pine Hills/Otter Pond Research (e.g., farm ponds, reservoirs), (2) the ability of Natural Area in Union County, Illinois. M.S. dispersing metamorphosed young to colonize Thesis, Southern Illinois University at new aquatic habitats rapidly, and (3) introduc- Carbondale. xix -i- 167 pp. tion of tadpoles by humans through fish

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Lake management reports. 1. Horseshoe Lake 64 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Appendix (LSUMZ 6569); 2.5 mi E of Aldridge (SIUC H-66 1-662); 3 mi NE Aldridge (INHS 6062- 6068); 5 mi N & 3 mi E Wolf Lake. Pine Hills, Recreation area (KU 63546); 5 mi SE Aldridge Locality data for Hyla avivoca, Hyla (INHS 3102 [lot of 2], 3312-3318.4175-4179); chrysoscelis, Hyla cinerea, and Hyla versicolor 5 mi SE Aldridge (UIMNH 1725-1727); complex examined. Vouchers are listed in Hogan's Bottoms on Dongola Road just W of parentheses after localities. When a voucher is Cypress Creek (SIUC H-52 16-52 17, H-5256); not listed, the individual(s) was (were) identi- La Rue Ecological Area (SIUC H-4387); La fied by mating call, photograph, and/or capture Rue Pine Hills Research Natural Area (SIUC (with subsequent release). H-5121); Pine Hills Recreation Area (ISM

626 1 6 1 -626 1 66. 626 1 7 1 ); Pine Hills, between Hyla avivoca: Alexander Co., IL: Cypress SIU field station and Route 3 (SIUC H-2 [lot of Creek National Wildlife Area, Cache River just 5]); swamp near Wolf Lake (UIMNH 95883- SE of Tamms (SIUC H-5379): Olive Branch. 95884; UMMZ 95800. 95883-95884. 95885 Horse Shoe Lake (FMNH 2201 [lot of 3]). [lot of 3] ); Wolf Lake (swamp at base of bluffs) Johnson Co., IL: 0.7 mi E of Grantsburg on (SAM 104. 262-263); Wolf Lake (UIMNH Route 146 (SIUC H-5259-5260): Belknap Road

1 3074. 1 3076- 1 308 1 . 705 1 6); Wolf Lake just W of Cache River (SIUC H-5255): Bell swamps (SIUC H-1267). Pond. W of Grantsburg on Route 146 (SIUC H- Hyla cinerea: Alexander Co.. IL: 1 .2 km 1 474 1 ): Cache River bottoms. .6 km W of West SSE of Horseshoe Lake spillway (SIUC H- Vienna (SIUC H-5257): Cache River State 5289); Big Cypress Swamp (SIUC H-5292); Natural Area, at Cache River and Route 37 Fayville Road 0.7 mi W of Olive Branch (SIUC (SIUC H-5254); Deer Pond Nature Preserve, H-4954-4955); Horseshoe Lake (INHS 1987, ca. 0.3 mi N of old Route 146 (SIUC H-5382); 5123; UIMNH 23897); Horseshoe Lake Heron Pond. Little Black Slough (SIUC H- Conservation Area (SIUC H-3398. H-466I. H- 5123-5124); North Cypress access N of parking 4786, H-4813, H-4866, H-4938-4939. H-4969- lot (SIUC H-5319): Route 37 at Cache River 4970. H-5007-5010, H-5I46, H-5272. H-5275- 0.9 km S of West Vienna (SIUC H-5248); 5276, H-5280-5281, H-5 191-5292, H-5294, H- Route 45 at Dutchman Creek bottoms ca. 1.25 5315-5316); Horseshoe Lake State Conserva- mi N of Belknap Road (SIUC H-5270-5271); tion Area on small cypress tree island, S side of W of West Vienna and E of Route 37 (SIUC H- West Side Dr. and Miller City arm of lake 5122); W side of Cypress Pond State Natural (INHS 128 18- 128 19); old channel of Cache Area (SIUC H-5258); Heron-Pond Little Black River just S of Tamms sewage disposal pond Slough, swamp south of Wildcat Bluff; Heron- (SIUC H-5226. H-5473); Olive Branch (FMNH Pond Little Black Slough, swamp west of 6290 [lot of 8]); 0.3 mi NE Olive Branch Wildcat Bluff; Heron-Pond Little Black (captured and released); 0.6 mi NE Olive Slough, swamp NE of Hairy Hill; W side of Branch (captured and released); 0.3 to 2 mi SW Little Black Slough. E of Cypress (SIUC H- Olive Branch on Miller City-Olive Branch 5262-5264). Pope Co.. IL: 0.8 mi SE of Road, many sites (captured and released; Reevesville along Bay Creek floodplain (SIUC DORs); Olive Branch. Horse Shoe Lake H-525 1 -5253, H-5383-5385); 1 .5 km NE of (FMNH 2199 [lot of 53]); Route 127 at S side Reevesville. along Sugar Creek (SIUC H-5543, of Mill Creek at Union County line (SIUC H- H-5598 [tadpoles]). Pulaski Co., IL: Cache 5162-5163); along Cache River SE of Frog River State Natural Area, 0.25 km SW of Route City. Jackson Co., IL: Carbondale, Evergreen 37 at Cache River bridge (SIUC H-5268); Terrace (MAM 826); Carbondale, SIUC Cache River State Natural Area. 2 km NW of campus (SIUC H-5 152-5 153); Etherton Karnak (SIUC H-5269); Limekiln Slough. 1/2- (swamp on east side of Hwy 127) (INHS way between parking lot and spring (SIUC H-

1 06 1 7- 1 06 1 8 ) ; Etherton Swamp on Route 1 27 5381). Union Co., IL; 2 mi E Aldridge September 1999 Bird-\oiced Treefrog and Green Treefrog in Southern Illinois 65

S of Murphysboro (SIUC H-3202-3205); Giant West Vienna (SIUC H-5 145); Route 146 along City State Park, north end near rappelling area Bell Pond, 0.6 km E of Grantsburg (SIUC H- (captured, photographed, released): Grimsby, 5213-5214); Vienna (INHS 2342 [lot of 4]); W intersection of Routes 3 and 149 (SIUC H- edge of Little Black Slough SE of Cypress

5150); Kinkaid Creek, first bend S Route 149 (SlUCH-5261). Massac Co., IL: Mermet (heard choruses on several nights in early Lake Conservation Area (SIUC H-3777, H- 1960s: now extinct due to habitat destruction 3814); SE edge of Mermet Lake Conservation from torrential overflow of Kinkaid Lake dam); area along Rt. 45; NE of Mermet Lake along Kinkaid Lake, south end near end of road Rt. 45; NW of Mermet Lake along Rt. 169; W

(captured, photographed, released by J. side of Fort Massac State park; E edge of Fort Karnes); Lake Indian Hills sewage lagoon near Massac State Park; 4 km SE Unionville in Ohio

Carbondale (SIUC H-5243); Kinkaid Lake River floodpain. Pope Co., IL: 1 km NW of

Johnson Creek Recreation Area access road 1.1 Renshaw along old railroad bed (SIUC H- km E of Route 151 (SIUC H-4439): Lake 5387); 1 mi W of New Liberty on the Murphysboro State Park (SIUC H-5147): Unionville Road (SIUC H-4450, H-5 126): 1.5 Logan Hollow Fish Farm, Gorham (SIUC H- km E of Reevesville (SIUC H-5376, H-5599);

5149); Lovetf s Pond (SIUC H-5223-5225, H- Barren Creek (SIUC H-4442); Bay Creek 0.8 5265-5267); north side of middle of Lovett's mi E of Reevesville (SIUC H-5378); Smithland Pond (captured, photographed, released); Lock and Dam road, 0.1-0.3 km W of Army Malibu Mobile Home Park, Carbondale (SIUC Corps building (SIUC H-4443-4447, H-5 127); H-5143); Oakwood Bottoms (MAM 691): farm pond 0.3 km S of Big Black Slough, along Oakwood Bottoms Waterfowl Management Rt. 145. Pulaski Co., IL: 50 mE of dock at

Area (SIUC H-5141-5142): Route 3, 2 km S of lower Cache River access area (SIUC H-5 1 28); intersection of Route 149 (SIUC H-5148); along Limekiln Slough, 5.3 mi E Ullin (INHS

1 1 1 - 1 1 ); Belknap Road 0.2 km of , 06 06 N Route 15 1 0.3 km N of Route 3 (SIUC H- 3 4711); S side of Makanda (SIUC H-5144); Karnak railroad crossing (SIUC H-5239); ca. 2 Touch of Nature Fisheries Ponds, S of mi SW Pulaski (MAM pers. comm.); Cache Carbondale (SIUC H-5352-5354); Town and River State Natural Area, 2 km NW of Karnak Country Trailer Court, S edge of Carbondale (SIUC H-5249): Post Creek Cutoff, NE side of (SIUC H-3349-3352, H-3965-3966, H-3968- Karnak (SIUC H-5377); Route 37 on S side of 3971, H-5218); Turkey Bayou area (MAM Cache River at Cache River State Natural Area pers. comm.); Wides Village, Murphysboro (SIUC H-5 1 29-5 131); Route 37, 0.2 mi S of (SIUC H-3974). Johnson Co., IL: farm pond 3 Cache River (SIUC H-5236); Route 51 at S km NW of Vienna; weedy bay on SW side of edge of Ullin, at Cache River bottoms (SIUC

Lake of Egypt; 0.4 mi N of Belknap Road on H-5 1 25); Route 5 1 on N side of town of Route 45 (SIUC H-5238): 0.7 mi N of Belknap Pulaski (SIUC H-5 132-5 133); Route 51, 3 mi on Route 45 (SIUC H-5234); 1.6 mi NE of N of Ullin (SIUC H-5 1 36-5 1 37); Sandusky to Karnak on Belknap Road (SIUC H-5240); Pulaski road just E of Cache River (SIUC H-

Belknap Road 1.1 km N of Karnak railroad 5138-5139); Shawnee College campus (SIUC crossing (SIUC H-5245); Cypress Pond State H-4438); Cache River floodplain W of Natural Area (SIUC H-5246-5247. H-5408- Mounds; large pond NE side of Mounds; along 5413); Dutchman Lake boat launch (SIUC H- Cache River 6 km W of Villa Ridge. Randolph 5242); Heron Pond State Natural Area (SIUC Co., IL: Route 3 at Rockwood (SIUC H-4710).

IL: 1 . 1 of intersection of Mt. H-5 1 5 1 ); old Cypress Road and 1 46, 1 .3 mi W Union Co., km S of Vienna (SIUC H-5241); old Route 146, 0.2 Pleasant Road and Route 146 (SIUC H-5 160- mi E of Dutchman Creek (SIUC H-5 140)): 5161); 2.0 mi. N Wolf Lake (INHS 3923): 5.0 Route 37, 0.1 mi N of Cache River (SIUC H- mi. SE Aldridge (INHS 3100); 5.0 mi. SE 5244); Route 37, 0.5 km S of Cache River Aldridge (INHS 3949): Aldridge (INHS 3149- along Cypress Creek (SIUC H-5215); (SIUC H-52 1 1 ); Route 37, 2.2 km SW of 3150); 66 Illinois Natural History Survey Bulletin Vol. 36 Art. 2

Hogan's Bottoms, just W of Cypress Creek [lot of 1 1]). Massac Co., IL: near Cagle Pond

(SIUC H-5154, H-5156; H-5158, H-5173- (SIUC H-3799). Pope Co., IL: ca. 1 mi W of 5180, H-5235): Morgan Pond (SIUC H-5168- New Liberty on Unionville Road (SIUC H- 5172); Mt. Pleasant, 0.3 mi S of Cache River 4951-4952); Ohio River Recreation Area near (SIUC H-5157); Mt. Pleasant, 100 m S of main Golconda (SIUC H-3796); One Horse Gap intersection (SIUC H-5 166-5 167); Old Cape (SIUC H-3798). Saline Co., IL: N Carrier Road 1.3 km E of Bluff Lake (SIUC H-4781- Mills (SIUC H-3803). Union Co., IL: Giant 4783); Old Cape Road at 3rd bridge E of Route City State Park (SIUC H-3793); Trail of Tears 127 (SIUC H-4722); Old Cape Road at Route State Forest (SIUC H-3795). Williamson Co., 127 (SIUC H-5 159); Pine Hills (SIUC H-62- IL: Crab Orchard National Wildlife Refuge 63, H-72, H-77-78, H-188, H-653, H-1 108, H- (SIUC H-3792). I676-I679, H-1900-1902, H-2673, H-3356- Hyla versicolor complex: Jackson Co., IL: 3365); Pine Hills Recre. Area (ISM 626071- Murphysboro (FMNH 25940). Madison Co.,

626078. 626098); Pine Hills Swamp (INHS IL: no specifics (USNM 57827). St. Louis

2503-2505; ISU 1639, 1643, 1662, 1684, Co., MO: St. Louis (USNM 3667 [lot of 4], 1706); swamp near Wolf Lake (UMMZ 95796 104553).

[lot of 3]); Union County Conservation Area Hyla sp. {H. avivoca or H. versicolor (SIUC H-4719-4721, H-5164-5165); Upper complex): Alexander Co., IL: Horseshoe Lake Bluff Lake along logging road on E side (SIUC (LSUMZ 51046). Union Co., IL: Ware, Union H-5 155); Wolf Lake (swamp at base of bluffs) Co. Wildlife Refuge (LSUMZ 51047). (SAM 105); Wolf Lake (UIMNH 13028, 13030-13032); retention pond along Rt. 127 at

Alto Pass; pond at Pine Hills campground; "State Pond" 3 km NW of Jonesboro; S half of Union County Conservation Area (large choruses throughout area); Cache River floodplain 1 .5 km NW of Mt. Pleasant; pond on NW side of Dongola; sink hole 2 km SE Dongola. Williamson Co., IL: Crab Orchard National Wildlife Refuge, spillway road (SIUC H-4102); Grassy Bottoms Swamp along Crab

Orchard Lake spillway road (SIUC H-5 1 34- 5135); Little Grassy Lake (SIUC H-4436); Crab Orchard Lake in weedy bay 2.5 km E of spillway; Crab Orchard Lake, in S half of grassy bay; W side of Little Grassy Lake at Touch of Nature Camp. Hyla chrysoscelis: Alexander Co., IL: 0.3 mi SE Promised Land Rd., S of Horseshoe

Lake (LEB 122 [lot of 3]); Horseshoe Lake Conservation Area (SIUC H-4807, H-5078). Franklin Co., IL: SW Zeigler (SIUC H-3802). Jackson Co., IL: Cedar Lake (SIUC H-3804); Giant City State Park (SIUC H-3794); Grand

Tower (SIUC H-3801 ); N of Big Muddy River,

W side of Route 3 (SIUC H-7 1 [lot of 7]). Johnson Co., IL: Cypress Pond State Natural Area (SIUC H-5707-5709); Feme Clyffe State Park (SIUC H-3800); Grantsburg (SIUC H-81

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