536.1 AMPHIBIA: ANURA: RANIDAE RANA BLAIRI Catalogueof America Amphibias ad Reptiles. Brow, L.E. 1992. Raa blairi. Raa blairi Mecham, Littlejoh, Oldham, Brow, ad Brow Plais Leopard Frog Raa blai Mecham, Littlejoh, Oldham, Brow, ad Brow, 1973: 3. Typelocality, '1.6 km. W New Deal, Lubbock Co.,Texas." Holotype, Uiv. Michiga Mus. Zool. (UMMZ) 131690,a adult male (i alcohol) collected o 6August 1971by Charles Everett (examied by author). Raablairi(sic): Brooks, 1976:89. R. Blai' (sic): Dulap, 1982:247. Cotet. No subspecies are recogized. m Defiitio. Raa blairi is a species of the Raapipies complex with oe or both dorsolateral folds iterrupted posteriorly ad displaced medially, o vestigial male oviducts, ad usually a complete, pale, supralabial stripe. Vocal sacs are exteral ad of moderate size. The dorsum is pattered with dark brow, roud spots, ofte surrouded by light, arrow borders. A dark sout spot isusually preset, ad the tympaumalmost always has a white spot i the ceter. The posterior surface of the thigh is mottled or reticulatedwithdarkbrowmarkigs. Thearea betweespots othe dorsum is buff, pale brow, sometimes reddishbrow (i Oklahoma), or sometimes dull gree (olivegree i Arizoa). The abdome is pale cream, except frequetly with some yellow posteriorly, as well as i the groi ad o the proximal part of the thigh. F&ure 1. Female Raa blai' from Sprig Lake. Tazewell Couty... ~ll%ois.photograph by D.W. Whiuah. Diagosis. Raa blairi ca be distiguished from other members of the R.pipiescomplex by the followig combiatio of morphological characteristics: oe or both dorsolateral folds iterrupted posteriorly ad displaced medially; o vestigial oviducts i Descriptios. The first extesive descriptive data for R. blairi were provided by McAlister (1962) before the species was foally described as ew. Littlejoh ad Oldham (1968) were the first todocumet the correspodece of a distictive matig call with males; ad usually a complete, pale, supralabial stripe. The distictive matig call has a low pulse umber (average less tha six pulses per call) ad a slow pulse rate (average less tha six pulses per sec below 24 C). Map. The large ope circle marks the typelocality; the solid triagle idicates a populatio that may have bee itroduced; solid symbols represet other locality records (localitiesthat are close together geographically are sometimes combied uder oe symbol).
L TIME IN SECONDS Flgure 2. Audiospectrogram (arrow bad) of matig calls of Raa bhiri from orth of Hudso, McLea Couty, Illiois, 24 April 1970; cloaca1 temperature 16.1 C, water temperature 16.0 C, air temperature 9.4"C, SVL 68 mm measured alive (L.E. Brow recordig). I differet morphology for the species. The type series (four males, four females, all adults) was described i depth by Mecham et al. (1973). Dulap ad Kruse (1976) ad Post ad Pettus (1966) icluded good, reliable descriptios. The differet "ierspot" (sic) coloratio (olivegree) of specimes from Arizoa was idetified by Frost ad Bagara (1977). I Oklahoma, Black ad Sievert(1989) reported iterspot coloratio of ta to reddish brow; red coloratio has also occasioally beeecoutered ithatstate (R.L. Lardie, pers. comm.). A statistical corparko of R. blairiad Raa berladie' for certai morphological traits was made by Platz (1972). Accouts i field guides, regioal hadbooks, etc., that give fair to good descriptios for idetificatioof adults idude Balligerad Lych(1983), Behler adkig(1979), Blackad Sievert(l989), Brow (19851, Chriitiase ad Bailey (1991), Clarke (1984), Collis (1982), Collis ad Colhs (1991), Coat ad Collis (1991), Garrett ad Barker (1987), Hammerso (1982a), Johso (1977, 1987), Lych (19851, Pace (1974), Smith (1978), ad Stebbis (1985). Detailed descriptios of tadpoles were give by Korky (1978) ad Scott ad Jeigs (1985); brief descriptios were provided by Frost ad Bagara (1977) ad Stebbis (1985); Hillis (1982) gave comparative commetary. Korky (1978) characterized tadpoles ear trasformatio. The egg was described by Frost ad Bagara (1977); the egg mass was described by Hammerso (1982a), Lych (1985), ad Stebbis (1985). Developig embryos, ewly metamorphosed stages, ad juveiles have ot bee described. The matig calls are typically groups of 24 wellspaced otes which may be described as "tuck tuck tuck tuck, tuck tuck tuck, tuck tuck tuck," etc. The first group of otes is usually loger tha subsequet groups. Matig call data were preseted by Brow ad Brow (1972), Dulap ad Kruse (1976), Frost ad Bagara (1976), Littlejoh ad Oldham (19681, Mecham et al. (19731, Axtell (1977), ad Frost ad Platz (1983). Characteristics of matig calls of 10 males recorded at the type locality i a temperature rage of 22.0240 C were summarized by Mecham et al. (1973) as follows: mea call duratio 0.70 sec (rage 0.361.00 sec); mea pulse rate 5.3 pulses/sec (rage 4.86.1 pulses/sec); mea pulse duratio 28.6 msec (rage 2434 msec); mea pulse rise time 9.7 msec (rage 716 msec). Rages for matig call characteristics of all 39 males (temperature rage 15.525.0 C) reported by researchers are: call duratio 0.351.00 sec; pulse rate 3.46.8 pulses/sec; pulse duratio 19.838.3 msec; pulse rise time 716 msec. Mecham (1971) described differet types of chuckle calls (grut, grid, low trill). Illustratios. A black ad white photograph of a dorsolatera1 view of the live, adult holotype was preseted by Mecham et al. (1973) ad Dubois (1977). Other illustratios iclude: color photographs of dorsolateral views of live adults (Behler ad Kig, 1979; Black ad Sivert, 1989; Brow, 1985; Collis, 19; Collis ad Collis, 1991; Hammerso, 1982a; Johso, 1982, 1987); a color photograph of a dorsolateral view of a live subadult (Garrett ad Barker, 1987); color illustratios of dorsolateral views of adults (Coat ad Collis, 1991; Stebbis, 1985); black ad white photographs of dorsolateral views of live adults (Bozema et al., 1987; Brow ad Morris, 1990; Brow et al., i press; Collis, 1974, 1982; Dulap ad Kruse, 1976; Johso, 1977; Mecham et al., [paratype], 1973; Platz, 1972); a black ad white photograph of a frotal view of a live adult (Brow ad Moms, 1990); a black ad white photograph of a dorsal view of a preserved adult (Creel, 1963) with a partly igested bat, a Easter Pipistrelle (Pipistrellus subflaous); other black ad white photographs of dorsal views of preserved adults (Pace, 1974; Smith, 1956); blackadwhitedrawigsofadults (Balliger ad Lych, 1983; Clarke, 1984; Johso, 1987); a closeup, blackad white photograph of a displaced dorsolateral fold (Post ad Pettus, 1966); a closeup, black ad white photograph of a lateral view of a collapsed vocal sac of a preserved male (Pace, 1974); a black ad white photograph of a lateral view of a live tadpole, Goser stage 39 (Scott ad Jeigs, 1985); black ad white photographs of dorsal ad lateral views of preserved tadpoles, Goser stage 40 (Korky, 1978); black ad white drawigs of lateral (Goser stage 38) ad dorsal views, mouthparts, ad iris of tadpoles (Scott ad Jeigs, 1985); photographs of chromosomes (Ward, 1977); photographs of isozyme pheotypes (Dulap, 1979,1982; Frost ad Bagara, 1977; Kruse ad Dulap, 1976; Platz, 1972); audiospectrograms of matig calls (Brow ad Morris, 1990; Frost ad Platz, 1983; Kruse, 1981; Mecham, 1971; Mecham et al., 1973); oscillograms of matig calls (Littlejoh ad Oldham, 1968; Mecham, 1971); ad audiospectrograms ad oscillogram of chuckle calls, e.g., grut, grid, low trill (Mecham, 1971). Distributio. Raa blairi occurs primarily i the Great Plais ad Prairie Peisula, with a few populatios i Arizoa. The species rages from southeaster ad southcetral South Dakota i the orth, to cetral Texas i the south, ad from southeaster Arizoa i the west, to cetral Idiaa i the east. I the west the rage exteds ito easter Colorado, easter ad southwester New Mexico, southeaster Arizoa, adwestertexas. Eastwardly, these frogs rage throughout the southeaster half of Nebraska, all of Kasas, much of Oklahoma (except the southeaster quarter), ad much of orther Texas (except i the ortheast). I the Prairie Peisula, R. blai' is foud i the southwester half of Iowa, the orther half of Missouri, ad ia wide bad across cetral llliois to cetral Idiaa. A extesio of the rage follows the Mississippi River south ito southeaster Missouri. Fiftee isolated populatios have bee foud i Cochise Couty, southeaster Arizoa. A populatio i orthcetral Arizoa (Ashurst Lake, Cocoio Couty; r*r
Platz, 1976) may be itroduced 0.E. Platz iclarkso ad Rorabaugh, 1989). May populatios i a umber of areas appear to be isolated. Major distributioal studies by regio iclude: Arizoa (Frost ad Bagara, 1977); Colorado (Hammerso, 1982a; Post ad Peus, 1966); Illiois (Brow ad Morris, 1990); Iowa (Christiase ad Bailey, 1991); Kasas (Collis, 1982); Missouri (Johso, 1987); Nebraska (Brooks, 1976; Lych, 1978,1985); NewMexico (Frittsetal., 1984); Oklahoma (Black, 1976; Lardie, 1982); Texas (Dixo, 1987); ortheradcetral Great Plais (Dulap adkruse, 1976); souther Great Plais (Hillis, 1981); ad geeral (Littlejoh ad Oldham, 1968; Pace, 1974). Most of these publicatios iclude rage maps. Fossil Record. Noe has bee positively idetified. However, may Pleistocee fossils (Kasas Eshelma [19751, Holma [1971,1972,1984,19871, Rogers [19821, Tihe [19541; Texas Holma [19691, Kasper ad Parmley [19M1, Parmley [19883) have bee assiged to R. pipies, R. pipies complex, or Raa sp. idet. from withi the preset rage of R. blairi, but allopatric to R.pipies, sesu slricto (Brow ad Morris, 1990) ad other leopard frog taxa. Raa pipies, sesu lalo, has bee the most cosistetly reported frog from the Pleistocee of the U.S.A. (Holma, 1972). However, Rogers (1984) could ot separate the boes of R. blairi, R.pipies, ad Raa spheocephala. Likewise, Holma (1977) could ot fmd ay cosistet differeces i the ilia of these three species ad R. berladieri, ad he cocluded that the boes of species of leopard frogs are so similar that the fossil record will ot clarify their relatioships. Cosequetly, distictio of fossilized specimes of differet siblig species of leopard frogs may ot readily be possible at preset. Thus, Rogers et al. (1985) reported "R. blairior R.pipieru" from the Middle Pleistocee of Colorado (withi the preset rage of R. pipies, just outside the preset rage of R. blai). Pertiet Literature. Before the advet of settlemet by Europeas, much of the habitat of R. blairiprobably was prairie ad adjacet areas (Brow ad Morris, 1990; Brow et al., i press). With the subsequet extesive alteratio of these eviromets by humas ad climatic chages, R. blairiow occurs i a wide variety of habitats. Muchofthe ladthroughoutthe rage ofr. blairiisdevoted to agriculture, but Brow ad Morris (1330) ever foud the species i "cultivated fields" i Illiois. This is probably due to a umber of types of severe modificatios for agriculture. Outside the breedig seaso Brow ad Morris (1990) foud the species ear breedig sites, i old fields, alog creeks, o bottomlads, ad i variable habitats that ofte showed past disturbace (oagricultural) by humas, but ot i mature uplad forests. Former prairie regios ad associated river floodplais, flatlads, rollig hills, ad areas ear aquatic habitats are ihabited i Missouri uohso, 1977, 1987). I Nebraska, Lych (1978) foud that R. blai'occurred o loess soils ad "i areas extesively used for agriculture." Rowcrop cultivatio icreases levelsofturbidity i streams, which the species ofte seems to prefer. Other workers (Black ad Sievert, 1989; Hillis, 1981; Sco adjeigs, 1985) metioedthe associatioof R. blairi withturbid pools, ad muddy taks ad rivers. Parasitological evidece from Nebraska (Brooks, 1976) idicated that R. blairi is quite aquatic. Lych (1985) also foud that both youg ad adult R. blairiwere easily ecoutered ithe fall alog streams ad rivers i Nebraska. I southwester Kasas, Collis ad Collis (1991) foud R. blai' abudat i grasslads. At Cheyee Bottoms i cetral Kasas, R. blairiprefers prairie streams ad marshes (Irwi ad Collis, 1987). I Colorado the species ihabits the easter foothills, shortgrass prairie ow utilized for pasture ad grai farmig, sady soils, the margis of aquatic habitats, plais grasslad, sadhills, rocky cayo bottoms, ad semiarid plais (Cousieau ad Rogers, 1991; Gillis, 1979; Hammerso, 1982a). The species occurs at elevatios below 1828 m i Colorado (Hammerso 1982a). Lardie ad Black (1981) characterized the habitat of R. blairi i the xeric Cimarro Gypsum Hills regio of orthwester Oklahoma as Mixedgrass Plais Biotic District, Grasslad Faual Regio, Shortgrass Plais Biotic District, ad grammabuffalo grasslad. The area also cotais escarpmets, badlads, sadstoe outcrops, sad dues, cavers, ad sikholes. Raa blai' has bee foud i a umber of caves i Oklahoma (Black, 1969, 1973; Bozema et al., 1987; Collis ad Collis, 1988). I Texas the species is adapted to dry prairies ad plais, i which it seeksout aquatic habitats (Garread Barker, 1987). J.E. Platz (pers. comm.) foud that R. blairiwas typically foud i Texas at shallow playa lakes, roadside ditches, ad more recetly at cattle stock taks. I New Mexico, Scott ad Jeigs (1985) reported that R. blairi is foud i a area that has umerous playa lakes of variable duratio. Specimes were take at rivers ad pods (humamade or atural) i temporary streambeds. I Arizoa, extreme aridity limits available habitat ad the species is mostly restricted to opermaet or semipermaet livestock taks ad irrigatio sloughs, as well as a occasioal permaet pool i a itermittet stream (Frost ad Bagara, 1977). Populatios of R. blairi were foud at elevatios from 12001800 m i Arizoa by Frost ad Platz (1983). I additio to some of the precedig, Stebbis (1985) metioed oak ad oakpie woodlad as habitat, ad he idicated that elevatios from 110 2590 mare ihabited. Breedig sites are quite variable ad have icluded the followig: roadside addraiage ditches, marshes, raior temporarypools, flooded areas, farm pods, lakes, livestock taks, streams, rivers, pools i itermittet streams, irrigatio sloughs, ad ashsettlig pods of a coalfired power plat (Black ad Sievert, 1989; Brow ad Morris, 1990; Caldwell ad Glass, 1976; Clarkso ad Rorabaugh, 1989; Frost ad Bagara, 1977; Hammerso, 1982a; Hillis, 1981; Johso, 1977, 1987; Kruse ad Fracis, 1977; Sco ad Jeigs, 1985). Although both letic ad lotic sites are utilized, the former seem to be preferred or more frequetly used. Male R. blairifrequetly call i a floatig positio at the water surface (Black ad Sievert, 1989; Hillis, 1981). Frost ad Bagara (1977) played tape recordigs of matig calls of R. blairito sexually active male R. blairi i the eveig before callig bega. Males respoded vocally to these recordigs, ad they also ofte tured to face (ad sometimes approached) the source of the recordig. However, playbacksof matig calls of Raa chiricahuesisfailed to evoke ay respose frommale R. blairiuder similar circumstaces. Breedig ca occur aytime from February through October, depedig o local evirometal coditios (Axtell, 1977; Axtell ad Haskell, 1977; Black ad Sievert, 1989; Caldwell ad Glass, 1976; Collis, 1982; Collis ad Collis, 1331; Dulap ad Kruse, 1976; Ehrlich, 1979; Frost ad Bagara, 1977; Frost ad Platz, 1983; Gray ad Stegall, 1986; Hammerso, 1982a; Hillis, 1981; Johso, 1977, 1987; Lych, 1978,1985; Post adpettus, 1967; Rudquist et al., 1978; Sco ad Jeias. 1985; Stebbis. 1985). Hammerso (1982a) sometimesfo~d~ewl~ laid eggs ad large tadpoles together, ad he suggested that breedig might occur more tha oce a year at a give site. Tadpoles ca metamorphose at ay time durig the summer depedet upo whe eggs were laid (Hammerso, 1982a). However, whe eggs are laid i late summer or fall, tadpoles overwiter ad trasform the ext sprig (Axtell ad Haskell, 1977; Collis, 1982; Collis ad Collis, 1331; Hammerso, 1982a; Johso, 1987; Lych, 1985; Sco ad Jeigs, 1985). Newly metamorphosed juveiles averaged 27 mm i legth i Missouri (Johso, 1987), ad the supralabial stripe is ofte well defied i the youg (Stebbis, 1985). The usual aual activity period was foud by Collis (1982), Collis ad Collis (1991), Hammerso (1982a), ad Johso (1987) to exted from early sprig (Feb.Apr.) ito fall (Oct.). Hammerso (1982a) oted that i the fall, large umbers of R. blairigathered at some pods used as overwiterig sites. Sometimes the species is active i witer (Collis, 1982; Collis ad Collis, 1991). Witer activity uder ice i Oklahoma was reported by Black et al. (1976). Smith et al. (1983) oted four froze R. blairi i December that had apparetly died after leavig a Missouripod through holes i the ice. Frost ad Bagara (1977) foud that after beig disturbed, R. blairiofte sought refuge i vegetatio surroudig bodies of water, whereas R. chiricahuesis almost always jumped ito the water. Gillis (1979) observed that whe R. blairi were subjected to dehydratio stress, they assumed a watercoservatio posture (legs tucked ear body, vetral surface pressed o substrate). Raapipimsoly adoptedthisposture whemuchmore body moisture was lost (Gillis, 1979). Reports of atural hybrids betwee R. blai'ad three other species of leopard frogs iclude: R. blairix R. berladieri, (Littlejoh ad Oldham, 1968; Platz, 1972, 1981); R. blairix R. pipies, (Black, 1976; Cousieau ad Rogers, 1991; Dulap, 1979; Dulap ad Kruse, 1976; Hammerso, 1982a; Kruse, 1981; Kruse ad Dulap, 1976; Lych, 1978,1985; Pace, 1974); R. blairix R. spheocephala, (Axtell, 1977; Black, 1976; Collis, 1982; Johso, 1977, 1987). Notable amog these were the studies of Platz (1972,1981) o atural hybrid R. blairix R. berladim.i Texas. I 1969, twelve of 138 idividuals were hybrids (eight F1, four iferred backcrosses), while i 1975, six of 94 frogs were iferred backcrosses. These aimals were primarily idetified o the basis of isozyme pheotypes which coflicted with idetificatio based o morphology ad color paer. Lych (19781, usig his ow data pooled with that of Dulap ad Kruse (1976) ad
Pace (1974), foud atural hybrid R. blairi x R. pipies at 13 of 39 sympatric localities i Nebraska. The pooled frequecy of hybrids i most localities i Iowa, South Dakota, adnebraska raged from 1.0 6.2% except i two areas where it was cosiderably higher (11.6%, 15.8%). Kruse ad Dulap (1976) foud that atural hybrid R. blairi x R. pipies could be idetified by albumi pheotype; likewise, Axtell (1977) idetified atural hybrid R. blairix R. spheocephala by LDH pheotype. Habitat alteratio by humas ad ustable ecological coditios have bee suggested as causes of the atural hybridizatio (Collis, 1982; Hammerso, 1982a; Hillis, 1981; Johso, 1987; Lych, 1978). Artificial laboratory hybridizatios have bee carried out betwee R. blairi ad eight other raid species: R. areolata, R. berladieri, R. chiricahuesis, R. megapoda, R. motezumae, R. palustris, R. pipies, ad R. spheocephala. The objectives of these hybridizatios were: to aid i the idetificatio of atural hybrids (Axtell, 1977; Dulapad Kruse, 1976; Kruse ad Dulap, 1976; Platz, 1972); to characterize isozyre pheotypes of hybrids ad backcrosses (Dulap, 1979, 1982); to determie degree of geetic divergece (Cuellar, 1971; Mecham, 1968,1969; Post ad Pettus, 1966); to study postmatig reproductive isolatio (Frost ad Bagara, 1977; Frost ad Platz, 1983; Post ad Pettus, 1966); ad to carry out likage aalyses (Dulap, 1982; Wright et al., 1983). The karyotype has 2 26, six metacetric ad seve submetacetric chromosomal pairs, ad dimorphic ucleolar orgaizer regios o chromosome pair o. 10 (Ward, 1977). Frost ad Platz (1983) foud abormal meiotic activity i six laboratoryproduced male hybrid R. chiricahuesis X R. blairi, with formatio of a average of six bivalets (chromosomal elemet umbers raged from 18.722.0). Eight of ie hybrid females failed to show sigificat oocyte developmet; the ith female cotaied oly a few eggs, all of which were immature. Seve hybrid males had mea sperm couts of less tha 6% of those of cotrols (Frost ad Platz, 1983). Major phylogeetic studies ivolved use of artificial laboratory hybridizatios (Cuellar, 1971; Mecham, 1969) ad cladistic aalyses of biochemical data (Hillis, 1988; Hillis ad Davis, 1986; Hillis et al., 1983). Zoogeography i the Great Plais, Prairie Peisula, ad alog the Mississippi River i the cetral Midwest was discussed by Brow ad Morris (1990) ad Brow et al. (i press). Frost ad Bagara (1977) commeted o zoogeography i the Chihuahua Desert. Reviews of historical views ad ewer evidece pertaiig to the taxoomy, speciatio, ad evolutio of R. blai ad other leopard frog species were preseted by Brow (1973), Dubois (19771, Hillis (19881, ad Pace(1974). ThediscoveryofR. blai ithe middle of Moore's (1946) proposed "orthsouth gradiet of hybrid iviability" through the Uited States i R. pipies, sew lato, provided importat evidece that Moore's (1944, 1946, 1949) siglespecies hypothesis was icorrect (Brow, 1973, Brow ad Brow, 1972). O a lesser scale, Smith (1961) cocluded that there was a wide, orthsouth zoe of itergradatio across cetral Illiois betwee two forms he recogized ("Raapipiespi>ies" i the orth ad "Raapipies spheocephala" i the south). Later, Brow ad Morris (1990) ad Morris et al. (1983) showed that the suspected itergrades were actually R. blairi. Comprehesive studies of reproductive isolatig mechaisms ivolvig R. blairiiclude Frost ad Platz (1983) i Arizoa ad New Mexico with three other species, ad Hillis (1981) i Texas ad Oklahoma with two other species. Both studies cocluded that prematig isolatio was well developed. Hillis (1988) summarized isolatig mechaisms amog R. blairi ad other sympatric, related, raid species. May other papers provided additioal data or gave pertiet commetary. Several papers have addressed predatio o R. blairi. Kruse ad Fracis (1977) foud that three predatory kh species cosumed tadpoles of R. blai' ad three other aura species much more readilythathe fishcosumed Bullfrogtadpoles (Raa catesbeiaa). Ehrlich (1979) observed that Bullfrog tadpoles uhesitatigly cosumedtheeggs ad ewly hatchedlarvae of R. blai. Large Bullfrogs were foud by Smith (1977) to readily oriet to distress calls of recetly metamorphosed R. blairi for purpose of cosumptio, ad Hammerso (l982a, 1982b) implicated Bullfrogs as extirpatig populatios of R. blai' through predatio or competitio. However, Hayes ad Jeigs (1986) questioed the importace of Bullfrog predatio i the declie of wester North America raids. At the Uiversity of Kasas Natural History Reservatio, Fitch (1982) foud that R. blairimade up a importat part of the diet of several species of sakes. Hammerso (1982a) oted the distress calls produced by R. blai' whe beig preyed upo by Wester Terrestrial Garter Sakes (Thamophk elegas). Raa bhiri were a sigificat food item foud i a est of the Mississippi Kite (Ictiia misisippiesk) by Robiso (1957). The remais of the carcasses of six small R. blai were foud by Parrley ad Tyler (1978) aroud the etrace of a burrow of a Blacktail Prairie Dog (Cyomys ludoviciaus) occupied by a Burrowig Owl (Athee cuicularia). Specific iformatio o the food habits of R. blairi is scaty. Black (1973) examied the alimetary caals of several R. blairi collected i total darkess i gypsum caves i orthwester Oklahoma. He foud the remais of crickets (Family Gryllacrididae), beetles (Carabidae ad Catopidae), ad flies (Heleomyzidae). The frogs ad ivertebrates occurred ear guao mouds. Hammerso (1982a) foud a R. blairi i Colorado that had its stomach full of grasshoppers, ad Creel (1963) captured a large R. blairi(svl 114 mr) i Texas cosumig a bat (Pipistrellussubflaous) with oly oe of its wigs protrudig from the frog's mouth. I cotrast to the uspotted dorsums foud at low frequecies i populatios of R. pipies (termed Bursi mutats) ad R. spheocephah, o dorsally uspotted frogs were foud by Brow ad Fuk (1977) amog 219 preserved R. blairi from Illiois. Details of timeefficiet, low cost procedures for laboratory rearig of tadpole ad postmetamorphic R. blairiwere preseted by Frost (1982). Platyhelmith parasites of R. blai i Nebraska were extesively surveyed by Brooks (1976) ad Brooks ad Welch (1976). Physiological effects of experimetal aaerobiosis were studied by Rose ad Drotma (1967). Gillis (1979) demostrated the highertolerace to water loss by adult R. blairi, whe compared to adult R. pipies. Isozyme studies ot heretofore metioed iclude Levy ad Salthe (1971,1974) ad Salthe (1969). The declie or extictio of populatios has bee oted by Christiase ad Bailey (1991), Clarkso ad Rorabaugh (1989), Cousieau ad Rogers(lWl), Frost (1983), Frost ad Bagara (1977), Hammerso (1982a, 1982b), Hayes ad Jeigs (1986), ad Platz (1981). Suggested causes icluded: water pollutio; predatio by itroduced game fishes; groudwater pumpig; itroductio of exotic fishes ad amphibias; agricultural developmet; icreased aridity/drought; habitat loss or alteratio; toxicats; competitio with R. berladieri; ad predatio by, competitio with, ad/or larval ihibitio by Bullfrogs (R. catesbeiaa). Remarks. I publicatios durig the seve years prior to the formal descriptio of R. blairi as a ew species, the followig commo ames were used for the frog: DF complex, DF morphotype, DF frogs, wester call type, wester call race, wester type, wester form, wester, souther plais type, souther plais form, souther plais, S. Plais, S. plais, plais type, souther plais ("wester") type, "wester" (souther plais) type, plais form, high plais frogs, cetral plais form, ad wester (plais) type. Some iformatio o R. blairiwas published uder the ame "R. pipies" before R. blairi was described as a ew taxo (see Brow, 1973). Coversely, some iformatio published uder the ame R. bhiri apparetly reprsets older data collected from R. pipies ad/or other leopard frog species. Etymology. The specific ame hoors the late Dr. W. Frak Blair, Professor of Zoology, The Uiversity of Texas at Austi. Literature Cited Axtell, C.B. 1976[19773. Comparisosof morphology, lactatedehydrogease, ad distributio of Raa blairiad Raa utricularia i Illiois ad Missouri. Tras. Illiois St. Acad. Sci. 693748. Axtell, R.W. ad N. Haskell. 1977. A iterhiatal populatio of Pseudacrisstreckerifrom Illiois, with a assessmet of its postglacial dispersio history. Nat. Hist. Misc. Chicago Acad. Sci. (202):l8. Balliger, R.E. ad J.D. Lych. 1983. How to kow the amphibias ad reptiles. Wm. C. Brow Co., Dubuque, Iowa. Behler, J.L. ad F.W. Kig. 1979. The Audubo Society field guide to North America reptiles ad amphibias. Alfred A. Kopf, New York. Black, J.H. 1969.A cavedwellig populatioofambystoma tigrium mamriium i Oklahoma. J. Herpetol. 3:183184..1973. A checklist of the cave faua of Oklahoma: Amphibia. Proc. Oklahoma Acad. Sci. 53:3337. 1976. Oklahoma leopard frogs. Bull. Oklahoma Herpetol. Soc. 1:610.
, T. Hukapiller, ad D. Dawso. 1976. Witer activity i Oklahoma frogs. Bull. Oklahoma Herpetol. Soc. 1:22. ad G. Sivert. 1989.A field guide to amphibias of Oklahoma. Oklahoma Dept. Wildl. Coserv., Oklahoma City. Bozema, S., K.S. Johso, B. Jagow, B. Baker, G. Kowalski, J. Looey, P. Ziegler, A. Dodds, S. Martiez,adP. Riley. 1987.The D.C. Jester Cave system. Oklahoma Udergroud 14:146. Brooks, D.R 1976. Parasites of amphibias of the Great Plais. Part 2. Platyhelmiths of amphibias i Nebraska. Bull. Uiv. Nebraska St. Mus. 106592. ad N.J. Welch. 1976. Parasites of amphibias of the Great Plais. I. The cercaria of C@balogoimus breuicius Igles, 1932 flrematoda: Cephalogoimidae). Pmc. Helmithol. Soc. Washigto 439293. Brow, L. (ot L.E. Brow). 1985. Grasslads. The Audubo Society Nature Guides. Alfred A. Kopf, New York. Brow, L.E. 1973. Speciatio i the Raapipiescomplex. Amer. Zo 01. 137379. ad J.R. Brow. 1972. Call types of the Raapipiemcomplex i Illiois. Sciece 176:92&929. ad RS. Fuk. 1977. Absece of dorsal spottig i two species of leopard frogs (Aura: Raidae). Herpetologica 33290293. ad MA. Morris. 1990. Distributio, habitat, ad zoogeography of the Plais Leopard Frog (Raa blai) i Illiois. Illiois Nat. Hist. Surv. Biol. Notes (136):l6.,, ad T.R Johso. I press. Zoogeography of the Plais Leopard Frog (Raa blaid. Bull. Chicago Acad. Sci. Caldwell, J. ad G. Glass. 1976. Vertebrates of the Woodso Couty State Fishig Lake ad Game Maagemet Area, p. 6276. I Prelimiary ivetory of the biota of Woodso Couty State Fishig Lake ad Game Maagemet Area. Report St. Biol. Surv. Kasas. Chriitiase. J.L. ad RM. Bailw. 1Wl.The salamaders ad frogs of Iowa. 1' Dept. Nat. ~ es. Nogame Tech. Ser. (3):l24. Clarke, RE. 1984. Frogs adtoads i Kasas. Kasas School Nat. 303 IS. * Clarkso, R.W. ad J.C. Rorabaugh. 1989. Status of leopard frogs (Raapipiescomplex: Raidae) i Arizoa ad southeaster Califoria. Southwest. Nat. 34531538. Collis, J.T. 1974. Amphibias ad reptiles i Kasas. 1st ed. Uiv. Kasas Mus. Nat. Hist. Publ. Educ. Ser. (1):ix + 283 p.. 1977. Kasas frogs &toads. Kasas Fish Game 34:1216. 1982. Amphibias ad reptiles i Kasas. 2d ed. Uiv. Kasas Mus. Nat. Hist. Publ. Educ. Ser. (8):xiii + 356 p. ad S.L. Collis. 1991. Reptilesad amphibias of the Cimarro Natioal Grasslads, Morto Couty, Kasas. U.S. Forest Service, Elkhart, Kasas. Collis, M. a d N. Collis. 1988. Widow Cave, Major Couty, Oklahoma (MJ042). Oklahoma Udergroud 15:&19. Coat, R ad J.T. Collis. 1991. A field guide to reptiles ad amphibias easter ad cetral North America. 3rd ed. Houghto Miffli Co., Bosto. Cousieau, M. a d K. Rogers. 191. Observatios o sympatric Raa pipies, R. blairi, ad their hybrids i easter Colorado. J. Herpetol. 25114116. Creel, G.C. 1963. Bat as a food item of Raapipies. Texas J. Sci. 15: 104106. Cuellar, H.S. 1971. Levels of geetic compatibility of Raa areolata with southwester members of the Raapipiescomplex (Aura: Raidae). Evolutio 25:399409. Dixo, J.R. 1987. Amphibias ad reptiles of Texas with keys, taxoomic syopses, bibliography, ad distributio maps. W.L. Moody, Jr., Nat. Hist. Ser. No. 8. Texas A&M Uiv. Press, College Statio. Dubois, A. 1977. Les problemes de I'esp5ce chez les amphibies aoures, p. 161284. IC. Bocquet, J. GCermot, ad M. Lamotte (eds.), Les probemes de I'esp5ce da le rege aimal. Tome 11. MCm. No. 39. Soc. Zool. Frace. Dulap, D.G. 1979. Hemoglobi pheotypes i the frogs Raapipies, Raa blairi, their hybrids ad a backcross. Comp. Biochem. Physiol. 62B:167173. 1982. Likage aalysis of the trasferri, albumi, ad hemoglobi loci i leopard frogs. J. Heredity 73247248. ad K.C. Ktuse. 1976. Frogs of the Raapipiescomplex i the ortherad cetral plais states. Southwest. Nat. 20:559571. Ehrlich, D. 1979. Predatio by Bullfrog tadpoles (Raa catesbeiaa) o eggs ad ewly hatched larvae of the Plais Leopard Frog (Raa blai). Bull. Marylad Herpetol. Soc. 152526. Eshelma, R.E. 1975. Geology ad paleotology of the early Pleistocee (late Blaca) White Rock faua from orthcetral Kasas. Claude W. Hibbard Mem. Vo1.4, Mus. Paleotol. Uiv. Michiga Pap. Paleotol. (13):iv + 60 p. Fitch, H.S. 1982. Resources of a sake commuity i prairiewoodlad habitat of ortheaster Kasas, p. 8397. I N.J. Scott, Jr. (ed.), Herpetological commuities, a symposium of the Society for the Study of Amphibias ad Reptiles ad the Herpetologists' League, August 1977. Wildl. Res. Report 13, U.S. Dept. Iterior, Fish Widl. Serv., Washigto, D.C. Fritts,T.H., RD. Jeigs,adN.J. Scott, Jr. 1984.Areviewoftheleopard frogs of New Mexico. Upubl. report submitted to the New Mexico Dept. Game ad Fish. Frost,D. 1983. Past occurrece of Acriscrepitas (Hylidae) i Arizoa. Southwest. Nat. 28:105. Frost, J.S. 1982. A timeefficiet, low cost method for the laboratory rearig of frogs. Herpetol. Rev. 137377. ad J.T. Bagara. 1976. A ew species of leopard frog (Raa pipiescomplex) from orthwester Mexico. Copeia 1976332 338. ad. 1977. Syrpatry betwee Raa blairi ad the southem form of leopard frog i southeaster Arizoa (Aura: Rai dae). Southwest. Nat. 22:443453. ad J.E. Platz. 1983. Comparative assessmet of modesof reproductive isolatio amog four species of leopard frogs (Raa pipies complex). Evolutio 376678. Garrett, J.M. ad D.G. Barker. 1987. A field guide to reptiles ad amphibias of Texas. Texas Mothly Press, Austi. Gillis, J.E. 1979. Adaptive differeces i the water ecoomies of two species of leopard frogs from easter Colorado (Amphibia, Aura, Raidae). J. Herpetol. 13:445450. Gray, P. ad E. Stegall. 1986. Distributio ad status of Suecker's Chorus Frog (Pseudacris streckeri strecke) i Kasas. Tras. Kasas Acad. Sci. 89:8185. Hammerso, G.A. 1982a.Amphibias ad reptiles i Colorado. Colorado Div. Wild., Dever. 1982b. Bullfrogelimiatigleopardfrogs icolorado? Herpetol. Rev. 13:115116. Hayes, M.P. ad M.R. Jeigs. 1986. Declie of raid frog species i wester North America: are Bullfrogs (Raa catesbeiaa) resposible? J. Herpetol. 20:49&509. Hillis, D.M. 1981. Prematig isolatig mechaisms amog three species of the Raapipiescomplex itexas ad souther Oklahoma. Copeia 1981:312319.. 1982. Morphological differetiatio ad adaptatio of the lar vae of Raa berladieri ad Raa spheocephala (Raapipies complex) i sympatry. Copeia 1982:168174. 1988. Systematics of the Raapipiescomplex: puzzle ad paradigm, p. 3963.1R.F. Johsto, P.W. Frak, adc.d. Micheer (eds.), A. Rev. Ecol. Syst., Vol. 19. Palo Alto, Califoria. ad S.K. Davis. 1986. Evolutio of ribosomal DNA: fifty millio years of recorded history i the frog geus Raa. Evolutio 40: 12751288., J.S. Frost, ad D.A. Wright. 1983. Phylogey ad biogeography of the Raapipiescomplex: a biochemical evaluatio. Syst. Zool. 32:132143. Holma, J.A. 1%9.The Pleistocee amphibias ad reptiles oftexas. Publ. Mus. Michiga St. Uiv. Biol. Ser. 4:161192.. 1971. Herpetofaua of the Sadahl local faua (Pleistocee: Illioia) ofkasas. Cotrib. Mus. Paleotol. Uiv. Michiga23 349355.. 1972. Herpetofaua of the Kaopolis local faua (Pleistocee: Yarmouth) of Kasas. Michiga Acad. 533798.. 1977. The Pleistocee (Kasa) herpetofaua of Cumberlad Cave, Marylad. A. Caregie Mus. 46:157172..1984. Herpetofauas of the Duck Creek ad Williams local fauas (Pleistocee: Illioia) of Kasas. Cotrib. Quat. Vert. Paleotol.: Vol. Mem. Joh E. Guilday, Spec. Publ. Caregie Mus. Nat. Hist. (8):2038. 1987. Climatic sigificace of a late Illioia herpetofaua from southwester Kasas. Cotrib. Mus. Paleotol. Uiv. Michiga 27:129141. Irwi, K.J. ad J.T. Collis. 1987. Amphibias ad reptiles of Cheyee Bottoms, p. 401432. I Cheyee Bottoms, a evirometal assessmet. Publ. Kasas Biol. Surv. ad Kasas Geol. Surv. Johso, T.R. 1977. The amphibias of Missouri. Uiv. Kasas Mus. Nat. Hist. Publ. Educ. Ser. (6):ix + 134 p.
. 1982. Toads ad frogs of Missouri [poster]. Missouri Dept. Coserv., Jefferso City.. 1987. The amphibias ad reptiles of Missouri. Missouri Dept. Coserv., Jefferso City. Kasper, S. ad D. Parmley. 1990. A late Pleistocee herpetofaua from the lower Texas Pahadle. Texas J. Sci. 42:2892%. Korky, J.K. 1978. Differetiatioof the lame of members ofthe Raa pipiescomplex i Nebraska. Copeia 1978:455459. Kruse, K.C. 1981. Phootactic resposes of female Norther Leopard Frogs (Raapipies) to Raa blairi, a presumed hybrid, ad cospecific matig trills. J. Herpetol. 15145150. ad D.G. Dulap. 1976. Serum albumis ad hybridizatio i two species of the Raapipiescomplex i the orth cetral Uited States. Copeia 1976:394396. ad M.G. Fracis. 1977. A predatio deterret i larvae of the Bullfrog, Raa catesbeiaa. Tras. Amer. Fish. Soc. 106:248 252. Lardie, R.L. 1982.A prelimiary checklist ofthe amphibias ad reptiles of orthwester Oklahoma (excludig the Oklahoma Pa hadle). Bull. Oklahoma Herpetol. Soc. 7:s78. ad J.H. Black. 1981. The amphibias ad reptiles of the Cimarro Gypsum Hills regio i orthwester Oklahoma. Bull. Oklahoma Herpetol. Soc. 576125. Levy, P.L. ad S.N. Salthe. 1971. Kietic studies o variat hearttype lactate dehydrogeases i the frog, Raapipies. Comp. Biochem. Physiol. 39B:343355. ad. 1974. Studies o the variability of muscletype lactate dehydrogease i the frog, Raa pipies. Comp. Biochem. Physiol. 48B:355377. Littlejoh, M.J. ad R.S. Oldham. 1968. Raapipiescomplex: matig call structure ad taxoomy. Sciece 162:10031005. Lych, J.D. 1978. The distributio of leopard frogs (Raa blairi ad Raapipies) (Amphibia, Aura, Raidae) i Nebraska. J. Herpetol. 12:157162.. 1985. Aotated checklist ofthe amphibias ad reptiles of Nebraska. Tras. Nebraska Acad. Sci. 133357. McAlister, W.H. 1962. Variatio i Raapipies Schreber i Texas. Amer. Midl. Nat. 67534363. Mecham, J.S. 1968. Studies o evolutioary effects of isolatio i the Raa pipies complex. Yr. Bk. Amer. Phiios. Soc. 1968:314 316.. 1969. New iformatio from experimetal crosses o geetic relatioships withi the Raa pipies species group. J. Exp. Zool. 170:169180. 1971. Vocalizatiosofthe leopard frog, Raapipies, ad three related Mexica species. Copeia 1971:505516., M.J. Littlejoh, R.S. Oldham, L.E. Brow, ad J.R. Brow. 1973. A ew species of leopard frog (Raapipiescomplex) from the plais of the cetral Uited States. Occ. Pap. Mus. Texas Tech Uiv. (18):l11. Moore, JA. 1944. Geographic variatio i RaapipiesSchreber of easter North America. Bull. Arer. Mus. Nat. Hist. 82345370.. 1946. Icipiet itraspecific isolatig mechaisms i Raa pipies. Geetics 31:304326..1949. Geographic variatio of adaptive characters i Ram pipiesschreber. Evolutio 5124. Morris, M.A., R.S. Fuk, ad P.W. Smith. 1983. A aotated bibliography ofthe Illiois herpetological literature 19601980, ad a updated checklist of species of the state. Illiois Nat. Hist. Surv. Bull. 33123138. Pace,A.E. 1974. Systematic ad biological studies of the leopard frogs (Raapipies complex) of the Uited States. Misc. Publ. Mus. 2001. Uiv. Michiga (148):l140. Parmley, D. 1988. Additioal Pleistocee amphibias ad reptiles from the Seymour Formatio, Texas. J. Herpetol. 228287. ad J.D. Tyler. 1978. Herpetological prey at a Burrowig Owl hole. Bull. Oklahoma Herpetol. Soc. 313. Platz, J.E. 1972. Sympatric iteractio betwee two forms of leopard frog (Raapipiescomplex) i Texas. Copeia 1972:232240..1976. Biochemical ad morphological variatio of leopard frogs i Arizoa. Copeia 1976660672.. 1981. Suture zoe dyamics: Texas populatios of Raa berladieriad R. blairi. Copeia 1981:733734. Post, D.D. ad D. Pettus. 1966. Variatio i Raa pipies (Aura: Raidae) of easter Colorado. Southwest. Nat. 11:476482. ad. 1967. Sympatry of two members of the Raapipies complex i Colorado. Herpetologica 23323. Robiso, T.S. 1957. Notes o the developmet of a brood of Mississippi Kites i Barber Couty, Kasas. Tras. Kasas Acad. Sci. 60:174180. Rogers, K.L. 1982. Herpetofauas of the Courlad Caal ad Hall Ash local fauas (Pleistocee: Early Kasas) of Jewell Co., Kasas. J. Herpetol. 16:174177. 1984. Herpetofauas ofthe Big Sprigs ad Horet's Nest quarries (ortheaster Nebraska, Pleistocee: late Blaca). Tras. Nebraska Acad. Sci. 12:8194., CA. Repeig, R.M. Forester, E.E. Larso, SA. Hall, G.R. Smith, E. Aderso, ad T.J. Brow. 1985. Middle Pleistocee (late Irvigtoia: Nebraska) climatic chages isouthcetral Colorado. Natl. Geograph. Res. 1535563. Rose, F.L. ad RB. Drotma. 1967.Aaerobiosis i a frog, Raapipies. J. Exp. Zool. 166:427432. Rudquist, E.M., E. Stegall, D. Grow, ad P. Gray. 1978. New herpetological records from Kasas. Tras. Kasas Acad. Sci. 81:73 77. Salthe, S.N. 1969. Geographic variatioofthe lactate dehydrogeases of Raapipies ad Raapalustris. Biochem. Geet. 2:271 303. Scott, N.J. Jr. ad R.D. Jeigs. 1985.The tadpoles of five species of New Mexica leopard frogs. Occ. Pap. Mus. Southwest. Biol. (3):l21. Smith, A.K. 1977. Attractio of Bullfrogs (Amphibia, Aura, Raidae) to distress calls of immature frogs. J. Herpetol. 11:234235. Smith, D.D., R. Powell, T.R. Johso, ad H.L. Gregory. 1983. Life history observatios of Missouri amphibias ad reptiles with recommedatios for stadardized data collectio. Tras. Missouri Acad. Sci. 17:3758. Smith, H.M. 1956. Hadbook of amphibias ad reptiles of Kasas. Uiv. Kasas Mus. Nat. Hist. Misc. Publ. @):I356. 1978. Aguide to field idetificatio amphibias of NorthAmerica. Golde Press, New York, WesterPubl. Co., Racie, Wiscosi. Smith, P.W. 1961. The amphibias ad reptiles of Illiois. Illiois Nat. Hist. Surv. Bull. 28:l298. Stebbis, RC. 1985. A field guide to wester reptiles ad amphibias. 2d ed.. Houghto MiMi Co., Bosto. Tihe, JA. 1954. AKasas Pleistocee herpetofaua. Copeia 1954217 221. Ward, O.G. 1977. Dimorphic ucleolar orgaizer regios i the frog Raa blairi. Ca. J. Geet. Cytol. 195157. Wright, DA., C.M. Richards, J.S. Frost, A.M. Camozzi, ad B.J. Kuz. 1983. Geetic mappig i amphibias, p. 287311. I M.C. Rattazzi, J.G. Scadalios, ad G.S. Whitt (eds.), Isozymes: curret topics i biological ad medical research. Geetics ad evolutio, Vol. 10. Ala R. Liss, New York. Iaure E. Brow, Departmet of Biological Scieces, Illiois State Uiversity, Normal, IL 61761. Primary editor for this accout, Joh D. Lych. Published 30 November 1992 ad Copyright O 1992 by the Society for the Study of Amphibias ad Reptiles. m