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Speciation Within the Chorus Frog Pseudacris Triseriata: Morphometric and Mating Call Analyses of the Boreal and Western Subspecies

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Speciation Within the Chorus Frog Pseudacris Triseriata: Morphometric and Mating Call Analyses of the Boreal and Western Subspecies Copeia, 1989(3), pp. 704-712 Speciation within the Chorus Frog Pseudacris triseriata: Morphometric and Mating Call Analyses of the Boreal and Western Subspecies JAMES E. PLATZ Data from mating call and morphological analyses of the boreal and western subspecies of the chorus frog Pseudacris triseriata were obtained along a transect from South Dakota to Oklahoma. Two call types are evident. At the northern end of the transect calls are long and low in pulse rate. Both traits had low variability relative to populations further south. The southern end of the transect contained populations with a much shorter call and pulse rates approximately twice those in the north. Identical long and short call types are evident in Col- orado as well. Limited data from Oklahoma indicates that the call of P. triseriata feriarum is more like that of P. t. maculata. Multivariate stepwise discriminant analyses performed on data sets containing only morphological measurements produced two clusters of populations which correspond on geographic grounds to two currently recognized subspecies, P. t. maculata and P. t. triseriata. When call data are included, the same two clusters are identified suggesting concor- dance between call type and morphology. The data indicate that the two entities represent separate species and that the zone of overlap is much more extensive than previously depicted. FSEUDACRIS triseriata (Wied) is one of the extending north along the Atlantic as far as 1 smallest North American hylids. Accord- New Jersey. Pseudacris t. kalmi, the New Jersey ing to Tordoff and Pettus (1977), some live Chorusin Frog, is restricted in range, occurring excess of 6 yr. In eastern Colorado, Spencer in parts of Delaware, Maryland, southern New (1964) found it to be an anuran of limited va-Jersey, New York and eastern Pennsylvania. Ac- gility with few individuals moving more than cording to Conant (1975) it intergrades with P. 750 m from the hatch site. It is also the most t.feriarum in eastern Pennsylvania and at Staten widespread species of chorus frog in North Island, New York. America, occurring from the Gulf of Mexico to The criteria used to distinguish these four New York and southern Ontario in the east, subspecies hinge, to a large degree, on variable and from parts of Texas and Arizona in the dorsal skin patterns. The boreal form has short- southwest, north to Alberta and the Northwest er legs and often a dorsal pattern of spots in Territory in Canada. three longitudinal rows distinguishing it from Conant (1975) recognized four subspecies of the Western form which has longer legs and P. triseriata. Three of these have extensive three well defined dorsal stripes. The Upland ranges. The Boreal Chorus Frog, P. triseriata Chorus Frog has thin broken stripes differen- maculata, occupies the northwestern limits tiating of it from the New Jersey Chorus Frog, the range: from northwestern Ontario, east which to has broad well defined stripes. Winnipeg in Manitoba, and south to New Mex- Although the present nomenclature has re- ico, and east including parts of Nebraska, South mained stable for the past 25 yr, the literature Dakota, Minnesota and Wisconsin. Pseudacris reveals t. an earlier history of repeated revision. triseriata, the Western Chorus Frog, replaces Since the original description of Hyla triseriata maculata in the south resulting in extensive sym-(now P. t. triseriata) by Wied (1839), four dif- patry in a broad arc extending through most ferent of generic names have been applied, as have Nebraska, eastern South Dakota, central Min- numerous specific epithets for this and other nesota and northern Wisconsin and Michigan. subspecies now recognized as P. triseriata. For Along its southern limit, the western formdetailed reviews see Smith (1956), Harper overlaps with and is then replaced by the (1955),Up- Schwartz (1957) and Cook (1964). land Chorus Frog, Pseudacris triseriataferiarum: The first serious efforts to understand geo- common over most of the Gulf Coast states, graphic variation within P. triseriata were those ? 1989 by the American Society of Ichthyologists and Herpetologists PLATZ-CHORUS FROG SPECIATION 705 of Smith and Smith (1952) and Smith lation variation. (1956). To provide such demic infor- At that time boreal, western and upland mation, forms I have conducted preliminary studies were considered subspecies of P. nigrita. over three Smith breeding seasons. The results indi- (1956) rediagnosed the Boreal Chorus cate that Frog the taxonomic (P. status of P. triseriata is nigrita septentrionalis) as P. n. maculata. not fully Based resolved. on differences in: 1) breeding habitat; 2) the dis- tinct nature of mating calls; and 3) the lack MATERIALS of AND METHODS intergradation among sympatric populations of P. n. nigrita and P. n. feriarum, Schwartz Call analysis.-In (1957) order to assess geographic recommended specific status for P. patterns n. feriarum for the boreal and western subspecies, as P. triseriata feriarum. He further 221 suggestedadult males representing 20 localities were that P. n. triseriata and P. n. maculata should be collected. Specific localities are given in Figure treated as subspecies of P. triseriata (i.e., P. t. 1. At the time of collecting, mating calls were triseriata and P. t. maculata, respectively). This obtained for 259 adults using Sony model TC- recommendation is based on evidence inter- 150 cassette recorders and microphones. Water preted as intergradation between P. t. feriarum temperature was recorded 2 cm posterior to an and P. t. triseriata reported by Smith (1956).individual as it called. Temperatures were de- Conant (1975) followed Schwartz (1957). termined For to the nearest 0.1 C using a Bailey convenience each of the four currently recog-BAT-12 fast reading telethermometer. All re- nized subspecies of P. triseriata will be referred cordings were made in night choruses during to in this paper as maculata, triseriata, feriarum, the breeding season (March-June). All calling or kalmi. individuals at a given locality were within my Smith's (1956) comparison of maculata hearing and distance. Each recording generally in- triseriata is critical to our current knowledge cluded of a minimum of one additional calling in- Pseudacris. It represents the only published dividual,study usually calling in a tandem mode. Af- of morphological variation, based on terlarge calls were recorded, each individual was cap- numbers of specimens collected over atured wide and placed in a labeled plastic bag coded geographic area. His conclusions rely onagainst ex- the taped calls and frozen in liquid ni- amination of over 800 individuals, including trogen 403 or placed on dry ice for later removal maculata from 112 localities in Canada and the of tissue samples. Specimens were subsequently United States, and 343 triseriata from 62 local- preserved in formalin and then transferred to ities (Smith and Smith, 1952) as well as 60%125 alcohol solution. presumed intergrades from 42 locations. Based A Kay Electrometrics Model 6061-B Sono- on his examination and measurements of these graph was used to generate sonograms. Three specimens, he differentiated present day macu- calls for each male were analyzed for dominant lata from triseriata in more specific terms frequencythan (DF), pulse number (PN), call dura- Conant (1975): maculata "differs by the tionpro- (CD), and pulse rate (PR). Because CD and portionately shorter tibia, femur, and foot; PR rel- are temperature dependent, recordings from atively shorter and narrower head; proportion- a test population composed of five adult male ately heavier body; and by the higher frequency maculata from an isolated pond (1 km south of of specimens with a spotted or mottled Glenpat- Cunningham Lake, Douglas Co., Nebras- tern." Examination of his table I indicates 343 ka) were made over a several week period in maculata and 430 triseriata were used to deter- order to obtain calls from a known group for a mine tibia-body length ratios. The resulting range of temperature values (10.5-17.7 C). mean values, given as percent of body length, These data (n = 39) were then used to deter- were 39.3 and 42.6, respectively. Computation mine the relationship between temperature and of confidence limits for the means (using his CD. The least squares regression was employed published SE of the mean times the appropriate and the resulting regression equation (Y = t value) indicates that the means are statistically -0.0974X + 2.277) was used to generate a different (P < 0.05). However, in this particular function, CD,, = CDamb. - (Tmb. - 14) comparison involving over 170 localities (sam- (-0.0974), to permit conversion of duration of ple sizes were not given), mean values and all SE subsequent calls to 14 C. Here "CD" refers were obtained by pooling all maculata vs all trise- to call duration in seconds and the subscript riata. This method resulted in very small "amb." SE refers to the temperature (T) of a re- values but precludes assessment of inter-popu- corded call or the duration (CD) from the son- 706 COPEIA, 1989, NO. 3 Fig. 1. Transect populations of Pseudacris triseriata maculata and P. t. triseriata. Populations connected by dashed lines represent P. t. triseriata. All others, except population 17, represent P. t. maculata. Sites collected are as follows: South Dakota: (1) Lake Oahe, Hwy. 1804, Potter Co., (2) Marsh Lake, Hamlin Co., (4) Springfield, Bon Homme Co. Iowa: (3) Lake Okoboji, Dickinson Co., (6) Wilson Island State Park, Potta- wattamie Co. Nebraska: (5) Beemer, Cuming Co., (7) Dodge Park, Omaha, Douglas Co., (8) 120th and Fort St., Omaha, Douglas Co., (9) Waterloo, Douglas Co., (10) Ogallala, Keith Co., (11) Cozad, Dawson Co. Kansas: (12) Concordia, Cloud Co., (13) Lawrence north, Douglas Co., (14) Lawrence south, Douglas Co., (15) Arkansas City, Cowley Co., (16) Hwy. 57, 15 km E St. Paul, Cherokee Co. Colorado: (18) Ft. Collins, Larimer Co., (19) 11 km W Fairplay, Park Co., (20) 3 km W Aroya, Lincoln Co.
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