The Journal of North American Herpetology
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ISSN 2333-0694 JNAHThe Journal of North American Herpetology Volume 2017(1): 19-27 29 March 2017 jnah.cnah.org SEASONAL INCIDENCE OF CAPTURE AND REPRODUCTION OF FIVE FOSSORIAL SNAKE SPECIES IN WEST VIRGINIA WALTER E. MESHAKA, JR.1, THOMAS K. PAULEY2, DANIELLE F. WRIGHT1, AND KELLI-MARIE HERRICK2 1Section of Zoology and Botany, 300 North Street, Harrisburg, Pennsylvania 17120, USA [author contact: wmeshaka@ pa.gov] 2Department of Biological Sciences, Marshall University, One John Marshall Drive, Huntington, West Virginia 25755 USA ABSTRACT — Museum specimens of five species of fossorial snakes collected in West Virginia during 1930–2000 were examined to determine monthly incidence of capture, adult body sizes, reproductive cycle, and clutch characteristics. Captures occurred over the shortest time in the year in the Eastern Earthsnake (Virginia valeriae valeriae) and Northern Red-bellied Snake (Storeria occipitomaculata occipitomaculata) and were longest in the Northern Brownsnake (S. dekayi dekayi). Male gonadal cycle conformed to the temperate pattern, whereas that of females tended towards a tropical pattern. Incidence of females nearing oviposition or parturition was highest during June–July for all species, and length of their reproductive seasons were generally in keeping with those of northerly populations of the respective species. Mean clutch sizes were largest in the Northern Brownsnake (mean = 20.5) and smallest in the single oviparous snake, the Eastern Wormsnake (mean = 2.8). Adult body sizes were similar to respective populations elsewhere within their ranges. The Mountain Earth Snake (V. pulchra) was the least represented species in this study. A meaningful degree of predictability existed in the life history traits examined in our study as they related to geographic trends of this Allegheny snake assemblage. INTRODUCTION Natural History and the Carnegie Museum of Natural Twenty-two species of snakes are native to West History. Body length was measured in cm as snout-vent Virginia (Green and Pauley, 1987). Five are small- length (SVL). Sex was determined by examination of bodied, predators of invertebrates, and fossorial in habits the gonads. Enlarged testes indicated sexual maturity (Green and Pauley, 1987). Three of these, the Eastern in males. The length and width in mm at the center of Wormsnake, Carphophis amoenus amoenus (Say, 1825), the right testis was each expressed as a percentage of Northern Brownsnake, Storeria dekayi dekayi (Holbrook, the SVL to provide a monthly distribution of testis size. 1836), and Northern Red-bellied Snake, Storeria Methods by Trauth et al. (1994) served as guidelines for occipitomaculata occipitomaculata (Storer, 1839), are size ranges associated with categories of ovarian follicles either widespread in or occur in approximately one and oviductal eggs and embryos. Follicles < 2 mm were half of West Virginia, and two species, the Mountain deemed immature. Vitellogenic ovarian follicles were Earthsnake, V. pulchra (Richmond, 1954) and the generally > 3 mm. Eastern Earthsnake, Virginia valeriae valeriae Baird and The largest ovarian follicle among those > 2.0 mm was Girard, 1853, are restricted to the northeastern corner used to provide a measure of monthly follicular growth. of the state (Green and Pauley, 1987; Powell et al., The largest oviductal eggs or embryos were measured for 2016). Basic life history data are lacking for all of them. a monthly estimate of respective growth rates. Clutch size Access to West Virginia specimens of these five species was estimated by counts of enlarged follicles, oviductal from the holdings of the Carnegie Museum of Natural eggs or counts of young. Mean values were followed by History and the herpetological collection of Marshall + 1 standard deviation. F-tests were performed to test University provided us with the opportunity to examine for statistical differences in variances around the means. certain life history traits. We were especially interested Two-tailed t-tests were performed to test for statistical in understanding their responses in relation to findings in differences between means. All statistical analyses were northern and southern latitudes. performed using Microsoft Excel version 2010. MATERIALS AND METHODS RESULTS Specimens of the five species of snakes were examined Species Accounts from the holdings of Marshall University Museum of Carphophis amoenus amoenus (Say, 1825) — One Journal of North American Herpetology 2017(1) 19 5 10 Male Female Percent testis length Percent testis width 4 Juvenile 8 3 6 2 4 No. Individuals 1 Percent (%) testis dimensions 2 0 0 0 2 4 6 8 10 12 0 2 4 6 8 10 12 Month Month Figure 1. Monthly incidence of captures of 58 male, 43 female, Figure 2. Monthly distribution of testis size as a percentage and 19 juvenile Eastern Wormsnakes (Carphophis amoenus of male snout-vent length (SVL) of 56 Eastern Wormsnakes amoenus) from West Virginia. (Carphophis amoenus amoenus) from West Virginia. 30 Male 30 Largest follicles Female Largest shelled egg length *Juvenile 26 25 24 16 18 15 14 * * SVL (cm) SVL * * * * * * 10 10 * * * * * * * 6 * Follicle and shelled egg size (mm) 5 2 0 0 0 2 4 6 8 10 12 0 2 4 6 8 10 12 Month Month Figure 3. Monthly distribution of follicle and ovum size in 37 Figure 4. Monthly distribution of snout-vent lengths (SVL) of Eastern Wormsnakes (Carphophis amoenus amoenus) from West 58 male, 43 female, and 19 juvenile Eastern Wormsnakes Virginia. (Carphophis amoenus amoenus) from West Virginia. 5 10 Male Percent testis length Female Percent testis width Juvenile 8 6 6 4 4 No. Individuals 2 Percent (%) testis dimensions 2 0 0 0 2 4 6 8 10 12 0 2 4 6 8 10 12 Month Month Figure 5. Monthly incidence of captures of 19 male, 19 female, Figure 6. Monthly distribution of testis size as a percentage and 13 juvenile Northern Brownsnakes (Storeria dekayi dekayi) of male snout-vent length (SVL) of 19 Northern Brownsnakes from West Virginia. (Storeria dekayi dekayi) from West Virginia. © The Center for North American Herpetology 20 35 8 30 y = 1.2401x - 10.278 R2 = 0.1991 6 25 p= 0.006 20 4 15 No. Young 10 Follicle diameter (mm) 2 5 0 0 0 2 4 6 8 10 12 15 16 18 20 22 24 26 28 30 Month SVL (cm) Figure 7. Monthly distribution of follicle size in 17 Northern Figure 8. Relationship between clutch size and female snout- Brownsnakes (Storeria dekayi dekayi) from West Virginia. vent length (SVL) of 17 Northern Brownsnakes (Storeria dekayi dekayi) from West Virginia. 20 Male 9 Female Male 16 *Juvenile 8 Female Juvenile 7 20 6 15 * * 5 * SVL (cm) SVL * * 4 * * * 10 * No. Individuals 3 * 2 5 1 0 0 0 2 4 6 8 10 12 0 2 4 6 8 10 12 Month Month Figure 9. Monthly distribution of snout-vent lengths (SVL) of 19 Figure 10. Monthly incidence of captures of 11 male, 17 male, 19 female, and 13 juvenile Northern Brownsnakes (Storeria female, and five juvenile Northern Red-bellied Snakes (Storeria dekayi dekayi) from West Virginia. occipitomaculata occipitomaculata) from West Virginia. 10 Percent testis length 6 Percent testis width 8 10 8 6 6 4 4 Follicle diameter (mm) Percent (%) testis dimensions 2 2 0 0 0 2 4 6 8 10 12 0 2 4 6 8 10 12 Month Month Figure 11. Monthly distribution of testis size as a percentage of Figure 12. Monthly distribution of largest follicle size in nine male snout-vent length (SVL) of 11 Northern Red-bellied Snakes Northern Red-bellied Snakes (Storeria occipitomaculata (Storeria occipitomaculata occipitomaculata) from West Virginia. occipitomaculata) from West Virginia. Journal of North American Herpetology 2017(1) 21 Table 1. Adult body sizes in cm SVL (mean followed by range) and clutch sizes of the Common Wormsnake (Carphophis amoenus) from various locations in the United States. Taxon/Location Male Female Clutch size Nesting season Carphophis a. amoenus Connecticut (Lamson, 1935; Klemens, 1993) ..... 22.2 (15.8–25.5) ............... 28.0, 29.3 .......... 4.3 (4–5) ...........May–July Pennsylvania (Hulse et al., 2001) ....................... 19.8 (14.6–21.9) ...... 24.1 (21.6–27.6) .................5 (5) ....................... Maryland (Clark, 1970) ................................................................................................. 3.5 (2–5) ....................... West Virginia (this study) .................................. 18.8 (14.1–23.2) ...... 20.9 (14.9–28.0) .......... 2.8 (1–5) ................. June Virginia- northern (Mitchell, 1994) .............................................................................................. 4.0 (2–6) ....................... Virginia (Mitchell, 1994) ............................ 18.2 (14.1–23.7) ...... 20.2 (16.6–27.5) .......... 3.6 (2–6) ................. June Virginia (Table 20 in Clark, 1970) ................................................................................. 2.9 (1–4) ....................... Maryland (Table 20 in Clark, 1970) ................................................................................. 3.5 (2–5) ....................... North Carolina (Palmer and Braswell, 1995) ............................................................................ 3.4 (1–6) .......July–August North Carolina- western piedmont (Brown, 1992)................................................................................................ 3.0 (2–5) .......... June–July North Carolina- western piedmont (Wilson and Dorcas, 2004) ............. 21.1 (17.0–24.4) ...... 23.0 (19.0–26.2) .......... 4.0 (3–5) ....................... Alabama (Mount, 1975) .......................................................................................