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Body Temperature Variations of the Louisiana Pine Snake (Pituophis Ruthvent) in a Longleaf Pine Ecosystem

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Body Temperature Variations of the Louisiana Pine Snake (Pituophis Ruthvent) in a Longleaf Pine Ecosystem Herpetological Natural History, 9(2), 2006, pages 117-1 26. 02006 by La Sierra University BODY TEMPERATURE VARIATIONS OF THE LOUISIANA PINE SNAKE (PITUOPHIS RUTHVENT) IN A LONGLEAF PINE ECOSYSTEM John G. Himesl and Laurence M. Hardy2 Museum of Life Sciences, Louisiana State University in Shreveport, One University Place, Shreveport, Louisiana 7 11 15-2399, USA D. Craig Rudolph and Shirley J. Burgdorfi Wildlife and Habitat Silviculture Laboratory, Southern Research Station, USDA Forest Service, Nacogdoches, Texas 75962, USA Abstract. The thermal ecology of the Louisiana pine snake, Pituophis ruthveni, was studied from 1993-97 in Louisiana and Texas. All snakes were implanted with temperature-sensitive radiotransmitters. Temperatures were recorded from snakes located above ground and underground and were compared between size and sex classes (juveniles, adult males, adult females). Associated air and substrate tempera- tures were also recorded. Collectively, body temperatures of snakes were lowest during winter (1 1.4"C), increased during spring (22.3"C), peaked during summer (27.7"C), and decreased during autumn (22.7"C). Seasonal body temperatures were most similar between juveniles and adult males, primarily because adult females had higher body temperatures during summer. During spring and autumn, snake, air, and substrate temperatures were generally 34°C higher above-ground than underground. Temperatures of snakes of all size and sex classes increased from morning through evening and most closely approximated the air and substrate temperatures at 1800 h and 1900 h, when snakes were located predominantly underground. Temperatures of juveniles and adult males showed no consistent relationship to air or substrate temperatures from 0700-1700 h. However, temperatures of adult females were significantly lower than air temperature from 1000-1600 h. Temperatures of juveniles and adult males were significantly higher above ground than underground from 1100-1400 h and 1100-1700 h, respectively, with no consistent relationship between hourly above-ground and underground temperatures of adult females. Overall, temperature patterns of i9 ruthveni and I? catenifer deserticola appear to be similar, except that T! c. deserticola maintains higher tem- peratures in the spring than does l? ruthveni. Key Words. Body temperature; Longleaf pine; Louisiana pine snake; Pituophis ruthveni; Thermal ecology. The body temperature of snakes has a profound snakes are physiologically capable of engaging) is effect on most, if not all, facets of their ecology. limited by the range of thermal conditions available Snakes are nearly exclusively ectothermic, deriving in the environment (Peterson et al. 1993). The ther- their body heat primarily from external (environ- mal environment itself is subject to daily and sea- mental) sources. Thus, body temperature in snakes sonal fluctuations and thus body temperature in (and thus the level and type of activity in which snakes may be expected to fluctuate accordingly. However, most snakes are very adept at behavioral 'Present address: Florida Fish and Wildlife Conservation thermoregulation- in an effort to offset (at least par- Commi~~i~n,Species Conservation Planning Section, tially) the potentially debilitating effects of subopti- 3911 2321, Panama City, Florida 32409, mal thermal conditions, resulting in a correspon- U.S.A. Email: [email protected] 2Please use for correspondence. ding change in the types of activities in which 3Present address: US Fish and Wildlife Service, 510 Snakes may engage (Heatwole and Taylor 1987). Desmond Drive, Suite 102, Lacey, Washington 98503, USA. Consequently, the timing of basic and essential Herpetological Natural History, Vol. 9(2), 2006 activities, such as those associated with foraging would suggest that gravid females spend more time and reproduction, is proximately determined by basking, thus gaining heat energy needed for the body temperature and ultimately determined by the physiological demands of vitellogenesis. On the thermal environment (Lillywhite 1987). other hand, this species is highly fossorial (Himes Peterson et al. (1993) posed five fundamental et al. 2005), which would seem to contradict the questions about thermal ecology and associated body higher thermal needs of egg development. temperature variation. These questions aim to assess Therefore, understanding more about the thermal (1) the range of possible body temperatures under relationship of the Louisiana pine snake to its envi- natural conditions; (2) the proximate factors that ronment might contribute to a better understanding determine which body temperatures a snake selects of the thermal ecology of species with extreme from that range of possibilities; (3) the variation of reproductive characteristics. body temperatures of individual snakes under natural Mean cloacal temperatures of other taxa of conditions over extended periods of time; (4) the Pituophis include 25.5"C in P. c. catenifer functional effects (i.e., developmental, physiologi- (Cunningham 1966), 28.0°C in f? c. sayi (Fitch cal, behavioral) of body-temperature variation; and 1956), and 28.6"C in f? c. deserticola (Diller and (5) the ecological consequences of body-temperature Wallace 1996). In all three studies, temperatures variation. Ideally, studies on the thermal ecology of were recorded from snakes only once and thus daily snakes should cover each of these five areas. and seasonal fluctuations in body temperature Our main objective was to assemble data to could not be determined. Daily fluctuations in body cover these five areas of Peterson et al. (1993) for temperature of single individuals of l? c. sayi (Dill the Louisiana pine snake, Pituophis ruthveni. This 1972) and l? c. deserticola (Parker and Brown species is a highly threatened endemic of the relict- 1980) varied from 2632°C and 19-35"C, respec- ual longleaf pine forests of western Louisiana and tively. However, snakes were monitored over a one- adjacent Texas (Jennings and Fritts 1983; Reichling to two-day period and thus seasonal fluctuations in 1988; Rudolph and Burgdorf 1997). The few body temperature could not be determined. remaining natural populations of I? ruthveni are Parker and Brown (1980) recorded 155 cloacal highly disjunct and occur at extremely low densi- temperatures from 19 c. deserticola that emerged ties. Until the 1990s, when R ruthveni was known from or retreated into a communal hibernaculum in the wild from only 100 specimens (Rudolph, during spring (n = 44) and autumn (n = 38), respec- unpubl. data) and the U.S. Forest Service initiated tively. Cloaca1 temperatures were significantly high- an intensive ecological project on this species, no er in spring (mean = 27.88"C) than in autumn (mean quantitative studies had been conducted on the nat- = 24.43"C). However, no temperatures were record- ural history of I! ruthveni. In fact, what little infor- ed from snakes during winter and temperature dif- mation is available on I! ruthveni (e-g., morpholog- ferences by size and sex class were not noted. ical variation, reproductive biology) was collected Parker and Brown (1980) also recorded 416 from snakes maintained in captivity (e.g., Reichling body temperatures from nine l? c. deserticola 1989), and nothing, to our knowledge, has been implanted with temperature-sensitive radiotransrnit- reported on the thermal ecology of I? ruthveni. ters. The mean body temperature of surface-active Pituophis ruthveni occurs in a unique habitat snakes from spring through autumn averaged that is distinctive from that of other species of 27.9"C. Mean inactive body temperatures of snakes Pituophis. In addition, clutch sizes of I! ruthveni (n underground and under rocks varied from 19.68"C = 4 eggs; no deviation from mean) are smaller than (autumn) to 27.0 1"C (summer) and approximately those of all other species of Pituophis, and its eggs 12°C (autumn) to 24OC (summer), respectively. (mean length = 11.94 + 0.88 [95% CI] cm, Body temperatures of snakes underground were 10.00-13.00 cm, n = 8; mean width = 3.56 + 0.33 below air and insolated substrate temperatures, sirni- cm, range 3.00-4.00 cm, n = 8) and hatchlings lar to shaded substrate temperatures, and significant- (mean total length = 54.4 + 2.5 cm, range 52.0-55.5 ly higher in summer compared to autumn. When cm, n = 4; mean mass = 106.7 -t 5.7 g, range snakes were under rocks, body temperatures were 104.0-108.0 g, n = 3) are the largest of all snakes in similar to air and shaded substrate temperatures, con- the United States (Reichling 1989, 1990). Thus, the sistently below insolated substrate temperatures, and energetics of growing the exceedingly large eggs significantly higher during cloudy days in summer Himes et a1.-Body Temperature of Pituophis ruthveni compared to spring. Again, no temperatures were MATERIALS AND METHODS recorded from snakes during winter and temperature differences by size and sex class were not noted. Study Areas Greenwald (1971) determined that the aerobic This study was conducted within the historic energy available for activity (metabolic scope) of I! longleaf pine (Pinus palustris) region of Bienville, catenifer afinis was maximum at a body tempera- Sabine, and Vernon Parishes in Louisiana, and in ture of 30°C. In 1974, Greenwald determined that Angelina, Jasper, Newton, and Sabine Counties in strike velocity and frequency of prey capture were Texas. Sandhills that are dissected by intermittent maximum at body temperatures of 27°C and 33"C, and small perennial streams characterize
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