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Overwintering in the Nest by Hatchling Map Turtles (Graptemys Geographica)

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Overwintering in the Nest by Hatchling Map Turtles (Graptemys Geographica) 1211 Overwintering in the nest by hatchling map turtles (Graptemys geographica) Roy D. Nagle, Clayton L. Lutz, and Andrew L. Pyle Abstract: We monitored 75 natural nests of Graptemys geographica (Lesueur, 1817) in central Pennsylvania to deter- mine the tactics and patterns of hatchling emergence. Following incubation, all hatchlings in 95% of nests delayed emergence from their natal nest cavities throughout autumn and winter until the following spring. Nests were con- structed in a variety of substrates ranging from loose sand or coal to hard-packed clay mixed with gravel. Time from egg deposition to natural hatchling emergence averaged 333 days. During winter, hatchlings tolerated subzero tempera- tures as low as –8 °C, which are lethal to hatchlings of some sympatric species. Emergence occurred from 10 April to 25 May, and most hatchlings were found during morning following rain. There was an interactive effect of minimum daily air temperature and rainfall level 1 day prior to emergence on the number of hatchlings emerging each day. One half of all hatchlings found dead were contained in nests in which autumn emergence had apparently occurred. We speculate that autumn emergence by hatchlings of some turtle species may be an adaptive response to nest conditions likely to provide poor environments for successful overwintering. Résumé : Nous avons suivi 75 nids naturels de Graptemys geographica (Lesueur, 1817) dans le centre de la Pennsyl- vanie afin de déterminer les stratégies et les patterns d’émergence des nouveau-nés. Après l’incubation, dans 95 % des nids, tous les nouveau-nés retardent leur émergence de la cavité du nid natal pendant l’automne et l’hiver jusqu’au printemps suivant. Les nids sont construits dans une variété de substrats, allant de sable ou de charbon non consolidé à de la glaise compacte mêlée de gravier. Le temps entre la ponte des oeufs et l’émergence naturelle des petits est en moyenne de 333 jours. En hiver, les petits supportent le gel jusqu’à –8 °C, des températures létales pour les nouveau- nés de quelques autres espèces sympatriques. L’émergence a eu lieu du 10 avril au 25 mai et la plupart des petits ont été retrouvés le matin après une pluie. Ilyauneffetinteractif entre la température minimale journalière de l’air et la quantité de précipitations la veille de l’émergence, d’une part, et le nombre de petits qui émergent chaque jour, d’autre part. La moitié de tous les nouveau-nés trouvés morts proviennent de nids où il s’est apparemment produit une émer- gence d’automne. Nous posons l’hypothèse que l’émergence à l’automne des nouveau-nés chez certaines espèces de tortues peut être une réaction adaptative à des conditions dans le nid qui vont vraisemblablement fournir un mauvais environnement pour survivre à l’hiver. [Traduit par la Rédaction] Nagle et al. 1218 Introduction of some species hatch during late summer or autumn yet re- main in nests throughout winter, delaying emergence and Turtles exhibit remarkable abilities to cope with adverse dispersal until the following spring. As a result, some hatch- conditions and commonly do so through prolonged periods lings spend 6–8 months in the nest post hatch, whereas their of inactivity. At early developmental stages, for example, incubation period is approximately 2 months (Tinkle et al. embryos of some species enter extended diapause in 1981; Christens and Bider 1987; Gibbons and Nelson 1978; response to chilling (Ewert 1979, 1985), whereas in other Lindeman 1991; Jackson 1994; Costanzo et al. 1995; DePari species embryos complete development but wait to hatch 1996; Tucker 1999). By utilizing the “proven sanctuary” af- coincidently with favorable conditions, such as the onset of forded by the nest, such a strategy may reduce uncertainties the wet season in Australia’s wet-dry tropics (Doody et al. associated with the fluctuating or declining resource levels 2001). In yet other species, offspring complete development of late autumn (Gibbons and Nelson 1978) or the vulnerabil- and hatch from their eggs but delay emergence and migration ity of hatchlings to predators during a period of low growth from terrestrial nests. potential (Wilbur 1975). Delayed emergence is the phenomenon in which hatchling Among North American turtles, hatchlings of five species turtles remain in or near their natal nest cavities for extended of the family Emydidae and two species of the family Kino- periods following hatching. At northern latitudes, neonates sternidae are reported to delay emergence from nests Received 16 February 2004. Accepted 12 July 2004. Published on the NRC Research Press Web site at http://cjz.nrc.ca on 7 October 2004. R.D. Nagle,1 C.L. Lutz,2 and A.L. Pyle.3 Environmental Science and Studies, Juniata College, Huntingdon, PA 16652, USA. 1Corresponding author (e-mail: [email protected]). 2Present address: Nongame Branch, Arizona Game and Fish Department, Phoenix, AZ 85023, USA. 3Present address: Programa Restauración de Tortugas Marinas, 1203-1100, Tibás, San José, Costa Rica. Can. J. Zool. 82: 1211–1218 (2004) doi: 10.1139/Z04-096 © 2004 NRC Canada 1212 Can. J. Zool. Vol. 82, 2004 Table 1. North American turtles that typically delay nest emergence, and locations where they were sampled. Family Species Location Reference(s) Emydidae Chrysemys picta (Schneider, 1783) Idaho Lindeman 1991 Michigan Wilbur 1975; Tinkle et al. 1981 Minnesota Woolverton 1963 Nebraska Costanzo et al. 1995 New Jersey DePari 1996 Quebec Christens and Bider 1987 Deirochelys reticularia (Latr., 1801) South Carolina Congdon et al. 1983; Buhlmann and Coffman 2001 Graptemys geographica (Lesueur, 1817) Indiana Baker et al. 2003 Terrapene ornata (Agassiz, 1857) Nebraska Costanzo et al. 1995 Trachemys scripta (Schoepff, 1792) Florida Jackson 1994 Illinois Tucker 1999 South Carolina Gibbons and Nelson 1978 Kinosternidae Kinosternon flavescens (Agassiz, 1857) Iowa Christiansen and Gallaway 1984 Nebraska Iverson 1991 K. subrubrum (Lacépede, 1788) South Carolina Gibbons and Nelson 1978 (Table 1). One factor common among such species is the abil- Materials and methods ity of females to produce multiple clutches during a given re- productive season. Owing to the lengthy egg-laying season, We located turtle nests from May 2000 through July 2002 hatching occurs over a broad temporal range and may include by searching terrestrial areas for nesting females along a periods highly unfavorable for growth and survival of hatch- portion of the Juniata River in Huntingdon County, Pennsyl- ling turtles (Gibbons and Nelson 1978). By delaying nest vania. The Juniata River is the second largest tributary of the emergence until spring, hatchlings are more likely to enter en- Susquehanna River and local discharge rates vary annually vironments favorable to the acquisition and allocation of re- from >10 000 ft3/s (1 ft3/s = 28.316 85 dm3/s) during April sources. Delayed emergence has also been suggested as a to <1000 ft3/s during August and September (Pluto and Bellis proximate response to impenetrable nest soil conditions 1986; Durlin and Schaffstall 2001). The area is near the (Cagle 1944; Hartweg 1944; DePari 1996), although the pro- western range limit of G. geographica in the central part of pensities of some turtle species to delay emergence across the state, where the larger rivers of Pennsylvania’s ridge and broad latitudinal and longitudinal ranges and among a variety valley region give rise to the Allegheny Front. Sympatric of substrates appear to counter a general interpretation of the species at the site include Chelydra serpentina (L., 1758), “nest integrity” hypothesis (Table 1). Clemmys insculpta (LeConte, 1830), and Sternotherus A more complete understanding of the adaptive signifi- odoratus (Latr., 1801). cance of delayed emergence by hatchling turtles requires Observed G. geographica nests were protected from pred- data on the relative costs and benefits, as well as the pres- ators using cylindrical 2.5 cm×5cmwire-mesh cages mea- ence and magnitude of geographic variation within and suring ~75 cm diameter × 25 cm high. We rotated cages into among species whose hatchlings overwinter in the nest. Fur- the soil to bury their edges and hold them in place. Thermis- thermore, documenting temporal aspects of delayed emer- tor probes from calibrated HOBO® temperature data loggers gence can elucidate important aspects of natural history and (Onset Computer, Pocasset, Massachusetts) were placed in a provide valuable information for wildlife managers and sample of nests each year to monitor thermal environments land-use planners. associated with incubation, overwintering, and hatchling Map turtles (Graptemys geographica) are a highly aquatic emergence. Protected nests were monitored at least twice emydid turtle endemic to the rivers and lakes of eastern and weekly throughout July and August for external signs of re- central North America (Ernst et al. 1994). They are noted for duced viability (erosion, subterranean predation, etc.) and frequent communal basking (Newman 1906; Flaherty and for hatchling emergence. Samples of nests were dug and in- Bider 1984; Pluto and Bellis 1986), extreme sexual size di- spected during September, October, and the following April morphism (Gordon and MacCulloch 1980; Gibbons and and May to determine hatching success and hatchling emer- Lovich 1990), and a molluscivorous diet (Newman 1906; gence strategies. Vogt 1980). In Pennsylvania, isolated populations of G. geo- During the first year of our study, nests were monitored graphica occur in the Delaware and Susquehanna river sys- daily during spring to determine patterns of hatchling emer- tems (Ernst et al. 1994). Only a few studies have described gence. In late March 2001, we buried rings of 15 cm high selected aspects of reproduction in G. geographica (Newman aluminum flashing to a depth of about 5 cm inside the pro- 1906; Gordon and McCullough 1980; Vogt 1980) and only tective wire cages to restrain hatchlings as they emerged one other study has examined aspects of early life stages from nests.
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