N35
2019 Northeastern Naturalist Notes Vol. 26, No. 3
S.M. Hartzell
Observation of a Cambarus bartonii bartonii (Common Crayfish)
Overwintering in a Terrestrial Winter Microhabitat
Sean M. Hartzell*
Abstract - In cool climates, crayfish that are not primary burrowing species are typically thought to
retreat into deeper waters in winter and may seek refuge in small aquatic burrows or interstitial spaces
associated with aquatic substrates. Herein, I report a novel observation of a juvenile Cambarus bartonii
bartonii (Common Crayfish) during winter within eastern Pennsylvania occupying a terrestrial
microhabitat consisting of moist, terrestrial leaf litter associated with a small depression in an upland
forest. I found the crayfish in association with a small aggregation of Plethodon cinereus (Eastern
Red-backed Salamander). Requirements of a hibernaculum by the salamanders with sufficient moisture
and buffered from freezing may have also provided sufficient conditions for the crayfish to utilize
the depression as a winter refugium.
During winter in cooler climates, crayfish that occupy surface waters (i.e., tertiary
burrowers; Hobbs 1981) are typically thought to respond by moving into deeper waters
for overwintering (Crocker and Barr 1968, Ortmann 1906, Prins 1968). Crayfish
may remain active in deeper waters throughout the winter, although a number of studies
have indicated an association between reduced activity in crayfish and cooler water
temperatures during winter, such as studies on crayfish in the genera Faxonius (formerly
Orconectes; Crandall and De Grave 2017) and Pacificastus (e.g., Bovbjerg 1952; Bubb et
al. 2002, 2004; Flint 1977; Prins 1968). When inactive during winter, crayfish have been
found to retreat into interstitial spaces below substrate or burrow into streambanks (Prins
1968). Conversely, some crayfish species could become more active during winter. For
instance, Cambarus thomai (Jezerinac) (Little Brown Mudbug), a primary burrowing species,
may increase activity during winter, as it has been reported to occupy surface waters
only during winter months (Loughman and Simon 2011). This latter observation suggests
the winter biology of certain crayfish species is presently understudied, particularly for
members of the genus Cambarus.
Few observations appear to have been published regarding winter behavior of Cambarus
bartonii bartonii (Fabricus) (Common Crayfish), a medium-sized, tertiary burrowing
crayfish that typically occupies springs, small to medium-sized streams, and occasionally
lentic habitats in eastern North America (Hobbs 1989). This species may retreat to deeper
waters in winter and/or burrow into the streambank or seek refugium within interstitial
spaces. For instance, Hamr and Berrill (1985) collected Common Crayfish during winter
within a larger river, but not within a smaller stream, suggesting that individuals at the latter
site may have burrowed or migrated to deeper waters during winter. Common Crayfish
often occupy springs and seeps (Hobbs 1989, Ortmann 1906) and may be collected in
these habitats during winter (S.M. Hartzell, pers. observ.). However, no previous studies
have reported observations of this species occupying a terrestrial refugium during winter,
Herein, I describe a novel observation of a young Common Crayfish found during winter in
eastern Pennsylvania occupying a terrestrial microhabitat associated with an overwintering
aggregation of Plethodon cinereus (Green) (Eastern Red-backed Salamander).
*Department of Biological and Allied Health Sciences, Bloomsburg University of Pennsylvania,
Bloomsburg, PA 17815; seanhartzell77@gmail.com.
Manuscript Editor: David Yozzo
Notes of the Northeastern Naturalist, Issue 26/3, 2019
2019 Northeastern Naturalist Notes Vol. 26, No. 3
N36
S.M. Hartzell
On 13 December 2016 at 1200 h, I collected a small (~4 cm total length) Common Crayfish
underneath moist leaf litter within a small depression along an abandoned roadway ~2
km south of Mechanicsville, Montour County, PA (40°57'05.5"N, 76°34'40.7"W). The feature
that contained the crayfish consisted of a depression approximately 1.5 m in length and
1 m in width along a hillside slope within an upland mixed-deciduous forest. The depression
did not appear to be immediately associated with any perennial or ephemeral water source
such as a stream, spring, or ephemeral runoff channel; however, a small spring was located
~20 m from the site of the depression. The depression contained a dense concentration of
moist leaf litter, but was not inundated with water during the observation reported herein.
The crayfish was located ~8–10 cm beneath the surface of the leaf litter and associated
with a small aggregation of 3 Eastern Red-backed Salamanders. The crayfish was alive but
torpid. The temperature of the leaf litter was 4 °C, as measured with a mercury thermometer.
Following these observations, I returned the crayfish and salamanders to their initial
positions in the leaf pile and covered them with leaves. Although weather data from the
general locality of the observation site during the period leading up to the observation are
unavailable, data from a regional weather station located ~43 km NW of the site indicate
that the weather the proceeding week consisted of mean temperatures varying from -2.2 °C
to 4.4 °C, with temperatures typically ≤0 °C (Weather Underground 2019). No precipitation
had occurred for 10 d prior to the observation reported herein, with the exception of ~5 cm
snowfall the day prior (Weather Underground 2019).
To my knowledge, the observation reported herein appears to be unique and may suggest
that the crayfish could have been utilizing the moist leaf litter as a refugium to overwinter.
This possibility may be supported by the fact that several Eastern Red-backed Salamanders
were associated with the leaf litter containing the crayfish. Eastern Red-backed Salamanders
are not freeze-tolerant and must hibernate in refugia that do not freeze (Casper 2005).
Many terrestrial or semi-terrestrial salamanders overwinter underneath leaf litter (Bishop
1941), suggesting that the Eastern Red-backed Salamanders were utilizing the leaf litter
that contained the Common Crayfish as a hibernaculum. Eastern Red-backed Salamanders
are lungless and require moist conditions for cutaneous respiration (Casper 2005, Petranka
1998). These requirements by Eastern Red-backed Salamanders may suggest that the leaf
litter was sufficiently moist and buffered from freezing to allow the Common Crayfish to
utilize this microhabitat as a terrestrial refugium during winter. Given the cold temperatures
and lack of heavy precipitation in the proceeding days, the crayfish may have occupied the
depression for at least some time prior to the date of its discovery reported herein. Due to
the occurrence of the depression in upland habitat and its distance from any permanent
streams or ephemeral channels, it is unlikely that the crayfish’s presence at this location
was the result of being washed there during a severe precipitation and/or high water event.
Further studies could survey similar habitats during winter to elucidate the frequency that
Common Crayfish or other crayfish species may occupy terrestrial refugium during winter.
Acknowledgments. I thank 2 anonymous reviewers for their time and comments which facilitated
the improvement of this manuscript. Additionally, I think the Bloomsburg University College of
Science and Technology and Katherine Burke for support during the preparation of this manuscript.
Funding towards this publication was generously provided by the Bloomsburg University Office of
Graduate Studies and Sponsored Research.
Literature Cited
Bishop, S.C. 1941. The Salamanders of New York. New York State Museum Bulletin 324:1–365.
Bovbjerg, R.V. 1958. Comparative ecology and physiology of the crayfish Orconectes propinquus and
Cambarus fodiens. Physiological Zoology 25:34–56.
N37
2019 Northeastern Naturalist Notes Vol. 26, No. 3
S.M. Hartzell
Bubb, D.H., M.C. Lucas, and T.J. Thom. 2002. Winter movements and activity of Signal Crayfish,
Pacifastacus leniusculus, in an upland river, determined by radio telemetry. Hydrobiologia
483:111–1119.
Bubb, D.H., T. J. Thom, and M.C. Lucas. 2004. Movement and dispersal of the invasive Signal
Crayfish, Pacifastacus leniusculus, in upland rivers. Freshwater Biology 49:357–368.
Casper, G.S. 2005. Plethodon cinereus (Green, 1818) Eastern Red-Backed Salamander. Pp. 796–800,
In M. Lannoo (Ed.). Amphibian Declines: The Conservation Status of United States Species. University
of California Press, Oakland, CA. 1115 pp.
Crandall, K.A., and S. De Grave. 2017. An updated classification of the freshwater crayfishes (Decapoda:
Astacidea) of the world, with a complete species list. Journal of Crustacean Biology
37:615–653.
Crocker, D.W., and D.W. Barr. 1968. Handbook of the Crayfishes of Ontario. University of Toronto
Press, Toronto, ON, Canada. 158 pp.
Flint, R.W. 1977. Seasonal activity, migration, and distribution of the crayfish Pacifastacus leniusculus
in Lake Tahoe. American Midland Naturalist 97:280–292.
Hamr, P., and M. Berrill 1985. The life histories of north-temperate populations of the crayfish Cambarus
robustus and Cambarus bartoni. Canadian Journal of Zoology 63:2313–2322.
Hobbs, H.H., Jr. 1981. The crayfishes of Georgia. Smithsonian Contributions to Zoology 318:1–549.
Hobbs, H.H., Jr. 1989. An Illustrated Checklist of the American Crayfishes (Decapoda: Astacidae,
Cambaridae, and Parastacidae). Smithsonian Institution Press, Washington, DC. 236 pp.
Loughman, Z.J., and T.P. Simon. 2011. Zoogeography, taxonomy, and conservation of West Virginia’s
Ohio River floodplain crayfishes (Decapoda, Cambaridae). Zookeys 74:1–78.
Ortmann, A.E. 1906. The crawfishes of the state of Pennsylvania. Memoirs of the Carnegie Museum
2:343–523.
Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Books, Washington,
DC. 592 pp.
Prins, R. 1968. Comparative ecology of Orconectes rusticus rusticus and Cambarus tenebrosis in Doe
Run, Meade County, Kentucky. Internationale Revue der gesamten Hydrobiologie und Hydrographie
53:667–714.
Weather Underground. 2019. Data from Williamsport Regional Airport, Pennsylvania, December
2016. Available online at https://www.wunderground.com/history/daily/us/pa. Accessed 28
March 2019.