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2018 Northeastern Naturalist Notes Vol. 25, No. 2
Z.J. Loughman, P. Woods, and D.A. Lieb
First Record of Creaserinus fodiens (Digger Crayfish) from
Pennsylvania
Zachary J. Loughman1,*, Peter Woods2, and David A. Lieb3
Abstract - Pennsylvania’s stream crayfishes have received considerable conservation attention and
survey efforts in the past decade. Burrowing crayfishes have begun to receive the same level of effort
as stream species, resulting in the discovery of Creaserinus fodiens (Digger Crayfish) in a state game
land in the Glaciated Plateau of Crawford County, PA. Herein, we describe the collection of this species
in Pennsylvania, and determine that C. fodiens is native to the state and is worthy of additional
survey efforts and associated conservation assessments.
There has been a surge in crayfish conservation efforts over the past 2 decades (Richman
et al. 2015, Taylor et al. 2007). Historically, most crayfish-survey efforts focused on
tertiary burrowing species—crayfishes that occur in permanent lotic and lentic waterways
(Hobbs 1981). Ecological studies and surveys of primary burrowing species—crayfishes
that inhabit a burrow for the bulk of their life history (Hobbs 1981)—were rare, due to
difficulties in collecting burrowers. Most recent regional- or watershed-survey efforts for
crayfishes have included these enigmatic and ecologically important species (Kilian et al.
2010, Loughman and Simon 2011, Loughman and Welsh 2013, Simmons and Fraley 2010).
These efforts have expanded the documented range of several burrowing crayfishes and
resulted in a better understanding of burrower ecology and conservation (Kilian et al. 2010,
Loughman and Simon 2011, Simons and Fraley 2010).
In recent years, more crayfish surveys have been conducted in Pennsylvania than any
other northeastern state (Lieb et al. 2008, 2011a, 2011b). The bulk of efforts have focused
on assessing the replacement of native species by invasive crayfish (Lieb et al. 2011a, b),
as well as the documentation of native species distributions and conservation status (Lieb
et al. 2008, 2011a, 2011b). All of the aforementioned efforts focused on tertiary burrowers;
primary burrowing crayfishes were largely ignored.
With the exception of a single recent publication by Loughman et al. (2017), few contemporary
data are available for Pennsylvania’s burrowing crayfish, though substantial
historical information does exist. The state’s burrowing crayfish fauna was first documented
by Arnold Ortmann, Curator of Crustacea and Mollusks at Carnegie Museum of Natural History
in Pittsburgh, PA, whose surveys in the late 1800s and early 1900s are the foundation for
all subsequent efforts. Ortmann (1905, 1906) collected 4 burrowing crayfish species from
Pennsylvania and determined that Cambarus dubius Faxon (Upland Burrowing Crayfish),
Cambarus monongalensis Ortmann (Blue Crawfish), and Cambarus thomai Jezerinac (Little
Brown Mudbug) were restricted to the Appalachian Plateau and Allegheny Mountains of
western Pennsylvania (Appalachian physiographic provinces), and Cambarus diogenes Girard
(Devil Crayfish) was restricted to the Coastal Plain of southeastern Pennsylvania.
Given the need for updated information on burrowing crayfish, we surveyed western
Pennsylvania’s burrowing crayfishes from 2014 to 2016 (Loughman et al. 2017). In the
1West Liberty University, Department of Natural Sciences and Mathematics, PO Box 295, West Liberty,
WV 26074. 2Western Pennsylvania Conservancy, 800 Waterfront Drive, Pittsburgh, PA 15222.
3Western Pennsylvania Conservancy and Pennsylvania Fish and Boat Commission, 595 East Rolling
Ridge Drive, Bellefonte, PA 16823. *Corresponding author - zloughman@westliberty.edu.
Manuscript Editor: David Yozzo
Notes of the Northeastern Naturalist, Issue 25/2, 2018
2018 Northeastern Naturalist Notes Vol. 25, No. 2
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Z.J. Loughman, P. Woods, and D.A. Lieb
spring of 2014, P. Woods received photographs documenting burrowing crayfish chimneys
in roadside ditches (RSD) on State Game Land (SGL) 101 in Crawford County, PA (Fig. 1).
This documentation is significant because (1) burrowing crayfishes had not previously
been found that far north in Pennsylvania, and (2) that portion of Crawford County is in
the Northwestern Glaciated Plateau, outside of Pennsylvania’s Appalachian physiographic
Figure 1. (A) Location (yellow dot) of the first documentation of Creaserinus fodiens in Pennsylvania,
(B) roadside ditch adjacent to Pennsylvania State Game Land 101 with C. fodiens burrows and associated
chimneys at site of capture, and (C) chimney in foreground of close-up view.
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2018 Northeastern Naturalist Notes Vol. 25, No. 2
Z.J. Loughman, P. Woods, and D.A. Lieb
provinces. Furthermore, none of the primary burrowing species that were known to occur
in Pennsylvania had previously been documented in this physiographic region in either
Pennsylvania, neighboring Ohio (Thoma and Jezerinac 2000), or New York (Crocker 1957).
On 14 August 2014, Z.J. Loughman investigated the SGL 101 burrows and collected a
single juvenile male Creaserinus fodiens (Cottle) (Digger Crayfish) via burrow excavation,
and procured an additional juvenile female by hand at 24:30 h from a portion of a RSD
with standing water at the same location. Burrows were plentiful in all RSD habitats in
the general vicinity where both animals were captured. Upland habitat consisted of mesophytic
forests dominated by Fagus grandifolia Ehrh. (American Beech), various Acer spp.
(maples), and Betula lenta L. (Black Birch).
The collection of C. fodiens from SGL 101 adds another species and genus to Pennsylvania’s
crayfish fauna. Given the extent of crayfish introductions globally, determination of
the status of C. fodiens (native or exotic) is critical in deciding if conservation or eradication
efforts are warranted for this species. We consider C. fodiens to be native to Pennsylvania.
Creaserinus fodiens is the dominant burrowing crayfish in neighboring Ashtabula County,
OH (Thoma and Jezerinac 2000), with records for the species extending back to the 1800s
and it has long been documented in ephemeral wetlands along Lake Erie (Crocker and Barr
1968), north of the Pennsylvania population in Ontario, Canada.
Furthermore, in the central and northern parts of its range, C. fodiens is an ephemeral
pool specialist, often associated with pools that occur within forested environments similar
to those that are plentiful in SGL 101(Crocker and Barr 1968, Loughman and Simon 2011,
Loughman et al. 2012, Thoma and Jezerinac 2000). Such environments are not typically
frequented by anglers; thus the pools are unlikely to be directly exposed to introduced crayfishes
via bait-bucket releases. We hypothesize that C. fodiens populations in Pennsylvania
immigrated from Midwestern populations in neighboring Ohio following retreating glaciers
at the end of the last glacial advance. Currently, the statewide distribution and conservation
standing of C. fodiens remains unknown; surveys are currently underway to determine the
true extent of its distribution in Pennsylvania.
Figure 2. Juvenile male Creaserinus fodiens collected at Pennsylvania State Game Land 101 on 14
August 2014.
2018 Northeastern Naturalist Notes Vol. 25, No. 2
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Z.J. Loughman, P. Woods, and D.A. Lieb
Acknowledgments. This work was only possible through funds provided by the Pennsylvania
Wild Resource Conservation Program and the Western Pennsylvania Conservancy. We acknowledge
2 anonymous reviewers whose comments on an earlier draft of this manuscript increased its scope
and quality.
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