2008 SOUTHEASTERN NATURALIST 7(3):493–504
An Annotated Checklist and Preliminary Designation of
Drainage Distributions of the Crayfishes of Alabama
Guenter A. Schuster1,*, Christopher A. Taylor2, and John Johansen3
Abstract - As a first step toward elucidating the current status of Alabama’s crayfish
fauna, museums known to have significant crayfish collections were queried
for their holdings from that state. A total of 4649 records of Alabama crayfishes
were obtained from seven museums. Three-hundred thirty of the records did not
have suitable information for geo-referencing. The largest holdings were in the
National Museum of Natural History (2544 records). Specimen identifications were
verified, and all records were geo-referenced. Geographic distribution strongly
favored upland drainages in the northern and central portions of the state. The Tennessee
River drainage was the most collected drainage (1018 records, 23.6% of
records). A total of 85 species of crayfishes are reported for the state of Alabama;
only Orconectes virilis (Virile Crayfish) is deemed to be non-native. Even with the
extensive collection of crayfish records in museums, there is a need for crayfish
inventory work in Alabama. A total of 3107 records (76.3%) were collected prior
to 1987, and 1379 (33.8%) were collected prior to 1970. In addition, there is a paucity
of records from the coastal drainages of SE Alabama. There is also an underrepresentation
of burrowing crayfishes, especially those classified as either primary
or secondary burrowers. Only 212 (4.9%) of all records are of burrowing species.
Lastly, a limited amount of fieldwork in Alabama has documented the presence of a
species previously unreported for the state.
Introduction
As pointed out by Schuster and Taylor (2004) and others (Boschung and
Mayden 2004, Crandall 1997, Lydeard and Mayden 1995), the aquatic fauna
of Alabama is extremely diverse, and the total number of taxa may exceed
that of any other state or province in North America. Recent distributional
and ecological information has been compiled for major aquatic groups such
as fishes (Boschung and Mayden 2004, Mettee et al. 1996), caddisfl ies (Harris
et al. 1991), and unionid mussels (Williams et al., in press). However,
such information for crayfishes is lacking.
Alabama has a relatively mild, stable climate, was spared from recent
glacial advances, and has a diverse geology and topology ranging from
high-gradient Appalachian streams to coastal fl oodplains, factors that have
undoubtedly contributed to the State’s high biodiversity.
1Department of Biological Sciences, Eastern Kentucky University, 521 Lancaster Avenue,
Richmond, KY 40475. 2Division of Biodiversity and Ecological Entomology,
Illinois Natural History Survey, 1816 South Oak, Champaign, IL 61820. 3Tulane University
Museum of Natural History, 3705 Main Street, Building A-3, New Orleans,
LA 70075. *Corresponding author - Guenter.Schuster@eku.edu.
494 Southeastern Naturalist Vol.7, No. 3
Ortmann (1905, 1931) and Hobbs (1969) speculated that the Cumberland
Plateau is the center of radiation for the crayfish genera Cambarus and
Orconectes, both of which are major components (combined total of >50%)
of the Alabama crayfish fauna. Whether or not that speculation is correct,
the Cumberland Plateau is the second largest physiographic province in the
state (Boschung and Mayden 2004), and its hydrology (containing both the
Tennessee and Black Warrior river drainages) and highly dissected topology
strongly infl uence aquatic faunal components for most of the northern section
of the state.
Certainly another important aspect of the evolution of aquatic organisms
in Alabama is the diversity of river systems in the state. Three major
basins drain Alabama: the Ohio River, Mobile Bay, and the Apalachicola
River. In addition, there are four relatively isolated Coastal Plain drainages:
Pascagoula River, Perdido Bay, Pensacola Bay, and Choctawhatchee Bay
(Boschung and Mayden 2004). The long-term isolation (until the recent
connection of the Tombigbee and Tennessee river systems) of the major basins
and the coastal drainages, undoubtedly resulted in divergent evolution
among populations leading to speciation.
To address the deficiency in baseline distributional information for Alabama
crayfishes, a study was undertaken with three main objectives. The
first was to determine the extent of Alabama crayfish holdings in various
museums in the United States. Secondly, museums with significant holdings
were visited and, where appropriate, specimens were identified or had their
identifications confirmed or corrected. Invalid taxonomic names were also
corrected. Thirdly, all museum specimens with adequate locality information
were geo-referenced and compiled into a single database. In addition to
museum work, a limited amount of field collecting was conducted between
2005 and 2007 using standard field techniques.
Methods
Museums known to have significant crayfish collections were queried
for their holdings of Alabama crayfishes. Electronic records from those
museums containing more than 50 records were then integrated into a single
Microsoft Excel database. A record is defined as a collection of one species
of crayfish from a single location. With two exceptions, museums with
more than 50 Alabama records that had not been previously examined by
the authors were then visted. A concentrated effort was made to identify
previously unidentified Alabama specimens and confirm identifications of
specimens that did not fit known distributions. Once confirmed, all records
in the Microsoft Excel database were geo-referenced using GEOLocate
(V 2.13, Tulane University), and entered into a Microsoft Access database
in a decimal degree format. All geo-referenced records were then visually
confirmed on county road or topographic maps or electronic maps in the
software package TopoUSA (DeLorme).
2008 G.A. Schuster, C.A. Taylor, and J. Johansen 495
Institutional acronyms are as follows: Auburn University of Natural History
(Auburn); University of Alabama Museum of Natural History (UA);
Eastern Kentucky University Branley A. Branson Museum of Zoology
(EKU); Museum of Comparative Zoology at Harvard University (MCZ);
Illinois Natural History Survey Crustacean Collection (INHS); National Museum
of Natural History, Smithsonian Institution (USNM); North Carolina
State Museum of Natural History (NCSM); Tulane University Museum of
Natural History (TU).
Figure 1. Distribution of known crayfish records for Alabama (n = 4319).
496 Southeastern Naturalist Vol.7, No. 3
Results
A total of 4649 records of Alabama crayfishes were obtained from seven
museums and recent fieldwork (Fig. 1). Of these, 330 (7.1%) records did not
have adequate information for geo-referencing. In addition, 249 records did
not contain date of collection. The largest holdings of records were found at
USNM with 2544 records (54.8%), TU with 804 (17.3%), and UA with 487
(10.5%). The temporal distribution of Alabama crayfish collections was very
uneven (Fig. 2). Sixty-three percent of all records were collected during the
1960s and 1970s, with 17.1% of collections made during the 1990s. Only
9.0% of collections were made prior to 1960.
The geographical distribution of Alabama records strongly favored
upland drainages in the northern and central portions of the state. The
Tennessee River drainage was the most represented drainage having 1018 records
(23.6%) (Fig. 3). The Tombigbee (428 records), Black Warrior (539),
Cahaba (217), Coosa (462), Tallapoosa (480), and Alabama (308) drainages
combined accounted for 56.3% of all records. The coastal drainages of
southeastern Alabama, including the Choctawhatchee, Conecuh, Escatawpa,
Perdido, and Yellow, accounted for only 9.3% of all records (Fig. 4).
In addition to museum work, recent fieldwork in Alabama has resulted in
the addition of one new species. Orconectes durelli (Saddle Crayfish) was
collected from Second Creek, Lauderdale County, on October 22, 2005.
The compilation and examination of museum records and fieldwork revealed
85 described crayfish taxa known to occur in Alabama (Appendix 1).
Of these, only O. virilis is deemed to be non-native. Taxa are presented in
alphabetical order by subfamily, genus and species with common name,
known range by drainage basin, and habitat category for each species.
Figure 2. Number of geo-referenced Alabama Crayfish records by decade of collections
(n = 4070). Number does not include records with incomplete locality data or
missing date of collection.
2008 G.A. Schuster, C.A. Taylor, and J. Johansen 497
Figure 3. Number of geo-referenced Alabama Crayfish records by major river drainage
(AL = Alabama, BW = Black Warrior, CH = Chattahoochee, CO = Coosa, MO =
Mobile, TA = Tallapoosa, TN = Tennessee, and TO = Tombigbee) (n = 4319).
Figure 4. Number of geo-referenced Alabama Crayfish records by minor river drainage
(BW = Blackwater, CA = Cahaba, CP = Chipola, CT = Choctawhatchee, CO =
Conecuh, ES = Escatawpa, PE = Perdido, and YE = Yellow) (n = 4319).
498 Southeastern Naturalist Vol.7, No. 3
All crayfish living in Alabama belong to the family Cambaridae, which
contains two subfamilies, Cambarellinae and Cambarinae. Representatives
of both subfamilies reside in Alabama. Cambarellinae has a single genus,
Cambarellus, of which three species are known from Alabama. The remaining
82 species belong to Cambarinae. Six genera of Cambarinae are present
in Alabama: Cambarus, Fallicambarus, Faxonella, Hobbseus, Orconectes,
and Procambarus. Ten subgenera and 27 species of Cambarus occur in
Alabama, making it one of the largest crayfish genera in the state. All five
species of Fallicambarus in Alabama belong to the subgenus Creaserinus.
Faxonella is represented by a single species, as is the genus Hobbseus. Orconectes,
also one of the largest crayfish genera in Alabama, is represented
in the state by seven subgenera and 19 species. Procambarus is the most
diverse genus in Alabama, with six subgenera and 29 species found within
its boundaries.
Discussion
History of astacology in Alabama
Until 1870, the crayfishes of Alabama went almost unnoticed. Hagen
(1870) provided the first checklist of crayfishes from Alabama and remarked
that “nearly the whole state remains unexplored.” His list included 5 species:
Cambarus advena (= Procambarus hagenianus hagenianus [Southeastern
Prairie Crayfish]), C. immunis (= Orconectes immunis (Hagen) [Calico
Crayfish]), C. acutus (= P. acutus [White River Crayfish]), C. lecontei (= P.
lecontei [Mobile Crayfish]), and C. versutus (= P. versutus [Sly Crayfish]).
Two of these species, P. lecontei and P. versutus, were the first Alabama
crayfishes described from Alabama specimens. The specimens of both were
collected by Louis Agassiz on a trip to Mobile in the early 1850s.
A century after Hagen’s work, Bouchard (1976) provided the first attempt
at a comprehensive checklist of Alabama crayfishes. He listed 58 species
for the state and estimated that approximately 75 will be known when the
fauna has been completely surveyed. In addition, he was the first to attempt
to determine the conservation status for the crayfishes of Alabama. He listed
14 species in need of protection. One was listed as threatened, while the
remaining 13 were listed as species of special concern.
Hobbs’ (1989) illustrated checklist of American crayfishes listed
72 species and subspecies from Alabama, with two species being listed
as questionable. One of these, Fallicambarus hedgpethi, has been synonymyzed
with Fallicambarus fodiens (Digger Crayfish), while the other,
Procambarus bivittatus (Ribbon Crayfish), is now considered to be a
member of the state’s fauna.
In a conservation review of North American crayfish, Taylor et al. (1996)
listed 78 taxa as occurring in Alabama. Crandall (1997) erroneously cited
Taylor et al. (1996) in reporting 76 taxa as occurring in Alabama. Most recently,
Taylor et al. (2007) presented an update on the conservation of North
American crayfish. They listed 86 taxa, including the subspecies Cambarus
2008 G.A. Schuster, C.A. Taylor, and J. Johansen 499
bartonii bartonii (Farbricius), in their list for Alabama, which we exclude
here. The marked increase in the number of taxa reported from Alabama
between 1976 and 2007 was largely due to taxonomic revisionary work and
new species descriptions.
Current status
The records reported herein do not reflect an up-to-date assessment
of the Alabama crayfish fauna. With a total of 3107 (76.3.0%) records
collected prior to 1987 and 1379 (33.8%) predating 1970, the ability to
determine current population trends and distributions is severely hampered.
More alarming is the under-representation of burrowing crayfishes
(Hobbs 1981, 1989). Of the 4649 records, only 212 (4.9%) are of species
that spend a significant portion of their lives in subterranean burrows.
This under-sampling prevents an assessment of both historical and current
population levels and distributions.
Our preliminary examinations reveal that at least one manuscript name
may be applicable to a previously described species (Orconectes jonesi
[Sucarnoochie River Crayfish]); however, others may represent undescribed
species. There remains a great need for additional taxonomic evaluation of
museum collections and field collecting. One noteworthy example is the
presence of Procambarus zonangulus (Southern White River Crayfish) in
Alabama. Procambarus zonangulus is a member of the P. acutus species
group and has been considered by some to be the form present in most of the
Gulf Coastal Plain, including Alabama (Huner 2002). We did not find specimens
assignable to P. zonangulus during our visits to museums; however,
we follow Huner (2002) in including it in our species list. Finally, in addition
to specimen-based taxonomic uncertainties, recent molecular work has suggested
the presence of at least two undescribed taxa in the genus Cambarus
from karst regions of northern Alabama (Buhay et al. 2007). Future molecular
analyses will surely reveal the presence of unique species in other regions
of the state.
Future efforts in Alabama should include: 1) taxonomic revisions of numerous
crayfish groups and descriptions of new taxa; 2) additional fieldwork
in under-sampled drainages, especially in coastal drainages of southeastern
Alabama; 3) additional fieldwork in regions where the last known collections
predate 1987; and 4) statewide collections of primary and secondary
burrowing species.
Acknowledgments
We gratefully acknowledge Jim McHugh and Steve Rider of the Alabama Division
of Wildlife and Freshwater Fisheries and the State Wildlife Grants program
for their financial support of crayfish work in Alabama. We also acknowledge Hank
Bark and Nelson Rios of Tulane University Museum of Natural History for their
assistance with the curation of the TU Collection and the administration of the
grant funds. We also thank Karen Reed and Rafael Lemaitre (UNSM), Phil Harris
and Bernard Kuhadja (UA), and Michael Gangloff (Auburn) for access to the
500 Southeastern Naturalist Vol.7, No. 3
respective collections under their care and assistance during our visits. Thanks also
to John Cooper (NCSM) and Ardis Johnston (MCZ) for providing collection records.
We would also like to thank Courtney Graydon, Emily Hartfield, and Dan Jones for
their help in the field. Finally, we thank the following for providing specimens: Paul
Freeman (The Nature Conservancy), Jim Godwin (Alabama Heritage Program),
Courtney Graydon (Alabama State Lands), Dennis DeVries and Emily Hartfield
(Auburn), Stuart McGregor and Pat O’Neil (Geologic Survey of Alabama), and Steve
Rider (Alabama Department of Wildlife and Freshwater Fisheries).
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502 Southeastern Naturalist Vol.7, No. 3
Appendix 1. List of Alabama crayfish species alphabetically by subfamily, Genus
and species with common names, habitat (C = caves, Le = lentic, Lo = lotic, PB =
primary burrower, and SB = secondary burrower), and known range by drainage
basin (AL = Alabama; BW = Black Warrior; CA = Cahaba; CH = Chattahoochee;
CO = Coosa; CT = Choctawhatchee; ES = Escambia: Blackwater, Conecuh, and Yellow;
MO = Mobile Bay: Mobile, Tensaw, and direct tributaries; PA = Pascagoula:
Escatawpa; PE = Perdido; TA = Tallapoosa; TN = Tennessee; and TO = Tombigbee);
UN=Uncertain range or species under taxonomic revision.
Species Common name Habitats Drainages
Cambarellus diminutus Hobbs Least Crayfish Le MO, PA
C. lesliei Fitzpatrick & Laning Angular Dwarf Crawfish Le AL ,MO, TO
C. shufeldtii (Faxon) Cajun Dwarf Crayfish Le MO
Cambarus acanthura Hobbs Thornytail Crayfish PB AL, BW, CA,
CH, CO, ES,
PA, PE, TO,
TN
C. bartonii cavatus Hay Appalachian Brook Crayfish Lo CH,CO
C. coosae Hobbs Coosa Crayfish Lo CA,CO
C. cracens Bouchard & Hobbs Slenderclaw Crayfish Lo TN
C. diogenes Girard Devil Crayfish PB,SB AL, BW, CO,
CT, ES, MO,
TA, TN, TO
C. distans Rhoades Boxclaw Crayfish Lo TN
C. englishi Hobbs & Hall Tallapoosa Crayfish Lo TA
C. girardianus Faxon Tanback Crayfish Lo CA, CO, TN
C. graysoni Faxon Twospot Crayfish Lo TN
C. halli Hobbs Slackwater Crayfish Lo TA
C. hamulatus (Cope) Prickly Cave Crayfish C BW, TN
C. howardi Hobbs & Hall Chattahoochee Crayfish Lo CH
C. jonesi Hobbs & Barr Alabama Cave Crayfish C TN
C. latimanus (Le Conte) Variable Crayfish Lo AL, BW, CA,
CH, CO, CT,
ES, TA, TN,
TO
C. longirostris Faxon Longnose Crayfish Lo CO, TN
C. ludovicianus Faxon Painted Devil Crayfish PB MO
C. manningi Hobbs Greensaddle Crayfish Lo CO
C. miltus Fitzpatrick Rusty Grave Digger PB MO, PE
C. obstipus Hall Sloped Crayfish Lo BW
C. parvoculus Hobbs & Shoup Mountain Midget Crayfish Lo TN
C. polychromatus Thoma et al. Paintedh& Mudbug PB, SB CT
C. rusticiformis Rhoades Depression Crayfish Lo TN
C. scotti Hobbs Chattooga River Crayfish Lo CO
C. striatus Hay Ambiguous Crayfish Lo, PB, SB AL, BW, CA,
CH, CO, CT,
ES, TA, TO,
TN
C. tenebrosus Hay Cavespring Crayfish C, Lo TN
C. unestami Hobbs & Hall Blackbarred Crayfish Lo TN
C. veitchorum Cooper & Cooper White Spring Cave Crayfish C TN
Fallicambarus burrisi Fitzpatrick Burrowing Bog Crayfish PB MO, PA
F. byersi (Hobbs) Lavender Burrowing Crayfish PB ES, PA, PE
F. danielae Hobbs Speckled Burrowing Crayfish PB MO, PA
2008 G.A. Schuster, C.A. Taylor, and J. Johansen 503
Species Common name Habitats Drainages
F. fodiens (Cottle) Digger Crayfish PB, SB AL, BW, CA,
CH, ES, MO,
PE, TA, TN,
TO
F. oryktes (Penn & Marlow) Flatwoods Digger PB MO
Faxonella clypeata (Hay) Ditch Fencing Crayfish Le AL, CH, CT,
ES, MO, PA,
TA, TO
Hobbseus prominens (Hobbs) Prominence Riverlet Crayfish Le AL, BW, CA,
TO
Orconectes alabamensis (Faxon) Alabama Crayfish Lo TN
O. australis australis (Rhoades) Southern Cave Crayfish C TN
O. chickasawae Cooper & Hobbs Chickasaw Crayfish Lo UN
O. compressus (Faxon) Slender Crayfish Lo TN
O. cooperi Cooper & Hobbs Flint River Crayfish Lo TN
O. durelli Bouchard & Bouchard Saddle Crayfish Lo TN
O. erichsonianus (Faxon) Reticulate Crayfish Lo BW, CA,
CO, TA, TN
O. forceps (Faxon) Surgeon Crayfish Lo TN
O. holti Cooper & Hobbs Bimaculate Crayfish Lo UN
O. jonesi Fitzpatrick Sucarnoochee River Crayfish Lo UN
O. lancifer (Hagen) Shrimp Crayfish Le AL
O. mirus (Ortmann) Wonderful Crayfish Lo TN
O. perfectus Walls Complete Crayfish Lo AL, BW, TA,
TO
O. placidus (Hagen) Bigclaw Crayfish Lo TN
O. putnami (Faxon) Phallic Crayfish Lo TN
O. sheltae Cooper & Cooper Shelta Cave Crayfish C TN
O. spinosus (Bundy) Coosa River Spiny Crayfish Lo BW, TN
O. validus (Faxon) Powerful Crayfish Lo BW, TN, TO
O. virilis Hagen Virile Crayfish Lo BW, CA,
CO, TA, TN
Procambarus acutissimus (Girard) Sharpnose Crayfish Le, Lo AL, BW, CA,
CO, CT, TA,
TN, TO
P. acutus (Girard) White River Crawfish Le, Lo AL, BW, CO,
CT, ES, MO,
PA, TN, TO
P. bivittatus Hobbs Ribbon Crayfish Le, Lo AL, CT, MO,
TO
P. capillatus Hobbs Capillaceous Crayfish Le, SB ES
P. clarkii (Girard) Red Swamp Crawfish Le, Lo AL, BW, CA,
CO, MO, PA,
TA, TN, TO
P. clemmeri Hobbs Cockscomb Crayfish Lo PA, TO
P. escambiensis Hobbs Escambia Crayfish Le, Lo, SB ES, PE
P. evermanni (Faxon) Panhandle Crayfish Lo AL, MO, PE
P. hagenianus hagenianus (Faxon) Southeastern Prairie Crayfish PB TO
P. hayi (Faxon) Straightedge Crayfish Le, Lo TO
P. hubbelli (Hobbs) Jackknife Crayfish Le, SB ES
P. hybus Hobbs & Walton Smoothnose Crayfish Le, SB BW, CA, TO
P. lagniappe Black Lagniappe Crayfish Lo TO
504 Southeastern Naturalist Vol.7, No. 3
Species Common name Habitats Drainages
Procambarus lecontei (Hagen) Mobile Crayfish Lo MO, PA, TO
P. lewisi Hobbs & Walton Spur Crayfish Le, Lo AL, CH, ES,
TA
P. lophotus Hobbs & Walton Mane Crayfish Le, Lo AL, BW, CO,
CT, ES, TA,
TN, TO
P. marthae Hobbs Crisscross Crayfish Le, Lo AL, BW, CA
P. okaloosae Hobbs Okaloosa Crayfish Le, Lo ES
P. paeninsulanus (Faxon) Peninsula Crayfish Le, Lo CH
P. pecki Hobbs Phantom Cave Crayfish C TN
P. penni Hobbs Pearl Blackwater Crayfish Lo AL, PA, TO
P. shermani Hobbs Gulf Crayfish Le, Lo AL, MO, PA,
TO
P. spiculifer (Le Conte) White Tubercled Crayfish Lo AL, BW, CA,
CH, CO, CT,
ES, MO, PA,
PE, TA, TO
P. suttkusi Hobbs Choctawhatchee Crayfish Lo CT
P. verrucosus Hobbs Grainy Crayfish Lo AL, CH, ES,
TA
P. versutus (Hagen) Sly Crayfish Lo AL, BW, CA,
CH, CO, CT,
ES, MO, PA,
PE, TA, TO
P. viaeviridis (Faxon) Vernal Crayfish Le, Lo BW, TN
P. vioscai paynei Fitzpatrick Payne’s Creek Crayfish Lo BW, TO
P. zonangulus Hobbs & Hobbs Southern White River Crawfish Le, Lo UN