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CRAYFISHES OF ALABAMA
Guenter A. Schuster, Christopher A. Taylor,
and Stuart W. McGregor

The University of Alabama Press / Tuscaloosa

CRAYFISHES OF
ALABAMA
 PUBLISHED WITH SUPPORT FROM

Alabama Wildlife Federation

Center for Biological Diversity

U.S. Fish and Wildlife Service’s Alabama Field Office

The University of Alabama Press

Tuscaloosa, Alabama 35487-0380 Tennessee Valley Authority
uapress.ua.edu

Copyright © 2022 by the University of Alabama Press WATERWAYS AL

KY
UC

All rights reserved.

LIA
KENT

NCE

 
Inquiries about reproducing material from this work
Pro

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should be addressed to the University of Alabama Press. in at

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R e st o ri n g K Y W

Typeface: Minion Pro / Myriad Pro The Kentucky Aquatic Resources Fund (KARF),
a cooperative partnership effort between
Cover image: Cambarus striatus from Bogue Chitto Creek, the U.S. Fish and Wildlife Service’s Kentucky
Perry County, Alabama; photo by Guenter A. Schuster Field Office and the Kentucky Waterways Alliance
Back cover image: Barbicambarus simmonsi from Shoal Creek,
Lauderdale County, Alabama; photo by Guenter A. Schuster
Cover design: Todd Lape / Lape Designs
All photos by Guenter A. Schuster, unless otherwise noted

Cataloging-in-Publication data is available

from the Library of Congress.
ISBN: 978-0-8173-2106-2
E-ISBN: 978-0-8173-9376-2 Geological Survey of Alabama
We dedicate this book to

Dr. Horton H. Hobbs Jr. (1914–1994)

While he studied multiple crustacean groups over his career,

Dr. Hobbs’s work with North American crayfishes has been
the most impactful. His professional career began at the Uni-
versity of Virginia at a time when few zoologists were study-
ing crayfishes, and his interest in taxonomy and systematics
allowed him to fill that void. He spent the last 20 years of
his career at the Smithsonian National Museum of Natural
History, where his contributions to science flourished. He
described 157 crayfish taxa from the United States, 38% of
the entire US fauna. In addition, he also described one fam-
ily and four genera of US crayfishes, as well as 104 ostracod,
eight shrimp, and six crab species. However, numbers tell only
half the story of Dr. Hobbs’s impact. His descriptions usually
contain detailed information on the life history, habitat, and
distribution of crayfishes, all critical for crayfish management,
conservation, and further ecological studies. Dr. Hobbs was
always willing to assist budding crayfish biologists. This, com-
bined with his publishing record, makes it easy to state that
every crayfish researcher over the past 50 years has benefited
from his life and his accomplishments.

and

Dr. Edward O. Wilson (1929–)

Dr. Wilson is a native-born son of Alabama who became

the world’s foremost ant expert. In his career he has pub-
lished many books and has twice won the Pulitzer Prize. He
is known, along with his collaborators, for introducing many
novel and important biological concepts such as communica-
tion in ants through pheromones, island biogeography, socio-
biology, character displacement, and biophilia. He has also
become the world’s leading spokesperson and promoter for
biodiversity and conservation efforts. Yet he has always kept
Alabama close to his heart. He is arguably the greatest natural-
ist in the modern era. He will always be remembered for how
he significantly changed our view of the living world.
County map of Alabama.
CONTENTS

ix Preface 133 10. Species Accounts

133 Genus Barbicambarus
xi Acknowledgments 138 Genus Cambarellus
151 Genus Cambarus
xv Abbreviations 248 Genus Creaserinus
263 Genus Faxonella
xvii List of Alabama Crayfish Taxa 268 Genus Faxonius
332 Genus Hobbseus
336 Genus Lacunicambarus
357 Genus Orconectes
1 1. Introduction 365 Genus Procambarus

8 2. Physiography 459 11. Species of Hypothetical Occurrence in Alabama

17 3. Inland Waters of Alabama

21 4. Crayfish Distributions and Habitat Associations 461 Glossary

43 5. Crayfish Biology and Ecology 465 Appendix A. Alabama Crayfish Species

Described after Book Submission
52 6. Crayfish Conservation
467 Appendix B. Crayfish Taxa by County
59 7. Crayfish Anatomy and Important Taxonomic
Characters 477 Literature Cited

81 8. Species Identification and Format of Accounts 489 Index

83 9. Key to the Crayfishes of Alabama

Procambarus clarkii;
Hagen, 1870.
PREFACE
In this book we provide distribution maps, photographs,
and information on the biology, ecology, and natural history
for each species. Furthermore, we discuss the need for con-
servation concern for each species. A taxonomic key is also
provided to assist in the identification of all known species.
This may also be useful for identifying species from common
drainages in the states abutting Alabama. We are certain that
with time and continued collecting effort, additional species,
especially cryptic new species, may be found in the state. We
hope that this book will draw attention to this very important
group of aquatic organisms and lead to additional research on
the crayfishes of Alabama.
ix
Procambarus
hagenianus;
Faxon, 1914.
ACKNOWLEDGMENTS
This project began in the spring of 2005 and has been ongoing
ever since. It has involved years of searching for funding, con-
ducting fieldwork, visiting natural history museums, spending
countless hours examining and photographing specimens,
making maps, maintaining the Alabama crayfish computer
database, and writing numerous manuscripts leading up to
this book. None of this could have been accomplished by the
three authors alone. Innumerable people helped us along the
way, and without them this project could never have come to
fruition. Below we list those who have been instrumental in
helping with various crucial aspects of this study. We hope
that we have recognized everyone who has contributed in
some way to this project, and we greatly apologize to anyone
we may have neglected.
We would like to give special thanks to Steve Rider and
Jim McHugh (retired), both from the Wildlife and Freshwater
Fisheries Division (WFFD) of the Alabama Department of
Conservation and Natural Resources (ADCNR), for suggest-
ing and supporting this project. It started with a phone call
and ended with a book!
FUNDING
This project was originally funded through the Alabama
WFFD with Section 6 funds from the Endangered Species
Act of 1973, as amended, with the help of Steve Rider and Jim
McHugh. This funding gave us the resources to visit museums
and begin our fieldwork. The result was an initial database for
Alabama crayfishes of over 4,000 records. This funding truly
was the impetus that got this project off the ground. Rider and
McHugh even came to the Tulane University Museum of Nat-
ural History to give us a hand for several days in identifying
and cataloging that collection.
Subsequent State Wildlife Grant (SWG) funding from
the WFFD, secured with the assistance of Steve Rider, Jim
McHugh, Traci Wood, Nick Nichols, and Fred Harders, per-
mitted us to execute additional field sampling statewide to
update the existing database with modern records. The Geo-
logical Survey of Alabama (GSA) provided additional finan-
cial support under the direction of Pat O’Neil and Nick Tew
that allowed continuation of field and museum work during
the latter half of this project, and their support is appreciated.
We would also like to thank Jeff Powell, US Fish and Wildlife
Service, Daphne, Alabama, for providing funding for three
status survey grants. These grants provided critical funding
for additional survey work on potentially imperiled species.
We would like to thank the following individuals and
organizations for financially supporting the publication of
this book: Tim Gothard, Alabama Wildlife Federation; Tierra
Curry, Center for Biological Diversity; Shannon O’Quinn
and Todd Amacker, Tennessee Valley Authority; Jeff Powell,
Ecological Services Field Office, US Fish and Wildlife Service,
Daphne, Alabama; Michael Floyd and Lee Andrews, Ecologi-
cal Services Field Office, US Fish and Wildlife Service, Frank-
fort, Kentucky; Ward Wilson, Kentucky Waterways Alliance
and the Kentucky Aquatic Resource Fund. We would also like
to thank Ronald Cicerello and Jonathan Miller for donating
personal funds to help with the publication costs of this book.
Their contributions and support are greatly appreciated.
MANUSCRIPT REVIEW
Reviewing manuscripts is a very important and often thank-
less job. We would like to thank Susan Adams, Mael Glon, Dan
Johnson, and Chris Skelton for reviewing and commenting
on various editions of the key. We also wish to greatly thank
Susan Adams and James Fetzner for reviewing the complete
manuscript of this book. All these reviews have made it a
much better product.
MUSEUM SUPPORT STAFF
Specimens housed in museums are critical for projects such
as this book. Without vouchered museum specimens, our
data set would be considerably smaller. To that end, we want
to thank all the museum curators and managers who made
our visits so worthwhile and enjoyable, and we would also like
to extend our appreciation to those who provided specimen
loans during the course of this study. These include staff of the
following museums: Smithsonian Institution, US National
Museum of Natural History, Washington DC—Rafael Lemai-
tre, Karen Reed, and Bill Moser; Tulane University Museum
of Natural History, New Orleans, Louisiana—Hank Bart, Nel-
son Rios, and John Johansen; Auburn University Museum
xi
xii Acknowledgments
of Natural History, Auburn University, Auburn, Alabama—
Mike Gangloff (now at Appalachian State University, Boone,
North Carolina) and Brian Helms (now at Troy University,
Troy, Alabama); University of Alabama Decapod Collec-
tion, University of Alabama, Tuscaloosa, Alabama—Bernie
Kuhajda (now at the Tennessee Aquarium Research Institute,
Chattanooga, Tennessee) and Phil Harris; Eastern Kentucky
University, Richmond, Kentucky—Dave Hayes; Mississippi
Museum of Natural Science—Bob Jones; Museum of Com-
parative Zoology, Harvard University, Cambridge, Massa-
chusetts—Adam Baldinger; Illinois Natural History Survey,
Champaign, Illinois—Dan Wylie.
SPECIMEN LOANS OR GIFTS
Over the years, many people provided us loans or gifts of
crayfish specimens, many of which turned out to be valu-
able additions to our study. A small army of individuals
across the state took the time to collect specimens. These
specimens were usually sent to us preserved, but some took
on the extra burden of shipping live specimens for photo-
graphs or extraction of DNA for systematic study. It cannot
be overstated how important these specimens were to our
study, and we are thankful to these individuals for help-
ing us in this way. It allowed us to see specimens that we
would not have seen otherwise. These kind friends include
Susan Adams, US Forest Service; Dave Armstrong, ADCNR
WFFD; Jeff Baker, Alabama Power Company; Zanethia
Barnett, US Forest Service; Rebecca Bearden, GSA; Paul
Freeman, formerly of The Nature Conservancy; Jeff Garner,
ADCNR WFFD; Nicole Garrison, postdoctoral fellow at
Auburn University; Jim Godwin, Alabama Natural Heritage
Program, Auburn University; Courtney Graydon Norman,
formerly of the ADCNR State Lands Division; Chris Greene,
ADCNR WFFD; Brian Holt, ADCNR State Lands Division;
Jon Hopper, former University of Alabama graduate student,
now an instructor at Shelton State Community College;
Alex Huryn, University of Alabama, Department of Biology;
Cal Johnson, formerly of the GSA; Carol Johnston, Auburn
University; Mike Kendrick, former University of Alabama
graduate student, now with the South Carolina Department
of Marine Resources Research Institute; Steve Krotzer, Ala-
bama Power Company (retired); Paul Moler, Florida Fish
and Wildlife Conservation Commission (retired); Parker
Nenstiel, formerly of the GSA; Ashley Peters, ADCNR State
Lands Division; the late Malcolm Pierson, Alabama Power
Company; Jeff Powell, US Fish and Wildlife Service; Mike
Sandel, former University of Alabama graduate student,
now professor at the University of West Alabama; Lindsay
Shannon, Auburn University; Jeff Simmons, Tennessee Val-
ley Authority; J. Brett Smith, formerly of the GSA; Michael
Stewart, Troy University (retired); Mike Venarsky, former
University of Alabama graduate student, now in Australia;
Ken Weathers, ADCNR WFFD; Shane Welch, Clemson Uni-
versity; Traci Wood, ADCNR; and E. Anne Wynn, GSA.
FIELD ASSISTANCE
Numerous individuals have joined and assisted us in the
field over the years, and we greatly appreciate their time and
effort in helping us haul dip nets and seines, flip rocks, set and
retrieve traps, and dig burrows. They include Susan Adams,
Rebecca Bearden, Mike Buntin, B. D. Carter, Ron Cicerello,
Mike Compton, Carl Couret, Andy Ford, Jeff Garner, Court-
ney Graydon Norman, Jennifer Grunewald, Emily Hartfield,
Brian Helms, Jesse Holifield, Alex Huryn, John Johansen,
Cal Johnson, Dan Jones, Mike Kendrick, Stephanie Kilburn,
Jim Lacefield, Evan Lawrence, Jim McHugh, Parker Nenstiel,
Ashley Peters, Chris Rice, Mike Sandel, Claire Schuster, Jeff
Simmons, Brett Smith, Eric Spadgenske, Sandi Stanley, Andy
Stites, Jim Stoeckel, Claire Thomas, Bronwyn Williams, Carl
Williams, Ben Wilson, and Traci Wood.
We would like to extend a very special thank-you to Jeff
Garner and his dive crew, Mike Buntin and Jesse Holifield, for
taking on the challenge of diving in often treacherous reser-
voirs and large rivers to look for and collect crayfishes. They
provided us with some very important records that we oth-
erwise would not have gotten. We also give a special thank-
you to Alex Huryn, Mike Kendrick, Bernie Kuhajda, and
Mike Venarsky for providing us with crayfishes from special
habitats where collection is difficult, such as caves and vernal
ponds. They provided us with valuable records and insights
on species that are not commonly encountered.
INFORMATION
Large, multiyear, multifaceted works such as this compilation
require more than just a few hands and minds to perform tasks
sufficiently to create a usable, viable product. To that end, we
reached out to numerous people who gladly provided their
own expertise, materials, supplies, and hands. Ashley Peters
of the Natural Heritage Section of the State Lands Division
of ADCNR provided information on and access to properties
under its control and provided information from its Natural
Heritage Database. Dr. Sandy Ebersole of the Geological Sur-
vey of Alabama was instrumental in helping us understand
the relationships of crayfish distributions to geology/physiog-
raphy and hydrology.
MAPPING AND TECHNICAL ASSISTANCE
A comprehensive book on such a large and diverse faunal
group as the crayfishes of Alabama, a state with equally diverse
geology and aquatic resources, would be greatly diminished
in value without clear and concise maps that reflect the ranges
and overlap of the fauna, underlying geology, and watersheds.
We were fortunate in having access to excellent sources to
provide those maps, and our heartfelt thanks go out to E.
Anne Wynn, Dr. Sandy Ebersole, and Gary Hastert of GSA for
their expertise in creating those maps and for their patience
throughout the process, not only with the book but with the

several projects leading up to its creation. Thanks also go to
Eric St. Clair of the State Oil and Gas Board of Alabama and
to Ashley Peters for their assistance with watershed maps. We
are very grateful to Sherry L. Bostick for providing much-
needed assistance with the tables and graphs in the book.
PHOTO USAGE
We would like to thank Susan Adams, Chris Lukhaup, and Jim
Lacefield for their generosity in allowing us to use their photos.
AUTHORS’ PERSONAL THANKS
(GAS) I am first and foremost eternally grateful to my fam-
ily, especially my wife, Claire, for her continual support over
the past 50 years. They have always supported my need to be
on field trips and at museums, conferences, and meetings. In
the process, I have missed birthdays, anniversaries, and other
family milestones. Without their continual understanding
and love, my career would not have been possible. I would
also like to thank my parents, Theresia and Franz Schus-
ter, who took me by the hand in Linz, Austria, in 1955 with
fifty dollars in their pocket and emigrated to Dayton, Ohio.
Because of World War II, neither had more than a sixth-grade
education, yet throughout my life they impressed upon my
sisters and me the importance of a good education. I know
they were often befuddled by my career choices and had little
understanding of things such as freshwater mussels, caddis-
flies, and crayfishes, but they never wavered in their support
of my choices.
This project began over 14 years ago and in many ways
represents the culmination of my career. I would be remiss if
I did not mention some of the most important professional
influences in my life. These, of course, are the outstand-
ing teachers and mentors I had in my early education as an
aquatic zoologist, naturalist, and teacher. These include Drs.
Ossi Lindqvist (University of Dayton, Dayton, Ohio), Don-
ald Batch and Branley Branson (Eastern Kentucky University,
Richmond, Kentucky), and David “Ets” Etnier (University of
Tennessee, Knoxville, Tennessee). Each left important lifelong
imprints on me. I am indebted to them for whatever success I
have had in my career. I would also like to thank Dr. Horton
H. Hobbs Jr., a consummate crayfish researcher who described
more crayfish species than anyone else. His contributions to
astacology cannot be overstated. I met him as a master’s stu-
dent on a visit to the Smithsonian. I will never forget his great
kindness to me in opening his cabinet of crayfish reprints and
allowing me to take whatever I wanted. More importantly,
he encouraged me to describe my first new crayfish species,
Cambarus batchi. It was that little push that brought me so
much joy and fulfillment in my professional life.
I would also like to thank colleagues Susie Adams,
Rebecca Bearden, Ron Cicerello, Arnie Eversole, Jeff Garner,
Zac Loughman, Barbara Ramey, Henry Robison, Jeff Sim-
mons, Chris Skelton, Brett Smith, Roger Thoma, Bronwyn
Acknowledgments xiii
Williams, and Carl Williams for their help and support over
the years. I would like to thank many of my former students
who became professional colleagues and valued friends, and
who provided me with continuing interest and support in my
work: Brett Billings, Bob Butler, Mike Compton, Zack Couch,
Bob DiStefano, Wendell Haag, John Johansen, Rebecca Blan-
ton Johansen, Dan Jones, Ellis Laudermilk, Mathew Patterson,
Greg Pond, and Matt Thomas. I would also like to thank the
Society of Foundation Professors of Eastern Kentucky Uni-
versity for awarding me the John D. Rowlett Award for Fac-
ulty Scholarship, Research, or Professional Development for
2020. This award came with a $5,000 endowment that helped
defray publication costs. This book would not have been pos-
sible without my coauthors Chris Taylor and Stuart McGre-
gor, who tirelessly worked on all aspects of this project. They
made me a better biologist, and I have loved working with
them. I will greatly miss our Alabama field trips as well as our
endless search for the best barbecue restaurants in the state.
(CAT) I am indebted to the sacrifices made by my wife, Rita,
and son Jason. Their understanding of my passion for cray-
fishes and desire to complete this project allowed me to spend
a significant amount of time away from home since 2005.
It was never easy, but they never complained. I thank my
graduate advisers, Drs. Brooks Burr and Lawrence Page, for
instilling in me the value of hard work and the importance
of disseminating science in peer-reviewed literature. I am
also thankful to the Illinois Natural History Survey (INHS)
and Prairie Research Institute for never questioning my time
commitment to this research. Cathy Bialeschki at INHS
was most helpful with all aspects of travel planning. Many
of my former graduate students, particularly Stephanie Kil-
burn, Cody Rhoden, Christopher Rice, Andy Stites, and Clair
Thomas, assisted with numerous aspects of data analysis and
fieldwork, and for that I am thankful. Finally, I would like to
thank my two coauthors. An endeavor such as this would not
be possible without close interaction with dedicated, knowl-
edgeable, and fun colleagues. I learned so much from both of
them and will be forever grateful.
(SWM) I wish to take this opportunity to thank the people
whose influences on my life led to my career path, and specif-
ically my participation in this work. Primarily my late father,
L. W. “Skeets” McGregor, and my mother, Ann M. McGregor,
who supported me through my formative years and education.
Daddy was especially instrumental in introducing me to the
wonders of nature and taught me to appreciate and value them.
Mom was instrumental in teaching me the value of learning.
Their respective influences led me to a career that allowed me
to affect and protect our beautiful world. My nephew, Josh
Ford, offered me a companion on many trips to the great out-
doors and gave me the opportunity to share my knowledge
and experience by teaching him many of the valuable lessons
I had learned from my parents. Now he and his wife, Parris,
have given me my great-nephew, Cahlan, and great-nieces,
xiv Acknowledgments
Ella and Evie, who I hope will afford me additional opportuni-
ties to share that knowledge. I also thank my sister, Kelly Ford,
for being a protective big sister to a bratty little brother, and
for giving me the gift of my nieces and nephews! Ashley Peters
deserves a special thank-you as well, not only for her support
of my work on this book as field help and in her considerable
mapping and database talents in the Natural Heritage Section
of the State Lands Division, but also for her patience with my
long absences, literally and figuratively!
Thanks also go to the many educators and colleagues I
met along the way, all of whom had some level of influence
on my path, but a few of whom deserve special acknowledg-
ment. The late Clara Yeatman (second grade) and the late
Niki Graham (eleventh grade) were especially influential
teachers who encouraged me to appreciate learning and to
broaden my horizons. The late Dr. Paul Yokley of the Univer-
sity of North Alabama was a leader in southeastern biology
and learning and was a very positive influence during my
undergraduate studies. During graduate school at Tennessee
Tech University I worked for the US Fish and Wildlife Service
under the capable direction of Tom Talley, Bob Bay, and the
late Doug Winford, and their leadership and friendship are
much appreciated.
During my three-plus decades at GSA I have been fortu-
nate to work with many capable and cooperative colleagues.
My gratitude is expressly given to Dr. Pat O’Neil, under whose
wing I served and learned during most of my career at GSA;
and to Dr. Scott Mettee, my original supervisor, who took a
chance and hired me as a naive but ambitious recent college
graduate. Tom Shepard and I spent many, many long days
and often nights together roaming the wilds of Alabama
collecting various taxa, and from very few people have I
learned more. Tom is, indeed, one of the most well-informed
and modest people I have ever known, and my thanks go
to him for his friendship. Nick Tew, the state geologist, and
Bennett Bearden, the assistant state geologist, are competent
leaders who guide with a firm and fair hand; they are also
good and trusted friends who continually supported our
project. Rebecca Bearden came to work at GSA during the
latter stages of research for the book and was instrumental in
gathering many of our field collections; her contributions are
greatly appreciated.
Some colleagues I have worked with over the years who
influenced my contribution to this project include the late Dr.
David Stansbery of Ohio State University, the late Dr. Hor-
ton H. Hobbs III of Wittenberg University in Springfield,
Ohio, and the late Dr. John C. Cooper of the North Carolina
Museum of Natural Sciences in Raleigh. Dr. Stansbery was a
mentor during my early professional years and was instru-
mental in teaching me about freshwater mussels; he instilled
an understanding of the challenges in identifying difficult
taxa. Horton and his students accompanied me on numerous
trips into caves in northern Alabama; he identified numer-
ous crayfish and taught me a lot about cave inhabitants. John
shared many details of his considerable cave-related work in
Alabama with me, mostly over the phone and in snail mail
and email exchanges when he edited past works of mine. Dr.
Jim Williams of Gainesville, Florida, selflessly provided infor-
mation from his considerable storehouse of knowledge of the
southeastern fauna throughout my career.
A very special thank-you is due Jeff Garner of the WFFD.
His extraordinary contributions to my career are eclipsed
only by his friendship. Over the past 25-plus years we have
gone on many adventures together and I have benefited
immeasurably from his vast knowledge, curiosity, sense of
humor, and culinary skills. Words alone cannot express the
respect and affection I have for Jeff, so a hearty “thank you!”
will have to suffice.
And finally, I would like to thank the two senior authors
for honoring me by inviting me to participate in this project.
Guenter was a favorite professor from graduate school and
has a sincere and genuine interest in and affection for his stu-
dents. He has been a cherished friend and colleague since the
summer of 1986. Chris has patiently shared with me his con-
siderable wealth of knowledge on crayfishes over the life of
this work, and I thank them both!
ABBREVIATIONS

The following abbreviations are used in the text for state and
federal agencies, institutions, and nongovernmental organiza-
tions, as well as standard technical terms and measurements.

ADCNR Alabama Department of Conservation and K knob

Natural Resources MCZ Museum of Comparative Zoology, Harvard
AL areola length University, Cambridge, Massachusetts
AP adventitious process MI first form male
AUMNH Auburn Museum of Natural History, Auburn, MII second form male
Alabama MMNS Mississippi Museum of Natural Science,
AW areola width Jackson, Mississippi
BLM bladelike mandibles MP mesial process
C carina NCSM North Carolina Museum of Natural Sciences,
CaK caudal knob Raleigh, North Carolina
CaP caudal process PCL postorbital carapace length
CB caudal bulge RL rostrum length
CeK cephalic knob RW rostrum width
CeP cephalic process S shoulder
CP central projection TCL total carapace length
ESA Endangered Species Act TUMNH Tulane University Museum of Natural History,
Fe female New Orleans, Louisiana
GSA Geological Survey of Alabama, Tuscaloosa, TVA Tennessee Valley Authority, Chattanooga,
Alabama Tennessee
INHS Illinois Natural History Survey, Champaign, UADC University of Alabama Decapod Collection,
Illinois Tuscaloosa, Alabama
IUCN International Union for Conservation of USNM US Museum of Natural History, Smithsonian
Nature, Cambridge, United Kingdom Institution, Washington, DC
I–V first pereiopod (walking leg) to fifth pereiopod VMS ventromedial spine on carpus

xv
ALABAMA CRAYFISH TAXA

The following list includes the genus and species names of 99

taxa, along with the author names and common names of the
crayfishes known to live in Alabama. The classification follows
that of Crandall and De Grave (2017).

Genus Species Authors Common Name

Barbicambarus simmonsi Taylor and Schuster, 2010 Tennessee Bottlebrush Crayfish

Cambarellus diminutus Hobbs, 1945 Least Crayfish

Cambarellus lesliei Fitzpatrick and Laning, 1976 Angular Dwarf Crawfish

Cambarellus rotatus Schuster and Kendrick, 2017 Twisted Dwarf Crayfish

Cambarellus shufeldtii (Faxon, 1884) Cajun Dwarf Crayfish

Cambarus andersoni Jones and Eversole, 2015 Florence Crayfish

Cambarus bartonii cavatus Hay, 1902a Appalachian Brook Crayfish

Cambarus clairitae Schuster and Taylor, 2016 Zebra Crayfish

Cambarus coosae Hobbs, 1981 Coosa Crayfish

Cambarus cracens Bouchard and Hobbs, 1976 Slenderclaw Crayfish

Cambarus distans Rhoades, 1944 Boxclaw Crayfish

Cambarus diupalma Jones and Eversole, 2015 Mountain Fork Crayfish

Cambarus englishi Hobbs and Hall, 1972 Tallapoosa Crayfish

Cambarus gentryi Hobbs, 1970 Linear Cobalt Crayfish

Cambarus girardianus Faxon, 1884 Tanback Crayfish

Cambarus graysoni Faxon, 1914 Twospot Crayfish

Cambarus halli Hobbs, 1968 Slackwater Crayfish

Cambarus hamulatus (Cope, 1881) Prickly Cave Crayfish

Cambarus howardi Hobbs and Hall, 1969 Chattahoochee Crayfish

Cambarus jonesi Hobbs and Barr, 1960 Alabama Cave Crayfish

Cambarus laconensis Buhay and Crandall, 2009 Lacon Exit Cave Crayfish

Cambarus latimanus (LeConte, 1856) Variable Crayfish

Cambarus lentiginosus Jones and Eversole, 2016 Speckled Crayfish

Cambarus longirostris Faxon, 1885c Longnose Crayfish

Cambarus manningi Hobbs, 1981 Greensaddle Crayfish

xvii
xviii Alabama Crayfish Taxa

Genus Species Authors Common Name

Cambarus obstipus Hall, 1959 Sloped Crayfish

Cambarus parvoculus Hobbs and Shoup, 1947 Mountain Midget Crayfish

Cambarus pecki Hobbs, 1967b Phantom Cave Crayfish

Cambarus pyronotus Bouchard, 1978 Fireback Crayfish

Cambarus rusticiformis Rhoades, 1944 Depression Crayfish

Cambarus scotti Hobbs, 1981 Chattooga River Crayfish

Cambarus speleocoopi Buhay and Crandall, 2009 Sweet Home Alabama Cave Crayfish

Cambarus striatus Hay, 1902a Ambiguous Crayfish

Cambarus tenebrosus Hay, 1902a Cavespring Crayfish

Cambarus unestami Hobbs and Hall, 1969 Blackbarred Crayfish

Cambarus veitchorum Cooper and Cooper, 1997a White Spring Cave Crayfish

Creaserinus burrisi (Fitzpatrick, 1987a) Burrowing Bog Crayfish

Creaserinus byersi (Hobbs, 1941) Lavender Burrowing Crayfish

Creaserinus danielae (Hobbs, 1975) Speckled Burrowing Crayfish

Creaserinus fodiens (Cottle, 1863) Digger Crayfish

Faxonella clypeata (Hay, 1899a) Ditch Fencing Crayfish

Faxonius alabamensis (Faxon, 1884) Alabama Crayfish

Faxonius compressus (Faxon, 1884) Slender Crayfish

Faxonius cooperi (Cooper and Hobbs, 1980) Flint River Crayfish

Faxonius durelli (Bouchard and Bouchard, 1995) Saddle Crayfish

Faxonius erichsonianus (Faxon, 1898) Reticulate Crayfish

Faxonius etnieri species complex Bouchard and Bouchard, 1976

Faxonius forceps (Faxon, 1884) Surgeon Crayfish

Faxonius holti (Cooper and Hobbs, 1980) Bimaculate Crayfish

Faxonius jonesi (Fitzpatrick, 1992) Sucarnoochee River Crayfish

Faxonius juvenilis (Hagen, 1870) Kentucky River Crayfish

Faxonius lancifer (Hagen, 1870) Shrimp Crayfish

Faxonius mirus (Ortmann, 1931) Wonderful Crayfish

Faxonius palmeri palmeri (Faxon, 1884) Gray-Speckled Crayfish

Faxonius perfectus (Walls, 1972) Complete Crayfish

Faxonius placidus (Hagen, 1870) Bigclaw Crayfish

Faxonius spinosus (Bundy, 1877) Coosa River Spiny Crayfish

Faxonius validus (Faxon, 1914) Powerful Crayfish

Faxonius virilis (Hagen, 1870) Virile Crayfish

Faxonius yanahlindus (Taylor et al., 2016) Spinywrist Crayfish

Hobbseus prominens (Hobbs, 1966) Prominence Rivulet Crayfish

Lacunicambarus acanthura (Hobbs, 1981) Thornytail Crayfish

 Alabama Crayfish Taxa xix

Genus Species Authors Common Name

Lacunicambarus dalyae Glon et al., 2019 Jewel Mudbug

Lacunicambarus erythrodactylus (Simon and Morris, 2014) Warpaint Mudbug

Lacunicambarus freudensteini Glon et al., 2020 Banded Mudbug

Lacunicambarus ludovicianus (Faxon, 1884) Painted Devil Crayfish

Lacunicambarus miltus (Fitzpatrick, 1978) Rusty Grave Digger

Lacunicambarus mobilensis Glon et al., 2020 Lonesome Gravedigger

Orconectes australis (Rhoades, 1941) Southern Cave Crayfish

Orconectes sheltae Cooper and Cooper, 1997b Shelta Cave Crayfish

Procambarus acutissimus (Girard, 1852) Sharpnose Crayfish

Procambarus acutus (Girard, 1852) White River Crawfish

Procambarus bivittatus Hobbs, 1942a Ribbon Crayfish

Procambarus capillatus Hobbs, 1971 Capillaceous Crayfish

Procambarus clarkii (Girard, 1852) Red Swamp Crawfish

Procambarus clemmeri Hobbs, 1975 Cockscomb Crayfish

Procambarus escambiensis Hobbs, 1942a Escambia Crayfish

Procambarus evermanni (Faxon, 1890) Panhandle Crayfish

Procambarus hagenianus hagenianus (Faxon, 1884) Southeastern Prairie Crayfish

Procambarus hayi (Faxon, 1884) Straightedge Crayfish

Procambarus holifieldi Schuster et al., 2015 Celestial Crayfish

Procambarus hubbelli (Hobbs, 1940b) Jackknife Crayfish

Procambarus hybus Hobbs and Walton, 1957 Smoothnose Crayfish

Procambarus lagniappe Black, 1968 Lagniappe Crayfish

Procambarus lecontei (Hagen, 1870) Mobile Crayfish

Procambarus lewisi Hobbs and Walton, 1959 Spur Crayfish

Procambarus lophotus Hobbs and Walton, 1960 Mane Crayfish

Procambarus marthae Hobbs, 1975 Crisscross Crayfish

Procambarus okaloosae Hobbs, 1942a Okaloosa Crayfish

Procambarus paeninsulanus (Faxon, 1914) Peninsula Crayfish

Procambarus planirostris Penn, 1953 Flatnose Crayfish

Procambarus shermani Hobbs, 1942a Gulf Crayfish

Procambarus spiculifer (LeConte, 1856) White Tubercled Crayfish

Procambarus suttkusi Hobbs, 1953 Choctawhatchee Crayfish

Procambarus verrucosus Hobbs, 1952 Grainy Crayfish

Procambarus versutus (Hagen, 1870) Sly Crayfish

Procambarus viaeviridis (Faxon, 1914) Vernal Crayfish

Procambarus vioscai paynei Fitzpatrick, 1990c Payne’s Creek Crayfish

Procambarus zonangulus Hobbs and Hobbs, 1990 Southern White River Crawfish
CRAYFISHES OF ALABAMA
1
Introduction
Alabama is known for its natural beauty, seemingly endless
supply of freshwater, and abundant wildlife species that are
often referred to as “charismatic megafauna.” Alabama is per-
haps less known for its tremendous diversity of nongame
species, species that to some are far less charismatic but are
nevertheless of great importance ecologically. In Alabama’s
waterways live more species of fishes, mussels, snails, and
crayfishes than in any other state in the United States.
If one were to travel the backroads of Alabama and engage
local citizens in conversation, one would probably learn a
great deal of important information about the immediate
area and its abundant riches: which local spring has the “best
water you ever drank”; where mountain lions and black bears
roam; where the “blue hole” in the local creek has no bot-
tom; what hollow had the most moonshine stills; and whose
grandpaw made the best whiskey. And you would also find
that nearly every stream in Alabama has abundant crayfishes
in it. Indeed, what child has not flipped over rocks in the local
creek, dragged a net through a pond or along a riverbank, or
tied a piece of bacon or chicken to a string and lowered it into
a hole to catch crawdads? These animals do seem to hold a
special place in the hearts and minds of children and adults
statewide, and for good reason: Alabama has a world-class
crayfish fauna. Almost any spring, creek, river, pond, lake,
marsh, swamp, ditch, bog, or cave has crayfish in it. If there is
water present, there likely are crayfish living in it.
Crayfishes, sometimes called crawfishes, crawdads, ditch-
bugs, yabbies, flusskrebs, or mudbugs, are a taxonomically
and ecologically diverse group of aquatic crustaceans. World-
wide there are over 600 known species, and new species are
described each year. They are common organisms in many
freshwater habitats, including lotic and lentic habitats as well
as caves and underground burrows that reach to the water
table. So, why are they important? What are they good for?
In their environments, they are usually the largest inverte-
brates and often represent the greatest amount of invertebrate
biomass. As crayfish biologists are fond of saying, “They eat
everything, and everything eats them.” This simple statement
underlines the great ecological importance of crayfishes.
Crayfishes are crustaceans that belong to the order
Decapoda, the infraorder Astacidea, and the superfamily Asta-
coidea. Freshwater crayfishes are native to all continents except
Africa and Antarctica. The crayfishes of the world represent a
monophyletic group of crustaceans that belong to five fami-
lies: Astacidae, Cambaridae, Cambaroididae, Cricoidoscelosi-
dae, and Parastacidae (Crandall et al. 2000; Crandall and De
Grave 2017). They are an old group that can be traced back
about 185 to 200 million years ago (mya) to the supercontinent
Figure 1.1. Changes in numbers of recog-
nized Alabama species according to year
and authors.
1
Figure 1.2. Number of Alabama crayfish
species in each genus. Two recently
described Lacunicambarus species not
included in totals.

Figure 1.3. Total crayfish distribution

records from across Alabama.

Pangaea (Crandall et al. 2000). The separation of Pangaea into
the northern Laurasia and the southern Gondwana about 185
mya resulted in an evolutionary split in crayfishes into North-
ern and Southern Hemisphere faunas. The family Parastacidae
is restricted to the Southern Hemisphere and is found in South
America, Madagascar, Australia, and New Guinea. There are
currently about 200 species in this family (Crandall and De
Grave 2017). The other four families are found in the North-
ern Hemisphere. The family Cricoidoscelosidae is a recently
described fossil family from China (R. S. Taylor et al. 1999).
The family Cambaroididae is restricted to Asia and consists of
six species (Crandall and De Grave 2017). In recent years, the
position of Asian species in the genus Cambaroides has been
debated. Crandall and De Grave (2017) elevated this group of
six species to their own family based on molecular data. The
family Astacidae is distributed in Europe, western Asia, and
western North America and comprises about 20 species (Cran-
dall and De Grave 2017). The largest family is Cambaridae,
which consists of approximately 400 species (Crandall and De
Grave 2017). It is restricted to North America, including parts
of Central America, Cuba, Mexico, the United States, and Can-
ada. The crayfishes of Alabama all belong to this family.
This book represents the most in-depth treatment of cray-
fishes for the state of Alabama to date. The number of species
recognized as living in Alabama has changed over the years.
Figure 1.1 shows these changes according to the authorities
and years the lists were published. A current list of all known
species for the state is provided (see “List of Alabama Crayfish
Taxa”). Alabama currently contains 99 species. The number of
species within each genus is provided in figure 1.2. The find-
ings in this book are based on over 9,000 records from more
than 2,600 collecting locations across the entire state (fig. 1.3).
METHODS
We compiled historical records for Alabama crayfishes by
requesting museum records from the following institutions:
Geological Survey of Alabama (GSA), Auburn University
Museum of Natural History (AUMNH), Illinois Natural
Figure 1.4. Number of Alabama crayfish
specimen lot records over decades of
collecting.
Introduction 3
History Survey (INHS), National Museum of Natural History
(USNM), Mississippi Museum of Natural Science (MMNS),
Museum of Comparative Zoology at Harvard University
(MCZ), Tulane University Museum of Natural History
(TUMNH), and the University of Alabama Decapod Col-
lection (UADC). This data query resulted in over 4,500
specimen lots (unique collections of a single species from a
unique location) ranging in dates from a few pre-1900 col-
lections to 2004 (fig. 1.4). We subsequently visited AUMNH,
INHS, USNM, TUMNH, and UADC and examined any spec-
imens for which the identification appeared dubious or the
species was not identified. The exact collection location (i.e.,
latitude and longitude) of some records could not be deter-
mined because of the scant locality information on the col-
lection label. After examination of museum specimens, our
list of historical records included 4,259 specimen lots from
approximately 2,200 unique locations. These records were
used to build a FileMaker Pro (FileMaker, Inc.) database that
contained all available specimen metadata, including but not
limited to species identification, number of individuals col-
lected, location, date of collection, collectors, identifier, and
museum. For those records that did not include latitude and
longitude, we georeferenced them if adequate common loca-
tion information was available. Approximately 425 historical
specimen lots lacked either common location information or
date of collection and were not used in our data analysis.
Fieldwork for this project was conducted over a 14-year
period, with various levels of collecting effort per year (fig.
1.5). During this period, a total of 4,487 lots were collected
from approximately 2,600 unique locations sampled for cray-
fishes across Alabama. The two senior authors usually made
biannual collecting trips to the state, mainly in the spring and
fall of each year. The timing of the trips was chosen to maxi-
mize the chances of obtaining MI (first form male) crayfishes
and finding workable water levels in target sampling habitats.
In addition to the authors, other personnel from the GSA and
other state and federal agencies, as well as private individuals,
were involved in collecting Alabama crayfishes and sending
specimens to the authors for verification and vouchers. These
4 Chapter 1
collaborators are listed in the acknowledgments. All records
for crayfishes collected by the authors and other collaborators
were added to our previously constructed database in File-
Maker Pro.
While we attempted to sample all areas of Alabama and
conduct fieldwork in a random fashion, after gathering data
from our initial museum research we were able to determine
which areas of the state might have been undersampled. We
also realized that certain habitat types around the state had
not been well sampled (e.g., swamps, marshes, vernal pools,
roadside ditches, large rivers, and reservoirs). With this in
mind, we used historical distribution data and habitat type
information to plan some field trips during this project.
The physiographic map used in this book was modified
from Ebersole et al. (2019). Features on the map were derived
from several online sources and compiled to create the map
using ArcGIS Desktop 10.5, then imported into Adobe Illus-
trator for manipulation. County boundaries were derived
from the US Census Bureau TIGER/Line County Shapefile
(US Census Bureau 2015), and state boundaries, rivers, and
other water bodies were derived from the National Atlas of
the United States (2005a, 2005b). Distribution maps in the
species accounts use the same base map. The map of water-
sheds in Alabama was created in ArcGIS and manipulated to
define major watersheds. The approximate location of the Fall
Line was manually added. The maps of dams on the major
rivers were similarly created.
We experimented with displaying species richness with
several different hydrologic units as defined by Seaber et al.
(1987). In their system, the United States is divided and sub-
divided into successively smaller hydrologic units classified
into four levels: regions, subregions, accounting units, and
cataloging units. The hydrologic units are nested within each
other, from the largest in area (regions) to the smallest in area
(cataloging units). Each successively smaller hydrologic unit
is identified by a unique hydrologic unit code (HUC) consist-
ing of two to eight digits based on the four levels of classifi-
cation in the hydrologic unit system. We experimented with
Figure 1.5. Number of Alabama crayfish
specimen lot collections during the
course of this study.
using HUC 4-, 8-, 10-, and 12-digit levels with varying degrees
of confidence. At the HUC 4 and 8 levels, significant portions
of too many HUCs lay outside the state, and without crayfish
data from adjacent states, no accurate scores could be deter-
mined. Conversely, the HUC 12 level presented rather small
geographic areas, and without uniform, standardized sam-
pling across all HUCs, a greater abundance of local sampling
presented disproportionately high richness in some cases,
with no confidence in the results. While it is not perfect by
any means, we found that the HUC 10 level offered the most
reasonable assessment of species richness statewide, with
fewer and smaller portions of HUCs falling outside the state
and less likelihood of local sampling density biasing results.
The HUC 10 map displaying crayfish species richness was cre-
ated in ArcGIS Desktop 10.5.
The collecting gear we used for this project included all the
gear described in the “Collecting Crayfishes” section. Once
crayfishes were captured, they were put into buckets with
water at the bottom. When a particular site was adequately
sampled, representative specimens were kept for preservation
and photography. Labels with the following detailed informa-
tion were placed into each individual container used for that
site: field number, state, county, water body name, description
of site location, latitude and longitude, date, and collector
names. Labels were placed into each individual container
used for that site. Representative individuals were chosen to
be kept alive for photography, and the remaining individuals
were preserved. Those that were to be preserved were put into
a 70% ethanol solution. Before preservation of specimens, if
necessary, samples of abdominal muscle tissue were removed
from selected specimens and preserved in 95% to 100% etha-
nol for later analysis.
Specimens that were kept alive for photographs were put
individually into slightly inflated, resealable plastic bags with
a very small amount of water (only enough to keep the inter-
nal relative humidity at 100%). All the individual bags from a
site were put into larger resealable plastic bags with one label
and put into a cooler with ice. A removable shelf in the cooler

separated the crayfishes from the ice, and the ice was regu-
larly replenished. The crayfishes stored in this fashion could
be kept alive in the field for longer than one week. In the lab-
oratory, each individual bag was cleaned with fresh tap water
and stored in the cooler until the crayfishes could be photo-
graphed. After photographing, specimens were preserved and
added to the original field collection.
COLLECTING CRAYFISHES
A typical field crew consisted of at least two, and often as many
as five or six individuals. The most useful method for collect-
ing crayfishes in streams was seining. A nylon seine with 0.12-
inch (0.3 cm) mesh and measuring 10 feet (3 m) in length
and 5 feet (1.5 m) in depth was used in a variety of habitats.
The seine had two brail poles and small lead weights along
the bottom. Seining was most effective in shallow riffles and
pools of streams and in lentic habitats without dense vegeta-
tion. Seining usually involved two people holding the brails,
while one or two others kicked and turned over rocks and
debris in front of the seine, washing crayfishes into the seine.
In lentic habitats, the seine was dragged along the bottom and
then pulled out of the water. Sometimes it was helpful to have
one or two people walking in front of the seine to disturb the
bottom. Dip nets were also employed in streams to capture
crayfishes from undercut banks or in leafy and woody debris.
Dip nets were also very useful in roadside ditches, marshes,
and swamps, especially in habitats with dense vegetation and
organic debris.
Burrowing species were captured mostly by digging out
the burrows with shovels or trowels. Extracting crayfishes
from burrows can be difficult, time consuming, and often
frustrating. It may not be clear from casual examination of a
burrow whether there is a crayfish in it. Fresh, wet mud balls
around the opening or on the chimney of the burrow may be
a good indication that there may be a crayfish inside. Some-
times capturing the crayfish required completely opening the
burrow, while other times opening the burrow to the water
level and agitating the water surface would entice the crayfish
to the surface where it could be grabbed. Burrowing crayfishes
were also caught on warm rainy nights when they were walk-
ing outside their burrows. Sometimes when crayfishes were
hunted with a flashlight at night, they were seen at the lip of
burrow openings. Although they tended to be quite skittish,
with some prowess by the collector they could sometimes be
nabbed. In addition, after heavy rains, burrowers were some-
times found in water-filled tire ruts or in low-lying pools in
fields where burrows were common.
Baited minnow traps with funneled openings on both
sides were also used. The bait was usually canned cat food,
and traps were left in place for several days before being
retrieved. These were especially useful in deep-water environ-
ments. Diving with a surface air source was also employed
in large rivers and reservoirs. These habitats are very difficult
to collect in otherwise. The diver would handpick exposed
Introduction 5
crayfishes from the bottom or from under debris and rocks
along the bottom.
TAXONOMY AND SYSTEMATICS
As currently recognized by Crandall and De Grave (2017),
the family Cambaridae consists of 14 genera, one of which,
Palaeocambarus, is known only from fossils. The taxonomy
of this family was relatively stable until the mid-1990s, mainly
because of the work of Horton H. Hobbs Jr., and Joseph F.
Fitzpatrick Jr. Thanks to their efforts, new genera (such as
Hobbseus Fitzpatrick and Payne 1968) were named, and the
most diverse genera, including Cambarellus (Fitzpatrick
1983), Cambarus (Hobbs 1969), Fallicambarus (Hobbs 1973),
Orconectes (Fitzpatrick 1987b), and Procambarus (Hobbs
1972b), were divided into subgenera. However, with the advent
of molecular phylogenetics, these classification schemes came
into question. Important papers have been published for each
of these genera questioning the validity of the recognized
subgenera within each (reviewed in Crandall and De Grave
2017). This has led to some dramatic changes in cambarid
taxonomy and systematics. Some of the changes recognized
by Crandall and De Grave (2017) that are significant to our
understanding of Alabama crayfishes are summarized below.
First, molecular data do not support the recognition of the
subfamily Cambarellinae, nor do they support the subgenus
Dirigicambarus within Cambarellus (Crandall and De Grave
2017). Hence, there are now two recognized subgenera: the
subgenus Cambarellus is restricted to Mexico, and the subge-
nus Pandicambarus is restricted to the Gulf Coastal Plain of
the southeastern United States.
Second, in the genera Cambarus and Procambarus, no
subgenera are currently recognized. In addition, within the
genus Cambarus, the former subgenera Lacunicambarus
and Tubericambarus have been elevated into a single genus,
Lacunicambarus (Glon et al. 2018). In Alabama, five taxa were
moved from Cambarus to this newly elevated genus.
Third, the genus Fallicambarus previously had two subge-
nera: Creaserinus and Fallicambarus. Each is now recognized
as a separate genus. All four of the Alabama species in the
former genus Fallicambarus are now relegated to the genus
Creaserinus. Most of the species in Fallicambarus, as currently
recognized, are distributed west of the Mississippi River.
Fourth, one of the most significant changes affecting Ala-
bama crayfish taxonomy occurred in the genus Orconectes.
None of the formerly recognized subgenera are currently
recognized. In addition, the genus was divided into two gen-
era based on molecular studies showing two distinct genetic
lineages (i.e., clades) in what was called Orconectes. The
clades basically represented an ecological split into species
restricted to living in caves (stygobites) versus those living
in surface waters (epigean species). The type species of the
genus Orconectes is O. inermis, a stygobite; thus the generic
name is restricted to the clade consisting of obligate cave
species. Because of this, the clade consisting of the epigean
6 Chapter 1
species required a different name. The oldest available name
was Faxonius, which was first used by A. E. Ortmann (1905).
For the Alabama crayfish fauna, this means that the genus
Orconectes, as now recognized, consists of two cave species:
O. australis and O. sheltae. All other Alabama species formerly
in Orconectes are now in the genus Faxonius.
Taxonomy, like all other fields of science, is in a constant
state of flux, which is the result of greater understanding
through new research findings. In terms of taxonomy and
systematics, the main goal is that the nomenclature and clas-
sification of all species reflect evolutionary events. In the last
20-plus years, a great deal of new knowledge about evolution-
ary relationships among not only crayfishes, but all species, has
been gained through the exploding field of molecular biology.
One of the more important things we have learned in cray-
fish taxonomy and systematics is that many morphological
characters that were thought to indicate crayfish evolutionary
relationships actually represent evolutionary convergence. It
has become incumbent upon crayfish taxonomists to look
beyond morphological similarities or differences to obtain a
true vision of past evolutionary events.
HISTORY OF CRAYFISH WORK IN ALABAMA
The first known crayfish collection from Alabama curated into
a scientific collection was that of specimens of Procambarus
lecontei (= Cambarus lecontei, sensu Hagen). It was collected
(as an undescribed species) by Louis Agassiz, the founder
of the MCZ at Harvard University. He collected a series of
specimens (one MI, two MIIs, and seven females) while in
Mobile, Alabama, in 1853 on a speaking tour. The specimens
were curated into the MCZ upon its establishment in 1859. In
1867, Herman Hagen, a German entomologist, joined the staff
of the MCZ at the invitation of Agassiz. Hagen’s specialty was
neuropteroid insects, in particular the order Trichoptera, and
Agassiz assigned him, among other things, to work on North
American crayfishes. Three years later, Hagen published his
first and only crayfish work, Monograph of the North Amer-
ican Astacidae (Hagen 1870). In this monograph he formally
described Agassiz’s Procambarus lecontei, as well as five other
species—Faxonius juvenilis, F. lancifer, F. placidus, F. virilis,
and Procambarus versutus—which are now known from Ala-
bama. Two of these six species, F. juvenilis and F. virilis, are not
native to the state but now reside here.
Prior to Hagen’s 1870 monograph, a total of 7 species had
been described that are now known from Alabama (Girard
1852; LeConte 1856; Cottle 1863). From Hagen’s publication
in 1870 until 1914, a total of 22 species now known to occur
in Alabama were described. Most of these (16 species) were
described by Walter Faxon (1884, 1885c, 1890, 1898, 1914);
W. P. Hay described four species (Hay 1899a, 1902a); and
Cope (1881) and Bundy (1877) each described one species.
So, between 1852 and 1914, a total of 35 species known from
Alabama were described. This number represents 36% of the
total known fauna. From 1914 to 1940, only a single species,
Faxonius mirus, was described (Ortmann 1931). The most
productive years were between 1940 and 2017, when 61 spe-
cies (63%) known to occur in Alabama were described. These
include the years that Horton H. Hobbs Jr. was actively study-
ing crayfishes. He described 33 species (33% of the fauna)
known from Alabama between 1940 and 1990. Since 1990, a
total of 15 species have been described that live in Alabama.
These include species described by Fitzpatrick (1992), Bouch-
ard and Bouchard (1995), J. Cooper and Cooper (1997a and
1997b), Thoma et al. (2005), Buhay and Crandall (2009), C.
Taylor and Schuster (2010), Jones and Eversole (2015), Schus-
ter et al. (2015), Jones and Eversole (2016), Schuster and Tay-
lor (2016), C. Taylor et al. (2016), and Schuster and Kendrick
(2017).
Bouchard (1976a) was the first to attempt a complete
checklist of Alabama crayfishes (fig. 1.1). He listed 58 species
for the state and estimated that as many as 75 species would
eventually be found. In the same publication Bouchard was
the first to attempt to evaluate the conservation status of Ala-
bama’s crayfish species. He suggested that 13 crayfish species
needed some sort of conservation protection. He listed one as
threatened and the remaining 12 as species of special concern.
Hobbs, in his Illustrated Checklist of the American Cray-
fishes (1989), listed 72 species and subspecies for the state and
presented two species as questionable (Creaserinus hedgpethi
and Procambarus bivittatus). Since that publication, C. hedg-
pethi has been synonymized with C. fodiens, and P. bivittatus
has been recorded for the state. Fitzpatrick (1990a) prepared
a list of only those species he considered in need of conser-
vation protection. C. Taylor et al. (1996) presented a compre-
hensive list for the state, and it also was the first extensive
review of the conservation of all North American crayfish
species. They listed a total of 83 species as occurring in Ala-
bama. A second review of North American crayfish species
(C. Taylor et al. 2007) listed 86 taxa, including species and
subspecies, for the state (fig. 1.1). The increase in the num-
ber of species from 1997 to 2007 was attributed to taxonomic
revisionary work and new species descriptions. Schuster et
al. (2008), based on over 4,000 database records accumu-
lated from various museum holdings and recent collections,
listed a total of 85 species. They also listed Faxonius virilis
as the only nonnative species. The first record of this species
in Alabama is a 1967 USNM record consisting of nine MIIs
from the Guntersville Reservoir (Tennessee River) system in
Marshall County.
A number of studies involving specifically Alabama cave
crayfishes have been published. These include the descrip-
tions of new species (Rhoades 1941; Hobbs 1967b; J. Cooper
and Cooper 1997a, 1997b; Buhay and Crandall 2008, 2009);
studies on the biology and ecology of cave species (M. Cooper
1965; J. Cooper 1969; J. Cooper and Cooper 1977, 2010; Huryn
2007; Huryn et al. 2008; Spanjer and Cipollini 2006; Venarsky
et al. 2012); and studies on the evolution, history, and phylo-
geography of the Alabama cave crayfish fauna (Buhay 2006;
Buhay and Crandall 2005; Hobbs and Barr 1960, 1972).

Several studies have surveyed selected Alabama drain-
ages to document the crayfish fauna. These studies include
Yarbrough (1973), Tallapoosa and Chattahoochee Rivers;
Buchanan (1992), Cahaba River; Ratcliffe and De Vries (2004),
Tallapoosa River; Miller (2009), Little Choctawhatchee River;
Heath et al. (2010), Choctawhatchee, Pea, and Yellow Rivers;
Hopper et al. (2012), Sipsey River; and Helms et al. (2015), Tal-
lapoosa River. Surveys have also evaluated the current con-
servation status of certain target crayfish species (Stites et al.
2014; Kilburn et al. 2014).
Introduction 7
Although the history of crayfish studies in Alabama spans
many years, most of this work has been done in recent years.
This book may represent a true beginning to the full realiza-
tion of the wealth of crayfish diversity in Alabama. We hope
this book will stimulate more research, and we are certain
that the number of Alabama crayfish species will continue to
rise with more intensive surveys. We also hope this book will
stimulate not only more crayfish inventories, but also research
into the biology, life history, and ecology of the species living
in Alabama.
 2
Physiography
Physiography relates to the structure and type of under-
lying geologic formations and local geologic and climatic
forces that shape the landscape. Physiography, including the
underlying geology and shallow soils, is one of several deter-
minants of watershed boundaries in Alabama and, in many
cases, provides a template for its diverse and abundant flora
and fauna. The complex relationships of physiography and
climate can affect speciation, isolation of some species, and
extinction events. Within the framework of geologic time,
mountain chains have been created, uplifted, and eroded by
wind, rain, and temperature. Rivers have migrated, intersect-
ing and capturing other streams and changing direction, and
the coastline has been altered by changes in sea level during
the advance and retreat of continental ice sheets, with the
shoreline sometimes reaching as far inland as current Muscle
Shoals, Tuscaloosa, Montgomery, and Phenix City during the
highest sea levels. Historical geologic and climatic events were
intimately involved in creating, changing, and maintaining
Alabama’s biodiversity. The varied physiography and present
moderate climate sustain Alabama’s diversity across numer-
ous unique aquatic habitats, ranging from coastal marshes to
high-gradient, boulder-laden mountain streams.
Continents are divided into large physiographic provinces
based on relationships of geomorphology, or the topographic
features created by near-surface physical, chemical, or even
biological processes that give a region a unique character.
These provinces can in turn be subdivided into progressively
smaller physiographic sections, districts, and subdistricts (see
Wikipedia’s “Physiographic Province” entry, https://en.wiki-
pedia.org/wiki/Physiographic_province, accessed June 22,
2019). Alabama occupies parts of five physiographic prov-
inces, listed here generally from north to south: Interior Low
Plateaus, Appalachian Plateaus, Valley and Ridge, Piedmont,
and Coastal Plain. In Alabama these provinces are divided
into physiographic sections, and some sections are further
subdivided into physiographic districts and subdistricts
(Ebersole et al. 2019). A brief discussion of the physiographic
sections, districts, and subdistricts in Alabama follows.
Six major physiographic sections are recognized within
Alabama (Ebersole et al. 2019). They include (generally
arranged from north to south) the Central Basin, High-
land Rim, Cumberland Plateau, Alabama Valley and Ridge,
8
Piedmont Upland, and East Gulf Coastal Plain (fig. 2.1). Fur-
ther delineation of districts and subdistricts within sections
recognizes additional unique physical areas, illustrating
the broad and diverse physiographic character of Alabama.
Accounts for each physiographic section follow; the infor-
mation presented for each unit is derived from Ebersole et
al. (2019) unless otherwise noted. Summary information on
physiography is given in table 2.1. Additional information on
the geology of Alabama can be found in Harper (1920, 1942),
Johnston (1930), Lacefield (2013), Osborne et al. (1989), and
Szabo et al. (1988).
The Fall Line is a significant feature that influences the
distributions of many plants and animals in Alabama. It is
found at the maximum inland intrusion of ancient seas and
essentially marks the boundary where the older, harder, more
consolidated rocks of the interior or “upland” physiographic
provinces meet the younger, looser, less consolidated sedi-
ments of the East Gulf Coastal Plain, or “lowland.” In some
portions of the Fall Line there is a clearly defined demarca-
tion, such as where the Tallapoosa River traverses the Pied-
mont Upland and then abruptly meets the East Gulf Coastal
Plain in an area marked by steep waterfalls where the Pied-
mont rocks plunge beneath the coastal sediments (fig. 2.2). In
other areas there is a more gradual and often broad transition
zone, with the harder rocks of the interior commingling with
the looser sediments of the Coastal Plain before ultimately
disappearing beneath them. The latter phenomenon is espe-
cially evident in the Transition Hills in extreme northwest
Alabama. In this area, the East Gulf Coastal Plain, Cumber-
land Plateau, and Highland Rim sections are very close to one
another, and elements of all three can be found quite closely
spaced and often intricately interwoven.
CENTRAL BASIN
The Central Basin section in Alabama is a continuation
of the Outer Central Basin physiographic division of Ten-
nessee (Bingham and Helton 1999; Raisz and Mills 1962;
Starnes 2009) and is part of the Interior Low Plateaus
province, which extends from the Tennessee River valley
in northern Alabama northward to the glacial boundary in
Indiana. The extent of the section in Alabama lies within a
Figure 2.1. The physiography of Alabama. Districts
in each section are arranged descending from
north to south. Alluvial deposits, which parallel the
major river channels in the East Gulf Coastal Plain,
also occur in the Alabama Valley and Ridge section.
Compiled by Sandy Ebersole, Gregory M. Guthrie,
and Dane S. VanDervoort.
Table 2.1. Physiographic division within Alabama

Elevation Approx. %
Province Section District Subdistrict
(m) of state

Interior Low Plateaus Central Basin 1

Outer Central Basin 154–286

Highland Rim 7

Tennessee Valley 122–383

Little Mountain 119–289

Moulton Valley 138–354

Appalachian Plateaus Cumberland Plateau 16

Jackson County Mountains 156–573

Warrior Basin 29–431

Sequatchie Valley 119–431

Murphrees Valley 143–427

Sand Mountain 162–613

Blount Mountain 169–470

Wills Valley 157–483

Lookout Mountain 152–589

Valley and Ridge Alabama Valley and Ridge 8

Armuchee Ridges 163–505

Birmingham–Big Canoe 74–464

Cahaba Ridges 62–377

Cahaba Valley 55–379

Coosa Ridges 135–481

Coosa Valley 97–416

Weisner Frontal Ridges 98–650

Piedmont Piedmont Upland 9

Northern Piedmont 38–735

Central Uplands 38–735

Gainesville Ridges 103–324

Southern Piedmont 54–297

Greenville Slope 54–297

Pine Mountain 77–292

Coastal Plain East Gulf Coastal Plain 60

Fall Line Hills 26–335

Transition Hills 124–331

Black Prairie 21–173

Chunnenuggee Hills 18–209

Southern Red Hills 30–218

Buhrstone Hills 12–162

Flatwoods Lowland 15–119

Lime Hills 6–177

Hatchetigbee Dome 6–177


Province Section District
Dougherty Plain
Southern Pine Hills
Alluvial-Deltaic Plain
Deltaic Plain
Coastal Lowlands
Alluvial Plain*
*The Alluvial Plain is not a true district but rather consists of recent alluvial
deposits, usually along the coast but also along larger rivers inland.
Figure 2.2. Little Uchee Creek at Moffitts
Mill, Lee County, where rocks of the Pied-
mont Upland quickly plunge under the
loose sediments of the East Gulf Coastal
Plain. Photo by Jim Lacefield.
single district, the Outer Central Basin district. It manifests
as relatively narrow, fingerlike outcrops of limestone along
the Elk River and a small portion of upper Limestone Creek
(Limestone County) and Shoal Creek (Lauderdale County)
and some of their tributaries. These outcrops enter the state
from Tennessee, plunge relatively quickly beneath the Ten-
nessee Valley district, and represent a minor portion of the
state’s area, just over 1%.
HIGHLAND RIM
The Highland Rim section of north-central and northwest
Alabama is part of the Interior Low Plateaus province and
reaches from the Alabama-Tennessee state line south to the
northern slope of the Cumberland Plateau section. Valley
floors are composed predominantly of level limestone beds,
and ridges are typically capped by sandstone. It represents
about 7% of the state’s area and is drained exclusively by the
Tennessee River. Three districts compose the Highland Rim
section—the Tennessee Valley, Little Mountain, and Moulton
Physiography 11
Elevation Approx. %
Subdistrict
(m) of state
23–168
< 5–159
< 3–35
< 3–28
0–53
1–237
Valley—all sloping to the west and plunging beneath Coastal
Plain deposits at a rate of about 9.5 meters per kilometer.
The Tennessee Valley district is the largest district within
the Highland Rim section in Alabama, extending from the
foot of the Little Mountain district north to the Alabama-Ten-
nessee state line. The level parts of the valley include numer-
ous karst features such as springs, ponds, lime sinks, and
caves formed by solution of the underlying limestone. Water
discharged from springs often sustains stream flows during
droughts, and some large springs, such as Brahan Spring in
Huntsville and Tuscumbia Spring in Tuscumbia, have long
served as municipal drinking water supplies.
The Little Mountain district is a series of low sandstone-
capped ridges with lower elevations underlain by limestones
(fig. 2.3). Several connected ridges make up the collective Lit-
tle Mountain district, which is a cuesta, or broad, often saw-
toothed ridge with a steep, north-facing slope and a gentle
slope on the southern face because of the dip of the under-
lying rocks. The Moulton Valley district is a relatively long,
narrow, flat valley with a limestone base. It lies to the south of
12 Chapter 2
and parallel to the Little Mountain district and just north of
the Cumberland Plateau section.
CUMBERLAND PLATEAU
The Cumberland Plateau section is part of the greater Appa-
lachian Plateaus province, which extends from central Ala-
bama northeast to eastern Kentucky. It is composed of an
undulating surface of sandstone and shale that is frequently
dissected by often steep-sided valleys, hollows, and coves,
and it represents about 16% of the state’s area. Eight districts
compose the Cumberland Plateau section in Alabama: the
Jackson County Mountains, Warrior Basin, Sand Mountain,
Sequatchie Valley, Blount Mountain, Murphrees Valley, Wills
Valley, and Lookout Mountain.
The Jackson County Mountains district is a plateau of
high relief, with level mountaintops capped by sandstones
with underlying limestones, and stream valleys cut through
limestone beds. This is a heavily karstic area because of the
limestones. The Warrior Basin district is the largest district
within the Cumberland Plateau section in Alabama and is a
broad, dissected plateau of sandstone and shale. Flow in larger
streams is usually sustained during summer months, but
many headwater tributaries go dry because of low recharge
from groundwater aquifers. The Sequatchie Valley district
is a pronounced topographic feature, expressed as a long,
level valley extending from Cumberland County, Tennessee,
southwest to Blount County, Alabama. The Murphrees Val-
ley district is a relatively small, tripartite valley lying on bed-
rock composed of limestone, sandstone, and shale. The Sand
Mountain district is a large sandstone-capped plateau with
underlying shale gradually merging with the Warrior Basin
district in the southwest and the Blount Mountain district in
the southeast. The Blount Mountain district is a remnant of
the Cumberland Plateau underlain by sandstones and shales.
Figure 2.3. Waterfall Creek in Colbert
County, plunging off the sandstone cap
of the Little Mountain district. Photo by
Jim Lacefield.
The Wills Valley district in Alabama is composed of three
limestone valleys separated by sandstone ridges of low relief.
The Lookout Mountain district is a flat, narrow remnant of
the Cumberland Plateau extending from near Chattanooga,
Tennessee, to Etowah County, Alabama (fig. 2.4). Lookout
Mountain is composed of sandstones and shales and is very
similar to Sand Mountain in topography and character.
ALABAMA VALLEY AND RIDGE
The Alabama Valley and Ridge section is the southernmost
section of the Valley and Ridge province, which ranges from
Alabama northeast to southern New England. The section is
composed of a series of folded and faulted parallel ridges and
valleys that trend northeast–southwest, and it represents about
8% of the state’s area. Seven districts are recognized within the
Alabama Valley and Ridge section: the Armuchee Ridges, Bir-
mingham–Big Canoe Valley, Cahaba Ridges, Cahaba Valley,
Coosa Ridges, Coosa Valley, and Weisner Frontal Ridges.
The Armuchee Ridges district consists of a small cluster
of northeast-trending chert and sandstone ridges and inter-
vening lowlands with limestone beds. The Birmingham–Big
Canoe Valley district is developed on soluble limestones and
dolomites and less soluble shales, sandstones, and chert. Many
springs originate in the carbonaceous terrain of the Birming-
ham–Big Canoe Valley, yielding unique aquatic habitats. It
is extensively developed and receives significant non-point-
source pollution such as nutrients and sediments from the
Birmingham metropolitan area.
The Cahaba Ridges district lies predominantly between
the Cahaba Valley and the Birmingham–Big Canoe Valley
districts and abuts the Fall Line Hills district at its southwest
terminus. It is characterized by several parallel ridges that
trend northeast–southwest, formed by massive sandstone and
conglomerate beds.

Figure 2.4. View from Sand Mountain
looking southeast to Lookout Mountain,
DeKalb County. Photo by Jim Lacefield.
Figure 2.5. Near-vertical rocks of the Coosa
Valley district along the Coosa River down-
stream of Weiss Dam, Cherokee County.
Photo by Jim Lacefield.
The Cahaba Valley district lies predominantly between
the Cahaba Ridges and Coosa Ridges districts, with a small
portion along the Coosa Valley and Weisner Frontal Ridges
districts at its northeast extent. At its southwest terminus it
commingles with the Fall Line Hills district. It is developed on
weathered, soluble limestones and dolomites, erodible shales,
and resistant chert beds.
The Coosa Ridges district consists of a folded parallel
ridge belt extending northeast–southwest and characterized
by parallel, linear sandstone ridges separated by shale valleys
(fig. 2.5). The Coosa Valley district, a plain of shale and lime-
stone with ridges of low relief, represents over half the total
area of the Alabama Valley and Ridge section.
The Weisner Frontal Ridges district consists of a series
of maturely dissected mountains of extreme relief with
intervening narrow carbonate valleys adjacent to the Valley
and Ridge section. It represents the westernmost extent of
metasedimentary rocks in Alabama.
Physiography 13
PIEDMONT UPLAND
The Piedmont Upland section, a part of the Piedmont prov-
ince of the Appalachian Highlands physiographic region, is
the nonmountainous section of the “older Appalachians,” as
described by Fenneman (1938). The Piedmont developed on
belts of metamorphic rocks trending northeast–southwest
and is the result of long-term degradation of the surface
rocks while the underlying rocks were severely deformed
and angled to the surface. The Piedmont Upland section in
Alabama is a wedge-shaped feature bounded to the south by
sediments of the East Gulf Coastal Plain, to the northwest by
the Alabama Valley and Ridge section, and to the east by the
Alabama-Georgia state line; it covers about 9% of the state.
The Piedmont Upland section is divided into two districts:
the Northern Piedmont and Southern Piedmont.
The Northern Piedmont district is a well-dissected upland
forming a prominent ridge system trending northeast–southwest
14 Chapter 2
and including Cheaha Mountain, the highest point in the state at
735 m. Numerous small springs and clear upland streams occur
throughout the Northern Piedmont district, providing unique
aquatic habitats. The Northern Piedmont district is further
subdivided into two subdistricts: the Central Uplands and the
Gainesville Ridges.
The Central Uplands subdistrict corresponds to the same
district in Georgia (Clark and Zisa 1976), including highly
deformed sedimentary and igneous rocks. Cheaha Mountain
is in this subdistrict. The Gainesville Ridges subdistrict con-
stitutes the southern extent of the Northern Piedmont dis-
trict and is underlain by quartzite gneiss and sheared igneous
rocks on ridges trending northeast–southwest.
The Southern Piedmont district consists of a rather flat
surface with little topographic relief. Some of the more inter-
esting aquatic habitats in the district are located along the Fall
Line, where streams cut through the edge of the Piedmont
and enter the East Gulf Coastal Plain, resulting in a series of
rapids or spectacular falls over the basal Piedmont rocks. The
Southern Piedmont district has two subdistricts: the Green-
ville Slope and Pine Mountain.
The Greenville Slope subdistrict is a continuation of the
Greenville Slope Piedmont district of Georgia (Clark and
Zisa 1976). It consists of high-grade igneous and sedimen-
tary rocks and is characterized by broad, flat ridges. The Pine
Mountain subdistrict lies south of the Greenville Slope sub-
district and is a continuation of the Pine Mountain district
of Georgia (Clark and Zisa 1976), but lower in elevation.
The subdistrict is composed of ridges of metamorphic rocks
trending northeast–southwest and lying along the Fall Line,
with prominent ridges of quartzite throughout (Sapp and
Emplaincourt 1975).
EAST GULF COASTAL PLAIN
The East Gulf Coastal Plain section in Alabama is part of the
Coastal Plain province. It is characterized by gently rolling
hills, sharp ridges, prairies, and broad alluvial floodplains. In
Alabama it extends from the extreme northwest in a gently
curving arc to the southeast to the Chattahoochee River and
southward. Rocks underlying the Coastal Plain are sedimen-
tary in origin and consist of sand, gravel, porous limestone,
chalk, marl, and clay. These strata dip to the southwest at
approximately seven to eight meters per kilometer and strike
generally in east–west belts. Some strata are more resistant
to erosion and underlie broad, saw-toothed ridges known
as cuestas that slope gently to the south and have steep
north-facing slopes. The East Gulf Coastal Plain section rep-
resents about 60% of the state’s land area and occurs south
and southwest of the Fall Line. Ten physiographic districts
are recognized in the East Gulf Coastal Plain section: the
Fall Line Hills, Black Prairie, Chunnenuggee Hills, Southern
Red Hills, Lime Hills, Dougherty Plain, Southern Pine Hills,
Alluvial-Deltaic Plain, Deltaic Plain, and Coastal Lowlands,
with some further subdivided into subdistricts. Much of the
section, especially south of the Fall Line Hills, has little to no
discernible slope, leading to often sluggish streams and exten-
sive areas of swamps, bogs, marshes, and vernal pools.
The Fall Line Hills district in Alabama is a wide, cres-
cent-shaped band extending from extreme northwest Ala-
bama near the Mississippi and Tennessee state lines to the
southeast across central Alabama to the Chattahoochee River
in eastern Alabama. The Fall Line Hills form a significant
boundary between the Highland Rim, Cumberland Plateau,
Alabama Valley and Ridge, and Piedmont Upland sections
and the lower Coastal Plain; the contact zone, while not
firmly recognizable, is generally referred to as the aforemen-
tioned Fall Line. Streams draining the Fall Line Hills district
are well sustained, even during the driest years, because of
extensive sand and gravel aquifers (Geological Survey of Ala-
bama 2018). Topography can be rugged and includes steep
slopes, particularly near streams. In its western portion, the
Fall Line Hills district exists as an irregularly shaped transi-
tion belt (the Transition Hills subdistrict), where rocks of the
Cumberland Plateau, Highland Rim, and East Gulf Coastal
Plain sections commingle. This contrasts with the eastern
portion of the district, where the Fall Line marks a very sharp
transition between unconsolidated Coastal Plain sediments
and harder Piedmont Upland rocks. The Fall Line essen-
tially divides Alabama into two distinct physical regions, the
upland and lowland, and is generally considered the most
significant physical feature in Alabama affecting the distribu-
tions of aquatic species. It is the zone of contact between the
harder rocks of the Appalachians and Interior Plateau with
the looser, unconsolidated sediments of the Coastal Plain.
Because of limitations due to life history traits, morpholog-
ical adaptations, or lack of invasion routes, many species are
limited to either above (upstream of) or below (downstream
of) the Fall Line, or nearly so, while other species are unre-
stricted by this barrier. Above the Fall Line, streams generally
flow swiftly because of their higher gradient, reflecting the
influence of harder base rocks, steeper slopes, and higher ele-
vations, whereas streams below the Fall Line generally flow
more slowly and have muddy or sandy substrates and a lower
gradient, indicating flatter topography. In this zone, streams
descend from resistant sedimentary and crystalline rocks to
the less resistant sands and clays of the Coastal Plain.
The Transition Hills subdistrict of the Fall Line Hills dis-
trict occupies extreme northwest Alabama. For many years,
researchers included it in the greater Fall Line Hills district,
but differences in topography and geology have led to its rec-
ognition as a distinct subdistrict. The Transition Hills sub-
district includes extensive upland areas of sands and gravels,
with valleys of limestone.
The Black Prairie district is an undulating, deeply weath-
ered plain developed primarily on chalk and marl and is
characterized by concentrations of soft, white-gray limestone
(fig. 2.6). Because of thin soils and impermeable rocks, the
Black Prairie represents a unique and clearly defined hydro-
logic region in the state. Many streams have eroded to chalk

Figure 2.6. Chalk outcrop along the Tom-
bigbee River in the Black Prairie district,
Sumter County. Photo by Jim Lacefield.
Figure 2.7. Cedar Creek, Butler County,
in the Southern Red Hills district. This
stream displays the typical slow-flowing,
sandy-bottomed nature of many streams
on the Coastal Plain. Photo by Jim
Lacefield.
bedrock and are noted for high rates of runoff and highly
variable flows during storm events. Small streams frequently
go dry while flow in larger streams is significantly reduced.
Harper (1943) noted that the Black Prairie is one of the driest
regions in the state and that natural prairie grasslands once
covered up to 10% of the area. The Black Prairie is a significant
distributional barrier for some aquatic species. The Chun-
nenuggee Hills district consists of a series of pine-forested
sand hills developed on hardened beds of chalk in its western
reach and on clay, sandstone, siltstone, and chalk farther east
(Sapp and Emplaincourt 1975). The rocks in this district are
somewhat more permeable than the chalk in the Black Prairie
district but are still relatively poor aquifers.
The Southern Red Hills district is a southward-sloping
upland of moderate relief in southwest Alabama (fig. 2.7).
Two subdistricts are found in the Southern Red Hills district:
the Buhrstone Hills and Flatwoods Lowland.
The Buhrstone Hills subdistrict is located along the south-
ern edge of the Southern Red Hills proper and is characterized
Physiography 15
by very rugged terrain developed on indurated, resistant sili-
ceous claystone and sandstone. The greatest topographic relief
on the Coastal Plain of Alabama occurs in the Buhrstone
Hills, where ridges locally rise 90 to 120 m above stream val-
leys. Streams in this area acquire upland characteristics, with
high gradients, hard-rock bottoms, and swifter flows. The Flat-
woods Lowland subdistrict is nestled between the Southern
Red Hills district proper and the Chunnenuggee Hills district.
It developed on dark clays and marls that weather to stiff,
plastic soils. Few ponds and springs are found in the area, and
potable groundwater is difficult to find. As its name suggests,
the rugged Lime Hills district occurs over resistant limestones.
In this area interesting habitats develop where streams cut
through alluvial sediments down to limestone features, result-
ing in streambeds with upland character as in the Buhrstone
Hills subdistrict. The Lime Hills district has one subdistrict,
the Hatchetigbee Dome subdistrict. The Hatchetigbee Dome
subdistrict is an anticline (flexure) that is oriented north-
west–southeast and fault bounded on its northern border by
16 Chapter 2
the southeastern end of the anticline merging with the Jack-
son fault (Raymond et al. 1988). Mineral springs occur in the
Hatchetigbee area, and some are saline because of salt domes
near the surface.
The Dougherty Plain district, or “wiregrass region,” is an
extension of an upland in Georgia and is underlain by lime-
stone, bedded sand and clay, and terrace materials (Ebersole
et al. 2019). Topography consists of low cuestas (Sapp and
Emplaincourt 1975), which are more dissected in south-central
Alabama than farther southeast. Karst topography is found in
some parts of the district because of underlying limestone,
and as a result many minor drainages in the district go under-
ground, especially in extreme southeastern Alabama. Active
solution of the underlying limestone produces many shallow,
flat-bottomed depressions, and small headwater streams are
noticeably absent from the Dougherty Plain as a result. The
name “wiregrass” originates from the common occurrence
of needlerush in the wet, shallow depressions. The Southern
Pine Hills district is an area of low relief with broad, rounded
ridges and V-shaped valleys with sand and clay sediments.
This region is not subject to solution like the Dougherty Plain,
and the boundary between the two districts is sometimes a
distinct escarpment. Flat uplands with shallow ponds, bogs,
and marshes occur throughout the district, and many of
the valleys are saucerlike and perpetually wetted by seepage
from nearby hills. The abundance of warm summer rains is
a major factor in leaching fertility from the soil and favoring
the growth of pines in this region. Streams are well sustained
Figure 2.8. The Mobile-Tensaw Delta
from the I-65 bridge, Baldwin and Mobile
Counties. In this vicinity the Mobile River
distributaries begin to separate. Photo by
Jim Lacefield.
by groundwater flows in summer and are commonly called
“blackwater” creeks, particularly those originating in the
Pine Hills proper, where the term refers to the natural color
imparted by dissolved and suspended organic matter. Younger
terrace deposits occur along major streams. The Coastal Low-
lands district is a flat to gently undulating plain that is often
locally swampy and is continuously altered by erosion and
deposition of sediments. The majority of the plain is indented
by numerous tidal streams and fringed by tidal marshes and
offshore barrier islands. The landward edge of the district is
defined by the base of the Pamlico marine scarp (Sapp and
Emplaincourt 1975). The Alluvial-Deltaic Plain district has
characteristics reminiscent of both freshwater alluvial depos-
its and marine deposits resembling the adjacent Deltaic Plain
district. This district includes lower elevations of floodplains
of the Alabama and Tombigbee River systems north of the
Deltaic Plain district. Because of the low elevation, this area
resides in hurricane surge zones (Zachry et al. 2015).
The Deltaic Plain district is the relatively flat, low-eleva-
tion floodplain of the Mobile-Tensaw Delta. At the northern
end of the district the Tensaw River (a distributary of the
Mobile River) separates and drains into Mobile Bay (fig. 2.8).
The Alluvial Plain is not technically a physiographic dis-
trict but is chiefly alluvium (sand, clay, and gravel) of river
and stream valleys. Because alluvium is found throughout
the state, elevations range significantly, from less than 1 m to
237 m, with lowest elevations generally closer to the coast and
higher elevations in the north and east.
3
Inland Waters of Alabama
Because of Alabama’s latitude (approximately 30°–35° north
latitude) and proximity to the Gulf of Mexico, its climate is
considered humid subtropical, with year-round precipitation
(Chaney 2013). Summers are dominated by warm, humid air
from the Gulf of Mexico, leading to frequent thunderstorms,
and air temperatures often reach daytime highs over 30°C
(90°F) and nighttime lows near 20°C (70°F). Winters are gen-
erally mild, with alternating cold fronts bringing precipitation
events interspersed with clear days, and average low tempera-
tures in January ranging from near 0°C (30°F) in the north
to near 4°C (40°F) in the south. Rainfall in Alabama averages
127–137 cm per year in the north and 152–165 cm per year in
the south.
Runoff of precipitation follows surface and groundwater
courses, with topography, soil type, and underlying geology
influencing discharge and retention rates. A recent assessment
using spatial technology documented that approximately
214,000 km of surface streams traverse the state, with about
61% permanently flowing and 39% flowing intermittently
during or after precipitation events. According to the South-
east Aquatic Resources Partnership (SARP), 17,424 dams have
been inventoried in Alabama, from small mill dams to large
run-of-the-river lock and dam structures, representing 12%
of the total for the southeastern United States and leading
to many disconnected river and stream reaches (Southeast
Aquatic Resources Partnership 2017). Gangloff et al. (2009)
reported that most of the small mill dams, especially on the
Coastal Plain, no longer exist.
Alabama leads the nation in navigable river length at 2,314
km, with lock and dam structures built on every major river to
facilitate navigation, and with frequent dredging to maintain
channel depths sufficient for navigation by commercial tows
(Geological Survey of Alabama 2018). Surface area of lakes,
ponds, and reservoirs totals 227,838 ha, with about 1,456,868
ha of freshwater and saline wetlands associated with Alabama
waters (Geological Survey of Alabama 2018).
MAJOR RIVER BASINS
All rivers in Alabama eventually drain into the Gulf of Mex-
ico via one of several conduits. The state is divided into five
major river basins and again into numerous subbasins (fig.
3.1). These major basins and subbasins are briefly described
here, arranged generally from north to south.
The Tennessee River is a major tributary of the Ohio River
basin of the greater Mississippi River basin, ultimately enter-
ing the Gulf of Mexico southeast of New Orleans, Louisiana.
Headwaters of the Tennessee River originate in the southern
Appalachian Mountains in North Carolina, Tennessee, and
Virginia, and the river enters Alabama in the extreme north-
east corner of the state in the Sequatchie Valley district of
the Cumberland Plateau section. From there the river flows
southwest until it breaches the Jackson County Mountains
district and meanders westward across the Tennessee Valley
district before exiting the state in the Transition Hills subdis-
trict of the Fall Line Hills district in the extreme northwest
corner of the state. It also drains a small portion of the War-
rior Basin district of the Cumberland Plateau section (fig. 2.1).
In Alabama, it drains about 17,612 km2, or about 13% of the
state’s area (Geological Survey of Alabama 2018). A series of
large run-of-the-river lock and dam structures on the Ten-
nessee River are operated by the Tennessee Valley Authority
(TVA) for navigation and power production and impound
78,995 ha in the state (fig. 3.2).
The headwaters of the Mobile River basin are found in the
Valley and Ridge section in northwest Georgia and south-
east Tennessee and in the East Gulf Coastal Plain section in
northeast Mississippi. In Alabama, it drains all or significant
parts of the Cumberland Plateau, Alabama Valley and Ridge,
Piedmont Upland, and East Gulf Coastal Plain sections (fig.
2.1). The Mobile River basin drains about 113,130 km2, includ-
ing 84,874 km2 in Alabama, or about 61% of the state’s area. It
has a greater yield of water per square kilometer of land than
any other basin in the United States, including the Mississippi
River (O’Neil 2013). Two major subbasins within the Mobile
River basin are the Tombigbee River system and the Alabama
River system. The Alabama and Tombigbee Rivers join to
form the Mobile River shortly before entering Mobile Bay via
the Mobile-Tensaw Delta. Other, smaller tributaries that enter
Mobile Bay without passing through the Mobile River are not
included in drainage totals here.
The western part of the Mobile River basin includes
the Black Warrior and Tombigbee River systems, and
about 33,184 ha of impoundments for hydroelectric power
17
Figure 3.1. Major river basins, drainages, and systems in Alabama.

Figure 3.2. Wheeler Dam, Tennessee
River, Lawrence County, Alabama.
Figure 3.3. Mitchell Dam, Coosa River,
Chilton County, Alabama.
Figure 3.4. West Point Dam, Chatta-
hoochee River, Chambers County,
Alabama.
generation, navigation, or flood control. The eastern part of
the basin includes the Alabama, Cahaba, Coosa, and Tall-
apoosa River systems and is heavily impounded with hydro-
electric and flood control reservoirs, with about 68,800 ha
of impoundments (fig. 3.3). The preponderance of large-vol-
ume reservoirs in the eastern part of the drainage can be
attributed to topography and geology. The western part of
the basin lies mainly on the loose sands and gravels of the
East Gulf Coastal Plain section and is of relatively low relief,
whereas the eastern part drains much of the Alabama Valley
and Ridge section, with more stable underlying rocks and
Inland Waters of Alabama 19
steeper topography making it more conducive to dam con-
struction (fig. 2.1).
Often lumped together as the “coastal drainages,” several
smaller river systems that flow into the Gulf of Mexico through
various water bodies drain the remaining approximately 26%
of the state. These include, from west to east, the Escatawpa
River of the Pascagoula River basin (Mississippi Sound), the
Perdido River (Perdido Bay), the Conecuh, Blackwater, and
Yellow Rivers (Pensacola Bay), the Choctawhatchee-Pea River
system (Choctawhatchee Bay), and the Chipola and Chatta-
hoochee Rivers (Apalachicola Bay) (fig. 3.4). All these systems
20 Chapter 3
lie entirely within the East Gulf Coastal Plain section except
for the Chattahoochee River system, whose headwaters are in
the Blue Ridge and Piedmont Upland sections in northern
Georgia (Geological Survey of Alabama 2018; US Geological
Survey, n.d.).
The Escatawpa River drains a small portion of extreme
southwest Alabama along the Alabama-Mississippi state line
from Choctaw County south to Mobile County. The Perdido
River drains parts of Baldwin and Escambia Counties and
forms the state line between Alabama and the western margin
of the Florida Panhandle. The Conecuh River system is the
largest of the coastal drainages and drains much of south-cen-
tral Alabama from Bullock County southwest to Escambia
County, where it enters Florida and becomes known as the
Escambia River. The Blackwater and Yellow River systems are
relatively small drainages, nestled between the much larger
Conecuh and Choctawhatchee River systems. The Choc-
tawhatchee River system is relatively large and drains much of
extreme southeast Alabama. The Chattahoochee River enters
Alabama in Chambers County in the Piedmont Upland sec-
tion; it forms much of the boundary between Alabama and
Georgia and is a tributary of the Apalachicola River and
ultimately Apalachicola Bay (Geological Survey of Alabama
2018). The headwaters of the Chipola River system are found
in extreme southeast Alabama, nestled between the Choc-
tawhatchee and Chattahoochee River systems, and drain into
the Apalachicola River and Apalachicola Bay.
MAJOR AQUIFERS
Of the surface water in Alabama, about 40% by volume
and 70% by geographic area originates from groundwater
sources (Geological Survey of Alabama 2018). Groundwa-
ter is widely available, especially in the southern half of
the state, because of the loose, unconsolidated sands and
gravels on the Coastal Plain. In the northern section of the
state, where Paleozoic carbonaceous rocks dominate and
numerous karstic springs occur, surface waters are more
extensively used as public or domestic water supply. Almost
all water used for “beneficial purposes” on the East Gulf
Coastal Plain originates from groundwater through pri-
mary porosity of geologic formations. In bedrock aquifers
above the Fall Line, water originates primarily in fractures,
joints, bedding planes, and shallow regolith (Geological
Survey of Alabama 2018).
Alabama has several major aquifers that are replenished
from recharge areas covering most of the state’s land surface.
Recharge areas are the basis for groundwater and surface-wa-
ter interactions and provide surface-water base flow that
supports stream discharge and reservoir levels, wetlands, and
aquatic habitats throughout the state. Based on the geologic
map of Alabama (Osborne et al. 1989), J. Moore (1998) rec-
ognized and mapped the significant aquifers in the state and
noted that major water-bearing geologic units underlying the
state vary in age from the geologically ancient Ordovician
system to the recent alluvial, coastal, and terrace deposits
of the Quaternary system. These geologic units vary signifi-
cantly in their water-bearing capacity, aquifer characteristics,
and rates of recharge. Groundwater use and development is
directly related to these characteristics as well as the econom-
ics of drilling wells to exploit the resource. Detailed infor-
mation on and maps of the groundwater resources of the
state can be found in a report by the Geological Survey of
Alabama (2018).
4
Crayfish Distributions and
Habitat Associations
GENERAL DISTRIBUTIONS
Given the warm, wet climate, diverse physiography, and abun-
dant hydrologic resources in Alabama, the state is home to a
correspondingly rich assortment of crayfish species, with 99
species currently recognized. Geologic and hydrologic barriers
often dictate the distributions of aquatic species within Ala-
bama, and perhaps the most significant feature is the Fall Line.
Evidence that the Fall Line creates a boundary to crayfish
distributions is apparent when comparing the distributions
of taxa found above the Fall Line versus below the Fall Line
as well as those that appear not to be affected by its presence.
Even at the generic level, the Fall Line influences the distri-
butions of members that share habitat patterns. The single
Barbicambarus species in Alabama is exclusively lotic that
dwells under large slab rocks in streams with swiftly flowing
water and is found only above the Fall Line. Faxonella and
Hobbseus, composed largely of burrowing species, and Cam-
barellus, whose species usually occupy bogs and other wet-
lands, in Alabama are restricted to the Coastal Plain. There
they take advantage of the loose sediments and generally flat
topography that readily provide necessary habitats. The two
members of the genus Orconectes in Alabama are restricted
to caves associated with limestone formations and are also
restricted to the abundant karst formations above the Fall
Line. Members of the four remaining genera known from
Alabama are found both above and below the Fall Line, but
each genus is inclined toward one or the other. Cambarus, a
very diverse genus made up largely of lotic species but with
some lentic and burrowing species, along with Faxonius, a less
diverse group whose members are also prone to occupying
lotic systems, are more focused above the Fall Line. Cambarel-
lus, Creaserinus, and Procambarus show just the opposite ten-
dency. Many of their members tend to be lentic or burrowing
species but often tolerate lotic environments, and members of
all three genera are concentrated below the Fall Line.
In Alabama, 45 species occur almost exclusively above the
Fall Line and 41 species almost exclusively below it; only 13
species have broad distributions both above and below (table
4.1). These distributional differences may reflect geologically
ancient dispersal patterns and the diversity and distribution
of habitats. The older interior rocks have had more time to
change and consolidate, leading to more opportunities for
speciation while at the same time providing a wider physio-
graphic diversity. In the Coastal Plain the younger, less stable
sediments have permitted fewer opportunities and less time
for speciation. In a few cases species have been moved to new
physiographic sections in Alabama by human means and
have prospered (e.g., Faxonius lancifer, F. palmeri palmeri, F.
virilis, Procambarus clarkii) (Schuster et al. 2008; Adams et al.
2015; McGregor et al. 2018).
As is the case with freshwater mussels and fishes (when
including marine and estuarine fishes), Alabama is the most
crayfish species–rich state in the country (Mettee et al. 1996;
Boschung and Mayden 2004; J. Williams et al. 2008). Ala-
bama’s diverse and ancient geology allowed sufficient time
and opportunity for speciation among crayfishes, as well
as ample opportunity for endemism. Currently there are
15 crayfish species endemic to Alabama, with 8 endemic to
the Tennessee River system and 7 to the Mobile River basin
(table 4.2). The Tennessee River system, again mimicking
the richness of mussel and fish species, is the most spe-
cies-rich basin in the state (table 4.3). The Tennessee River
basin in Alabama is home to 47 crayfish species, represent-
ing almost half the state’s list, despite draining only 13% of
the state’s area. Furthermore, 27 species (28% of the state’s
crayfish fauna) are endemic to the basin. The Mobile River
basin drains 61% of the state and hosts 58 species, with 20
species (21%) endemic to that basin. The collective Coastal
Plain drainages found to the east and south of Mobile Bay
and the Mobile River basin drain about 22% of the state’s
area and host 29 species, with 8 species endemic there.
Coastal Plain tributaries in Alabama from Mobile Bay and
the Mobile-Tensaw Delta west, including Mississippi Sound
tributaries as denoted in table 4.3 (Escatawpa River/Mis-
sissippi Sound/Mobile Bay), are generally represented by a
crayfish fauna somewhat distinct from that of other Coastal
Plain drainages to the east and are therefore addressed sep-
arately here. This area hosts 25 species, with 3 endemic spe-
cies, and drains 4% of the state.
21
Table 4.1. Species found either above or below of the Fall Line, or not restricted by the Fall Line

Above the Fall Line (45)

Barbicambarus simmonsi Cambarus jonesi Faxonius compressus

Cambarus acanthura Cambarus laconensis Faxonius cooperi

Cambarus andersoni Cambarus lentiginosus Faxonius durelli

Cambarus bartonii cavatus Cambarus longirostris Faxonius erichsonianus

Cambarus clairitae Cambarus manningi Faxonius forceps

Cambarus cracens Cambarus obstipus Faxonius juvenilis

Cambarus distans Cambarus parvoculus Faxonius mirus

Cambarus diupalma Cambarus pecki Faxonius palmeri palmeri

Cambarus englishi Cambarus rusticiformis Faxonius placidus

Cambarus gentryi Cambarus scotti Faxonius spinosus

Cambarus girardianus Cambarus speleocoopi Faxonius validus

Cambarus graysoni Cambarus tenebrosus Faxonius virilis

Cambarus halli Cambarus unestami Faxonius yanahlindus

Cambarus hamulatus Cambarus veitchorum Orconectes australis

Cambarus howardi Faxonius alabamensis Orconectes sheltae

Below the Fall Line (41)

Cambarellus diminutus Lacunicambarus freudensteini Procambarus lagniappe

Cambarellus lesliei Lacunicambarus ludovicianus Procambarus lecontei

Cambarellus rotatus Lacunicambarus miltus Procambarus lewisi

Cambarellus shufeldtii Lacunicambarus mobilensis Procambarus marthae

Cambarus pyronotus Procambarus bivittatus Procambarus okaloosae

Creaserinus burrisi Procambarus capillatus Procambarus paeninsulanus

Creaserinus byersi Procambarus clemmeri Procambarus planirostris

Creaserinus danielae Procambarus escambiensis Procambarus shermani

Faxonella clypeata Procambarus evermanni Procambarus suttkusi

Faxonius holti Procambarus hagenianus hagenianus Procambarus verrucosus

Faxonius jonesi Procambarus hayi Procambarus versutus

Hobbseus prominens Procambarus holifieldi Procambarus vioscai paynei

Lacunicambarus dalyae Procambarus hubbelli Procambarus zonangulus

Lacunicambarus erythrodactylus Procambarus hybus

Not Restricted by Fall Line (13)

Cambarus coosae Faxonius lancifer Procambarus lophotus

Cambarus latimanus Faxonius perfectus Procambarus spiculifer

Cambarus striatus Procambarus acutissimus Procambarus viaeviridis

Creaserinus fodiens Procambarus acutus

Faxonius etnieri species complex Procambarus clarkii

Another random document with
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 CHARITY UNDERTOOK TO DIVIDE EVERYTHING WITH EQUAL
FAIRNESS.

Up to now they had seen very few people. A boy driving

cattle in the distance was an Indian chasing buffaloes; an
old man with a dog was a chief with his wolf hound.

But when they took to their raft again, their sharp eyes
spied a fisherman some distance away with his line across
the stream.
 "Now you'll hear him swear," said Charlie, with a
delighted chuckle. "I know what fishermen are like. I've
passed them before."

"He isn't a fisherman," said Hope; "he's a pirate looking

out for ships from his island."

"It's a pity we have no gun to blow him to pieces! My

dear husband, put a bit more strength into that old punt
stick of yours! Let's rush down the stream and pull hold of
his line—perchance we may pull in the pirate to his death!"

So all four got hold of their oars, and by dint of

prodding the banks in punt-like fashion, the raft began to
quicken its pace. The fisherman saw them before they
reached him and pulled in his line. He did not swear but
laughed heartily as the raft approached.

"Who the dickens is this?" he asked. "Here, hi, don't

pass me by! Is there room for me on board?"

He was a tall, broad-shouldered young man.

Charlie threw a ferocious look at him.

"You're a pirate, let us pass!"

The young man had stopped the raft with his foot.
Charlie was rather exhausted with his efforts, and the little
girls were panting for breath.

"If I'm a pirate, I beg to tell you that this water is mine,
and that you are my prisoners. You'll land at once, and
forfeit your ship."

With a quick, dexterous stroke, he had seized hold of

the rope, and drawn the raft close to the bank.
 Winding it round a tree stump, the discomfited voyagers
found their passage stopped.

Charity looked up into the pirate's face. She saw that

his eyes were twinkling, and she felt reassured.

"The ocean is free to all," she said boldly.

The young man pointed to a board close by.

"Private water. Trespassers will be prosecuted."

And then Charlie saw that he had brought his raft along
the wrong bend of the stream. He had seldom before come
as far as this.

For a moment the little Captain looked perplexed.

"We are on a peace voyage," he said, "otherwise with

cutlasses and guns we would make short work of you. We
are searching for an island called 'Tarjak,' and for treasure
thereon."

"Ah," said the young man in a mysterious tone, "now

you've come to the right party. I know the very spot and
will lead you to it, if I may share in the booty."

"He means treachery!" said Charity in a loud whisper.

"I've got you in my power!" said the young man sternly.

Charlie and the little girls hastily consulted together.

"We'll just let him join us," was Charlie's decision.

Then he turned to the stranger.

"Now will you lead us?"

 "I'll tow you along," was the cheerful reply. "You have
spoilt my fishing, but I'll take you straight to a treasure
island such as you have never seen before."

In a very short time, this amazing young man was

marching along the banks rope in hand, and the raft was
being towed along without any effort on the part of the
crew. They gave themselves up to the delight of it, for all
small backs and wrists were aching, and it was delicious to
be towed along so swiftly, without any effort.

And presently the stream widened considerably and a

veritable small island appeared. The Pirate used the oars
now, and stood in the middle of the raft himself. He brought
it to the edge of the island and told the children to get out.

To their delight there was a tiny thatched hut in the

middle of it.

"Years ago," said the pirate, "I landed here in a small

boat, having been shipwrecked, and having lost all I
possessed. I built myself this hut and lived here in peace,
quitting it for rougher waters and sea fishing occasionally.
For the most part I lived on fish. One day I was digging for
bait, when I stumbled on a—a cache, and in a certain spot
on this island is one hidden now. Treasure is in it. If you
have brought spades, dig away till you find it. I will give you
a clue. Four paces from Security, an arm's length to the
right, dig for two feet down!"

"We haven't any spades," said the little girls. "The

Captain has the only one."

The young man went into the hut, and soon appeared
with a pitchfork and two spades of very small proportions.
 "And now," said Charity, "where is Security? It must be
the hut, of course!"

In a very few minutes each child was digging four paces

from the hut. But Charlie began to flag, whereupon the
young man whispered something to him.

"Your Captain is ill," he said; "he's going to rest. I'll take

his place."

Charlie sat down on the step leading to the hut and

watched the others with a rather bitter face. It was hard to
be bowled over so soon, just when he would like to prove
his strength to be superior to the girls.

Digging went on steadily, but the three little girls made

slow progress.

The young man dug too, but he presently said, "I'll give
another clue:"

"By the side of a singer's home, a hand's span from the

base."

The little girls were completely puzzled. Charlie's bright

eyes roved to and fro. At last his face lightened.

"I have it. A singer's home is a bird's tree, and the base
is the trunk of it. There's only one tree which it can mean,
and it's the ash tree between Bolt and Ben."

Charity made a rush to the spot, and Charlie sprang up

declaring he was quite rested. He and the three girls all
attacked the ground round the ash tree, and the young man
quietly slipped into the hut, leaving them at work.
 It was not long before Charity's spade hit against
something hard. Then four eager pairs of hands dragged it
to light. It was a rusty tin box tied with string and sealed.

Charlie took command as Captain, and cut the string

with his clasp knife. His face was solemn as he did it, but
the little girls' faces bubbled all over with curiosity and
delight.

It was hard to open, but at last the lid gave way, and
then Charlie very carefully lifted out the contents.

Four black farthings, a blue marble, and two peach

stones.

Faith's face fell; she was dreadfully disappointed. She

had really expected to find it full of precious stones.

Charity danced up and down.

"Golden sovereigns, a blue jewel worth a million

pounds, and seeds of a pomegranate that grew in
Paradise!" she cried.

Charlie turned to her with an approving face.

"You are right, my wife, wonderful treasure indeed!

Does the Pirate mean to let us carry it off?"

"The Pirate invites you to a meal. He is tired of a very

lonely life and welcomes treasure seekers to his home."

It was the young man who spoke. The children dashed

into the hut, Charlie clutching his tin box. There another
surprise awaited them.
 A kettle was boiling merrily. A cake was on the table,
and some ham sandwiches. The Pirate had cups and
saucers, and was measuring tea into a brown teapot.

"We have no cow on the island, but we have sugar and

good tea. Let us fall to!"

It seemed like magic. The children sat up round a little

table and they had a merry meal. After it was over, the
treasure box was produced and the treasures divided. The
Pirate took one of the peach stones. Charity took the other.

"I as Captain's wife have first choice," she said. "I am

going to plant this wonderful seed, and perhaps it will
spring up into a magic tree which will reach the sun."

Charlie gave Hope and Faith a farthing each.

"A guinea for Bolt and Ben," he said. "I being Captain
keep the blue diamond and a guinea, my wife can have the
other."

Then the Pirate pulled out his pipe, and sitting cross
legged on the ground told them the most wonderful story of
how the treasures had been obtained and hidden away. The
children listened breathlessly, but at last the Captain said it
was getting late, and they must go. The Pirate took them
back to their raft, and then he surprised them again. He got
out a very small boat from under the willows, tucked
himself into it, and fastening the rope of the raft to his
painter, rowed gaily off down the stream, towing the
children back to the spot where he found them. Then he
bade them good-bye.

"We shall never meet again most likely," he said, "but I

warn you to keep to your own waters. There are other
pirates who would make short work of you should they find
you where I did!"

The children waved their hands gaily to him. Charlie

was supremely happy, and content at the result of his
voyage after treasure, and all their tongues wagged fast as
they made their way down the stream towards Charlie's
home. It was nearly six o'clock when they got there, so the
little girls bade their new friend a hasty good-bye.

"It has been perfect—simply perfect!" said Charity, and

the others echoed her words.

They ran home then as fast as they could, and told

Granny all of their adventures.

"Who do you think the Pirate can be?" Faith asked. "He
has such nice kind eyes, but a very grave face."

Granny said she could not possibly tell, and Aunt Alice
could not help them.

But the next day the rector's wife paid a long call.
Charity happened to be in the room, and though she was as
quiet as a little mouse she kept her ears wide open, and
when she was alone with Hope and Faith she was quite
excited.

"I believe Mrs. Webster was talking about the Pirate, I

believe she was! She told Aunt Alice she had a little
grandson staying with her last week, and she wished she
had asked us over to tea with him, as he was so lonely. And
then she said that Fred Cardwell had been so good to him.
He had taken him off fishing with him several days, and had
entertained him on an island which the little boy had loved.
She asked Aunt Alice if she knew the Cardwells, and I
pricked up my ears and listened hard. Aunt Alice said no,
and she said Fred Cardwell lived with a very cross, ill father
—I think he's a squire, like Sir George, and they live about
five miles from here. The father is parry—something—a long
word, and Fred had to come home and look after him, and
he's no mother or brothers and sisters, and Mrs. Webster
said it was a terrible life for him, and it made him gloomy,
and he doesn't go anywhere or won't know anybody, but he
likes children and she said her little grandson loved him.
Don't you think Fred may be our pirate? Because there can't
be lots of islands about, and perhaps that was why he had
cake and tea in the hut, he had put them there when the
little boy was with him!"

Charity paused for breath.

"I wish we could see him again," said Faith.

"We'll keep a sharp look-out along the roads when we

walk," said Hope. "Mrs. Cox wasn't at all right about the
country. We do meet people very often, and we may meet
him."

"We'll ask Lady Melville if she knows him when we see

her next," Charity said; "and now I'm going to plant my
magic seed. Come and see me do it."

So Hope and Faith accompanied her to the little garden,

and she planted it just below their bedroom window.

"Perhaps it will be like Jack's beanstalk, and grow so

high that we can step out of our window on to its branches,"
said Hope.

"It's sure to be different from any other tree, for it has

been hidden away for years, I'm sure," said Charity.
 They did not see Charlie for some time after this. They
heard that he was ill, and one day a letter came from him
addressed to "My Wife and Crew."

The little girls opened it with great delight. It was very

short.

"This is to tell you that youre Captin lies

dangirosly wunded, and his sickness is suvere.
His next voyage will not take plaice, and when
he gets better Ben is to come to him to delever
messages to his wife and to Bolt.

"Charles, Captain of
the 'Success.'"

Faith was Ben. She wondered why she was especially

invited. Charity tossed her head.

"He knows he can do what he likes with Faith. I expect

none of us will be able to go with him now, for Aunt Alice
said we were to begin lessons. This new governess, Miss
Vale, is coming next Monday."

"Well, anyhow," said Hope, "I'm glad it isn't school, and

Aunt Alice says she will be very nice."

"I must answer his letter," said Faith.

But she was not very fond of letter-writing and put it

off. She left Charity and Hope playing in the orchard that
afternoon and went off to visit her friend the shepherd.
There was nothing she enjoyed so much as creeping into his
little cottage and sitting on a small stool in the chimney
corner with the old man. Sandy would come and rest his
nose in her lap, and she and Timothy found plenty to say to
each other. She told him all about Charlie, and the raft and
the Pirate.

"'Tis a pity the little laddie enjoys such poor health,"

said Timothy to her. "The doctor be such a hearty man, but
there—the Lord have a way of His Own with each o' us, and
'tis ordained for him to be weakly. I often sits and thinks o'
strength. 'Tis misused in the body, and if so be the soul is
strong, 'tisn't so much odds about the body!"

"I wonder how strong my soul is," said Faith. "I'm not
very strong in my body, Granny says. Can I make my soul
strong, Timothy?"

"Ask the Comforter," said the old man. "He'll strengthen

the weak. We are told 'He helpeth our infirmities.'"

"What do infirmities mean?"

"Our weakness and ailments, surely. The Book says, 'We

can be strengthened with might in the inner man' by Him."

"Is our inner man our souls?" asked Faith.

"I suppose I have an inner woman—I'm not a man."

"'Tis just a figure o' speech. 'Tis your little soul you
want to be made strong."

"I like the Comforter very much," said Faith softly and
reverently; "He came and comforted me this morning,
Timothy. Aunt Alice scolded me because she told us not to
leave the front door open, there was such a wind. And Hope
left it open; she came out last, and the wind knocked a
china vase off the table, and broke it, and Aunt Alice was
very angry and scolded me, because she thought I'd gone
out last. And I went away and cried, and then I distinctly
felt the Comforter near me, and I asked Him to comfort me.
I almost felt He took me in His Arms. He was so close. And I
kept quite still, and then I couldn't be sorry any more, for I
knew He knew I hadn't done it. He was so kind!"

Faith heaved a sigh of happiness, and Timothy nodded

his head.

"'Tis just so!" he said simply.

And then they began to talk about Sandy and the

sheep, and when she left the cottage, Faith's little face was
radiant.

"I feel when I'm talking to you," she said as she bade
the old man farewell, "that I'm getting happier every
minute. I shan't be able to come and see you so often when
we do lessons, but I'll come whenever I can!"

Charity and Hope could not understand her friendship

with the old man. But Faith paid no heed to them. She was
a quiet, old-fashioned child, and loved to go her own way
without any interference of other people.

CHAPTER VI
CHARLIE STILL IN COMMAND
 MISS VALE arrived on Monday, and the little girls fell in
love with her. She was very pretty, with bright, dark eyes,
and a quick, cheerful manner. But they found she was very
firm and strict in some things, and lessons could not be
trifled with.

"I shall not give you any lessons you cannot prepare,
and when I come I expect to find them done. If they are
not, I shall conclude it is idleness that is the cause and will
deal with it accordingly."

This sounded very alarming, but the children found that

she was right, and that there was no excuse for their
lessons not being learnt.

She came from nine o'clock to one, and they had an

hour every afternoon in which they did what they called
their "prep" for her.

Charity and Hope did everything together, Faith could

not keep up with them. She was slow and persevering, but
not very clever at books. Yet Miss Vale, if she had any
preference, liked to teach her the best of the three, for her
whole heart was in what she did, and she was extremely
conscientious.

In a few days' time, Faith was allowed to go to see

Charlie. His mother met her at the door.

"My boy did too much with you that day. He has been in
bed ever since. His father says there must be no voyages
down the stream for a long time to come, so don't
encourage him to talk about it."

Faith was taken into the sitting-room, where Charlie lay

on the couch, looking very white and frail, but he greeted
her most cheerfully:
 "Come on, Ben. Isn't it hard lines for me?"

"What's been the matter with you?" asked Faith in a

sympathetic tone.

"Oh temperature; it's always that. My head nearly

bursting and I'm hot as fire! But I'm all right now. What
have you been doing?"

Faith gave an account of their days since last they met,

and from being very bright Charlie's spirits sank, and he
began to talk most gloomily.

"It's no use my trying to do anything like other fellows.

If God had made me a girl, I shouldn't have minded half so
much, but boys are meant to be strong, and I think it's a
shame I shouldn't be. I quite hate myself sometimes. If I
was meant to be weak and ill, I oughtn't to have been born
a boy. It was a big mistake."

"But," said Faith with rather a shocked look, "God

borned you, and He can't make mistakes. And I don't see
why girls should be ill and not boys. Besides, Charlie, if you
aren't strong outside, you can be inside. I was talking with
my friend Timothy about you. He said:"

"'It wasn't any odds about the body, it's the soul that
really matters.'"

"Don't you like people with strong souls?"

"I don't know what they're like," Charlie said, staring at

her.

Faith's eyes grew big and glowed with light as she

replied:
 "Oh, they're heroes!—Always smiling at difficult days
and going straight on with their heads up, even if they're
hurt. I've thought about them a lot. And I'm hoping the
Comforter will make my soul grow big and strong. I should
like never to cry when I'm hurt, and never grumble when
things are horrid! And keep smiling even if the whole world
turned against me and trampled me down!"

Faith spoke with such fervour that Charlie was much

impressed.

She added:

"We all think you have a strong soul because you're so

cheerful."

"Oh, I'm not!" said Charlie. "I've been beastly to Mother

I'm so awfully disappointed that Dad won't let me go on the
raft. It's the one thing I really enjoy! There's nothing else I
can play at. It does seem a shame."

"We play lots of games," said Faith, "and we've no raft."

"No, but you are girls, and you can run about, and climb
trees. I've had a miserable time shut up here all alone."

"I suppose," said Faith shyly, "you wouldn't like the

Comforter to stay with you? I think you would feel better if
He did. I don't know Him very well yet, but Timothy talks to
me about Him. And I think it's so wonderful that He likes
coming and living with boys and girls and making them
happy and good and strong!"

"What do you mean?" asked Charlie.

"Timothy told me about Him first. If you're ill and

unhappy, He would like to come to you and comfort you.
That's why He's called the Comforter. Wherever He goes He
comforts, and He does it perfectly, because, you know, He
is God."

Faith's voice sank to an awed whisper.

"What a funny girl you are!" said Charlie. "You seemed

as if you were talking about a real person!"

"But the Comforter is real."

"Well, a real person is somebody you can see."

"If you can't see Him, you can hear Him," said Faith
gravely. "There are lots of real things we can't see. The
wind—"

"Oh, I know, but you're preaching a sermon."

"Am I?"

Faith subsided into silence. Presently she said:

"Then I can't tell you how to be happy and comforted, if

you won't believe what Timothy says."

"Can you find out this puzzle?"

Charlie would have no more serious talk; and produced

a little puzzle box with a padlock which opened itself in a
wonderful way; Faith was interested at once. She paid him
quite a long visit, and when the time came for her to go,
Charlie produced two packets which he charged her to give
to the wife and to Bolt.

"They're my wishes," he said. "You signed on to do what

I told you for a year, and I'm still in command, though I'm
ill!"
 "Aye, sir, aye!" said Faith, saluting him in best style.

Then she went home, and Charity and Hope opened

their packets eagerly.

There was a great deal of paper but not much writing.

Charity's was as follows:

"Captain Charles sends greetings to his wife.

He wishes her to find the Pirate's Haunt, and let
me know his house and his riteful name. She
must make the journey by land, but she must
not fale to do it. For a lot hangs on the Finding!
And she must rite the name in sekrecy and seal
her letter with a red seal and send it through
the post. And if she does not keep it sekret
death will o'ertake her.

"(Signed) CAPTAIN
CHARLES."

Hope opened hers and found a strange map, drawn in

red ink by Charlie. This was the letter accompanying it.

"Captain Charles to his humble and devotted

servant Bolt. I charge you to studey this map,
and to find on your walks the place that is
called Boggy Glen. There is a wonderful herbe
therein, called Wild peppermint, whichesame
will releave the Captain of his mortal sicknes,
and is to be sent to me in a sealed letter by
poste with no derlay.
 "(Signed) CAPTAIN
CHARLES."

"What fun!" cried Hope. "You and me, Charity, will have
to be busy!"

"And he's given me nothing to do," said Faith, feeling

aggrieved; "nothing at all."

"You're his messenger," said Charity, "you must take

back answers to these notes. A King's messenger is most
important, and so is a Captain's messenger."

Faith's face brightened. She took back two notes the

next day to the doctor's house, but did not see Charlie.
These were the notes:

"MY DEAR HUSBAND,

"I'm sorry you're sick, but I'll find the Pirate

in a jiffy. I wish you had told me to hang him or
something difficult like that, but after all I don't
want him hanged, because he was very kind. I
mustn't let him know of course. I'll criep about
like a spy and follow his tracks. I think I've
already found out his name. But I'll write it
hidden in a sentence and you'll have to find it
out. Good-bye and good luck.

"Your sooperior
WIFE."
 "MY DEAR CAPTAIN,

"I will find the heeling herb, no fear about

that! And I'll make a potion of it to be
swallowed at midnight in the dark. And you will
rise up early a hale and harty man. So three
cheers for our Captain and his crew.

"Yours as signed,
BOLT."

Faith had these letters read aloud to her and she

thought them wonderfully clever.

The very next day, Charity and Hope set off on their
different quests. Faith wandered out alone. She called at
Timothy's cottage, but he was out; and then she rambled
on through some fields, and made her way along a strange
lane, which had banks of primroses on each side. Presently
she saw a hole through the hedge; she crept through, and
then she started, for two men were talking together. They
were standing by a rick of hay, and she heard one man say
with passion in his tone:

"I tell you, Fielding, I'm so dead sick of rotting here like
a vegetable, when I might be up and doing with the rest of
the working world, that at times I feel inclined to make a
bolt for it!"

"It's hard lines," murmured the other man, and then the
two separated.

One went on across the field, the other turned and

faced Faith, and she saw it was the Pirate. Just for a
moment he looked as if he were going to pass her without
speaking, and his brows were knitted so fiercely, and his
face so gloomy, that Faith was frightened. He stood still, as
he saw her shrinking into the hedge, and then his brow
cleared.

"Why, it's one of the treasure seekers, isn't it?" he said

in his pleasant voice.

Then Faith held out her small hand.

"I'm so glad you will speak to me," she said, "I should
have been so disappointed, if you hadn't."

"Would you? And what are you doing alone here?"

"Just walking about and trying to amuse myself,"

answered Faith. "Charity and Hope have gone off to do
errands for Charlie, but I've none to do."

For a moment she was tempted to tell him that Charity

was looking for him, then she remembered that Charlie said
it was to be kept a secret, so she shut her lips determinedly.

"Well, I'm an idle person too. We'll take a walk together,

I want to go to Dapperton Bridge to see if there's a chance
of any salmon there. Will you come with me?"

"Is it very far? I should love to."

"No, it is only half a mile further on."

He took hold of her hand, and chatted about different

things till they came to the bridge; then he leant his arms
on it, and as he gazed at the river flowing beneath, his face
became gloomy again, and absorbed. Faith stood by his side