• TABLE OF CONTENTS
HIDE
 Front Cover
 Frontispiece
 Table of Contents
 Acknowledgement
 Material studied
 Methods of collecting
 Field notes
 Previous work on the crayfishes...
 Derivation of the fauna
 Endemism
 Principal migration routes
 Barriers to crayfish migration
 Ecology
 Ecological habits and habitats
 The subfamily cambarinae
 The chief characters used in keys...
 Key to the Florida crayfishes
 Genus procambarus
 Barbatus group
 Barbatus subgroup
 Procambarus barbatus
 Procambarus pubischelae
 Procambarus escambiensis
 Procambarus econfinae
 Procambarus apalachicolae
 Procambarus rathbunae
 Shermani subgroup
 Procambarus shermani
 Kilbyi subgroup
 Hubbelli subgroup
 Alleni group
 Advena section
 Procambarus advena
 Procambarus geodytes
 Procambarus pygmaeus
 Rogersi group
 Procambarus rogersi rogersi
 Procambarus rogersi campestris
 Blandingii section
 Procambarus blandingii acutus
 Procambarus bivittatus
 Clarkii subgroup
 Procambarus okaloosae
 Procambarus paeninsulanus
 Evermanni subgroup
 Fallax subgroup
 Procambarus leonensis
 Procambarus pycnogonopodus
 Spiculifer group
 Procambarus versutus
 Pictus group
 Procambarus pictus
 Procambarus youngi
 Lucifungus subgroup
 Procambarus lucifungus alachua
 Procambarus pallidus
 Seminolae subgroup
 Genus troglocambarus
 Genus cambarellus
 Genus orconectes
 Genus faxonella
 Genus cambarus
 Cambarus latimanus
 Cambarus floridanus
 Cambarus cryptodytes
 Diogenes section
 Cambarus byersi
 Cambarus (species incertis)
 A county list of the Florida...
 Literature cited
 Index






Group Title: University of Florida publication
Title: The Crayfishes of Florida
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00090901/00001
 Material Information
Title: The Crayfishes of Florida
Alternate Title: Biological science series - University of Florida ; vol. 3, no. 2
Physical Description: 1 p. l., 179 p., 1 p. : illus. (incl. maps) XXIV pl. (incl. front.) ; 25.5 cm.
Language: English
Creator: Hobbs, Horton Holcombe Jr., 1914-
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1942
Copyright Date: 1942
 Subjects
Subject: Crayfish   ( lcsh )
Crustacea -- Florida   ( lcsh )
Genre: non-fiction   ( marcgt )
 Notes
Statement of Responsibility: by Horton H. Hobbs, Jr.
Bibliography: "Literature cited": p. 172-176.
General Note: "November, 1942."
 Record Information
Bibliographic ID: UF00090901
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 01245795
lccn - 42036584

Table of Contents
    Front Cover
        Front cover
    Frontispiece
        Frontispiece
    Table of Contents
        Page i
        Page ii
        Page iii
        Page iv
        Page v
    Acknowledgement
        Page 1
        Page 2
    Material studied
        Page 3
    Methods of collecting
        Page 4
        Page 5
        Page 6
    Field notes
        Page 7
    Previous work on the crayfishes of Florida
        Page 8
        Page 9
    Derivation of the fauna
        Page 10
        Page 11
    Endemism
        Page 12
    Principal migration routes
        Page 13
    Barriers to crayfish migration
        Page 14
        Page 15
    Ecology
        Page 16
        Page 17
        Page 18
        Page 19
    Ecological habits and habitats
        Page 20
        Page 21
        Page 22
    The subfamily cambarinae
        Page 23
        Page 24
    The chief characters used in keys and descriptions
        Page 25
        Page 26
        Page 27
    Key to the Florida crayfishes
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
    Genus procambarus
        Page 33
        Page 34
    Barbatus group
        Page 35
        Page 36
        Page 37
    Barbatus subgroup
        Page 38
    Procambarus barbatus
        Page 39
        Page 40
    Procambarus pubischelae
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
    Procambarus escambiensis
        Page 46
        Page 47
        Page 48
    Procambarus econfinae
        Page 49
        Page 50
        Page 51
    Procambarus apalachicolae
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
        Page 57
        Page 58
    Procambarus rathbunae
        Page 59
    Shermani subgroup
        Page 60
    Procambarus shermani
        Page 61
        Page 62
        Page 63
    Kilbyi subgroup
        Page 64
        Page 65
        Page 66
    Hubbelli subgroup
        Page 67
        Page 68
    Alleni group
        Page 69
        Page 70
        Page 71
        Page 72
    Advena section
        Page 73
        Page 74
    Procambarus advena
        Page 75
        Page 76
        Page 77
        Page 78
        Page 79
    Procambarus geodytes
        Page 80
        Page 81
        Page 82
    Procambarus pygmaeus
        Page 83
        Page 84
        Page 85
        Page 86
        Page 87
    Rogersi group
        Page 88
    Procambarus rogersi rogersi
        Page 89
    Procambarus rogersi campestris
        Page 90
        Page 91
        Page 92
    Blandingii section
        Page 93
    Procambarus blandingii acutus
        Page 94
        Page 95
    Procambarus bivittatus
        Page 96
        Page 97
    Clarkii subgroup
        Page 98
        Page 99
    Procambarus okaloosae
        Page 100
        Page 101
        Page 102
        Page 103
    Procambarus paeninsulanus
        Page 104
        Page 105
        Page 106
    Evermanni subgroup
        Page 107
        Page 108
        Page 109
        Page 110
    Fallax subgroup
        Page 111
        Page 112
        Page 113
    Procambarus leonensis
        Page 114
        Page 115
        Page 116
    Procambarus pycnogonopodus
        Page 117
        Page 118
    Spiculifer group
        Page 119
        Page 120
        Page 121
        Page 122
        Page 123
        Page 124
        Page 125
    Procambarus versutus
        Page 126
        Page 127
        Page 128
    Pictus group
        Page 129
    Procambarus pictus
        Page 130
    Procambarus youngi
        Page 131
        Page 132
        Page 133
    Lucifungus subgroup
        Page 134
        Page 135
    Procambarus lucifungus alachua
        Page 136
        Page 137
        Page 138
    Procambarus pallidus
        Page 139
        Page 140
        Page 141
    Seminolae subgroup
        Page 142
        Page 143
        Page 144
        Page 145
    Genus troglocambarus
        Page 146
        Page 147
        Page 148
    Genus cambarellus
        Page 149
        Page 150
        Page 151
        Page 152
    Genus orconectes
        Page 153
    Genus faxonella
        Page 154
        Page 155
    Genus cambarus
        Page 156
        Page 157
    Cambarus latimanus
        Page 158
        Page 159
        Page 160
    Cambarus floridanus
        Page 161
    Cambarus cryptodytes
        Page 162
        Page 163
    Diogenes section
        Page 164
        Page 165
        Page 166
    Cambarus byersi
        Page 167
    Cambarus (species incertis)
        Page 168
        Page 169
    A county list of the Florida crayfishes
        Page 170
        Page 171
    Literature cited
        Page 172
        Page 173
        Page 174
        Page 175
        Page 176
    Index
        Page 177
        Page 178
        Page 179
        Page 180
        Page 181
        Page 182
        Page 183
        Page 184
        Page 185
        Page 186
        Page 187
        Page 188
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        Page 197
        Page 198
        Page 199
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        Page 207
        Page 208
        Page 209
        Page 210
        Page 211
        Page 212
        Page 213
        Page 214
        Page 215
        Page 216
        Page 217
        Page 218
        Page 219
        Page 220
        Page 221
        Page 222
        Page 223
        Page 224
        Page 225
        Page 226
Full Text

UNIVERSITY OF FLORIDA PUBLICATION
NOVEMBER, 1942
BIOLOGICAL SCIENCE SERIES


THE CRAYFISHES


OF FLORIDA





BY

HORTON H. HOBBS, JR.


Vol. III


No. 2


Published by the University of Florida under the auspices of the
Committee on University Publications


















































Plate I. Troglocambarus maclanei Hobbs









TABLE OF CONTENTS

PAGE

THE CRAYFISHES OF FLORIDA.-..............-......................-------- --------------------- 1
Acknowledgments ........................---....--------.................................... 1
Material Studied ..........................................................---..------------- 3
Methods of Collecting ........................--------- ---------------- 4
Field Notes ............................... ..........--................--------- ----. 7
Methods of Preservation and Study .................................. ........................ 7
Previous Work on the Crayfishes of Florida ...................................................... 8

FACTORS AFFECTING THE DISTRIBUTION OF CRAYFISHES IN FLORIDA........................ 9
Florida as a Faunal Region .................................---........ ----------- 9
Derivation of the Fauna ................---.----.... ----------.--......... 10
Endemism .......................--......-------------..-----.. ------- -- 12
Principal Migration Routes ....................... ..-------- ------- ......... 13
Barriers to Crayfish Migration ................................................ 14
Ecology ...........................---------------------------------................. 16
Ecological Habits and Habitats .................................................... .................. 20

THE SUBFAMILY CAMBARINAE ................................---- -------........------- 23
The Evaluation of Taxonomic Characters in the Cambarinae ............................ 23
The Chief Characters Used in Keys and Descriptions ........................................ 25
t Key to the Florida Crayfishes .............................-.................-------.. .. ...- 28

GENUS PROCAMBARUS ................------------- ---------------------------------------- 33
SBarbatus Section ....................-..................----- ---------------.......... 33
Barbatus Group .......................--.................... ----. --------------- 35
Barbatus Subgroup ....... ........ ...........----------............................ 38
Procambarus barbatus ............................ ..... ................ ............... 39
Procambarus barbatus .............................................................................. 39
Procambarus pubischelae ................................ ...........................------- 41
Procambarus escambiensis .....-............-...-------------------46
Procambarus econfinae ............................-------- --.......................... 49
Procambarus latipleurum .............................---........... .......................... 52
Procambarus apalachicolae ........................................-----............................-- 55
Procambarus rathbunae .................................------------------.................................... 59
1i







TABLE OF CONTENTS-Continued

PAGE
Shermani Subgroup ...................................................................................... 60
Procambarus shermani ................................................ ...................... 61
Kilbyi Subgroup .......................................................................................... 64
Procambarus kilbyi .................................................................................. 64
Hubbelli Subgroup ..................................................................................... 67
Procambarus hubbelli ...............................-.............................. ........... 67
Alleni Group ...................... ........................ ............................ 69
Procambarus alleni ................................................................................. 69
Advena Section .............................. .... ............... ......................... ......... 73
Advena Group ..................... .... ..... ....... ........................................... 73
Procambarus advena ................................ .................. 75
Procambarus geodytes ...................................... ................................ 80
Procambarus pygmaeus .......................... ................................... 83
Rogersi Group ...................... .. ....... ................................ ........ 88
Procambarus rogersi rogersi ............................ .................................... 89
Procambarus rogersi ochlocknensis ........................................................ 89
Procambarus rogersi campestris ............................................................. 90
Acherontis Section ----------- ----------------...................... ............................. 91
Procambarus acherontis ........................ .......................................... 91
Blandingii Section ........................................................................................ 93
Blandingii Group .............................................................................................. 93
Blandingii Subgroup ................................................................ ........... 93
Procambarus blandingii acutus ............................................................. 94
Procambarus bivittatus .............................................. .......................... 96
Clarkii Subgroup .......................................................................................... 98
Procambarus okaloosae ............................................................................ 100
Procambarus paeninsulanus .................................................... ........... 104
Evermanni Subgroup ........................................................... ................... 107
Procambarus evermanni ................................................. ..................... 107
Fallax Subgroup ................................. .......... .. ....................... .... 111
Procambarus fallax .................... .......................... ........... 111
Procambarus leonensis .......................................................................... 114
Procambarus pycnogonopodus ........................... ............................... 117
ii






TABLE OF CONTENTS-Continued

PAGE
Spiculifer Group ........................................ .... ........................ 119
Procambarus spiculifer ............................................................... 119
Procambarus versutus ...................................... 126
Pictus Group ........... ....................................... 129
Pictus Subgroup ................................................ 129
Procambarus pictus ... .... ....................... ....................... 130
Procambarus young ....................................................... 131
Lucifugus Subgroup ...................................... ...................... 134
Procambarus lucifugus lucifugus ...................................................... 134
Procambarus lucifugus alachua ............ ........ .................... 136
Procambarus pallidus ...................... ... ..................... 139
Seminolae Subgroup .. ................................................. 142
Procambarus seminolae ......................... ............. ....................... 142

GENUS TROGLOCAMBARUS .......................................................................... 146
Troglocambarus maclanei ............................................... 146

GENUS CAMBARELLUS ... ........... ........................... ........................ 149
Cambarellus schmitti ................... ..... ....... ............................ 149

GENUS ORCONECTES .......... ........................ ....... ............................................ 153
Subgenus Faxonella ..................................................... 154
Orconectes clypeata .......... ...................... ........................... 154

GENUS CAMBARUS ... ....................................................... 156
Bartonii Section ............................... ................... ......................... 157
Cambarus latimanus .......................... ...... ............. 158
Cambarus floridanus .......................................................................... 161
Cambarus cryptodytes ........................... ..... ................................... 162
Diogenes Section ........................ .................................................................. 164
Cambarus diogenes ....................................... 164
Cambarus byersi ....................................... 167
Cambarus (species incertis) ................................................................. 168

A COUNTY LIST OF THE FLORIDA CRAYFISHES ........................................................ 170

LITERATURE CITED ........ .... ........... .......................................... 172

INDEX ............................................... ... .................................................................... 177
iii






TABLE OF CONTENTS-Continued


TEXT FIGURES
PAGE
a-Specimens of Field and Species Catalogue Cards .................................................. 6
b-Terminology and Methods of Taking Measurements ............................................ 24
c-Hypothetical First Pleopod of Male ...................................................................... 26

MAPS

1-County Map of Florida Showing Areas in Which Field Work Has Been Done.... 2
2-Continuous North-South Extents of Well Drained Soils That Appear to Act as
Barriers to Crayfish M igration ........................................................ ............... 14
3-Migrations of the Barbatus Group ........................................................................ 34
4--Distribution of the Barbatus Group .................................................................... 36
5-Distribution of the Advena Section, Procambarus alleni, and Procambarus
blandingii acutus ............................................................................................... 74
6-Distribution of the Clarkii Subgroup .................................................................. 99
7-Distribution of the Evermanni and Fallax Subgroups ........................................ 110
8-Distribution of the Spiculifer Group and Procambarus acherontis .................. 120
9-Distribution of the Pictus Group and Procambarus bivittatus .......................... 130
10-Distribution of the Genera Troglocambarus, Cambarellus, and Orconectes in
Florida .................................................................................................................. 148
11-Distribution of the Genus Cambarus in Florida ................................................ 156

PLATES

I-Frontispiece-Troglocambarus maclanei
II-Procambarus barbatus, P. pubischelae, P. escambiensis, P. econfinae
III-Procambarus latipleurum, P. apalachicolae, P. rathbunae, P. shermani
IV-Procambarus kilbyi, P. hubbelli, P. alleni, P. advena
V-Procambarus geodytes, P. pygmaeus, P. rogersi rogersi, P. rogersi och-
locknensis
VI-Procambarus rogersi campestris, P. acherontis, P. blandingii acutus, P.
bivittatus
VII--Procambarus okaloosae, P. paeninsulanus, P. evermanni, P. fallax
VIII-Procambarus leonensis, P. pycnogonopodus, P. spiculifer, P. versutus
IX-Procambarus pictus, P. young, P. lucifugus lucifugus, P. lucifugus
alachua






TABLE OF CONTENTS-Continued


X-Procambarus pallidus, P. seminolae, Troglocambarus maclanei, Cambar-
ellus schmitti
XI-Orconectes clypeata, Cambarus latimanus, C. floridanus, C. cryptodytes
XII-Cambarus diogenes, C. byersi, C. species incertis
XIII-Procambarus pubischelae
XIV-Procambarus escambiensis
XV-Procambarus econfinae
XVI-Procambarus latipleurum
XVII-Procambarus apalachicolae
XVIII-Procambarus shermani
XIX-Procambarus geodytes
XX-Procambarus pygmaeus, Cambarellus schmitti
XXI-Procambarus bivittatus
XXII-Procambarus okaloosae
XXIII-Procambarus young
XXIV-Procambarus seminolae








THE CRAYFISHES OF FLORIDA


Although the crayfishes of the subfamily Cambarinae are among the
most familiar of all invertebrate animals, many of the species and the group
as a whole present a number of interesting biological problems. Wholly Nearc-
tic in origin and distribution, the subfamily now occupies a large area in North
America east of the Continental Divide in spite of the low vagility of most of
the species and a marked tendency to be limited by topographical and ecologi-
cal barriers. Here in the vast extent of latitude, topography, and climate be-
tween south-central Canada and Guatemala the crayfishes of this group have
encountered and shown numerous adaptations to a wide range of ecological
conditions.
Perhaps it is for these reasons that the crayfishes have attracted a number
of able students, who have not only laid a good basis for taxonomic and geo-
graphic work but, at least in the case of Ortmann, utilized the group for classi-
cal researches in geographic distribution. Most of this work, however, has
dealt with regions other than the southeastern coastal plain. Here, except for
the pioneer work of LeConte, the group has been neglected, and there is little
evidence that any of the older workers suspected that this region and especial-
ly Florida would prove to be exceptionally rich in species and present so many
striking instances of recent migration, speciation, and adaptation to special-
ized habitats.
My own work, begun ten years ago, was at first visualized as the compar-
atively simple task of locating some eight or ten species that had been reported
for Florida, learning to recognize them in the field, and determining their
local geographic and ecological ranges. It soon became apparent, however,
that Florida did not present a simple problem. As the collections grew in
numbers of specimens and in the regions and habitats represented, it became
evident that I had to deal with an almost uniquely rich and varied crayfish fau-
na. Today the Florida list has grown to 42 species and subspecies, the largest
list for any state in the Union, and the details of their occurrence present a
wealth of data on the comparatively recent spread of the stock into the region,
on its encounters with ecological highways and barriers to migration, and the
consequent development of a high proportion of endemic races and species.

ACKNOWLEDGMENTS
The present investigation has been carried out with the active encourage-
ment of Professor J. Speed Rogers. His suggestions have been invaluable in
working out the details of the problem, and I have gone to him freely for criti-
cism of the manuscript. For this aid, for the loan of equipment, and for the
personal interest he has shown in this work on the Florida crayfishes, I wish
to express my sincerest thanks.
Especial thanks are due Dr. Waldo L. Schmitt of the United States Na-
tional Museum and Dr. Fenner A. Chace of the Museum of Comparative Zo-






2 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2


Map 1.-County Map of Florida showing Areas in which Field Work has been done.






HOBBS--THE CRAYFISHES OF FLORIDA


ology who have kindly made comparisons of my specimens with typical mater-
ial and through whom loans were secured from their respective museums. I
wish also to express my gratitude to Dr. E. P. Creaser formerly of the Museum
of Zoology, University of Michigan, for the identification of specimens when
this work was first begun.
To Professors T. H. Hubbell, C. F. Byers, H. B. Sherman, and M. D.
Cody I wish to express my appreciation for many valuable suggestions and
criticisms throughout the course of the present study. Thanks are also due Mr.
G. Robert Lunz of the Charleston Museum and Mr. K. E. Goellner of the Mu-
seum of Zoology, University of Michigan, for the loan of specimens.
For their contribution of specimens and for companionship on collecting
trips I am indebted to Dr. H. K. Wallace, Dr. A. F. Carr, Dr. Lewis Berner, Dr.
Frank N. Young, Dr. A. M. Laessle, Messrs. Lewis J. Marchand, W. M. Mc-
Lane, Joel M. Martin, John D. Kilby, Coleman J. Goin, .James J. Friauf, R.
E. Bellamy, Glendy Sadler, G. W. Van Hyning, R. W. Williams, Ray Boles,
Lee Stanton, and many others.
Finally I wish to acknowledge my great indebtedness to my parents, Mr.
and Mrs. H. H. Hobbs, who have made this study possible through financial
aid and by providing transportation on numerous collecting trips, and to my
wife, Georgia Blount Hobbs, who has checked many of the detailed compila-
tions necessary in the preparation of the manuscript.

MATERIAL STUDIED

By far the greatest amount of the material on which this paper is based
is contained in my own collection, nearly all of which has been collected by
myself and my colleagues of the University of Florida. This collection com-
prises some ten to twelve thousand Florida specimens.
I have also examined an additional three to five hundred specimens in
the collections of the various museums. These included all of the extant types
of the species that have been described from Florida by other workers.
In 1935 and again in 1937 I visited the United States National Museum
in order to study the type specimens of all of the species of the subfamily Cam-
barinae contained in that collection. Further typical material was examined at
the Museum of Comparative Zoology and at the Philadelphia Academy of Sci-
ences during February 1937. While at these museums I took the opportunity
of examining many other specimens which had been collected in Florida,
Georgia, and Alabama.
The United States National Museum, the University of Michigan Mu-
seum of Zoology, and the Museum of Comparative Zoology have kindly lent
me numerous specimens including some Florida materials. These have been
extremely useful in making comparisons and in giving me additional locality
records for the state.






4 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Unless otherwise specified all of the specimens mentioned in this paper
are in my personal collection at the University of Florida.1
In the beginning I had planned to do intensive collecting throughout the
state of Florida; however, it was soon apparent that some regions required
much more detailed study than others. The crayfish fauna of southern Flor-
ida is restricted to two widely distributed species, but the northern and eastern
portions of the state have no fewer than 40 species or subspecies, and so re-
quired intensive collecting in order to decipher relationships and ascertain
the limits of the ranges involved.
The accompanying map shows that although collections have been made
in all of the 67 counties of Florida, western and northern Florida have re-
ceived a much greater emphasis than the southern two-thirds of the peninsula.
In addition collections have been made in the following counties of Alabama
and Georgia: ALABAMA-Baldwin, Butler, Conecuh, Dale, Elmore, Escambia,
Houston, Lee, Lowndes, Mobile, Montgomery, and Russell; GEORGIA-Ap-
pling, Baker, Ben Hill, Brooks, Bryan, Bulloch, Burke, Camden, Clarke,
Clinch, Colquitt, Cook, Decatur, Dooly, Dougherty, Early, Echols, Effingham,
Emanuel, Floyd, Glynn, Grady, Gray, Greene, Habersham, Houston, Jeffer-
son, Jenkins, Jones, Lanier, Liberty, Long, Lowndes, McDuffie, McIntosh,
Mitchell, Murray, Oconee, Pierce, Rabun, Screven, Sumter, Tattnall, Thomas,
Toombs, Ware, Washington, Wayne, and Wilkes.

METHODS OF COLLECTING
One of the best collecting implements is a fine meshed minnow seine. In
open water, even in small ditches or puddles where vegetation is not too dense,
thorough seining is quickly completed and, if care is taken to drag the bottom,
is highly efficient. Seines also proved successful in those underground pools
in which the bottom was not strewn with rocks.
Another very useful device is the "D"-ring or semicircular dip net. This
consists of a semicircular frame, twelve inches in diameter, made of one-quar-
ter inch spring-steel rod with the ferrule in the middle of the rounded side, and
fitted with a five foot hickory handle. A variety of bags have been used with
this net, all equipped with a strong canvas rim. A bag of one-quarter or one-
half inch netting permits the swiftest movement through the water and is parti-
cularly useful in subterranean water where the crayfishes are likely to be very
wary and require rapid manipulation of the net. Where C. schmitti or other
very small species are sought for it is necessary to use a much smaller mesh,
and a bag of strong scrim with about one-sixteenth inch mesh is excellent. A
useful net for general work has a short canvas bag with a bottom of strong wire
screen (one-sixteenth inch mesh).

'The holotype and allotype of all of the species I have described or am describing in this paper
are deposited in the United States National Museum. Paratypes of several of them have been
deposited in the Museum of Comparative Zoology, University of Michigan Museum of Zoology, the
Philadelphia Academy of Sciences, and the Carnegie Museum.






HOBBS-THE CRAYFISHES OF FLORIDA 5

An ordinary garden rake is often very effective in roadside ditches and
in streams thickly grown with vegetation. Masses and mats of vegetation when
torn loose and brought out on the banks generally contain many of the cray-
fishes that live in these habitats. In many of the lakes and ponds of Florida
crayfishes are abundant in the roots of the floating water hyacinths. By rapidly
rolling the hyacinth mats or lifting the tangled mass onto the bank, many cray-
fishes may be captured as they crawl back toward the water. Some of the more
sluggish species may readily be captured without either nets or seines, and in
swift water where stones make a dip net cumbersome, it is often best to turn the
stones and capture the crayfish by hand.
Probably the most satisfactory of all collecting methods is to hunt at
night with the aid of a headlight. Then many of the crayfishes have left their
daytime hiding places and are to be found venturing into relatively open sit-
uations. When the bright beam of the light strikes the eyes of a crayfish two
brilliant red spots are reflected, and at fairly close range the entire animal is
clearly visible.
A large number of the specimens in my collection were dug from bur-
rows, and many species can rarely be collected in any other way. Since the size
of the burrow, and especially the size of the chimney pellets, gives some indica-
tion of the size of the crayfish that made the burrow, it is possible to concen-
trate on mature specimens and largely avoid laborious delving for immatures.
After digging out a number of crayfishes one learns the habits of the var-
ious species and can often identify them before he begins to dig, and once the
burrowing habits of a species are learned, the task of digging becomes less ar-
duous. For those which construct simple burrows a shovel may be used to dig
almost straight down to the bottom of the excavation, and little work with the
hands is necessary. When the mouth of the burrow opens below the water table
it is almost impossible to secure the specimen unless one keeps his hand in the
burrow to prevent the escape of the crayfish. The digging out of all primary
burrowers is more of a task. Since the burrows are complex and ramify in any
direction it is necessary to keep one hand in the burrow so that none of the
tunnels will be lost. Collecting is thus much easier if two persons work together.
When one must work alone a garden trowel may be used to loosen the soil
around the burrow so that the hand in the burrow may proceed downward un-
til the crayfish is cornered. In some instances, where the burrows are decidedly
more complex, one must dissect the entire structure; more often the crayfish
seeks the deepest portion, but occasionally it retreats to one of the side pas-
sages. The specimen should be grasped by the carapace, but often it is the cray-
fish which seizes the collector's finger. In the latter case the crayfish can often
be drawn to the surface or far enough up the burrow to be readily caught. For
certain of the primary burrowing species the extraction of the crayfish from its
burrow is much simpler. After the water table is reached, the water in the bur-
row is stirred vigorously and then left undisturbed for a short time; soon the
antennae of the crayfish will be seen whipping to and fro at the surface, and if








6 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2
p


3-1839-2a, 2b
Tallahassee


IMroh 18, 1939
Leon County, Florida


12 miles west of Tallahassee on St. Hy. 19. Crypto-
labis Ravine a deep ravine leading into the Ochlocknee
River. Water issues from boggy seepage area and flows
into a small clear stream with a coarse sandy bottom.
4 PE. s&culifer was taken from the creek, under logs and
debris while C. floridanus was taken with considerable
difficulty from burrows of the stream beds. The burrows
are extensive, and the thick mat of roots makes digging
a task. (Other pertinent data ar included here).
P. spiculifer (LeConte)
2. floridanus (ms. name)
Rogers, Young, Berner, and Hobbs


Detrm. by
Hobbs


Procambarus spiculifer


2-20- 0


1 66 I
.2A6Q if
IQar



W younSemg
cithAysodEs:
Cambarus floridanus


Number
3-1839-2a
Stte
Florida


12 miles west of
Tallahassee on
St. HY. 19.
Cryptolabis Ravine


Hobbs
Hobbs


Deam. by
ke-l.


Cambarus florida s


2 n2 tea E


2 1"




_ Am as.

Procambarue aeiolofer


NumbeT
-1899Q-2h


Ste
Fl iA


county
Loon
fASS?-~--
12 miles west of
Tallahassee on
St. Hy. 19.
Cryptolabia Ravine


Hobbs

Hobbs


Text Figure a.-Specimens of Field and Species Catalogue Cards.


I c----


.. . ... floride- "d, .


obbs ...... ... *... s or a






HOBBS-THE CRAYFISHES OF FLORIDA


one makes a quick grab the crayfish may be easily captured. If the first grab
is unsuccessful the crayfish will return to the surface in a short time.
The cavernicolous species, of course, present special collecting problems
due to limited and varied access to their subterranean habitats. This is empha-
sized in the case of Troglocambarus maclanei, which has the very unusual and
apparently specific habit of clinging to the ceilings of wholly submerged por-
tions of underground waterways. These various collecting methods are de-
scribed under the individual discussions of the cavernicolous species.
Although trapping has not been sufficiently tested, the few attempts that
I have made proved very unsatisfactory. An ordinary minnow trap made of
screen wire with a funnel turned inward at one end worked fairly well in a
cave, but I have had little success in trapping any of the surface species.

FIELD NOTES

Detailed field notes with precise data as to locality and habitat are partic-
ularly pertinent in Florida. The abrupt changes of soil types and ground wa-
ter often make an apparent discontinuity in the range of a certain species very
hard to verify, and it is often difficult or impossible to discover without de-
tailed habitat data just what factors in the environment are correlated with a
species' ecological distribution.
The kind of data recorded in my field catalogue which is kept on 3" x
5" library cards is illustrated in figure a-1. The catalogue number "3-1839-
2a, 2b" is written in the upper left-hand corner; the figure "3" represents the
month (March); the next (one or) two digits "18" denote the day of the
month; the next two indicate the year (1939). The figure "2", following the
second dash indicates that this was the second collection made on March 18,
1939. (The "a" and "b" following the "2" show that two different species
were collected from this station). Text figures a-2 and a-3 are the references
to this field catalogue entry in the species catalogue.

METHODS OF PRESERVATION AND STUDY
After trying several methods of killing and preserving crayfishes the
following has been adopted as the most satisfactory standard process: as soon
as the specimens are caught they are dropped into 80% ethyl alcohol; after
several days all except a quarter of an inch (left to avoid losing any branchiob-
dellid worms) is poured off and enough fresh alcohol of the same percentage
is added to well cover the specimens. This, if the container is tightly sealed,
will last for several years. All of my specimens except large series are kept in
eight ounce, clamp-top, glass jars. This method of preservation has the disad-
vantage that the color of the animal disappears almost immediately. A four
percent solution of formaldehyde is less rapid in destruction of color, and in
some instances I have had specimens preserved in formaldehyde in which the
color pattern, but not the color, was evident even a year after preservation, but






8 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

formaldehyde makes the specimens extremely brittle, so that a great deal of
breakage is inevitable. The 80% alcohol, on the other hand, leaves all of the
joints pliable so that the specimen may be handled with little danger of being
broken, and to work with such specimens, of course, is much more pleasant.
For studies of the first pleopod of the male or annulus ventralis of the female
it is best to remove the part in question, dehydrate with alcohol, clear it with
xylene, and after allowing it to dry, mount it on an insect pin with a numbered
label that permits one to unquestionably associate the pinned part with the
proper specimen. I have never found any shrinkage in such cleaned and pinned
mounts.

PREVIOUS WORK ON THE CRAYFISHES OF FLORIDA
The earliest published record of a crayfish taken from Florida was that
of R. Gibbes in 1850. Since this record was for C. affinis the determination
was evidently an error, and what species was actually seen is left for conjec-
ture.2
Hagen (1870) described C. fallax and recorded C. lecontei. The speci-
mens on which the latter determination was made were collected at Pensacola,
and although I have not seen them, it is probable that they are either versutus
or spiculifer, but hardly lecontei.
Faxon (1884) recorded alleni, clarkii, fallax, and versutus from Flori-
da but stated "three young specimens from Pensacola, Fla. (M. C. Z., No.
249), are also placed here [in C. lecontei] by Hagen, but the justice of the
determination seems doubtful. They do not agree with the types from Mobile."
Then (1885a) in the Revision of the Astacidae he gave additional locality rec-
ords for fallax and clarkii and stated that "a species of the C. bartonii group
also inhabits Florida." In 1890 he added an additional locality for alleni,
gave a brief description of its female, and described C. evermanni from Flo-
maton which is on the Alabama-Florida line. From the same locality he record-
ed barbatus (see p. 40).
Lbnnberg (1894a) made a preliminary report on his collecting fallax
and alleni from Florida and on the discovery of C. acherontis. In his second
publication (1894b) he cited additional localities for fallax and alleni and de-
scribed C. acherontis, the first of the cavernicolous species known from Flor-
ida.
Faxon (1898) recorded additional localities for fallax, alleni, and an-
other species that he took to be LUnnberg's acherontis.3
Harris (1903) in his An Ecological Catalogue of the Crayfishes Belong-
ing to the Genus Cambarus listed five species, fallax, clarkii, versutus, alleni,
and acherontis, from Florida.

'Hagen (1870: 101) "Lewis R. Gibbes quotes C. affinis from Florida, but his determinations
are not at all trustworthy."
'This is not acherontis but lucifugus lucifugus (Hobbs 1940: 389).






HOBBS-THE CRAYFISHES OF FLORIDA


Ortmann (1905a) stated that six species are known from Florida and
listed evermanni and barbatus as occurring in the western part of the state, as
well as in Alabama and Mississippi, although he cited no Florida records for
these two species.
Faxon (1914) gave additional localities for fallax and alleni and de-
scribed the peninsula specimens heretofore referred to clarkii as clarkii pae-
ninsulanus. The additional locality cited for acherontis4 (Eustis, Lake County,
Florida) is again based on an erroneous determination.
Subsequent to 1914 no further references to Florida crayfishes were pub-
lished until I described Cambarus rogersi in 1938. Thus in 1932 when my
own work on Florida crayfishes began ten species had been recorded for Flor-
ida: affinis, fallax, lecontei, alleni, clarkii, versutus, acherontis, barbatus, ev-
ermanni, and clarkii paeninsulanus. Of these, one, affinis, must be deleted;
two others, clarkii and barbatus, should in all probability be now referred to
closely allied but distinct species,5 and a fourth, described under the trinomial
Cambarus clarkii paeninsulanus, is distinct from clarkii (Procambarus pae-
ninsulanus, Hobbs and Marchand, in press).
Since 1938 my own descriptions of Florida specimens have added 13 spe-
cies and subspecies to the published list of Florida crayfishes: 1940-Cam-
barus pallidus, C. lucifugus lucifugus, C. lucifugus alachua, C. hubbelli, C.
kilbyi, C. rathbunae, and C. pictus. 1941-C. cryptodytes, C. floridanus, and
C. byersi. I also have two papers which are now in press describing a new gen-
us, Troglocambarus, and three new species: T. maclanei, Procambarus pycno-
gonopodus, and P. leonensis.

FACTORS AFFECTING THE DISTRIBUTION OF CRAYFISHES
IN FLORIDA
FLORIDA AS A FAUNAL REGION
Cooke and Mossom (1929), Cooke (1925, 1939), Leverett (1931), and
Stubbs and Hubbell as quoted by Carr (1940) have discussed or summarized
what is known or conjectured about the geological history of Florida, and I
have drawn freely from them in my discussion of the possible migration routes
of the several invading crayfish stocks. A great difficulty lies in the dearth of
conclusions or of even a preponderance of evidence for the sequence of the
minor advances, retreats, and embayments of the gently rising and sinking
Florida coasts. That such events took place is evident enough, but the time
relationship between the coastal terraces and other minor changes is not clear;
nor is it definitely known whether the early Pleistocene submergences in-
volved all of the peninsula of Florida or whether certain islands in the penin-

'These specimens are near lucifugus lucifugus (Hobbs 1940: 393).
"The specimens from the Pensacola region referred to clarkii are herein described as a new
species, P. okaloosae. The Florida barbatus mentioned by Ortmann (1905a) is probably escambien-
sis. Specimens determined lecontei, as stated above, were probably confused with spiculifer or ver-
sutus.






10 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

sula region have had a continuous terrestrial existence since sometime in the
Pliocene. This much at least is certain, that Florida, as well as the adjacent
parts of Georgia and Alabama, is geologically recent and ecologically young,
and that the migration of crayfish into and within the state must have largely
taken place subsequent to early Pleistocene and is still in progress.
It seems best to qualify the recentness of migration with a "largely" be-
cause of the marked structural changes and distribution shown by the four
peninsular cavernicolous forms. St. John (1936), Hubbell (1939), and Carr
(1940) have presented good biogeographical evidence for the persistence of
an Ocala Island since late Pliocene, in spite of strong geological evidence that
it must have been submerged. Certainly the existence of such an island and
the survival there of an earlier wave of migration from which the cavernico-
lous forms, and probably Procambarus pictus as well, were derived would
give a far more comfortable margin of time for the development of the several
cavernicolous species that now occupy the site or periphery of the Ocala Is-
land. Indeed it is not unthinkable that adaptation to subterranean waters which
reached the "Island" by way of strata from the Piedmont fitted the crayfish to
utilize this freshwater refuge during a Pleistocene submergence, if Cooke's
contention is to be accepted.
On the other hand, cavernicolous adaptations may not require more time
than has been available since the end of the early Pleistocene. Certainly lime-
stone solution in Florida is rapid and extensive and must have begun soon af-
ter Florida appeared above the sea. In the light of the marked tendency to de-
velop spelean adaptations shown by the crayfish as a group, and in the light
of recent indications from genetics and Kinsey's conclusions (1929, 1936) on
evolution in Cynips it may well be that there has been sufficient time since
Pleistocene to provide for the invasion and adaptations of all the Florida
fauna.
Here in the southeastern coastal plain topographic relief, although slight,
is accentuated by an extensive variation in soil types; upland soils are irreg-
ularly interspaced with poorly drained flatwoods; extensive areas of swamps
and marsh-are broken or wholly segregated by well drained sands; and the re-
markable development of subterranean drainage with its development of cav-
erns, huge springs, sink holes, and thousands of solution ponds and lakes has
also in many regions accentuated the xeric condition of many wide extents of
surface soil.
The chief extensive areas of well drained soils that appear to have been
the most generally important barriers to migration are shown on Map 2. Other
local barriers as well as the chief highways for dispersal are best discussed
under the particular species or groups concerned.
DERIVATION OF THE FAUNA
The crayfish fauna of Florida comprises five genera which have had a
relatively long and complex history. According to Ortmann the Cambarinae






HOBBS-THE CRAYFISHES OF FLORIDA


(his genus Cambarus) "originated in Mexico, and immigrated, probably at
the beginning of the Tertiary, into the southwestern and southern United
States, originally occupying only the southwestern Cretaceous plain, the Ozark
Mountains, and the southern extremity of the Appalachian System."
He cites the following centers of origin for the various genera: Procam-
barus (Ortmann's subgenera Cambarus and Procambarus) originated in Mex-
ico and following the above mentioned path reached the foot of the Appalach-
ian Mountains in the lowlands of Alabama and Georgia where a "secondary
center" was developed. "Here the more advanced forms of this subgenus took
their origin, and spread all over the Atlantic and Gulf coast plain, and further
up the Mississippi valley" (Ortmann 1905a: 124).
Cambarellus (Ortmann's subgenus Cambarellus) "originated in the
southerrrn-Uited States and immigrated into Mexico, first into the central pla-
teau, then into the Pacific slope" (Ortmann 1906a: 24).
Orconectes (Ortmann's subgenus Faxonius) "developed in the central
basin oTtThIthree great rivers, spreading over almost all of the Mississippi
Drainage, and crossing over into the Hudson Bay, Great Lakes, and even into
the Atlantic drainages, probably by the aid of shifting divides" (Ortmann
1905a: 24).
,Cambarus (Ortmann's subgenus Bartonius) came into existence "in the
mountainous region of the southern Appalachians, probably including also the
Ozark region, and from here it spread chiefly over the Appalachian chain in a
northeasterly direction as far as New Brunswick" (Ortmann 1905a: 124).
The fifth genus, Troglocambarus, probably had its origin in the caverni-
colous waters of the Florida peninsula, and is possibly of much more recent
origin than the others.
Some representatives of Procambarus or its progenitors migrating into
the southeast pushed southward and then followed the Gulf coast as the sea re-
treated, so that when the elevation of Florida left vast stretches of poorly
drained lowlands, traversed by numerous streams, a broad highway was
opened to them. Thus the Procambarids reached Florida from the north along
several paths. These paths are pointed out below in the section devoted to Prin-
cipal Migration Routes.
The genus Cambarellus is represented in Florida by only one species,
which occupies a large range in the panhandle and south and east as far as
the drainage of the Suwannee River. This stock undoubtedly came into Flori-
da from the west, making its way along streams and through the coastal flat-
woods.
Likewise only one species of the genus Orconectes occurs in Florida, and
the available data indicate that this species is a migrant from the west, which
has taken a path similar to that followed by Cambarellus. However, Orconectes
took a more northern route and penetrated into the state only in the West Flor-
ida Lime-Sink or Cypress Pond Region (Harper 1914: 201).






12 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

The genus Cambarus reached Florida along at least three different
routes: one of them from the west along the coastal region, the others from the
north, one in the region of the Escambia River, and the other along the Apa-
lachicola River. None of the members of this genus are found east of the Och-
locknee River in Gadsden County.
Several authors, notably, Hubbell (1940), Rogers (1933), Carr (19-
40), and Kurz (1933), have been impressed by the fauna and flora of the
remarkable ravines along the east bank of the Apalachicola River in Liberty
and Gadsden counties. These authors have pointed out that several animals
and plants which are common in the Appalachian and Piedmont regions reach
the southern limits of their ranges in this area, and in addition that several
forms, endemic in the ravines, have their closest affinities with northern
groups. At least two plants, Tumion taxifolium and Taxus floridana, are prob-
ably Tertiary relicts, while the katydid Hubbellia marginifera (Walker) has
relatives only in Europe, the Near East, and Madeira. Carr (1940: 4) pre-
sents a very plausible explanation of why the Apalachicola River supports
such a large assemblage of animals and plants having their closest affinities
with Appalachian and Piedmont species.

Endemism
Twenty-five of the 42 Floridian Cambarinae are, as far as known, en-
demic to the state. After thorough collecting is done in southern Alabama this
probably will be reduced to not more than 17. Of these 17 six are cavernicolous
forms.
The following are now recorded only from Florida but probably occur in
southern Alabama: P. escambiensis, P. rathbunae, P. hubbelli, P. shermani,
P. okaloosae, P. pycnogonopodus, P. bivittatus, C. species incertis. The follow-
ing 17 are probably confined to Florida: P. apalachicolae, P. econfinae, P.
latipleurum, P. kilbyi, P. alleni, P. geodytes, P. rogersi rogersi, P. rogersi
campestris, P. rogersi ochlocknensis, P. acherontis, P. young, P. pictus, P.
lucifugus lucifugus, P. lucifugus alachua, P. pallidus, C. cryptodytes, T. mac-
lanei.
The Floridian cavernicoles are representatives of at least four indepen-
dent advances into the subterranean waters. Three of these were probably cor-
related with the early insular development in central Florida. It seems most
probable that the stock which gave rise to Procambarus acherontis reached the
Pliocene islands at an early time but was destroyed by subsequent emergence
leaving acherontis as a relict in the subterranean waters of the area. The cav-
ernicolous species of the pictus group (P. lucifugus lucifugus, P. lucifugus
alachua, P. pallidus) and Troglocambarus maclanei (the latter probably of
earlier origin) have evidently been derived from a stock of stream dwellers
which occupied the southeastern coastal plain and migrated into the peninsula
when the Suwannee Straits were closed. The only spelean species of Cambarus
(C. cryptodytes) found in Florida is probably a derivative of an earlier Ap-






HOBBS-THE CRAYFISHES OF FLORIDA


palachian and Piedmont stock which moved southward along the Apalachicola
River.
At least two surface dwelling species of Procambarus are probably relicts
of a once widely distributed stock. These are P. alleni, occupying an extensive
area in the southern part of the peninsula, and P. pictus, confined to a few
small tributaries of the St. Johns River in Clay County. The latter is particu-
larly interesting in that its closest affinities are found among the peninsular
cavernicoles.
The other endemics are derived from three stocks which have apparently
migrated into the region in more recent times. One of these, the barbatus stock,
(illustrated on Map 3) reaching Florida along several migration paths, has
given rise to four species in the panhandle: P. apalachicolae, P. econfinae, P.
latipleurum, and P. kilbyi. The pictus stock, which probably had its origin in
the southeastern part of Georgia, migrated southwestward along the Ochlock-
nee River system into the coastal flatwoods, crossed the Apalachicola and gave
rise to P. young, which appears to be extremely localized in a few small
streams around Weewahitchka. The advena stock following the same route
produced P. pygmaeus and the three subspecies of P. rogersi.
Principal Migration Routes
I believe there is sufficient data to postulate the principal migration ,
routes by which the crayfishes have reached Florida and I have indicated the
species, or their forerunners, which appear to have utilized each route.
I. Early Migrations During Former Bridgings of the Suwannee Straits
It seems probable that the Straits were reopened at least twice after insu-
lar Florida was first connected with the mainland. The first openings of the
Straits with the probable subsequent submergence of the whole peninsula
would provide for the origin of the cavernicolous P. acherontis. The second
bridging of the Straits provided for the entrance of the stocks that gave rise to
T. maclanei, P. pictus, and the subterranean species: P. lucifugus lucifugus
and P. lucifugus alachua. Perhaps the alleni stock made its way into the penin-
sula at the same time. These stocks were thus left to occupy the Florida island
after the third opening of the Straits barred it for a time to further immigra-
tion.
II. Migrations Since the Last Closing of the Suwannee Straits
1. Migration Southward in the Flatwoods Region along Trail Ridge.
Four Florida species have apparently entered or been derived from stocks that
entered the state along this path. These are the endemic P. geodytes, and P.
seminolae, P. pubischelae, and P. advena.
2. Immigrations Via River Systems and Adjacent Flatwoods.
(a) Perdido River: P. escambiensis, P. evermanni (?), P. spiculifer,
P. versutus, and C. byersi.






14 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

(b) Escambia River: P. bivittatus, P. shermani, P. blandingii acutus,
P. evermanni, P. okaloosae, P. spiculifer, P. versutus, and C.
species incertis.
(c) Apalachicola and Chipola rivers: C. latimanus, C. floridanus, P.
apalachicolae, P. latipleurum, P. spiculifer, and P. versutus.
(d) Ochlocknee River which acts as a connecting link between the
lowland of southeast Georgia and the Apalachicola flatwoods:
P. young, P. pygmaeus, P. rogersi rogersi, P. rogersi ochlock-
nensis, P. rogersi campestris, and P. kilbyi.
(e) Choctawhatchee River and the Marianna Lowlands: P. hubbelli
and 0. clypeata.
Several of the stream inhabitants are so widely disseminated through-
out the streams of the state that I cannot reconstruct their paths of entry into
the Florida waters with any feeling of certainty, and indeed a number of these
stream inhabitants have undoubtedly entered the state by several paths.















Map 2.--Continuous North-South Extents of Well Drained Soils that appear to act as
Barriers to Crayfish Migration.


CONTINUOUS NORTH-SOUTH EXTENTS OF WELL-DRAINED SOILS THAT APPEAR
To ACT As BARRIERS TO CRAYFISH MIGRATION
(Explanation to Text Map 2)

A. (Norfolk-Greenville, and Norfolk-Orangeburg Soil Areas)
Western barrier to: P. paeninsulanus, P. hubbelli, P. pycnogono-
podus, P. apalachicolae.
Eastern barrier to: P. byersi, P. okaloosae, P. evermanni (part).






HOBBS-THE CRAYFISHES OF FLORIDA


B. (Norfolk-Greenville, and Norfolk-Orangeburg Soil Areas)
Western or northern barrier to: P. rogersi rogersi, P. latipleurum,
P. kilbyi (part).
Eastern barrier to: P. hubbelli (part), P. latipleurum.
C. (Norfolk-Greenville, Blanton-Norfolk, and Norfolk-Orangeburg Soil
Areas, reinforced at the south by the broad mouth of the Apalachi-
cola)
Western barrier to: P. rogersi ochlocknensis, P. leonensis.
Eastern barrier to: P. seminolae, P. pycnogonopodus, P. young.
D. (Blanton-Norfolk Soil Areas)
Western barrier to: P. fallax, P. alleni (?).
Eastern barrier to: P. leonensis.
E. (Blanton-Norfolk, and Fellowship-Gainesville Soil Areas)
Southern barrier to: P. pubischelae, P. seminolae, P. pygmaeus
(part), P. advena.
Northern barrier to: P. alleni.
Eastern barrier to: C. schmitti, P. spiculifer (part).
Western barrier to: P. pictus.

Species That Traverse Such Barriers or Occur on Either Side of Them

A number of species are found on both sides or within the limits of the
barriers in question. Their means of traversing them are indicated by: N-the
present range appears to extend north of the northern limits of the barrier;
S-the range has been extended into coastal flatwoods south of the southern
extent of the barrier; ?-means of traversing barrier definitely unknown.


Barrier A-
P. rathbunae-? (N)
P. spiculifer-N
P. versutus-? (N)
C. schmitti-?
C. diogenes-N

Barrier B-
P. paeninsulanus-?
P. pycnogonopodus-S
P. spiculifer-N
P. versutus-?
C. schmitti-?
0. clypeata-N
C. diogenes-N


Barrier C-
P. kilbyi-S
P. paeninsulanus-?
P. pygmaeus-S
P. rogersi (complex)-S
P. spiculifer-N
C. schmitti-?
Barrier D-
P. kilbyi-S
P. paeninsulanus-?
P. spiculifer-N
C. schmitti-?
Barrier E-
P. kilbyi-S
P. fallax-?
P. paeninsulanus-?
P. spiculifer (part)-N






16 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

ECOLOGY
In general, it may be said that crayfishes are found in any body of fresh-
water and in almost any poorly drained soil. Small local areas of apparently
suitable soils are occasionally found to be unoccupied, but these areas will
usually prove to be completely surrounded by well drained soils and thus iso-
lated from any natural stocking.
In attempting to classify and describe the various types of situations in-
habited by the several species of crayfishes, one is confronted with the diffi-
culty that the actual distinctions between the situations occupied by different
species are complex, and often not at all easy to discern. The differences and
likenesses between situations that are most apparent to the student are by no
means always the distinctions that appear to be most important to the crayfish.
Nevertheless the desirability of attempting to describe and classify habitats in
terms that can be recognized by the field naturalist is very real. In the discus-
sion that follows, the ecological subdivisions are frequently more detailed than
the actual habitat discrimination shown by the crayfishes would warrant and
result in many species being listed in two or more distinct situations. So de-
tailed a subdivision, however, at least facilitates the generalizations that are
necessary if one is to attempt a summary of the habitat correlations of the
whole crayfish fauna.
Most of the difficulty in defining a clear-cut ecological distribution of
the whole fauna is due to two circumstances: that Florida comprises several
regions that are actually quite diverse in topography, soil, and drainage, and
that for so many species the geographic range does not coincide with any major
ecological subdivision of the state.

AQUATIC HABITATS
Lotic Situations
Several large rivers, such as the Apalachicola and Choctawhatchee, which
cross the panhandle of Florida have their origins in the agricultural sections
of Alabama and Georgia and carry heavy loads of silt and clay which give
them the characteristic red color of streams that rise in the Piedmont. For the
most part the beds and margins of these streams are nearly barren of aquatic
plants, but extensive hardwood swamps are developed in their flood plains.
Such lower streams undoubtedly have a larger crayfish fauna than is indicat-
ed by the number of specimens recorded from them, but I believe that they do
not support so abundant a fauna as their smaller tributaries. However, the dif-
ficulties of collecting have prevented as thorough a survey of the large lower
streams as has been possible in the smaller water courses.
Several of the smaller rivers, for example, the Chipola and Santa Fe, are
partly spring-fed and flow through lime rock outcrops along part of their
courses. Only after heavy rains do these streams become turbid or take on the
dark red or brown color of swamp water. In places they flow over limestone






HOBBS-THE CRAYFISHES OF FLORIDA


outcrops and produce shoals and riffles, and in many of these such aquatic
vegetation as Vallisneria, Sagittaria, Philotria, Riccia, and Chara is abundant.
A more complex description of this type of stream is given on page 123. Al-
though the number of species of crayfishes is small, the population is dense,
especially in the riffles and vegetation zones.
The Hillsborough, Manatee, and Withlacoochee rivers represent another
class of the larger streams. These rivers arise in the swampy lake regions of
the interior of the peninsula, and for the most part are devoid of aquatic plants.
In some of the broader areas, however, Nymphaea, Piaropus, and other aqua-
tics associated with lenitic situations are abundant, and in shallow reaches
some of these streams gain momentum and form riffles which support as lux-
uriant a flora as that of the spring-fed rivers of the panhandle and northern
Florida. The crayfish population in the shallow and broad regions of these
streams is extremely large.
The flatwoods streams of the panhandle and northern Florida are numer-
ous and in general have many characters in common. Most of them are small
and relatively shallow, supporting dense growths of Myriophyllum, Potamoge-
ton, Juncus, Pontederia, Sagittaria, and filamentous algae. Most of these
streams are sluggish, with dark water indicating an abundance of organic
acids. Many are semi-permanent and flow only part of the year. The crayfish
population reaches a maximum in this type of stream. If the water ceases to
flow and the stream begins to dry up the crayfishes burrow or live in the deep-
er pools where the water is more permanent.
Scattered throughout the state, particularly in the rolling uplands, many
small, clear, sand bottomed streams occur. This type of stream is abundant
along the north shore of the Choctawhatchee Bay, in the rolling country east
of Crestview, Okaloosa County, and in the Central Highlands. These streams
originate in seepage areas and small springs and flow through sandy and clay
beds over most of their course. Aquatic vegetation consisting of Orontium,
Nymphaea, and Potamogeton is sparsely scattered along the margins. If one
collects in these streams during the day when the crayfishes have retreated in-
to their burrows in the banks or remain concealed in the roots of the marginal
semiaquatic vegetation, he is left with the impression that the crayfish fauna
is poor; however, at night the animals appear in large numbers.
Numerous large springs are present throughout the state. Most of them
flow into large basins which support a rich aquatic flora that includes Nym-
phaea, Riccia, Cabomba, Potamogeton, Isnardia, Cardamine, Myriophyllum,
Utricularia, Chara, and filamentous algae. Many of these springs (viz., Blue
Springs, Jackson County, Wakulla Springs, Wakulla County, Silver Springs,
Marion County, etc.) form the origin of long "spring runs" where aquatic
plants, especially Vallisneria, Naias, Sagittaria, and Ceratophyllum are abun-
dant. Crayfish are numerous in the vegetation both in the springs and in the
runs. A number of the large springs in the peninsula flow from or through sul-
fur- and salt-bearing strata and deposit dense, cream-colored sludges and pre-






18 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

cipitates on the vegetation in them. Even here Procambarus fallax is often com-
mon.
In the deep ravines along the east side of the Apalachicola River in Liber-
ty County, small streams that have their origins in seepage areas and small
springs flow over barren sand and clay bottoms and are often dammed by fal-
len limbs and leaf drifts. These streams are all well shaded and cool and sup-
port a surprisingly rich crayfish fauna.

Lenitic Situations
Temporary ponds are very common in all parts of the state, both in the
rolling uplands and in the flatwoods areas. In both regions the ponds vary
widely as to the time and regularity of becoming dry. Some are dry except
during the rainy season, and seldom have a crayfish population; others which
become dry for only a short period during the year, support a large popula-
tion. When these ponds do become dry the crayfishes construct burrows. Some
of these temporary ponds are within dense swamps and are barren of aquatic
plants, while in the more open ponds Sagittaria, Panicum, Juncus, Utricularia,
and filamentous algae are often abundant.
Permanent but fluctuating ponds are also common in most regions of the
state. Such ponds rise and fall markedly with the rainy and dry seasons and
usually have an abundance of rooted and floating vegetation. Some of the
commoner plants are Nymphaea, Castalia, Piaropus, Pontederia, and Cerato-
phyllum. These ponds usually support a very large crayfish population.
Deep, well-like sink-hole ponds are common in certain regions of Florida
and many are inhabited by crayfish, but ponds of this type do not have nearly
so abundant a crayfish fauna as the shallower ponds which support a more
luxuriant growth of hydrophytes.
In the larger lenitic situations the crayfish population appears very def-
initely to be correlated with the kind and amount of rooted aquatic vegetation.
Although from the standpoint of the limnologist the lakes of Florida fall into
several rather distinct types, ranging from the deeper, clear lakes of the sand
hill region to the shallow and usually acid and discolored waters of the flat-
woods lakes, for the crayfish it appears to be only the character of the shore-
line that is important.
The deep and often extensive Nymphaea-Nelumbo-Utricularia marshes,
that are locally known as "Prairies", are common in many parts of Florida.
They usually have a depth of about six feet or less, and almost the entire area
is crowded with a dense growth of both emergent and submergent vegetation,
with Nymphaea, Piaropus, Nelumbo, Typha, Juncus, and grasses especially
common. Here a few species of crayfishes are often extremely abundant.
Roadside ditches and canals in which water stands most of the year also
develop a rich growth of submergent, emergent, and floating aquatic plants.
Commonly found in these plant associations are Myriophyllum, Ceratophyl-






HOBBS--THE CRAYFISHES OF FLORIDA


lum, Potamogeton, Utricularia, Persicaria, Ludwigia, Pontederia, Sagittaria,
Nymphaea, Castalia, Piaropus, Globifera, and grasses. These ditches which
form a very typical and characteristic part of much of the Florida landscape
are one of the most productive of all crayfish habitats.
In the everglades freshwater marshes are extensively developed with saw
grass as the characteristic dominant. Although there are numerous pot holes
which dot the rock ridge bounding the everglades along the east coast, the most
extensive bodies of fresh water are the abandoned rock pits and the extensive
canals. In these P. alleni often occurs in large numbers.
Subterranean Situations
Much of both the peninsular and panhandle regions have extensive un-
derground water systems, which in many cases have been exposed by the for-
mation of sink holes and small limestone caverns. In a number of places where
these cavernicolous waters are accessible various spelean species of crayfishes
have been taken. These underground waters, at least throughout the peninsula,
show very little variation in either temperature or pH, being around 700 and
7.1 respectively. Data on the temperature and pH of the underground waters
of the panhandle is very meager but indicate that these waters are very close to
those of the peninsula. One subterranean species occurs in a sulfur spring
(Palm Springs) near Orlando, in Seminole County. A description of this
spring is given on p. 92.

Flatwoods and Seepage Areas
Perhaps the most interesting members of the crayfish fauna of Florida
are the burrowing species, which are largely confined to the flatwoods and
seepage areas. Extensive areas of flatwoods occur in the coastal regions and
portions of the interior. Here the water table in many places is almost at the
surface, and permanently wet areas which support a large number of semi-
aquatic plants are common. Conspicuous among these plants are Sarracenia,
Ericaulon, Xyris, Drosera, Pinguicula, Frimbristylis, Rynchospora, and a
large variety of grasses. The soils of these wet areas usually belong to the
Plummer and Portsmouth groups. In some of these areas the ground is literal-
ly riddled with crayfish tunnels, and chimneys are a conspicuous feature of the
surface. In much of the flatwoods region characterized by Leon soils the water
table periodically descends much farther below the surface, but only occa-
sionally does it drop to a depth greater than three or four feet. While the cray-
fish are not so abundant in Leon soils they are present in large enough num-
bers to be classified as common. Large colonies of many of the flatwoods spe-
cies are often found burrowing in roadside ditches; particularly is this true
in the vicinity of the numerous flatwoods ponds or bays.
Seepage areas along hillsides or stream slopes are, particularly in the
panhandle, usually thickly populated by burrowing crayfishes. The hillside
conditions are for the most part not markedly different from those found in






20 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

the wet flatwoods areas, but in the seepage areas along streams the crayfishes
construct a maze of intricate tunnels among the thick tangle of tree and shrub
roots. The soils in these seepage areas show considerable variation, although
most of them are dark and contain a high concentration of decayed organic
matter.

ECOLOGICAL HABITS AND HABITATS

A-Species That Occur In Burrows
These burrowing forms show three more or less distinct degrees of
burrowing habit, indicated by: P-primary burrowers (restricted
to burrows); S-secondary burrowers (generally occupying bur-
rows but wandering into open water during rainy seasons) ; T-ter-
tiary burrowers (burrowing only in periods of drought or occasion-
ally, but not necessarily, during the breeding season).

I. Burrowing in Seepage Areas
P. advena-P P. rogersi ochlocknensis-P
P. geodytes (around sulfur C. latimanus-S
springs)-P C. floridanus-P
P. rogersi rogersi-P C. diogenes-P or S
P. rogersi campestris-P C. byersi-P
II. Burrowing in Flatwoods
P. advena-P P. kilbyi-S
P. pygmaeus-? P. hubbelli-S
P. rogersi rogersi-P P. alleni-S
P. rogersi campestris-P P. seminolae-S-T
P. rogersi ochlocknensis-P P. paeninsulanus-T
P. pubischelae-S P. okaloosae-T
P. escambiensis-S P. fallax-T
P. econfinae-S P. leonensis-T
P. latipleurum-S P. pycnogonopodus-T
P. apalachicolae-S 0. clypeata-T (?)
P. rathbunae-S C. byersi-P

III. Burrowing in Floodplains of Large Streams
C. floridanus-P C. species incertis-S (?)
C. diogenes-P-S P. bivittatus-T
P. shermani-S P. blandingii acutus-T


IV. Burrowing in Banks of Streams
P. hubbelli-S
P. kilbyi-S
P. paeninsulanus-T


P. spiculifer-T
P. versutus-T






HOBBS-THE CRAYFISHES OF FLORIDA


V. Burrowing in Drying Ponds
P. alleni-S
P. hubbelli-S
P. kilbyi-S
P. latipleurum-S
P. seminolae-S-T


P. fallax-T
P. leonensis-T
P. paeninsulanus-T
P. pycnogonopodus-T


B-Species Occasionally or Usually Found Outside of Burrows
I. Inhabiting Ponds, Lakes, and Ditches. L-permanent lakes or
ponds; T-temporary or fluctuating ponds; D-semipermanently
inundated ditches.


P. okaloosae-D
P. pubischelae--D
P. pycnogonopodus-D
P. rathbunae-D
P. econfinae-D-T
P. kilbyi-D-T
P. latipleurum-D-T


P. seminolae-D-T
P. alleni-T-D
P. hubbelli-T-D
0. clypeata-T-D
P. paeninsulanus-L-T-D
P. fallax-L-T-D
P. leonensis-L-T-D


II. Inhabiting More or Less Lotic Situations. A-rills and ravine
brooks (of Apalachicola drainage); I-intermittent streams; B-
brooks, creeks, and small rivers.


C. latimanus-A
P. kilbyi-I
P. okaloosae-I
0. clypeata-I
P. blandingii acutus-B
P. pictus-B
C. schmitti-B
P. spiculifer-B
III. Inhabiting Sluggish Streams and Slou
P. kilbyi
P. hubbelli
P. alleni'
P. fallax
P. leonensis
P. pycnogonopodus


P. versutus-B
P. bivittatas-I-B
P. fallax-I-B
P. leonensis-I-B
P. paeninsulanus-I-B
P. pycnogonopodus-I-B
P. seminolae-I-B

ghs
P. bivittatus
P. paeninsulanus
P. okaloosae
P. evermanni
P. seminolae
0. clypeata


C-Species Confined to Subterranean Waters


P. acherontis
P. lucifugus lucifugus
P. lucifugus alachua


'Found extensively in drainage canals of southern Florida.


P. pallidus
T. maclanei
C. cryptodytes





SUMMARY OF THE GENERIC AND SUBGENERIC CHANGES
(1870-1941) (Hobbs, in press-c)


Hagen 1870 Faxon 1885 Ortmann 1905-6 Fowler 1911 Faxon 1914 Creaser 1933 Lyle 1938 Hobbs (in press-c)
(Generic Names)


Procamnbarus


Group I


Group II


Procambarus Group I


rGroun II


Girardiella Procambarus


.-------_ ___ Paracambarus

Group V Cambarellus

Group IV Faxonius

Group III Bartonius


Paracambarus Group IV

Cambarellus Group V

Faxonius Group VI

Cambarus Group VII


Faxonius, as a
generic name.


Paracambarus

Cambarellus

Orconectes

Cambarus


------------------ .- ------- ... .. .... ----- Troglocambarus


Group III,
in part

Group III,
in part

Group I,
in part


Group II

Group III,
in part






HOBBS--THE CRAYFISHES OF FLORIDA


THE SUBFAMILY CAMBARINAE
All of the North American crayfishes east of the Rocky Mountains be-
long to the subfamily Cambarinae of the Family Astacidae. As here recognized
this subfamily comprises six genera. One of these, the monotypic Paracam-
barus, is found only in Mexico; all of the other five occur in the United States
east of the Rocky Mountains, and of these, two extend northward into Canada;
one is found in Cuba and Mexico, and another in Mexico. Florida has at least
one representative of each of these five genera. One of them, Troglocambarus,
is endemic; Procambarus is represented by 33 species and subspecies, Troglo-
cambarus by one, Cambarellus by one, Orconectes by one, and Cambarus by
six. The chart on page 22 presents the progression of taxonomic distinctions
within this subfamily made by various students since 1870.
Definition.-The first abdominal segment of the male bears a pair of
variously shaped appendages with from two to five terminal elements, one of
which always serves as a sperm conduit [the corresponding appendages of
the female are reduced or absent]. The podobranchiae of the second and third
maxillipeds and of the first three pairs of pereiopods possess "a broad bilobed
plaited lamina; the epipodite of the first maxilliped is destitute of branchial
filaments; the coxopoditic setae are acute, not hooked, at the end; the telson
is commonly divided more or less completely by a transverse suture" (Faxon
1885a: 2). Gills are absent on the last thoracic somite, and there is no bilobed
lamina on the podobranchiae of the fourth pair of pereiopods.
THE EVALUATION OF TAXONOMIC CHARACTERS IN THE CAMBARINAE
Considerable emphasis has in the past been given the presence or absence
of hooks on the ischiopodites of the second, third, and fourth pereiopods of
the male in arriving at taxonomic groupings. These hooks are used by the male
in copulation, and are much more strongly developed in the breeding individ-
uals. While this character is extremely useful in many instances, it cannot
always be taken as indicative of kinship. Procambarus pubischelae, for exam-
ple, shows marked variation in this regard. In certain portions of its range
the individuals have hooks on the ischiopodites of the third and fourth pereio-
pods, while in other regions hooks are present on only the third pair. While so
extreme a variation is known only in the case of this one species, there are sev-
eral instances in which it is approximated. The following may be taken as an
example. Procambarus geodytes which is beyond any doubt closely allied to
P. advena (hooks present on the ischiopodites of only the third pereiopods)
has hooks on the ischiopodites of the third and fourth pereiopods, and if some
of the former groupings (Faxon 1914, Groups II and III) were followed, this
species would be entirely separated from advena, and placed in a group where
it very obviously does not belong. Both Ortmann (1905a: 95) and Creaser
(1931b: 10) have pointed out the independent development (or primitive re-
tention) of hooks on the ischiopodites of the fourth pereiopods in certain spe-
cies, the relatives of which have hooks on the third pereiopods only.






24 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2


dacryl


pa/m o fAand


'r/4.,


'ee* &?
-whio1pod/floe


Text Figure b.-Terminology and Methods of Taking Measurements. LH-length of
outer margin of hand; LD-length of dactyl; LP-length of palm; LCa-length of
carapace; LCe-length of cephalic section of carapace; LR-length of rostrum;
LA-length of areola.






HOBBS-THE CRAYFISHES OF FLORIDA


Other characters that have been appealed to for establishing species
groups appear to be clearly adaptive rather than phylogenetic. The length-
breadth ratio of the areola shows a definite correlation with the type of habitat,
and in some instances is probably as indicative of habitat as of relationship.
For instance most of the crayfishes that occur in flowing water (presumably
with a high oxygen content) have a broader and shorter areola, and hence a
smaller gill chamber, than do those living in stagnant waters or in lenitic sit-
uations. This correlation of gill chamber and habitat reaches its greatest de-
velopment in the primary burrowers where the areola tends to be extremely
narrow or obliterated. Again, the spines on the rostrum, postorbital ridges,
and along the cervical groove tend to be reduced or obsolete in the burrowing
colonies of species which have more strongly developed spines in populations
living in open water. P. alleni and P. paeninsulanus are excellent examples.
Other conspicuous structural characters such as the shape of the chelae and
body proportions show evidence of this same adaptive modification.
The first pleopod of the male and the annulus ventralis of the female
have proved to be the most reliable of all taxonomic characters, for they ap-
pear to show little variation within the species and to be little affected by adap-
tation to environment.
The adult male crayfish exhibits two distinctly different and usually al-
ternating morphological forms. Previous authors have referred to these as the
"first" and "second" form males or as "form I" and "form II", respectively.
These two forms are definitely associated with the reproductive cycle. The
"first form" is in the breeding stage and may be distinguished by the presence
of the corneous condition of the well defined terminal processes of the first
pleopod, while in the "second form" male the terminal processes of the first
pleopod are not so well defined, are usually blunt, and are never corneous.
Moreover, the hooks on the ischiopodites of the pereiopods are strongly de-
veloped and sometimes corneous in the first form, whereas in the second form
they are usually reduced and seldom corneous.
The new subdivisions and groupings made necessary by the rich acces-
sions of Florida species have consequently been based upon the structure of
the first pleopod and the annulus ventralis supplemented by such special char-
acters as have proved to be apropos and useful in grouping or distinguishing
between the various species.

THE CHIEF CHARACTERS USED IN KEYS AND DESCRIPTIONS

The first pleopod of the male (Text Fig. c) is of paramount taxonomic
importance, and it is necessary to clearly differentiate and distinguish its
parts. The anatomical position which is assumed in the following discussion
is with the distal end of the appendage extending ventrad. When the appendage
is in resting position, i. e., held against the thorax, the side in contact with the
thorax is the cephalic surface, and the side viewed in ventral aspect is the cau-






26 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2










n "









C













Text Figure c.-Hypothetical First Pleopod of Male. C-caudomesial margin; I-me-
sial process; II-cephalic process; III-centrocaudal process; IV-caudal process;
V-Centrocephalic process; III and V together make up the central projection.
. o










V--Centrocephalic process; III and V together make up the central projection.






HOBBS-THE CRAYFISHES OF FLORIDA


dal surface. The other anatomical directions are obvious when the caudal and
cephalic directions are established.
Although there are considerable differences in forms and proportions of
the terminal processes of the various species they can all apparently be homol-
ogized with a primitive four-parted arrangement (Hobbs, in press-a). Distal-
ly the appendage is made up of a rolled plate-like structure which has spines,
tubercles, or plate-like projections extending from its distal edge and has its
lateral edges folded tightly against the middle portion. The margin "C" is on
the caudomesial face of the appendage, and the mesial process "I" extends
distad from this margin. As one follows around the perimeter of the append-
age he finds a spine projecting from its cephalodistal margin; this is the ce-
phalic process "II"; another process, the centrocaudal "III", is given off from
the distolateral margin, and it makes up the caudal part of the central projec-
tion. The caudal process "IV" projects from the caudolateral, caudal, or cau-
domesial margin of the appendage, and the cephalic part of the central pro-
jection, the centrocephalic process "V", extends from the inner edge of the
roll and projects distally from the center of the appendage.
The terminal processes show considerable variation. Sometimes they are
well developed, sometimes one may be reduced, and occasionally one or more
may be absent. There is some shifting of the positions of these processes; for
example, in most of the members of the barbatus (Plates II and III) group
the cephalic process arises from the cephalomesial margin of the appendage
(Hobbs, in press-a).
The rostrum also shows considerable variation. It may bear lateral
spines, the margins may be interrupted without bearing spines, or they may be
smooth with no indication of a break between the acumen and the rest of the
rostrum. The subrostral ridges are the swollen ridges along the ventrolateral
margins of the rostrum, and in some species they extend forward to form the
cephalic tip while the rostral ridges fade out before reaching the tip.
The length of the carapace (Text Fig. b) is taken from the tip of the ros-
trum to the caudal margin of the areola. The length of the areola is taken along
the middorsal line, and its width taken at the narrowest part.
There are five pairs of pereiopods (walking legs) of which the chelipeds
are the first pair. In the male, hooks are present on the ischiopodites (third
segment from the proximal end) of the second and third, third, or third and
fourth pereiopods.
The secondary sexual characters of the male which are not mentioned
when contrasting the two sexes in description consist of the presence of the
above-mentioned hooks on the ischiopodites of the second, third, and fourth
pereiopods, and the outgrowths from the bases of the coxopodites proximall
segments) of the fourth and fifth pereiopods.






28 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

KEY TO THE FLORIDA CRAYFISHES
(Based on the first form male)
1 Ischiopodite of third maxilliped without teeth on mesial margin.
(Only one species of this genus known) -7 Troglocambarus
maclanei Hobbs __---------------------- p. 146
1' Ischiopodite of third maxilliped with teeth on mesial margin-- 2
2 (1') First pleopod of male with two terminal apices that are either
straight, divergent, or gently curved; never bent so much as at
a right angle to the main shaft. (Only one species, which belongs
to the monotypic subgenus Faxonella, known in Florida)_
Orconectes clypeata (Hay) -- ------ ---- p. 154
2' First pleopod of male terminating in two or more parts; if two,
these are bent caudad at right angles to the main shaft .----. 3
3 (2') Hooks on ischiopodites of second and third pereiopods of male.
(Only one species of this genus known in Florida) _____-Cam-
barellus schmitti sp. nov. ---------------p. 149
3' Hooks on ischiopodites of third, or third and fourth pereiopods_4
4 (3') First pleopod of male terminating in two blade-like or bulbous
processes that are always bent at right angles to the main shaft
..---- .. Genus Cambarus ------ ---..------------_ 40
4' First pleopod of male terminating in three or more processes that
are corneous, spiniform, plate-like, or blunt ---- Genus Pro-
cambarus ---- ---------_ ------ -----5
5 (4') Hooks usually present on ischiopodites of third pereiopods only;
if present on both third and fourth, then margins of rostrum
uninterrupted (interrupted in some specimens of alleni, an oc-
casional specimen of latipleurum, and all specimens of the
cavernicolous acherontis; in alleni and acherontis the areola is
long and narrow, and the hooks on the ischiopodites of the fourth
pereiopods are bituberculate; in latipleurum the spines on the
rostrum are minute and almost at the apex of the rostrum)--- 6
5' Hooks on ischiopodites of both third and fourth pereiopods; mar-
gins of rostrum interrupted and often bearing lateral spines
____------ (Blandingii Section) ------ ---_----- ---24
6 (5) Eyes reduced; carapace without pigment (Acherontis Section with
a single species) .----- P. acherontis (L6nnberg) -----.. p. 91
6' Eyes well developed; carapace pigmented ------------7
7 (6') Chela flattened; inner margin of palm bearing a single, well de-
fined, cristiform row of tubercles_ ..--- (Advena Section) -19
7' Chela subovate, not strongly compressed; inner margin of palm
never bearing a single cristiform row of tubercles, either bar-
bate or bearing several irregular rows of tubercles which are
usually small___ -- (Barbatus Section) _- ---_--- ---8
8 (7') Hooks on ischiopodites of third and fourth pereiopods; hooks on
fourth bituberculate (Alleni Group with a single species)_.._-.
P. alleni (Faxon) __-- ____ ____.----------- p. 69






HOBBS--THE CRAYFISHES OF FLORIDA


8' Hooks on ischiopodites of third, or of third and fourth pereiopods;
hooks on fourth simple ..-- (Barbatus Group) .------ 9
9 (8') Hooks on ischiopodites of third and fourth pereiopods ..____- 10
9' Hooks on ischiopodites of third pereiopods only ------------___17
10 (9) Palm of chela barbate along inner margin ------_--16
10' Palm of chela not barbate along inner margin .-------_--- 11
11 (10') Cephalic process of first pleopod arising from cephalic surface
_-------- P. shermani sp. nov....----- --...------- .....-.---.. p. 61
11' Cephalic process of first pleopod arising from mesial surface _12
12 (11') Mesial process of first pleopod conspicuously large, subspatulate,
bent caudad at an angle of 45-90 degrees to the main shaft of
the appendage -------- P. kilbyi (Hobbs) ----- -- p. 64
12' Mesial process of first pleopod never subspatulate; directed dis-
tad or caudodistad but not so much as at a 45 degree angle to
the main shaft_- ..-- ---.---------- -----. 13
13 (12') Caudal process of first pleopod thumb-like, or if not thumb-like
the cephalic process curved caudad -___-.. ____ --__14
13' Caudal process of first pleopod not thumb-like, forming a later-
ally compressed blade-like structure along the caudolateral tip
of the appendage; cephalic process straight or curved cephalad

14 (13) Cephalic process of first pleopod straight; mesial process acute
and directed caudodistad_ ----__-P. apalachicolae sp. nov. p. 55
14' Cephalic process of first pleopod curved caudad; mesial process
subspiculiform and directed distad but curved cephalad .----
P. latipleurum sp. nov .....-----.--.._ -----------------p. 52
15 (13') Coxopodite of fourth pereiopod with no prominent outgrowth;
outgrowth on that of fifth pereiopod simple. Opposable mar-
gin of dactyl of chela with only the faintest indication of an
excision. Inner margin of palm of chela never barbate--- -
P. econfinae sp. nov. ...._..---.--------------- ------- p. 49
15' Coxopodite of fourth pereiopod with a single large tubercle; out-
growth on that of fifth with two or three emarginations. Oppos-
able margin of dactyl of chela with a distinct excision. Inner
margin of palm of chela usually barbate -_--_P. escambiensis
sp. nov.....--.----------- ----- --- --- p. 46
16 (10) Cephalodistal surface of first pleopod with a rounded prominence;
cephalic process small and acute and arising from the mesial
surface of the appendage; caudal process rounded and non-
corneous ----... P. pubischelae sp. nov.. ..------.----._ p. 41
16' Cephalodistal surface of first pleopod without a rounded promi-
nence, cephalic process blade-like and arising from the cephalic
surface of the appendage; caudal process with sharp corneous
edges .---- P. shermani sp. nov ............------------- p. 61






30 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

17 (9') Inner margin of palm of chela not barbate _-----P. rathbunae
(Hobbs) .-------------------___ ----.. -------- p. 59
17' Inner margin of palm of chela barbate-------------- 18
18 (17') Caudal process of first pleopod forming a corneous blade-like
structure along the caudolateral tip of the appendage__--
P. hubbelli (Hobbs) _-- --_____------------ ----- p. 67
18' Caudal process of first pleopod rounded and non-corneous------
P. pubischelae sp. nov-........._....- ------- --.. p. 41
19 (7) Hooks on ischiopodites of both third and fourth pereiopods--- -
P. geodytes sp. nov ..- ------------------------ p. 80
19' Hooks on ischiopodites of third pereiopods only __--20
20 (19') First pleopod of male with the plate-like, corneous fan formed by
the central projection directed aross the cephalic surface of the
tip -- ------------------------- ------22
20' First pleopod of male with the plate-like, corneous fan formed by
the central projection directed cephalocaudad ---_ ---_ 21
21 (20') Cephalic process of first pleopod present as a vestige ----P. ad-
vena (LeConte) .------------ -------.. --.-----__.. p. 75
21' Cephalic process of first pleopod absent_ -- P. pygmaeus sp.
nov.-----------_------------------_____ ------------p. 83
22 (20) The plate-like central projection of the first pleopod directed
obliquely caudad across the cephalic border of the tip; cephalic
process present --__---. P. rogersi ochlocknensis Hobbs -- p. 89
22' The plate-like central projection of the first pleopod directed lat-
erad across the cephalic border of the tip; cephalic process
present or absent___-----___-----------__-------- 23
23 (22') Caudal process of first pleopod bent, but not so much as at a right
angle to the main shaft of the appendage ---- P. rogersi cam-
pestris Hobbs ----------------------------- .p. 90
23' Caudal process of first pleopod bent at a right angle to the main
shaft of the appendage .----- P. rogersi rogersi (Hobbs) p. 89
24 (5') Eyes reduced; carapace without pigment ----- (Pictus Group, in
part) -------------------_------ -----------29
24' Eyes well developed; carapace pigmented -- --------- 25
25 (24') First pleopod bearing a hump on cephalic margin (Clarkii Sub-
group of Blandingii Group) ---- -----_---------- 35
25' First pleopod never bearing a hump on cephalic margin -__- 26
26 (25') Chela bearing a row of seven or less tubercles along mesial margin
of palm (if more, then a median carina is present on upper sur-
face of rostrum) ..--- Spiculifer Group_ __--------_- 39
26' Chela bearing a row of eight or more or several rows of tubercles
along mesial margin of palm, or inner margin of palm smooth.
Never with a median carina present on rostrum ....--- ------27
27 (26') Areola broad and short. Cephalic process of first pleopod spini-
form and clearly extending beyond the main body of the ap-
pendage ----- (Pictus Group, in part) ------_------ 29






HOBBS-THE CRAYFISHES OF FLORIDA


27' Areola narrow and long. Cephalic process of first pleopod some-
times spiniform, but if so extends barely distad of the main
body of the appendage (Blandingii Group) ---------- 28
28 (27') First pleopod with all four terminal apices strongly developed,
the cephalic and caudal process and the central projection all
corneous and blade-like -----Blandingii Subgroup _- -----34
28' First pleopod never with all four terminal apices strongly devel-
oped, and the cephalic and caudal processes and the central
projection never all blade-like___ -(Fallax-Evermanni Sub-
groups) -----.....---------- ----------------------- ----36
29 (24 and 27) Carapace pigmented; eyes well developed .-.--- ------32
29' Carapace without pigment; eyes reduced ---------------30
30 (29') Eye with small pigment spot -_-. P. lucifugus alachua (Hobbs)
._----------- ---------- -------------- -----p. 136
30' Eye without small pigment spot ... -------------- ---- 31
31 (30') Rostrum broadest distad of base ----- P. lucifugus lucifugus
(Hobbs) ...--- ----------------..----- -----p. 134
31' Rostrum broadest at base --- __P. pallidus (Hobbs)___ p. 139
32 (29) Length of inner margin of palm of chela greater than length of
dactyl; acumen of rostrum as long as or longer than rest of
rostrum ---- _P. young sp. nov..------.------ --- ---- p. 131
32' Length of inner margin of palm of chela less than length of dactyl;
acumen of rostrum much shorter than rest of rostrum ----33
33 (32') Central projection of first pleopod extends distad to tip of cepha-
lic process; both spiculiform; cephalic process directed distad
.-------- P. seminolae sp. nov.------------------p. 142
33' Central projection of first pleopod extends distad but never so far
as tip of cephalic process; neither spiculiform; cephalic process
directed at about a 45 degree angle to the main shaft ----
P. pictus (Hobbs) _......------ ------- ----- p. 130
34 (28) Length of rostrum greater than length of areola; a dark stripe on
either side of thoracic portion of carapace that persists even in
alcohol--___ -_ P. bivittatus sp. nov. -.---.------------------ p. 96
34' Length of rostrum less than length of areola; no dark stripe on
thoracic portion of carapace ------ P. blandingii acutus (Gir-
ard) __-- _----- --------- --.. ..- ----------- --- p. 94
35 (25) Mesial process of first pleopod directed distad. Caudal process
forms an undulating blade across the caudolateral surface (in
ventral view). Cephalocaudal dimension of cephalic process
greater than length--___--P. okaloosae sp. nov.-.----p. 100
35' Mesial process of first pleopod directed caudad. In ventral view
caudal process forms a narrow blade closely applied to the
central projection. Cephalocaudal dimension of cephalic pro-
cess less than length ----- P. paeninsulanus (Faxon) -- p. 104
36 (28') Mesial process of first pleopod blade-like --- P. fallax (Hagen)
.------------ ...---...------------------- p. 11






32 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

36' Mesial process of first pleopod spiniform .- _.---------------_ 37
37 (36') Caudal process of first pleopod long and blade-like, extending
distad almost to tip of central projection ---- P. evermanni
(Faxon) _--------.-----------___ .---------__.p. 107
37' Caudal process of first pleopod vestigial, hardly discernible; cen-
tral projection reduced to a minute corneous spine or blade
extending from the center of the appendage --------- 38
38 (37') Distal end of first pleopod bent at about a 45 degree angle to the
main shaft; shoulder on cephalodistal surface rounded ---_-
P. leonensis Hobbs __--------___.--_-....-....p. 114
38' Distal end of first pleopod only slightly bent; shoulder on cepha-
lodistal surface angular-.....-P. pycnogonopodus Hobbs --_-
----------- ------------ ------------ p. 117
39 (26) Rostrum with slight median carina --- P. versutus (Hagen)
--.----. .-------...-- --------- ...p. 126
39' Rostrum without median carina --_--P. spiculifer (LeConte)-"
---- ------------ ------------------------------ 119
40 (4) Areola obliterated at midlength_ -- (Diogenes Section) --.--43
40' Areola broad or narrow, but never obliterated at midlength .------
(Bartonii Section) --__--- -.__---....41
41 (40') Eyes reduced; eyes and carapace without pigment ---- C. cryp-
todytes Hobbs ___ __ -------------- -- p. 162
41' Eyes well developed; both eyes and carapace pigmented .-___- 42
42 (41') Areola with three or four punctations in narrowest part ---__
C. latimanus (LeConte)--...- ..-----.--..-.... -- .p. 158
42' Areola with one or two punctations in narrowest part .----C. flo-
ridanus Hobbs ------ --.----- ---- -- -----.. p. 161
43 (40) Rostrum lanceolate or with acumen distinct, well set off, deeply
excavate. No cristiform row of tubercles along inner margin
of palm of chela _---- C. diogenes diogenes (Girard) ---p. 164
43' Rostrum subovate, acumen never sharply set off, deeply or shal-
lowly excavate. A more or less distinct cristiform row of tuber-
cles along inner margin of palm of chela ----------44
44 (43') Rostrum short; directed ventrad; margins converging from base.
The cristiform row of tubercles along inner margin of palm
acute and well marked. Cephalomesial margin of antennal
scale slanting or gently rounded __----- C. byersi Hobbs p. 167
44' Rostrum long; directed ventrad, but not so greatly as in byersi;
margins subparallel for some distance at base. The cristiform
row of tubercles along inner margin of palm of chela less acute
than in byersi. Cephalomesial margin of antennal scale angu-
late ------ C. species incertis ..._--------_ ------ p. 168






HOBBS-THE CRAYFISHES OF FLORIDA


GENUS PROCAMBARUS ORTMANN
Groups I and III ( in part) of Cambarus, Hagen 1870, Ill. Cat. Mus. Comp.
Zool., Harvard Coll., No. 3: 32-55, 74, 84, 87-88.
Groups I and II of Cambarus, Faxon 1885a, Mem. Mus. Comp. Zool., Har-
vard Coll., 10 (4): 17, 47.
Subgenus Cambarus, Ortmann 1905a, Proc. Amer. Philos. Soc., 44 (180):
96-106.
Subgenus Procambarus, Ortmann 1905c, Ann. Carnegie Mus., 3 (3): 437.
Subgenus Ortmannicus, Fowler 1911, Ann. Rep. New Jersey Mus. 1911,
Part II: 340.
Groups I, II, III of Cambarus, Faxon 1914, Mem. Mus. Comp. Zool., Harvard
Coll., 40 (8): 410-414.
Subgenus Cambarellus, Creaser 1933b, Occ. Pap. Mus. Zool., Univ. Mich.,
(275): 21.
Genus Procambarus, Hobbs (in press-c), Amer. Mid. Nat.
A historical summary of the taxonomic treatment of the group by former
authors has been published elsewhere (Hobbs, in press-c). A table from this
paper (p. 22) shows the synonomy involved.
Diagnosis.-"First pleopod of first form male terminating in from two
to five distinct parts which may be truncate, plate-like or spiniform. Shoulder
present or absent on cephalic surface of distal third. If the pleopod terminates
in only two parts this shoulder is always present. Hooks present on the ischiop-
odites of the third or of the third and fourth pereiopods in the male. Third
maxillipeds of normal size bearing a row of teeth along the inner margin of
the ischiopodite" Hobbs (in press-c).
The geographic range of this genus is undoubtedly more extensive than
that of any of the others in the Subfamily Cambarinae, extending from Wis-
consin and Colorado to Guatemala and eastward to the Atlantic seaboard from
New York to Florida.
Except in the mountainous regions one or more representatives of the
genus can be found in any body of freshwater, or nearly any low, poorly
drained area, but it reaches its greatest abundance in the coastal plains of the
southern and southeastern United States.
In Florida Procambarus is represented by 33 species and subspecies
and extends throughout the state from the western extremity of the northwest-
ern panhandle to Big Pine Key, off the southern tip of the mainland.

THE BARBATUS SECTION
Diagnosis.-Cephalodistal surface of the first pleopod of first form
male terminates in a ridge (sharp or truncate) or a knob-like prominence
which is distinctly not a part of one of the terminal processes; mesial process
always directed distad unless spatulate; cephalic process, when present, al-
ways extends distad from mesial surface (except in shermani); central pro-







34 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2


Map 3.-Migrations of the Barbatus Group. The dot-dash line shows the Sunderland
Shore Line of early Pleistocene; the shaded portions indicate the areas inhabited by the
ancestral stock during early Pleistocene. The earliest wave of migration probably came
from the general region designated by I; later migrations appear to have been derived
from the more circumscribed centers, shown as II and III. The letters indicate the
general regions in which the modern species occur, and the lines represent the probable
paths taken by their ancestors. The species A-escambiensis, B--econfinae, C-barba-
fus, D-pubischelae, E-atipleurum, F-apalachicolae, and G-rathbunae are mem-
bers of the barbatus subgroup, while K-kilbyi, L-shermani, and M-hubbelli belong
to monotypic subgroups, designated respectively by the names of the species.






HOBBS-THE CRAYFISHES OF FLORIDA


jection never decidedly the most conspicuous terminal element. Hooks pres-
ent on ischiopodites of third, or third and fourth pereiopods.
Fourteen of the known species of Procambarus are referable to the bar-
batus section: barbatus, pubischel.e, escambiensis, econfinae, latipleurum,
apalachicolae, rathbunae, shermfAi, kilbyi, hubbelli, simulans, gracilis, ha-
genianus, and alleni. Ten of these occur in Florida, and one or more has been
recorded from New Mexico, Colorado, Texas, Oklahoma, Kansas, Arkansas,
Missouri, Iowa, Wisconsin, Illinois, Mississippi, Alabama, and Georgia.
The Barbatus Section may be divided into the following groups and sub-
groups:
BARBATUS SECTION
Barbatus Group
Barbatus Subgroup
Shermani Subgroup
Kilbyi Subgroup
Hubbelli Subgroup
Alleni Group
THE BARBATUS GROUP
Diagnosis.-First pleopod of first form male terminates in four dis-
tinct parts and bears no decided hump or shoulder on cephalic margin. Areola
relatively short and broad; no lateral spines present on rostrum nor are mar-
gins interrupted (except occasionally in latipleurum); hooks present on
ischiopodites of third, or third and fourth pereiopods.
The barbatus group comprises an assemblage of secondary burrowing
species which are almost confined to the flatwoods of the coastal terraces (or
adjoining flatwoods) from the Perdido River drainage system eastward into
South Carolina, and southward to Flagler, Alachua, and Levy counties in
Florida. It should be noted that within these boundaries the actual range of
the group is broken into distinctly discontinuous areas. (Map 4).
Of the ten species which belong to the barbatus group barbatuss, pubis-
chelae, escambiensis, econfinae, latipleurum, apalachicolae, rathbunae,
shermani, kilbyi, and hubbelli) all except the first occur in Florida, and the
last seven are thus far known only from this state. It is probable, however,
that further collecting will extend the range of escambiensis, rathbunae, sher-
mani, and hubbelli into Alabama.
The barbatus subgroup appears to me to be a natural assemblage of re-
lated species, and the relationships indicated by morphological criteria are
directly correlated with the existing pattern of geographic distribution. It is
thus possible to postulate the paths of migration by which the progenitors of
the present barbatus subgroup occupied the areas where they are now estab-
lished.
Although shermani, kilbyi, and hubbelli are closely related to the mem-
bers of the barbatus subgroup, their anatomical peculiarities are so distinct
from the other members of the barbatus group and from each other that it






36 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2


Map 4.-Distribution of the Barbatus Group.






HOBBS--THE CRAYFISHES OF FLORIDA


seems best to erect separate subgroups for their reception. As will be pointed
out below they probably originated from different stocks of the barbatus
group, independently.
The present distribution of the barbatus group is most acceptably ex-
plained on the assumption that its forerunners occupied the gulf coastal region
(I) as indicated (Map 3), comparatively early in the Pleistocene emergence
of the coastal plain; a southward and eastward migration took place that re-
sulted in the isolation of at least three groups: a western (A), a central (B),
and a northeastern (C and D) which occupied respectively the regions of the
drainages of the Perdido, Flint-Chattahoochee (or Choctawhatchee), and
Altamaha rivers. Perhaps a fourth group (L) which took a different south-
ward path along the Escambia should be considered here; however, I am in-
clined to believe that this migration should be referred to a later period.
The western group progressed southward along the Perdido River where
it was isolated from the central group by the high, rolling country in Okaloosa
and Walton counties. This stock gave rise to P. escambiensis (A).
The central stock progressed southward either along the Flint-Chatta-
hoochee or along the Choctawhatchee drainages giving rise to econfinae (B).
The third stock, moving eastward, diverged, part (C) to cross the Alta-
maha and Savannah rivers and part (B) to extend southward into Florida.
The part that moved north of the Altamaha gave rise to P. barbatus while that
which moved southward was ancestral to P. pubischelae in Georgia south of
the Altamaha River and in northeastern Florida.
Following the first wave of migration which resulted in the isolation of
escambiensis, econfinae, barbatus, and pubischelae, a second, and possibly
a third, wave of migration is strongly indicated. In the second region the mi-
gration came from two separate but more restricted regions (labelled II and
III on the map), one in the vicinity of the headwaters of the Escambia River
in southern Alabama and the other in southeastern Alabama and southwestern
Georgia. The stock from the more western area migrated southward along
the Escambia River to give rise to P. shermani (E).7
The group which occupied Area II in southeastern Alabama and south-
western Georgia extended southward along three separate routes. The more
western one (G) extended in a southwesterly direction and reaching the Black-
water and Yellow river systems gave rise to P. rathbunae. Another line of
migration followed the west side of the embayment of the Flint River into the
coastal region, and there segregated to form P. apalachicolae (F) and P. lati-
pleurum (E). Still farther east another stock pushed southward to give rise
to P. kilbyi (K) which followed the coast southward as far as Levy County,
and westward across the Apalachicola River into Calhoun, Franklin, and Gulf
counties.

'This species is particularly interesting in that it possesses characters which are remarkably
like those of the blandingii section. Further consideration of this species and its relationships is
given below.






38 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

The origin and time of migration of the stock from which P. hubbelli was
derived are more questionable. On morphological grounds hubbelli is evident-
ly close to the original Pro-barbatus stock and perhaps should be referred to
the earliest migration across the Sunderland Terrace, but the relationship of
its present range to that of other species in the central portion of the panhandle
seems to indicate a comparatively late, delayed migration from the region of
Area II.
All of the species derived during the first period of migration share a
number of peculiar characters, and with the exception of econfinae, which al-
most certainly has its closest affinities with escambiensis, all have first form
males that show a heavy beard along the inner margin of the palm of the chela.
Except for shermani, those species derived from subsequent migrations have a
naked chela, and the pleopods of all of this group (except for shermani and
kilbyi) are remarkably similar. P. shermani is unique in so many ways in this
assemblage that it may well have been derived from a stock already differen-
tiated from that which provided the main body of later migrants. P. hubbelli
is likewise a disjunct species in this group, although it has barbate chelae.
It is interesting to note how definitely the present ranges of the members
of the barbatus group illustrate Matthews' "Center Fire" hypothesis of distri-
bution. There is little doubt that this group had its distribution from the older
region of the southeastern coastal plain, and now we find the periphery of the
present range occupied by the earliest segregates from the original stock, while
the region between the center of dispersal and the periphery occupied by later
migrants from the center of dispersal. The older species are barbatus and pu-
bischelae along the eastern seaboard, econfinae in a small area around Pana-
ma City (completely surrounded by members of the later migration), and es-
cambiensis on the western periphery of the range, along the Perdido River; the
more recent forms, particularly rathbunae, apalachicolae, and latipleurum
have filled in the gaps, and are established farther inland. The latter two par-
ticularly indicate a more recent advent into the Panama City region as evi-
denced by their complete encirclement of P. econfinae.
The first pleopods of the barbatus group, especially those of P. barbatus,
very closely approximate the form of the appendages in P. mexicanus, and it
is possible that barbatus more nearly represents the ancestral stock than do
any of the other members of the group.
THE BARBATUS SUBGROUP
Diagnosis.-Cephalodistal surface of first pleopod of first form male
generally compressed and bears a row of setae; mesial process typically spin-
iform, occasionally flattened, but more often cylindrical, extends distad be-
yond the rest of the appendage; cephalic process, small and corneous, has
shifted mesially and arises from the distomesial margin (except in shermani
where it arises from the cephalic surface); central projection markedly var-
iable, corneous, and very small, generally forms a triangular plate that is
flattened cephalocaudad; caudal process usually large and terminates in






HOBBS--THE CRAYFISHES OF FLORIDA


either a knob, a distally flattened prominence, a corneous blade-like structure
directed cephalocaudad, or a subtriangular, non-corneous process. In most in-
stances the distal portion of the appendage is bent more or less caudad. The
areola is broad or relatively broad; the margin of the rostrum is toothless and
uninterrupted (except in occasional specimens of latipleurum) ; the ischiopo-
dites of the third pereiopods bear hooks while those of the fourth may be with
or without hooks; the inner margin of the palm of the chelae of the male may
or may not be barbate.
The members of the barbatus subgroup commonly inhabit flatwoods, and
generally live in temporary bodies of water. When the pools dry up the cray-
fishes dig into the wet soil of the old pool bottoms and construct rather simple
burrows which may range from six inches to three feet in depth, depending on
the soil type and the length and severity of the period of drought. Specimens
are occasionally found in the smaller streams of the flatwoods, but these stream
dwellers also construct burrows at the breeding season. I have never collected
specimens from a large body of water, and there is some evidence that large
rivers act as a barrier to certain of the species.
The range of the barbatus subgroup is confined to the southeastern coas-
tal plain where it is separated into an eastern and western portion. The eastern
portion extends northward into the southeastern part of South Carolina; on the
northwest the boundary is marked by the edge of the Tifton Upland; on the
west (south of the Tifton Upland) by the comparatively high, well drained re-
gion in Hamilton, Columbia, Union, and Alachua counties, Florida. The south-
ern part of the range extends into northern Florida, southward to Alachua and
Flagler counties. The western portion of the range has its eastern boundary
along the Apalachicola River; its northern boundary is the northern limit of
the coastal flatwoods in Gulf, Bay, Walton, Okaloosa, Santa Rosa, and Escam-
bia counties, Florida; its western boundary is in the vicinity of the Perdido
River.
Generally in Florida these flatwoods extend hardly more than 25 miles
from the coast, although Flomaton, Escambia County, Alabama, where P. es-
cambiensis has been reported, lies approximately 30 miles from the coast.

Procambarus barbatus (Faxon)
Plate II, Figs. 1-5; Maps 3, 4

Astacus penicillatus LeConte (not Astacus penicillatus Olivier 1791) 1856,
Proc. Acad. Nat. Sci. Philad., 7: 401.
Cambarus penicillatus Hagen 1870: 16, 31-33, 53-54, 55, 97, 100, 106, 107,
108, P1. I, figs. 93, 94; P1. III, fig. 149; Faxon 1884 (part): 138; Faxon
1885a (part): 18, 36-38, 39, 158, 173.
Cambarus barbatus Faxon 1890: 621, new name to replace penicillatus Le-
Conte 1856, preoccupied by penicillatus Olivier 1791 (among Faxon's
neotypes are specimens from the Escambia River at Flomaton, Alabama,






40 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

herein described as P. escambiensis Hobbs); Harris 1903 (part): 58,
72, 137, 138, 150, 152, 153, 154; Hobbs 1940a: 389, 410, 414, 418;
Hobbs 1940b: 3; Hobbs (in press-a).
Procambarus barbatus Hobbs (in press-c).
Diagnosis.-Rostrum without lateral spines; areola relatively broad
with three to five punctations in narrowest part; male with hooks on ischiopo-
dites of third and fourth pereiopods; inner margin of palm of chela of first
form male usually bearded within (bearing a heavy growth of plumose setae);
postorbital ridges terminating cephalad without spines; no lateral spines pres-
ent on carapace. First pleopod of male, form I, reaching coxopodite of third
pereiopod and terminating in four parts; mesial process, the largest of the
four, corneous, subspiculiform, and extending distad and slightly recurved
cephalad; cephalic process small, corneous, and arising from mesial surface
of appendage, is curved and projects from beneath distal portion of central
projection; caudal process, hardly discernible from the truncate portion, some-
what flattened, and forming the caudodistal portion of the outer part; central
projection small, corneous, triangular, and arising from central portion of tip,
is flattened cephalocaudad and directed cephalodistad, somewhat overhanging
the basal portion of the cephalic process. Cephalic margin of main shaft near
tip bearing a cluster of setae which forms a single row as it approaches the
terminal processes. Annulus ventralis subrectangular with prominent tubercles
on each cephalolateral margin; sinus originating near center of annulus,
slightly dextrad of midventral line, curves gently dextrad, then sharply sinis-
trad to cross midventral line where it makes a hairpin turn almost back to mid-
ventral line; thence caudad to caudal margin.
Remarks.-Faxon (1885a: 38) states, "LeConte does not specify from
what part of Georgia his specimens come, nor is the locality of the Georgia
specimens in the Museum of Comparative Zoology any more precisely indicat-
ed." LeConte's types of this species are unknown. According to Faxon (1890:
621) ... LeConte's specific name penicillatus must be rejected, since it has
been used previously by Olivier (Encyc. Meth., Hist. Nat. des Insectes, VI,
1791, p. 343), in combination with the same generic name, for another animal
(Palinurus penicillatus of recent authors)." In this paper Faxon proposed the
name, Cambarus barbatus, to replace Cambarus penicillatus. Faxon (1914:
414) cites a first form male from Georgia, but without further data, (M. C. Z.
no. 279) as his type specimen, and several specimens from the Escambia River,
Flomaton, Alabama (M. C. Z. no. 3845) as paratypes. My assignment of the
name barbatus is based on study of Faxon's Neotype (M. C. Z. no. 279); his
"paratypes" belong to another species, Procambarus escambiensis, herein de-
scribed.
I have made camera lucida drawings of Faxon's neotype and "paratypes"
at M. C. Z., and since that time have collected in several counties in southeastern
Georgia and South Carolina. Among my specimens from Liberty County, Geor-
gia, there is a first form male which resembles the neotype very closely. In ad-






HOBBS-THE CRAYFISHES OF FLORIDA


edition, I also have a large number of specimens collected north of the Altamaha
River in Georgia and South Carolina which belong to this species. Collections
made in Georgia south of the Altamaha River have disclosed a very closely
related form but not P. barbatus. I believe it likely that Faxon's neotype was
collected from the region between the Altamaha and Savannah rivers in the
coastal plain.
Although this species does not occur in Florida it is included here for two
reasons. It has been confused with a very closely allied species which occurs in
Florida, and it is also the only known non-Floridian member of the barbatus
subgroup, unless Procambarus pearsei (Creaser) is also a member of this
group.
Distribution.-GEORGIA-Bulloch, Effingham, Jenkins, Liberty, Long,
and Screven counties. SOUTH CAROLINA-Hampton County (Hobbs 1940b: 3).
Faxon cites the following records with a query: South Carolina (?); Charles-
ton (?) ; Mississippi (?); Eastern (?).
All of the records cited above, except where otherwise indicated, are new;
heretofore the only data given has been "Lower Georgia" or Georgia.

Procambarus pubischelae, sp. nov.
Plate II, Figs. 6-10; Plate XIII; Maps 3, 4.

Diagnosis.-Rostrum without lateral spines; areola relatively broad
with three to five punctations in narrowest part; male with hooks on ischio-
podites of third only, or third and fourth pereiopods; palm of chela of first
form male usually bearded; postorbital ridges terminating cephalad without
spines; no lateral spines present on carapace. First pleopod of male, form I,
reaching coxopodite of second pereiopod and terminating in four distinct
parts; mesial process corneous and subspiculiform; cephalic process corneous,
small, and spiniform; caudal process noncorneous and truncate; central pro-
jection corneous, subtriangular, and compressed cephalocaudad. Cephalo-
distal setae-bearing prominence subangular. Annulus ventralis subrectangu-
lar with a triangular lateral projection on each side; excavate cephalad with
tuberculate prominences laterad; sinus originating dextrad of midventral
line, bends sharply sinistrad, and runs laterad to cross the midventral line
where it forms an arc and terminates just before cutting the caudal margin
sinistrad of middle.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen nar-
rower than thorax (1.21-1.35 cm. in widest parts respectively).
Width and depth of carapace subequal in region of caudodorsal margin of cervical
groove. Greatest width of carapace slightly caudad of caudodorsal margin of cervical
groove.
Areola of moderate width (4.68 times longer than wide); with four punctations
in narrowest part; sides not parallel. Cephalic section of carapace about 1.99 times
as long as areola (length of areola 33.5% of entire length of carapace).
Rostrum flattened above, almost reaching base of distal segment of peduncle of
antennule; margins converging to tip; no lateral spines present. Upper surface of ros-







42 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

trum punctate; marginal ridges rather high and sharp. Subrostral ridge well defined
but evident in dorsal view for only a short distance from base.
Postorbital ridges well defined, merging rather abruptly cephalad into carapace
without forming a tubercle or spine. Suborbital angle rounded, only moderately prom-
inent; branchiostegal spine well developed; no lateral spines on sides of carapace, a
few small tubercles present instead. Surface of carapace punctate dorsad, weakly gran-
ulate laterad.
Abdomen longer than thorax (3.0-2.6 cm.).
Cephalic section of telson with two spines in each caudolateral corner.
Epistome subovate with a single emargination cephalad.
Antennule of usual form; a spine present on ventromesial side of basal segment.
Antennae reaching caudad to fourth abdominal segment. Antennal scale broad,
broadest in middle; inner margin evenly rounded; spine on outer margin moderately
strong.
Chela subovate, compressed dorsoventrally, of moderate length, and broad. Hand
entirely tuberculate except on ventrolateral surface which bears setiferous punctations.
Inner margin of palm with a heavy beard of plumose setae. A distinct ridge present on
both fingers. Fingers not gaping. Opposable margin of dactyl with a row of 11 round-
ed tubercles, and with sparsely scattered denticles along distal half. Lateral margin of
dactyl with a row of six subsquamous, ciliated tubercles along basal half. Upper surface
with a single submedian ridge at the base of which are a few small scattered tubercles,
flanked with scattered setiferous punctations distad. Lower surface also with a dis-
tinct median ridge and scattered setiferous punctations. Opposable margin of immov-
able finger with six rounded tubercles and with minute denticles scattered along entire
length. Lateral margin with a weak ridge. Upper surface tuberculate proximad and
punctate distad with a submedian ridge. Lower surface with a distinct submedian
ridge and scattered punctations. A single row of punctations bearing long, stiff setae
present below opposable margins of both fingers.
Carpus of first pereiopod longer than wide (.80-.56 cm.); shorter than inner mar-
gin of palm of chela (.88 cm.); a distinct longitudinal groove above; dorsomesial and
mesial surfaces tuberculate; otherwise sparsely punctate. Two larger tubercles along
midmesial surface, and two along cephalic dorsomesial margin. Three large tubercles
present in a row running obliquely from lower cephalomesial surface to cephalic mar-
gin of ventral surface.
Merus of first pereiopod punctate mesiad and laterad. A group of scattered
tubercles present on upper surface, more numerous distad. Lower surface with two
rows of tubercles, an outer one of about nine which are smaller in size than the inner
row of about 14, most of which are spike-like.
Hooks present on ischiopodites of third and fourth pereiopods. Bases of coxo-
podites of fourth and fifth pereiopods with strong outgrowths directed caudoventrad
and cephaloventrad respectively.
First pleopod reaching base of second pereiopod when abdomen is flexed; termi-
nal portion of appendage with a tuft of setae on cephalic surface. Appendage ending
in four distinct parts. Mesial process subspiculiform and corneous, extends beyond the
rest of the terminal processes in a cephalodistal direction. Cephalic process, small and
corneous, arises from the mesial surface and extends almost parallel to the mesial
process. Caudal process truncate, noncorneous, heavy and slightly excavate distad,,
makes up the caudolateral portion of the tip. Central projection small, corneous, sub-
triangular, flattened cephalocaudad, and directed cephaloventrad. Cephalodistal por-
tion of the appendage while rounded in lateral aspect is somewhat dome-shaped in
contrast to the gradual sloping appearance seen in Procambarus barbatus.
Male, Form II.-Differs from the first form male in the following respects: inner







HOBBS-THE CRAYFISHES OF FLORIDA


margin of palm of chela never bearded; all spines and tubercles greatly reduced in
size and somewhat in number; hooks on ischiopodites of third and fourth pereiopods
very small; no distinction between the caudal process and the central projection of the
first pleopod, no parts corneous; dome-shaped appearance of cephalodistal portion
not nearly so well marked as in first form male.
Allotypic Female.-The outstanding difference between the first form male and
the female is in the chela which is never bearded within and always weaker (see
measurements).
Annulus ventralis subrectangular in shape with a subtriangular projection on
both lateral surfaces; deeply excavate cephalad with tuberculate prominences laterad;
caudal margin slightly excavate; sinus originates dextrad of midventral line, bends
sharply sinistrad, and runs laterad to cross the midventral line where it forms an are
and terminates just before it cuts the caudal margin.
Measurements.-Male (form I) Holotype: carapace, height 1.34, width 1.35,
length 2.60 cm.; areola, width .19, length .87 cm.; rostrum, width .46, length .53 cm.;
abdomen, length 3.0 cm.; right chela, length of inner margin of palm .88, width of
palm .83, length of outer margin of hand 2.23, length of movable finger 1.14 cm. Fe-
male Allotype: carapace, height 1.41, width 1.45, length 2.80 cm.; areola, width .24,
length .91 cm.; rostrum, width .53, length .60 cm.; abdomen, length 3.10 cm.; right
chela, length of inner margin of palm .64, width of palm .73, length of outer margin
of hand 1.78, length of movable finger 1.02 cm.
Type Locality.-A cypress pond and roadside ditch 9.4 miles north (State High-
way 82) of Lake City, Columbia County, Florida. This locality is in the flatwoods
area which extends westward and southwestward from the Okefenokee Swamp. Most
of the specimens collected here were taken from simple burrows about one to one and
one-half feet in depth and marked by low, poorly formed chimneys.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form I), another male (form II), and one female are deposited
in the Museum of Comparative Zoology; one male (form I), another male (form II),
and a female in the University of Michigan Museum of Zoology; one male (form I),
one male (form II), and a female in the Academy of Natural Sciences at Philadelphia.
Fourteen males (form I), 18 males (form II), 56 females, four immature males, and
four immature females are in my personal collection at the University of Florida.
Remarks.-My series of approximately 250 specimens does not represent
a homogeneous population. Although all are probably conspecific the speci-
mens from each locality tend to show small but distinguishable variations that
enable one to recognize fairly accurately the region from which they came. In
general some five variants may be recognized, and in the discussion that fol-
lows each is designated by the name of the town nearest to its place of capture.
One distinct variant occurs in the vicinity of Baxley, (Appling County)
Georgia. The male is provided with hooks on the ischiopodites of only the third
pair of pereiopods, but in one or two specimens there is a rudiment of a hook
on the ischiopodite of the fourth. The areola is relatively broad with about three
punctations in the narrowest part, and the rostrum is broader than in specimens
from some of the other localities; the annulus ventralis bears no tubercles.
The males of the Jacksonville, Florida, specimens possess hooks on the is-
chiopodites of the third and fourth pereiopods. The areola is narrower and
longer than in the Baxley specimens, and the annulus ventralis bears a group
of three to five acute tubercles along the lateral margins.






44 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

In the males of the Waycross, Georgia, specimens hooks are present on the
ischiopodites of both the third and fourth pereiopods. The rostrum as well as
the areola is broader than that in the Jacksonville specimens, and the annulus
ventralis, although similar to that of the Baxley females, has the lateral mar-
gins comparatively higher.
Like the Baxley males, those from Adel, (Cook County) Georgia, possess
hooks on the ischiopodites of only the third pereiopods and can be distinguished
from the Baxley specimens only by the structure of the first pleopod. The areo-
lae are also similar, but the annuli ventralis of the Adel females bear an irreg-
ular group of three or four tubercles along each lateral surface.
The Lake City, Florida, specimens are most closely allied to those from
Jacksonville but may be recognized by the difference in structure of the first
pleopod. The surface of the annulus ventralis, though irregular, does not bear
acute tubercles.
The specimens from Nassau County, Florida, are most like those from
Jacksonville; those from Alachua, Columbia, and Union counties are for the
most part similar to the Lake City specimens. The annuli ventralis in the three
females from Flagler County are without tubercles, but until a first form male
has been collected I hesitate to state what the relationships of the Flagler Coun-
ty specimens are.
Procambarus pubischelae probably has its closest affinities with P. barba-
tus and P. escambiensis. It is possible that P. pubischelae will prove to be a sub-
species of P. barbatus, but present data indicate that they are geographically
isolated in the region of the Altamaha River. On Georgia State Highway 38,
between Hinesville and the river, I took specimens of P. barbatus. In this region
flatwoods occur for a mile or two immediately north of the Altamaha, but the
southern bank consists of a broad upland which extends southward almost to
Jesup. About one-half mile north of Jesup P. pubischelae was collected. Other
locality records strengthen the evidence that P. barbatus occurs only north of
the Altamaha while P. pubischelae occurs only south of it.
Specimens Examined.-I have examined a total of 245 specimens of Pro-
cambarus pubischelae, from the following counties in Georgia and Florida:
GEORGIA-Appling, Camden, Clinch, Colquitt, Cook, Lanier, Lowndes, Ware,
and Wayne. FLORIDA-Alachua, Baker, Columbia, Duval, Flagler (?), Nas-
sau, and Union.
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
dI 6 4 9 18 1 3
" II 3 5 3 1 1 19
9 2 10 12 4 2 51 2
9 (eggs) 1 2 11
9 (young) 9
6 (immature) 1 6 8
9 (immature) 11 5 17






HOBBS-THE CRAYFISHES OF FLORIDA


Geographical and Ecological Distribution.-Procambarus pubischelae
is probably confined to southeastern Georgia and northeastern Florida, where
it is abundant in flatwoods situations. The northern boundary of its range
seems to be in the vicinity of the Altamaha River, and while my collections in
central Georgia are too inadequate to cite the western boundary in that state,
this species is probably, with few exceptions, confined to the regions of the
Coastal Terraces (specimens have been collected in Appling, Colquitt, Cook,
Lowndes, Clinch, and Ware counties). In Florida the western boundary of the
range lies in Columbia County, and probably follows the western limits of the
flatwoods [shown in Harper's Generalized Soil Map of Florida (1925)] in
Hamilton, Columbia, Bradford, and Alachua counties. The southern limit of
the range seems to be approximately a line drawn from Lake Butler to Orange
Heights, then due east to the Putnam County line. (No representatives of this
species have been found in Putnam, Clay, or St. Johns counties, although sev-
eral females, which if not P. pubischelae are very closely related to it, were
collected 11 miles southeast of San Mateo, Flagler County.) The range prob-
ably extends eastward to the salt marshes and brackish inlets of the ocean in
Georgia, while the eastern limit in Florida is still questionable.
Procambarus pubischelae was collected with P. alleni in Flagler County,
with P. seminolae in Alachua, Baker, Duval, Union, and Nassau counties, with
P. fallax in Alachua County, and with P. paeninsulanus in Baker County.
P. pubischelae is a characteristic secondary burrower. It is found abun-
dantly (usually in groups) in the roadside ditches of the flatwoods. For the
most part the burrows are extremely simple, consisting of a single tunnel which
leads almost straight downward. Sometimes a burrow is slightly angulated or
curved and terminates in a slight enlargement. The mouth is generally marked
by a low, crudely formed chimney; occasionally, however, carefully built
chimneys, in which the pellets are arranged in similar fashion to those of P.
advena, are constructed. The burrows seldom exceed two and one-half feet in
depth and have been observed in sandy clay soils, in black muck, in sandy
mud, and in mottled red and blue clay.
I saw scores of open burrows scattered over the bottom of a shallow, tem-
porary flatwoods pond west of Jacksonville shortly after a rainy season. Each
burrow was about one and one-half feet deep and was marked by a small patch
of yellow sandy clay around its opening. At the mouths of many of the burrows
the crayfish could be seen with their chelae extended toward the opening, and,
upon being disturbed, retreated into their holes. Occasionally, as I waded
through the pond a crayfish would dart from a small clump of vegetation,
scurry for a distance of one to four feet, and then disappear into one of the
burrows. Some of these burrows had two openings, but all of them were other-
wise very simple.
It is likely that most of the life of this species is spent in and about the
mouth of the burrow, the crayfish leaving it only in search of food or in search
of a mate. During the day I have seen this species in open water only once, and
then only a few feet away from the mouth of a burrow. At night, however, speci-
sns are often found in open water several feet from a burrow.






46 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Occasionally a male and female have been found in the same burrow,
and in each of these instances the male was in the first form. Generally there
is an inch or two of water in the burrows, but I have often dissected burrows
which, although moist, contained no water. In the latter cases the crayfish
were apparently in good condition, not visibly affected by the lack of standing
water.
Procambarus escambiensis, sp. nov.
Plate II, Figs. 11-15; Plate XIV; Maps 3, 4
Cambarus barbatus Faxon 1890 (in part), Proc. U. S. Nat. Mus. 12 (785):
621.
Diagnosis.-Rostrum without lateral spines; areola relatively broad
with three to five punctations in narrowest part; male with hooks on ischiopo-
dites of third and fourth pereiopods; palm of chela of first form male bearded
within; postorbital ridges terminating cephalad without spines; no lateral
spines present on carapace. First pleopod of male, form I, reaching coxopo-
dite of second pereiopod and terminating in four distinct and corneous parts;
mesial process long and spiniform (extending beyond the other terminal pro-
cesses) and directed caudodistad; cephalic process small and subspiculiform;
caudal process blade-like, compressed laterally; central projection also blade-
like, but compressed obliquely cephalocaudad. Annulus ventralis subovate;
longitudinal excavation along midventral line; high multituberculate ridges
on either side of cephalic half, flattened caudad; sinus originates dextrad of
midventral line about one-third of the total length of annulus from cephalic
border; curves sharply laterad to cross midventral line and then gently caudad
and finally dextrad to cut the caudal margin.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen nar-
rower than thorax (1.25-1.54 cm. in widest parts respectively). Width and depth of
carapace subequal in region of caudodorsal margin of cervical groove. Greatest width
of carapace slightly caudad of caudodorsal margin of cervical groove.
Areola of moderate width (4.36 times longer than wide) with three punctations
in narrowest part; sides parallel for a short distance in middle. Cephalic section of
carapace about 2.36 times as long as areola (length of areola 29.7% of entire length
of carapace).
Rostrum shallowly excavate above, reaching base of distal segment of peduncle
of antennule; margins converging to tip; no lateral spines present. Upper surface of
rostrum polished with a few scattered punctations. Marginal ridges high and sharp.
Subrostral ridges well defined and evident in dorsal view for over half the length of
the rostrum.
Postorbital ridge well defined, merging abruptly cephalad into the carapace
without forming a tubercle or spine. Suborbital angle obtuse but prominent; branchio-
stegal spine well developed. No lateral spines or tubercles on sides of carapace. Sur-
face of carapace with conspicuous punctations dorsad, granulate laterad.
Abdomen and thorax subequal in length.
Cephalic section of telson with two spines in each caudolateral corner.
Epistome subtrapezoidal in shape with a strong cephalomedian spine.







HOBBS-THE CRAYFISHES OF FLORIDA


Antennules of usual form. A spine present on ventral side of basal segment; ven-
tral surface conspicuously hirsute.
Antennae reaching caudad to fifth abdominal segment; antennal scale broad;
broadest slightly distad of middle. Spine on outer margin strong.
Chela subovate, compressed dorsoventrally, broad and moderately long. Hand
entirely tuberculate. Inner margin of palm densely bearded with plumose setae. A
distinct ridge present on upper surface of both fingers. Fingers slightly gaping. Op-
posable margin of dactyl with a distinct excision at midlength, proximad of which
are three dome-shaped tubercles, distad of which are six; interspersed between these
are minute denticles. Lateral margin of dactyl with a proximal row of about eight
subsquamous ciliated tubercles which terminates slightly distad of excision on opposa-
ble margin. Distad of this row of tubercles is a row of about seven setiferous puncta-
tions. Upper surface of dactyl with a submedian ridge flanked proximad by a group
of several squamous tubercles and distad by scattered setiferous punctations along
upper opposable surface and by a row of similar punctations laterad. Lower surface
with a weak ridge flanked distad by a row of setiferous punctations and proximad by
small tubercles mesiad and punctations laterad. Opposable margin of immovable
finger with eight dome-shaped tubercles and a single large one projecting from lower
surface at base of distal fifth. Interspersed among these are scattered minute denticles.
Lateral margin with a poorly defined rounded ridge, most prominent near midlength
of finger. Proximal third of upper surface with scattered squamous ciliated tubercles;
distal portion with scattered setiferous punctations. Lower surface of immovable fin-
ger similar to that of dactyl.
Carpus of first pereiopod longer than wide (1.06-.80 cm.); shorter than inner
margin of palm of chela (1.20 cm.); a distinct longitudinal groove above. Dorso-
mesial surfaces tuberculate, otherwise sparsely punctate. Six prominent tubercles on
mesial surface and three on dorsal mesiodistal margin.
Merus of first pereiopod punctate laterad and mesiad. A group of scattered tuber-
cles present on upper surface, more numerous distad. Lower surface with an irregular
outer row of 18 spike-like tubercles and an inner row of 15.
Maxillipeds conspicuously heavily bearded.
Hooks present on ischiopodites of third and fourth pereiopods. Both pairs of
hooks extending proximad over basiopodites. Bases of coxopodites of fourth and fifth
pereiopods with strong outgrowths directed caudoventrad and cephaloventrad respec-
tively. Outgrowth on fourth strongest but simple; outgrowth on fifth bituberculate.
First pleopod reaching second pereiopod when abdomen is flexed. Terminal por-
tion of appendage with cephalodistal setae-bearing prominence rounded. Tip of
pleopod ending in four distinct parts. Mesial process corneous, long, and spiniform,
directed caudodistad and extending distad beyond the other terminal processes. Ce-
phalic process which is small, corneous, and subspiculiform arises from mesial sur-
face and directed distad. Caudal process corneous and blade-like, compressed later-
ally and lies at the caudolateral margin of the tip. Central projection also blade-like
and corneous, is compressed cephalocaudad, and directed obliquely (cephalomesiad
to cephalolaterad) across the center of the tip.
Male, Form II.-Differs from the male of the first form in the following respects:
all spines and tubercles greatly reduced in size and number; palm of chelae not beard-
ed; fingers not gaping, nor is there an excision on dactyl; all four terminals of pleopod
evident, but none are corneous, and all blunt; central projection and caudal process
indistinctly separated.
Allotypic Female.-Differs from the first form male in the following respects:
palm of chela not bearded and weaker; no excision on dactyl.
Annulus ventralis subovate; longitudinal excavation along midventral line; high
multituberculate ridges on either side of cephalic half; flattened caudad; sinus orig-







48 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

inmates dextrad of midventral line about one-third of the total length of annulus from
cephalic border; curves sharply laterad to cross midventral line and gently caudad
and finally dextrad to cut caudal margin.
Measurements.-Male (form I) Holotype: carapace, height 1.64, width 1.54,
length 3.23 cm.; areola, width .22, length .96 cm.; rostrum, width .48, length .60 cm.;
abdomen, length 3.21 cm.; right chela, length of inner margin of palm 1.20, width of
palm 1.10, length of outer margin of hand 2.82, length of movable finger 1.57 cm.
Female Allotype: carapace, height 1.16, width 1.12, length 2.42 cm.; areola, width
.17, length 1.73 cm.; rostrum, width .40, length .46 cm.; abdomen, length 2.60 cm.;
right chela, length of inner margin of palm .62, width of palm .55, length of outer
margin of hand 1.50, length of movable finger .81 cm.
Type Locality.-About 100 yards east of the Perdido River on U. S. Highway 90,
Escambia County, Florida. All of my specimens of this species were taken from bur-
rows in the type locality which is a small seepage area and drainage ditch. The soil
consists of clay overlain by about one to two feet of black muck.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes, one male (form I) and one female are deposited in the Museum of Com-
parative Zoology; one male (form I) and one female in the University of Michigan
Museum of Zoology. One male (form I), three females, and one immature female are
in my personal collection at the University of Florida.
Specimens at the Museum of Comparative Zoology, No. 3845, including one male,
form I, three females, three immature males, and one immature female, are also desig-
nated paratypes of P. escambiensis.
Remarks.-Faxon (1890: 621), under the heading "Additional local-
ity", listed the following for C. barbatus: "Escambia River at Flomaton, above
Pensacola, Florida. D. S. Jordan, B. W. Evermann, and C. H. Bollman (M.
C. Z.). One male, form I; five females, five young." In Faxon (1914: 414)
these specimens are listed as "Paratypes" of Cambarus barbatus. I have exam-
ined these "Paratypes" and found that they belong to Procambarus escambien-
sis. This species is closely related to Procambarus barbatus and might easily
be confused with it; however, a careful examination of the first pleopod of the
male readily distinguishes Procambarus escambiensis from all other known
species. In an attempt to get this crayfish in the Escambia River, I have col-
lected at several points along its course in Florida and Alabama, but have
been unable to find it either in the river or burrowing in its flood plains. In-
stead I found another closely related form, Procambarus shermani. It would
thus seem questionable as to whether the Flomaton, Alabama, labels on the
specimens of the species referred to barbatus by Faxon represent an actual
collection in this vicinity.
In its pubescent chelae and other general body structures, P. escambien-
sis most closely resembles P. pubischelae. The structure of the first pleopod
of the male, however, indicates that its closest relative is P. econfinae.
As was pointed out in the discussion of the distribution of the barbatus
group, P. escambiensis probably came into this region from the north and
west by a route that paralleled the movement of the econfinae and shermani
stocks east of it.
Specimens Examined.-I have examined a total of 11 specimens of Pro-






HOBBS-THE CRAYFISHES OF FLORIDA


cambarus escambiensis, from Escambia County, Florida, and all of my speci-
mens were taken during the month of April [4 d d (form 1), 1 d (form II),
6 9 9, and 1 9 (immature) ]. I have also seen and made camera lucida draw-
ings of the specimens, mentioned above, in the Museum of Comparative Zoolo-
gy.
Geographical and Ecological Distribution.-Procambarus escambiensis
is known from only two localities, on the western periphery of the range of
the barbatus group. In my collection I have specimens from a single locality,
a low wet place at the Perdido River on U. S. Highway 90. The specimens at
the Museum of Comparative Zoology were said to have been taken from the
Escambia River. Even if these specimens were taken from the Escambia River
region, which I questioned above, I do not believe that they came from the
river itself but probably from an overflow pool or nearby marshy area. All
of the species of this subgroup are burrowers, and although occasionally I
have seen them in open water I have never observed them in a large or swift
stream.
Cambarus byersi was taken from burrows along with P. escambiensis in
the type locality.
P. escambiensis is probably a secondary burrowing species. The simple
burrows were one and one-half to two feet deep in clay and muck; the water
table was from six inches to a foot below the surface. The chimneys over the
burrows were somewhat demolished by recent rains but were probably poorly
constructed.
Procambarus econfinae, sp. nov.
Plate II, Figs. 16-20; Plate XV; Maps 3, 4
Diagnosis.-Rostrum broadly lanceolate; areola broad and short with
space for about six punctations in narrowest part; however, sparsely punctate;
male with hooks on ischiopodites of third and fourth pereiopods; palm of che-
la of first form male not bearded along inner margin; postorbital ridges
terminating cephalad without tubercles or spines; no lateral spines present on
carapace. First pleopod of male, form I, reaching coxopodite of third pereio-
pod and terminating in four distinct, corneous parts; mesial process subspic-
uliform, directed distad, and extending distad of the other terminal process-
es. Cephalic process small and directed cephalodistad but hidden in lateral
view by the terminal setae. Caudal process compressed laterally, broadly
rounded, and forming a high ridge across the distal caudolateral margin of
the appendage. Central projection small and blade-like, and directed cephalo-
laterad. Cephalodistal setae-bearing prominence somewhat dome-like. Annu-
lus ventralis subovate; sinus originates on midventral line slightly caudad of
cephalic margin; turns gently sinistrad forming a broad arc back to the mid-
ventral line along which it proceeds almost to the caudal margin of the annulus.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen longer
than thorax (2.45-2.30 cm.). Height of carapace greater than width (1.17-1.15 cm.).
Greatest width of carapace slightly caudad of caudodorsal margin of cervical groove.




A







50 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Areola broad and short (2.52 times longer than wide); sparsely punctate, how-
ever, with three or four in narrowest part; sides parallel in middle. Cephalic section
of carapace about 2.7 times as long as areola (length of areola 27.4% of entire length
of carapace).
Rostrum broad-lanceolate, flattened above with low lateral ridges; margins con-
verging to tip; no lateral spines present. Upper surface of rostrum punctate. Sub-
rostral ridges evident in dorsal view for only a short distance near base.
Postorbital ridges well defined terminating cephalad without spines or tubercles.
Suborbital angle moderately well developed, rounded; branchiostegal spine small. No
lateral spines on sides of carapace. Surface of carapace punctate dorsad, granulate
laterad.
Abdomen longer than thorax (2.45-2.3 cm.).
Cephalic section of telson with two spines in each caudolateral corner.
Epistome broadly oval with slightly undulant cephalolateral margins.
Antennules of the usual form, not bearded. A spine present on ventral surface
of basal segment.
Antennae reaching caudal margin of first abdominal segment (apparently in-
jured but would probably extend no farther back than the third abdominal segment);
antennal scale broad; broadest slightly distad of middle. Spine on outer margin small.
Chela broadly ovate, compressed, heavy, entirely tuberculate, and with slender
fingers. Inner margin of palm not bearded but bearing an irregular row of about 10
tubercles. A well defined ridge present on upper surface of both fingers. Fingers
slightly gaping. Opposable margin of dactyl with 16 rounded tubercles between which
are a few scattered denticles. Lateral surface of dactyl ridge-like, bearing five tuber-
cles on basal half and about 10 setiferous punctations on distal half. A few scattered
punctations on upper proximolateral surface. Upper surface with a distinct submedian
ridge flanked by setiferous punctations. Lower surface of finger with a longitudinal
ridge near outer margin; otherwise with setiferous punctations. Opposable margin
of immovable finger with 12 rounded tubercles, and in addition a larger one extending
from lower surface at base of distal fourth. Minute denticles interspersed among these
tubercles. Outer margin of immovable finger with a distinct excavate ridge bearing
setiferous punctations. Upper and lower surfaces with submedian ridges flanked prox-
imally by tubercles and distally by setiferous punctations.
Carpus of first pereiopod longer than broad (.63-.51 cm.); shorter than inner
margin of palm of chela (.81 cm.); well defined longitudinal groove above. Dorso-
mesial and mesial surfaces studded with tubercles. Only four prominent ones on mesial
surface. Upper mesiodistal surface with three prominent tubercles.
Merus punctate mesiad and laterad with a few tubercles on upper distal surface.
Lower surface with an irregular outer row of about 12 tubercles and a more regular
inner one of about 14.
Hooks present on ischiopodites of third and fourth pereiopods. Hooks on fourth
small. Bases of coxopodites of fourth pereiopods with no prominent outgrowths;
corresponding position on fifth pereiopod with a small recurved plate-like hook.
First pleopod reaching base of third pereiopod. Cephalodistal setae-bearing
prominence somewhat dome-like. Mesial process long, subspiculiform, corneous, and
directed distad and extending distad of the other terminal processes. Cephalic process
long and corneous, directed cephalodistad but hidden in lateral view by the terminal
setae. Caudal process compressed laterally, broadly rounded and corneous, and form-
ing a high ridge across the distal caudolateral margin of the appendage. Central projec-
tion, blade-like, corneous, and directed cephalolaterad.
Male, Form II.-The description of the first form male will suffice for the male
of the second form if it is kept in mind that the tubercles are smaller but more spini-






HOBBS--THE CRAYFISHES OF FLORIDA


form. First pleopod with all of the processes evident and occupying the same relative
position; however, none of them are corneous, and all are much reduced in size.
Allotypic Female.-Likewise the description of the first form male is applicable
to the female except in the general form of the chelae, which are smaller and appar-
ently weaker.
Annulus ventralis subovate. Sinus originates on midventral line slightly caudad
of cephalic margin; turns gently sinistrad forming a broad arc back to the midventral
line along which it proceeds almost to the caudal margin of the annulus.
Measurements.-Male (form I) Holotype: carapace, height 1.17, width 1.15,
length 2.30 cm.; areola, width .25, length .63 cm.; rostrum, width .50, length .48 cm.;
abdomen, length .25 cm.; right chela, length of inner margin of palm .81, width of
palm .81, length of outer margin of hand 2.04, length of movable finger 1.17 cm.
Female Allotype: carapace, height 1.18, width 1.14, length 2.40 cm.; areola, width .21,
length .65 cm.; rostrum, width .44, length .49 cm.; abdomen, length 2.65 cm.; left
chela, length of inner margin of palm .62, width of palm .64, length of outer margin
of hand 1.50, length of movable finger .90 cm.
Type Locality.-Flatwoods in the northern part of Panama City, Bay County,
Florida. Here a large colony of this species occupies a small wet area lying between
the railroad and U. S. Highway 231 near the northern city limits. This area is subject
to flooding in wet weather; however, at the time I collected here the water table was
about one foot beneath the surface. All of my specimens were dug from burrows.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form I) and one female are deposited in the Museum of Compara-
tive Zoology; one male (form I) and one female in the University of Michigan Museum
of Zoology. Twelve males (form I), one male (form II), and 28 females are in my
personal collection at the University of Florida.
Remarks.-Procambarus econfinae is an extremely localized species ap-
parently occupying a total area of less than 200 square miles. My series of
specimens shows little variation. Its nearest relatives are P. escambiensis, P.
apalachicolae, and P. latipleurum. The first pleopod very closely approxi-
mates that of P. escambiensis while the palm of the chela is naked as in the
other species just mentioned.
Specimens Examined.-I have examined a total of 48 specimens, from
two localities in the vicinity of Panama City, Bay County, Florida [northern
part of city: June 1938-10 3 & (form I), 1 (form II), 139 9, 39 9
with young, 1 9 with eggs; the same locality: April 1938-2 S & (form I),
1 & (form II), 2 9 9 ; 7.9 miles northeast of Panama City on U. S. Highway
231: June 1938--3 6 (form I), 89 9, 29 9 with eggs, 29 9 with
young].
Geographical and Ecological Distribution.-Procambarus econfinae is
confined to a small area of the flatwoods on the small peninsula on which Pan-
ama City is located. The northern boundary of its range is marked by North
Bay and by a band of well drained soils of the Blanton-Norfolk group; the
western boundary is marked by a portion of the bay and soils of a similar
type along with the Lakewood-St. Lucie group on the southwest, while to the
south East Bay acts as a barrier. Only to the east is there no apparent barrier
delimiting the range of this species unless the brackish nature of the small
streams which cut across this area, and the marshy Wetappo Intercoastal






52 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Canal prevent an eastward migration. A discussion of the possible origin of
this species was taken up in the discussion of the barbatus group.
P. econfinae was collected along with P. pycnogonopodus in Bay County.
Almost certainly this species, like its relatives, is a secondary burrower.
Both of the localities from which it was taken are in the flatwoods where the
water table is only a foot or so below the surface, and standing water is present
in them during the rainy season. Here the crayfish construct simple burrows
consisting of a single passage downward, which never branches more than
once. The burrows are from one to three feet deep and are marked by low,
somewhat crude chimneys. In several instances I found a male and a female in
the same burrow. In one of these burrows a female carrying young was accom-
panied by a male; in another burrow having two branches a male and female
were taken from one branch and a lone female from the other. In still another
branched burrow a male was found in one pocket and a female in the other.
This is the only species I have collected in which a female carrying young was
accompanied by a male.

Procambarus latipleurum, sp. nov.
Plate III, Figs. 21-25; Plate XVI; Maps 3, 4

Diagnosis.-Rostrum narrow and acute lanceolate with or without lateral
spines; if spines are present they are small and close to the tip; areola relative-
ly broad with about six punctations in narrowest part; males with hooks on is-
chiopodites of third and fourth pereiopods, hooks on fourth bulbiform; palm
of chela of first form male not bearded within; chela slender and weak with
no (except sometimes one on immovable finger) well developed tubercles on
opposable surfaces of fingers; chela slender and weak; postorbital ridges
terminating cephalad in a small tubercle or spine; no lateral spines present on
carapace. First pleopod of male, form I, reaching coxopodite of third pereio-
pod and terminating in four distinct parts; mesial process corneous, extreme-
ly long, spiculiform, and directed distolaterad, distinctly curved; cephalic
process small and corneous, forming a hook which somewhat overhangs the
central projection; caudal process somewhat truncate but having a corneous
caudodistal edge; a distinct depression on lateral surface of main body of ap-
pendage at base of this process; central projection small and corneous, com-
pressed cephalocaudad and directed cephalodistad, somewhat overhung by the
cephalic process; cephalodistal setae-bearing surface rounded; pleura of ab-
domen conspicuously broad. Annulus ventralis almost square in outline, lat-
eral sides rounded; small acute tubercles present on cephalolateral margins;
sinus originates on cephalic margin along midventral line and runs caudad
slightly dextrad of the midventral line to middle of annulus where it makes a
sharp turn across the midventral line forming an arc and terminates just be-
fore it reaches the caudal margin, slightly sinistrad of the midventral line.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen and
thorax subequal in width. Height of carapace slightly greater than width (1.55-1.42







HOBBS-THE CRAYFISHES OF FLORIDA


cm.). Greatest width of carapace about midway between caudodorsal margin of cervi-
cal groove and caudal margin of carapace.
Areola of moderate width (4.65 times longer than wide) with six punctations in
narrowest part; sides parallel in middle. Cephalic section of carapace about 1.96 times
as long as areola (length of areola 33.8% of entire length of carapace).
Rostrum acute-lanceolate, flattened above with rather high marginal ridges; mar-
gins converging to tip; no lateral spines present. Upper surface of rostrum punctate.
Subrostral ridges well defined, barely visible in dorsal view almost to tip.
Postorbital ridge well defined, terminating cephalad in a small tubercle. Suborbi-
tal angle obtuse and rounded; branchiostegal spine well developed. No lateral spines
on sides of carapace. Surface of carapace punctate dorsad, weakly granulate laterad.
Abdomen much longer than thorax (3.42-2.75 cm.).
Cephalic section of telson with two spines in each caudolateral corner.
Epistome fan-shaped with a faint cephalomedian projection.
Antennules of usual form, but heavily bearded. A spine present on ventral surface
of basal segment.
Antennae reaching caudal margin of fourth abdominal segment; antennal scale
broad; broadest in middle. Spine on outer margin moderately strong.
Chela subovate, compressed, long and slender, entirely tuberculate; tubercles
thick. Inner margin of palm with two rows of about nine tubercles each. A very poorly
defined submedian ridge on upper surface of each finger. Fingers not gaping. Oppos-
able margin of dactyl with an upper and lower row of very small tubercles bordering
a wide band of minute denticles covering the entire margin. Upper surface of both fin-
gers bearing setiferous punctations except near base where there are a few scattered tu-
bercles. Median ridges poorly defined. Lateral and ventral surfaces with a few tubercles
at base, otherwise with setiferous punctations. The outstanding feature of the chela of
this species is the absence of well developed tubercles on opposable*margins of fingers.
Carpus of first pereiopod longer than wide (.72-.46 cm.); shorter than inner mar-
gin of palm of chela (.82 cm.); faint indication of a longitudinal groove above. Dor-
somesial, mesial, and ventral surfaces studded with tubercles, otherwise with setiferous
punctations. No acute tubercles present on ventral surface and about four prominent
ones on mesial surface.
Merus of first pleopod punctate laterad, tuberculate dorsad and mesiodistad. A
group of tubercles present on dorsodistal surface. Lower surface with a very irregular
outer row of about 15 tubercles and an inner row of about 16.
Hooks present on ischiopodites of third and fourth pereiopods, hooks on fourth
pereiopod bulbiform. Coxopodites of fourth and fifth pereiopods with distinct out-
growths; those on the fourth heavy and directed ventrolaterad; those on the fifth small
and slender and directed ventrad.
First pleopod reaching base of third pereiopod when the abdomen is flexed. Ter-
minal portion of appendage with the cephalodistal setae-bearing surface rounded. Me.
sial process corneous, extremely long, spiculiform, and directed distolaterad; distinct-
ly curved. Cephalic process small and corneous, forming a hook which somewhat over-
hangs the central projection. Caudal process somewhat truncate but having a corneous
caudodistal edge; a distinct depression on lateral surface of main body of appendage
at base of this process. Central projection small and corneous, compressed cephalocau-
dad and directed cephalodistad, somewhat overhung by the cephalic process.
Pleura of abdomen distinctly long and broad.
Male, Form 11.-Differs from the first form male in the following respects: small
lateral spines are present on rostrum near tip; tubercles reduced in size; postorbital
ridges terminate cephalad in small spines; epistome with less regular margins. First
pleopod of male with scarcely a distinction between any of the terminal processes ex-







54 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

cept the mesial which is large and claw-like, curved cephalodistad, and directed slightly
laterad.
Allotypic Female.-Differs from the first form male in the following respects:
epistome with an acute strong cephalomedian projection; chela broader and shorter.
Annulus ventralis almost square in outline; lateral sides rounded; small acute
tubercles present on cephalolateral margins; sinus originates on cephalic margin along
midventral line and runs caudad slightly dextrad of the midventral line to middle of
annulus where it makes a sharp turn across the midventral line forming an arc and ter-
minates just before reaching the caudal margin, slightly sinistrad of the midventral line.
Measurements.-Male (form I) Holotype: carapace, height 1.55, width 1.42,
length 2.75 cm.; areola, width .20, length .93 cm.; rostrum, width .40, length .58 cm.;
abdomen, length 3.43 cm.; right chela, length of inner margin of palm .82, width of
palm .64, length of outer margin of hand 2.06, length of movable finger 1.0 cm. Fe-
male Allotype: carapace, height 1.58, width 1.45, length 2.82 cm.; abdomen, length
3.41 cm.; areola, width .21, length .87 cm.; rostrum, width .42, length .70 cm.; right
chela, length of inner margin of palm .55, width of palm .61, length of outer margin of
hand 1.62, length of movable finger .97 cm.
Type Locality.-Roadside excavation and intermittent stream in flatwoods, 5.8
miles west of Weewahitchka on State Highway 52, Gulf County, Florida. Most of my
specimens were taken with a seine from the excavation in which the water was from one
to three feet deep. A few specimens were dug from simple burrows, never with more
than two branches, nor more than two feet deep. The chimneys of most of these burrows
were neatly formed.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form II) and a female are deposited in the Museum of Compara-
tive Zoology; one male (form II) and one female in the University of Michigan Mu-
seum of Zoology. One male (form I), seven males (form II), two females, three imma-
ture males, and two immature females are in my personal collection at the University
of Florida.
Remarks.-All of my specimens were taken within a radius of 13 miles
of Weewahitchka.
This species has its closest affinities with P. apalachicolae, P. rathbunae,
P. pubischelae, and P. barbatus. It is indeed surprising to find this local form
in the Apalachicola Flatwoods without apparent barriers to separate it from
P. apalachicolae. The break between these two species occurs somewhere in
the 25 mile wide extent between Weewahitchka and Port St. Joe. Westward
it is probably limited by the well drained area of Blanton-Norfolk soils.
Specimens Examined.-I have examined a total of 54 specimens of Pro-
cambarus latipleurum, from Gulf County, Florida, collected during the
months of April, October, and November. [April-21 d' (immature) and
119 9 (immature); October-2 2 d (form I), 9' 6 (form II), 29 9,
3 S 5 (immature), and 2 9 9 (immature); November-1 6 (form II) and
39 9.]
Geographical and Ecological Distribution.-Procambarus latipleurum
has been taken only in the flatwoods around Weewahitchka, and judging from
the fact that a closely related species has been collected in abundance both
south and west of its range and that ample barriers bar its migration north-
ward and eastward, it is probably confined to a small area in this region.






HOBBS-THE CRAYFISHES OF FLORIDA


This species was collected along with P. pycnogonopodus, P. pygmaeus,
and P. kilbyi.
This species is most likely a secondary burrower and is apparently con-
fined to flatwoods conditions where it occupies temporary bodies of water.
Its burrows are simple, never having more than two branches, usually consist-
ing of a single vertical tunnel which penetrates the water table within two feet
below the surface of the ground. The chimneys vary in height from one to four
inches and are neatly formed, having vertical rather than sloping outer walls.

Procambarus apalachicolae, sp. nov.
Plate III, Figs. 26-30; Plate XVII; Maps 3, 4

Diagnosis.-Rostrum broad, short, and without lateral spines; areola
relatively broad with about four punctations in narrowest part; male with
hooks on ischiopodites of third and fourth pereiopods; palm of chela of first
form male not bearded within; chela heavy with relatively short fingers; post-
orbital ridges terminating cephalad without lateral spines or tubercles; no
lateral spines present on carapace. First pleopod of first form male reaching
coxopodite of third pereiopod and terminating in four distinct parts; mesial
process acute with a corneous tip (extending caudodistad beyond the other
terminal processes); cephalic process small, corneous, blade-like, and trun-
cate distad; caudal process large, non-corneous, and thumb-like; central pro-
jection corneous, small, subtriangular, compressed cephalocaudad, and situat-
ed obliquely cephalomesiad to caudolaterad; cephalodistal setae-bearing sur-
face rounded. Annulus ventralis with cephalic and caudal margins concave,
lateral margins convex; sinus originates on cephalic surface on midven-
tral line, curves sinistrad forming a wide arc and terminates just cephalad of
midcaudal margin; a few small tubercles on each cephalolateral surface.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen narrow-
er than thorax (.89-1.0 cm. in widest parts respectively). Height of carapace slightly
greater than width in region of caudodorsal margin of cervical groove (1.05-1.0 cm.).
Greatest width of carapace slightly caudad of caudodorsal margin of cervical groove.
Areola moderately broad (3.7 times longer than wide) with only two punctations
in narrowest part (however, there is room for four or five, were they spaced as they are
elsewhere in the areola); sides not parallel. Cephalic section of carapace about 2.48
times as long as areola (length of areola about 28.7% of the entire length of carapace).
Rostrum flattened above, almost reaching base of distal segment of peduncle of
antennule; margins converging to tip; no lateral spines present. Upper surface of ros-
trum punctate. Marginal ridges strongly developed and terminate before reaching tip
of rostrum. Subrostral ridges prominent and evident in dorsal view to tip of rostrum.
Postorbital ridges well defined, terminating cephalad without forming a tubercle
or spine. Suborbital angle obtuse; branchiostegal spine small. No lateral spines or tu-
bercles on sides of carapace. Surface of carapace punctate dorsad, weakly granulate
laterad.
Abdomen longer than thorax (2.23-1.95 cm.).
Cephalic section of telson with four spines in the sinistral and three in the dextral
caudolateral corners.






56 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Epistome broadly ovate with a small cephalomedian projection; margins slightly
undulate.
Antennule of usual form with a small spine present on ventral side of basal seg-
ment.
Antennae reaching caudad to third abdominal segment; antennal scale moderately
broad; broadest in middle. Spine on outer margin well developed.
Chela subovate, compressed dorsoventrally, moderately heavy, and entirely tuber-
culate. Inner margin of palm not bearded but bearing an irregular row of about 11 tu-
bercles. A distinct ridge present on upper surface of both fingers. Fingers not gaping.
Opposable margin of dactyl with six rounded tubercles on proximal three-fifths and
several rows of minute denticles distad of these; denticles are also interspersed between
tubercles. Upper distal margin with a few minute tubercles above the denticles. Lateral
surface of dactyl with a row of eight tubercles and several setiferous punctations. Up-
per and lower surfaces each with a submedian longitudinal ridge flanked by scattered
tubercles proximad and setiferous punctations distad. Opposable margin of immov-
able finger with ten rounded tubercles and a larger one projecting from lower margin
at base of distal fifth; interspersed between these tubercles is a large number of minute
denticles; lateral surface with an excavate ridge bearing setiferous punctations; upper
and lower surfaces with a submedian ridge flanked proximally by scattered tubercles,
distally by setiferous punctations.
Carpus of first pereiopod longer than wide (.63-.43 cm.); a poorly defined longi-
tudinal groove above. Dorsomesial and mesial surfaces tuberculate, otherwise punc-
tate. Mesial surface with several scattered tubercles. Cephalomesial and dorsomesial
margins with a continuous row of about seven tubercles.
Merus of first pereiopod punctate on lateral and proximomesial surfaces. Upper
surface with scattered tubercles over entire length. Lower surface with an irregular out-
er row of about 15 tubercles and an inner row of about 17.
Hooks present on ischiopodites of third and fourth pereiopods. Coxopodites of
fourth and fifth pereiopods with outgrowths. Those on the fourth heavy, subtriangular,
and directed ventrolaterad; those on the fifth thin, somewhat rounded, and similarly
directed.
First pleopod reaching coxopodite of third pereiopod when the abdomen is flexed.
Terminal portion of appendage with cephalodistal setae-bearing prominence rounded.
Mesial process acute, with corneous tip, and extending distad beyond the other termi-
nal apices. Cephalic process, small, slender, truncate and corneous, arises from the me-
sial surface and is directed distad. Caudal process thumb-like, non-corneous, and di-
rected caudodistad. Central projection corneous, small, triangular, compressed cepha-
locaudad, and extends obliquely, cephalomesiad to caudolaterad.
Male, Form II.-Differs from the first form male in the following respects: tu-
bercles fewer in number; many, however, more spiniform; subrostral ridges evident
in dorsal aspect for only a short distance from base and at tip of rostrum. First pleopod
with all terminals evident, but much reduced and non-corneous.
Allotypic Female.-Differs from the first form male in the following respects:
ridges of rostrum extending to tip; subrostral ridges evident in dorsal aspect along
basal half of rostrum; chela proportionately smaller; tubercles somewhat reduced in
size.
Annulus ventralis with cephalic and caudal margins concave, lateral margins con-
vex. Sinus originates on cephalic surface on midventral line; curves sinistrad forming a
wide are and terminates just cephalad of midcaudal margin. A few small tubercles on
each cephalolateral surface.
Measurements.-Male (form I) Holotype: carapace, height 1.05, width 1.0, length
1.95 cm.; areola, width .15, length .56 cm.; rostrum, width .31, length .62 cm.; abdo-






HOBBS-THE CRAYFISHES OF FLORIDA


men, length 2.23 cm.; right chela, length of inner margin of palm .62, width of palm .54,
length of outer margin of hand 1.57, length of movable finger .86 cm. Female Allotype:
carapace, height 1.30, width 1.21, length 2.34 cm.; areola, width .23, length .69 cm.;
rostrum, width .38, length .50 cm.; abdomen, length 1.95 cm.; right chela, length of
inner margin of palm .53, width of palm .63, length of outer margin of hand 1.42,
length of movable finger .85 cm.
Type Locality.-Roadside ditch in wire grass flatwoods, 11.1 miles west of Bea-
con Hill, U. S. Highway 98, Bay County, Florida. All of the specimens were dug from
simple burrows in a sandy, mucky soil.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form I) and one female are deposited in the Museum of Compara-
tive Zoology; one male (form I) and one female in the University of Michigan Museum
of Zoology. Eight males (form I), 12 females, and one immature male are in my per-
sonal collection at the University of Florida.
Remarks.-Procambarus apalachicolae occupies a rather peculiar range
when the range of its relatives, P. econfinae and P. latipleurum, are taken into
consideration. While the ranges of these three species are not entirely separ-
ated by apparent physiographic barriers, I find no instance of intergradation.
The nearest relatives of this species are almost certainly P. latipleurum
and P. rathbunae, and it is possible that P. apalachicolae will prove to be a
subspecies of the latter.
Specimens Examined.-I have examined a total of 85 specimens, from
the following counties in Florida: Bay, Franklin, Gulf, and Walton. All of
these localities were in the flatwoods along the coast from the Apalachicola
River westward to the mouth of Choctawhatchee Bay.

SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
6I 6 12 9 1
C II 3 1
9 12 13 6
9 (eggs) 1 2
9 (young) 1
S (immature) 7 1
9 (immature) 14

Geographical and Ecological Distribution.-As was pointed out above
Procambarus apalachicolae, P. econfinae, and P. latipleurum present an intri-
cate pattern of distribution in the Apalachicola flatwoods. The ranges of the
last two have already been discussed, but it is necessary to recall the limits of
their ranges in connection with that of P. apalachicolae. The three species to-
gether occupy practically all of the available flatwoods between the Apalachi-
cola River on the east and the Choctawhatchee River and Bay on the west. P.
latipleurum occupies the northeastern part of the area, and there is no known






58 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

barrier to the south to separate it from P. apalachicolae. In collecting along
State Highway 6 between Weewahitchka and Port St. Joe the most northern
locality for P. apalachicolae was 5.6 miles north of Port St. Joe, while the
most southern record for P. latipleurum was 12.6 miles south of Weewahitch-
ka; hence, along this road there is a zone of six miles from which neither spe-
cies has been taken. The frequently inundated marshes bordering the Wetappo
Intercoastal Canal occur within this area and may be a limiting factor in the
distribution of these two species. Thus, on the basis of the material at hand it
seems as though the northern limit of the range of P. apalachicolae is along
the Wetappo Intercoastal Canal and the East Bay. It is also consistent with the
evidence at hand that this same marsh bordered canal to the west in Bay Coun-
ty may also mark the limits of the ranges of apalachicolae and econfinae.
Apparently the range of P. apalachicolae is completely broken by St. An-
drews and North bays and a strip of well drained soils to the north of North
Bay. In western Bay County and in Walton County apalachicolae seems to be
scattered throughout the flatwoods as far as 12 miles east of Destin. The south-
ern boundary of the range is apparently marked by the salt marshes and dunes
along the coast.
P. apalachicolae was taken along with P. pycnogonopodus, P. rogersi
rogersi, and P. rogersi (intergrades) in Bay County, and with P. rogersi (in-
tergrades) in Franklin County as well as in Gulf County.
Like the other members of the barbatus group, P. apalachicolae is a sec-
ondary burrower. Typically, it is a flatwoods species living in depressions in
which water is present during some part of the year. When the depression be-
comes dry the crayfish construct simple burrows with small chimneys. These
chimneys show a large variety of patterns; some are apparently very carefully
constructed, with straight vertical walls; others are low and have the appear-
ance of small mud piles that show little resemblance to a typical chimney. In
most of the stations where this species has been collected the water table was
about one foot below the surface, but in several instances it was as deep as
three feet. Burrows have been found in a wide variety of soils: in sands, clays,
and muck. In most instances wire grass has been present in the vicinity where
this species has been taken, and in at least one locality gallberries and palmet-
tos were abundant.
In a small roadside ditch having a vertical bank and in which water was
standing several specimens of this species had dug burrows into the bank,
with openings both above and below the water table; several openings were
as high as three feet above the existing water level, and most of these had tun-
nels which were about one foot below the level of the water.
In two instances specimens of apalachicolae were taken from standing
water; several specimens were dipped from a bed of Juncus repens in a small
swamp pool, and others were taken in the isolated pools of a drying swamp
stream.






HOBBS-THE CRAYFISHES OF FLORIDA


Procambarus rathbunae (Hobbs)
Plate III, Figs. 31-35; Maps 3, 4

Cambarus rathbunae Hobbs 1940a, Proc. U. S. Nat. Mus. 89 (3097): 414-
418, Fig. 21; also 387, 389.
Procambarus rathbunae Hobbs, Amer. Mid. Nat. (in press-c).
Diagnosis.-Rostrum without lateral spines; areola relatively broad
with three or four punctations in narrowest part; male with hooks on ischiopo-
dites of third pereiopods only; palm of chela not bearded along inner margin;
postorbital ridges terminating cephalad without spines; no lateral spines pres-
ent on carapace. First pleopod of male reaching coxopodite of second perei-
opod and terminating in four distinct parts; mesial process extends distad
beyond the rest of the terminal elements in a caudodistal direction; cephalic
process small and corneous and arises from mesial surface of the appendage
and is directed distad; caudal process, making up the caudolateral surface of
the tip, forms a distinct corneous ridge which is directed cephalodistad; cen-
tral projection small, subtriangular, corneous, and somewhat compressed
cephalocaudad with a small rounded plate laterad. Cephalodistal setae-bear-
ing prominence sloping. Annulus ventralis broadly subovate with the smaller
ends in the longitudinal axis; however, truncate both cephalad and caudad;
excavate along midventral line with a transverse ridge on either side at mid-
length which is abrupt caudad but gradually sloping cephalad; cephalic por-
tion on either side of excavation tuberculate; sinus originates about one-third
of the length of the annulus from the cephalic border, slightly dextrad of the
midventral line, runs caudosinistrad barely crossing the midventral line where
it abruptly turns and forms a straight line to the midcaudal margin.
Remarks.-Procambarus rathbunae is most nearly related to P. apala-
chicolae; in fact a few specimens of the latter species from the southwestern
part of Bay County and the southeastern part of Walton County are structur-
ally intermediate between the two species. It seems doubtful that they repre-
sent intergrades, however, because continuous barriers of salt water and sand
hills appear to completely isolate the two. The possibility of intergradation
cannot be dismissed, however; a thorough search should be made in Walton,
Okaloosa, and Washington counties before it can be finally determined wheth-
er the specimens from the Destin Region are really a variant of Procambarus
apalachicolae, or whether they are apalachicolae x rathbunae. P. pubischelae
and P. escambiensis are also near relatives of this species.
Specimens Examined.-I have examined a total of 27 specimens of Pro-
cambarus rathbunae, from Okaloosa and Holmes counties, Florida.
Seasonal Data.-I have collected this species during the months of April,
May, and October. In April 1938 two males (form I), five males (form II),
and nine females were taken from burrows and a ditch at Milligan. In May
1942 one male (form I) and one female were collected eight miles west of
Ponce de Leon on U. S. Highway 90. In October 1941 two males (form II),






60 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

four females (one of which was carrying young), and three immature females
were taken 3.6 miles south of Crestview on State Highway 54.
Geographical and Ecological Distribution.-It is likely that this species
is widespread in the flood plains of the Yellow River and seepage areas and
flatwoods along its tributaries in Santa Rosa and Okaloosa counties. It is pos-
sible that it will be found in the Blackwater River drainage, for there are
continuous low marshes between the mouths of the two rivers which may pos-
sibly form a highway for this species. I am unwilling at present to make any
statement concerning its possible distribution in the Choctawhatchee drain-
age, for I have it from only one locality, while P. hubbelli, another member
of the barbatus group occupying similar habitats, has been found to be wide-
spread in this river system in Florida.
P. okaloosae was captured in the type locality of P. rathbunae both in the
aquatic vegetation of a roadside ditch and from burrows adjacent to those of
P. rathbunae.
All except one of the specimens of P. rathbunae taken at Milligan were
dug from simple burrows in the roadside ditches. This single specimen was
caught in a dip net dragged along the ditch. The burrows ranged in depth from
six inches to a foot below the surface of the ground. Several of the burrows
were open, though most of them were marked by closed chimneys. The soil is
a sand-clay mixture, and the ground is covered with a thick mat of grass. Some
sections of the ditch held water, but at this time most of it was dry.
Near Crestview and Ponce de Leon P. rathbunae was taken from a slop-
ing seepage area along the margins of a bay around which wire grass and
pitcher plants were common. The burrows were extremely complex, closely
resembling those of the rogersi subspecies. They were from one to three feet
in depth and branched several times. Some of them had several chimneys, most
of which were poorly formed, and in several places where excavations had
been made along the hillside open burrows were numerous in their walls. In
no place in the area was the water table more than a foot below the surface.
While collecting at Milligan I encountered an open burrow in which a
male was either attempting to enter or else being driven out by the female. I
could see both crayfish since the opening of the burrow was unusually large.
The male seemed to be attempting to enter, and the female, facing him, was
apparently blocking the burrow.
On the basis of my limited observations the habits of P. rathbunae are
similar to those of the other better known species of the barbatus group.
THE KILBYI, SHERMANI, AND HUBBELLI SUBGROUPS
The three remaining species of the barbatus group are so distinct from
the members of the barbatus subgroup and from one another that each necessi-
tates a separate subgroup to indicate its isolated position.
THE SHERMANI SUBGROUP
A separate shermani subgroup is necessitated to receive the very disjunct
Procambarus shermani.






HOBBS-THE CRAYFISHES OF FLORIDA


Procambarus shermanis, sp. nov.
Plate III, Figs. 36-40; Plate XVIII; Maps 3, 4

Diagnosis.-Rostrum without lateral spines; areola relatively narrow
(narrower than in any other species in the barbatus group) with two puncta-
tions in the narrowest part; male with hooks on ischiopodites of third and
fourth pereiopods; palm of chela of first form male usually bearded; postor-
bital ridges terminating cephalad in small tubercles; a lateral spine present on
carapace. First pleopod of first form male reaching coxopodite of second perei-
opod and terminating in four distinct parts; mesial process acute and corne-
ous; cephalic process and central projection blade-like, corneous, and com-
pressed laterally; both directed caudodistad; caudal process subspatulate and
directed laterodistad with a small corneous ridge arising on the lateral surface
and flanking it along its caudal margin; no cephalodistal setae-bearing promi-
nence; setae borne on the slightly curved cephalodistal margin. Annulus ven-
tralis subovate with the greatest length in the longitudinal axis; deep sub-
median furrow flanked on either side by high multituberculate wall, deepest
portion of furrow in center of annulus; sinus originates in bottom of furrow
slightly dextrad of midventral line, runs gently caudosinistrad, crossing mid-
ventral line where it turns caudodextrad terminating just before reaching cau-
dal margin, slightly dextrad of midventral line.
Holotypic Male, Form I.-Body subovate, compressed laterally. Abdomen nar-
rower than thorax (1.25-1.40 cm. in widest parts respectively). Width and depth of
carapace subequal in region of caudodorsal margin of cervical groove. Greatest width
of carapace slightly caudad of caudodorsal margin of cervical groove.
Areola relatively narrow (12.9 times longer than wide) with two punctations in
narrowest part; sides parallel in middle. Cephalic section of carapace about 1.89 times
as long as areola (length of areola 34.6% of entire length of carapace).
Rostrum acute lanceolate, flattened above but with conspicuously high lateral
ridges, reaching midlength of distal segment of peduncle of antennule; margins con-
verging to tip; no lateral spines present. Upper surface of rostrum with shallow mo-
saic furrows. Subrostral ridges weakly developed and not evident in dorsal view.
Postorbital ridge very prominent and terminating cephalad in a tubercle. Subor-
bital angle obtuse, rounded; branchiostegal spine well developed. A single lateral
spine present on sides of carapace, flanked by several small tubercles. Surface of cara-
pace punctate dorsad, strongly granulate laterad.
Abdomen and thorax subequal in length.
Cephalic section of telson with three spines in the right and four in the left caudo-
lateral corners.
Epistome broadly oval with a small, acute cephalomedian projection.
Antennule of usual form. A spine present on ventral side of basal segment.
Antennae reaching caudad to third abdominal segment; antennal scale of mod-
erate length, widest slightly distad of middle. Spine on outer margin strong.

'This species is named for Professor H. B. Sherman, Department of Biology, University of
Florida, who has frequently aided me in my work on the Florida crayfishes, not only in the loan of
personal equipment but also in invaluable criticisms and suggestions.







62 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Chela subovate, flattened dorsoventrally, of moderate length and width. Hand
entirely tuberculate. Inner margin of palm bearded with plumose setae. A distinct ridge
present on upper surface of both fingers. Fingers not gaping. Opposable margin of
dactyl convex, with a faint indication of an excision at end of proximal third, proximad
of which are five dome-shaped tubercles, distad of which are 14, all of which are inter-
spersed with minute denticles. Lateral margin of dactyl with a row of eight prominent
tubercles. Upper surface of dactyl with a submedian ridge at the base of which is a
group of small squamous tubercles, more numerous laterad; a single row of setiferous
punctations along lateral margin of ridge, likewise a single row along upper surface
of opposable margin. Lower surface with a prominent, though not distinctly defined,
ridge flanked proximad by a few scattered tubercles and distad by several rows of setif-
erous punctations. Opposable margin of immovable finger with 20 dome-shaped tu-
bercles and a single large, more acute one projecting from lower surface at base of
distal fourth. Interspersed among these are scattered minute denticles. Lateral margin
of immovable finger with a ridge flanked proximally by ciliated squamous tubercles
and distally by setiferous punctations. Lower surface of immovable finger with a prom-
inent ridge, and ventromesial surface distinctly excavate. A row of setiferous puncta-
tions runs the entire length of the finger just below opposable margin, and a few tuber-
cles are present on inner side of ridge at base.
Carpus of first pereiopod longer than wide (.73-.59 cm.); shorter than inner
margin of palm of chela (.89 cm.); a distinct longitudinal groove above. Dorsomesial
and mesial surfaces tuberculate. Eight prominent tubercles on mesial surface, twelve
on dorsomesial surface, and an additional very strong one on distal dorsomesial mar-
gin. Ventral and ventromesial surfaces each with a single spine on distal margin.
Merus of first pereiopod punctate laterad and proximomesiad, otherwise tubercu-
late. Two very prominent spines on upper surface near distal end; one prominent spine
on same surface at distal margin, below which are two prominent tubercles. Lower
surface with an irregular outer row of about 20 tubercles and an inner row of about 16
spike-like tubercles.
Maxillipeds bearded.
Hooks present on ischiopodites of third and fourth pereiopods. Bases of coxopo.
dites of fourth and fifth pereiopods with strong outgrowths directed caudoventrad and
cephaloventrad respectively.
First pleopod reaching base of second pereiopod when abdomen is flexed; with
no cephalodistal setae-bearing prominence; setae borne on the slightly curved cephal-
odistal margin. Tip ending in four distinct parts. Mesial process acute, corneous, and
does not extend beyond the other terminal processes. The other three elements are sub-
equal in length; cephalic process blade-like and corneous, as is the central projection;
both are compressed laterally and directed slightly caudodistad. Caudal process sub-
spatulate and corneous, directed distolaterad with a small corneous ridge arising on the
lateral surface and flanking it around its caudal margin. The cephalic and caudal pro-
cesses and the central projection together form a thin corneous blade across the lateral
tip of the appendage.
Male, Form 11.-Differs from the male of the first form in the following respects:
all tubercles greatly reduced in size and number; however, all spiny portions accen-
tuated; palm of chela lightly bearded; postorbital ridges bearing spines. First pleopod
with truncate terminals; three small terminals representing the cephalic and caudal
processes and the central projection; mesial process shortened and less acute.
AUotypic Female.-Differs from the first form male in the following respects:
palm of chela not bearded and weaker; cephalic margin of epistome broken by several
tubercles.
Annulus ventralis subovate with the greatest length in the longitudinal axis; deep
submedian furrow flanked on either side by a high multituberculate wall; deepest por.






HOBBS-THE CRAYFISHES OF FLORIDA


tion of furrow in center of annulus; sinus originates in bottom of furrow slightly dex-
trad of midventral line; runs gently caudosinistrad, crossing midventral line where it
turns caudodextrad terminating just before reaching caudal margin, slightly dextrad of
midventral line.
Measurements.-Male (form I) Holotype: carapace, height 1.40, width 1.40,
length 2.98 cm.; areola, width .08, length 1.03 cm.; rostrum, width .50, length .40 cm.;
abdomen, length .30 cm.; right chela, length of inner margin of palm .89, width of
palm .78, length of outer margin of hand 2.30, length of movable finger 1.27 cm. Fe-
male Allotype: carapace, height 1.90, width 1.84, length 4.02 cm.; areola, width .06,
length 1.48 cm.; rostrum, width .60, length .90 cm.; abdomen, length 4.00 cm.; right
chela, length of inner margin of palm .82, width of palm .80, length of outer margin of
hand 2.46, length of movable finger 1.47 cm.
Type Locality.-About 12 miles southwest of Jay, Santa Rosa County, Florida, in
the Escambia River swamp along McCaskill's Mill Creek. The crayfish were taken from
flood plain pools among dead leaves and other debris. Burrows were numerous in the
bottoms of these pools, and at night the crayfish could be seen at the mouths of them. I
collected here both at night and during the day, using a headlight and dip net at night
and a seine during the day. A larger catch was obtained during the daylight hours. At
the time of my visit to the swamp the river was flooded, and even during my brief stay
of less than 24 hours the water had risen so that practically all of the isolated pools
were connected and a swift current was flowing through many of them.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States NemRal Museum. Of the remaining
paratypes one male (form I) and one female are dep eiiria the Museum of Compara-
tive Zoology; one male (form I) and one female in the Uunwhety of Michigan Museum
of Zoology. Two males (form I), one male (form II), fewr femai ,22 iNm nature
males, and 21 immature females are in my personal collection at the Uzlva of Flr-.
ida.
Remarks.-The first specimens of this species that I saw were canuht by
Mr. Ray Boles in swamp pools in the river bottom a few miles south of the type
locality. One second form male and three immature females were taken along
with a series of P. okaloosae and P. bivittatus. It was not until April 1942 that
I succeeded in getting a series of this species.
P. shermani somewhat bridges the gap between the barbatus section and
the section of blandingii. In the structure of the chelae and body conformation,
with the exception of the presence of lateral spines on the carapace, it fits
nicely into the barbatus section. The pleopod, however, is intermediate be-
tween that found in the two sections. The cephalic process has moved forward
and extends from the cephalic margin rather than the mesial margin. Also the
caudal process is corneous and very similar to that found in the blandingii
subspecies and hayi.
Procambarus shermani like the rest of the species of the barbatus group
probably invaded this region from the north, paralleling the migration of
escambiensis in the Perdido River system.
The closest relatives of this species seem to be Procambarus escambien-
sis, P. latipleurum, P. rathbunae, P. econfinae, P. apalachicolae, and P. pubis-
chelae.
Specimens Examined.-I have examined a total of 57 specimens of Pro-
cambarus shermani, from Santa Rosa County, Florida, and all of them were






64 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

taken within three miles of the type locality. Four specimens were collected
in May 1941: 1 d (form II) and 3 9 (immature). In April 1942 the fol-
lowing were taken: 5 & & (form I), 1 d (form II), 7 9 22 d & (imma-
ture), and 18 9 (immature).
Geographical and Ecological Distribution.-Procambarus shermani has
been taken only from the Escambia River swamp. The fact that a closely allied
species occurs in the Perdido River drainage system and another in the Yellow
River drainage system, both of which occupy the same general type of habitat
as that from which shermani was taken, would seem to indicate that the latter
is confined to the Escambia River drainage.
P. shermani was collected along with P. blandingii acutus, C. diogenes,
and C. species incertis in Santa Rosa County.
Practically nothing is known of the habits of this species. It is apparently
confined to swamp pools or sluggish water, for it was not taken in McCaskill's
Mill Creek which is a clear sand-bottom, moderately flowing stream, but was
relatively abundant in the small pools near its margins. It apparently burrows
in these pools, but even in the middle of the day specimens may be taken from
leaf mats and other debris.
As was pointed out under the heading "Type Locality," the Escambia
River is subject to considerable fluctuation in water level. In October 1941 I
was able to walk all over the swamp, even down to the river's edge, but in April
1942 the p T'r was so high that I could not even get into the swamp at the
place I visited before, and it was only with considerable difficulty that col-
lecting was carried out in the swamp pools farther upstream along McCaskill's
Mill Creek. The creek itself was completely obliterated in the mass of water in
the swamp. Collecting could be successfully carried out in only a few isolated
pools.
THE KILBYI SUBGROUP
A separate kilbyi subgroup is necessitated to receive the very disjunct
Procambarus kilbyi.

Procambarus kilbyi (Hobbs)
Plate IV, Figs. 41-45; Maps 3, 4

Cambarus kilbyi Hobbs 1940a, Proc. U. S. Nat. Mus. 89 (3097): 410-414,
fig. 20; also 387-389, 418.
Cambarus kilbyi Hobbs (in press-a).
Procambarus kilbyi Hobbs (in press-c); Hobbs (in press-d); Hobbs (in
press-b).
Diagnosis.-Rostrum without lateral spines; areola relatively broad;
male with hooks on ischiopodites of third and fourth pereiopods; palm of
chela not bearded along inner margin; postorbital ridges terminating ce-
phalad without spines; no lateral spines present on carapace. First pleopod of






HOBBS-THE CRAYFISHES OF FLORIDA


first form male reaching coxopodite of third pereiopod and terminating in
four distinct parts; mesial process, the largest of the four, corneous, subspat-
ulate, and bent near midlength at about a 50 degree angle to the main shaft of
the appendage; cephalic process arises from the mesial surface of the append-
age and is slender, subspiniform, and truncate; caudal process somewhat
rounded, compressed laterally, and extends in a ventrocaudal direction from
the caudolateral margin of the appendage; central projection small, corneous,
triangular, and compressed cephalocaudad; cephalodistal setae-bearing
prominence knob-like. Annulus ventralis subovate; sinus originates on cephal-
ic border slightly sinistrad of midventral line, curves gently dextrad of mid-
ventral line, then somewhat more sharply sinistrad across the midventral line,
and finally to the midcaudal margin.
Remarks.-Although P. kilbyi is decidedly variable and occupies a rela-
tively large area, unlike several of the other species of the barbatus group, it
is not on the basis of the material at hand divisible into local variants. While
the general appearance of any specimen enables one to tell with a reasonable
degree of certainty from what locality it was taken, I do not have sufficient ma-
terial to determine the extent of these variants in a single local population.
There is a tendency for the outline of the annulus ventralis to become
more angulate in specimens from the southeastern portion of the range. The
two extremes in the shape of this structure are found in specimens from Cal-
houn and Levy counties. In Calhoun County the annulus is subovate with the
angles rounded, while in the Levy County females it is decidedly angulate.
The interesting fact is that there is a somewhat gradual transition between the
two types (although not without exceptions) in the intervening region.
While the first pleopod of the male shows considerable variation, all of
these differences seem to be individual ones.
Specimens Examined.-I have examined a total of 305 specimens of
Procambarus kilbyi, from the following counties in Florida: Calhoun, Frank-
lin, Gulf, Jackson, Jefferson, Lafayette, Leon, Levy, Liberty, Madison, Tay-
lor, and Wakulla.
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
I 5 4 10 9 3 2 2
acII 5 15 4 7 7 5 6
9 5 9 11 16 5 2 1 6 7 5
9 (eggs) 3 1
9 (young) 4 2
6 (immature) 8 4 37 1 1 5 2 4
9 (immature) 11 7 46 4 2 9 4 4
Geographical and Ecological Distribution.-Procambarus kilbyi is a
Florida endemic and seems to be confined to the coastal flatwoods from Levy
County westward into Calhoun and Gulf counties. Exactly what, if anything,
has checked the westward migration of this species has not been determined;







66 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

there is no apparent barrier to the west of the most western known record. The
Apalachicola River probably formed a temporary barrier, but now that kilbyi
has established itself west of the river I can see nothing which can restrict its
farther westward migration unless P. apalachicolae proves too much of a
competitor. It is possible that P. kilbyi will migrate farther into Bay, Gulf,
and Walton counties in the future. The eastern boundary of its range is limited
by the high, dry soils which extend from Hamilton County southward through
Marion County into the center of the state. The southern limit of the range is
not definitely known, though the record from Levy County represents the most
southern locality (about 6 miles southwest of Bronson [State Highway 19]).
It is very possible that this species extends farther south along the coast; cer-
tainly, however, it goes no farther south than Hernando County. The south-
western limit of the range is apparently marked by the dunes and salt marsh-
es along the gulf.
P. kilbyi has been collected with P. rogersi campestris in Leon and Wa-
kulla counties, with P. rogersi ochlocknensis in Liberty County, with P. rogersi
rogersi in Calhoun County, with P. rogersi intergrades in Franklin County,
with P. pycnogonopodus in Gulf, Jackson, and Calhoun counties, with P. spic-
ulifer in Gulf and Jackson counties, with P. apalachicolae and P. latipleurum
in Gulf County, with P. leonensis in Jefferson, Liberty, Wakulla, and Madison
counties, with P. pygmaeus in Liberty County, and with P. paeninsulanus in
Madison and Taylor counties.
Although P. kilbyi probably should be classified as a secondary burrow-
ing species it is not unusual to find it in open water, in temporary puddles, or
even in small temporary streams. The type specimens were taken from a small,
temporary creek by means of a dip net which was pushed through the vege-
tation.
In Leon County about 12 miles south of Tallahassee the chimneys of P.
rogersi campestris were common around a typical sour-gum and ti-ti bay
which is very low and subject to flooding in wet weather. These burrows had
chimneys which ranged from two to six inches in height. Near by at the edge of
a small bay in sandy soil and in a region where the water table was about one
to three feet below the surface, I found burrows which were less complex than
the majority of the burrows I had been digging, and in them were the first
specimens of P. kilbyi that I had seen. Wire-grass, palmetto, gallberry, and
small ti-ti were characteristic of the flora. Pitcher plants and sundews were
also common.
In Gulf County about 13 miles south of Weewahitchka specimens were
taken from burrows which were marked by neatly formed chimneys about
five inches high. In the ditch close by smaller specimens of P. kilbyi, P. lati-
pleurum, and P. pycnogonopodus were taken.
In Taylor County about six miles north of Perry I waded through a road-
side excavation which was about a foot deep in water. Scattered drowned bur-
rows were observed and dug, but no crayfish were found in them. However, by






HOBBS-THE CRAYFISHES OF FLORIDA


wading through the grass which covered the bottom I startled an occasional
crayfish which would dart some ten or twelve feet ahead, where if I was care-
ful to keep from disturbing it a second time it could be bagged with a dip net.
P. kilbyi is probably the fastest swimmer of any of the Florida crayfishes, and
because it blends so perfectly with the environment, is one of the hardest spe-
cies to catch in open water.
In a small temporary stream near Blountstown I scooped up hundreds of
specimens of very young kilbyi, and upon uprooting some of the vegetation
found that the bottom of the stream was riddled with scores of open burrows.
Many other collecting records give similar ecological data. All point to
the fact that P. kilbyi is a coastal flatwoods species and apparently is just as
well suited to open water as to the burrowing habit. All of the stations in which
this species has been taken from open water are subject to becoming dry, and
this makes it necessary that the crayfish occupy burrows for a part of the year.
All of the females with eggs or young that I have seen have been taken from
burrows.
THE HUBBELLI SUBGROUP
A separate hubbelli subgroup is necessitated to receive the very dis-
junct P. hubbelli.

Procambarus hubbelli (Hobbs)
Plate IV, Figs. 46-50; Maps 3, 4

Cambarus hubbelli Hobbs 1938b (nomen nudum), Proc. Fla. Acad. Sci. 2:
90.
Cambarus hubbelli Hobbs 1940a: 406-410, fig. 19; also 387, 389, 414, 418.
Procambarus hubbelli Hobbs (in press-c).
Diagnosis.-Rostrum without lateral spines; areola relatively broad
with three punctations in narrowest part; male with hooks on ischiopodites of
third pereiopods only; palm of chela of first form male usually bearded
along inner margin; postorbital ridges terminating cephalad without spines;
no lateral spines present on carapace. First pleopod of first form male reach-
ing coxopodite of third pereiopod and terminating in four distinct parts; mesi-
al process spiniform and extends in a distal caudomesial direction; cephalic
process consists of a small, corneous, triangular structure which somewhat
shields the central projection cephalad; caudal process large, corneous, fan-
like, and situated along the distal caudolateral surface; central projection
small, corneous, laterally flattened, and lying somewhat beneath the cephalic
process; cephalodistal setae-bearing prominence knob-like. Annulus ventralis
subovate with the greatest length in the transverse axis; sinus originates on the
cephalic margin slightly dextrad of the midventral line, curves rather abrupt-
ly sinistrad to cross the midventral line where it curves gently caudodextrad
and terminates just before reaching the midcaudal margin.






68 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Remarks.-None of the mostly small variations shown by my specimens
indicates any correlation with any definite locality. The mesial processes of
the first pleopods of several specimens from the southeastern part of Walton
County are directed more mesiad than in specimens from other localities, but
even there specimens are found in which the first pleopod is almost typical.
Procambarus hubbelli is probably the most disjunct species of the barba-
tus group. The first pleopod of the male gives no clue to its relationships except
that it indicates rather distant affinities with the other species of the barbatus
group. It seems most closely related to P. escambiensis, and this relationship
is indicated by both morphological and geographical evidence. Both of these
species have barbate chelae and show several other morphological similari-
ties, and the proximity of the range of P. escambiensis is consistent with the
morphological evidence.
Specimens Examined.-I have examined more than 300 specimens of
Procambarus hubbelli, collected from Holmes, Jackson, Walton, and Wash-
ington counties, Florida.
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
C 1 1 1 16 12
II 1 2 82 3
9 7 4 102 17 1
9 (eggs) 1
9 (young)
c (immature) 18
9 (immature) 44
Geographical and Ecological Distribution.-Procambarus hubbelli is
known from the western portion of Jackson County, from scattered localities
in the southern part of Holmes County, from the southeastern part of Walton
County, and from a few scattered localities in the northern and southwestern
parts of Washington County. It is probable that this species is confined to the
Choctawhatchee River drainage where it inhabits the flatwoods adjoining this
river and Holmes Creek. Undoubtedly, this species is locked in, so to speak,
by the sand ridges that run north and south in Walton County and northeast to
southwest in the southeastern part of Washington County. It thus appears that
the only highway for further active migration is along the coast in Walton and
Bay counties.
Procambarus hubbelli was collected with P. pycnogonopodus in Walton,
Washington, and Holmes counties, and with P. paeninsulanus in Holmes and
Washington counties.
Like other members of the barbatus group, P. hubbelli is a secondary
burrowing, flatwoods species. Although most of my specimens were taken
from burrows, some were found to occur in sluggish creeks and temporary
ponds in the flatwoods. They are common in roadside ditches and have been






HOBBS--THE CRAYFISHES OF FLORIDA


taken from burrows in low areas along creeks where wire-grasses, bog-inhabit-
ing orchids, and pitcher plants are characteristic elements of the flora. The
burrows are generally simple, and have only one or two main passages, which
are almost invariably vertical. They have been found in almost pure clay, in
coarse sand, and often in black mud. The chimneys are usually crudely con-
structed and have the appearance of a small pile of mud. An occasional chim-
ney, however, seems to have been carefully built. I have seen the bottom of
a small temporary pond in Holmes County literally riddled with burrows over
an area of more than 100 square feet; here many of the crayfish were crouched
in the mouths of the burrows, and upon being disturbed darted backward into
their holes. Occasionally I have taken specimens from a submerged mud and
leaf drift in the quiet reaches of a stream.
It is common to find a first form male and a female occupying the same
burrow. In my collection there is only one female with eggs, and she was the
only occupant of the burrow from which she was taken.
I have no reason to believe that this species has habits widely different
from those of the other members of the barbatus group, so what has already
been said concerning the habits of P. pubischelae, escambiensis, kilbyi, and
rathbunae is in general applicable to P. hubbelli.
THE ALLENI GROUP
A separate alleni group is necessitated to receive the very disjunct Pro-
cambarus alleni.

Procambarus alleni (Faxon)
Plate IV, Figs. 51-55; Map 5
Cambarus alleni Faxon 1884, Proc. Amer. Acad. Arts and Sci. 20: 110-112,
also 138.
Cambarus alleni Faxon 1885a: 18, 19, 35-36, 158, 173, PI. I, fig. 1, P1. VIII,
figs. 2, 2'; Faxon 1890: 619-620, 621; Lonnberg 1894a: 125; LUnn-
berg 1894b: 3, 10; Faxon 1898: 646; Ortmann 1902: 277; Harris
1903: 58, 70, 97, 137, 143, 152, 166; Ortmann 1905a: 100, 102, 105,
127, 129; Ortmann 1905b: 403; Ortmann 1906: 18, 19; Faxon 1914:
370, 371, 415; Creaser 1931b: 4; Creaser 1934: 4.
Procambarus alleni Hobbs (in press-c).
Diagnosis.-Rostrum with or without lateral spines; areola relatively
narrow (7-14 times as long as broad); hooks on ischiopodites of third and
fourth pereiopods, those on fourth bituberculate; palm of chela not bearded
along inner margin but bears from 9-11 tubercles; postorbital ridges terminat-
ing cephalad with or without spines; one lateral spine usually present on sides
of carapace. First pleopod of male, form I, reaching coxopodite of third perei-
opod and terminating in four parts; mesial process corneous, subspiculiform,
S-shaped, and extends distad beyond the rest of the appendage; cephalic pro-
cess arising from mesial side of appendage, is corneous and blade-like and its






70 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

distal tip lies proximad of the other terminal apices; central projection a
small, corneous blade, borne along mesial side of caudal process; caudal pro-
cess large and finger-like, extends distad from caudolateral margin; cephalo-
distal setae-bearing surface forming an obtuse angle. Annulus ventralis subo-
vate with a lateral wing-like projection on either side; sinus originates on mid-
cephalic margin, runs caudad for two-thirds the length of the annulus and then
bends sinistrad, making a hairpin turn to the midventral line where it curves
caudad to cut the midcaudal margin of the annulus.
Remarks.-Procambarus alleni has been recorded as one of the several
species in which the rostrum, postorbital ridges, and the cervical groove are
without lateral spines. Most of the specimens in the museums, no doubt, ex-
hibit these features, but in my collection which includes specimens from local-
ities over a large portion of the range, all variations, from a complete lack of
spiny parts to a condition equally as spiny as found in P. fallax, are repre-
sented. Because of this some of the females of the two species are almost in-
distinguishable.
P. alleni exhibits numerous minute variations, and although most of
them are not radical changes, practically every structure generally used for
a taxonomic character shows some degree of variation. Some of these are cor-
related with the habitat of the crayfish while others have no discernible cor-
relations either with habitat or with locality.
Some local, probably inbred, populations are recognizable on a rather
circumscribed combination of several of these variations, and although col-
lections from other regions may exhibit the same list of variations, here they
will rarely be shown in the phenotype of any one individual.
In general specimens of P. alleni from a stream or other permanent body
of water are more spiny than are those collected from burrows or from pools
that are definitely temporary.
Previous authors in discussing the relationships of P. alleni have as-
signed it to a wide variety of different groups, but at least one opinion has been
shared by all of them-that it is an aberrant species. Although I do not think
alleni is so aberrant as some of these authors have contended, it is geographi-
cally far removed from P. simulans which I believe to be its nearest relative.
Ortmann (1905a: 100, 102) erected a separate group for P. evermanni, P.
barbatus, P. wiegmanni, and P. alleni, selecting alleni as the type of the group.
I agree that alleni and barbatus are allied; however, P. evermanni and P.
wiegmanni are not correctly placed with the other two species. In view of the
following similarities between alleni and simulans I do not think they should
be referred to separate sections, but on the basis of the first pleopod I have
placed them in separate groups. In both species the areola is relatively narrow,
and the rostrum in many of the specimens of alleni is devoid of spines, as is the
case in simulans. The general shape of the carapace is similar, and while the
chelae are different, there are certain resemblances in them. The annulus ven-
tralis of the female of alleni is definitely of the simulans type, and finally, the
first pleopod of the male of alleni is closer to that of simulans than to that of






HOBBS-THE CRAYFISHES OF FLORIDA


any other member of the barbatus section. As would be expected the first pleo-
pods of the second form males of the two species agree in detail to a much
greater extent than do those of the first form males. Faxon has emphasized
that in simulans only the third pair of pereiopods bears hooks, while in alleni
hooks occur on both the third and fourth pereiopods. I do not think that this
distinction need be given a great deal of weight, for it has been noted that the
number of walking legs with hooks is variable-even within certain single
species. It is noteworthy that simulans, gracilis, and alleni occupy respective-
ly the western, the north-central, and the southeastern extremities of the range
of the genus.9
The alleni stock probably migrated into Florida from the northwest
shortly after the closing of the Suwannee Straits, to be followed at a later date
by an influx of other species, fallax, paeninsulanus, pubischelae, etc.
The apparent geographical-ecological relationship of P. alleni to P. fal-
lax is well marked outside of the range of alleni. The more ubiquitous fallax
is found in habitats very similar to those occupied by alleni within the latter's
own range. Where their ranges overlap both species have occasionally been
taken from the same station, but generally fallax appears to have taken sole
possession of the streams and the more permanent bodies of water. Since fallax
occupies the area between the northwestern and northeastern extremities of the
range of alleni, it seems at least possible that some competition is going on
between these two species, and that fallax is the more successful competitor.
In the Withlacoochee River, which cuts across the range of alleni, fallax seems
to be the only crayfish inhabitant, and this is also true for the Hillsborough and
Aliphia rivers and several streams in Hardee and Indian River counties, well
within the range of alleni.
Specimens Examined.-I have examined approximately 620 specimens,
from the following counties in Florida: Brevard, Broward, Charlotte, Citrus,
Collier, Dade, Flagler, Glades, Hendry, Hillsborough, Indian River, Lake,
Lee, Levy, Manatee, Martin, Monroe, Okeechobee, Orange, Osceola, Palm
Beach, Pasco, Pinellas, Polk, Putnam, Sarasota, Seminole, St. Johns, St. Lu-
cie, Sumter, and Volusia.
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
CI 8 9 5 38 3 1 7 2 19 18 29 10
CII 6 5 3 19 4 2 4 29 8 17 2
9 17 8 10 51 8 1 14 11 47 53 39 24
9 (eggs) 4 4 1 1
9 (young) 1
J (immature) 4 9 7 9 16 5
9 (immature) 2 1 5 1 2 5 3 12

'The only other species of the simulans group, hagenianus, formerly reported from South Caro-
lina (type locality cited as near Charleston) has not been taken from South Carolina since that time.
Mr. G. Robert Lunz of the Charleston Museum has collected rather diligently for this species and
has never succeeded in obtaining a specimen from South Carolina, and he doubts that the types came
from that state. It has been definitely recorded from Alabama and Mississippi.






72 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

Geographical and Ecological Distribution.-The extent of the range of
P. alleni probably exceeds that of any other species of crayfish in Florida. The
species is abundant in certain localities east of the St. Johns River and has
been collected at intervals along the east coast to Big Pine Key. The sinuous
northern limit of the range of this species may be approximated by a line
drawn along the St. Johns River from its mouth to northern Seminole County,
thence westward to Bushnell in Sumter County, and northwestward through
Lecanto and Citronelle in Citrus County, and northward through Rosewood in
Levy County, and again northwestward to the mouth of the Suwannee River.
The range probably extends eastward to the dunes and brackish waters along
the Atlantic while the southern and western boundaries are marked by similar
situations along the gulf.
Throughout the northern extent of its range P. alleni is associated with
other species, and in at least one instance, it occurs in the same habitat with
paeninsulanus at the boundaries of their ranges. A series of sand ridges cuts
across State Highway 13 between the sixth and eighth mile posts north of
Cedar Keys. In a collection made between the two most western ridges (about
six miles north of Cedar Keys) all of the specimens were P. alleni; likewise
a collection between the next ridges to the east, 6.3 miles northeast of Cedar
Keys, gave only P. alleni. Between a still more eastward pair of dunes, 6.7
miles northeast of Cedar Keys, both P. alleni and P. paeninsulanus were tak-
en. This is the most eastern record for P. alleni in Levy County. Within one-
half mile to the east of this locality the sand dunes give way to low flatwoods
that extend eastward to Bronson. This flatwoods region has been frequently
collected, and although paeninsulanus was abundant, no specimens of alleni
were found.
Faxon (1914) records P. alleni from Lake Butler, near Tarpon Springs,
in Hillsborough County, and from ponds near Tampa. This species is abun-
dant in the roadside ditches throughout the northern part of its range, and the
cypress ponds in St. Johns County are thickly populated by it.
Cypress ponds seem to be a favorable habitat for this species. Often in
wading through the shallow margins of these ponds which usually have a bot-
tom of black silty mud, one observes the small mounds of lighter soil which
mark the openings of the crayfish burrows. If one approaches one of these
mounds cautiously the crayfish may be seen at the mouth of the burrow with
its chelae extended. Some individuals dig their burrows beneath felled logs,
and by moving submerged logs one often uncovers several burrows. (Occa-
sionally I have seen Amphiuma means and Siren intermedia in small tunnels
under such logs, and it is possible that these tunnels were constructed by cray-
fish.) I have seen alleni hiding in the submerged vegetation of roadside ditch-
es; usually, however, if disturbed, at least some of the crayfish rapidly re-
treated into open burrows in the bottom of the ditch.
Although this species has been taken in flowing water, the streams
seemed to be either temporary or very sluggish. The preferred habitat of alleni
appears to be somewhat temporary, still bodies of water in a region where the
soil will support a burrow.






HOBBS-THE CRAYFISHES OF FLORIDA


The burrows of alleni are simple, vertical, or slanting, and range in depth
from one to three feet. The chimneys appear as mounds of small pellets of
mud or clay, and only occasionally does one have a definitely cylindrical
structure.
About seven miles south of Dunnellon, Citrus County, I found burrows
in pure sand, and when I attempted to dig the animals out many were lost be-
cause the sand sloughed off the walls of the burrows faster than I could dig
it out. The water table in this case was only a few inches below the surface,
and just how these burrows remained open I cannot understand. Alleni is the
common flatwood species throughout the extent of its range and is the only
species I have seen from the everglades.
THE ADVENA SECTION
Diagnosis.-The cephalodistal surface of the first pleopod of the first
form male never terminates in a ridge or a knob-like prominence but in a cor-
neous, reduced cephalic process, or if the cephalic process is absent, then the
cephalodistal surface is almost flush with the centrocephalic process of the
central projection. The mesial process is slender, spiniform or blade-like, and
generally directed distad; the central projection is decidedly the most conspic-
uous terminal element, and is either laterally compressed or directed across
the cephalodistal tip of the appendage. The caudal process is present as a large
bump or thumb-like process. The rostrum is broad and short and without later-
al spines; the areola is narrow or obliterated; the male has hooks on the ischi-
opodites of the third, or the third and fourth pereiopods; the chelae are com-
pressed and bear a cristiform row of tubercles along the inner margin of the
palm.
The advena section includes six species and subspecies (advena, geody-
tes, pygmaeus, rogersi rogersi, rogersi ochlocknensis, and rogersi campestris).
P. advena is with little doubt the most primitive member of the section.
The ancestral stock probably migrated from the northwest. The easternmost
branch of this stock then gave rise to the present day advena while the western
branch advanced along the Ochlocknee River to the Apalachicola flatwoods
where it gave rise to the rogersi group. It was along this latter path that pygmae-
us (a member of the advena group) later moved into the same region. A por-
tion of the advena group probably diverged to migrate southward into Flori-
da along the eastern side of Trail Ridge. Part of this stock continued along the
St. Johns River, and another turned southwestward into the region of Alachua
County. Certainly P. advena has its closest affinities with P. geodytes and P.
pygmaeus.
The advena section has been divided into two groups: the advena group
and the rogersi group.
THE ADVENA GROUP
Diagnosis.-The mesial process of the first pleopod of the male is well
developed, spiniform, or slightly compressed; the cephalic process is either







74 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2


*"ol le/ei-
P eodudej
p zdvenae


P./ogerJ/ roger-J
togePri camp ELY/f is
,ogercs/ ocA/ockmenmi/g

.69e/-/ rn/er ra de.r
*~p 4/a'~d/rn/i' zrcu/uJ


Map 5.-Distribution of the Advena Section, Procambarus alleni, and Procambarus
blandingii acutus.






HOBBS-THE CRAYFISHES OF FLORIDA


lacking or is represented by a vestige on the cephalodistal surface; the caudal
process is present only as a small swelling-the distinct rim or a corneous
process is lacking; the central projection is relatively large and blade-like,
compressed laterally, and is directed caudodistad. Hooks are present on the
ischiopodites of the third, or the third and fourth pereiopods of the male.
Three species belong to this group, namely, P. advena, P. geodytes, and
P. pygmaeus. The range of the group in eastern Florida extends from Nassau
County, southward along the St. Johns River and its tributaries to the south-
western part of Seminole County; westward it is found abundantly in the flat-
woods in Clay County and is known from one isolated locality in Alachua
County. In Georgia the range swings westward into Echols, Lowndes, Cook,
Colquitt, and Thomas counties, and thence southward into Liberty and Gulf
counties in western Florida. Northward in Georgia it extends into Ben Hill
County, and eastward into Appling, Wayne, and Bryan counties.
All of the species of the advena group are closely related, but each seems
to be either geographically or ecologically isolated from the others. P. geody-
tes is apparently geographically isolated from the other two while P. advena
and P. pygmaeus appear to be ecologically separated.
The barrier, if any, which exists between advena and geodytes lies in the
area along the St. Johns River between Green Cove Springs and the Welaka
regions. P. advena was collected four miles south of Green Cove Springs, Clay
County-[U. S. Highway 17]-and P. geodytes was taken 14 miles southeast
of Palatka, Putnam County-[State Highway 308]-and at Orange Springs,
Marion County.
P. advena is a primary burrowing species while P. pygmaeus lives in
open water and is probably a secondary burrower; hence, even though the
ranges of the two species overlap they occupy different habitats. This more
or less complete ecological isolation is moreover probably reinforced by the
morphological isolation due to the very small size of pygmaeus.

Procambarus advena (LeConte)
Plate IV, Figs. 56-60; Map 5
Astacus advena LeConte 1856, Proc. Acad. Nat. Sci. Philad. 7: 402.
Cambarus carolinus Hagen 1870: 87-88, P1. I, figs. 51-54, P1. III, fig. 5.
Cambarus advena Faxon 1884: 113, 140, 141; Faxon 1885a: 8, 12, 47-48,
49, 54-56, 58, 158, 173; Faxon 1885b: 358; Ortmann 1902: 277, 279;
Harris 1903: 58, 68, 129, 150; Ortmann 1905a: 98, 100, 101, 104;
Ortmann 1905c: 438; Faxon 1914: 412; Hobbs 1938a: 65; Hobbs
1940a: 389, 393; Hobbs (in press-a); Hobbs (in press-d).
Procambarus advena Hobbs (in press-c).
Diagnosis.-Rostrum subovate, acute, without lateral spines; areola nar-
row; male with hooks on ischiopodites of third pereiopods (hooks in some
specimens bituberculate); chela of first form male never bearded but bears a






76 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

cristiform row of large tubercles along inner margin of palm; postorbital
ridges terminating cephalad without lateral spines or tubercles; no lateral
spines present on carapace. First pleopod of male reaching coxopodite of
second pereiopod and terminating in four parts; mesial process slender and
acute, directed caudodistad; cephalic process vestigial; caudal process not dis-
tinguishable from bulbiform portion of outer part; central projection by far
the most conspicuous, is corneous, blade-like, compressed laterally, and di-
rected caudodistad. Annulus ventralis subovate with the greatest length in
the longitudinal axis; cephalolateral margins raised; fossa disappearing be-
neath caudosinistral wall; sinus, not evident cephalad of fossa, arises from
sinistral side of fossa, curves gently to midventral line, and cuts the midcaudal
margin of the annulus.
Remarks.-For several years I have attempted to associate the type spec-
imen of this species at the Museum of Comparative Zoology with some of my
own specimens, hoping that the type locality might be more specifically
known. I have examined the type, made measurements, drawings, and photo-
graphs of it, and through the kindness of Dr. Fenner A. Chace of the Museum
of Comparative Zoology and Mrs. A. F. Carr, Jr. of the University of Florida,
most of my specimens have been compared with it, but still, I am uncertain
as to where LeConte's specimen was collected.
While there are a number of local variants in the Florida specimens as-
signed to advena the variation is so slight that the locality from which any
given specimen has been taken can be determined only if series of the whole
region are available for comparison; however, when these variants are com-
pared with Georgia specimens the differences are more pronounced.
Probably the most decided variation is to be noted in the shape of the ros-
trum. This is comparatively broader in specimens collected near Baxley, Ap-
pling County, Georgia, than in specimens from elsewhere over the range. The
nearest approach to the Baxley type of rostrum is shown by two specimens
collected near Waycross, Ware County, Georgia. Their rostra are not exactly
alike, but neither is so broad, and each is more acute than those of the Baxley
forms. Specimens from Clinch, Cook, and Echols counties, Georgia, are acute
and not markedly different from the Waycross forms. In Nassau and Duval
counties, Florida, the rostral ridges of the specimens are gently convergent up
to near the tip where they bend more or less sharply together to form an acute
apex (the bend may be either angular or slightly rounded). In specimens from
Clay County, Florida, the subrostral ridges extend laterad of the rostral
ridges almost to the apex, and the rostrum is almost triangular. A few speci-
mens from Putnam County, Florida, show a radical difference in the general
trend of the rostral ridges in that they exhibit a mesial invagination just distad
of the base, and the rostrum is definitely not triangular. The Alachua County,
Florida, forms have rostra which are very much like the Clay County speci-
mens except that the subrostral ridges do not extend so far laterad as do those
of the latter.






HOBBS-THE CRAYFISHES OF FLORIDA


The areola is fairly constant over the entire range except in the speci-
mens from Putnam County. In this population it is practically obliterated.
First form males are known from only five localities, and in two of these
from only a single specimen. The main differences in the few I have seen seem
to be in the development of the cephalic process and the direction in which
the central projection is extended. These are possibly individual variations.
A rather careful analysis was made of the annulus ventralis of the ad-
vena females in my collection, and I find at least four types which are clear-
ly distinct: one type from Appling County, Georgia; another from Alachua
County, Florida; another from Putnam County, Florida; and finally, the
largest group, including all of the others. Naturally the latter group shows a
wide range of minor variations. A single female from Echols County, Geor-
gia, has a very distinct annulus ventralis, but since I have only the one speci-
men from that locality it is possible that this is again an individual variation.
Although these and a few other characters are definitely variable, the
great majority of characters are remarkably uniform throughout the species.
This is more noteworthy in that this species does not have a high frequency
over its range but tends rather to occur in small local concentrations of pre-
sumably inbred populations and appears to have comparatively poor powers
of dispersal.
Specimens Examined.-I have examined a total of 81 specimens of Pro-
cambarus advena, only 27 of which were collected in Florida. This species
has been found in the following counties in Georgia and Florida: GEORGIA-
Appling, Ben Hill, Bryan, Clinch, Colquitt, Cook, Echols, Lowndes, Pierce,
Thomas, Ware, and Wayne. FLORIDA-Alachua, Clay, Duval, Nassau, and
Putnam.
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
lI 1 3 3
ll 2 1 6 3 5 7 3
9 2 10 1 4 1 4 11 2 6
9 (eggs) 2 1
9 (young)
6 (immature) 2
9 (immature) 1
Geographical and Ecological Distribution.-The southern limit of the
range of P. advena appears to be in Alachua and Putnam counties where spec-
imens have been taken from two dissimilar ecological situations. The eastern
limit is marked by the St. Johns River, and the western, except at the extreme
south end of the range, by Trail Ridge. The Alachua County records are all
from a single locality, a seepage area in a garden and woods along Hogtown
Creek about two miles north of Gainesville [U. S. Highway 441].
I have collected over a greater part of Alachua County but have not en-
countered this species elsewhere. Neither do I have specimens from Bradford






78 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

County, but I strongly suspect that numerous isolated colonies are widespread
in the eastern part of Alachua County and in Bradford County.
In Georgia the northern limits of the range reach into Ben Hill, Appling,
and Bryan counties; here it extends almost a hundred miles farther west than
it does in Florida, for I have taken this species 9.5 miles southeast of Meigs in
Thomas County. It seems probable that P. advena is confined to the coastal
terraces and the marshy and wet hillsides of the Tifton Upland along the
courses of its small streams.
In Nassau County, Florida, P. advena was taken from burrows in a road-
side ditch in which there were also burrows of P. pubischelae and P.-semino-
lae. In Georgia it was also taken with the same two species.
P. advena is a primary burrower, spending most of its life in its burrow.
The burrows are beautifully constructed and, though rambling, are very
elaborate with numerous galleries. They are made either in some plastic soil
or in sand underlain by plastic material, and in the latter case the passage
through the upper sand is plastered with mud brought up from the deeper
part of the burrow. Usually there are several large chambers, some that are
interspersed along the runways and others placed at the terminals of the sev-
eral passages. It is not uncommon to find a chamber with three or four pas-
sages leading away from it. In every burrow there is at least one passage
which spirals downward a few inches to two or three feet to a "cellar" below
the water table. This passage is usually near one of the chimneys of the rami-
fying structure. I have often found nuts, large pebbles, grass, and sticks in the
passages and chambers.
The chimneys of these burrows are always carefully constructed and in
their excellent masonry often approach the chimneys of C. diogenes. I have
seen them extending five or six inches above the ground level and consisting of
neatly formed, round pellets about five-sixteenths of an inch in diameter.
Often a single crayfish will construct three or four of these chimneys from its
single complex of passageways. Some burrows extend from six to eight feet
horizontally with as many as four chimneys, and downward to a depth of three
feet to the "cellar." Occasionally I have dug burrows in which a chamber
about six inches high and six to ten inches in diameter was located above the
cellar passageway.
A single soil class does not seem to determine the distribution of the spe-
cies, for I have found it in clay, sandy clay (so hard that it felt like sandstone
to my hands), sandy muck, loam, and even in seepage areas in which a black
soil was predominant. Available subsurface groundwater seems to be a sine
qua non in determining the distribution; streams and permanent open bodies
of water seem to be as barren of this species as do the "Black-jack" ridges.
Specimens are sometimes frequent along creeks but are confined to seepage
areas well above the normal stream level. The water table in most of the lo-
calities I have collected is probably variable, and at times may cover the
mouth of the burrows. In some localities where this is obviously the case, the
groundwater in dry seasons may descend more than two feet below the surface.






HOBBS--THE CRAYFISHES OF FLORIDA


Although P. advena occurs chiefly in the flatwoods, it cannot be regard-
ed as a typical flatwoods species; it is apparently just as successful in seep-
age areas along small streams in a region of high pine and rolling hammock
lands.
In the single Alachua County locality this species occurs in comparative-
ly large numbers in the lower corner of a garden bordering Hogtown Creek.
At one time this plot was a swampy seepage area. The owner constructed
small drainage ditches, cleared away the undergrowth of vegetation, and this
one-time swamp has been converted into a very rich garden-spot where the
crayfish still occur. There are probably a hundred chimneys in this plot, and
fresh ones may be seen almost every morning, especially after a rain. It seems
that the crayfish are most active at night. They do not harm the plants except
by throwing up chimneys on top of young seedlings. There is another inhabit-
ed seepage area about fifty to one hundred yards upstream, and occasional
chimneys may be seen at other spots along the creek bed for several hundred
yards.
The habitat described above is markedly different from the locality from
which this species was taken in Putnam County-a pine flatwoods, 11 miles
north of Palatka, in which wire grass and deer's-tongue (Trilisa) are com-
mon. The soil is a sandy muck underlain by a very plastic clay so that after
a heavy rain the whole area is water-logged.
This species appears to be most inquisitive; almost any sort of commo-
tion at the mouth of the burrow seems to attract it to the surface of the water
table. Recently I have found that in most instances, particularly in dry weath-
er, when the burrow is disturbed the crayfish either retreats into the deep
chamber or was already there. If the passage to this chamber is located, the
water thoroughly agitated, and then allowed to remain still, the crayfish usu-
ally climbs up the passage to the surface of the water. Here its two antennae
may be seen whipping to and fro near the surface, and the crayfish can readily
be captured. In other instances it has been necessary to dissect the burrow in
order to find its occupant. Then, even though the greatest care is exercised,
some of the numerous side passages are lost, and the chances are about even
that the crayfish will escape. I have found that a frequently successful and
time-saving method of collecting is to open the passage to the deeper chamber
of several burrows and thoroughly agitate the water in the mouth of each. If
each opened burrow is then cautiously approached, the crayfish is likely to be
in the process of repairing or inspecting the damage and may be caught with
little difficulty. It is necessary to approach the burrow carefully, however,
because the slightest sound will cause the crayfish to retreat into the deeper
passages. In the rainy season when the crayfish is frequently in one of the
smaller chambers or tunnels nearer the surface rather than in the deepest pas-
sage, a careful dissection of the entire burrow is often necessary.
In a flatwoods, in Clay County, I found an area, populated by P. advena,
in which the burrows possessed no chimneys but opened directly into a ditch
that had several inches of water in its bottom. The mouths of the burrows were






80 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

above the water, but since the water was then quite low I believe that the
mouths would be below the surface in rainy seasons. This is the only place
where I have encountered such burrows for this species.
Numerous copepods occupy the burrows made by advena, and occasion-
ally amphipods of the genus Crangonyx are found clinging to the crayfish's
swimmerets. Further examination of the crayfish reveals many ostracods (six-
legged forms, genus Entocythere, authority Ward and Whipple 1918) cling-
ing to the pubescent regions of the ventral surface.
It is likely that the burrows of advena form a definite microhabitat,
characterized by a small but special fauna in which the other species associat-
ed with advena have found a stable niche.

Procambarus geodytes, sp. nov.
Plate V, Figs. 61-65; Plate XIX; Map 5

Cambarus advena geodytes Hobbs 1937: 154 (nomen nudum).
Diagnosis.-Rostrum without lateral spines, broad, short, and acute;
areola linear or obliterated; male with hooks on ischiopodites of third and
fourth pereiopods; palm of chela of first form male not bearded but bears
a cristiform row of tubercles; postorbital ridges terminating cephalad with-
out spines or tubercles; no lateral spines on sides of carapace. First pleopod
of male, form I, reaching coxopodite of second pereiopod when the abdomen
is flexed and terminating in four parts; mesial process corneous, slender and
blade-like, and directed caudodistad; cephalic process corneous, very small,
and located on the cephalolateral margin near tip of appendage; caudal pro-
cess not present as a corneous ridge or corneous prominence, but made up of
the swollen caudodistal terminal of the appendage; central projection, the most
conspicuous of the four terminals, corneous, compressed laterally, acute, and
directed caudodistad. Annulus ventralis subcylindrical with cephalic margin
deeply cleft and high tuberculate lateral ridges; sinus originates on midven-
tral line about one-fourth the length of the annulus from the cephalic margin,
turns gently dextrad, somewhat more sharply sinistrad to cross the midventral
line, and finally caudodextrad to the midventral line terminating just before
cutting the caudal margin of the annulus.
Holotypic Male, Form I-Body subcylindrical, slightly depressed. Abdomen much
narrower than thorax (.96-1.40 cm. in widest parts respectively). Width of carapace
greater than depth in region of caudodorsal margin of cervical groove. Greatest width
of carapace about midway between cervical groove and caudal margin of carapace.
Areola linear, almost obliterated. Cephalic section of carapace about 1.56 times
as long as areola (length of areola 39% of entire length of carapace).
Rostrum deeply excavate, almost reaching penultimate segment of peduncle of
antennule; margins converging to tip; no lateral spines present. Upper surface of ros-
trum with a row of setiferous punctations bordering the high marginal ridges. Subros-
tral ridges well defined and evident in dorsal view almost to tip.
Postorbital ridge well defined, terminating cephalad in small tubercles. Subor-







HOBBS--THE CRAYFISHES OF FLORIDA


bital angle absent; branchiostegal spine heavy but short. No lateral spines on carapace.
Surface of carapace punctate dorsad, tuberculate laterad.
Abdomen shorter than carapace (2.60-2.95 cm.).
Cephalic section of telson with one spine in the sinistral and two in the dextral cau-
dolateral corners.
Epistome subovate with a single cephalomedian acute tubercle, and a caudolater-
al one on either side at base.
Antennules of usual form. No spine present on ventral side of basal segment.
Antennae extending caudad to first abdominal segment; antennal scale small
with mesial and lateral sides subparallel, cephalic margin of blade portion sloping.
Spine on outer margin very strong.
Chela distinctly depressed, of moderate length and width. Hand with squamous,
ciliated tubercles except on upper proximal and lower surfaces. Inner margin of palm
with a cristiform row of seven tubercles. A distinct submedian ridge present on upper
surface of both fingers. Fingers slightly gaping. Opposable margin of dactyl with five
large, corneous tubercles along proximal three-fifths and a row of minute denticles
along distal two-fifths, a few scattered ones between the large tubercles. Lateral margin
of dactyl with three squamous tubercles at base and a row of setiferous punctations
distad of them. Upper surface of dactyl with a submedian ridge flanked by setiferous
punctations. Lower surface with sparsely scattered setiferous punctations. Opposable
margin of immovable finger with five prominent tubercles on proximal three-fifths
and with a large strongly cornified one extending from lower margin at base of distal
fourth. A row of minute denticles broken only by the large tubercles extends along en-
tire margin. Lateral margin of immovable finger with a row of about seven setiferous
punctations. Upper surface with a distinct submedian ridge flanked by setiferous punc-
tations. Lower surface with setiferous punctations; setae along lateral portion extreme-
ly long.
Carpus of first pereiopod one-half as broad as long; a well defined longitudinal
groove above, punctate except on mesial surface which is tuberculate-about six prom-
inent ones.
Merus punctate on mesial and lateral surfaces. Upper surface with an irregular
row of small tubercles, two somewhat larger ones near distal end. Lower surface with
an outer row of seven tubercles and an inner row of ten, inner row more spike-like.
Hooks on ischiopodites of third and fourth pereiopods; hooks on third bitubercu-
late. Bases of coxopodites of fourth pereiopods with very strong compressed out-
growths directed caudoventrad; those on fifth much smaller but acute and directed
cephaloventrad.
First pleopod extending to base of second pereiopod when abdomen is flexed. Tip
terminating in four distinct parts. Mesial process corneous and blade-like, and directed
caudodistad. Cephalic process small, corneous, acute, and situated on the cephalo-
lateral margin near tip. Caudal process not present as a corneous ridge or prominence
but made up of the swollen caudodistal portion of the main shaft. Central projection,
the most prominent terminal element, corneous, acute, and directed caudodistad. The
whole distal portion of the appendage is bent somewhat caudodistad.
Male, Form II.-Differs from the male, form I, only in the reduction of the second-
ary sexual parts, and the first pleopod has no indication of a cephalic process; inner
margin of palm of chela has eight tubercles; spines on telson are mirrored images of
those in holotype; there are also other differences in tubercle counts.
Allotypic Female.-Differs from the first form male chiefly in tubercle count on
the chela and other spiny portions.
Annulus ventralis subcylindrical with a single small tubercle on either side at
middle; cephalic margin deeply cleft and with high tuberculate lateral ridges; sinus






82 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

originates on midventral line about one-fourth the length of the annulus from the ce-
phalic margin, turns gently dextrad, somewhat more sharply sinistrad to cross the
midventral line and finally caudodextrad to the midventral line just before cutting the
caudal margin of the annulus.
Measurements.-Male (form I) Holotype: carapace, height 1.17, width 1.40,
length 2.95 cm.; areola, width .01, length 1.15 cm.; rostrum, width .42, length .43 cm.;
abdomen, length 2.60 cm.; right chela, length of inner margin of palm .55, width of
palm .88, length of outer margin of hand 1.85, length of movable finger 1.33 cm. Fe-
male Allotype: carapace, height 1.52, width 1.56, length 3.20 cm.; areola, width .01,
length 1.23 cm.; rostrum, width .50, length .50 cm.; abdomen, length 2.95 cm.; right
chela, length of inner margin of palm .55, width of palm .96, length of outer margin of
hand 1.86, length of movable finger 1.31 cm.
Type Locality.-Orange Springs, in the northeastern part of Marion County, Flor-
ida. All of the specimens taken here were dug from complex burrows around the mar-
gin of the swimming pool.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form I) and one female are deposited in the Museum of Compara-
tive Zoology; one male (form I) and one female in the University of Michigan Museum
of Zoology. Four males (form I) and nine females are in my personal collection at the
University of Florida.
Remarks.-Here again I have too few specimens to determine which, if
any, variations are peculiar to local populations. Of the six localities repre-
sented in my collection, the specimens from near Welaka seem to represent the
most highly inbred population. The rostra of these specimens are broader and
resemble those of the Baxley specimens of P. advena. The shape of the chelae is
also distinct.
The pleopod of the male from Palm Springs seems to be rather distinct;
however, since I have only the one specimen it is possible that this is only an
individual difference. (The cephalic process is present as a small acute spine
at the cephalodistal margin of the gonopod; in the other specimens the process
is entirely lacking, or may be represented by a rudimentary bump along the
distal cephalolateral margin.)
There is little doubt that P. geodytes has its closest affinities with P.
advena. For some time I considered it as a subspecies of advena, but upon fur-
ther study, I am convinced that my intergradation data is too indefinite to call
geodytes a race of advena. (The form which I have ascribed to P. advena col-
lected in Putnam County appears to be somewhat intermediate between the
two species; however, since I have so few specimens from this region and no
first form male, and the habitat from which they were taken is more similar
to that occupied by P. advena, those specimens in question will be considered
a variant of the latter unless further collecting and study will prove them to be
otherwise.)
Specimens Examined.-I have examined a total of 42 specimens of Pro-
cambarus geodytes, from Marion, Putnam, and Seminole counties. In Marion
County it was taken from burrows in the swampy area around Salt Springs in
the Ocala National Forest and at Orange Springs; in Putnam County it was






HOBBS-THE CRAYFISHES OF FLORIDA


found around the edge of a small sulfur spring flowing into the St. Johns
River about three miles north of Welaka, and along the shore of Lake Crescent,
3.5 miles north of Crescent City; in Seminole County it was dug from burrows
around Palm Springs and at Shepherd Springs just north of Sanlando.
My 42 specimens were collected in April, October, November, and De-
cember. First form males were taken during October and November. A single
mature male, form II, was taken in December, and one female with young and
two immature females in April.
Geographical and Ecological Distribution.-This species is endemic to
Florida and appears to be confined to the St. Johns River drainage. Although
it is known to occur in only Putnam, Seminole, and Marion counties, I believe
that subsequent collecting will reveal its presence in Flagler, Lake, Orange,
and Volusia counties.
Procambarus geodytes is not associated with any other crayfish, al-
though both fallax and acherontis have been taken from springs and spring-
runs around which geodytes burrows.
Like P. advena, geodytes is a primary burrower, and I have no evidence
that its habits are different from those of advena. Although ecologically simi-
lar, geodytes is almost confined to regions around sulfur or mineral springs
where the water table is available only a few inches to a foot below the surface.
The burrows of geodytes are equally as complex and elaborate as those of ad-
vena. Along the bank of a small sulfur spring in Putnam County numerous
chimneys were crowded together in a small seepage area. I dug into the midst
of a group of these and struck a buried log. When this was moved three cray-
fish were uncovered. Several times I have noticed that members of this species
construct runways beneath the entire length of a felled log. One of the largest
burrows of this nature I have observed was at Palm Springs; here a board was
lying on the ground, and under it was a crayfish tunnel about twelve feet long.
At intervals there were vertical passages leading downward from the tunnel.
The chimneys constructed by geodytes are much like those of advena, and they
are usually found in regions where the soil is a sandy muck containing an
abundance of decaying organic matter.
From burrows at both Orange Springs and Welaka specimens of this
species were collected on which amphipods of the genus Crangonyx were
holding to the abdomens of the crayfish.

Procambarus pygmaeus, sp. nov.
Plate V, Figs. 66-70; Plate XX; Map 5
Diagnosis.-Rostrum without lateral spines, sublanceolate; areola nar-
row; male with hooks on ischiopodites of third pereiopods only; palm of che-
la of first form male not bearded but bears a cristiform row of tubercles; post-
orbital ridges terminating cephalad without spines or tubercles; no lateral
spines on sides of carapace. First pleopod of first form male reaching coxop-
odite of second pereiopod when abdomen is flexed, and terminating in three







84 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

distinct parts; mesial process slender and blade-like, extends caudodistad
slightly beyond the central projection; cephalic process absent; caudal pro-
cess forming a sharp corneous ridge along caudolateral margin; central pro-
jection large, thin, plate-like, and directed distad; distal portion of outer part
of appendage hardly inflated, whereas in advena it is greatly swollen. Annu-
lus ventralis subovate, with a submedian furrow; sinus originates along mid-
ventral line about one-third of the total length from the cephalic margin;
curves gently sinistrad, somewhat sharply dextrad to midventral line where
it turns caudad to cut the midcaudal margin; fossa not so plainly seen as in
advena.
Holotypic Male, Form I-Body subovate, compressed laterally. Abdomen narrow-
er than thorax (.68-.81 cm. in widest parts respectively). Width and depth of carapace
subequal in region of caudodorsal margin of cervical groove. Greatest width of cara-
pace slightly caudad of caudodorsal margin of cervical groove.
Areola narrow (32 times longer than wide) with a few scattered punctations, only
one in narrowest parts; sides parallel for some distance in middle. Cephalic section of
carapace about 1.92 times as long as areola (length of areola 34.2% of entire length
of carapace).
Rostrum flattened above reaching middle of distal segment of peduncle of anten-
nule; margins converging to tip; no lateral spines present. Upper surface of rostrum
with scattered punctations. Marginal ridges high. Subrostral ridges well defined and
evident in dorsal view to tip of rostrum.
Postorbital ridge well defined, merging abruptly cephalad into carapace without
forming tubercles or spines. Suborbital angle absent; branchiostegal spine moderate-
ly developed. No lateral spine on carapace, a few small tubercles instead. Surface of
carapace punctate dorsad, granulate laterad.
Abdomen slightly longer than thorax (1.95-1.87 cm.).
Cephalic section of telson with two spines in each caudolateral corner.
Epistome broadly ovate but slightly angular cephalad and cephalolaterad.
Antennules of usual form. No spine present on ventral side of basal segment.
Antennae extending caudad to second abdominal segment; antenna scale small and
narrow with cephalic margin of blade portion sloping. Spine on outer margin very
strong.
Chela distinctly depressed, of moderate length and breadth. Hand tuberculate ex-
cept on ventromesial and ventrolateral surfaces. Lower surface of palm with a group
of tubercles in two irregular rows running obliquely proximolaterad to mesiodistad.
Inner margin of palm with a cristiform row of eight tubercles. A distinct ridge present
on upper surface of both fingers. Fingers gaping. Opposable margin of dactyl with
six tubercles along basal three-fifths; a single row of minute denticles between these,
and continuous to tip. Lateral margins of dactyl with six large tubercles along basal
half, distad of which is a row of setiferous punctations. Upper surface of dactyl with a
submedian ridge flanked proximad by a group of tubercles, distad by setiferous punc-
tations. Lower surface setose with setiferous punctations. Opposable margin of im-
movable finger with four tubercles on proximal half and one tubercle extending from
lower surface at base of distal third. Crowded denticles interspersed between these.
Lateral margin with a row of setiferous punctations. Upper surface with a median ridge
flanked by setiferous punctations. Lower opposable surface with a distinct excavation
and setiferous punctations.
Carpus of first pereiopod longer than wide (.50-.41 cm.); a shallow longitudinal
groove above. Punctate except on mesial surface which is tuberculate. Two tubercles







HOBBS--THE CRAYFISHES OF FLORIDA


larger than others; a row of three or four tubercles on cephalomesial margin; two tu-
bercles on cephaloventral margin.
Merus punctate on mesial and lateral surfaces. Upper surface with an irregular
row of tubercles. Lower surface with an irregular outer row of about 14 tubercles and
an inner row of about 12 spike-like tubercles.
Hooks on ischiopodites of third pereiopods only. Bases of coxopodites of fourth
pereiopod with a large tubercle directed ventrad; that on fifth very small.
First pleopod extending to base of second pereiopod when abdomen is flexed. Tip
terminating in three distinct parts. Mesial process slender and blade-like, directed cau-
dodistad and extends slightly beyond central projection. Cephalic process absent. Cau-
dal process forming a distinct corneous ridge on caudolateral margin at base of central
projection. Central projection large and plate-like, directed distad. Distal portion of
appendage only slightly inflated.
Male, Form II.-Differs from the male, form I, only in the reduction of the second-
ary sexual parts and in the first pleopod. Caudal process of first pleopod absent; both
the others are somewhat truncate, heavier, and non-corneous.
Allotypic Female.-Differs from the first form male in tubercle count on cheliped,
and in proportions of chela; antennae extend caudad to fifth segment of abdomen.
Annulus ventralis subovate, with the greatest length in the longitudinal axis; a
longitudinal furrow present along midventral line with high lateral walls along cephalic
half; sinus originates on midventral line about one-third of total length from cephalic
margin, curves gently sinistrad, then somewhat more sharply caudad to midventral line,
where it turns caudad to cut the midcaudal margin of the annulus.
Measurements.-Male (form I) Holotype: carapace, height .78, width .81, length
1.87 cm.; areola, width .02, length .64 cm.; rostrum, width .28, length .34 cm.; abdo-
men, length 1.95 cm.; right chela, length of inner margin of palm .40, width of palm
.52, length of outer margin of hand 1.12, length of movable finger .60 cm. Female Al-
lotype: carapace, height .72, width .80, length 1.80 cm.; areola, width .01, length .61
cm.; rostrum, width .25, length .32 cm.; abdomen, length 1.93 cm.; right chela, length
of inner margin of palm .40, width of palm .52, length of outer margin of hand 1.02,
length of movable finger .63 cm.
Type Locality.-About 16 miles north of Fargo on Georgia State Highway 89,
Clinch County, Georgia. A small swamp stream flowing through a cypress bay in the
flatwoods region. Pontederia sp., and Juncus repens were abundant in the edge of the
stream and in the adjoining roadside ditch. Most of the specimens were taken by push-
ing a dip net through the dense Juncus beds.
Disposition of Types.-The male holotype, the female allotype, and a second form
male paratype are deposited in the United States National Museum. Of the remaining
paratypes one male (form II) and one female are deposited in the Museum of Compara-
tive Zoology; one male (form II) and one female in the University of Michigan Mu-
seum of Zoology. One male (form I), four males (form II), 15 females, and 35 imma-
ture specimens are in my personal collection at the University of Florida.
Remarks.-With the few specimens available it is difficult to determine
the extent of the local variations in this species. The rostra of the specimens
from Clinch County, Georgia, seem to be more lanceolate than those from
Wayne County, Georgia, or those from Florida. The basal segments of the an-
tennule of the Clinch County and Florida specimens bear no spine on the ven-
tral surface, while a well developed one is present on that of some of the speci-
mens from Wayne County. These two differences are the most pronounced;
however, other and rather minute differences occur, and perhaps with the ac-







86 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

quisition of more specimens throughout the range, definite local variants can
be recognized.
Procambarus pygmaeus has its closest affinities with Procambarus ad-
vena. This is particularly interesting since the ranges of these two species over-
lap considerably. The two are somewhat different, however, in their habits
as is pointed out in the discussion of the geographic and ecological distribu-
tion of the two species.
Specimens Examined.-I have examined a total of 110 specimens of Pro-
cambarus pygmaeus, from the following counties in Georgia and Florida.
GEORGIA--Clinch and Wayne counties. FLORIDA-Gulf and Liberty counties.
The exact localities are: GEORGIA, Clinch County-7.6 miles north of Fargo
[State Highway 89]; 15.7 miles north of Fargo [State Highway 89]; 5 miles
northeast of Homerville [U. S. Highway 84]. Wayne County-.5 mile north
of Jessup [State Highway 38]. FLORIDA, Gulf County-11.7 miles west of
Weewahitchka [State Highway 52]; 6.6 miles east of the Gulf-Bay County
line [State Highway 52]; 6.1 miles south of Weewahitchka [State Highway
6] ; 1.7 miles east of the Gulf-Bay County line [State Highway 52] ; 4.1 miles
west of Weewahitchka [State Highway 52]. Liberty County-5.4 miles south
of Telogia [State Highway 135]; 11.3 miles south of Telogia [State Highway
135].
SEASONAL DATA
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
dI 1 1 2
d II 5 7 2
9 3 1 8 1 17 12 1
9 (eggs) 1
9 (young)
S (immature) 1 5 2 21 1 1
9 (immature) 1 3 12 1
Geographical and Ecological Distribution.-Procambarus pygmaeus is
undoubtedly much more widely distributed than is indicated by the few local-
ities from which it has been taken. It is probably scattered throughout the
southeastern part of Georgia south of the Altamaha River and in the northern
part of Florida in the Okefenokee region. Although this species has not actual-
ly been recorded from the northern part of Florida it is very likely that collec-
tions made in the northern part of Baker and Columbia counties will disclose
its presence there. In the Florida panhandle it is probably common in local
areas in Liberty, Franklin, Gulf, and Bay counties.
The range of P. pygmaeus as indicated on Map 5 is interesting in that
it provides additional support for the conjectured path of migration for at
least two other stocks migrating into the Florida panhandle through the Och-
locknee drainage system. It has been postulated (Hobbs, in press-d) that the
forerunner of the rogersi subspecies had its origin in southeastern Georgia and
migrated along the Ochlocknee River into Florida. Further, it is highly prob-






HOBBS-THE CRAYFISHES OF FLORIDA


able that the precursor of P. young utilized this same path. Since pygmaeus
occurs both in southeastern Georgia and in the western part of Florida, and
the apparent connections between these two portions of the range are in the
Ochlocknee region, it would seem to add some degree of validity to the above
mentioned postulated migration route taken by the rogersi and young stocks.
In Georgia P. seminolae and P. pubischelae were taken with P. pygmaeus
in both Clinch and Wayne counties, and in Wayne County P. advena was
found burrowing near the ditch where P. pygmaeus occurred. In Florida P.
kilbyi, P. pycnogonopodus, P. latipleurum, and P. paeninsulanus were col-
lected with P. pygmaeus in Gulf County, and P. kilbyi and P. leonensis were
taken with it in Liberty County.
This species was first taken by Dr. Frank N. Young and myself while col-
lecting at night with headlights, in a swamp stream about seven miles north
of Fargo, Clinch County, Georgia. Dr. Young was using a coffee sieve for
catching water beetles, and in one scoop dipped up a small crayfish which
proved to be an adult female of P. pygmaeus. It was about an inch long and
marked with brilliant red splotches over a green basal color After this speci-
men was bagged, we spent some time trying without success to secure a first
form male. However, about 16 miles north of Fargo we found the species
more abundant and succeeded in capturing about 50 specimens, two of which
were first form males.
A glance at a specimen of pygmaeus would give the impression that it was
a very small, highly colored specimen of P. advena. Finding this new species
was a surprise, for it had been only a short distance back that I had dug P.
advena from burrows in a roadside ditch, and to find two so nearly related
forms so close together was at least unusual. Even more astounding was the
fact that pygmaeus was not dug from burrows but was taken from flowing wa-
ter. Judging by the method used in collecting them, they must have been out
crawling under the vegetation and over the bottom of the ditch and stream.
Since most of my collecting was done at night I was unable to ascertain whether
these specimens had burrows in the bottom of the ditch and stream or whether
they were true open water forms.
A number of specimens of pygmaeus has been added to my collection
since that time, but all of them have come from similar situations. It has been
noted that every locality from which this species has been taken is in swampy
terrain, and whether in quiet or flowing water Juncus repens is abundant. In
most of these localities this brilliantly red and green colored plant forms thick-
ly matted beds over large areas of the pool or stream bottoms.
For some time I was unable to find this species burrowing, and this
seemed unusual in view of the fact that its relatives are all primary burrowers.
Specimens have now been taken from burrows in nearly all of the localities
cited. P. pygmaeus apparently is a secondary or tertiary burrower. These
burrows are fully as complex as those of the other members of the advena
section, having a number of side passages and sometimes several openings
over which are moderately well constructed chimneys. Most of the burrows I






88 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

have seen were in soft muck very close to the edge of the water or in recently
dried up ditches.
THE ROGERSI GROUP
Diagnosis.-Rostrum broad, short, and devoid of lateral spines; areola
very narrow or obliterated; male with hooks on ischiopodites of third pereio-
pods only; chela compressed and bearing a cristiform row of tubercles along
inner margin of palm; postorbital ridges terminating cephalad without later-
al spines; no lateral spines present on sides of carapace. First pleopod of first
form male terminating in three or four parts; mesial process well developed,
spiniform, or slightly compressed; cephalic process present or absent, if
present consists of a reduced spine on cephalodistal surface; caudal pro-
cess large and thumb-like, bent mesiad at a 15-90 degree angle to the main
shaft; central projection large and plate-like, extending across cephalodistal
surface or obliquely distad in a cephalomesial to caudolateral direction.
A summary of the relationships and distribution of this group given by
Hobbs (in press-d) is quoted in part: "The rogersi group consists of three dis-
tinct though intergrading races, the ranges of which cover a considerable area
in the eastern and central parts of the panhandle of Florida. The extreme east-
ern and western limits of the range of the complex are almost one hundred
miles apart, while the most northern and southern limits are separated by a
distance of about fifty miles. Within these limits are portions of Bay, Gadsden,
Gulf, Calhoun, Franklin, Leon, Liberty, and Wakulla counties.
"All of the members of the complex are primary burrowers with presum-
ably poor powers of dispersal. Their ranges extend through a monotonous
flatwoods, broken only here and there by small, scattered areas unsuitable
for habitation by them. If rogersi had a higher vagility such a range would
seem to favor maintenance of a homogeneous population over the entire area;
but instead the region is inhabited by small, local populations, and when spec-
imens from several of these are compared, it becomes evident that the rogersi
complex, especially in the zone of intergradation, is very heterogeneous. In a
series of specimens from the region of intergradation, transitional forms be-
tween the three subspecies may be beautifully traced.
"The ranges of the three races . [are briefly indicated below]. The
zone of intergradation consists of a large area in Franklin, Gulf, Bay and the
southern parts of Liberty and Wakulla counties. Toward the eastern side of this
area, in eastern Franklin and western Wakulla counties, specimens show a
definite approach to rogersi campestris, which occurs in Leon and Wakulla
counties; west of the Apalachicola River, in northern Gulf County, material
is closer to rogersi rogersi; while in the southern part of Liberty County the
intermediates in most characters more closely resemble rogersi ochlocknensis.
In general, the nearer one approaches the ranges of the three well defined sub-
species, the more nearly do the intermediates resemble typical material of
these subspecies. The actual situation in respect to the intergrades is more
complex than is here indicated ... "






HOBBS--THE CRAYFISHES OF FLORIDA


Procambarus rogersi rogersi (Hobbs)
Plate V, Figs. 71-75; Map 5

Cambarus rogersi Hobbs, Journ. Wash. Acad. Sci. 28 (2): 61-65, figs. 1-11.
Cambarus rogersi Hobbs 1940a: 410; Hobbs (in press-a).
Procambarus rogersi Hobbs (in press-c).
Procambarus rogersi rogersi Hobbs (in press-d).
Diagnosis.-Rostrum without lateral spines; areola very narrow, almost
obliterated; male with hooks on ischiopodites of third pereiopods only; chela
compressed and bearing a cristiform row of tubercles along inner margin of
palm; postorbital ridges terminating cephalad without spines or tubercles;
no lateral spines present on carapace. First pleopod terminating in three or
four parts; mesial process well developed, spiniform; cephalic process usual-
ly absent, if present consisting of a small spine on cephalodistal surface; cau-
dal process non-corneous, thumb-like, and bent caudomesiad at a 90 degree
angle to the main shaft; central projection corneous, plate-like, and directed
across the cephalic tip of the appendage. Annulus ventralis subovate, broader
than long with cephalic margin entire; lateral walls without tubercles.
Remarks.-This subspecies is confined to a relatively narrow strip of
flatwoods in Calhoun County, where each of the local populations constructs
a group of complex burrows.
Specimens Examined.-I have examined a total of 36 specimens of this
subspecies, collected during April, May, June, and November. First form
males were collected in April and May, and one female carrying eggs was
taken in April.

Procambarus rogersi ochlocknensis Hobbs
Plate V, Figs. 76-80; Map 5

Procambarus rogersi ochlocknensis Hobbs (in press-d).
Diagnosis.-Rostrum without lateral spines; areola obliterated; male
with hooks on ischiopodites of third pereiopods only; chela compressed and
bearing a cristiform row of tubercles along inner margin of palm; postorbital
ridges without spines or tubercles; no lateral spines present on carapace. First
pleopod terminating in four parts; mesial process well developed, spiniform;
cephalic process forming a spine on cephalodistal surface; caudal process
heavy, non-corneous, somewhat inflated and directed mesiodistad; central
projection corneous, thin, plate-like, and directed obliquely caudolaterad
from the cephalodistal surface. Annulus ventralis subovate, distinctly longer
than broad, no cephalic wall cephalicc margin deeply cleft); lateral walls
bearing small tubercles.
Remarks.-P. rogersi ochlocknensis is found in seepage areas along trib-
utaries of the Ochlocknee River in Gadsden County, and from similar situa-






90 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

tions and flatwoods in the northern parts of Liberty County. Like the other
two subspecies it seems to be largely restricted to local areas where ground wa-
ter is available within two or three feet below the surface at all times of the
year. All of the burrows of this subspecies are extremely complex, consisting
of intricate tunnels and at least one deep passage which spirals downward.
Specimens Examined.-I have examined a total of 49 specimens of P.
rogersi ochlocknensis, which were collected in March, April, May, August,
and December. First form males were taken in March, April, May, and De-
cember, and two females with eggs were found in March.
Procambarus rogersi campestris Hobbs
Plate VI, Figs. 81-85; Map 5

Procambarus rogersi campestris Hobbs (in press-d).
Diagnosis.-Rostrum without lateral spines; areola obliterated; males
with hooks on ischiopodites of third pereiopods only; chela compressed and
bearing a cristiform row of tubercles along inner margin of palm; postorbital
ridges without spines or tubercles; no lateral spines present on carapace. First
pleopod terminating in four parts; mesial process well developed, spiniform;
cephalic process forming a spine on cephalodistal surface; caudal process
thumb-like and extends mesiodistad at a 45 degree angle to the main shaft;
central projection forming a large corneous fan across cephalic side of tip and
bent laterodistad at about a 45 degree angle to the main shaft. Annulus ven-
tralis subcylindrical with cephalic wall deeply cleft; lateral walls tuberculate.
Remarks.-Typical specimens of this subspecies are found in the south-
western part of Leon County and in the northwestern part of Wakulla County.
All of my specimens have been taken from complex burrows in the low flat-
woods areas in this region.
Specimens Examined.-I have examined a total of 37 specimens of P.
rogersi campestris, which were collected in May, June, August, and Novem-
ber, and first form males were taken in June and November.
The Rogersi Intergrades
Intergrades between the three subspecies of Procambarus rogersi occupy
an area larger than the combined ranges of the three defined races and extend
considerably to the west of the range of P. rogersi rogersi. The area of inter-
gradation extends from the southern part of Liberty County westward through
Franklin, Gulf, and Bay counties west of St. Andrews Bay.
As has been pointed out by Hobbs (in press-d) the main characters which
furnish the intergradation data are the first pleopod of the male and the an-
nulus ventralis of the female.
Specimens Examined.-I have examined a total of 192 rogersi inter-
grades, which were collected during the months of April, May, June, Septem-
ber, October, November, and December. First form males were found in May







HOBBS-THE CRAYFISHES OF FLORIDA


and December, and six females with eggs in May and one female with young
in June.
THE ACHERONTIS SECTION
A separate acherontis section is necessitated to receive the very disjunct
Procambarus acherontis.

Procambarus acherontis (Linnberg)
Plate VI, Figs. 86-90; Map 8

Cambarus acherontis Lonnberg 1894b, Bihand till K. Svenska Vet.-Akad.
Handlingar, Vol. 20, Pt. 4, No. 1: 6-12, Figs. 1-5b.
Cambarus acherontis Lonnberg 1894a: 125-127; Hobbs 1940a: 387-394,
401, fig. 15.
Procambarus acherontis Hobbs (in press-c).
Diagnosis.-Rostrum with small lateral spines; areola narrow (about
20 times as long as broad) ; male with bituberculate hooks on third and fourth
pereiopods; palm of chela not bearded within; postorbital ridges terminating
cephalad with spines; lateral spines present on sides of carapace; eyes re-
duced and without pigment, and the body albinistic. Cephalodistal surface of
first pleopod of male not terminating in a ridge or a knob-like prominence but
almost flush with the central projection; cephalic process absent; mesial pro-
cess lies along the caudomesial margin and terminates one and one-half times
its own length proximad of the tip 7 central projection strongly developed, sub-
triangular, corneous, flattened laterally, and extends beyond the rest of the
appendage distally. Annulus ventralis with cephalic portion hidden beneath
two projections from the sternum just cephalad of it; subovate in general out-
line with a lyre-shaped prominence above; sinus originates on midventral line
near base of the above mentioned prominence, curves gently dextrad until
it approaches the caudal margin of the annulus where it turns sharply sinistrad
and finally caudad where it cuts the caudal margin just sinistrad of the mid-
ventral line. (This structure shows considerable variation, especially in the
bending of the sinus, the general outline, and the presence or absence of tuber-
cles on the cephalolateral surfaces.)
Remarks.-Among the specimens I have at hand there is considerable
variation. All spiny parts show considerable diversity; the rostrum has two or
three extra spines in addition to the usual two, and the telson also bears a vari-
able number of spines. The annulus ventralis is the most non-stable structure I
observed; not only is it different from specimen to specimen in shape, but the
sinus takes a number of different forms. With all of these variations, however,
none are so great as to confuse acherontis with any other known species.
This species has no close affinites with any other crayfish, but I believe
it to be more nearly related to the members of the advena group than to any
other. In some respects it appears intermediate between the spiculifer and






92 UNIVERSITY OF FLORIDA BIOLOGICAL SERIES, VOL. III, No. 2

advena groups. The lateral spines on the rostrum and bituberculate hooks on
the fourth pereiopods are like those of P. versutus, but the other characters are
definitely more like those of advena.
As a result of the fact that the type specimens of P. acherontis were no
longer extant at the time that Faxon examined troglodyte forms from Gum
Cave, Citrus County, Florida, he mistook them to be specimens of P. acheron-
tis. Later authors accepted Faxon's identifications, and this precluded the pos,
sibility of Ortmann's arriving at a correct evaluation of the true relationships
of acherontis as well as leading to a series of misidentifications of Florida's
cavernicolous crayfishes. In at least two instances these misidentifications
have been recorded in literature. L5nnberg's Cambarus acherontis was lost
from the time it was described in 1894 until November 1938 (Hobbs 1940a:
338).
Specimens Examined.-I have examined a total of 44 specimens, from
Seminole County, Florida. All of these specimens were taken from Palm
Springs, about 12 miles north of Orlando on November 11, 1938. [Among
them are 3 d d (form I), 13 d c (form II), 12 9 9,9 $ (immature),
and 7 9 9 (immature).]
Geographical and Ecological Distribution.-As has been pointed out
elsewhere, this species is probably a relict of a once more widely spread stock
which migrated into Florida soon after the first closing of the Suwannee
Straits. Then with submergence of the land bridging the Straits and the pos-
sible inundation of the whole island, a portion of this stock retreated into sub-
terranean freshwater. Whether or not the whole peninsula was submerged af-
ter the reopening of the Suwannee Straits, it does seem highly probable in
light of its aberrant structures and distribution that acherontis was derived
from the earliest surviving stock to arrive in the peninsular region of the state.
No published observations other than those of L6nnberg have been
made on the habits of P. acherontis except those I made at Palm Springs in
1938. "We found more than a score of white crayfish lying in the algae over
the bottom of a pool formed by the spring. This pool is about 60 by 20 feet
and over the most part approximately six feet deep. The walls and bottom
were covered with a thick algal growth, and deposited on it was a sediment
characteristic of sulfur springs. The water had a pH of 7.6. Mr. Marchand
caught most of the forty-four specimens which were secured by diving to the
bottom and capturing them with his hands. They were extremely sluggish,
many lying in the algae on their backs with their feet turned up toward the
surface as though dead. Even after they were bagged, there was appreciably
little sign of life" (Hobbs 1940a: 338).
L6nnberg's account points out the fact that this species does not confine
itself to the mouths of springs or sinks but occurs in at least one underground
stream, evidenced by his specimens found in a well dug near Lake Brantley.
"At first they were fairly numerous, but later on, when I had heard about it
and tried to obtain some specimens, I could only procure two males" (L6nn-
berg 1894b: 4).






HOBBS-THE CRAYFISHES OF FLORIDA


THE BLANDINGII SECTION
Diagnosis.-Cephalodistal margin of first pleopod of male never bears a
ridge or knob-like prominence unless it is a part of one of the terminal pro-
cesses; distad the appendage may be directed straight or caudad; mesial
process generally bent (either caudodistad or caudolaterad); a crescentric
terminal protuberance never present; cephalic process when present arises
from cephalic or cephalolateral margin, never from mesial surface; hooks
present on ischiopodites of third and fourth pereiopods.
Members of this section are found throughout the coastal regions from
Texas to New Jersey and throughout the middle west. They are found in al-
most any type of aquatic situation and frequently construct burrows.

BLANDINGII SECTION
Blandingii Group
Blandingii Subgroup
Clarkii Subgroup
Evermanni Subgroup
Fallax Subgroup
Spiculifer Group
Pictus Group
Pictus Subgroup
Lucifugus Subgroup
Seminolae Subgroup
THE BLANDINGII GROUP
The blandingii group consists of four subgroups (the blandingii, clarkii,
fallax, and evermanni) which constitute a very heterogeneous assemblage.
While the former three subgroups have rather distant affinities with each
other, apparently having been developed along different lines, they are tied
together by definite affinities with evermanni. Because of this a definition of
the blandingii group would involve a combination of the descriptions of the
four subgroups. The only characters common to all of the members of the
blandingii group are: hooks on the ischiopodites of the third and fourth perei-
opods; a relatively narrow or obliterated, long areola and a complex of gen-
eral similarities that are easier to see than to precisely define. Among these is
the structure of the rostrum which may or may not bear lateral spines; if
lateral spines are absent, then the margins are always interrupted. The are-
olae of the spiculifer and pictus groups are broad and short except in the
cavernicolous species of the latter.
THE BLANDINGII SUBGROUP
Diagnosis.-The first pleopod terminates in four well developed parts;
no hump is present on the anterior margin of the appendage; mesial process
subspiculiform or narrowly blade-like; cephalic process never spiculiform, is




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