HORTON H. HOBBS,

and THOMAS C. BARR, Origins and Affinities

of the Troglobitic

Crayfishes of

W North America

' (Decapoda: Astacidae) i

II. Genus Orconectes

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 105

Horton H. Hobbs, jr. Origins and Affinities

and Thomas C. Ban. Jr. r *_\ T I I_ V

J

 or the Iroglobitic

Crayfishes of

North America

(Decapoda: Astacidae)

II. Genus Orconectes

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
ABSTRACT

Hobbs, Horton H., Jr., and Thomas C. Barr, Jr. Origins and Affinities of the

Troglobitic Crayfishes of North America (Decapoda: Astacidae). II. Genus

Orconectes. Smithsonian Contributions to Zoology, number 105, 84 pages,

1972.—The troglobitic members of the genus Orconectes, comprising six species

and subspecies, are here assigned to the Pellucidus Section of the genus. Orconectes

australis australis frequents spelean habitats along the eastern edge of the Cumberland Plateau from Alabama to Kentucky, where it intergrades with Orconectes

australis packardi, which ranges as far north as Rockcastle County, Kentucky. Orconectes incomptus, new species, is known from only three localities at the base of

the Highland Rim in Jackson County, Tennessee. Orconectes pellucidus ranges

along the western flank of the Cincinnati Arch from Trigg County, Kentucky, and

Montgomery County, Tennessee, northeastward to Hart County, Kentucky. The

two subspecies of O. inermis occupy cave systems of the north Pennyroyal and

Mitchell Plain in Kentucky and Indiana, the nominate subspecies occurring in the

southern portion of the range and intergrading with the northern O. inermis testii

from near the state line to Monroe County, Indiana. The four species are believed

to have taken their origins from a wide-ranging, pre-Pleistocene, epigean stock

which frequented streams of low gradient. With subsequent uplift of the area, the

epigean derivatives of the stock for the most part became extinct, whereas the three

or four derivatives that invaded subterranean habitats gave rise to the modern troglobites which occupy much of the original range of their ancestors. Complete

bibliographic citations to these crayfishes and summaries of our present knowledge

of them preceed a listing of all of the caves from which troglobitic Orconectes have

been found, together with the identities of the crayfishes and previously reported

epizootic ostracods.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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Contents

Page

Introduction 1

Pellucidus Section 2

Systematic Position 3

Key to Genera of Troglobitic Crayfishes 4

Distribution and Phylogeny 4

Range of Pellucidus Section 4

Relationships of the Troglobitic Orconectes 5

The Ancestral Stock 5

Origin of the Pellucidus Section 5

Faunistic Associations 10

Key to Troglobitic Species of the Genus Orconectes 11

Orconectes australis australis 11

Orconectes australis packardi 22

Orconectes incomptus 32

Orconectes inermis inermis 36

Orconectes inermis testii 49

Orconectes pellucidus 58

Literature Cited 73

Appendix 81

List of Caves 81

Collectors 83

Horton H. Hobbs, Jr.

and Thomas C. Barr, Jr.

Origins and Affinities

of the Troglobitic

Crayfishes of

North America

(Decapoda: Astacidae)

II. Genus Orconectes

Introduction

The first mention of the existence of blind crayfishes

seems to be a reference in the minutes of the "Stated

Meeting of May 24, 1842" of the Academy of Natural

Sciences at Philadelphia (Anonymous 1843), in which

W. T. Craige, M.D., was credited with the donation

of "A white eyeless crayfish (Astacus Bartoni?) and

a small white fish, also eyeless (presumed to belong

to a sub-genus of Silurus), both taken from a small

stream called the 'River Styx' in the Mammoth

Cave, Kentucky, about 2J/2 miles from the entrance."

This crayfish occurring in Mammoth Cave was designated Astacus pellucidus by Tellkampf the following

year.

Not until 1871 did Cope describe the second

troglobitic species, Orconectes inermis, from Wyandotte Cave, Crawford County, Indiana, basing his

new genus on the albinistic character of the species.

The next troglobitic relative to be named was one

supposedly collected from a cave in Jugoslavia and

described in 1880 by Joseph as Cambarus typhlobius.

In the following year he mentioned the same species

under the names Cambarus coecus and Cambarus

stygius; all three names are almost certainly based

on a mislabeled specimen of Orconectes pellucidus

(see Holthuis 1964). The type is no longer extant

and repeated efforts to secure additional specimens

from the type-locality have been unsuccessful; to cast

further doubt on the existence of a member of the

Horton H Hobbs, Jr., Department of Invertebrate Zoology,

Smithsonian Institution, Washington, D.C. 20560. Thomas

C. Barr, Jr., Department of Zoology, University of Kentucky,

Lexington, Kentucky 40506.

NOTE AND ACKNOWLEDGMENTS

Part I of this series (Hobbs and Barr 1960)

treated the genus Cambarus, and since its publication only one additional troglobite, Cambarus

zophonastes Hobbs and Bedinger 1964, from

Arkansas, has been added to the genus. No

summary of the remaining troglobitic crayfishes

is planned by us. A key to the seven Floridian

species, together with summaries of their

ranges, was recently prepared by Hobbs

(1971b), and only three additional species are

known: Procambarus niveus Hobbs and Villalobos 1964, from Pinar del Rio, Cuba; Procambarus pecki Hobbs 1967b, from northwestern Alabama; and Procambarus rodriguezi

Hobbs 1943, from Veracruz, Mexico.

For their assistance in amassing most of the

collection on which this study has been based,

we express our appreciation to the many persons

cited in the appropriate places. Particular thanks

are due the following for the large number of

specimens and data which they made available

to us: Martha R. and John E. Cooper, H. H.

Hobbs III, Thomas C. Jegla, Russell M. Norton, and Stewart B. Peck.

We should also like to thank Rendell Rhoades

for calling to our attention the earliest reference

of Tellkampf to Orconectes pellucidus, and we

are indebted to Jack F. Marquardt for his

patience and assistance in obtaining many of

the references cited.

For assistance and suggestions during the

preparation of the manuscript and for criticisms

of the final draft, we are deeply grateful to

Fenner A. Chace, Jr., Mr. and Mrs. John E.

Cooper, Margaret A. Daniel, Joseph F. Fitzpatrick, Jr., Georgia B. Hobbs, and Mary H.

Moulton.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

subfamily Cambarinae in Europe is the fact that the

only other representative living on the continent is

Orconectes limosus which was introduced from the

United States.

Hay described Cambarus pellucidus testii from

Mayfield's Cave in Monroe County, Indiana, in 1891,

and not until 50 years later did Rhoades apply the

name Cambarus pellucidus australis to a crayfish

from caves in northern Alabama. In 1944, he named

the subspecies O. pellucidus packardi from caves in

the Cumberland watershed in Kentucky. An additional species, Orconectes incomptus, is described

herein.

Generic stability for the American troglobitic crayfishes has existed since Hobbs (1942a), and since that

time, all the forms mentioned above have been assigned to the genus Orconectes by all American crayfish systematists.

Thus, in the 100 years from 1844 to 1944, all

except one of the presently recognized troglobitic

crayfishes of the genus Orconectes had received

names, and agreement had been reached on their

generic designation. There were still some doubts on

the part of some of us as to the affinities of these

troglobites with each other and with epigean forms,

so that an amassing of collections throughout the

range of the complex had continued.

As a result of considerable field work by one of us,

Barr (together with that of others interested in cave

faunas), to determine the ranges and the limits of

variation of the several populations, conclusions not

in agreement with those of previous students of the

complex have been reached concerning the relationships of these crayfishes. We are proposing the recognition of a Pellucidus Section of the genus, comprised

of four species, two of them with geographic races

(subspecies).

PELLUCIDUS SECTION

Diagnosis: Albinistic; areola 3.7 to 6.7 times longer

than broad and constituting 34.0 to 46.0 percent of

total length of carapace; terminal elements of first

pleopod of first form male never constituting more

than

 J/s total length of appendage; males with hooks

on ischia of third or third and fourth pereiopods;

females with small uniramous pleopods on first abdominal segment; annulus ventralis of female slightly

movable, separated from sternum immediately cephalic to it by distinct sulcus.

The morphological bases for the above treatment,

the ranges of each of the taxa, and postulates concerning their origins and relationships constitute the

principal segment of this study.

The species and subspecies of the Pellucidus Section

of the genus Orconectes are:

O. australis australis (Rhoades)

O. australis packardi Rhoades

O. incomptus, new species

O. inermis inermis Cope

O. inermis testii (Hay)

O. pellucidus (Tellkampf)

A discussion of the distribution and phylogeny of

the newly erected Pellucidus Section is followed by

a key to its six members. In the treatments of the

individual species and subspecies, the synonymies are

believed to be complete, and, excluding references

to many newspaper articles, popular accounts, and

brochures, the bibliographic citations have been

searched and evaluated with reasonable thoroughness.

Following these citations are chronologically arranged

abstracts of the contributions made by each of the

authors with emphasis being placed on systematics,

distribution, ecology, and habits; comments, where

appropriate, follow the summaries of previous contributions. Diagnosis of the taxon is amplified with

full descriptions of type-specimens (or topotypes) and

illustrations. The specimens examined are cited together with locality, collector, and date. We have not

been consistent in presenting locality data, chiefly because we have attempted to follow the system of references utilized in the cave directories of the respective states (Barr 1961, Powell 1961, and Tarkington

et al. 1965). For Alabama and Indiana, Public Land

Survey coordinates are utilized; for Tennessee, latitude and longitude are employed; and for Kentucky

for which no cave atlas exists, we have given simple

directions for locating the caves. Lists are provided

giving names of the collectors (page 83) and, by state

and county, all the caves (page 81) in which the

species treated here have been reported, with references to the crayfishes and the symbiotic ostracods

which occur on them.

The troglobitic species which comprise the Pellucidus Section of Orconectes are most commonly

found in larger, permanent underground streams.

The largest populations occur in deep subterranean

lakes, the bottoms of which are covered with fine silt,

and through which water moves very slowly. When
NUMBER 105

cave passages flood—a common phenomenon in late

winter and early spring—the crayfishes quickly move

into the flooded areas, where they are sometimes

stranded in small pools as the flood waters recede.

Although crayfishes are rarely observed moving overland, their occasional presence in isolated, upper level

pools substantially above flood marks clearly indicates

that they can and do migrate from pool to pool.

Sudden and sharp disturbances of the water results

in rapid, backward darting of the cave Orconectes,

but with a less pronounced disturbance they slowly

crawl toward the deeper portions of the pools, sometimes disappearing beneath a ledge or pile of stones.

If a crayfish is stranded in a small pool or on land,

continued disturbance provokes a defensive stance,

the animal rearing up on the ambulatory pereiopods

and extending the open chelae. Some published statements to the contrary, troglobitic Orconectes definitely exhibit a negative phototaxis. Very dim, diffuse

illumination, of an intensity comparable to the candlelight of the early cave explorers usually has little

effect, which probably explains the older statements

that these animals are insensitive to light. If a strong

flashlight beam is focused on a crayfish, however, it

invariably moves away, after an initial delay of about

five to 10 seconds. The selective advantage of negative phototaxis is obvious.

An insect struggling on the surface of a quiet pool,

or gentle disturbance of the surface with a stick or

the fingers, soon attracts a nearby crayfish, which

orients beneath the disturbance and slashes upward

rapidly with the chelipeds. If a cave cricket or other

insect is siezed in this manner it is immediately

dragged beneath the surface and eaten. Crayfishes

are also easily attracted by meat or fish baits, and

can be caught readily in minnow traps.

Most of the available information on reproduction

in troglobitic Orconectes is applicable only to O.

inermis (Jegla 1966, Jegla and Poulson 1970). Although there is evidence that reproduction can occur

at any time during the year, there are apparently

definite seasonal peaks in the abundance of form I

males, ovigerous females, and the hatching of juveniles.

Systematic Position

Although epigean crayfishes occur on all of the continental masses except Africa, only in North America

are troglobitic species represented in the cave fauna.

Furthermore, in only one subfamily, the Cambarinae,

of the Holarctic Astacidae have some members become thoroughly adapted to a subterranean environment.

The taxonomic outline presented below indicates

the relationships of the troglobitic Orconectes to other

crayfishes occurring in the Northern Hemisphere.

Those taxa with troglobitic representatives are indicated by an asterisk. Because no up-to-date list of

these crayfishes is available, the described troglobitic

species, together with original bibliographic citations,

are listed under the appropriate genera.

Parastacidae.—Southern Hemisphere

*Astacidae.—Northern Hemisphere

Astacinae.—Europe, western Asia, and western

North America

Cambaroidinae.—Eastern Asia and Japan

Cambarellinae.—Gulf Coast and Mississippi Valley in the United States southward to the

Cordillera Volcanica Transversal in Mexico

•Cambarinae.—North America east of the Rocky

Mountains, southward to Guatemala, Honduras, and Cuba

*Cambarus.—United States east of the Rocky

Mountains and eastern Canada

C. (Aviticambarus) hamulatus (Cope 1881:

879).—Alabama and Tennessee

C. (Aviticambarus) jonesi Hobbs and Barr,

1960:19.—Alabama

C. (Erebicambarus) hubrichti Hobbs, 1952:

689.—Missouri

C. (Jugicambarus) cryptodytes Hobbs,

1941:110.—Florida and Georgia

C. (Jugicambarus) setosus Faxon, 1889:

237.—Missouri

C. (Jugicambarus) zophonastes Hobbs and

Bedinger, 1964:11.—Arkansas

[C. (Erebicambarus) cahni Rhoades, 1941:

146, is not a troglobite.—Alabama]

Fallicambarus.—Ontario, Michigan, and Illinois southward to Florida and Texas

Faxonella.—Oklahoma and Texas eastward to

South Carolina

Hobbseus.—Alabama and Mississippi

*Orconectes.—East of the Rocky Mountains

from southern Canada to Georgia and Texas

O. australis australis (Rhoades, 1941:142).

—Alabama and Tennessee
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

O. australis packardi (Rhoades, 1944a:

121).—Kentucky

O. incomptus, new species.—Tennessee

O. inermis inermis Cope, 1872a: 419.—Indiana and Kentucky

O. inermis testii (Hay, 1891:148).—Indiana

O. pellucidus (Tellkampf, 1844b:383).—

Kentucky and Tennessee

Paracambarus.—State of Puebla, Mexico

*Procambarus.—Southern New England and

Great Lakes southward to Honduras, Guatemala, and Cuba

P. acherontis (Lonnberg, 1895:6).—Florida

P. lucifugus lucifugus (Hobbs, 1940:398).—

Florida

P. lucifugus alachua (Hobbs, 1940:402).—

Florida

P. milieu Hobbs, 1971b: 115.—Florida

P. niveus Hobbs and Villalobos, 1964:342.

—Cuba

P. pallidus (Hobbs, 1940:394).—Florida

P. pecki Hobbs, 1967b: 2.—Alabama

P. rodriguezi Hobbs, 1943:203.—Veracruz,

Mexico

*Troglocambarus.—Peninsular Florida

T. maclanei Hobbs, 1942a: 345.—Florida

Key to Genera of Troglobitic Crayfishes

1. Ischium of third maxilliped without teeth on opposable border.

Troglocambarus Hobbs (1942b: 345)

Ischium of third maxilliped with teeth on opposable border 2

2. First pleopods of male terminating in two elements bent at no less than 90 degrees to principal axis of appendage Cambarus Erichson (1846:88)

First pleopod of male terminating in two or more elements; if only two, then both never bent

at angle as great as 90 degrees to principal axis of appendage 3

3. First pleopod terminating in two conspicuous elements, sometimes with minute rudiment of

third (caudal process); cephalic surface of appendage either lacking shoulder or with

shoulder adjacent to base of central projection Orconectes Cope (1872a: 409)

First pleopod terminating in two or more elements; if only two, then cephalic surface with

strong, angular shoulder never contiguous with base of central projection.

Procambarus Ortmann (1905b: 43 7)

Distribution and Phylogeny

RANGE OF THE PELLUCIDUS SECTION.—As can be

seen in Figures 1 and 2, the ranges of the troglobitic

crayfishes of the genus Orconectes are allopatric and

encompass, in three somewhat distinct geographic

areas, much of the karst terrane from northern Alabama to southern Indiana. In caves in Mississippian

limestone along the western margin of the Cumberland Plateau, Orconectes australis has been reported

from 61 localities from Jackson and Madison counties in Alabama to Rockcastle County in Kentucky.

Two geographic races of australis have been recognized, O. a. australis in the southern part of the

range and O. a. packardi in the northern portion.

A third member of the group, O. incomptus, is known

from three caves in Ordovician limestones at the

base of the Highland Rim in Jackson County.

Tennessee.

Along the western flank of the Cincinnati Arch, in

caves in Mississippian limestones of the Pennyroyal

Plateau, Orconectes pellucidus has been found in 26

localities. Its range is a narrow one, apparently confined to the gently tilted outcrops extending from

Trigg County, Kentucky, and Montgomery County,

Tennessee, northeastward to Hart County, Kentucky.

Within Hart County, apparently a barrier exists along

an east-west sandstone ridge somewhat paralleling

the Green River, which almost completely separates

the range of O. pellucidus from that of O. inermis

inermis. This ridge apparently resulted from deposition of sandstones in a pre-Pennsylvanian river channel (Burroughs 1923), and separates the Mammoth

Cave fauna from that of the northern Pennyroyal

(Barr 1967a).

Two subspecies of Orconectes inermis, together

with intergrade populations, frequent the cave systems of the north Pennyroyal and Mitchell Plain between Hart County in Kentucky and Monroe County

in Indiana, where they have been found in 44 localities. The nominate spiny race occupies the southern

part of the range, with typical members occurring
NUMBER 105

as far north as Crawford and Orange counties, Indiana. The spineless O. inermis testii is the troglobitic

form found in the northernmost part of the range

in Monroe County, Indiana; intergrading populations occur between Meade County in Kentucky and

the southern counties in Indiana.

RELATIONSHIPS OF THE TROGLOBITIC ORCONECTES.

—Hobbs (1969:119-126) has discussed the relationships of these troglobites to the members of other genera of the subfamily in which he cited several reasons

for believing that the troglobitic Orconectes, which

are here assigned to the Pellucidus Section, have

retained some of the most primitive characteristics

to be found among modern representatives of the

genus. Whether some of these characteristics are fixed

atavistic ones that reappeared when more recent ancestral forms "returned" to an environment resembling that of their ancestors or whether they represent

primitive retentions cannot be properly assessed in

the absence of a fossil record, but the fact that these

appear in what are considered to be primitive members of several genera of epigean crayfishes suggests

a primitive retention rather than products of convergent evolution.

One of the most regrettable facts related to the

present study is that the primitive facies of the first

pleopod and annulus ventralis of the Pellucidus Section have not been preserved in any of the epigean

species of the genus; they are present with slight

modification, however, in the not-too-distantly related

Cambarus (Veticambarus) pristinus Hobbs, 1965,

and to a lesser degree in C. (V.) bouchardi Hobbs,

1970, in which even the annulus ventralis bears a

marked resemblance to those of the members of the

Pellucidus Section. It can only be assumed that the

pattern in the epigean stock of the genus Orconectes

was destroyed during the Pleistocene and remains as

a relict in the present members of the Pellucidus

Section, the ancestors of which, having found refuge

in subterranean waters, escaped the fate of their

surface relatives.

T H E ANCESTRAL STOCK.—Three of the more generalized members of the Pellucidus Section (O. a.

australis, O. i. inermis, and O. pellucidus) seem to

us to be remnants of an ancient, probably no younger

than late Miocene, epigean stock which have undergone few modifications other than those channelized

by the successful occupation of a spelean environment. To paraphrase Hobbs (1969:120), had these

crayfishes retained a pigmented body, well developed

eyes, a shorter areola, and a more robust body and

appendages, they would exhibit essentially all of the

characteristics attributed by him to the hypothetical

"Adorconectid Stock," the supposed ancestral form

of the genera Orconectes, Hobbseus, Faxonella, Cambarus, Fallicambarus, and of the graciloid and mexicanoid lines of Procambarus. Not only do these

troglobites share more features in common with

extra-generic members of the subfamily Cambarinae

than do other members of the genus but also they

are most similar to those species of Orconectes that

are considered to be generalized.

ORIGIN OF THE PELLUCIDUS SECTION.—Postulates

concerning the origin of the genus Orconectes—hence

the Pellucidus Section—have been recently recounted

and evaluated by Hobbs (1967b, 1969). It is indeed

unfortunate that freshwater crayfishes have left so

few fossils. To our knowledge, not a single Tertiary

fossil of a member of the family Cambarinae has

been discovered, and the few Quaternary fossils available are almost certainly members of extant species.

Consequently, a reconstruction of the evolutionary

history of any cambarine group must be based upon

data derived from morphology, zoogeography, and

upon what is known of the geological history of the

area involved. Helpful, also, are data which permit

inferences concerning availability of suitable ecological conditions in areas supposedly occupied by ancestral stocks.

In a reconstruction of evolution one of the most

tantalizing factors is that of time. In this discussion,

consideration is given to the time during which the

ancestral stock invaded the subterranean waters, and

further, to whether there was a single invasion or

several of them. Obviously, in the absence of a fossil

record, no absolute timetable can be established for

the evolution of these crayfishes, but evidence exists

that the general facies of the ancestral Adorconectoid

Stock (Hobbs 1969:119, 1971a: 11) had been established prior to the formation of the transverse volcanic ridge in Mexico. Only south of this ridge does

the rather primitive Pilosimanus Group (Villalobos

1954:306) of the Mexicanus Section of the genus

Procambarus occur, and there can be little doubt that

the ancestors of this group were isolated from their

northern relatives with the volcanic upheaval north

of the Isthmus of Tehuantepec in early Pliocene

times. If time can be allowed for the ancestral pilosi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 1.—Ranges of Orconectes inermis inermis and intergrades (open circles), Orconeetes

inermis testii (stars), and Orconectes pellucidus (closed circles).
NUMBER 10 5

FIGURE 2.—Ranges of Orconectes australis australis and intergrades (open circles), Orconectes

australis packardi (stars), and Orconectes incomptus (closed circles).
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

manus stock to have reached Veracruz in a southward migration from the central part of the United

States, it is conceivable that the level of evolution

represented by the pilosimanus-pellucidus ancestors

(Adorconectoid Stock) had been accomplished not

later than the middle Tertiary. If this dating is accepted, then in all probability the ancestral pellucidus

or Archiorconectoid Stock (Hobbs 1969:119) was

lurking in the area to the north and east of the

Mississippi embayment at that time.

The Cincinnati Arch had its origin in the middle

Ordovician and was uplifted several times during

the Paleozoic and Mesozoic eras. As a result, considerable erosion must have been accomplished, so

that by the middle of the Tertiary the development of

a karst topography in the central Ordovician and

flanking Mississippian sediments had progressed to

a stage that invited the invasions of subterranean

streams by a number of epigean stocks. Thus it seems

entirely possible that as early as the Miocene the

subterranean waters were available to the surfacedwelling Orconectoid Stock (Hobbs 1969:119). For

the existence of a possible pre-Quaternary stock from

which the cave fauna might have taken its origin,

evidence may be had from an extrapolation of what

must have occurred in the dispersal of the ancestors

of Orconectes limosus, one of the closest extant relatives of the troglobitic forms and one which is restricted to the Atlantic slope east of the Appalachian

Mountains. This spiny crayfish is apparently most at

home in streams with a gentle gradient and has

never been found in cascading streams or in those

with a rapidly shifting bottom; furthermore, all its

closer relatives occur in similar habitats west of the

Allegheny Mountains and none to the south of the

Mountains. These facts strongly suggest that the

ancestors of O. limosus reached the Atlantic seaboard

through migrations to the east across and/or north

of the region presently occupied by the Alleghenies.

Although Rhoades (1962) proposed that it was the

Illinoian glacier that was responsible for impounding

the Ohio River and for reversing the flow of the

segment east of Cincinnati to the Chesapeake Bay,

carrying with it the limosus ancestors, an earlier

arrival of this stock seems far more likely to us.

This conclusion is based on the fact that (1) the

isolated range of limosus, which abuts the Atlantic

Ocean, is otherwise surrounded by that of more progressive members of the genus, and (2) limosus

shares only the upland streams (marginal in respect

to its range) with these more divergent forms (members of the Propinquus Group; for ranges see Crocker

and Barr 1968, Fitzpatrick 1967, Meredith and

Schwartz 1960, and Ortmann 1906). As to how

early in the Tertiary the migration eastward occurred

cannot be determined on the basis of available data,

but it must have been at a time that a low-lying

stream flowed from the West Virginia-Ohio region

into the Atlantic, or involved stream piracy in low

gradient headwaters. Either might well have existed

in pre-Pleistocene times, for there existed eastward

flowing rivers (for example, the old Erigan River),

and the Alleghenies were peneplained several times

during the Tertiary. Perhaps there was a deflection

of a segment of the upper Teays River during one of

the periods of peneplanation. If it could be tentatively assumed that the limosus ancestors reached the

east coastal waters in the Tertiary, then the relationship of its range to those of the eastern members of

the Propinquus Section could be explained by the

latter having reached their present ranges after

limosus had arrived, perhaps during the Illinoian

and Wisconsin glacial periods as proposed by Fitzpatrick (1967:166).

In attempting to determine the place of origin of

the Pellucidus Section, consideration has been given

to the intra-group relationships in order to ascertain

which assemblage within the Section is probably the

most generalized. On the basis of the pleopod of the

male, it is obvious that two types are represented:

(1) that found in O. pellucidus, which in lateral

view exhibits a slender distal portion, with the distally

directed mesial process distinctly longer than the

central element, a feature that seems to be unique

in the genus; and (2) the type found in O. australis,

O. incomptus, and O. inermis, in which the distal

portion of the appendage is comparatively broader

and the two terminal elements are of approximately

the same length, usually with the mesial process directed caudodistad. In the conformation of the pleopod, the australis-inermis type is more like those of

the epigean forms than is the pellucidus type. Also,

it is in O. a. australis and O. incomptus that the

primitive caudal element of the pleopod is present;

furthermore, the only indication of a cephalic process

in the entire genus is the reduced one observed on

both appendages of a single male of O. a. packardi

from Hydens Cave, Pulaski County, Kentucky. Many

populations of australis are quite spiny, as strongly

so as typical forms of pellucidus and inermis, and like
NUMBER 10 5

them, either occasionally (in a. australis) or typically

(in a. packardi), possess hooks on the ischia of the

third and fourth pereiopods. Except for the absence

of the caudal process on the first pleopod of inermis

and no indication of a cephalic one, O. i. inermis

is apparently equally as primitive as O. australis.

Therefore, we have concluded that the subspecies

of O. australis, inhabiting subterranean streams of

the Cumberland Plateau, approaches more nearly

the hypothetical adorconectoid stock than do any

of the others, and that incomptus and inermis are

more closely allied to australis than is pellucidus.

Hobbs (1967b, 1969) postulated that the ancestral

genus Procambarus had its center of distribution in

the southeastern part of the United States and that

the adorconectid segment became differentiated in

pre-Pliocene times. It spread through "an area extending from Alabama northward to Kentucky, on

and near the Cumberland plateau." This stock was

believed to have been a pioneering one, inhabiting

small to large streams flowing over a gentle to

moderate gradient, and, although able to negotiate

riffle areas, was not able to become established in

them. Whereas the most primitive Orconectes and

some of the more generalized members of the genera

Cambarus and Procambarus are still largely restricted

to such habitats, other stocks invaded the sluggish

areas of streams, lakes, ponds, swamps, and subterranean waters, and some eventually sought groundwater by burrowing. Puzzling is the fact that even

though the early Orconectoid stock gave rise to

several lines that resulted in comparatively broad

adaptive radiations (Hobbs, 1969), the Orconectes

line has, in general although not without exception,

remained more conservative in maintaining its adaptations to a lotic environment. Perhaps this apparent

conservatism, particularly from Tennessee and Missouri northward, was not actually real, for radiation

which might have occurred in the north could well

have been terminated during the Pleistocene; however, the fact remains that Cambarus diogenes and

Fallicambarus fodiens are burrowing species throughout an area which might well have been taken over

by burrowing species of the genus Orconectes. Regardless of what might have occurred prior to the

Pleistocene, no primary or secondary burrowing

(sensu Hobbs 1942b:20) Orconectes is known to

exist today, and there are fewer than a dozen of the

approximately 70 described species and subspecies

which typically frequent lentic habitats. Thus we

visualize the early Orconectes stock as resembling in

most of its facies a combination of the characteristics

exhibited by O. limosus and the troglobitic O. a.

australis, and differing only slightly, if at all, from

the former in its ecological requirements. By the

Mid-Cenozoic, the most primitive segment probably

inhabited streams in the gently rolling terrane of a

broad belt extending from Alabama northward to

the east of the old Mississippi embayment to, and

perhaps beyond, the present Ohio River. More progressive segments had, by this time, migrated around

the northern end of the embayment occupying other

virginal, in respect to crayfish habitation, streams

to the west.

Because of the marked similarities between O.

inermis and O. australis as compared with the striking

differences which exist between closely related epigean

allies having no more distant ranges, it is perhaps

difficult to conceive of the current troglobitic fauna

representing products of parallel development in

three segments of a primitive, wide-ranging surface

species. Equally incredible, however, would be an

assumption of the existence of east-west subterranean

aquatic corridors across the Cincinnati arch in Kentucky; even the supposition of a temporal continuity

is beset with difficulties in the region of the now

barren (in respect to troglobitic crayfishes) axis of

the Cincinnati Arch.

Should one be reasonably confident (which we

are not) of the previous existence of an aquatic

corridor through the dome of the Arch connecting

the spelean waters of the eastern edge of the Cumberland Plateau with those of the Pennyroyal Plateau,

this would allow the hypothesis that the Adorconectoid ancestor invaded the subterranean water system

of the Cumberland Plateau in Tennessee during the

Middle or Late Tertiary. From this center it could

have migrated along underground channels to the

northwest and south, gradually spreading throughout

an area that cumulatively is even larger than that

frequented by the extant descendants. The stock

would have had to cross the dome of the Cincinnati

Arch, cross the Pennyroyal Plateau, and ultimately

colonize the Mitchell Plain. Subsequent degradation

of the dome of the Arch, destroying the continuous

subterranean passageways in it, would account for

the isolation of the ancestors of australis-incomptus

from those of inermis-pellucidus; likewise, the previously mentioned sandstone ridge along the Green

River could well have disrupted communicating chan-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

nels along it, thus segregating the ancestors of inermis

from those of pellucidus. Further erosion of the underground passages of the Cumberland Plateau probably

would account for the segregation of incomptus as

a relic along the base of the Eastern Highland Rim

from ancestral australis at the edge of the Plateau.

An alternative hypothesis (which we consider more

probable) involves multiple colonizations of the subterranean waters by populations of a single widely

ranging ancestral epigean species: one on the Cumberland Plateau, ancestral to O. australis and O.

incomptus; a second on the Pennyroyal Plateau, precursor of O. pellucidus; and a third on the Mitchell

Plain, the antecedent of O. inermis. The close morphological resemblance between inermis and australis

could be accounted for by postulating a dual invasion

of the Teays and upper Cumberland drainages by a

common epigean ancestor. The barriers mentioned

above would have been equally effective should this

hypothesis prove to be more consistent with data

obtained in the future.

As has been pointed out above, and indirectly by

Barr (1968:85), there are no epigean remnants of

the Section present in the area occupied by these

troglobites. An explanation for their absence, according to Barr, lies in the fact that with the regional

uplift at the end of the Pliocene or beginning of

the Pleistocene, the low-gradient streams in which

the ancestral stock lived were transformed to rapidlyflowing ones with steeper gradients, thus destroying

the habitat of this stock and thereby bringing about

its extinction at the surface. It seems at least possible,

if not probable, that one or more of the colonizations

of the subterranean waters coincided with the extinction of the ancestral stock.

Subspeciation in both O. australis and O. inermis

has probably been a product of Pleistocene and postPleistocene evolution with greater diversity in body

proportions and spination in inermis and in spination

and pleopodial diversity in australis. Morphologically,

incomptus bears the same relationship to O. a.

australis as does O. i. testii to the nominate subspecies; however, the ranges of australis and incomptus, while being nearly contiguous, seem to be

completely isolated.

To recapitulate the major features of the hypotheses proposed to account for some of the factors

involved in the origin and evolution of the troglobitic

crayfishes of the genus Orconectes, it has been postulated that the ancestral stock of the Pellucidus Section

was derived from a primitive Procambarus stock in

the early or middle Tertiary; that it occupied streams

of low gradient in the area of the dome of the

Cincinnati Arch; that in the late Tertiary, three

segments invaded subterranean channels of the Cumberland Plateau, Pennyroyal Plateau, and Mitchell

Plain where they were isolated from one another,

respectively, by the belt of Middle Ordovician shales

along the crest of the Cincinnati Arch and by a

sandstone ridge which parallels the Green River in

Hart County, Kentucky; and finally, that subspeciation in O. australis and O. inermis and the isolation

of incomptus from australis occurred in Pleistocene or

Recent times. To account for the apparent closer

affinities between australis and inermis than of either

to pellucidus, the range of which is somewhat interposed between their respective ones, it is suggested

that the ancestral pellucidus stock, located in the

western part of the area, became differentiated from

the common stem first. Later, stream piracy between

the Cumberland and Teays systems resulted in there

being dispersed in the two basins a more recent common stock from which on the Cumberland Plateau

arose australis, and from which on the Mitchell Plain

was derived inermis. The ancestral surface stock, unlike that of the Floridian troglobitic members of the

Pictus Group of the genus Procambarus (see Hobbs

1958), did not survive to the present but was destroyed with the destruction of their habitat—the

result of a regional uplift at the close of the Pliocene.

FAUNISTIC ASSOCIATIONS.—Barr (1967a: 184) presented a sketch of the regional cave faunas of the

Interior Low plateaus. Within these faunal groups

associations of the troglobitic Orconectes are as

follows:

O. i. testii.—Northern Bedford fauna (I-A)

O. i. inermis.—Southern Bedford, Corydon, and

Breckenridge faunas (I-A,B,C)

O. pellucidus.—Mammoth Cave and Hopkinsville faunas (I-D,E)

O. a. packardi.—Rockcastle fauna (V-c)

O. a. australis.—Caney Fork and Huntsville

faunas (V-D,E)

O. incomptus.—Not characteristic of any regional fauna, but in the northeast corner of

the Central Basin faunal region

Troglobitic crayfishes appear to be absent from regions II (Bluegrass), III (Cumberland Saddle), IV

(Central Basin) except for O. incomptus, and V- F

(Guntersville).
NUMBER 10 5 11

Key to Troglobitic Species of the Genus Orconectes

(Based on First Form Male)

1. First pleopod without shoulder at cephalic base of central projection (Figures 10a, lla), or

distal portion of appendage much slenderer than that proximal to it (Figure 14a) 2

First pleopod with shoulder at cephalic base of central projection (Figures 3a, 4a, 9a) .... 4

2. Cephalo-caudal thickness of first pleopod immediately proximal to base of central projection

less than half maximum cephalo-caudal thickness (Figure 14a).

Orconectes pellucidus (Tellkampf)

Cephalo-caudal thickness of first pleopod immediately proximal to base of central projection

more than half maximum cephalo-caudal thickness (Figures 10a, lla ) 3

3. Rostrum usually without marginal spines or tubercles, and areola constituting at least 43

percent of entire length of carapace (Figure lie ) Orconectes inermis testii (Hay)

Rostrum with marginal spines and areola constituting less than 43 percent of entire length

of carapace (Figure 10c) Orconectes inermis inermis Cope

4. Rostrum without marginal spines or tubercles (Figure 9c); mesial process of first pleopod

comparatively heavy (Figure 9a, e) Orconectes incomptus, new species

Rostrum with marginal spines or tubercles (Figures 3c, 4c); mesial process of first pleopod

slender or not so heavy as in O. incomptus (Figures 3a, 4a) 5

5. Hooks usually present on ischiopodites of fourth pereiopods; shoulder at cephalic base of

central projection of first pleopod angular; caudal process of first pleopod absent but

caudal element prominently inflated (Figure 4e). Orconectes australis packardi Rhoades

Hooks usually absent on ischiopodites of fourth pereiopods; shoulder at cephalic base of central projection of first pleopod rounded; caudal process usually present as fine spiniform

tubercle, remainder of caudal element not inflated (Figure 3e).

Orconectes australis australis (Rhoades)

Orconectes australis australis (Rhoades)

FIGURES 2, 3, 5o-v, 6, 7, Si-cc

Cambarus (Faxonius) pellucidus australis Rhoades, 1941:

141-148, fig. 35 [Type-locality: Shelta Cave*, Madison

County, Alabama].

Cambarus pellucidus australis.—Hobbs, 1942a: 353.—Jeannel and Henrot, 1949:84.—Rhoades, 1959:399.

Orconectes pellucidus australis.—Hobbs, 1942a: 353.—

Rhoades, 1944:117, 121.—Hobbs, 1948a: 16, 19, 20, figs.

6, 13,; 1948b:85.—Eberly, 1958:3.—Nicholas, 1960:133.

—Eberly, 1960:30.—Fingerman and Mobberly, 1960:44,

45.—Barr, 1961:32, 33, fig. 10.—Hart and Hobbs, 1961:

175, 176, 178.— Rhoades, 1962:65, 69, 79, 92.—Jegla et

al., 1965:639.—Larimer et al., 1966:409-413.—Hart and

Hart, 1966:8.—Hobbs, 1967b:8, 9, 12, 15.—Cooper,

1967:14.—Jones and Varnedoe, 1968:2, fig. 1.—Cooper,

M., 1969:203, 204.—Cooper and Cooper, 1969a: 28;

1969b: 22-23.—Nicholas, 1969:14.—Hobbs, 1969:120,

121.

Orconectes (Orconectes) pellucidus australis.—Hobbs,

1942b: 154 [by implication].

Cambarus.—Jeannel and Henrot, 1949:24.

Orconectes pellucidus.—Pennak, 1953:458 [in part].—

Hobbs, 1967b: 12 [in part].—Poulson, 1964: 757.—Cooper,

1966:97.

•This cave has been purchased by the National Speleological Society and has been designated a Nature Preserve

(NSS News, 26(2) :28, 1968).

Orconectes (Orconectes) pellucidus.—Hobbs, 1959:890 [in

part].

Orconectes pellucidus pellucidus.—Hart and Hobbs, 1961:

176, 178, 180, 184.—Hart and Hart, 1966:9.

Orconectes australis australis.—Barr, 1967a: 161 [by implication].

Colorless crayfish.—Tarkington et al., 1965, map 6.

Orconectes.—Barr, 1968:85 [in part].

Blind crayfish.—Jones and Varnedoe, 1968:11, 18, 31.—

Nicholas, 1970:22, illus.

White crayfish.—Jones and Varnedoe, 1968:45, 79.—

Graham, 1969:4.

Cave crayfish.—Cooper, 1968:34.

Orconectes Pellucidus.—Cooper and Poulson, 1968:30, fig.

8.

Crayfish.—Torode, 1968:152; 1969:16.

REVIEW OF LITERATURE.—Cambarus pellucidus

australis was described by Rhoades (1941:141) from

Shelta Cave, SE

 l/4 NE % sec. 27, T.3S, R.1W, north

of Huntsville, Madison County, Alabama. He also

recorded its presence in five additional caves in

Madison and Jackson counties, Alabama. While his

description of the subspecies is brief, he emphasized

its important features, pointed out the principal variations, and contrasted it with the typical subspecies.

Included were brief notes on the caves from which

the crayfish was reported.

Hobbs (1942a) added no new data but in his
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

generic revision of the Cambarinae referred Rhoades'

subspecies to the genus Orconectes. Hobbs (1942b),

by recognizing the subgenus Faxonella, implied that

this subspecies was a member of the typical subgenus.

Rhoades (1944) assigned the subspecies of O. pellucidus to his new "Group rafinesquei" and contrasted

his new subspecies, O. p. packardi, with the previously described ones stating that it "differs from O.

pellucidus australis (Rhoades) (1941) in having a

much shorter areola and thicker, longer tips on the

gonopods. Also O. p. australis has hooks only on the

third walking legs."

Hobbs (1948a) figured the first pleopods of each

of the described subspecies of O. pellucidus except

O. p. testii, questioned the propriety of referring them

to Rhoades' Group rafinesquei, and indicated that

"if any division of the Limosus section is made then

it would seem that O. inermis and the various subspecies of O. pellucidus would constitute a natural

group that should receive a status equivalent to that

of the other subdivisions." He also included a key

for the separation of the species and subspecies belonging to the Limosus Section of the genus.

Hobbs (1948b), in remarking on the range of his

new Orconectes wrighti, indicated that "Among the

members of the Limosus Section only the cavernicolous Orconectes pellucidus australis . . . has been

collected farther south—in several caves in northern

Alabama—and no species belonging to this section

has been taken farther southwest."

Jeannel and Henrot (1949) reported this crayfish

from Shelta Cave and, as "Cambarus," from Crystal

Cave. Pennak (1953) simply listed O. pellucidus

from Alabama caves. Eberly (1958 and 1960) reiterated the presence of this subspecies in northern

Alabama.

Rhoades (1959) in discussing the status of Orconectes inermis indicated that "With the description of

Cambarus pellucidus australis Rhoades (1941) from

Alabama caves, it became evident that the hooks on

the fourth walking legs represent a significant character for generic determination. In light of this new

evidence, Hobbs (1942a) redefined the genus." Hobbs

(1959) included the subspecies of O. (O.) pellucidus

in his key to the North American crayfishes.

Nicholas (1960) included this subspecies in his

checklist of macroscopic troglobitic organisms but

erred in indicating that it was known only from the

type-locality.

Fingerman and Mobberly (1960) found that the

"eyestalks, supraesophageal ganglia, and circumesophageal connectives of this subspecies contain a

red pigment-concentrating substance and a distal

retinal pigment light adapting one." This is cited

by Poulson (1964).

Barr (1961) stated that "Two species of white,

eyeless crayfishes inhabit Tennessee caves. Orconectes

pellucidus australis (Rhoades) is found along the

western margin of the Cumberland plateau from

Kentucky into Alabama." He also included a photograph of this crayfish.

Hart and Hobbs (1961), in describing new entocytherid ostracods, listed this subspecies as a host of

Entocythere barri ( = Sagittocythere barri) in Cave

Spring Cave and Shelta Cave, Madison County, Alabama, and Blind Fish Cave, Putnam County, Tennessee. Entocythere ungulata (= Dactylocythere ungulata) was found on O. p. australis in Big Mouth

Cave, Grundy County, Tennessee, and in Blind Fish

Cave, Putnam County, Tennessee. Entocythere

steevesi (= Dactylocythere steevesi) and Entocythere

tuberosa (= Donnaldsoncythere tuberosa) were also

found to be infesting this crayfish in Blind Fish Cave.

The crayfish in Blind Fish Cave was mistakenly

identified as O. p. pellucidus by them.

Rhoades (1962) had the following to say about

O. p. australis: "The armature of the carapace and

chelae much reduced from that of typical O. pellucidus. Areola the longest of all the O. pellucidus

subspecies. Hooks on the third walking legs only.

Gonopods with very short tips and the outer ramus

tends to clasp the inner ramus. Annulus ventralis

with high hemispherical area close to the anterior

margin.

"This blind cave subspecies appears to be widely

distributed and in considerable abundance in the

large underground drainage of the Mussel Shoals

region." Later, he cited Jackson and Madison counties, Alabama.

Rhoades maintained the assignment of the pellucidus subspecies to his Group rafinesquei, discussed

the origins of the four subspecies recognized by him,

and concluded his treatment of O. p. australis with

"The southern subspecies of the blind crayfish seems

rather abundant in the well developed subterranean

drainage of northern Alabama. Dr. Walter B. Jones,

in outlining the ecology of Alabama caves (Rhoades,

1941), mentions that the presence of blind crayfishes

is associated with the presence of cave fishes and

aquatic insects. Individuals seem to be most numerous
NUMBER 105 13

where surface water enters caves during the wet

season. Debris washed in tends to initiate the food

chains. Once begun, food chains may continue among

animals present. Bat guano and the molds that grow

from these deposits are important sources of food for

cave species."

Jegla et al. (1965) indicated that O. p. australis

in two Alabama caves [Shelta Cave and Sauta Cave,

personal communication, M. R. Cooper] "differ by

20 mm in maximum length attained, by 20 mm in

length at maturity, and in inflection point for cheliped

allometric growth" and stated that "these variations

may be a result of different levels of food supply

and interspecific competition."

Tarkington et al. (1965) simply noted having observed both green and colorless crayfishes in Hering

Cave, Madison County, Alabama. All colorless ones

from this locality observed by us were O. a. australis.

Cooper (1966) indicated that this crayfish is the

largest "and by far the most numerous macroscopic

organism in the aquatic community" of Shelta Cave.

Hart and Hart (1966), in citing locality records

for entocytherid ostracods which had been garnered

from the national crayfish collection, cited new locality records for this crayfish as host to the ostracods.

Larimer et al. (1966), investigating the caudal

photoreceptors of this crayfish from Shelta Cave,

found that while sensitivity to variations in wave

length is not markedly different from that of certain

epigean species, "there is a uniform shift of the

spectral sensitivity toward shorter wavelengths, placing the peak at about 497 m/x. as compared to 502

mfi for the epigean species."

Barr (1967a: 161) utilized the combination Orconectes australis packardi, thus implying that Rhoades'

Orconectes pellucidus australis was the nominate subspecies, Orconectes australis australis. (See review of

literature for O. australis packardi.) Hobbs (1967b)

discussed the affinities of this crayfish with allies in

other genera. Cooper (1967 and 1968) contributed

no new information.

Barr (1968:85) discussed the range of the troglobitic Orconectes, commenting on their possible origins.

Jones and Varnedoe (1968) listed this crayfish

from several caves in Madison County, Alabama,

and included a photograph of a specimen from Shelta

Cave. Cooper and Poulson (1968) illustrated a crayfish from the same cave. Torode (1968:152) reported

the observation of two ovigerous females in Canyon

Cave; the same data were reprinted in 1969.

M. Cooper (1969), in comparing this crayfish with

the related epigean O. limosus, found that not only

does it possess chelae of greater relative length than

does the epigean species but also exhibits "an increased rate of elongation" in development. Furthermore, the remaining pereiopods are comparatively

longer and narrower. Whereas the total length of the

antennule is no greater than that of the epigean

species, the flagella are proportionately longer, and

the external flagellum bears "a greater number of

segments, greater number of asthetasc-bearing segments, and .. . a strikingly greater length of the

chemoreceptive asthetascs." The internal flagellum of

the antennule and the antenna, which bear mechanoreceptors only, "are considerably longer" in the

troglobite.

Cooper and Cooper (1969a) indicated that this

crayfish occurs in many caves in Madison and Jackson counties, Alabama. Cooper and Cooper (1969b)

noted the association of this crayfish with the salamander Gyrinophilus palleucus in Alabama. Nicholas

(1969) added no original data.

Hobbs (1969:120), in discussing the origins of

the genera Orconectes, Hobbseus, Faxonella, Fallicambarus, and Cambarus, pointed out the primitive

characteristics of O. a. australis.

Graham (1969:4) reported having collected two

white crayfish from Indian Rock Cave, Jackson

County, Alabama.

Nicholas (1970:22) included a color photograph

of this crayfish from Turkey Scratch Cave, Warren

County, Tennessee.

COMMENTS ON PREVIOUSLY RECORDED DATA.—(See

also comments on O. australis packardi and O. pellucidus.) Of Rhoades' (1941) remarks concerning

this subspecies, only a few of them need to be modified in light of specimens that have been subsequently

acquired. Some first-form males do have rudimentary

hooks on the ischiopodites of the fourth pereiopods.

He did not mention, nor did he figure, the short, very

slender caudal process that is present on the pleopods

of all of the first form males we have seen from the

type-locality as well as on those from several other

localities. The inner and outer rami ( = mesial process

and central projection) of the first pleopod of the

first form male are as widely separated in australis

as in any of the other species. Thus Rhoades' statement that "Outer ramus with corneous tip, curved

tightly around the inner ramus" (1941:142) and

"outer ramus tends to clasp the inner ramus" (1962:
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

69) probably refers to the more laterally directed

central projection in australis. Among our specimens

from Shelta Cave, the length of the areola ranges

from 36.9 to 40.5 percent of the entire length of the

carapace; from the nearby Hering Cave, also in

Madison County, 34.0 to 38.8 percent; and specimens

from other caves within the range of the subspecies

have areolae that constitute percentages within these

extremes (34.0 to 40.5 percent).

In his 1962 account, Rhoades indicated that the

areola of australis is "the longest of all the O. pellucidus subspecies" (page 69). While in some individuals this is true (because individuals of the subspecies are larger), the ratio of the areola length

to carapace length is not. That it is longer than that

of O. a. packardi was claimed earlier by Rhoades

(1944:121). Remeasurements of the types of the

latter indicate areolae constituting 42.4, 41.3, and

41.7 percent of the respective carapaces—percentages

exceeding that of most specimens of O. a. australis

examined by us. Rhoades (1941:144) cited a range

of 33.3 to 39.5 percent for australis and indicated

that in O. pellucidus the range was 36 to 41 percent.

Furthermore, the areolae of O. inermis testii sometimes constitute as much as 45.0 percent of the

carapace length.

Hobbs (1948b: 85) indicated that the range of

australis was situated farther south than that of any

other member of the Limosus Section, to which

australis and its allies were assigned and which embraces its closest epigean Orconectes relatives. In

1952, however, Penn described Orconectes hathawayi

(now considered to be a member of the Section)

from Rapides Parish, Louisiana (and it, or a closely

allied species, has been collected subsequently in

Clarke County, Alabama), thus extending the known

range of the Section more than 300 miles to the

southwest. And if O. lancifer (Hagen) should be assigned to the Limosus Section as suggested by Fitzpatrick in Fitzpatrick and Prins (1965:146), then

the range of the section extends some 600 miles to

the southwest in eastern Texas.

Rhoades' (1959:399) statement concerning Hobbs'

basis for redefining the genus is not clear to us; the

hooks on the fourth walking legs have generic significance only to the extent that a crayfish with

such hooks might belong to any of the cambarine

genera except Cambarus, Faxonella, and Hobbseus;

even various combinations of hooks on the ischia of

the second, third, and fourth pereiopods are not

really diagnostic for any of the crayfish genera

(Hobbs 1962:274).

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum with marginal spines or tubercles

delimiting base of acumen, margins subparallel, convex or converging, upper surface concave and without

median carina; postorbital ridges terminating cephalically in spines or tubercles; hepatic area often with

number of spines; at least one, often several, cervical

spines present; areola 3.9 to 6.0 times longer than

broad and constituting 34.1 to 41.9 percent of entire

length of carapace; chelae not conspicuously setose

but with ciliated tubercles, mesiodorsal surface of

palm with several irregular rows of tubercles; hooks

on ischiopodites of third, and sometimes fourth, pereiopods. First pleopod of first-form male with greatest cephalocaudal axis of pleopod less than twice that

immediately proximal to base of central projection,

always terminating in two elements, and frequently

bearing vestigial spinelike caudal process; non-corneous mesial process moderately narrow, subacute, directed caudomesially, and reaching distally approximately same level as central projection, partially

obscuring latter in caudal aspect; central projection

corneous, flattened in cephalocaudal plane and directed somewhat laterally; shoulder at cephalic base

of central projection rounded. Annulus ventralis only

slightly broader than long, and as illustrated in

Figure 3/.

Holotypic Male, Form I: Body (Figure 3c, i)

subovate, depressed. Abdomen narrower than thorax

(14.7 and 19.7 mm in widest parts, respectively).

Width of carapace greater than depth in region of

caudodorsal margin of cervical groove (19.7 and

15.0 mm). Areola moderately broad (5.1 times longer

than wide), with 6 minute punctations across narrowest part. Cephalic section of carapace 1.5 times

as long as areola; length of areola 40 percent of

entire length of carapace. Rostrum almost twice as

long as broad, excavate, and with acumen 5/6 as

long as width of rostrum at base; cephalic extremity

reaching distal end of antennular peduncle; margins

not swollen, only slightly elevated, and with small,

acute, corneous marginal spines at base of acumen;

upper surface with evenly spaced minute setiferous

punctations; subrostral ridges very weak and evident

for only short distance along basal rostral margins.

Postorbital ridges poorly developed, short, with
NUMBER 105 15

shallow dorsolateral grooves, and terminating cephalically in small corneous acute tubercles. Suborbital

angle lacking. Branchiostegal spines represented by

small acute tubercles. Seven or eight small cervical

spines or tubercles present on each side of carapace

immediately caudal to cervical groove. Carapace

punctate dorsally and granulate laterally; hepatic area

with few small acute tubercles but without spines.

Abdomen shorter than carapace (41.5 and 44.2 mm).

Cephalic section of telson with 2 strong spines in each

caudolateral corner, mesial ones movable.

Projecting portion of epistome (Figure 3A) subtriangular, with rounded basal extremities and with

small cephalomedian extension; concave surface with

crowded setae. Eyes much reduced, completely hidden

beneath rostrum in dorsal aspect and extending

cephalically only halfway between margin of orbit

and marginal spines of rostrum. Antennules of usual

form, with small spine on ventral surface of basal

segment. Antennae broken but probably extending

beyond caudal margin of telson. Antennal scale (Figure 3g) broadest distal to midlength, distinctly less

than half as broad as long; outer thickened portion

narrower than lamellar area and terminating in

prominent corneous-tipped spine. Third maxillipeds

extending almost to distal end of peduncle of antenna.

Chela (Figure 3/) slender and only slightly inflated; mesial margin of palm 1.5 times longer than

width of palm; dorsal surface of palmar area largely

tuberculate in mesial half and punctate in lateral

half, both punctations and distal bases of tubercles

with fine setae; ventral surface of palm entirely

tuberculate, with prominent corneous-tipped spine

opposite base of articulation of dactyl; tubercles along

mesial portion arranged in irregular longitudinal

series, innermost row of about 14 tubercles; lateral

margin of propodus costate along almost its entire

length. Fingers not gaping; dorsal and ventral surfaces of both with longitudinal ridges flanked by

setiferous punctations; opposable margin of immovable finger with row of 17 rounded, corneous tubercles, sixth from base largest; at lower level of same

margin, larger tubercle present between eleventh and

twelfth tubercles of upper row; single row of minute

denticles between fifth and eleventh tubercles and

similar denticles crowded distally to base of broken

tip; opposable margin of dactyl with 19 tubercles,

seventh from base largest, single row of minute denticles between first and fourteenth tubercle, distally

minute denticles becoming crowded and extending

to base of broken tip of finger; mesial surface of

dactyl with tubercles along proximal half and punctations along distal. Carpus longer than broad, with

proximal two-thirds of mesial, dorsal, and lateral

surfaces tuberculate; ventral surface punctate; mesial

surface with 2 large corneous-tipped spines, smaller

one ventral to larger; similar spine at mesioventral

distal angle, another adjacent to boss of ventrolateral

articulation with propodus. Merus mostly tuberculate

except ventrolaterally; tubercles present along almost

entire dorsal surface, and 2 corneous-tipped, subapical ones; ventrolateral margin with row of 12

spikelike tubercles and ventromesial margin with

approximately 16; scattered tubercles closely flanking

these rows, a contributing cause of inexact counts.

Ischium with rows of tubercles along outer and opposable margins, row of 7 on opposable margin,

with additional ones flanking it.

Ischia of third pereiopods (Figure 3k) with strong

simple hooks broadly rounded apically and projecting

proximad beyond distal margin of basis. Coxae of

fourth pereiopods with caudomesially projecting

prominences; coxae of fifth pereiopods without prominences except for small mesioventral projections at

bases of phallic papillae.

First pleopod (Figure 3a, e, j , k) symmetrical,

reaching bases of third pereiopods when abdomen

is flexed and only shallowly situated in sternal groove;

tip ending in two or three parts as described in

Diagnosis.

Allotypic Female: Differs from holotype in following respects (acumen broken so that some proportions and other details cannot be determined):

cervical tubercles and spines small but 9 on one side

and 11 on other; cephalic section of telson with 3

spines in caudodextral corner; third maxillipeds extending cephalically to distal end of penultimate

segment of peduncle of antenna; mesial row of 12

tubercles on palm of chela, opposable margin of

fixed finger with row of 16 (fifth from base largest),

and opposable margin of dactyl with row of 21

(sixth from base largest); mesial surface of carpus

of cheliped with 2 corneous-tipped tubercles, smaller

situated proximoventral to larger; upper distal margin

of merus with 1 spiniform tubercle, others as in holotype, lower surface with lateral row of 9 tubercles

and mesial row of 17. (See measurements.)

Annulus ventralis (Figure 3/) rather shallowly situ-
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Orconectes australis australis: a, mesial view of first pleopod of holotypic male,

form I; b, mesial view of first pleopod of morphotypic male, form II; c, dorsal view of carapace

of holotypic male, form I; d, lateral view of first pleopod of morphotypic male, form II; e, lateral view of first pleopod of holotypic male, form I; /, dorsal view of distal podomeres of cheliped of holotypic male, form I; g, antennal scale of holotypic male, form I; h, epistome of holotypic male, form I; i, lateral view of carapace of holotypic male, form I; ;', caudal view of first

pleopods of holotypic male, form I; k, ventral view of caudal thoracic region of holotypic male,

form I; /, annulus ventralis and portion of sternum of allotypic female; m, lateral view of carapace of male from Blind Fish Cave, Putnam County, Tennessee.
NUMBE R 10 5 17

ated in sternum and not firmly fused to sternal plate

immediately cephalic to it; outline subovate, 1.2 times

broader than long with median portion elevated

ventrally and forming broad arc having its highest

segment slightly cephalic to midlength; arched portion, bearing shallow longitudinal groove, ending

caudally on gently sloping plateau-like rim about

one-fourth length of annulus from caudal margin;

sinus originating at caudal base of arch sinistral to

median line, and after extending dextrally to median

line, turning suddenly caudally and ending on caudal

margin of annulus.

Morphotypic* Male, Form II: Differs from holotype in following respects: acumen, slightly longer

than width of rostrum at base, extending beyond

distal end of peduncle of antennule; third maxillipeds extending cephalically slightly beyond base of

ultimate segment of peduncle of antenna. Hooks

on ischia of third pereiopods unusually prominent for

second-form male, but boss on coxa of fourth pereiopod not so well developed.

First pleopod (Figure 3b, d) with no trace of

caudal process and mesial process extending slightly

beyond tip of central projection, latter especially

reduced and less well defined than in holotype.

Chelae lacking in morphotype but in second form

paratype differs from holotype as follows: mesial

row of only 10 or 11 tubercles on inner margin of

palm; opposable margin of fixed finger with row of

15 small rounded tubercles, progressively smaller

distally, with 3 somewhat larger ones forming row

below level of long row, 2 of them situated between

ninth and tenth tubercles of upper row and 1 between tenth and eleventh; carpus as in allotype;

merus with 1 spiniform tubercle near upper distal

end and with lower mesial row of 17 tubercles and

lateral one of 13.

MEASUREMENTS (in millimeters).—Orconectes australis australis:

Holotype Allotype Morphotype

Carapace:

Height 15.0 12.7 10.8

Width 19.7 16.6 13.6

Length 44.2 31.2

*Rhoades did not specifically designate a morphotype but

labeled one of the second form males as "allotype," as he

also designated the allotypic female. This "allotypic male" is

considered equivalent to the morphotypic male of other

authors.

Holotype Allotype Morphotype

Rostrum:

Width 6.0 5.5 4.4

Length 11.4 acumen 8.9

broken

Areola:

Width 3.5 3.3 2.3

Length 17.7 15.0 12.5

Chela:

Length of mesial margin

of palm 15.9 11.0 no

chela

Width of palm 10.3 7.3

Length of lateral margin

of chela 41.4 28.3

Length of dactyl 22.5 15.1

TYPES.—Holotype, allotype, and morphotype,

USNM 79363, 79364, 79365 (tfl, ? , rfH); Paratypes, Academy of Natural Sciences, Alabama Museum of Natural History, collections of Leslie Hubricht and Rendell Rhoades.

TYPE-LOCALITY.—Shelta Cave (SE %, NE y4, sec.

27, T.3S, R.1W), Huntsville, Madison County,

Alabama.

SPECIMENS EXAMINED.—Specimens from Alabama

and Tennessee were examined as follows:

ALABAMA: Jackson County: (1) Limrock Blowing Cave,

SW J4, NW l/4, sec. 11, T.4S, R.4E, 33II, 29 , S. B.

Peck, XII/29/65. (2) Doodlebug Hole (= Blowing Cave),

NW y4, NE }4, sec. 30, T.1S, R.4E, 131, John Stade,

11/14/60. (3) Saltpeter Cave, SW '/i> SW J4, sec. 16,

T.3S, R.3E, 1^1, 13II, 15 , W. B. Jones, VI/9/40. (4)

Kennamer Cave, NW l/4, SE y4, sec. 27, R.4S, T.3E, 1$ ,

Bret Blosser, X/20/63. (5) Salt River Cave, NW J4, NW

54, sec. 2, R.1S, T.6E, 1 9, T. C. Barr, Jr., X/17/54. (6)

McFarland Cave, SW «4, NW l/4, sec. 22, T.3S, R.3E,

2 31, 1J3, W.B.J., 11/29/40; 131, 2 3II, S.B.P., XII/

29/65. (7) Paint Rock Cave, SE y4, SW »4, sec. 29,

R.4S, T.3E, 1$ , S.B.P., IX/6/65; 19, S.B.P., XII/20/65.

(8) Guess Creek Cave, SW y4, SE y4, sec. 22, T.3S, R.4E,

2 31, 23II, 3 2 , T.C.B. and S.B.P., IX/5/65. (9) Jess

Elliott Cave, NW l/4, NE »/4, sec. 31, T.1S, R.6E, 131, 19

with eggs, S.B.P., IX/3/65. (10) Doug Green Cave, SW

l

/4, SW y4, sec. 9, T.2S, R.4E, 19, S.B.P., 111/14/66.

(11) Bell Spring Cave, NW 54, NE }4, sec. 9, T.2S, R.6E,

1 3 II, J. E. and M. R. Cooper, IV/9/66. (12) Fern Cave,

SE 54, SE }4, sec. 21, T.4S, R.3E, 231 , 49 , J.E.C. and

M.R.C., VIII/26/63; 19, H.H.H. Ill, VI/6/70. (13)

Borderline Cave, NE J4, NW 54, sec. 17, T.2S, R.3E, 1? ,

William Torode, XI/7/67. (14) Larkins Cave, NW l/4,

NE

 l/4, sec. 27, T.4S, R.4E, Ij3 , 1J9, S.B.P., IX/14/68.

(15) Sauta Cave (=Blowing Cave), SE '/4, SE y4, sec. 7,

T.5S, R.5E, 131, 13II, L. G. Conrad, VI/18/63. (16)
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Canyon Cave, NE 54, NE 54, Sec. 36, T.4S, R.3E, 1 5 with

eggs and young, Richard Graham and W.T., XI/29/68.

Madison County: (1) Shelta Cave (=Shelta Cavern), SE

54, NE 54, sec. 27, T.3S, R.1W, 131, 1 3 II, 19 (holotype,

allotype, morphotype), A. R. Cahn, III/1/38; 1 $ II, 29 ,

A.R.C., VI/1936; 13II, 19, Leslie Hubricht, VIII/5/39;

13 II, H. Henrot, VIII/16/46; 1$ I, 13II, 1$, Donald

Blair, VI/15/57; 2^1 , 13II, 4 9, T.C.B. and W.B.J.,

V/3/59; 131, 23 II, L.G.Co., VI/19/63; 2 411, 2 9, S.B.P.,

1965. (2) Sadler Spring Cave, SE 54, NE 54, sec. 3, T.4S,

R.1E, 13II, 1J9, W.B.J., VI/14/40. (3) Hering Cave

(=Cave Spring Cave), NE 54, NE 54, sec. 10, T.5S, R.2E,

2 9, Ij3 , W.B.J., IX/26/39; 3 3 II, 3 9, 1J3, 3j 9, W.B.J.,

XI1/1/39; 131, 4 3 II, 7 9, 4j $, 4j9, B. D. Valentine,

X/14/48; 2 3 II, 39 , 3j 9, B.D.V., X/21/48; 131, S.B.P.,

VIII/24/68. (4) Aladdin Cave, NE 54, SW J4, sec. 30,

T.2S, R.3E, 19, L.H., VI/18/57. (5) Burwell Cave, SW

54, NW 54, sec. 36, T.2S, R.2W, 131, S.B.P., 111/13/66;

6 31, 6 3II, 49 , Arthur W. Dobson, Jr., 11/10-11/68.

(6) Huntsville Spring Cave (-Big Spring Cave), SE J4,

SW 54, sec. 36, T.3S, R.1W, 19, W.B.J., X/6/39. (7)

Fuqua Spring Cave, NE 54, NW 54, sec. 15, T.1S, R.2E,

2j 3 , 2j 9, S.B.P. and A. Fiske, VI/25/67. (8) Byrd Spring

Cave, SW 54, NE 54, sec. 25, T.4S, R.1W, 29 , S.B.P., and

A.F., VII/5/67. (9) Matthews Cave, SE 54, SE # , sec.

12, T.4S, R.2W, 7 3 I, 2 3 II, 11 9, 2j 3 , S.B.P., VIII/26/68.

(10) Cold Spring Cave, NE 54, SW 54, sec. 28, T.3S,

R.1E, Ij3 , 1J9, S.B.P., VIII/20/68.

TENNESSEE: Fentress County: (1) Buffalo Cave, lat.

36°22'35"N, long. 84°57'36"W, 19, R. M. Norton, IX/

30/63. (2) Sells Cave, Lat. 36°33'25"N, long. 85°00'

24"W, 19, T.C.B., III/7/59. (3) Wolf River Cave, lat.

36°31'58"N, long. 84°56'38"W, 1 9 with eggs, J. E. Crouch

and T.C.B., VII/27/54. Franklin County: (1) Wet Cave,

Rowark Cove, lat. 35°13'47"N, long. 85°55'13"W, 131,

T.C.B., VII/15/54; 13II, T.C.B., X/14/56. (2) Walker

Spring Cave, lat. 35°13'36"N, long. 85°54'57"W, 19,

T.C.B., X/10/55; 13II, T.C.B., X/15/55. (3) Partin

Spring Cave, lat. 35°15'56"N, long. 85O52'51WW, 13II,

S.B.P. and A.F., VII/19/67. (4) Caroline Cove Cave, lat.

35°03'54"N, long. 86°07'41"W, 131, 1 3II, 4 9, 3j 9 ;

S.B.P. and A.F., VII/11/67. Grundy County: (1) Big

Mouth Cave, lat. 35°19'58"N, long. 85°34'48"W, 19, J.

N. Dent, VIII/2/55; 2 3 I, 6 3 II, 79, 1J3 , 5j 9, 1 9 with

eggs, R. B. Cumming, VIII/23/52. (2) Wonder Cave,

lat. 35°16'24"N, long. 85°50'59"W, 2 31, 1 9, J. M. Valentine, 111/17/31; 231, J.M.V. and J. C. Beakley, IV/19/35;

3 31, 13II, 29 , Charles E. Mohr, VI/30/37; 131, Nat.

Spel. Soc, VIII/30/41; 1 9, 2j 3 , H.H., VIII/13/46. (3)

Bear Cave, lat. 35°27'07"N, long. 85°34'48"W, 1 $ I, C. E.

McCary, X/12/55. (4) Crystal Cave, lat. 35°15'25"N.

long. 85°51'15", 19, H.H., VIII/13/46. Overton County:

(1) Raven Bluff Cave, lat. 36°29'33"N, long. 85o2r36wW,

19 with eggs, S. R. Gorin, VI/7/48; 13II, 19, T.C.B.,

11/28/59; 131, 13II, T.C.B., 111/19/61. (2) Sheep

Cave, lat. 36°12'41"N, long. 85°11'38"W, 19, T.C.B.,

VII/23/55. Putnam County: (1) Bridge Creek Cave, lat.

36°02'12"N, long. 85°37'55"W, 1 5II, T.C.B., VIII/4/54;

2 31, 4 3 II, T.C.B., 111/15/59. (2) Blind Fish Cave, lat.

36°03'19"N, long. 85°20'31"W, 1131, 2 311, 29 , T.C.B.,

C. E. Bush, and J. Harris, X/16/58; 131, T.C.B., IX/24/

60; 7 31, 2 3II, T.C.B., H. R. Steeves, Jr. and H. R. Steeves

III, XI/8/59. (3) Johnson Saltpeter Cave, lat. 36°04'

19"N, long. 85°47'38"W, 131, T.C.B., XI/5/55. Warren

County: (1) Cumberland Caverns, lat. 35°40'09"N, long.

85°40'51"W, 19, D. Egbert, VI/10/55; 131, Bert Denton,

II/7/54. (2) Turkeyscratch Cave, lat. 35°44'23"N, long.

85°36'00"W, 19, T. C. B., 1/1/58. White County: (1)

Indian Cave, lat. 35°49'01"N, long. 85°31'14"W, Ij9 ,

T.C.B., VI/19/55. (2) Wildcat Cove Cave, lat. 35°56'

38"N, long. 85°25'25"W, 2 9, 1J3, T.C.B. and L.H., XII/

24/56. (3) Ward Cave (=Dairy House Cave), lat. 35°

56'32"N, long. 85°27'03"W, 2 31, 3 3II, 2 9, T.C.B., X/

14/58; 13II, 19, R. Baroody and J. R. Holsinger, X/

28/69. (4) Ross Cave (exact location not known), 1 9, T.

E. Simpson, 1966.

RANGE.—Orconectes a. australis has a comparatively large range extending from the northern tributaries of the Tennessee River in Jackson and Madison

counties, Alabama, north-northeastward on the

Cumberland Plateau to the area of Fentress County,

Tennessee, and adjacent Wayne County, Kentucky,

where it intergrades with O. a. packardi.

VARIATIONS.—While no significance is attached to

political boundaries, they do furnish a convenient

means of associating variations in populations from

differing caves throughout the range of the subspecies with readily identifiable geographic regions.

For this reason, and the fact that the type-locality

is situated in the southernmost portion of the range,

the treatment is arranged by counties from the southwest to the northeast.

Not discussed below are the variations that occur

in the mesial process and central projection of the

first pleopod of the male, but some of them, including the extremes, are illustrated in Figure 8.

Madison County, Alabama: Specimens from the

type-locality (Shelta Cave) (Figure 7;) possess rostra

the margins of which may be subparallel, convex, or

convergent, and which, except in length, encompass

the limits of variation observed in the subspecies.

The marginal spines may be reduced to tubercles, but

the acumen is usually moderately long. The postorbital spines are quite variable in length, usually moderately long but occasionally reduced to tubercles. The

number of cervical spines, or tubercles, varies from

7 to 9, and those on the hepatic region from 2 to 5

(spines are usually rather heavy and short but more

spinelike in smaller individuals). The areola ranges
NUMBER 10 5 19

from 4.2 to 5.5 times longer than broad and constitutes 37.5 to 40.5 percent of the entire length of

the carapace. Hooks are invariably present on the

ischia of the third pair of pereiopods in males but

may be present, but never well developed, or absent

on the fourth pair. The caudal process of the first

pleopod of first-form males is always small, and in

some specimens is minute. The relative height of the

median longitudinal elevation of the annulus ventralis

is rather uniform, but the median furrow on it may

be very shallow or deep; the characteristic caudal

shelf is constant although it may be sloping or gently

curved.

In specimens from Hering Cave, the rostrum is

provided with longer marginal spines and acumen;

the cervical and hepatic spines are conspicuously long

and slender, and in number vary, respectively, from

4 to 8 and 0 to 7. The areola varies from 4.9 to 5.9

times longer than wide and comprises 34.6 to 38.8

percent of the length of the carapace. Hooks are

lacking on the fourth pair of pereiopods in the secondform males (no first-form males are available). The

median longitudinal elevation of the annulus is somewhat depressed and its longitudinal furrow almost

obsolete. The spines on the cheliped are comparatively

more prominent than in specimens from the typelocality.

In representatives of the populations in Aladdin

(Figure le) and Fuqua Springs (Figure 7g) caves,

the marginal spines on the rostrum are reduced to

tubercles and the acumen is only moderately long.

The number of cervical tubercles varies from 3 to 5,

and the hepatic spines are represented by low tubercles. The dimensions of the areola are within the

range of variation noted in topotypic specimens, and

the annulus ventralis is similar to that occurring in

specimens from Cave Spring Cave.

Specimens from Burwell (Figure 7i) and Matthews

caves are quite similar and comparatively large, and

an individual from Burwell Cave is the largest specimen of the species available to us. Its rostrum has

very short, heavy tubercles and the acumen is short.

The cervical spines are reduced in size and in number

(1 to 3), and there are no hepatic spines. The areola

varies from 4.1 to 6.6 times longer than broad and

comprises 38.5 to 42.7 percent of the carapace

length. All traces of hooks on the fourth pereiopod

in males from Burwell Cave are lacking, but in those

from Matthews Cave they are vestigial or well developed. The caudal process of the first pleopod is

well developed, and the annulus ventralis resembles

that in specimens from Shelta Cave.

The single female from Big Spring Cave (Figure

Id) has a unique rostrum (probably resulting from

an injury) in that the sinistral border bears 2 marginal spines and the dextral none. There are 6 cervical spines and 1 or 2 very small hepatic spines;

otherwise, the specimen is much like those from the

type-locality. In the Byrd Spring specimens (Figure

Ik) most spines are long but there are fewer cervical

(4 to 6) and hepatic (0 to 1) spines than in topotypes.

Jackson County, Alabama: Two of the most distinctive populations of this subspecies occur in Fern

(Figure 6p) and Saltpeter caves (Figure 6w). In

them, all the pereiopods seem to be more robust than

in specimens from elsewhere; the tubercles on the

chelipeds are heavy and short, and those on the

opposable margins of the fingers of the chela are

conspicuously strong. The marginal spines on the

rostrum are reduced to tubercles, and in one female,

almost obsolete; the cervical tubercles (occasionally

one spine) are quite small as are those on the hepatic

region, the latter tubercles sometimes lacking. The

areola is 4.2 to 5.3 times longer than broad and

comprises 38.5 to 39.6 percent of the carapace length.

Hooks are present only on the third pair of pereiopods in the male.

Similar to the specimens from Fern and Saltpeter

caves are the two crayfish from Jess Elliot Cave

(Figure 6v), but in them the pereiopods are not so

robust, and the tubercles on the opposable margins

of the fingers of the chelae are not nearly so strong.

Their areolae are 4.0 and 4.4 times longer than broad

and constitute 37.8 and 39.6 percent of the entire

length of the carapace. The first pleopod lacks a

caudal process, and the median longitudinal elevation of the annulus is somewhat depressed but has

a deep longitudinal furrow.

In sharp contrast to the populations just discussed,

those in Guess Creek Cave (Figure 6u) and Sauta

Cave (Figure 6r) have strong marginal spines on the

rostrum, a long acumen, as many as 8 slender, long

cervical spines and 7 hepatic spines. The pereiopods

are more like those in specimens from the typelocality, but the spines on the chelipeds are also long

and slender. The areola varies from 4.1 to 5.3 times

longer than broad and, reflecting the long rostra,
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

constitutes 36.2 to 39.3 percent of the total length of

the carapace (the three males, no more than 37.5

percent). In the specimens from Guess Creek Cave,

hooks are present on only the third pair of pereiopods,

but in the male from Sauta Cave, rudimentary hooks

are also borne on the fourth. The first pleopods of

the males lack the caudal process.

Somewhat intermediate in spination between the

Fern and Guess Creek specimens are those from McFarland (Figure 6$) and Doug Green (Figure 6q)

caves, but considerable variation exists in those from

the former; some have a few (one with 5) long

cervical spines and others have none—all reduced

to tubercles. The areolae are approximately 4.5 times

longer than wide and comprise 37.6 to 38.7 percent of

the carapace length. Hooks are present on only the

third pereiopods in the male and the first pleopods

lack or have very small caudal processes.

In addition to being almost aspinous (the rostrum

has been broken and partially regenerated), our

specimen from Doodlebug Hole (Figure 7 c) has an

unusual mesial process on the first pleopod of the

male (Figure 8a;); its laterodistal margin is emarginate, rendering the impression of a subterminal tooth

on that margin. The pleopod is devoid of a caudal

process, and hooks are restricted to the third pair of

pereiopods. Also almost spineless are the two females

from Paint Rock Cave (Figure 7a) and the single

females from Borderline, Kennamer (Figure 7 b),

and Salt River (Figure 6t) caves. The areolae vary

from 4.4 to 5.8 times longer than broad and constitute

37.0 to 37.7 percent of the carapace length.

Specimens from Limrock Blowing Cave (Figure

6o) are moderately spiny to smooth. Only secondform males and females are available, but the males

have no hooks on the fourth pereiopod. The areolae

are 3.9 to 5.0 times longer than broad and comprise

37.7 to 39.0 percent of the length of the carapace.

Franklin County, Tennessee: Specimens from the

four localities in Franklin County (Figure 6j-n) are

rather uniform. All are comparatively aspinous, possessing short marginal spines on the rostrum and

postorbital ridges, and having reduced cervical and

hepatic spines. The areolae range from 3.7 to 5.0

times longer than broad and constitute 36.4 to 41.0

(average 38.3) percent of the entire length of the

carapace. Hooks are present on only the third pereiopods, and the pleopods of the male lack a caudal

process. The annuli ventrales are typical of specimens

from the type-locality.

Grundy County, Tennessee: Three of the four

populations known to occur in Grundy County exhibit considerable variation. From Bear Cave (Figure

6g), Caney Fork drainage, the single specimen has

few and small spines. The areola is 4.3 times longer

than broad and constitutes 38.7 percent of the carapace length. It has hooks on only the third pair of

pereiopods, and the first pleopod is provided with a

caudal process.

In contrast, the population in Wonder Cave (Figure 6i), Elk River drainage, has both spiny and

aspinous members. Cervical spines are usually few

in number but are very prominent; hepatic spines

may or may not be present, but, if evident, are

usually small. The areola is 4.2 to 5.7 times longer

than broad and comprises 36.0 to 37.8 percent of

the total length of the carapace. Hooks are present

only on the third pereiopods of the male, and the

first pleopod may or may not bear a caudal process.

One specimen has a single cervical tubercle, no

hepatic spines, and the marginal spines on the rostrum are reduced.

In Big Mouth Cave (Figure 6h), Elk River drainage, most of the individuals have few, poorly developed spines, although one specimen could be described as very spiny. The areola is 4.7 to 5.7 times

longer than broad and occupies 36.8 to 39.8 percent

of the carapace length. In the males, hooks are present only on the third pair of pereiopods, and the

caudal process is absent.

Warren County, Tennessee: Only three specimens

from two localities are available from Warren County,

and the two from Cumberland Caverns (Figure 6e)

are similar to those from Bear Cave, Grundy County,

as would be expected, since both caves are close

together in the head of the Caney Fork basin. The

areolae are 4.7 and 4.8 times longer than broad, and

constitute 39.0 and 41.8 percent of the carapace

length. The single female from Turkey Scratch Cave

(Figure 6/), however, has few, but more strongly

developed, cervical spines; moreover, almost no trace

of hepatic spines is evident. The areola is proportionately broader and shorter than in those specimens

from Cumberland caverns—4.5 times longer than

broad and constituting 37.5 percent of the carapace

length. This specimen is morphologically and geographically intermediate between the other Warren

County specimens and those from White County.

White County, Tennessee: White County is represented in collections from four localities (Figures
NUMBER 105 21

6a-d), in three of which the specimens possess very

small rostral and cervical tubercles and the hepatic

ones are obsolete. The areola ranges from 3.5 to 5.0

times longer than broad and comprises 36.9 to 39.8

(average 37.8) percent of the carapace length. Welldeveloped hooks are present on the third pair of

pereiopods of the males, rudimentary ones occur on

the fourth, and the first pleopod bears a small caudal

process. In the single female from Ross Cave (Figure

6b), the rostral spines are long, as are the postorbitals, two cervicals, and one hepatic. The longitudinal

elevation of the annulus ventralis is somewhat

depressed.

Putnam County, Tennessee: With a single exception, the Putnam County populations typically

have long rostra. This is reflected in the fact that,

omitting two specimens, the areolae constitute from

35.7 to 38.5 percent (average 36.6) of the carapace

length, and are 3.8 to 5.0 times longer than broad.

In the two exceptions, one from Blind Fish Cave

(Figure 5t) and the other from Johnson Saltpeter

Cave (Figure 5v), the percentages are 39.6 and 39.3,

respectively. Much greater variation is apparent in

the relative development of the cervical and hepatic

spines. In Bridge Creek (Figure 5u) and Johnson

Saltpeter caves, they are characteristically small, as

they are in a few specimens from Blind Fish Cave,

but most of the latter have very long spines. Although

all these caves are rather close together in the Calfkiller Valley and its tributaries, Blind Fish Cave

differs markedly in the sort of habitat provided.

The crayfish occur in a wide, deep, and slowly moving

stream, rather than a small underground brook of

the type found in the other caves. Conceivably the

development of spininess is partly ecophenotypic, so

that individuals from deep lakes and pools are likely

to have longer spines than those from brook habitats.

In the males, there are hooks on the third pereiopods, and in a few, rudiments on the fourth. In one

second-form male, a hook is also present on one of

the second pereiopods. The caudal process of the

first pleopod may or may not be present.

Overton County, Tennessee: Only six specimens

from two rather widely separated localities are available from Overton County (Figure 5r, s) in the Obey

River drainage. These crayfish have very weak or

obsolete postorbital and hepatic spines, and the cervical spines are reduced to 1 to 3 small tubercles. The

areolae range from 3.7 to 4.9 times longer than

broad and comprise 38.7 to 41.9 (average 40.1) percent of the carapace length. In the single first-form

male, the hooks on the third pereiopods are well

developed and very weak ones are present on the

fourth; in the second-form males, there is no trace

of them on the fourth. The first pleopods possess a

minute caudal process, and, in the females, the longitudinal elevation of the annulus is somewhat depressed.

Fentress County, Tennessee: A single female from

each of three caves (Figure 5o, p, q) are the only

specimens available to us. Like the specimens from

Overton County, they have very reduced spiny elements. The areola constitutes 40.2 to 42.7 percent of

the carapace length, actually reflecting the short

rostrum, and is 4.1 to 4.8 times longer than wide.

The annulus ventralis also resembles that of specimens from Overton County.

Insofar as we have been able to determine, none

of the variations indicated are clinal; rather, they

occur in a mosaic pattern and, with few exceptions,

most are encompassed in the range of variations observed in the comparatively large series of specimens

from the type-locality. Were larger series available

throughout the range of the subspecies, certain combinations of characters might prove to be preponderant in some local populations. The only somewhat

distinctive ones recognized by us, however, are those

from Jackson County, Alabama, with robust pereiopods, and those from White County, with long rostra.

Populations which are interpreted to represent intergrades between O. australis packardi and the nominate subspecies are discussed under the treatment of

the former.

SIZE.—The largest specimen available is a second

form male, collected in Burwell Cave, Madison

County, Alabama, which has a carapace length of

48.0 mm. The largest first-form male (46.2 mm)

was found in the same cave, and the smallest firstform male was collected in Wonder Cave, Grundy

County, Tennessee (17.8 mm).

LIFE HISTORY NOTE.—First-form males have been

found during every month of the year except January, and a lack of collections made during that

month (only one specimen) is undoubtedly responsible for this gap. Our records fail to indicate a distinct preponderance of males of the first form during

any season. Ovigerous females were taken in June,

July, August, September, and November. William

Torode (1968:152; 1969:16) reported that on 29

November 1968 he and members of his party, in
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

exploring Canyon Cave, "found two female crayfish

carrying eggs, one of which was collected for John

Cooper. We watched the other one while little crayfish were crawling out of the eggs attached under the

abdomen." The preserved specimen has 66 first-instar

young and 14 eggs that had not hatched.

SEASONAL COLLECTIONS OF SPECIMENS EXAMINED

Not

Sex Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. known Total

6*1

9

juv. $

juv. $

ovig. $

Totals

_

_

1

-

-

1

10

7

5

1

-

_

23

7

6

4

-

-

17

2

1

_

_

-

-

3

2

1

4

-

-

7

7

8

8

2

4

1

30

2

2

8

3

1

18

13

11

26

5

6

1

63

11

4

6

2

1

1

25

17

15

18

4

7

-

61

1

2

-

1

4

1

8

8

2

3

-

19

_

2

4

-

-

6

73

65

94

16

24

5

277

Orconectes australis packardi Rhoades

FIGURES 2, 4, 5a-n, Ba-h

Orconectes pellucidus packardi Rhoades, 1944:113, 115,

117, 118, 121-122, figs. 3a-f [Type-locality: Cumberland

Crystal Cave (—Sloans Valley Cave) at Alpine, Pulaski

County, Kentucky].—Hobbs, 1948a: 16, 19, 21, figs. 8, 11.

—Eberly, 1958:3.—Cole, 1959:81.—Eberly, 1960:30.—

Hart and Hobbs, 1961:180.—Rhoades, 1962:65, 68, 79,

90.—Hart and Hart, 1966:8, 9.

Cambarus pellucidus.—Jillson, 1954:23.

Orconectes pellucidus.—Eberly, 1958:1-6 [in part].

Orconectes (Orconectes) pellucidus.—Hobbs, 1959:890

[in part].

Orconectes pellucidus packardii.—Nicholas, 1960:133.

Orconectes australis packardi.—Barr, 1967a: 161, 190.

Orconectes.—Barr, 1968:85 [in part].

REVIEW OF LITERATURE.—Orconectes australis

packardi was described by Rhoades (1944:121). His

description, consisting of comparisons with O. pellucidus and O. australis australis together with six

figures and measurements of the primary types, is

quoted in full: "O. pellucidus packardi differs from

O. pellucidus pellucidus by having a shorter acumen,

smoother carapace, longer areola, more ovate antennal scale, and more strongly curved tips of the gonopod. O. p. pellucidus always has hooks on both the

third and fourth walking legs but they are not always

present on the fourth walking legs of O. p. packardi.

"O. p. packardi differs from O. pellucidus australis

(Rhoades) (1941) in having a much shorter areola

and thicker, longer tips on the gonopods. Also O. p.

australis has hooks only on the third walking legs."

He also assigned the subspecies of O. pellucidus to

his new "Group rafinesquei." In addition to the

type-locality, Rhoades listed a sight record for Eureka

Cave, six miles west of Parker Lake, McCreary

County, Kentucky.

Hobbs (1948a) presented a key to the species belonging to the Limosus Section of the genus Orconectes, depicted certain diagnostic features, and questioned the advisability of recognizing the subdivision

of the Limosus Section into the Limosus and Rafinesquei groups as proposed by Rhoades (1944:113).

Jillson (1954) reported the presence of this crayfish in the type-locality.

Eberly (1958 and 1960), primarily concerned with

competition between O. inermis ( = his O. pellucidus)

and Cambarus laevis ( = C. bartonii laevis), presented

speculations concerning the relationships and origins

of the troglobitic members of the genus but presented

no additional information concerning O. australis

packardi.

Hobbs (1959), in his key to the crayfishes, referred

to this subspecies as one of four of Orconectes (O.)

pellucidus occurring between southern Indiana and

northern Alabama.

Nicholas (1960) included this crayfish in his checklist of macroscopic troglobitic organisms of the United

States and indicated that its range included "Caves

of Pulaski Co., Kentucky."

Hart and Hobbs (1961) indicated that Orconectes

pellucidus packardi serves as host for Dactylocythere

prionata (= Entocythere prionata) which they described from Wind Cave, Pulaski County, Kentucky.

Rhoades (1962) presented a short diagnosis of this

subspecies and offered a postulate concerning its

origin, essentially assuming that it was derived by a

"separation of a segment of O. pellucidus in the

caves of southeastern Kentucky. . . ."
NUMBER 105 23

Hart and Hart (1966) cited three new locality

records in reporting the hosts of the entocytherid

Sagittocythere barri which were obtained from the

national collection of crayfishes.

Barr (1967a) utilized the new combination O.

australis packardi, anticipating the completion of the

present study prior to that of his ecological treatment

of Mammoth Cave. In 1968, he indicated that "Three

or four separate colonizations are postulated to account for the known troglobitic species of Orconectes

and their present distribution."

COMMENTS ON PREVIOUSLY RECORDED DATA.—As is

obvious from the above summary of our previous

knowledge of this subspecies, almost nothing was

known about it, and the available descriptions and

figures are brief and incomplete. Except for confusion

that might arise from the statement that the areola

of this subspecies is shorter than that of O. a. australis, Rhoades' list of differential characters are clear

and concise. Whereas most populations of O. a.

australis attain a greater length than O. a. packardi,

and the absolute length of the areola is therefore

greater, there is an overlap in the range of variation

in respect to the ratio of the areola length to carapace

length. Most members of O. a. packardi, however,

exhibit a proportionately longer areola than do most

O. a. australis, the former ranging from 37.1 to

43.8 percent, and the latter from 34.1 to 41.9 percent

of the entire length of the carapace. As pointed out

above, usually this difference is a reflection of rostral

length rather than one in the length of the areola.

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum with marginal spines or tubercles

delimiting base of usually short acumen, margins

converging, its upper surface concave and lacking

median carina; postorbital ridges terminating cephalically in small spines or tubercles; hepatic area with

or without small spiniform tubercles; at least one

cervical spine or tubercle present, occasionally as

many as five; areola 4.2 to 5.9 times longer than

broad and constituting 37.1 to 43.8 percent of total

length of carapace; mesial surface of palm of chela

with several irregular rows of tubercles; well-developed hooks on ischiopodites of third pereiopods and

small or rudimentary ones often on those of fourth.

First pleopod of first-form male with length of greatest cephalocaudal diameter of pleopod less than twice

that immediately proximal to base of central projection, and always terminating in two terminal elements, caudal process absent or extremely vestigial;

broad, non-corneous mesial process directed somewhat caudally and distolaterally so that corneous

central projection, extending distally only slightly

beyond mesial process, almost completely visible in

caudal aspect; cephalodistal margin with distinct

angle at base of central projection. Annulus ventralis

approximately 1.7 times broader than long, and as

illustrated in Figure 4k.

Holotypic Male, Form I: Body (Figure 4c, I)

subovate, depressed. Abdomen narrower than thorax

(9.8 and 11.1 mm in widest parts, respectively).

Width of carapace greater than depth in region of

caudodorsal margin of cervical groove (11.1 and

9.2 mm). Areola broad (about 4 times longer than

wide), with widely scattered minute punctations,

only 3 or 4 across narrowest part. Cephalic section of

carapace about 1.5 times as long as areola; length

of areola 42.4 percent of length of carapace. Rostrum

approximately 1.6 times as long as broad, excavate,

and with acumen (although broken) slightly more

than 3/5 as long as width of rostrum at base; cephalic

extremity reaching almost to distal end of peduncle

of antennule; margins not swollen, only slightly elevated, and with small corneous tubercles at base of

acumen; upper surface with widely spaced minute

setiferous punctations; subrostral ridges moderately

strong and, in dorsal aspect, evident to almost midway between caudal margin of orbit and marginal

tubercles.

Postorbital ridges rather weak, short, with shallow

dorsolateral grooves, and terminating cephalically in

minute tubercles. Suborbital angle lacking. Branchiostegal spines acute. Five or six small, spiniform cervical tubercles present on each side of carapace immediately caudal to cervical groove. Carapace punctate dorsally and granulate laterally; hepatic area

with few very small acute tubercles but lacking spines.

Abdomen longer than carapace (29.0 and 26.2 mm).

Cephalic section of telson with 2 strong spines in

each caudolateral corner, mesial ones movable.

Epistome (Figure 4j) broadly rounded cephalically, with prominent subacute cephalomedian extension, surface concave (dorsally), with crowded setae.

Eyes much reduced, completely hidden beneath rostrum in dorsal aspect and extending cephalically

slightly less than halfway between caudodorsal margin

of orbit and marginal tubercles of rostrum. Antennules of usual form with prominent spine near distal

end of ventral surface of basal segment. Antennae

extending caudally almost to caudal margin of telson.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Orconectes australis packardi: a, mesial view of first pleopod of holotypic male,

form I; b, mesial view of first pleopod of morphotypic male, form II; e, dorsal view of carapace

of holotypic male, form I; d, lateral view of first pleopod of morphotypic male, form II;

e, lateral view of first pleopod of holotypic male, form I; f, dorsal view of distal podomeres of

cheliped of holotypic male, form I; g, caudal view of first pleopod of holotypic male, form I;

h, basal podomeres of third and fourth pereiopods of holotypic male, form I; i, antennal scale

of holotypic male, form I; j , epistome of holotypic male, form I; k, annulus ventralis and portion of sternum of allotype; I, lateral view of carapace of holotypic male, form I.

Antennal scale (Figure 4i) broadest near midlength,

almost half as broad as long; outer thickened portion

much narrower than lamellar area and terminating

distally in prominent corneous-tipped spine. Third

maxillipeds extending slightly beyond proximal end

of distal segment of peduncle of antenna.

Chela (Figure 4/) moderately heavy and somewhat

inflated; mesial margin of palm about 1.2 times

longer than width of palm; dorsal surface of palmar

area tuberculate mesially and proximally, otherwise

punctate, both tubercles and punctations with fine

setae; ventral surface of palm heavily tuberculate

mesially and punctate laterally, and without spine at

base of articulation with dactyl; tubercles along

mesial portion of palm arranged in irregular longitudinal series, innermost row of about 14 tubercles.

Fingers not gaping; dorsal and ventral surfaces of

both with rounded longitudinal ridges flanked by

setiferous punctations; opposable margin of immovable finger with row of 7 rounded corneous tubercles,
NUMBER 10 5 25

fourth from base largest, and several minute ones

continuing row distally; at lower level of same margin,

large tubercle present just distal to seventh tubercle

of upper row; several rows of minute denticles extending distally from seventh tubercle in upper row to

corneous tip of finger; lateral margin of finger subcostate; opposable margin of dactyl with row of 16

tubercles, sixth from base largest, those beyond eighth

exceedingly small; two or three rows of minute denticles extending distally from eighth tubercle to corneous tip of finger; mesial surface of dactyl with small

tubercles proximally and punctations distally. Carpus

longer than broad, with proximal two-thirds of mesial,

dorsal, and lateral surfaces tuberculate; ventral surface tuberculate proximally and mesially, otherwise

punctate; mesial surface with 3 prominent corneoustipped spines, largest near midlength, another proximal to it, and third immediately ventral to largest;

ventrodistal margin with prominent spine mesially

and conspicuous tubercle laterally, latter articulating

with socket on proximoventral surface of propodus.

Merus mostly tuberculate except ventrolaterally; tubercles along entire dorsal surface generally increasing in size distally, but none corneous-tipped; ventrolateral margin with somewhat irregular row of 11

spikelike tubercles and ventromesial margin with approximately 12; scattered tubercles closely flanking

both rows. Ischium with tubercles along outer and

opposable margins, opposable margin with row of

7 flanked by additional ones.

Ischia of third pereiopods only (Figure 4h) with

strong, simple, tapering hooks projecting proximally

beyond distal margin of basis. Coxae of fourth pereiopods with caudomesially projecting prominences;

coxae of fifth pereiopods without prominences except

for small mesioventral projections at base of phallic

papillae.

First pleopods (Figure 4a, e, g) symmetrical, barely

reaching level of caudal margins of coxae of third

pereiopods when abdomen is flexed, and shallowly

situated in sternal groove; tip ending in two parts

as described in Diagnosis.

Allotypic Female: Differs from holotype in following respects: tip of rostrum extending slightly

beyond peduncle of antennule; cervical tubercles

smaller; epistome, injured in previous instar, with

distinct subangular emargination on cephalodextral

border; chela with 10 tubercles along inner margin

of palm, tubercles more conspicuous and more linearly arranged on dorsomesial surface of palm, ventral

surface of palm less tuberculate, most setae on chela

longer, single row of minute denticles along almost

entire opposable margin of immovable finger interrupted by row of 18 very small corneous tubercles

along proximal three-fourths, opposable margin of

dactyl similar but with 21 small tubercles; carpus of

cheliped with large mesial spine bifurcate; spine ventral to it not conspicuously larger than other adjacent

ones. (See measurements.)

Annulus ventralis (Figure 4k) roughly subovate in

outline, with high, longitudinal, median elevation

devoid of shallow longitudinal trough; elevation highest (ventrally) along caudal half; subangular sinus

low on midcaudal face.

Morphotypic Male, Form II: Differs from holotype in following respects: rostral margins convex

laterally; only single cervical tubercle present; cephalic section of telson with three spines in caudodextral corner; chela closely resembling that of allotype but much smaller and slenderer.

First pleopods (Figure 4b, d) essentially similar to

that of holotype but terminal elements shorter with

central projection more rounded and non-corneous;

cephalic surface lacking angle at base of central

projection.

MEASUREMENT S (in millimeters).—Orconectes

australis packardi:

Holotype Allotype Morphotype

Carapace:

Height 9.2 10.5 7.8

Width 11.1 12.6 9.1

Length 26.2 30.0 22.3

Rostrum:

Width 3.6 4.3 3.3

Length 5.7 7.4 5.7

Areola:

Width 2.5 2.7 1.9

Length 11.1 12.4 9.4

Right Chela:

Length of inner margin

of palm 9.9 8.3 4.6

Width of palm 8.1 6.4 3.1

Length of outer margin

of chela 25.5 24.0 14.9

Length of dactyl 14.1 14.0 8.2

TYPES.—Holotype, allotype, and morphotype,

USNM 81310, 81312, 81331 (tfl, ? , cfH); Paratypes, Museum of Comparative Zoology, USNM, and

collection of Rendell Rhoades.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 10 5 27

TYPE-LOCALITY.—Cumberland Crystal Cave

(Sloans Valley Cave) at Sloans Valley, Pulaski

County, Kentucky.

SPECIMENS EXAMINED.—Specimens from Kentucky

were examined as follows:

McCreary County: (1) Eureka Cave, 0.6 mile NNW of

Nevelsville, 131, 19 , T.C.B., 11/14/59. (2) Steele Hollow Cave, 1.7 miles WNW of Bell Farm, 131, T.C.B. and

Russell M. Norton, IX/26/64. Pulaski County: (1) typelocality, 131, 19, G. H. Ehlers, X/8/41; 2 5II, G.H.E.,

date unknown; 131, 19, R. Rhoades, X/7/41 ; 1 $ II, 19 ,

G.H.E., 1/25/42; 2 31, \$U, 19 , T.C.B., VIII/9/63;

431, 33II, 4 9, 1J3, 1J9, S.B.P., W. M. Andrews, and

R.M.N., XI/5/64; 1 3 I, R. Walker, J. Purcell, and C. Harrell, 11/20/65. (2) Hydens Cave, 1.3 mi. NE of Blue

John, about 6 miles from Sloans Valley, 2 31, 2 3II, 3 9,

T.C.B., S.B.P., and W.M.A., VI/10/64. (3) Old Kentucky

Cave, 6 miles S of Somerset, 2 31, 1311, 8 9, 1J9, 19

with young, James R. Reddell and T.C.B., 1/29/67; 131,

13II, 2 9, Ij3 , Terrence G. Marsh, R.M.N., and L.

Merkle, III/4/67. (4) Pourover Cave, 0.8 mile ENE of

Colo in Happy Hollow, 4 9, T.C.B. and W.M.A., VI/2/65.

(5) Wind Cave, 5.0 miles SE of Somerset, 19 , T.C.B.,

XII/16/56; 23II, 19, T.C.B., IX/3/59; 13II, Jerry H.

Carpenter and T.G.M., II/8/69. (6) Baker Cave, near

Plato, 1 31, David P. Beiter and T.G.M., III/2/68; numerous small specimens stranded on gravel after flood [sight

record], T.C.B., 11/1962. Rockcastle County: (1) Duvalt

Cave, 3 miles SE of Mt. Vernon, near head of East Fork,

1 3II, 19 , S.B.P., VIII/30/64. (2) Fletchers Spring Cave,

1.0 mile N of Sand Springs on Dry Fork, 2 3II, T.C.B.

and R. A. Kuehne, VI/14/62; 131, 19 , T.G.M., 11/

20/68. (3) Pine Hill Cave, at Pine Hill on U.S. hwy. 25,

131, S.B.P., IX/7/64; 49 , J.R.R., IV/7/67; 1311, J- PVoigt, IX/9/65; 19 , L. G. Carr, date unknown. (4)

FIGURE 5.—Orconectes australis packardi (a-n), O. australis

australis (o—v), and intergrades between them illustrating

variation in spination: a, Duvalt Cave, Rockcastle Co., Ky.

( 9 ); b, Teamers Cave, Rockcastle Co., Ky. ( 9 ); c, Pine

Hill Cave, Rockcastle Co., Ky. (9) ; d,e, Cumberland

Crystal Cave, Pulaski Co., Ky. (31,311); /, Wind Cave,

Pulaski Co., Ky. ( 3 II); g, Pourover Cave, Pulaski Co., Ky.

(9) ; h, Hydens Cave, Pulaski Co., Ky. (31); i, Eureka

Cave, McCreary Co., Ky. (31); ;, Steele Hollow Cave,

McCreary Co., Ky. (31); k, Horse Hollow Cave, Wayne

Co., Ky. ( 9 ) ; /, Kogers Cave, Wayne Co., Ky. (31); m,

Blowing Cave, Wayne Co., Ky. ($1); n, Johnson Fork Cave,

Wayne Co., Ky. ( 9 ) ; o, Sells Cave, Fentress Co., Tenn.

( 9 ); p, Wolf River Cave, Fentress Co., Tenn. ( 9 ); q, Buffalo Cave, Fentress Co., Tenn. ( 9 ); r, Raven Bluff Cave,

Overton Co., Tenn. (311); s, Sheep Cave, Overton Co.,

Tenn. ( 9 ) ; t. Blind Fish Cave, Putnam Co., Tenn. ( 31);

u, Bridge Creek Cave, Putnam Co., Tenn. (31); v, Johnson

Saltpeter Cave, Putnam Co., Tenn. (31).

Teamers Cave, 1.2 miles NE of Mullins, 19, T.C.B., V/

11/63; 19, T.C.B., VI/18/63. Wayne County: (1) Blowing Cave, 0.75 mile SE of Sunnybrook at head of Carpenters Fork, 131, T.G.M. and Andrew R. Boone, VIII/21/67;

2 9, R.A.K., VII/25/65. (2) Horse Hollow Cave, 0.75

mile NW of Parmleysville in Horse Hollow, 1 3II, 6 9, 1 j 9,

S.B.P., VII/15/64. (3) Kogers Cave, 2.0 miles N of Hidalgo on west side of Shearer Valley, 131, T.C.B. and

W.M.A., II/5/67; 13II, J.H.C. and T.G.M., VII/13/68.

(4) Johnson Fork Cave. 0.4 mile E of Burfield on north

side of Johnson Fork, 19, T.C.B., VII/10/64.

Two females of this species in the Museum d'Histoire Naturelle in Geneve bear the label "grottee des

montagnes de Cumberland"; paratypes are in the collection of the Museum of Comparative Zoology, Harvard University, and that of Rendell Rhoades. The

latter paratypes have not been examined by us.

RANGE.—This subspecies seems to be confined to

subterranean passages in the upper Cumberland

drainage system in southeastern Kentucky. In the

southernmost localities in the State and in those in

the northern part of Tennessee, the characteristics indicate that packardi intergrades with the nominate

subspecies.

A geographic parallel exists between the australis

australis—australis packardi contact zone and the

peripheral distribution of certain troglobitic beetles

(Carabidae, Trechini) which one of us (Barr) is

currently investigating. The comparatively large (6-7

mm), eyeless beetles of the genus Nebonites Valentine

(1952) occur in many of the same caves. Nebonites

jonesi Valentine and Darlingtonea kentuckensis Valentine occupy roughly the same range as O. a. packardi in the Kentucky Cumberland Plateau, extending

a little farther north in the Kentucky River drainage

into a few caves which packardi has not attained.

Nelsonites walteri Valentine, on the other hand, occurs in Tennessee from Fentress to Van Buren

County, thus coexisting with O. a. australis throughout the northern half of its range. The range of the

large (7-8 mm) and very abundant Darlingtonea

kentuckensis stops with a single Tennessee cave record

just south of the Kentucky border. The TennesseeKentucky boundary is the approximate dividing line

separating still other elements of the regional trechine

fauna. To the north (Wayne, Clinton, and McCreary

counties, Kentucky) are Ameroduvalius jeanneli Valentine and a series of undescribed species of the

robustus and pubescens groups of Pseudanophthalmus
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 10 5 29

FIGURE 7.—Cephalic portion of carapace of Orconectes australis australis illustrating variation

in spination (all localities in Alabama) : a, Paint Rock Cave, Jackson Co. ( 9 ); b, Kennamer

Cave, Jackson Co. ( 9 ) ; c, Doodlebug Hole, Jackson Co. ( $ I) ; d, Big Springs Cave, Madison Co. ( $ ) ; e, Aladdin Cave, Madison Co. ($);/ , Sadler Spring Cave, Madison Co. ( $ II) ;

g, Fuqua Spring Cave, Madison Co. ( 5 ) ; h, Cold Spring Cave, Madison Co. ( $ II); i, Burwell

Cave, Madison Co. ( <JI) ; ;, Shelta Cave, Madison Co. ( 5 II) ; k, Byrd Spring Cave, Madison

Co. ($) .

FIGURE 6.—Orconectes australis australis, illustrating variation in spination: a, Wildcat Cove Cave, White Co., Tenn.

( 9 ) ; b, Ross Cave, White Co., Tenn. ( $) ; e, Indian Cave,

White Co., Tenn. ( 9 ) ; d, Ward Cave, White Co., Tenn.

($1); e, Cumberland Caverns, Warren Co., Tenn. ( 9) ;

/, Turkey Scratch Cave, Warren Co., Tenn. ( 9 ) ; g, Bear

Cave, Grundy Co., Tenn. ($1); h, Big Mouth Cave,

Grundy Co., Tenn. ($1); i, Wonder Cave, Grundy Co.,

Tenn. ($1); j , Wet Cave, Franklin Co., Tenn. (£11);

k, Walker Spring Cave, Franklin Co., Tenn. ( $U) ; I, Wet

Cave, Franklin Co., Tenn. ( $ II); m, Partin Springs Cave,

Franklin Co., Tenn. ($11); n, Caroline Cove Spring,

Franklin Co., Tenn. (SI); o, Limrock Blowing Cave, Jackson Co., Ala. ( 9 ) ; p, Fern Cave, Jackson Co., Ala. ($1);

q, Doug Green Cave, Jackson Co., Ala. ( 9 ); r, Sauta Cave,

Jackson Co., Ala. ( $1) ; s, McFarland Cave, Jackson Co.,

Ala. ( $ I) ; t, Salt River Cave, Jackson Co., Ala. ( 9 ) ;

u, Guess Creek Cave, Jackson Co., Ala. (Si); v, Jess Elliott Cave, Jackson Co., Ala. ( 9 ); w, Saltpeter Cave, Jackson Co., Ala. ($1).

(T. Barr, manuscript in preparation). To the south

(Overton, Fentress, and Pickett counties, Tennessee)

are Pseudanophthalmus beaklei Valentine, P. valentinei Jeannel, and one or more species of the intermedius group of Pseudanophthalmus. This general

zoogeographic phenomenon is thus expressed in two

very different groups of troglobitic arthropods. No

extrinsic barrier is postulated as the cause, since the

various trechine ranges show slight overlap and our

interpretation of australis s. str. and packardi as

geographic races of polytypic australis implies at least

a limited amount of gene flow.

VARIATIONS.—In Wayne County, the single female

specimen from Johnson Fork Cave (Figure 5n) has

two strong cervical spines, and the acumen of the

rostrum is distinctly longer than that of those speci-
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 10 5 31

mens from the other two localities. In a number of

respects, including the extreme development of

spines, the slender cheliped with longer fingers, and

the short areola, it resembles specimens from Eureka

Cave, McCreary County.

A single small first-form male from Blowing Cave

(Figure 5m), except for a reduction in the sizes of

the cervical and hepatic tubercles, seems fairly typical

of the subspecies. The first pleopod has a slight

prominence in the area on which the caudal process

is so well developed in O. a. australis.

The populations in Horse Hollow (Figure 5k)

and Kogers (Figure 5/) caves are quite similar,

having most spines on the carapace reduced to tubercles and possessing weak marginal spines on the

rostrum. The single first-form male has hooks on

both the third and fourth pereiopods, and the first

pleopod has a rounded shoulder on the cephalic surface and a very small caudal process (Figure Se).

Among the specimens from McCreary County, all

are small, the rostrum is short with convergent margins, and the cervical and hepatic spines are strongly

developed in the two from Eureka Cave (Figure 5J)

but practically obsolete in that from Steele Hollow

FIGURE 8.—(Abbreviations: cau, caudal process; cep, cephalic process; cpn, central projection; mes, mesial process.)

Distal portions of first left pleopods of males, form I, of

Orconectes australis packardi (a-d), Orconectes australis

australis (i-cc), and intergrades (e—h) to show variation in

the terminal elements. Orientation of all figures except d,

which is a mesial view, are primarily caudolateral aspects,

slightly altered to illustrate best the caudal process (clearly

evident in Figure aa): a, Pine Hill Cave, Rockcastle

Co., Ky.; b, Cumberland Crystal Cave, Pulaski Co., Ky.;

c,d, Hydens Cave, Pulaski Co., Ky.; e, Kogers Cave, Wayne

Co., Ky; f, Blowing Cave, Wayne Co., Ky.; g, Steele Hollow

Cave, McCreary Co., Ky.; h, Eureka Cave, McCreary Co.,

Ky.; i, Raven Bluff Cave, Overton Co., Tenn.; ;, Blind Fish

Cave, Putnam Co., Tenn.; k, Bridge Creek Cave, Putnam

Co., Tenn.; /, Johnson Saltpeter Cave, Putnam Co., Tenn.;

m, Wards Cave, White Co., Tenn.; n, Cumberland Caverns,

Warren Co., Tenn; o, Wonder Cave, Grundy Co., Tenn.;

p, Bear Cave, Grundy Co., Tenn.; q, Big Mouth Cave,

Grundy Co., Tenn.; r, Caroline Cove Cave, Franklin Co.,

Tenn.; s, McFarland Cave, Jackson Co., Ala.; t, Saltpeter

Cave, Jackson Co., Ala.; u, Limrock Blowing Cave, Jackson

Co., Ala.; v, Guess Creek Cave, Jackson Co., Ala.; w,

Doodlebug Hole, Jackson Co., Ala.; x, Jess Elliott Cave,

Jackson Co., Ala.; y, Fern Cave, Jackson Co., Ala.; z, Hering

Cave, Madison Co., Ala.; aa, Shelta Cave, Madison Co.,

Ala.; bb, Matthews Cave, Madison Co., Ala.; cc, Burwell

Cave, Madison Co., Ala.

Cave (Figure 5;). The areola is proportionately

shorter in these two localities (constituting 37.1 to

38.4 percent of the total length of the carapace)

than in any other known for the subspecies except

the one specimen from nearby Johnson Fork Cave

(Figure 5n), Wayne County (37.6 percent). The

merus of the cheliped bears unusually long spines

in the Eureka Cave specimens, and hooks are present

only on the ischia of the third pair of pereiopods.

Eureka Cave is seasonally inundated by waters of

Lake Cumberland, an impoundment of the Cumberland River, and the long spines may be related to

the deep quiet pools in which these individuals live.

In specimens from the type-locality (Figure bd, e),

the rostrum is usually short, with an acumen of

variable length; a few small hepatic spines are generally apparent. The length of the areola ranges from

38.5 to 43.1 percent of the entire length of the

carapace. The spines on the merus of the cheliped

are usually well developed, but never so long as in

specimens from Eureka Cave, and small or rudimentary hooks are more frequently present on the ischiopodites of the fourth pereiopod than not.

Among other specimens from Pulaski County, those

from Old Kentucky Cave possess hooks on both the

third and fourth pereiopods but cervical and hepatic

spines are reduced to rounded tubercles. Those specimens from other caves in the county exhibit hardly

more variations than are noted among individuals

from the type-locality, except those from Pourover

(Figure bg), Hydens (Figure bh), and Baker caves

have fewer spines on the carapace, and have areolae

that constitute no less than 40.3 percent of the carapace length. In one male from the latter locality, the

major spine on the mesial surface of the carpus of

the chela has four corneous tips, and on that of the

other male there are three. The latter male possesses

an accessory "projection" (Figure Sc, d) on the

mesiodistal surfaces of both first pleopods that could

be interpreted as a vestigial "cephalic process," but

comparable prominences are lacking in the pleopods

of the other male.

The two first-form males from Rockcastle County

have hooks only on the third pereiopod. In most of

the specimens, the cervical and hepatic spines are

reduced both in number and size; in some, only a

single small cervical spine is evident, and the female

from Teamers Cave has only a vestige of a cervical

spine. The four specimens from Duvalt (Figure 5a)
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and Teamers (Figure 56) caves have proportionately

longer areolae (40.5 to 43.7 percent) than do three

of the four from Pine Hill (Figure 5c) and Fletcher

Spring caves.

In the apparent absence of clines, and so few specimens from the assumed area of intergradation, our

evidence for the admixture of the gene pools of these

two taxa are limited; the continuity of the karst belt

occupied by them, however, enhances the supposition

that the two represent geographic races of a single

species.

SIZE.—The largest specimen available is a female

possessing a carapace length of 33.3 mm. This crayfish was collected from Pourover Cave, Pulaski

County, Kentucky. The largest first-form male (28.0

mm) was found in Hydens Cave, Pulaski County,

and the smallest (13.8 mm), in Baker Cave, Pulaski

County.

LIFE HISTORY NOTES.—First-form males have been

collected in January, February, March, June, August,

September, and October. No ovigerous females have

been found, but a single female carrying young was

taken in Old Kentucky Cave on 29 January 1967.

SEASONAL COLLECTIONS OF SPECIMENS EXAMINED

Not

Sex

$ I ...

$ II

$

juv. $

juv. $

9 with

young

Totals

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. known Total

1

15 14 12 17

21

22

42

2

3

1

91

Orconectes incomptus, new species

FIGURES 2 and 9

Orconectes pellucidus australis.—Hart and Hobbs, 1961:176

[not Rhoades 1941].

Orconectes pellucidus pellucidus.—Hart and Hart, 1966:9

[not Tellkampf 1844a].

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum devoid of marginal spines or tubercles delimiting base of acumen, margins converging,

its upper surface weakly concave and lacking median

carina; postorbital ridges terminating cephalically,

with or without small tubercles; hepatic area without

spiniform tubercles; three or four small cervical tubercles present; areola 4.1 to 5.2 times longer than

broad and constituting 39.6 to 43.0 percent of total

length of carapace; chelae conspicuously inflated and

setose; mesial surface of palm of chela with several

irregular rows of tubercles; well-developed hooks on

ischiopodites of third pereiopods only. First pleopod

of first-form male with length of greatest cephalocaudal diameter of pleopod less than twice that immediately proximal to base of central projection, and

terminating in three elements; short, broad, somewhat flattened mesial process directed caudodistad

and partially obscuring central projection in caudal

aspect; corneous central projection directed laterad;

minute caudal process situated between mesial process

and central projection; cephalodistal margin with

rounded prominence at base of central projection.

Annulus ventralis approximately 1.4 times broader

than long and as illustrated in Figure 9k.

Holotypic Male, Form I: Body (Figures 9c, I)

subovate, depressed. Abdomen narrower than thorax

(8.9 and 9.9 mm in widest parts, respectively). Width

of carapace greater than depth in region of caudodorsal margin of cervical groove (9.9 and 7.5 mm).

Areola broad (4.6 times longer than wide) with

scattered punctations, 4 or 5 across narrowest part.

Cephalic section of carapace 1.5 times as long as

areola. Length of areola 39.6 percent of length of

carapace. Rostrum approximately 1.5 times as long

as broad, shallowly excavate, and with acumen, although indistinctly set off at base, 1.5 times longer

than width of rostrum at base; cephalic extremity

reaching almost to distal end of peduncle of antennule; margins converging, not swollen, only slightly

elevated, and devoid of spines or tubercles; upper

surface with setiferous punctations; subrostral ridges

moderately well defined and evident, in dorsal aspect,

from caudal margin of orbit to almost half distance

to base of acumen.
NUMBER 105 33

Postorbital ridges moderately well developed,

grooved dorsolaterally, and terminating cephalically,

one bluntly and other in minute corneous tubercle.

Suborbital angle lacking. Branchiostegal spine small

and acute. Four small cervical tubercles present on

each side of carapace immediately caudal to cervical

groove. Carapace punctate dorsally and granulate

laterally; hepatic area with number of small tubercles

but lacking spines. Abdomen shorter than carapace

(21.7 and 22.2 mm). Cephalic section of telson with

2 strong spines in each caudolateral corner, mesial

ones movable.

Epistome (Figure 9;) broadly rounded cephalically,

with prominent cephalomedian extension and with

cephalosinistral margin slightly undulate. Surface

subplane and without setae. Eyes much reduced, not

completely hidden beneath rostrum in dorsal aspect

and extending cephalically about halfway between

caudodorsal margin of orbit and base of acumen.

Antennules of the usual form, with prominent spine

on ventral surface distal to midlength of basal segment. Antennae extending caudally beyond telson.

Antennal scale (Figure 9i) broadest distal to midlength and almost half as broad as long; outer

thickened portion much narrower than lamellar area

and terminating distally in prominent spine. Third

maxillipeds extending almost to midlength of distal

segment of peduncle of antenna. Chela (only left

one present, and it regenerated) with palm inflated

and finger long and comparatively heavy; mesial

margin of palm approximately 1.2 times longer than

width of palm; dorsal surface of palm tuberculate

mesially, otherwise punctate; both tubercles and

punctations with fine setae; ventral surface with

tubercles proximomesially, otherwise punctate; most

punctations and tubercles bearing conspicuously long

setae; spine lacking from surface opposite base of

dactyl; tubercles along mesial portion of palm sublinearly arranged, innermost row consisting of about

8 tubercles; lateral margin of palm subserrate, with

8 emarginations. Fingers not gaping; dorsal and

ventral surfaces with poorly defined longitudinal

ridges flanked by punctations bearing long setae:

opposable margin of immovable finger with row of

9 small tubercles along proximal third and with 2

somewhat larger ones distal to and below row;

crowded minute denticles present between tubercles

and extending distally to base of corneous tip of

finger; lateral margin of finger costate, grooved, and

bearing setiferous punctations; opposable margin of

dactyl with row of approximately 13 small tubercles

along proximal half, with crowded minute denticles

between tubercles and extending to corneous extremity of finger; setal tufts on both fingers conspicuously

long. Carpus longer than broad, with mesial surface

bearing one prominent spine and several tubercles,

one of which spiniform; tubercles extending onto

dorsomesial and ventromesial surfaces, otherwise

punctate; ventrodistal margin with prominent spine

mesially and another on boss articulating with ventral

socket of propodus. Merus tuberculate dorsally and

bearing two rows of spiniform tubercles ventrally,

outer row consisting of 9 and inner of 15; additional

small tubercles flanking both rows. Ischium with

several tubercles along opposable margin but not

arranged in discreet row.

Ischia of third pereiopods (Figure 9h) with strong,

simple hooks; hooks not extending proximal to distal

margin of basis. Ischia of fourth pereiopods without

hooks. Coxae of fourth pereiopods swollen caudomesially but lacking prominent caudomesially projecting prominences. Coxae of fifth pereiopods with small

mesially projecting prominences along cephaloventral

base of phallic papilla.

First pleopods (Figures 9a, e, g) symmetrical,

reaching coxae of third pereiopods when abdomen

is flexed, and shallowly situated in sternal groove.

Tip ending in three parts as described in diagnosis.

Allotypic Female: Differs from holotype in following respects: margins of rostrum with distinct

interruptions at base of acumen; subrostral ridges

visible in dorsal aspect for only a short distance

cephalic to caudal margin of orbit; postorbital ridges

ending cephalically without spines or tubercles; cervical tubercles extremely small and hepatic tubercles

almost obsolete; abdomen distinctly longer than carapace (23.4 and 22.2 mm) ; epistome with cephalolateral angles and ventral surface bearing setae; antennae extending caudad almost to end of telson;

chela with mesial row of 10 tubercles along margin

of palm; ventral surface of palm with small tubercle

at base of articulation with dactyl; opposable margin

of immovable finger with row of 7 tubercles (fourth

from base largest) along proximal two-thirds and

large one below row at base of distal two-fifths, between sixth and seventh tubercles; opposable margin

of dactyl with row of 4 tubercles along proximal
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 9.—Orconectes incomptus: a, mesial view of the first pleopod of holotypic male, form I;

b, mesial view of first pleopod of morphotypic male, form II; c, dorsal view of carapace of

holotypic male, lorm I; d, lateral view of first pleopod of morphotypic male, form II; e, lateral

view of first pleopod of holotypic male, form I; /, dorsal view of distal podomeres of cheliped of

morphotypic male, form II; g, caudal view of first pleopods of holotypic male, form I; h, basal

podomeres of third and fourth pereiopods of holotypic male, form I; t, antennal scale of holotypic male, form I; ;, epistome of holotypic male, form I; k, annulus ventralis and portion of

sternum of allotypic female; /, lateral view of carapace of holotypic male, form I.

two-fifths; ventral surface of merus with two rows of

9 tubercles.

Annulus ventralis (Figure 9A;) subovate, with

median longitudinal elevation somewhat depressed

and terrace-like caudal shelf, so obvious in O. australis

australis, not delimited from general curvature;

median longitudinal trough broad and shallow;

cephalic end of sinus originating slightly dextral to

median line, forming arc to median line and continuing caudally, ending on caudal margin of annulus.

Morphotypic Male, Form II: Differs from holotype in following respects: subrostral ridges scarcely

visible in dorsal aspect; postorbital ridges very weak

and terminating bluntly; cervical tubercles much reduced and hepatic tubercles almost obsolete; epistome

more similar to that of allotype, with cephalolateral

angle; antennae barely reaching cephalic margin of

telson; right chela (Figure 9/) with mesial half of

upper surface of palm tuberculate (11 in mesial row),

otherwise punctate, lower surface of palm with tu-
NUMBER 10 5 35

bercle at base of dactyl, outer margin of palm not

serrate, opposable margin of fixed finger with row of

8 tubercles (fourth from base largest) along proximal

two-thirds of finger and with single large tubercle

below row near midlength of finger, opposable margin

of dactyl with 9 tubercles (fourth from base largest)

along proximal four-fifths of finger; ventral surface

of merus of cheliped with mesial row of 8 tubercles

and lateral one of 12; hooks on ischia of third

pereiopods strong but not quite so well developed as

in holotype, absent from fourth pereiopods; caudomesial prominences on coxae of fourth pereiopods

and prominences on those of fifth obsolete.

First pleopod (Figure 9b, d) lacking trace of caudal

process but with short tuberculiform mesial process

and broad, prominent central projection; neither

element corneous.

TYPE-LOCALITY.—Cherry Cave, lat. 36°28'09"N.,

long. 85°36'28"W, Jackson County, Tennessee. A

description of the cave is presented in Barr, 1961:

272-273.

MEASUREMENT S (in millimeters).—Orconectes

incomptus:

Holotype Allotype Morphotype

Carapace:

Height 7.5 7.4 6.5

Width 9.9 9.5 9.1

Length 22.2 22.4 20.6

Rostrum:

Width 3.2 3.4 3.2

Length 4.8 3.5 5.1

Areola:

Width 1.9 2.3 1.9

Length 8.8 9.4 8.3

Chela:

Length of inner margin

of palm 6.2 7.6 7.4

Width of palm 5.2 6.4 6.6

Length of outer margin

of chela 16.5 18.2 17.0

Length of dactyl 10.2 9.6 9.1

DISPOSITION OF TYPES.—Holotype, allotype, and

morphotype, USNM 130299, 130300, and 130301

(cfl, ? , c? II) - Of the paratypes, 1 male, form II,

and 1 9 are in the collection of H. H. Hobbs III;

6 males, form II, and 8 females are in the Smithsonian Institution.

SPECIMENS EXAMINED.—Specimens from Tennessee

were examined as follows:

Jackson County: (1) type-locality, 1$ I, T.C.B., VIII/

8/59. (2) Carter Cave, lat. 36°16'53WN, long. 85°44'

10"W, 14II, T.C.B., VI/9/55; 19 , T.C.B., IX/1959;

5 3II, 6 9, J.E.C. and M.R.C., IX/21/68; 2$II, 2 9,

H.H.H. Ill, III/31/7O. (3) Haile Cave, lat. 36°20'00"N,

long. 85o43'08wW, 19 , T.C.B., V/6/59.

See Barr, 1961, for detailed locations and descriptions of these caves.

VARIATIONS.—The range of variation among the

available specimens is negligible, and the differences

pointed out in the accounts of the primary types seem

adequately to encompass those of any significance.

RELATIONSHIPS.—Although Orconectes incomptus

has very close affinities with O. a. australis and O. a.

packardi, we have no indication that intergrades exist

between it and either of the subspecies of australis.

Inasmuch as the differences between incomptus and

australis are so similar to those that distinguish O. i.

inermis from O. i. testii, we are tempted to treat

incomptus as a subspecies of O. australis. Despite this

similarity, there exists what we consider to be an

important difference in that there is abundant evidence of intergradation between the southern inermis

and the northern testii. In view of available data, it

seems most improbable that the subterranean passages

in Mississippian limestones on the eastern edge of

the Cumberland Plateau (the range of australis)

communicate with those of Ordovician limestones at

the base of the Eastern Highland Rim (the range

of incomptus), and if such connections do not exist,

then it is highly unlikely that intergrades between the

two might be found. The fact that the epizootic

Dactylocythere ungulata Hart and Hobbs, the only

entocytherid ostracod known to infest O. incomptus,

occurs on both australis and incomptus does not

necessitate the conclusion that such underground

connections are present, for the same ostracod also

occurs on Cambarus tenebrosus Hay which ranges

from the Nashville Basin eastward to the Tennessee

Valley, and it could have been responsible for transporting this ostracod via an epigean environment into

the two subterranean systems.

SIZE.—The largest specimen available is a female

having a carapace length of 24.5 mm. The only firstform male that we have seen is the holotype, in which

the carapace length is 22.2 mm.

LIFE HISTORY NOTES.—The single first form male

was collected on August 9; no ovigerous females or

females with young have been observed.

REMARKS.—The two bibliographic citations listed

are based upon a tentative determination made by
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the senior author, Hobbs, prior to a thorough study

of the crayfishes utilized in the current study. Hart

and Hart (1966:9) misinterpreted the identity of the

crayfish cited by Hart and Hobbs (1961:176) as

occurring in Carter Cave, Jackson County, Tennessee. Thus, O. incomptus should be included among

the hosts to the entocytherid ostracod, Sagittocythere

barn.

Orconectes inermis inermis Cope

FIGURES 1, 10, \2c-z, 13

Astacus pellucidus.—Cope, 1871a:4.—Cope 1871b:368;

1872c:297.

Orconectes inermis Cope, 1872a: 409, 410, 419, 420, fig.

116 [type-locality, Wyandotte Cave, Crawford County,

Indiana].—Cope, 1872b: 161, 162, 173, 174, fig. 116.—

Hagen, 1872:495.—Packard, 1873:94.—Faxon, 1884:

139; 1885a:40, 42-43.—Underwood, 1886:371.—Packard, 1888:39.—Sloan, 1888:[3 letters to Packard; see

Packard, 1888:15-16].—Lonnberg, 1894:126.—Lonnberg,

1895:4, 5.—Hay, 1896:484-485.—Blatchley, 1897:171.—

Hay, 1897:208.—Faxon, 1914:415.—Hobbs, 1942a:335,

350, 352, 354, 355, fig. 12; 1948a: 16, 19, 20, figs. 10, 14.

—Eberly, 1955:281, 282.—Holthuis, 1956:113, 116.—

Crocker, 957:13.—Eberly, 1958:3.—Rhoades, 1959:399,

400, 401.—Eberly, 1960:30.—Creaser, 1962:2, 3.—

Rhoades, 1962:65.—Fitzpatrick, 1963:60.—Barr, 1967a:

186; 1967b: 481.—Fitzpatrick, 1967:141, 142.—Poulson

and White, 1969:974, 975.

Cambarus pellucidus.—Packard, 1872a:30; 1873:94 [in

part].—Smith, 1873:639 [in part].—Collett, 1874:305.—

Smith, 1875:477 [in part].—Faxon, 1884:139 [in part];

1885a:42, 158, 168 [in part].—Sloan, 1888:15, 39-41, 82

[in part].—Packard, 1888:15, 16, 19, 39, 40, 41, 42, 82, 86,

118, 119, 122, 123 [in part].—Faxon, 1890:621, 626, 628

[in part].—Packard, 1890:393.—Hay, 1891:147, 148.—

Ortmann, 1892:11 [in part].—Hay, 1893:283-286, figs.

1, 3, 4, 7, 8, 9, 13, 14.—Lonnberg, 1894:126 [in part].—

Hay, 1896:478, 482-485, 489, fig. 3.—Blatchley, 1897:

138, 142, 144, 170, 171, 174 [in part].—Hay, 1897:208,

209, 210.—Faxon, 1898:647 [in part].—Hay, 1899:959,

966 [in part]; 1902a:235 [in part].—Ortmann, 1902:277,

278, 279 [in part].—Steele, 1902:16 [in part].—Eigenmann, 1903:169 [in part].—Harris, 1903b: 77, 112, 117—

118, 152 [in part].—Ortmann, 1905a: 92, 107, 108 [in

part].—Graeter, 1909:470.—Osborn, 1912:923.—Faxon,

1914:392.—Spurgeon, 1915:385-394 [in part].—Ortmann, 1918: 838, 848 [in part].—Spandl, 1926:95, 141

[in part].—Pope, 1926:170.—Chappuis, 1927:91, 120 [in

part].—Ortmann, 1931:64 [in part].—Bolivar et Jeannel,

1931:302.

Orconectes pellucidus inermis.—Smith, 1873:639.—Faxon,

1885a: 83.—Packard, 1888:41.—Rhoades, 1959:401.—

Nicholas, 1960:133.—Hart and Hobbs, 1961:176 [in

part].—Rhoades, 1962:68, 79, 81, 90.—Jegla et al., 1965:

639.—Jegla, 1965a:647; 1966:345-357.—Hart and Hart,

1966:8.—M. Cooper, 1969:203, 204, 206-207.—Jegla

and Poulson, 1970: 347-355.

Cray fish [sic.].—Shaler, 1875:362.

Crayfishes.—Collett, 1874:303.—Collett, 1879:362.

Orconectes.—Hovey, 1882:223 [in part].

Blind crayfish.—Packard, 1888:14.—Pope, 1926:163, 169.

—Jackson, 1953:31; 1955:57.

Eyeless crayfishes.—Sloan (in Packard), 1888:16.

Orconectes pellucidus.—Cope, 1879:492, 494, 495, 503,

505, 506 [in part], fig. 116.—Packard, 1888:128, 155

[in part], pl.-figs. 21:5, 22:5, 6, 7.—Pennak, 1953:458

[in part].—Eberly, 1955:281, 282; 1958:1-6 [in part];

1960:29, 30, 31, 32 [in part].—Huheey, 1961:43-45.—

Penn and Fitzpatrick, 1963:793.—Poulson, 1964:759.—

Vandel, 1964:509, 570, 575 [in part].—Jegla, 1964:81.—

Jegla, 1965b: 1421.—Mohr and Poulson, 1966:119, 120,

122, 123.—Poulson and Jegla, 1966:88.—Jegla and Poulson, 1968:273-282.—Jegla, 1969:135-137.

Cambarus (Orconectes) pellucidus.—Hovey, 1882:222 [in

part].

Cambarus pellucidus inermis.—Packard, 1888:41.

Cambarus pellucidus variety inermis.—Hay, 1893:284.

Cambarus inermis.—Faxon, 1898:647.

Orconectis inermis.—Harris, 1903b: 113 [erroneous spelling].

Cambarus (Faxonius) pellucidus.—Ortmann, 1905a: 107,

108 [in part]; 1905b:435 [in part]; 1931:64 [in part].—

Fage, 1931:373 [in part].

Blind crawfish.—Pope 1926, fig. 157.

Cambarus.—Bolivar et Jeannel, 1931:298, 299 [in part].

Cambarus pellucidus var. testii.—Wolf, 1934-38:105 [in

part].

Orconectes (Orconectes) inermis.—Hobbs, 1942b: 154 [by

implication].—Hobbs, 1959:890.

Cambarus (Cambarus) pellucidus testii.—Balss, 1955:1311.

Orconectes pellucidus pellucidus.—Eberly, 1960:30 [in

part].—Minckley, 1963:47, 51, 74.—Hart and Hart,

1966:8 [in part].—Hart and Hart, 1969:167.

Orconectes pellacious.—Jegla, 1965b: 1421 [erroneous spelling in abstract].

Orconectes.—Mohr and Poulson, 1966:123.—Barr, 1968:

85, 95 [in part].

Orconectes inermis testii.—Barr, 1966:17.

Orconectes sp.—Hart and Hart, 1966:8.

Cave crayfish.—Mohr and Poulson, 1966:91-92, 120, 121,

123.

Orconectes inermis inermis.—Barr, 1967a: 161.—Prins,

1968:672.

Orconectes "probably O. inermis".—Barr. 1968:90.

REVIEW OF LITERATURE.—The first record of the

existence of this crayfish was that of Cope (1871a: 4)

who listed "Astacus pellucidus" among the species

obtained in Wyandotte Cave in the Indianapolis

Journal on 5 September 1871. Later the same year he

(1871b: 368) again referred to it as he did in the

newspaper. "We descended a wall to the water,

some twenty feet below the surface, and found it to
NUMBER 10 5 37

communicate by a side opening, with a long, low

channel, through which flowed a lively stream of

very cool water. Wading up the current in a stooping

posture, we soon reached a shallow expansion or

pool. Here a blind crawfish was detected crawling

around the margin, and promptly consigned to the

alcohol bottle." This description has led us to conclude that this was not Wyandotte Cave; instead, it

is the cave nearby known as Sibert's Well Cave. The

following year, in identical papers Cope (1872a and

1872b), quoted parts of the original newspaper account but described, figured, and assigned the crayfish

to a new genus and species, Orconectes inermis, indicating that "The blind crawfish above mentioned is

specifically distinct from that of Mammoth Cave,

though nearly related to it .. . separating it generically from Cambarus, or the true crawfishes, on account of the absence of visual organs." He also assigned the albinistic pellucidus from Mammoth Cave

to his new genus. Cope (1872c) reiterated the presence of this crayfish in Wyandotte Cave.

Early in 1872, Packard (1872a) referred to Cope's

species as Cambarus pellucidus.

In the same year, Hagen (1872) ridiculed Cope's

assigning the two crayfishes to a new genus on the

basis of rudimentary eyes, and indicated that "the

description of the single specimen does not give any

character by which to separate it from the old species,

C. pellucidus."

Packard (1873) also relegated Cope's species to

synonymy with Cambarus pellucidus but Smith

(1873) utilized the combination O. pellucidus inermis

which in 1875 was apparently rejected in favor of

Cambarus pellucidus. Collett (1874) reported this

crayfish from Donaldson's and Hamer's caves, Lawrence County, Indiana. Shaler (1875) in describing

the cave fauna of Indiana mentioned the crayfish in

Wyandotte and other caves, and Collett (1879)

referred to those in Rhodes Cave, Harrison County,

Indiana. Cope (1879) published an account of this

species which is almost identical with those which appeared in 1872, but the crayfish is referred to as

Orconectes pellucidus, and the descriptions are

omitted.

Hovey (1882) utilized a new combination, Cambarus (Orconectes) pellucidus but contributed no

new data. Faxon (1884) included Cope's O. inermis

as a synonym of C. pellucidus.

Faxon (1885a) compared Cope's type, which was

a second form male, with specimens from Mammoth

Cave and he was of the opinion that this specimen

exhibited some of the variations of the typical form

of C. pellucidus. He commented on relationships and

included locality records from Indiana.

Underwood (1886) also treated Cope's species as

synonymous with C. pellucidus. Sloan (1888), in a

letter to Packard, reported eyeless crayfishes in caves

at Clifty which Packard indicated as being located

in Bartholomew County, Indiana. The location of

the caves is in error; actually they are 3.5 miles north

of Campbellsville, in Washington County. Packard

(1888) cited several new locality records for "Cambarus pellucidus," but in commenting on the food of

the crayfish (page 24) refers to it as O. pellucidus,

and in discussing the eye and vision there assigned

the species to both genera! He indicated that "the

food of the blind crayfish appears to consist of living

Caecidotea" [ — Asellus], an observation made by Mr.

Moses N. Elrod, on a crayfish taken from a well in

Orleans, Orange County, Indiana (page 24). In his

explanation of plate-27, figure 5 (page 156), he indicated that it was "after Cope" but it is obviously

the same figure utilized in Packard (1871) "after

Hagen" and is an illustration of O. pellucidus rather

than O. i. inermis.

Faxon (1890) summarized the Indiana localities

from which C. pellucidus had been reported, and

discussed the similarities between the species and

C. setosus and C. hamulatus. In the same year, Packard discussed the possible origin of C. pellucidus and

indicated that the "optic lobes and nerves" persist

but that there was a "total atrophy of the rods and

cones, retina (pigment) and facets." Packard (1890)

referred to the occurrence of C. pellucidus in Bradford Cave (also known as Stierstaedter Cave, located

0.5 mile east of Bradford on the east side of Corn

Creek), Harrison County, Indiana.

Hay (1891) indicated that Cambarus pellucidus

"occurs in many of the caves of southern Indiana.

They are usually small, the largest I have ever seen,

among 40 specimens, being barely 2 inches in length.

They are kept from extinction only by the inaccessibility of their home. They are very conspicuous when

in the water, and are very easily caught. When

startled they are utterly at loss where to go, and

often dart out upon the shore. I think it may be

safely said that as a rule they grow spinier as one

advances southward, although there are exceptions.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

A female collected in Wyandotte Cave is almost

without spines, but three specimens from a small cave

near there are exceedingly spiny."

Ortmann (1892) simply repeated known locality

records. Hay (1893) presented perhaps the best observations made on this species in describing his

experience in catching members of the intergrade

population in Shiloh Cave which he designated Cambarus pellucidus inermis. "When first observed, the

crayfish were generally, I might almost say always,

resting quietly in some shallow part of the stream

on one of the banks of clay. They lay with all their

legs extended and their long antennae gently waving

to and fro. Once or twice I saw them on the shore

a foot, at least, from the water, and one of these

appeared to have been digging in the soft mud.

When in the water I found it almost impossible to

catch them with the net, and after a few trials threw

it aside as useless. A much surer method was to

approach them slowly with the hand and then suddenly seize them. When once touched they started off

in great haste for some protecting rock, but often in

their alarm would dart out upon the banks where

they would lie unable to get back to the water. They

did not appear to be at all sensitive to the light. I

have often tried the experiment of slowly passing my

candle back and forth a few inches above them, or

of suddenly removing the light and then bringing it

close again, but with no effect whatever.

"Noise has no effect; a loud call or a shrill whistle

they do not notice. Nor does disturbing the waters

seem to affect them, and it is only when they are

touched that they manifest fear.

"It may also be stated that, as a rule, Cambarus

pellucidus is smoother the further north it occurs.

The material which I have collected myself, and all

that I have been able to obtain from others, will

uphold me in this statement.

"The small cave near Wyandotte produces individuals of exceeding spininess, it being the exception

to find there a comparatively smooth one.

"Coming further north, to Paoli, we can find much

smoother specimens, and at Shiloh they are smoother

still, while at Mayfield's Cave, in Monroe County,

occurs a form entirely without spines. So constant is

this feature of smoothness in the Mayfield Cave

specimens and so different is its appearance from

the typical pellucidus that I think it is worthy of

being characterized as a subspecies."

Hay also cited all the known localities in Indiana

in which troglobitic crayfishes had been found.

Lonnberg (1894, 1895) treats O. inermis as a

synonym of C. pellucidus and compares it with his

new troglobite from Florida, which is described in

the latter paper as Cambarus acherontis.

In his "Crawfishes of the State of Indiana," Hay

(1896) described and figured "C. pellucidus" indicating that "the form 'inermis' is now hardly recognized even as a variety of pellucidus." Again, he

summarized the range of the species in Indiana, citing

caves in Lawrence, Bartholomew, Orange, Crawford,

Harrison, and Jefferson counties.

Hay (1897) presented a synonomy of the species,

reviewed the localities from which the Indiana pellucidus had been collected, and Blatchley (1897:209)

included the following annotation: "The blind crayfish inhabits shallow pools with muddy bottom rather

than rapid flowing water. It moves slowly with its

antennae spread out before it, and gently waving to

and fro, feeling, as it were, every inch of its way.

It is wholly non-sensitive to light and seemingly so

to sound, but when disturbed by any movement in

the water it is extremely active, much more so than

ordinary terrestrial forms, leaping upward and backward with quick, powerful downward blows of its

abdomen." Blatchley (1897:171) compared in tabular form Cope's (1872a, 1872b) list of species occurring in Wyandotte Cave with his own.

Faxon (1898) commented that C. pellucidus in

Shiloh, Down's, and other caves in Lawrence County

and in a small cave near Paoli, Orange County,

Indiana, "belong rather to the form described by

Cope as C. inermis (Wyandotte Cave) than to the

typical form commonly found in the Mammoth Cave

of Kentucky." He also indicated that Hay has shown

a "transition" from typical pellucidus in southern

Indiana to the more northerly occurring form in

which there is a reduction of the spines.

In his "Synopsis of North American Invertebrates,"

Hay (1899) combined inermis and pellucidus under

the latter name in his key and indicated the range

to encompass Kentucky and Indiana. In a discussion

of the crustacean fauna of the Mammoth Cave region, he referred to the occurrence of C. pellucidus

in Indiana. Hay (1902a), in discussing the relationships of O. inermis testii, indicated that "this subspecies is found in a very small area in Indiana at

the very northern limits of the range of C. pellucidus."
NUMBER 10 5

Neither Ortmann (1902) nor Steele (1902) added

any additional information concerning their C. pellucidus, and Eigenman (1903) simply indicated that

pellucidus from caves near Mitchell, Indiana have

"an eye structure much more degenerate than specimens of the same species from Mammoth Cave."

Harris (1903b) summarized the habitat and range

of the species, combining, as had most of his predecessors, the Indiana and Kentucky "Cambarus pellucidus."

Ortmann (1905a, 1905b) discussed the affinities

of C. pellucidus (including the Indiana representatives) and indicated its closer relationship with crayfishes now assigned to Orconectes than to those belonging to the genus Procambarus. Osborn (1912)

added no original data.

Graeter (1909) briefly summarized the literature

on cave-dwelling species from 1900. Faxon (1914)

repeated (page 392) the fact that Hay and Banta

had found C. pellucidus associated with C. bartonii

laevis in caves in southern Indiana and included in

his check-list (page 415) O. inermis as a synonym

of C. pellucidus.

Spurgeon (1915) compared the eyes of C. setosus

and his C. pellucidus [ — 0. i. inermis and O. i. testit\

and found no important differences. He expressed the

opinion that the eyes of these troglobites represent

arrested development rather than degeneracy. He

also cited a new locality record (which is for the

intergrade population inermis x testii) and indicated

that the Clifty Caves are located in Washington

County, Indiana.

Ortmann (1918) mentioned the "blind cavespecies, C. pellucidus" from Kentucky and Indiana.

Pope (1926) included observations on individuals

in captivity and a photograph of a topotype.

Spandl (1926) simply referred to certain previous

works. Chappuis (1927) added nothing new but

again reported C. pellucidus from Indiana. Bolivar

and Jeannel (1931) and Fage (1931) indicated the

presence of this crayfish in "Donnelson's Cave"

[ = Donaldson Cave, Spring Mill State Park, Lawrence County, Indiana], and the former added Wyandotte Cave. Ortmann (1931) cited this crayfish within the range of "pellucidus."

Wolf (1934-1938) cited localities for Cambarus

pellucidus in Indiana together with several erroneous

records for "C. pellucidus var. testii"; these latter

records are for O. i. inermis and intergrades between

the two subspecies.

Hobbs (1942a), in revising the genus Cambarus,

reinstated Cope's generic name, Orconectes, expressed

the opinion that inermis was a valid species, and illustrated the first pleopod of the male. Hobbs (1942b)

recognized the subgenus Faxonella, thereby implying

that Orconectes inermis belonged to the typical subgenus.

Rhoades (1944) indicated that the range of O.

pellucidus pellucidus "with its subspecies" ranges

"from southern Indiana to northern Alabama," and

proposed the recognition of two groups of the Limosus

Section of the genus Orconectes.

Hobbs (1948a) illustrated the pleopods of topotypes of O. inermis along with those of its closest relatives, presented a key to the members of the Limosus

Section, and questioned the subdivision of the Section

as proposed by Rhoades. In addition, he indicated

that the status of O. inermis remained unsolved and

stated: "Perhaps it will be shown to be a subspecies

of O. pellucidus; however, until future work will indicate intergradation between the two, it seems advisable to retain its specific status" (page 19).

Pennak (1953) cited Indiana as being within the

range of O. pellucidus. Jackson (1953) presented

observations on this crayfish in Wyandotte Cave.

Jackson (1954, 1955) mentioned that blind crayfish and blind fish are plentiful in the stream in

Donaldson's Cave.

Balss (1955) erroneously reported "Cambarus

(Cambarus) pellucidus testii" from Wyandotte Cave.

Eberly (1955) cited a new locality record for O.

inermis, but relegated most of the Indiana locality

records to Cambarus pellucidus.

Holthuis (1956), in an application to the International Commission on Zoological nomenclature,

requested that the genus Orconectes be placed on the

"Official List of Generic Names in Zoology," the typespecies being given as O. inermis.

Crocker (1957) added no additional information

relative to the species, simply citing it as the typespecies of the genus.

The 1958 paper of Eberly is a mimeographed, almost identical version of Eberly (1960). Although

this study is centered around competition between

C. laevis and Orconectes inermis (probably an intergrade population), other pertinent data are presented, including the fact that C. laevis consumes 1.2
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

to 2.8 times the amount of oxygen per gram of body

weight as does the troglobite. Among them is an

expression of "the opinion that all of these cave

crayfish [subspecies of pellucidus and inermis] in the

Indiana-Kentucky-Alabama cave region are descended

from a once wide-spread surface population that was

relatively undifferentiated at that time. After entering the cave, the cave adaptations mentioned earlier

developed while certain features which are now used

in taxonomic determinations, e.g., the form of the

male genitalia, remained relatively unchanged." This

work is cited by Poulson (1964).

Hobbs (1959) designated the two subspecies of

Indiana troglobites as members of the subgenus Orconectes: Orconectes (Orconectes) inermis.

Rhoades (1959) reviewed the status of O. inermis

and gave an account of the events transpiring in

Cope's describing the species. He analyzed the descriptions of Cope's species and that of Hay's O. pellucidus testii, and concluded that Hay's subspecies is

synonymous with Orconectes inermis. He further

pointed out that "O. inermis intergrades freely with

O. pellucidus pellucidus (Tellkampf) (1844) in the

southern counties of Indiana." In view of this conclusion, he proposed the name, "Orconectes pellucidus inermis, for the smoother, less spinous subspecies

of south central Indiana." Of the affinities and distribution Rhoades stated that "There can be little

doubt of the affinity of the blind crayfish of Indiana

with those of the solution caverns of Kentucky. In

fact, the blind crayfishes from the caves of Harrison

and Crawford counties on the Ohio River are decidedly similar to the Mammoth Cave species with

only an occasional smooth spineless specimen. The

underground waters of Orange, Washington and

Lawrence counties a few miles to the north have

populations of blind crayfish which are truly transitional between the species and subspecies. Smoothness, spine reduction and other inermis characteristics

reach a climax in the cave populations in Monroe,

Brown, and Bartholomew counties on the northern

border of the range."

Nicholas (1960) included O. p. inermis in his

checklist.

Utilizing the observations of Eberly (1958, 1960),

Huheey (1961) proposed a model for the cyclical

evolutionary process in a subterranean habitat frequented by a troglobitic and a troglophilic crayfish.

He proposed that following the extinction of the

troglobite (A) [O. pellucidus = O. i. inermis], the

population of the troglophile (B) [C. bartonii = C.

laevis] will be greatly reduced because of a reduction

in food supply (A); B, in turn, will become better

adapted to cave life, thus coming to resemble more

closely the previously existing A; and "At some point

it will be possible for another epigean form (C),

although poorly cave-adapted, to compete successfully

with B and eventually replace it by virtue of its

greater robustness."

Hart and Hobbs (1961) cited O. p. inermis as

one of the hosts of Entocythere barri ( = Sagittocythere barri).

Creaser (1962) contended that the genus Orconectes should be composed of only three known

species: lancifer, inermis, and pellucidus "(with subspecies—if they are subspecies)," and refuted the

supposed relationships to other members of the genus

as was proposed by Hobbs (1942a).

Rhoades (1962) in his discussion of the evolution

of the Limosus Section of the genus Orconectes retained the Limosus and Rafinesquei groups he had

proposed earlier (1944), assigning the several subspecies of O. pellucidus to the latter group, but in

characterizing O. pellucidus pellucidus, he stated

(page 68) "Pleopods of the form I male short with

typically divergent tips," that is, possessing the sole

character (see page 69) on which he based his

"Group limosus." Unfortunately, the characters of

the two groups are reversed on page 75. Rhoades

postulated "that the genus originated on the northwest flank of the Mississippi embayment particularly

in the rocky streams of the Ozarkian Highlands."

From there the stock moved eastward from "the

Springfield Plateau of central Missouri through

southern Illinois to western Kentucky" to occupy

"the lower reaches of the Pliocene Teays and Ohio

rivers," with populations becoming "isolated into two

groups on the basis of: 1) occupation of the Pliocene

Teays River, the discharge of which poured into the

head of the Mississippi embayment, and 2) continued

migration along the escarpment streams to the Pliocene Ohio River." Rhoades concluded that "There

can be little doubt that the Group rafinesquei has

occupied the streams of the Cumberland Plateau since

very ancient times" and that the Pliocene surface

species perhaps "entered rock-fissure springs and lived

in subterranean water courses." Although he did not

hazard a guess in which part of the Plateau the in-
NUMBER 10 5 41

vasion took place, he stated that "It appears that a

specific population of Orconectes pellucidus was

widely distributed in the labyrinth of Cumberland

caves by the beginning of Pleistocene time. Perhaps

their wide distribution occurred in the late Pliocene

when cave corrosion was at its deepest stage and

all channels were free from filling. Isolation, which

was a factor in subspeciation of O. pellucidus, came

about as a function of river entrenchment and subsequent aggradation. . . . With glacial filling of the

[Ohio] river channel and sedimentation in the interrupted end of cave tunnels, especially in Illinoian

time, a barrier was formed which isolated a population north of the Ohio River" thus isolating the

ancestors of inermis. "Similar entrenchment and sedimentation in the upper Cumberland River could lead

to the separation of a segment of O. pellucidus in the

caves of southeastern Kentucky" isolating the ancestors of packardi. "The blind Orconectes in the

subterranean channels of the Mussel Shoals region

were similarly isolated . . ." Fitzpatrick (1963) only

mentioned pellucidus in discussing Creaser's concept

of generic assignments.

Penn and Fitzpatrick (1963) and Vandel (1964)

referred to Eberly's work, and the latter designated

"Orconectes pellucidus [ = O. i. inermis, in part] "un

cavernicole ancien." Minckley (1963) reported this

species as O. p. pellucidus from Doe Run, Meade

County, Kentucky.

Jegla (1964), in studying an integrade population

of inermis x testii in Shiloh Cave, found a preponderance of first-form males in the fall and the fewest

numbers in the summer months; females apparently

lay their eggs during the spring and summer months,

and young were found as early as the "first part of

May." The adult males undergo two major molting

periods each year, during the fall and late winter to

spring, the former increasing the number of first-form

males and the latter, the number of those in the

second form. The same author (1965a) reported that

although the eyes of this crayfish have lost, through

degeneration, some of the optic elements, "the sinus

gland is present and neurosecretory cell groups are

located on the surface of the medulla terminalis

neuropile in approximately the same position as they

are in a normal-eyed crayfish." Differences noted in

time-lapse of molting following eyestalk ablation were

thought perhaps to be due to "a difference in concentration of molt-inhibiting hormone in the blood

of crayfish" at the time the eyestalks were removed.

Jegla et al. (1965) in comparing interpopulation

variations in cave crayfishes found that in selected

caves three subspecies of "O. pellucidus" (inermis,

pellucidus, and australis) differ by 20 mm in maximum length attained, by 20 mm in length at maturity.

They concluded that interpopulation size differences

are not due to basic differences in physiology but

probably to differences in available food and perhaps

"interspecific competition resulting in character displacement." Jegla (1965b), in an abstract of his

doctoral dissertation, summarized part of the above

and the following citations by him. The combination,

Orconectes pellacious, was also introduced inadvertently.

Jegla (1966) enlarged upon and detailed the evidence supporting his abstract published in 1964.

Barr (1966) referred to Jegla's ecological studies

"on the crayfish O. i. testii."

Poulson and Jegla (1966) published an abstract

concerning the reproductive cycle and the circadian

rhythm of oxygen consumption in this crayfish.

Hart and Hart (1966), primarily interested in the

entocytherid ostracods occurring on crayfish hosts,

cited several new locality records for O. i. inermis.

Mohr and Poulson (1966) presented excellent

photographs of this crayfish, comparing it with epigean species, and recounted seeing it in "BronsonDonaldson Cave." They outlined the reproductive

cycle in Shiloh Cave, associating it with "biological

clocks."

Barr (1967a) introduced the combination Orconectes inermis inermis, anticipating that this manuscript would appear in print prior to his ecological

summary, and indicated that this subspecies "replaces O. pellucidus immediately north of Mammoth

Cave in the Pennyroyal (Hobbs and Barr, in preparation) . ... " In discussing the Pennyroyal, Barr

(1967b) pointed out that the extent of this plateau

limits the range of many of its endogenous cavernicoles, including O. inermis.

Fitzpatrick (1967) discussed the supraspecific

structure of the Cambarinae that was proposed by

Creaser, and, in so doing, referred to O. inermis.

Barr (1968) cited data from Eberly and Jegla, and

discussed the possible origin of the troglobitic members of the genus Orconectes.

Jegla and Poulson (1968), investigating circadian

rhythms in O. i. inermis from McCubbin's Cave,
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Breckenridge County, Kentucky, and O. inermis x

testii from Shiloh Cave, Lawrence County, Indiana,

reported that mechanisms for circadian rhythms have

apparently persisted in some of these crayfishes despite the period of time they have been restricted to

an environment in which daily light cycles do not

exist. The minimum and maximum oxygen consumption of the crayfishes were approximately 0.015 and

0.085 ml/g/hour. Their discussion of the observations

of Park et al. (1941) and the later analysis by Brown

(1961) applies to O. pellucidus rather than to O.

inermis.

Prins (1968) recorded the presence of this crayfish

in a surface stream, Doe Run, Meade County, Kentucky, and indicated that individuals are found in

the stream frequently, "especially during high discharges" from the headwater spring.

Jegla (1969) continued his study of the intergrade

population in Shiloh Cave and found that whereas

molting occurs throughout the year, two periods of

approximately two-months duration constitute the

maximum molting periods, February-March and

August-September. The adult male population consists of a preponderance of those in form II in the

summer months (60 to 70 percent) and those in

form I during the winter months (77 to 97 percent).

An annual ovarian cycle was observed in the females,

beginning during the early fall, with oviposition occurring during the summer. Although the incubation

period is not precisely known, Jegla estimated a duration of approximately two months. Young crayfish

are found throughout the year but are most abundant

in the summer and autumn, and "most conspicuous"

in August.

M. Cooper (1969) compared the length of time

required "for detecting, actively seeking, and locating

food stimuli" by this crayfish with that of the epigean

O. limosus. She found that O. i. inermis detected a

moving worm much more quickly than it did introduced glycine, and whereas the epigean species sensed

the presence of the latter more quickly, once the

troglobite "started seeking the chemical, it located

the source in much less time than did O. limosus."

The time required for the capture of the worm by

the troglobite was one-half that utilized by the epigean

crayfish.

Poulson and White (1969), in their discussion of

cave environments, pointed out the lack of morphological differentiation in O. inermis on either side

of the Ohio River, and indicated that "the troglophilic crayfish Cambarus bartoni [ = C. laevis] is much

rarer in the deep cave when the troglobitic Orconectes is present."

Hart and Hart (1969) expressed the opinion that

either this crayfish or Cambarus tenebrosus [actually

C. laevis] served as a host to Entocythere donnaldsonensis [ = Donnaldsoncythere donnaldsonensis].

Jegla and Poulson (1970) found a circannian

rhythm with respect to reproduction in O. inermis,

with periodicity of 338 to 396 days. They hypothesized

that "some event associated with heavy run-off from

surface precipitation triggers egg-laying and synchronizes the individuals' circannian rhythms in the cave."

COMMENTS ON PREVIOUSLY RECORDED DATA.—On

the basis of information available to Hagen (1872),

his criticism of Cope's (1872a) designating Orconectes inermis as a species distinct from Tellkampf's

Astacus pellucidus was entirely justified. Not until

Hobbs (1948a) made comparisons of the first pleopods of the males of the two did it become questionable that the two were, in all probability, conspecific.

Through Cope's folly of basing his genus Orconectes

on the eyeless condition of his new species, he established a generic name, which although largely relegated to synonomy for 70 years, had to be resurrected in 1942 when Hobbs revised the genus Cambarus. Most of the references within the 70 year span

consist of the addition of new locality records and

brief statements indicating the belief that Cope's

species was a synonym of pellucidus. Outstanding,

however, were some of Hay's observations on the

habits and reactions of the intergrade population

(inermis x testii) in Shiloh Cave, Lawrence County,

Indiana. The observations of Eberly (1958, 1960) are

noteworthy, particularly the conclusions concerning

cave adaptations and competition between this species

and the pigmented Cambarus laevis Faxon.

Rhoades' (1959) belief that O. inermis intergrades

freely with O. p. pellucidus does not coincide with

our interpretations. His discussion of reasons for recognizing testii as a synonym of inermis was not without

merit, in view of the fact that the males of the Kentucky segment of the latter (which Rhoades presumably considered to be O. pellucidus) possess pleopods

typical of inermis rather than being intermediate between inermis and pellucidus. We believe it appropriate, however, to recognize inermis and pellucidus as

distinct species and to retain the subspecific designa-
NUMBER 105 43

tion O. inermis testii for those aspinous populations

occurring in the northernmost part of the range. The

only reservation we have in suggesting this treatment

is based on the observations of Banta (1907:70) who

stated that "some individuals from Mayfield's [Cave,

Monroe County, Indiana] show some tendency toward rostral spines, while a single specimen taken in

Mayfield's was fully as spiny as the average individual

from Mitchell." We have seen no spiny individuals

among the material available to us from Monroe

County (see under O. i. testii), and our specimens

from Donaldson's Cave [ = Banta's Cave at Mitchell]

include moderately spiny to almost aspinous individuals.

The hypotheses of Rhoades (1962) relative to the

evolutionary history of these troglobites are not in

complete agreement with those outlined above; however, they are most worthy of consideration.

Among recent studies, the contributions to our

knowledge of the biology of the species by Jegla

(1964-1969) are outstanding. Despite the many more

references in the literature to O. pellucid us than to

O. inermis, much more is known of the biology of

the latter than that of O. pellucidus. Chiefly through

the work of Jegla and Hay, more data have been

accumulated on the intergrade population in Shiloh

Cave than for any other troglobitic crayfish.

Barr (1967a) introduced the combination Orconectes inermis inermis, and his hypotheses concerning

the origins of the troglobitic members of the genus

Orconectes (1968) are essentially those which are

detailed elsewhere in this study.

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum with marginal spines, tubercles, or

at least angular emargination at base of acumen;

margins convergent, upper surface without median

carina; postorbital ridges terminating cephalically

in spines or tubercles; hepatic area often with two

to many spines; at least one, often several, cervical

spines present; areola 4.5 to 6.7 times longer than

broad and constituting 37.0 to 42.5 percent of entire

length of carapace; chelae not conspicuously setose

but with ciliated tubercles, mesiodorsal surface of

palm with several irregular rows of tubercles; hooks

on ischiopodites of third and fourth pereiopods. First

pleopod of first-form male with greatest cephalocaudal

diameter of pleopod less than twice that immediately

proximal to base of central projection, always terminating in only two elements; non-corneous mesial

process broad basally, suddenly contracting to form

tapering subacute distal portion, latter directed caudodistally and slightly exceeding tip of central projection

distally; central projection corneous, flattened in

cephalocaudal plane, slightly concavoconvex with

convexity extending mesially; shoulder lacking at base

of central projection but cephalic surface with convexity along distal third of appendage. Annulus ventralis is illustrated in Figure 10/.

Topotypic Male (Form I) : Body (Figure 10c, h)

subovate, depressed. Abdomen narrower than thorax

(8.5 and 10.3 mm in widest parts, respectively).

Width of carapace greater than depth in region of

caudodorsal margin of cervical groove (10.3 and 7.2

mm). Areola moderately broad (5.1 times longer than

wide) with 5 or 6 minute punctations across narrowest part. Cephalic section of carapace 1.5 times longer

than areola; length of areola 40 percent of entire

length of carapace. Rostrum slightly longer than twice

width at base, excavate, and with acumen about 1.2

times longer than maximum width; cephalic extremity reaching beyond antennular peduncle and almost

as far as distal end of peduncle of antenna; margins

not swollen, little elevated, and with pair of welldeveloped, corneous-tipped marginal spines at base

of acumen; upper surface with regularly spaced,

minute, setiferous punctations; subrostral ridges moderately well developed and evident in dorsal aspect

along basal third of rostrum.

Postorbital ridges comparatively low, short, with

dorsolateral grooves, and terminating cephalically in

strong, corneous-tipped, acute spines. Suborbital angle

lacking. Branchiostegal spines acute and well developed. Cervical spines represented by row of 3 or 4

tubercles on each side of carapace immediately caudal

to cervical groove; tubercles progressively larger ventrally, and ventral two spiniform. Carapace punctate

dorsally and granulate laterally; hepatic area with

three or four small spiniform tubercles on each side.

Abdomen longer than carapace (25.5 and 23.0 mm).

Cephalic section of telson with 2 strong spines in

sinistral caudolateral corner (mesial one movable)

and 1 in dextral.

Protruding portion of epistome (Figure 10?)

almost twice as broad as long and resembling silhouette of shortened minaret; surface flattened, with

fine setae, and with fovea present caudally. Eyes much

reduced, without pigment, completely hidden beneath

rostrum in dorsal aspect, and extending cephalically
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Orconectes inermis inermis: a, mesial view of first pleopod of topotypic male, form

I; by mesial view of first pleopod of topotypic male, form II; c, dorsal view of carapace of topotypic male, form I; d, lateral view of first pleopod of topotypic male, form II; e, lateral view

of first pleopod of topotypic male, form I; /, dorsal view of distal podomeres of cheliped of

topotypic male, form I; g, epistome of topotypic male, form I; h, lateral view of carapace of

topotypic male, form I; i, antennal scale of topotypic male, form I; ;', caudal view of first pleopods of topotypic male, form I; k, ventral view of caudal thoracic region of topotypic male,

form I; I, annulus ventralis and portion of sternum of topotypic female.

about half way between margin of orbit and marginal

spines of rostrum. Antennules of usual form with

strong spine on ventral surface of proximal podomere

at base of distal fifth. Antennae extending caudally

slightly beyond caudal margin of telson. Antennal

scale (Figure lOt) broadest slightly distal to midlength, distinctly less than half as broad as long; outer

thickened portion much narrower than lamellar area
NUMBER 105 45

and terminating distally in very prominent corneoustipped spine. Third maxillipeds extending slightly

beyond midlength of distal podomere of peduncle of

antenna.

Chela (Figure 10/) slender and little inflated;

mesial margin of palm 1.1 times longer than width

of palm; dorsal surface of palmar area tuberculate

mesially and punctate laterally, punctations and distal

bases of tubercles setose; ventral surface of palm tuberculate and with corneous-tipped spine opposite

base of dactyl; more mesial tubercles larger and

arranged sublinearly, with 13 or 14 comprising most

mesial row of palm; lateral surface tuberculate. Fingers not gaping; dorsal and ventral surfaces with

broad, rounded, submedian, longitudinal ridges

flanked by setiferous punctations; opposable margin

of immovable finger with row of 7 tubercles, fifth

from base largest, along proximal two-fifths, single

tubercle below level of row slightly proximal to midlength of finger, and single row of minute denticles

between proximal five tubercles; distally, crowded

denticles forming slightly broader series to base of corneous tip of finger; lateral surface of immovable finger

subcostate. Opposable margin of dactyl with row of

5 tubercles along proximal two-fifths, distalmost largest, with denticles arranged as on immovable finger;

mesial surface of dactyl with small tubercles, decreasing in size distally, along proximal half and punctations along distal half. Carpus longer than broad,

with mesial, dorsomesial, and ventromesial surfaces

tuberculate, otherwise punctate; mesial surface with

3 tubercles somewhat larger than others, largest spiniform; distoventral margin with 2 spines, 1 adjacent

to base of ventrolateral articulation with propodus

and 1 near median line. Merus tuberculogranulate

dorsally, with group of 3 corneous-tipped spines

slightly proximal to distal end; except for tubercles,

weakly scabrous; ventrolateral margin with row of 9

spikelike tubercles and ventromesial margin with 11;

few scattered tubercles flanking two rows; distolateral

extremity with corneous-tipped spine. Ischium with

ventromesial row of 5 small tubercles and additional

smaller ones laterally; dorsal margin with subserrate

row of 7 or 8 small tubercles.

Ischia of third and fourth pereiopods (Figure 10k)

with hooks; those on third stout and recurved to

level of cephalic articulation with basis; those on

fourth much smaller and, while slightly recurved,

leaving broad gape between tip and corresponding

basis. Coxae of fourth pereiopods with caudomesially

projecting prominences; coxae of fifth pereiopods

without prominences except for small mesioventral

projection partially ringing base of phallic papilla.

First pleopods (Figure 10a, e, j , k), symmetrical,

reaching bases of third pereiopods when abdomen is

flexed, and comparatively shallowly situated in sternal

groove; tip ending in two parts as described in

Diagnosis.

Topotypic Female: Differs from the male in the

following respects: subrostral ridges evident almost

to base of marginal spines; 4 cervical spines on right

side and 2 on left; hepatic areas with 4 spines on right

and 5 on left; antennal scale broadest slightly proximal to midlength; third maxillipeds extending to

distal end of antennal peduncle; ventral surfaces of

both chelae with crowded small tubercles, right with

prominent median tubercle near midlength; immovable finger of chela with row of 14 tubercles along

mesial four-fifths of finger (sixth from base largest)

and prominent tubercle below tenth tubercle from

base; opposable margin of dactyl of chela with row

of 19 tubercles along proximal four-fifths (sixth from

base largest) ; carpus of chelipeds tuberculate with

single prominent spine on mesial surface of right and

row of 3 on left, latter tubercles increasing in size

distally, ventral surface with 3 prominent spines,

1 adjacent to base of ventrolateral articulation with

propodus and 2 mesial to it; merus of cheliped with

mesial row of 14 tubercles and lateral row of 7 on

right and 9 on left; ischia of chelipeds with row

of 7 small tubercles on right and 5 on left. (See

Measurements.)

Annulus ventralis (Figure 10J) shallowly situated

in sternum and not firmly fused to sternal plate immediately cephalic to it; outline subovate, 1.5 times

broader than long with median portion elevated

ventrally and forming asymmetrical arc with highest

segment caudal to midlength; arched portion bearing

shallow longitudinal groove ending caudally on steep

slope of arc; sinus originating on slope dextral to

median line, making gentle arc to median line and

extending caudally to midcaudal margin of annulus.

Topotypic Male, Form II: Differing only slightly

from first-form male: 3 cervical spines on each side

of carapace; telson with 2 spines in each caudolateral

corner of cephalic section; right antennal scale with

strong accessory spine on middle third of lateral margin; both chelae regenerated, left larger than right,
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

but neither well developed; hooks on ischia of third

and fourth pereiopods comparatively more reduced,

but boss on coxa of fourth only slightly smaller. (See

measurements.)

First pleopod (Figure 10b, d) reaching base of

third pereiopod when abdomen is flexed and markedly resembling pleopod of first-form male, except

central projection much shorter, more swollen, and

noncorneous.

MEASUREMENT S (in millimeters).—Orconectes

inermis inermis, topotypes:

Male Male

Form I Female Form II

Carapace:

Height 7.2 11.0 8.0

Width 10.3 14.4 10.6

Length 23.0 34.3 25.5

Rostrum:

Width 3.5 4.8 3.5

Length 6.0 9.1 7.0

Areola:

Width 1.8 2.5 2.1

Length 9.2 14.0 10.4

Chela: (no chela)

Length of inner margin

of palm 7.5 11.0

Width of palm 6.6 7.5

Length of outer margin

of chela 19.3 29.1

Length of dactyl 10.6 16.6

TYPES.—Not extant.

TYPE-LOCALITY.—Wyandotte Cave, Indiana. Actually Sibert's Well Cave, just below Wyandotte Cave,

Crawford County, Indiana (Hobbs, 1942a: 335).

SPECIMENS EXAMINED.—Specimens from Kentucky

and Indiana were examined as follows (those localities considered to be frequented by weakly spined

testii-like populations are preceded by an asterisk) :

KENTUCKY—Breckenridge County: (1) Cave and stream,

5 miles W of Big Spring, 6 $ II, 5 9 , W. L. Minckley, VI/

5/58. (2) Bandy Cave, 3.0 miles S of Irvington, 131,

8 3 II, 6 $ , T.C.B., IX/20/64. (3) Thornhill Cave, 3 miles

W of Big Spring, 7*1 , 13II, 5$ , T.C.B., R. McAdams,

X/18/64. (4) Lockard Cave, 3 miles SE of Bewleyville,

3 31 , 33II, 7 9, T.C.B., IX/20/64. (5) Bat Cave [location not determined], 19 , Ij9 , J. Rhines, 1961. Greene

County: (1) Brush Creek Cave, 0.8 mile E of Lobb, 4 3 II,

T.C.B., IX/28/63. (2) Scott Cave, 1.9 miles ESE of Eve,

Ij3 , T.C.B., IV/2/66; 13II, 19 , T.C.B. and T. C. BanIll, XI/1/70 . Hardin County: (1) Bland Cave, 7 miles

W of Sonora, 131, 8 311, 13 9, 1J3, T.C.B., R.A.K., and

R. Taylor, XII/8/62 ; 2 31, 19 , Hugh Thomas, XII/27/62.

(2) Nelson Cave, 1.5 miles W of Star Mills, 1 3 I, 1 3 II, 49 ,

Ij9 , T.C.B. and R.A.K., XI/2/63 . Hart County: (1)

Turner Cave, 3.7 miles SE of Magnolia, 1 3 H, T.C.B., VIII/

25/63. (2) Cooch Webb Cave, 0.4 miles N of Bear Wallow, 2 31, 13II, 9 9, 1J3, 2j9 , T.C.B. and R.A.K., XI/

2/63. (3) Riders Mill Cave, 2.5 miles N of Priceville, 6 31 ,

13II, 2 9 , T.C.B. and R.M.N., X/5/63. (4) Cub Run

Cave, near town of Cub Run, 2 9, T.C.B. and W.M.A.,

XI/28/64; 19 , L.H., 11/29/56. Meade County: (1) *Joe

Jones Cave, near lower reaches of Doe Run, 131. 3 9 ,

W.L.M., VII/16/60; 1J3, 1J9, J.R-, VII/25/61; 2j9 ,

R. Prins, III/2/63 ; 19 , Rudolph Prins, 111/16/63. (2)

*Rockhaven Cave in Otter Creek State Park, 2 3H . 3 9 ,

T.C.B. and R.A.K., XI1/2/61. (3) »Shackletts Cave, SW

of Garrett, 29 , T.C.B. and R.A.K., XII/2/61. (4) *Lime

Kiln Cave, 1.35 mi. NW of Battletown, T.C.B. and W.M.A.,

33II, 2 9, VI/29/65.

INDIANA: Crawford County: (1) Archibald Cave, SE 54,

NE 54, SE 54, sec. T.3S, R.1E, 1 3II, R.M.N., III/8/64.

(2) Crawfish Spring, Wyandotte Cave, NW J4, SW }4,

NW 54, sec. 27, T.3S, R.2E, 1 9 , W. P. Hay, date ?. (3)

Stream in Sibert's Well Cave, NE 54, SE 54, sec. 28, T.3S,

R.2E, 131, 1311, 19 , L.H., IX/1/39. (4) Wyandotte

Cave, "From Mr. Palmer's room," 1 3 H, coll. and date?. (5)

Small cave near Wyandotte [-Sibert's Well Cave ?], Ij9 ,

W.P.H., date ?. (6) *Marengo Cave, SW «/4, NE 54, NW

54, sec. 31, T.3S, R.1E, Ij9 , R.R, date ?. Harrison County:

(1) •Bradford Cave, SE '/i, SE / 4 , SE 54, sec. 3, T.2S,

R.4E, 131, T.C.B., VIII/17/57. (2) »Small cave, 4 miles

NE of Mauckport, 131, 19 , Sherman Minton, date?. (3)

King's Cave, NE 54, SE ]/4, SW 54, sec. 34, T.3S, R.4E,

13II, 19 , T.C.B., VIII/17/57; 131, L.H., VIII/17/57.

Lawrence County: (1) *Blue Spring Cave, SW J4, SE 54,

SW 54, sec. 6, T.4N, R.1W, 2 31, 13II, H.H.H. Ill, X /

25/69; 131, H.H.H. Ill, VII/7/70. (2) *Shiloh Cave,

NW 54, SE 54, NW 54, sec. 18, T.5N, R.1W, 13H, 3 9 ,

W.P.H., date?; 13II, 29 , W.P.H., date?; 83H , 9$ ,

W.P.H., date ?; 131, 19 , T.C.B., 1/7/56; 19 , Thomas C.

Jegla, 1/27/59; 19 , T.C.J., 1/29/59; 131 , 19 , T.C.J.,

IX/26/59; 1 3 II, 19 , T.C.J., VI/4/60 ; 19 , T.C.J., XII/

10/60; 131, 13II, T.C.J., IX/23/61; 2 9 , T.C.J., XI/

19/61; 13H, T.C.J., 111/12/62; 19 , T.C.J., VI/30/62 ;

131, T.C.J., XI/17/62; 131 , 2 3II, T.C.J., 11/24/63;

2 3II, T.C.J., 111/31/63; 13II, T.C.J., IV/20/63 ; 131 ,

Ij3 , T.C.J., V/?/63 ; 13I.T.C.J., VI/20/63 ; 131, 1 3 II,

T.C.J., VI/30/63. (3) *Sullivan Cave, NW 54, SW 54,

SE 54, sec. 20, T.6N, R.2W, 131, T.C.J., XII/13/59 ;

1 3 II, H.H.H. Ill, 1/23/70. (4) •Donaldson's Cave, NW

54, NE 54, NE 54, sec. 4, T.3N, R.1E, 131 , 19 with eggs,

F. N. Blanchard, date?. (5) *Stream flowing from Donaldson's Cave, 1 3II, C. W. Hart and Dabney G. Hart,

VI/26/64. (6) • Wagoner Cave, SW 54, NE 54, NE

54, sec. 19, T.5N, R.2W, 19 , H.H.H. Ill, 11/21/70. (7)

•Harrison Cave, NW 54, NE 54, SW 54, sec. 15, T.3N,

R.1W, 19 , H.H.H. Ill, V/2/70. (8) *Pless Cave, SE 54,

NE 54, SE 54, sec. 5, T.4N, R.1W, 131, H.H.H. Ill, X /

4/70. Orange County: (1) »Murray Spring Cave, SE 54,

NW 54, NW 54, sec. 6, T.1N, R.1E, 1 3II, H.H.H. Ill,

H/28/70. Washington County: Series from each of the
NUMBER 105 47

two Clifty Caves—(1) "River Cave, NE

 lA, SE lA, SE

•4, sec. 14, T.3N, R.2E, 1$, J.R.R., VI/3/65. (2) •Endless Cavern, NE }4, SE J4, SE y4, sec. 14, T.3N, R.2E,

131, 13 H, 1$ , H.H.H. Ill, 11/28/70; 1 3 I, H.H.H. Ill,

VI/28/70. (3) »Fredericksburg Cave, SE J/4, NW J4. SE

!4, sec. 4, T.1S, R.3E, 231 , 1$ , H.H.H. Ill, IX/27/70.

RANGE.—Orconectes inermis inermis ranges from

Green County, Kentucky, northward into Crawford

County, Indiana, intergrading with O. i. inermis

northward to Monroe County, Indiana.

VARIATIONS.—Orconectes inermis is, with little

doubt, the most highly variable of the four troglobitic

members of the genus. Not only are there variations

associated with restricted portions of its range but

also the variations that occur within a single locality

far exceed those that might be anticipated. The most

conspicuous of the variations noted are those of the

armature of the carapace and chelipeds, and there is

no evidence of sexual dimorphism in the relative

numbers or degrees of development of the spines and

tubercles. In most other respects, the variations within

a single population are neither greater nor less than

those which occur in most epigean and other troglobitic crayfishes. Generally, there is a marked direct

correlation between the relative development of spines

and tubercles on the carapac and chelipeds, and

although there are exceptions to the generalization

that the southern members of the species have more

and larger spines than do the northern members, it

is apparently true that none of the Indiana forms

approach in their armature that of the most spiny

individuals from Kentucky. By selecting individuals,

one could demonstrate an almost uninterrupted dine

between the very spiny individuals from Hart and

Hardin counties, Kentucky, and the almost spineless

members from Monroe County, Indiana. Although

inadequate series of specimens from most portions of

the range of the species prevent our being able to

make meaningful ratios of the numbers of spiny to

aspinous members occurring in the various populations, it seems clear that, whereas in the extreme

north there are no spiny individuals, in many if not

in most of the southern localities the spiny individuals

far outnumber the aspinous ones. In the area from

Lawrence County, Indiana, southward to Meade and

Breckenridge counties, Kentucky, there is more nearly

a balance between the two, with the most conspicuous

breaks occurring between the populations of Monroe

and Lawrence counties and between those of Meade

and Breckenridge counties.

To illustrate the limits of variation in the armature

of the carapace among the specimens available to us,

a series of camera lucida sketches of individuals from

throughout the range of the species are presented in

Figures 12 and 13.

Four localities are represented in our collections

from Hart County, Kentucky. In both Riders Mill

Cave (Figure 13c, d) and Cooch Webb Cave (Figure

13^, /, g) there are considerable variations in the

development of the marginal spines of the rostrum,

the postorbital, cervical, and hepatic spines. Most individuals have moderately prominent spines and approach the maximum development shown in Figure

13c, but fewer numbers more nearly resemble the

individual depicted in Figure 13a*. The two specimens from Cub Run Cave (Figure 13a, b) are vastly

less ornamented, and that from Turner Cave has,

at least, as poorly developed spines as the smoother

specimen from Cub Run Cave. Among the specimens

from Hart County, the areola ranges from 37.4 to

41.5 (average 39.2) percent of the entire length of

the carapace and is from 4.4 to 6.7 times longer

than wide.

Strongly developed spines are characteristic of the

populations in the two caves in Hardin County (Figures \2y, z and 13i), and among them are the most

prominently armored members of the species. Even

the least spiny individual has well-developed cervical,

postorbital, and rostral spines together with two

hepatic spines on each side of the carapace. The development of the long acumen seems to be responsible

for the apparently short areola which constitutes 38.4

to 40.7 (average 39.3) percent of the length of the

carapace and is 4.7 to 6.7 times longer than wide.

In Breckenridge County, the population in Lockard

Cave (Figure \2v, w) has the most spines. In

Thornhill Cave (Figure 12*, u) and the cave near

Big Spring (Figure \2q), the range of variation is

greatest. Although the crayfish in Bandy Cave (Figure

12r, x) are variable, most have weakly developed

spines, and in some even the marginal spines on the

rostrum are more tuberculiform than spiniform. In

Bat Cave the animals have an almost smooth carapace

with spines no more strongly developed than the least

spiny of those from Bandy Cave. In Breckenridge

County, the areolae constitute from 37.0 to 42.4

(average 40.0) percent of the carapace length and

range from 4.7 to 6.3 times longer than broad.
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The populations from Meade County (Figure 12n,

o, p) may be categorized as almost aspinous; not

even the single small cervical spines and marginal

spines on the rostrum are consistently acute, and none

exhibits a single moderately well-developed hepatic

spine. Nevertheless, there is always at least a tubercle

at the base of the acumen. The areolae range from

39.0 to 42.4 (average 40.8) percent of the carapace

length and is 4.7 to 6.4 times longer than broad.

Across the Ohio River in Crawford County, Indiana, variation is much more pronounced than in

Meade County, Kentucky. Those specimens from

Sibert's Well Cave (Figure 10c, h), Archibald Cave

(Figure \2g), and the small cave near Wyandotte

all have moderately well-developed rostral and as

many as three cervical spines, and the Sibert's Well

specimens all have hepatic spines as well. In sharp

contrast, the specimens from Marengo Cave (Figure

I2h) and one of those from Blue Spring Cave not

only lack rostral spines or tubercles but also possess

an areola constituting more than 44 percent of the

length of the carapace, thus possessing the characteristics of O. i. testii. The other specimen from Blue

Spring Cave, that from Crawfish Spring, and one

from Wyandotte Cave are somewhat intermediate

between these two types and their areolae constitute

41.9 to 42.3 percent of the carapace length. The

areolae of the Crawford County specimens range from

38.0 to 44.5 (average 41.3) percent of the length of

the carapace and is 5.1 to 6.6 times longer than wide.

Our single specimen from Orange County agrees

rather well with those from Sibert's Well Cave; its

areola constitutes 41.5 percent of the carapace length

and is 6.8 times longer than broad.

Among our four specimens from Harrison County,

the two from Kings Cave (Figure 12/) have small

rostral spines and cervical spines, but those from the

other two localities (Figure 12m) are devoid of

rostral spines, and that from the small cave northeast

of Mauckport lacks even a cervical spine. None of

the Harrison County specimens have hepatic spines.

The areola of these specimens ranges from 42.2 to

43.6 (average 43.0) percent of the carapace length

and from 6.3 to 7.1 times longer than broad.

The four specimens from Washington County (Figure \2k) exhibit almost as much variation as those

from Shiloh Cave in Lawrence County (see below).

All have small cervical spines; the rostrum of one

specimen lacks marginal spines, that of two has weak

tubercles, one has small spines, and their areolae

constitute 41.2 to 41.8 percent of the carapace

length and are 5.3 to 7.1 times longer than broad.

The much larger series from Lawrence County,

mostly from Shiloh Cave, is quite variable, but not

nearly to the extent of those from Crawford County.

The extremes of variation are depicted in Figure

I2c-e in which it may be seen that all specimens bear

small cervical tubercles or spines, and the rostral margins frequently bear short spines or tubercles, and

most are angular at the base of the acumen. Some,

however, have rostra with uninterrupted margins, in

this respect resembling typical O. i. testii. Only one

specimen (Figure 12/) from Sullivan Cave has moderately strong cervical and hepatic spines. Nevertheless, only one of the specimens measured has an

areola constituting less than 40 percent of the carapace length, the range of which is 39.2 to 44.0 (average 42.3) percent and its length 4.3 to 6.5 times its

width.

All the specimens from Monroe County (Figure

12a, b) consistently lack rostral and hepatic spines,

and the cervical spines are reduced to spiniform or

blunt tubercles of which generally there is but one.

The areola constitutes from 42.3 to 45.9 (average

44.6) percent of the length of the carapace and

ranges from 4.9 to 6.4 times as long as broad.

Were not the Monroe specimens so uniformly less

spinous than those forms in the more southern localities, and did they not possess areolae which are generally distinctly proportionately longer than those of

almost all of their close relatives that range to the

south, we would be inclined to agree with Rhoades

(1962) and treat Hay's Cambarus pellucidus testii

as a synonym of Orconectes inermis. There is the

further disqueting fact that the type-locality of O.

inermis is situated near the middle of the range of

the species. In spite of this fact, it seems more desirable to emphasize the uniform divergence of the

Monroe County populations from the more southern

ones; therefore, we recognize them as constituting

the subspecies O. inermis testii. In summary, we view

the specimens from Sibert's Well Cave and the populations from Breckenridge, Hardin, and Hart counties,

Kentucky, as typical O. i. inermis, those from Monroe

County, as typical O. i. testii, and those from the

intervening area as intergrade populations.

That an occasional specimen of what might appear

to be typical of one of the two subspecies should be
NUMBE R 10 5 49

found within the range of intergradation may, perhaps, cause some concern, but if there is a gene flow

between the two extremities of the range of the

species, then the presence of such individuals in the

area might well be anticipated. With respect to the

development of the spines on the carapace and chelipeds, if it is under the control of some genetic mechanism rather than an ecophenotypic effect, then the

flow from north to south seems to have been greater

than that in the reverse direction.

Such an hypothesis would help to explain the

apparent semi-isolation of the more spiny individuals

in Sibert's Well and Archibald caves (perhaps also

the population in Murray Spring from which we

have only one specimen). If it could be demonstrated

that they represent small isolated populations off the

mainstream of gene flow, then one might postulate

that, in a sense, in them are preserved the primitive

spiny characteristic which is gradually being replaced

by the more advanced aspinous one in the populations along the path of the mainstream of gene flow.

As has been pointed out above, it is our belief that

O. i. inermis is distinctly more primitive than O. i.

testii and that the invasion of the Indiana caves was

from an inermis stock moving northward in subterranean channels from Kentucky. Somewhere in its

trek across southern Indiana, a segment of the stock

lost its spines and came to dominate the northernmost

portion of the range. That the innovation was a

successful one can hardly be questioned, and our

materials indicate that the gene, or combinations of

genes, responsible for the reduced spination is now

moving in a southerly direction, invading the ancestral range of the species.

SIZE.—The largest specimen among those that we

have examined is a female having a carapace length

of 36.9 mm, and was collected in Bat Cave, Breckenridge County, Kentucky. The largest first-form

male, 32.9 mm, was found in Lockard Cave, also in

Breckenridge County, and the smallest first-form male,

18.2 mm, in Blue Spring Cave, Lawrence County,

Indiana.

LIFE HISTORY NOTES.—Among the specimens examined in this study, first-form males were present in

collections made throughout the year except in March

and April, months represented by a total of only

seven specimens. A single ovigerous female was collected in Donaldson's Cave in June, 1924; this specimen has a carapace length of 23.9 mm, and 27 eggs

are either attached to her pleopods or are in the

bottom of the container. Jegla (1969) observed four

females carrying eggs during his study in Shiloh Cave:

one on June 30, two on August 16, and one on

August 20. These crayfish carried an average of

45 eggs each.

SEASONAL COLLECTIONS OF SPECIMENS EXAMINED

Not

Sex Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. known Total

SI

<$II

9

juv.

juv.

$ ....

9

Totals

2

3

3

-

-

8

1

4

1

_

-

6

-

3

1

-

2

6

-

1

-

-

-

1

1

_

_

1

-

2

2

12

10

-

-

24

1

3

_

1

1

6

1

2

1

-

-

4

7

14

13

-

-

34

13

2

7

-

-

22

4

5

16

1

2

28

4

11

19

1

-

35

11

12

14

-

-

37

38

72

84

4

3

206

Orconectes inermis testii (Hay)

FIGURES 1, 11, \2a-b

Cambarus pellucidus.—Packard, 1888:16 [in part].—Faxon,

1890:621 [in part].—Blatchley, 1897:127.—Ortmann,

1902:277, 278, 279 [in part]; 1905a:92 [in part].—

Harris, 1903b: 167.—G r a e t e r , 1909:470.—Spurgeon,

1915:387-394.—Ortmann, 1918:838, 848 [in part].—

Hobbs, 1942a:352 [in part].

Orconectes pellucidus inermis.—Packard, 1888:41.—

Rhoades, 1959:401-402 [in part].—Nicholas, 1960:133

[in part],—Hart and Hobbs, 1961:176 [in part].—

Rhoades, 1962:68 [in part].

Cambarus pellucidus var. testii Hay, 1891:148.—Spandl,

1926:95.—Wolf, 1934-1938:105 [in part].

Cambarus pellucidus testii Hay, 1893:283, 285, 286, pis.

44, 45, figs. 2, 5, 6, 10, 11, 12; 1896:478, 484, 485, fig.

4; 1897:209.—Faxon 1898:647.—Hay, 1899:959,966.—

Harris, 1903a:606; 1903b:58, 112, 118, 151, 152, 162.—

Banta, 1907:69-71, 87, 90.—Faxon, 1914:415.—Spurgeon, 1915:385-394 [in part].—Creaser, 1932:336.—

Rhoades, 1941:144; 1959:400, 401.

Cambarus pellucidus testi.—Hzy, 1902a: 233, 235.

Blind cray-fish.—Banta, 1905:853.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cambarus (Faxonius) pellucidus.—Ortmann, 1905a: 97,

107, 108 [in part]; 1905b:435 [in part].

Cambarus (Faxonius) pellucidus testi.—Ortmann, 1931:64

[by implication].

Orconectes pellucidus.—Spandl, 1926:95.—Pennak, 1953:

458 [in part], 459, fig. 286.—Eberly, 1958:1-6 [in part].

Orconectes pellucidus testii.—Hobbs, 1942a: 352 [by implication] .—Rhoades, 1944:117.—Hobbs, 1948a: 19, 20.—

Eberly, 1954:59; 1955:281, 282; 1958:3; 1960:30.—

Holthuis, 1964:43.—Hart and Hart, 1966:8.—Hobbs,

1967b:12.

Orconectes (Orconectes) pellucidus testii.—Hobbs, 1942a:

352 [by implication].

Cambarus (Cambarus) pellucidus testii.—Balss, 1955:1311.

Orconectes (Orconectes) pellucidus.—Hobbs, 1959:890 [in

part].

Blind crayfish.—Powell, 1961:82, 89.

Photograph.—Stenuit and Jasinski, 1964:75; 1966:75.

Orconectes.—Barr, 1968:85 [in part].

REVIEW OF LITERATURE.—The earliest reference

to this crayfish is that of Packard (1888) who indicated that Mr. C. H. Bollman had visited Mayfield's Cave and found Cambarus pellucidus there.

Faxon (1890) repeated the localities given by Packard and Hay (1891) proposed the varietal name

testii for the crayfishes in Mayfield's and Truett's

caves.

Hay (1893) elevated his varietal name, employing

the combination Cambarus pellucidus testii, pointed

out its diagnostic features and presented illustrations.

In 1896, he included this crayfish in his key to the

crayfishes of Indiana, outlined its outstanding features, presented three figures, and made the statement that the "first abdominal appendages of the

male do not differ in any respect from those of

typical C. pellucidus."

Blatchley (1897) gave a brief description of Mayfield's Cave, and indicated that C. pellucidus was one

of its inhabitants. Hay (1897) mentioned the species

but added no new information.

Faxon (1898) added no additional information

but made the following statement: "So this form

comes to bear a close likeness to C. bartonii and

suggests the possible derivation of C. pellucidus from

C. bartonii."

Hay (1899) included this crayfish in his list and

key to the Astacidae of North America. In a later

observation he (1902a) added no additional data

but, in commenting on its relationships, stated that

"Regarding the relationship of C. pellucidus testi

Hay, I will say that so far as is known this subspecies

is found in a very small area in Indiana at the very

northern limits of the range of C. pellucidus. C.

bartoni [-C. laevis] occurs in the same cave, but it

does not resemble the blind species in any way, and

has not even characters by which we can mark it as

a permanent resident. Were the conditions reversed

and C. bartoni tenebrosus [ = C. tenebrosus] found

anywhere in company with C. pellucidus testi there

might be some grounds for regarding them both as

possible intermediates between C. pellucidus and C.

bartoni, but under conditions as they exist such a view

is untenable."

Ortmann (1902, 1905a, 1905b), in discussing the

affinities of C. pellucidus, did not mention Hay's subspecies, but his remarks about the species were presumably intended to apply to it. The species was assigned to his subgenus Faxonius (1905a).

Harris (1903b) catalogued the information published by the above authors, but most of the natural

history data (pages 113-118) included under Cambarus pellucidus testii are based on observations on

O. pellucidus or those of Hay on the intergrades

inermis x testii in Shiloh Cave. Banta (1905) added

no new data.

Banta (1907) adopted Hay's subspecific designation for the troglobite in Mayfield's and Truett's

caves but indicated that the "subspecies is not clearly

defined." Inasmuch as his observations represent the

only significant account of observations on this crayfish in its native habitat, most are quoted here.

"This crayfish is usually seen quietly resting on the

bottom of a pool. Rarely one is observed walking

slowly. When roughly disturbed it acts and swims

much as other crayfish do when excited; that is, it

swims without regard to the edge of the pool or even

the direction of the bank. However, if there is a

disturbance of the water and the crayfish becomes

aware of the pursuer while at a distance or before

being touched, it in nearly every case, swims or crawls

toward protecting rocks shelving over the edge of the

pool, or to some other such means of concealment.

If there is no such protection it moves toward the

opposite side of the pool. It occasionally retreats to

a hole under a rock. C. bartoni [ = C. laevis] is quite

often found in such holes. Possibly C. bartoni alone

forms these holes and C. pellucidus makes use of

them when deserted by C. bartoni. Often when

slightly disturbed by an object close at hand C. pellucidus backs off, then turns around, and crawls forward. However, when disturbed it usually starts to
NUMBER 10 5 51

swim immediately. If crowded when crawling forward toward a place of concealment, it begins to

swim caudal end foremost without first turning about

and really swims toward its pursuer. But the moment

it begins to swim it either turns squarely over ventral

side up or turns to one side so as to move in the

direction in which it was crawling. After changing

the direction of its course it rights itself and soon

disappears if there is ready means of concealment.

In its swimming and crawling motions this crayfish

is not less active than other crayfish.

"This species is sensitive to a jar in the water at a

distance of several feet if the disturbance is quite

pronounced, like that produced by dropping a pebble

into the pool. But considerable rippling or slow swishing about in the water often fails to produce any

effect upon individuals at a little distance. It seems

insensible to sound, although a heavy jar on the

bank of the pool may cause it to move. Light often

fails to have any apparent effect, but on two occasions when a bright light was suddenly flashed upon

perfectly quiet individuals they moved immediately,

swimming rapidly from the lighted area. In these

two cases there could have been no jar or other

disturbance, for I had quietly crept to near the

individuals from a distance and then suddenly thrown

the light full upon them. Sometimes when the light

was held upon individuals for several minutes they

failed to respond at all; usually, however, they moved

after two or three minutes."

On the basis of laboratory observations, Banta

stated the belief that "they moult two to four or

five times a year depending upon the size, the smaller

or younger ones molting oftener." "Very young individuals were seen during February and March, the

earliest date being February 17."

Graeter (1909) summarized the observations of

Hay and Banta, referring to this crayfish as

"Varietat." Faxon (1914) only included this subspecies in his checklist.

Spurgeon (1915), in studying the eyes of two

troglobitic species, utilized specimens of both O. i.

inermis and O. i. testii. (See Review of Literature for

O. i. inermis, page 39.) Ortmann (1918) mentioned

that C. pellucidus occurs in caves in Indiana.

Spandl (1926) added no new data but mentioned

Hay's variety testii "in den Mayfield- und Tuett [sic]-

Hohlen." Similarly, Ortmann (1931) and Creaser

(1932) only recorded the crayfish from Indiana; the

former, however, discussed the ranges of C. pellucidus testii and its relatives.

Wolf (1934-1938) misinterpreted distribution records for this crayfish in previous literature, thus adding several erroneous records.

Rhoades (1941) stated that "no doubt C. pellucidus australis of the South bears the same affinity

to C. pellucidus pellucidus as does Cambarus pellucidus testii Hay (1893) of the North."

In his generic revision, Hobbs (1942a) assigned

Cambarus pellucidus, including its subspecies, to the

genus Orconectes, and in recognizing the subgenus

Faxonella (1942b) implied that these crayfishes were

members of the typical subgenus.

Rhoades (1944) assigned O. p. testii to his "Group

rafinesquei" because "the gonopods are short and

the tips are separated for only a short distance" and

"the tips are both recurved in the same direction."

Hobbs (1948a) discussed the affinities of the crayfishes of the Limosus Section, questioning the validity

of Rhoades' Limosus and Rafinesquei groups but did

not contribute any new data for this subspecies.

Pennak (1953) recorded O. pellucidus from Indiana, and his figure 286 is redrawn from Hay's (1893)

illustration of O. p. testii.

Eberly (1954) discussed the association of this

crayfish with Cambarus laevis, and (1955) added two

new locality records in Monroe County, Indiana.

Later (1958 and 1960) he cited the occurrence of

the subspecies in the same county. Balss (1955) presented no original information and erroneously cited

this crayfish from Wyandotte Cave. Hobbs (1959)

did not refer to the subspecies by name, but in his

key indicated that four subspecies of O. (O.) pellucidus ranged from Indiana to Alabama.

Rhoades (1959) compared the descriptions of

Cope's O. inermis and Hay's O. pellucidus testii and

concluded that the latter is synonymous with O.

inermis and adopted the combination O. pellucidus

inermis for both. Although he mentioned populations

occurring in Brown and Bartholomew counties, Indiana, he cited no specific localities.

In his checklist of troglobites in North America,

Nicholas (1960) followed Rhoades (1959) in recognizing O. p. inermis as the only troglobitic crayfish

other than O. p. pellucidus occurring in Indiana.

Powell (1961) reported "blind crayfish" from two

caves in Monroe County, Indiana.

Hart and Hobbs (1961) cited O. pellucidus inermis
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in May's and Eller's caves as a host of Entocythere

barri ( = Sagittocythere barri).

Rhoades (1962) again treated 0. p. testii as a

synonym of O. p. inermis in his discussion of the

evolution of the crayfishes of the Limosus Section of

the genus.

Stenuit and Jasinski (1964 and 1966) published a

photograph of this crayfish.

Holthuis (1964) mentioned the subspecies in comparing the descriptions of O. pellucidus with Joseph's

Cambarus stygius. Hart and Hart (1966) cited this

crayfish as a host of the commensal entocytherid,

Sagittocythere barri, in May's Cave, Monroe County,

Indiana. Hobbs (1967b), in discussing parallel evolution in relatives of Procambarus pecki and O.

limosus, referred to this subspecies.

The most recent reference to this crayfish was that

of Barr (1968) who alluded to it in briefly discussing

the origin of the troglobitic Orconectes fauna.

COMMENTS ON PREVIOUSLY RECORDED DATA.—

Along with Packard's (1888) citation of the occurrence of this crayfish in Mayfield's Cave, he also

quoted from a letter from Dr. John Sloan who reported that "three eyeless crayfishes" had been collected in "Caves at Clifty," a locality which Packard

reported as being located in Bartholomew County.

Actually the caves referred to are in Washington

County. Spurgeon (1915) mentioned collecting crayfishes in Clifty Cave in the latter County. We have

specimens from River and Endless caves (the Clifty

caves) and they belong to what we consider to be

the intergrade population, inermis x testii.

Hay's (1893) statement that the first pleopods "do

not differ in any respect from those o f O. pellucidus

obviously is incorrect (cf. Figures lla, e, g and

14a, e, ;) .It seems highly probable to us that Hay's

comparison was made with specimens which were

from the more southern Indiana counties and which

he considered to be conspecific with O. pellucidus but

were either O. i. inermis or intergrades between O. i.

testii and the latter.

Faxon's (1898) statement that O. i. testii might

represent an intermediate form between O. pellucidus

and C. bartonii, suggesting a possible derivation of

O. pellucidus from the latter, was refuted by Hay

(1899) and has received no further consideration.

Our reasons for not adopting the treatment of this

subspecies proposed by Rhoades (1959) have been

discussed under O. i. inermis. His statement (1962:

68), however, that Orconectes pellucidus inermis "is

the blind crayfish of cave waters of Monroe, Brown,

and Bartholomew counties in south central Indiana"

leaves some doubt as to his concept of Cope's inermis,

for the type-locality of the latter is in Crawford

County. Rhoades' reference to specimens from Brown

County is the only one that we have encountered,

and, unfortunately, he cited no specific locality. If

his reference, without specific locality, to specimens

in Bartholomew County is based on the "Clifty

Caves" record of Packard, then there are no records

of this or any other troglobitic crayfish for that

county. None of the Indiana speleologists, including

Mr. Richard L. Powell of the Indiana Geological

Survey, consulted by us is aware of the existence of

caves in either Bartholomew or Brown counties.

See also the comments on previously recorded data

under Orconectes i. inermis (page 42).

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum without marginal spines or tubercles, acumen not delimited at base from remainder

of rostrum, margins convergent, upper surface without median carina; postorbital ridges terminating

cephalically without tubercles or spines; hepatic area

devoid of spines; at most, cervical spines represented

by very small tubercles; areola 4.6 to 6.4 times longer

than broad and constituting 42.3 to 45.8 percent of

entire length of carapace; chelae not conspicuously

setose but with ciliated tubercles and punctations

and a few long stiff setae; mesiodorsal surface of palm

with several irregular rows of tubercles; hooks on

ischiopodites of third and fourth pereiopods. First

pleopod of first-form male with greatest cephalocaudal diameter of pleopod less than twice that immediately proximal to base of central projection, always terminating in only two elements; non-corneous

mesial process broad basally, suddenly contracting to

form tapering subacute distal portion, latter directed

caudodistally and extending distally to about level of,

or slightly exceeding tip of, central projection; central

projection corneous, flattened in cephalocaudal plane,

slightly concavoconvex with convexity extending

mesially; shoulder lacking at base of central projection but cephalic surface with convexity on distal

half of appendage. Annulus ventralis as illustrated

in Figure Ilk.

Topotypic Male (Form I): Body (Figure lie, I)

subovate, depressed. Abdomen narrower than thorax

(8.8 and 9.6 mm in widest parts, respectively). Width
NUMBER 10 5 53

FIGURE 11.—Orconectes inermis testii: a, mesial view of first pleopod of topotypic male, form

I; b, mesial view of first pleopod of topotypic male, form II; c, dorsal view of carapace of topotypic male, form I; d, mesial view of first pleopod of topotypic male, form II; e, lateral view of

first pleopod of topotypic male, form I; /, dorsal view of distal podomeres of cheliped of topotypic male, form I; g, caudal view of first pleopods of topotypic male, form I; h, basal podomeres of third and fourth pereiopods of topotypic male, form I; i, antennal scale of topotypic

male, form I; ;', epistome of topotypic male, form I; k, annulus ventralis and portion of sternum

of topotypic female; /, lateral view of carapace of topotypic male, form I.

of carapace greater than depth in region of caudodorsal margin of cervical groove (9.6 and 7.5 mm).

Areola moderately broad (5.5 times longer than wide)

with 5 or 6 minute punctations across narrowest part.

Cephalic section of carapace 1.3 times longer than

areola; length of areola 43.1 percent of entire length

of carapace. Rostrum with length about 1.5 times

width at base, excavate, and with acumen approximately 0.65 times as long as maximum width; cephalic extremity reaching only to end of antennular

peduncle; margins not swollen, little elevated, and

uninterrupted by marginal spines or tubercles; upper

surface with regularly spaced, minute, setiferous

punctations; sub rostral ridges moderately well developed and evident in dorsal aspect along basal third

of rostrum.

Postorbital ridges weakly developed, short, with

shallow dorsolateral grooves, and terminating cephalically without spines or tubercles. Suborbital angle

lacking. Branchiostegal spines acute but small. Cervi-
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cal spines represented by row of 2 or 3 very small

tubercles on each side of carapace immediately caudal

to cervical groove, tubercles progressively larger ventrally. Carapace punctate dorsally and very weakly

granulate laterally; hepatic area with inconspicuous

granuliform tubercles but lacking spines. Abdomen

longer than carapace (25.0 and 21.7 mm). Cephalic

section of telson with 2 well-developed spines in each

caudolateral corner (mesial ones movable).

Protruding portion of epistome (Figure 11;') approximately twice as long as broad and resembling

silhouette of shortened minaret; surface flattened and

with fine setae; fovea present posteriorly. Eyes much

reduced, without pigment, almost completely hidden

beneath rostrum in dorsal aspect and extending

cephalically slightly more than half distance between

margin of orbit and base of acumen. Antennules of

usual form with moderately well-developed spine on

ventral surface of proximal podomere at base of distal

fifth. Antennae extending caudally beyond end of

telson for distance equivalent to length of telson.

Antennal scale (Figure Hi) broadest distal to midlength, about half as broad as long; outer thickened

portion much narrower than lamellar area and terminating distally in prominent corneous-tipped spine.

Third maxillipeds extending to about midlength of

distal podomere of peduncle of antenna.

Chela (Figure 11/) comparatively slender and not

much inflated; mesial margin of palm approximately

1.4 times longer than width of palm; dorsal surface

of palmar area tuberculate mesially and punctate

laterally, punctations and distal bases of tubercles

setose; ventral surface of palm with weak tubercles

and with small corneous-tipped spiniform tubercle

opposite base of dactyl; more mesial tubercles on

palm larger and sublinearly arranged, with 9 or 10

comprising most mesial row; lateral surface tuberculate. Fingers not gaping; dorsal and ventral surfaces

with broad, rounded, submedian, longitudinal ridges

flanked by setiferous punctations; opposable margin

of immovable finger with row of 7 tubercles, fifth

from base largest, along proximal two-fifths, single

tubercle below level of row slightly proximal to midlength of finger, and single row of minute denticles

between proximal five tubercles, distally forming two

rows extending almost to base of corneous tip of

finger; lateral surface of immovable finger subcostate.

Opposable margin of dactyl with row of 5 tubercles

along proximal two-fifths, distalmost largest, with

denticles arranged as on immovable finger; mesial

surface of dactyl almost smooth. Carpus longer than

broad, with mesial, dorsomesial, and ventromesial

surfaces tuberculate, otherwise mostly punctate; mesial

surface with 2 tubercles somewhat larger than others,

largest distinctly acute; distoventral margin with 2

acute tubercles, 1 adjacent to base of ventrolateral

articulation with propodus and 1 near median line.

Merus tuberculate dorsally, with 2 corneous-tipped

tubercles slightly proximal to distal end; mostly

punctate mesially and laterally; ventrolateral margin

with row of 9 prominent tubercles, and ventromesial

margin with 12; few scattered tubercles flanking both

rows; distolateral extremity with small corneoustipped spine. Ischium with ventromesial row of 6

small tubercles; dorsal margin scabrous but with only

2 well-defined tubercles.

Ischia of third and fourth pereipods (Figure 11 h)

with hooks; those on third stout and recurved to or

beyond level of cephalic articulation with basis; those

on fourth much smaller and, although recurved,

leaving broad gap between tip and corresponding

basis. Coxae of fourth pereiopods with caudomesially

projecting prominences; coxae of fifth pereiopods

without prominences except for small mesioventral

projection partially surrounding base of phallic

papilla.

First pleopods (Figure 11 a, e,g) symmetrical, reaching bases of third pereiopods when abdomen is flexed,

and comparatively shallowly situated in sternal groove;

tip ending in two parts as described in Diagnosis.

Topotypic Female: Differs from male in following respects: apex of rostrum reaching only midlength

of ultimate podomere of peduncle of antennule; cervical spines represented by 4 minute tubercles, ventralmost largest but scarcely larger than other tubercles

on hepatic and branchiostegal regions; cephalic section of telson with only 1 spine in caudodextral corner;

opposable margin of immovable finger of chela with

row of 11 tubercles along basal three-fifths, fourth

from base largest; opposable margin of dactyl of chela

with row of 15 small tubercles and with 2 larger

tubercles below level of row and between fourth and

fifth tubercles.

Annulus ventralis (Figure 11 k) shallowly situated

in sternum and not firmly fused to sternum immediately cephalic to it; outline subovate, 1.6 times

broader than long, with median portion elevated

ventrally forming asymmetrical arc with highest seg-
NUMBER 105 55

ment approximately at midlength; arched portion

bearing shallow longitudinal groove ending caudally

on steep slope of arc; sinus originating on slope

dextral to median line, making gentle arc to median

line and extending caudally to midcaudal margin of

annulus.

Syntypic Male, Form II: Differing from first-form

male in following respects: rostrum reaching almost

to distal end of peduncle of antennule; cervical tubercles exceedingly minute, only one at all larger than

granulations cephalic and caudal to it; branchiostegal

spines almost obsolete; lower surface of palm of chela

with few tubercles on lower proximomesial surface;

opposable margin of immovable finger of chela with

row of 20 small tubercles along proximal five-sixths of

finger and with 2 larger ones situated below level of

row between fourteenth and sixteenth tubercles; opposable margin of dactyl of chela with row of 26

along proximal six-sevenths of finger and with 2

larger ones situated below level of row between

seventh and ninth tubercles. Hooks on ischia of third

and fourth pereiopods comparatively more reduced,

but boss on coxa of fourth only slightly smaller.

(See measurements.)

First pleopod (Figure life, d) reaching base of third

pereiopod when abdomen is flexed and with most

features of first-form male; however, central projection non-corneous, shorter, and more inflated.

MEASUREMENT S (in millimeters).—Orconectes

inermis testii:

Topotypic Topotypic Syntypic

Male Male

Form I Female Form II

Carapace:

Height 7.5 9.8 9.2

Width 9.6 13.0 12.3

Length 21.7 27.8 26.7

Rostrum:

Width 3.2 3.8 4.1

Length 4.9 6.0 5.5

Areola:

Width 1.7 2.3 2.3

Length 9.4 12.8 11.9

Chela:

Length of inner margin

of palm 6.8 7.6 9.2

Width of palm 5.0 6.2 6.7

Length of outer margin

of chela 16.6 19.5 25.6

Length of dactyl 9.2 10.6 14.6

TYPES.—Syntypes, USNM 17702 (2c?II, 1?) ,

Museum of Comparative Zoology 7431 (1 cf II, 1 ? ) •

TYPE-LOCALITY.—Mayfield's Cave, SW ]/4, NE

 l/4,

SW y4, sec. 26, T.9N, R.2W, Monroe County, Indiana.

SPECIMENS EXAMINED.—Specimens from Indiana

were examined as follows:

Monroe County: (1) Carmichael Cave, NW }4, NW */4,

SW ]/4, sec. 19, T.7N, R.1W, 1 $ I, H.H.H. Ill, X/4/69.

(2) Eller's Cave, NW / 4 , NW }4, SW J4, sec. 15, T.8N,

R.2W, 3<JI, 1^ II, 1$, W. R. Eberly, 1/10/53; I $11,

1$, H.H.H. Ill, X/21/69; "probably from Eller's Cave,"

2 51, 2 5II, 19, date and collector unknown. (3) Mayfield's Cave, SW l/4, NE

 l/4, SW l/4, sec. 26, T.9N, R.2W,

2 5 II, Ij9 [syntypes], W.P.H., date unknown; 2 5 II, 3$ ,

Ij5, W.P.H., date unknown; \$l, 19, T.C.J., X/17/59;

1 5 I, T.C.J., 11/19/60; 1 5II, H.H.H. Ill, IX/20/69. (4)

May's Cave, SW */4, SE / 4 , NE «4, sec. 24, T.8N, R.2W,

15H, 19, 1951, collector unknown; 19, C.E.B., 1/18/51;

15II, 19, T.C.B., VIII/19/57; 19, H.H.H. Ill, IX/

26/69; 15II, collector and date unknown. (5) Reeve's

Cave, SW l/4, SW J4, NE '/4, sec. 34, T.8N, R.2W, 19.

H.H.H. Ill, X/3/69. (6) Salamander Cave, SW »/4, SW

54, NW y4, sec. 9, T.8N, R.2W, 19, H.H.H. Ill, X/8/69.

(7) Shaft Cave, NE >/4, NE

 l/4, SE */4, sec. 8, T.8N, R.2W,

151, 19, H.H.H. Ill and K. Burdsall, XII/5/69.

This crayfish was reported by Hay (1891) and others

to occur in Truett's [sic] Cave in Monroe County,

and it is probable that the "blind crayfish" observed

in Goode's Cave and Ranard School Cave by Powell

(1961:82, 89) were member of this subspecies. No

specific localities are known for Bartholomew and

Brown counties.

RANGE.—Orconectes inermis testii is thus restricted

to the subterranean waters of Monroe County, Indiana, but intergrades with the typical subspecies in the

more southern counties of the State.

The restricted gene flow between inermis testii and

inermis inermis is geographically paralleled in the

troglobitic beetle Pseudanophthalmus shUohensis

Krekeler (Carabidae). Nominate shUohensis occurs

in most Lawrence County caves north of the East

Fork of the White River, but the Monroe County

caves are occupied by P. s. mayfieldensis Krekeler

(T. Barr, manuscript in preparation). The biogeographic similarities in the distribution of these two

polytypic species of arthopods suggest a limited and

only partially effective extrinsic barrier between the

Bedford area caves and those of the Bloomington area.

VARIATIONS.—Although minor variations are abundant in our comparatively few specimens of this crayfish, all of them seem to be individual ones, rather

insignificant, and, except for those mentioned below,
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBE R 10 5 57

FIGURE 13.—Cephalic portion of carapace of Orconectes inermis inermis illustrating variation

in spination (all localities in Kentucky): a, b, Cub Run Cave, Hart Co. (9 , 2)\ c, d, Riders

Mill Cave, Hart Co. (31 , 31) ; e-g, Cooch Webb Cave, Hart Co. (31 , 9 , 9 ) ; h, Brush

Creek Cave, Green Co. ($11); i, Bland Cave, Hardin Co. ( 9 )•

scarcely exceed those variations noted in the above

descriptions.

In the single first-form male from Salamander

Cave, the carapace of which is damaged, the rostrum

FIGURE 12.—Cephalic portion of carapace of Orconectes

inermis testii (a, b), O. i. inermis (c, z,), and intergrades

between them, illustrating variation in spination: a, b,

Eller's Cave, Monroe Co., Ind. (9 , 31); c-e, Shiloh Cave,

Lawrence Co., Ind. (311, 9, 311); /, Sullivan Cave,

Crawford Co., Ind. (31); g, Archibald Cave, Crawford

Co., Ind. (311); h, Marengo Cave, Crawford Co., Ind.

(9) ; i, Wyandotte Cave, Crawford Co., Ind. (311); ;,

Donaldson's Cave, Lawrence Co., Ind. ( 31); k, River Cave,

Washington Co., Ind. ( 9 ) ; /, King's Cave, Harrison Co.,

Ind. (311); m, Bedford Cave, Harrison Co., Ind. (31);

n, Lime Kiln Cave, Meade Co., Ky. ( 9 ) ; o, Doe Run,

Meade Co., Ky. ( 9 ); p, Rockhaven Cave, Meade Co., Ky.

( 9 ); q, Cave, 5 mi. E of Big Spring, Breckenridge Co.,

Ky. ( 3 II) ; r, Bandy Cave, Breckenridge Co., Ky. ( 2 ) 5 s,

Same as q ($11); t, u, Thornhill Cave, Breckenridge Co.,

Ky. (31 , $1); v, w, Lockard Cave, Breckenridge Co., Ky.

( 9 , 31); x, Same as r ( 9 ); y, z, Nelson Cave, Hardin

Co., Ky. (9 , 9) .

bears a pair of minute corneous tubercles. The rostral

margins are strongly convergent and testii-like, and

the tubercles are so small as to be evident only on

close examination. This is the only specimen of this

subspecies that we have seen that has marginal ornamentation of any kind on the rostrum.

Only three of our specimens, two from Mayfield's

Cave and one from Eller's Cave, have areolae that

constitute less than 44 percent of the carapace length:

those from Mayfield's Cave comprise 43.3 (cTI) and

43.6 ( cT II) percent, and that from Eller's Cave, 42.3

( $ ) percent. Their carapace lengths are 21.7, 23.6,

and 19.4 mm, respectively. All three are spineless

and have strongly convergent rostral margins; thus,

except for the proportionately shorter areolae, they

are typical of the subspecies.

Perhaps the most conspicuous variations are in

the chelipeds, some being more robust than others,

some with the tubercles along, immediately above,

or below the mesial margin of the palm arranged in

almost perfect rows, and others with scarcely any
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sort of alignment evident. The number and arrangement of tubercles on the opposable surfaces of the

fingers of the chelae are likewise exceedingly variable

(see the descriptions of the fingers of the chelae of

the two males described herein). We strongly suspect

that much of this variation will eventually be correlated with the loss of the original appendage and

subsequent regeneration.

Hay (1893:285) discussed the hooks on the ischia

of the third and fourth pereiopods in his C. pellucidus

from Indiana. In all of the first-form males of both

subspecies of O. inermis that we have examined, hooks

are present on both pairs of legs. In young secondform and juvenile males, they are exceedingly small,

reduced to tubercles, or apparently lacking.

SIZE.—The largest specimen that we have is a female from Eller's Cave which has a carapace length

of 33.9 mm. The largest first-form male, carapace

length 30.9 mm, was collected in Mayfield's Cave,

and the smallest, which was taken from the same

cave, has a carapace length of 22.9 mm.

LIFE HISTORY NOTES.—Collections available to us

were made during the months of January, February,

and August through November, and first-form males

are among those made in January, February, October, and November. No ovigerous females or those

carrying young have been reported. Banta (1907)

indicated that "very young individuals" had been

observed in February and March and "at no other

time of the year."

SEASONAL COLLECTIONS OF SPECIMENS EXAMINED

Not

Sex Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dee. known Total

$1 31-------2- - 4 10

$ \ \ 1-1----11-- - 5 9

9 2-1---- 1 1 3- 1 5 1 4

juv . $ .... ----------- - 1 1

juv . 9 .... ----------- - 1 1

Total s 612----225-11 6 3 5

Orconectes pellucidus (Tellkampf) 1888:8, 10, 12, 19, 24, 25, 38, 39, 40, 41, 42, 82, 86,

110, 111, 112, 119, 122, 123, 125, 127 [in part].—GarFIGURES 1, 14-16 man, 1889:235, 236.—Faxon, 1890:626, 628.—Packard,

1890:393 [in part].—Parker, 1890:153, 154, 155, 157-

Krebse.—Anonymous, 1843a:49. 161, pi. 1, figs. 2, 3, 4, 5, 6.—Hovey, 1891:72.—Ortmann,

Astacus Bartoni?.—Anonymous [not Fabricius], 1843b: 1892:11 [in part].—Cunningham, 1893:537.—Stebbing,

175.—Putnam [not Fabricius], 1872:10. 1893:208.—Lonnberg, 1894:126; 1895:4-6, 9.—Hay,

Astacus pellucidus Tellkampf, 1844a:684; 1844b:383.— 1896:485 [in part].—Call, 1897:103, 104.—Hay, 1897:

Thompson, 1844: 111.—Tellkampf, 1845:85, 93.—Erich- 208 [in part].—Faxon, 1898:647 [in part].—Eigenmann,

son, 1846:87, 89, 95.—Gibbes, 1850:195.—Dana, 1852: 1899:60.—Hay, 1899:959, 966 [in part].—Call, 1901:

522.—Newport, 1855:164.—Lucas, 1864:iv.—Hagen, 103, 104.—Ortmann, 1902:227, 278, 279 [in part].—

1870:6, 7, 11, 55.—Smith, 1873:639.—Faxon, 1885a: 10. Hay, 1902a:226, 228, 230, 232, 234, 235 [in part];

—Rhoades, 1944:112; 1959:399. 1902b:436.—Steele, 1902:7, 16, 18 [in part].—EigenAstacus (Cambarus) pellucidus.—Erichson, 1846:95,96.— man, 1903:169 [in part].—Harris, 1903a:602, 606;

Rabe, 1890:9. 1903b:58, 67, 70, 80, 112-118, 151, 153, 157, 162, 167

Craw-fish.—Silliman, 1851:336. [in part].—Ortmann, 1905a:92, 95, 96, 97 [in part].—

Cambarus pellucidus.—Girard, 1852:87, 88.—Hagen, 1870: Bell, 1906:300, 304, 305.—Banta, 1907: 6, S9, 70, 71,

8, 27, 30, 31, 32, 33, 34, 55, 56, 97, 101, pi. 1, figs. 68- 72, 73, 102, 103 [in part].—Graeter, 1909:470 [in part].

71, pi. 3, fig. 148, pi. 6.—Packard, 1871:750, 751, fig. —Pearse, 1910:10.—Hovey, 1912:80, 81, 108, 109, 115,

131.—Cope, 1872a: 410, 419.—Packard, 1872b: 17, 18, 119, 124, 2 figs.—Osborn, 1912:923 [in part].—Faxon,

fig. 131.—Hagen, 1872:494, 495.—Packard, 1873:94.— 1914:415, pi. 7, fig. 2a-c [in part].—Spurgeon, 1915:

Smith, 1873:639 [in part].—Packard, 1874:209.—Put- 385, 386 [in part].—Pratt, 1916:391, 392, fig. 625 [in

nam, 1875a:222; 1875b: 191, 198.—Smith, 1875:477 [in part].—Ortmann, 1918:838, 848 [in part].—Garman,

part].—Putnam, 1877:16-19.—Packard, 1879:315, 316, 1924:88, 89.—Spandl, 1926:95, 141-142, 148 [in part].

317, figs. 268, 269.—Hubbard, 1880:38.—Leydig, 1883: —Stiles and Hassell, 1927:219 [in part].—Chappuis,

38, 39, 40.—Faxon, 1884:139, 140 [in part]; 1885a:4, 1927:91, 120 [in part].—Creaser, 1931:10; 1932:336.—

5, 7, 8, 9, 11, 16, 18, 19, 40, 41, 42, 43, 44, 45, 46, Giovannoli, 1933a:618-619, fig. 90; 1933b:236-237, fig.

59, 82, 83, 84, 111, 158, 169, 174, 178, 179 [in part]; 90.—Wolf, 1934-38:104 [in part].—Turner, 1935:876.—

1885b:358.—Underwood, 1886:371 [in part].—Packard, Park, 1938:209.—Fleming, 1939: 304, 305.—Bouvier,
NUMBER 10 5 59

1940:68 [in part].—Park et al., 1941:154-171, 5 figs.—

Rhoades, 1941:141, 142, 144.—Hobbs, 1942a: 335, 338,

342, 351, 352 [in part].—Jeannel and Henrot, 1949:21.—

Bott, 1950:25.—Dearolf, 1953:229.—Hobbs and Barr,

1960:19.—Vandel, 1964:448, 453, 461, 501, 502, fig. 76.

—Hobbs, 1967a: 125.

Crabs.—Darwin, 1859:137.

Eyeless Crabs.—Binkerd, 1869:86.

Cambarus pelulcidus.—Hagen, 1870:106 [Erroneous spelling].

Orconectes pellucidus.—Cope, 1872a:409, 410, 419; 1872b:

161, 162, 173, 174; 1879:492, 494, 495, 505, 506 [in

part].—Cope and Packard, 1881:879, 881, 882.—Joseph,

1882:12.—Wright, 1884:272, 273.—Faxon, 1884:139.—

Underwood, 1886:371.—Packard, 1888:24, 126, 140, 155,

pi. 21, fig. 2, pi. 22, fig. 7.—Packard, 1894:735, 742.—

Hobbs, 1948a: 19.—Pennak, 1953:458 [in part].—

Eberly, 1954:59; 1958:1, 2, 3 [in part].—Wells, 1959:5-

7.—Eberly, 1960:29, 30, 31 [in part].—Brown, 1961:

929, 930.—Creaser, 1962:3 [in part].—Rhoades, 1962:

68, 79, 94.—Fitzpatrick, 1963:60.—Holthuis, 1964:42,

43, 45, 47.—Vandel, 1964:509, 570, 575 [in part].—

Mohr, 1964:828.—Poulson, 1964:752, 756, 757, 759,

762, 764.—Frey, 1965:623, 624.—Fitzpatrick, 1967:141,

142.—Mohr and Poulson, 1966:166, 204.—Barr, 1967a:

160, 161, 186, 187, 192, pi. 46, fig. 10; 1967b:480.—

Hobbs, 1967a: 130; 1967b: 12 [in part].—Thompson,

1967:46, 47.—Barr, 1968:65, 91, fig. 18; 1971:71, 72.

Astacidae.—Shaler, 1875:361; 1876:10.

Cray fish.—Shaler, 1875:362, 363; 1876:11, 12.

Cambarus typhlobius Joseph, 1880:202.—Faxon, 1884:139;

1885a: 7, 45.—Underwood, 1886:373.—Hay, 1896:477.—

Harris, 1903b: 131, 151.—Faxon, 1914:427.—Bouvier,

1940:68— Van Straelen, 1942:2.—Bott, 1950:25.—Villalobos, 1953:348.—Villalobos, 1955:11.—Holthuis, 1964:

42, 43, 44, 45, 46, 47.

Cambarus stygius Joseph, 1881:241, 249 [nomen nudum];

1882:12 [homonym].—Underwood, 1886:373.—Packard,

1888:86, 123.—Faxon, 1914:427.—Spandl, 1926:95.—

Stammer, 1932:608.—Wolf, 1934-1938:105.—Holthuis,

1964:42, 43, 44, 45, 47. [Not Cambarus stygius Bundy,

1876.]

Cambarus coecus Joseph 1881:237 [nomen nudum].—

Faxon, 1884:139; 1885a:7, 45; 1914:427.—Holthuis,

1964:42, 45.

Blind crayfish.—Semper, 1881:77.—Call, 1901:101.

Cambarus (Orconectes) pellucidus.—Hovey, 1882:222 [in

part].

Orconectes.—Hovey, 1882:223 [in part].—Barr, 1968:85;

1971:72, 85, 86.

Cambarus Stygius.—Faxon, 1884:139; 1885a: 7, 45, 46.—

Bott, 1950:25.

Camtarus (Orconectes) pellucidus, form inermis.—Packard,

1888:156, pi. 27, fig. 5. [Erroneous spelling of Cambarus.']

Astacus Cambarus Stigius.—Rabe, 1890:9.

Cambarus.—Apfelbeck, 1895:24.—Bolivar et Jeannel, 1931:

306, 307, 309.—Vandel, 1964:494, 495, 512.

Cambrus pellucidus.—Price, 1900:155 [erroneous spelling].

Crayfish.—Call, 1901:100.

Cambarus (Faxonius) pellucidus.—Ortmann, 1905a: 107,

108, 111, 114, [in part]; 1905b:435 [by implication];

1931:64, 65 [in part].—Fage, 1931:373 [in part].—

Turner, 1935:876.

Cambarus caecus.—Bouvier, 1940:68 [erroneous spelling of

coecus].

Cambarus pellucidus pellucidus.—Rhoades, 1941:144.

Cambarus pelludicus.—Dearolf, 1942:50 [Erroneous spelling].

Crawfish.—Dearolf, 1942:52.

Cambarus Pellucidus.—Jackson, 1942:4.

Cambarus (Cambarus) pellucidus.—Balss, 1944:402; 1955:

1311, 1312.

Orconectes pellucidus pellucidus.—Rhoades, 1944:112, 113,

115, 117, 120, 121.—Hobbs, 1948a: 16, 19, 20.—Rhoades,

1959:401.—Cole, 1959:81.—Eberly, 1960:30.—Hobbs

and Barr, 1960:19.—Nicholas, 1960:133.—Barr, 1961:32.

—Rhoades, 1962:68, 90, 91.—Wolfe and Cornwell, 1964:

1467, 1468.—Jegla et al., 1965: 639.—Hart and Hart,

1966:8, 9.—Jegla, 1966:346, 347, 353.—Hobbs, 1967a:

131; 1967b: 7.—Barr, 1968:60.

Orconectes pellucidus pelluicdus.—Hobbs, 1948a: 16 [erroneous spelling].

Cambarus Coecus.—Bott, 1950:25.

Gambarus typhlobius.—Croizat, 1958:908 [Erroneous spelling].

Orconectes (Orconectes) pellucidus.—Hobbs, 1959:890 [in

part].

Crayfishes.—Barr, 1964:79; 1966:15.

REVIEW OF LITERATURE.—The first reference to this

crayfish that has come to our attention is an anonymous account of the occurrence of a white crayfish

in Mammoth Cave published in "Das Ausland" on

13 January 1843. In the same year, an anonymous

record of a gift of "a white eyeless crayfish (Astacus

Bartoni ?) " from Mammoth Cave, Kentucky, by

W. T. Craige to the Academy of Natural Sciences

appeared. In 1844, Thompson reported a blind "crayfish" from Mammoth Cave and considered it to be

conspecific with A. bartonii. Tellkampf (1844a), however, disagreed with Thompson as to the identity of

the crayfish occurring in Mammoth Cave and referred to it in a footnote in "Das Ausland," on 19

June as follows: "Astacus pellucidus, unserem A.

fluviatilis verwandt, dessen Grosse er jedoch nicht

erreicht. Die Scheren des ersten Fusspaares, sind

schlank und zart, des vorderste Glied wenig gebogen,

schwach gezahnt. Die Augen liegen unter dem

Kopfschilde versteckt." In the text referring to this

footnote he stated that "Nur wenige Schritte waren

wir gegangen, als ich einen kleinen, ungefahr 1^4

Zoll longen Krebs*) in dem seichten Wasser erblickte,
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

den ich ohne Muhe fing. Er war weiss und seine

Schale so durchsichtig, das man die Bewegung der

Kiemen und die innern Thiele durchscheinen sah."

Whereas in the same year (Tellkampf 1844b), a more

complete description appeared that has consistently

been cited as the original description, it bears no

specific date of publication. Therefore, it seems to us

that the description quoted above must take precedence. The more complete description is as follows:

"Der Astacus pellucidus Nob. ist unserem A. fluviatilis verwandt, erricht aber nicht dieselbe Grosse.

Die Scheeren des ersten Fusspaares sind schlank und

zart, des vorderste Glied wenig gebogen, schwach

gezahnt. Die Augen stehen nicht vor, sondern sind

unter dem Kopfschilde verstechkt. Die Fuhler lang

und sehr grazil. Die ersten Glieder der zwei vorleitzien Fusspaare haben auf ihrem vorderen Rande ein

nach vorn und inner gerichtetes, etwas gebogenes

Horn, das Horn des letzten Fusspaares ist nur angedeutet. Thomson sagt, dass der Krebs miit Astacus

Bartoni ubereinstimme, allein er ist davon bestimmt

verschieden." The following year, an English translation of Tellkampf's paper appeared.

Erichson (1846) redescribed the species, adding

several additional characters, and assigned it to his

new subgenus Cambarus. Gibbes (1850) and Silliman (1851) referred to specimens from and in

Mammoth Cave.

Girard (1852) elevated the subgenus Cambarus to

generic rank and was the first person to mention the

"anterior pair of abdominal legs" of male crayfishes,

indicating that the tips are "twisted in C. pellucidus."

In the same year, Dana discussed the relationships of

the American crayfishes to those occurring elsewhere

and to the lobsters and other Reptantia.

In 1855, Newport recorded the first detailed observations on the eyes of this crayfish, and Lucas

(1864) indicated that organs of vision were lacking.

Darwin (1859) referred to the eyes of blind crabs in

Carniole and Kentucky but added no original data.

Binkerd (1869) referred to pellucidus as an eyeless

crab which was rather scarce and seems to be "destitute of the power of hearing, but any motion imparted to the water seemed to create alarm. They

probably have a high nervous sensibility, which is not

inconsistent with their fair, soft, gelatinous appearance, in which they do not differ from the fish." He

further supposed that they were viviparous.

Hagen (1870), in his monograph of the American

Astacidae, presented a description of the species in

Latin together with illustrations. He discussed its affinities with his Group I [ = Procambarus] and Group

II [ = Orconectes], and pointed out that it is the most

aberrant species of the genus.

Cope (1872a and 1872b, essentially identical articles) compared the fauna of Wyandotte and Mammoth caves and erected the genus Orconectes to

receive the albinistic crayfishes occurring in them,

defending the rationale of proposing a new genus to

receive the two troglobites. Packard (1871 and 1872b)

are essentially identical. In discussing the inhabitants

of Mammoth Cave and their evolution, he emphasized Hagen's observations on the atrophied eyes and

presented a dorsal view of the animal which was

redrawn from Hagen's figure.

Hagen (1872) not only pointed out the folly of

Cope's erecting the genm Orconectes but also added

that Cope had not given "any character by which

to separate it [O. inermis] from the old species, C.

pellucidus."

Putnam (1872) simply quoted a sentence from the

anonymous article (1843). Packard (1873) and Smith

(1873 and 1875) agreed with Hagen that Cope's O.

inermis was not specifically different from C. pellucidus and that the species should be retained in the

genus Cambarus.

Packard (1874), at an entomological meeting of

the Boston Society of Natural History, exhibited drawings of the supraoesophageal ganglia of this crayfish

and of an epigean species; the differences noted were

mainly in the enlargement of the "sides of the ganglion in the blind species." He showed further that

the pigment cells are white and stated that the whole

eye was "in a state of arrested development."

Putnam (1875a) demonstrated several living pellucidus from Mammoth Cave at the 2 December 1874

meeting of the Boston Society of Natural History,

and (1875b) the same specimens had been exhibited

at the Essex Institute on 25 November, 1874. He

added another locality for the species—a cave several

miles down the Green River from Mammoth Cave

and on the opposite bank (perhaps Ganter's Cave).

Shaler (1875) referred to the Astacidae in Mammoth Cave and speculated on its origin with reference to Pleistocene glaciation. He expressed the view

that the cave fauna had been derived from, and was

being reinforced by interbreeding with, the epigean

fauna.
NUMBER 10 5 61

Putnam (1877) recounted observations of feeding

behavior, molting, and regeneration of appendages

among specimens retained in the laboratory from

13 November 1874 until 7 August 1875. One female

molted twice, 28-29 January and again on 20 April

with almost complete regeneration of legs and antennae following the second molt. During this time

the animals withstood temperatures varying from a

heated room to those which froze "the water in

their jars."

Cope's (1879) reference to the crayfish is only a

slightly modified version of his 1872 contribution.

Packard (1879) discussed and figured the eyes and

brain of this crayfish, comparing them with those of

an epigean crayfish from Iowa. The figure of the entire crayfish is the same as that published in 1871.

Hubbard (1880) simply recorded having collected

"good specimens" of pellucidus in Mammoth Cave.

Joseph (1880), in a brief note, proposed the name

C. typhlobius for a cave crayfish ostensibly inhabiting

a cave in Yugoslavia. The following year (1881) he

referred to the species as both C. coecus and C.

stygius, and in a third paper (1882) compared the

specimen, to which he referred as C. stygius, with

Packard's description of Cambarus pellucidus. Hovey

(1882) utilized a new combination, Cambarus (Orconectes) pellucidus, but contributed no original

information.

Cope and Packard (1881), in discussing Cope's

O. hamulatus, indicated that it and O. pellucidus

probably arose from different species of Cambarus

and Cope contrasted the two species. Semper (1881)

added no additional information, simply stating that

the blind crayfish of Mammoth Cave was well

known!

Leydig (1883) presented a histological study of the

eyes and antennae of specimens of pellucidus from

Mammoth Cave, extending the earlier observations

of Newport and Packard.

Faxon (1884) considered Cope's O. inermis to be

conspecific with Cambarus pellucidus. He cited Joseph's work and indicated that "until a more satisfactory account of this discovery is published, one

may well hesitate to admit the Carniola Cambarus

[typhlobius] into the list."

Wright (1884) reviewed Leydig's findings and compared the "olfactory cones" of O. pellucidus with

those of C. propinquus [ = O. propinquus], reporting

many more in the former.

Faxon (1885a) presented an excellent summary

of, and commented on, the contributions of previous

authors concerning this species. He discussed its affinities, and, in so doing, assigned it to his Group I

(=Procambarus, in part). He considered Cope's O.

inermis a synonym of Cambarus pellucidus from the

Mammoth Cave region and, erroneously, from Wyandotte Cave and a cave in Bradford, Harrison County,

Indiana. He also recorded Joseph's work and quoted

his description of C. stygius in full, lamenting the fact

that a fuller description had not followed. In a later

paper (1885b), Faxon cited a new locality for O.

pellucidus, White Cave, Kentucky.

Underwood (1886), in his list of North American

freshwater crustaceans, presented a partial synonomy

of C. pellucidus and recorded it from Kentucky and

Indiana, following Faxon in considering Joseph's C.

typhlobius and C. stygius doubtful species.

Packard (1888) constitutes the broadest treatment

in existence of the then known troglobitic crayfishes.

He quoted extensively from and summarized the contributions of previous investigators, treating taxonomy,

distribution, food and feeding habits, auditory, optic,

and olfactory senses, ecology, and evolution. While

he considered O. inermis to be a synonym of C.

pellucidus, he relegated Cope's genus to subgeneric

rank and indicated that the two cave species, pellucidus and hamulatus, constituted its members (page

42). In considering the origin of C. pellucidus, he

stated (page 39) that "it either is derived, with C.

affinis [-O. limosus Rafinesque], from a common

ancestor; or .. . what seems more probable, it is a

modification of C. affinis or an allied species, e.g.,

rus'icus. The characteristics which separate C.

pellucidus from C. affinis or C. bartonii or any outof-door species are those which have been induced

by its life in total darkness and the diminution of

its food-supply." Perhaps through a printer's error,

he utilized the combination, "Camtarus (Orconectes)

pellucidus, form inermis" to designate plate-figure

27:5, which is identical to the figure to which he

referred earlier (1871) as Cambarus pellucidus, one

that had been redrawn from Hagen, 1870.

Garman (1889) was impressed by the fact that

C. pellucidus is more closely allied to the Missouri

C. virilis than is C. setosus which, in turn, is more

similar to the Kentucky C. bartonii than is C. pellucidus, and he suggested the derivation of C. setosus

from C. bartonii, concluding (page 236) that, "Such
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

close affinities as exist between C. bartonii and C.

setosus do not permit their separation into different

genera, and the retention of the latter in the genus

Cambarus cannot but be followed by the disestablishment of the genus Orconectes and the return to the

older genus of the two species heretofore included

in the latter."

Faxon (1890) compared the arrangement of the

olfactory setae of this species with that of C. setosus;

he further postulated that "The closer superficial likeness between C. pellucidus and C. hamulatus, belonging to different sections of the genus, than between C.

hamulatus and C. setosus belonging to the same section, may be explained by the longer period of time

during which the subterranean influences have probably been exerted upon the first two species." In the

same year, Rabe indicated that Astacus Cambarus

pellucidus occured in Mammoth Cave and Astacus

Cambarus Stigius [sic] had been found in the caves

of Carniole.

Packard (1890) in his discussion of the effect of

cave life on animals and its bearing on the evolutionary theory referred to the eyes of pellucidus.

Parker (1890) compared the eyes of pellucidus with

those of C. setosus and found those of the former

to be without pigment, shaped like a blunt cone,

lacking facets but possessing an optic ganglion and

nerve, of which the terminus of the latter was "not

discoverable." At the apex of the cone he found a

"lenticular thickening of the hypodermis, in which

there exist multinuclear granulated bodies."

Hovey (1891) stated that pellucidus feeds on

"Crangonyx and other minute Crustacea." Nothing of

significance was added by Ortman (1892), Cunningham (1893), or Stebbing (1893). Packard (1894)

noted that the "eyes of the young are perceptibly

larger in proportion to the rest of the body than in

the adult," and quoted Tellkampf (1844), "the eyes

are rudimentary in the adults, but are larger in the

young." Packard also stated that "the blind crayfish

of Mammoth Cave . . . have, as we have ascertained

by anatomical investigation, degenerate ears, so that

the sense of hearing is with little doubt, nearly, if not

quite obsolete."

Lonnberg (1894 and 1895) contrasted Procambarus acherontis with pellucidus and suggested that

the latter is probably an older species. In the same

year Packard referred to pellucidus but contributed

no additional information relative to it.

Apfelbeck (1895) stated that specimens of an

eyeless Cambarus from the subterranean waters of

Herzegovina were in the Musee de Vienne.

Hay (1896) contrasted his C. pellucidus testii with

the nominate subspecies and reiterated the fact that

C. typhlobius was the only member of the genus

which occurs beyond the limits of the North American continent.

Call (1897) cited specific areas in Mammoth Cave

where pellucidus occurs. Hay, in the same year, presented a synonomy for C. pellucidus, including inermis

as a synonym.

Faxon (1898) referred to Hay's (1893) Indiana

records for pellucidus [ = inermis] and indicated that

the specimens from them, while transitional between

pellucidus and testii, are more like O. inermis (Wyandotte Cave) than like the typical form around Mammoth Cave, pointing out the reduction of spines in

northern specimens.

Eigenmann (1899) on 23 November 1898 collected

in Mammoth Cave a single female carrying young,

the only record, to our knowledge, of a female with

young. In the same year, Hay included pellucidus

in his key to the crayfishes, and, in citing its range,

obviously considered inermis a junior synonym.

Price (1900) is simply a title without text (see

bibliography). Call (1901) contributed no new data

relating to this crayfish. Ortmann (1902) believed

pellucidus to be related to species presently assigned

to the genus Procambarus and treated inermis as a

synonym of pellucidus.

Hay (1902a), in relating observations on pellucidus in Mammoth Cave stated that "when first

observed they were usually on the bottom, resting

quietly with their legs and antennae fully extended.

Unless they were disturbed they would remain in

this position for several minutes, and then with no

apparent reason start off at a rapid gait, move to

another spot and take up the same position," gently

moving the antennae to and fro. When disturbed,

they move "slightly about in various directions" before swimming away, but "there seemed to be no

ability on the part of the animal to select a safe

haven of refuge from a distance." When cornered,

they would rise high on their forelegs, waving their

chelae in the direction of danger. In drying pools,

they burrowed or crawled under a stone where, he

concluded, they were vulnerable to the cave rat.

Disturbances at the surface of the water seemed not
NUMBER 10 5 63

to frighten the animals, and he believed that "senses

of sight and hearing have entirely disappeared." On

the basis of his observations he concluded that copulation occurs in the early fall and stated that the

females "are said" to lay their eggs during the winter.

In a later paper (1902b), Hay contrasted pellucidus

and C. hamulatus and stated that ". . . C. pellucidus

and C. acherontis [probably Procambarus I. lucifugus

(Hobbs)], which are very dissimilar in general appearance, are closely related."

Steele (1902), Eigenmann (1903), and Harris

(1903a) contributed no additional information about

this crayfish, but Harris (1903b) summarized the

conclusions and observations of others.

Ortmann (1905a, 1905b) expressed the opinion

that pellucidus was more closely related to the members of the "fourth group" than to those of the

"first," and assigned it to his subgenus Faxonius

[ = Orconectes], defending this opinion on morphological and zoogeographic grounds, and placed it in

the most primitive section of the subgenus, the

Limosus Section. Bell (1906) simply reviewed the

observations made by others on the sense organs.

The references to pellucidus by Banta (1907),

Graeter (1909), and Pearse (1910) include summaries or observations made by others cited above.

Osborn (1912) added no original data. Hovey*s

(1912) contribution consists of photographs of a

male and female pellucidus together with the statement that the species "feeds on aquatic Crustacea

which it deftly extracts with its pincerlike claws from

under flat stones.*1

Faxon's (1914) checklist included O. inermis as a

synonym of C. pellucidus, and in it C. stygius, C.

coecus, and C. typhlobius are cited as "doubtful

species."

The account of the eye by Spurgeon (1915) is

based on observations of O. i. inermis, and he made

no additional contribution to our knowledge of O.

pellucidus. Pratt (1916) and Ortmann (1918) included pellucidus in their keys but contributed no

new data. Ortmann (1931) stated that C. sloani and

C. indianensis [both now assigned to the genus

Orconectes] are the species most closely allied to

pellucidus and "probably are the last remnants of

the surface-stock from which C. pellucidus descended."

The following based their statements about pellucidus on data cited above or on other that are essentially identical: Garman (1924), Spandl (1926),

Stiles and Hassall (1927), Chappuis (1927), Creaser

(1931, 1932), Bolivar and Jeannel (1931), and Fage

(1931).

Stammer (1932) expressed the opinion that Joseph's C. stygius was identical with Astacus fluviatilis

[ = Astacus astacus Linnaeus].

The observations of Giovannoli (1933a, 1933b)

on pellucidus in Mammoth Cave are essentially identical to those of Hay and others. "Usually it is

observed at rest but occasionally it is seen walking

along as if it knew where it was going. They seem

to lack entirely the inclination so characteristic of

above-ground species to retreat under rocks and

planks. They pay no attention to such shelters either

when undisturbed or when frightened. When merely

suspicious of danger they walk slowly towards deeper

water, but if startled they swim blindly backward

just as any crayfish does." Wolf (1934-1938) contains

no new information about pellucidus.

Turner (1935), in recording aberrant secondary

sexual characteristics in crayfishes, reached the same

conclusion as did Ortmann that the presence of hooks

on the ischia of the fourth pair of pereiopods in the

male represents a derived, rather than primitive, condition. Park (1938), concerned with periodicity of

activity in various animals, found pellucidus to be

arhythmic. Fleming (1939) and Bouvier (1940) made

no original contributions, and Rhoades (1941) simply

compared pellucidus pellucidus with his C. p. australis.

Park et al. (1941) reported pellucidus to be photonegative and presented data to indicate an arhythmic

activity pattern. Neither Dearolf (1942) nor Van

Straelen (1942) added to our knowledge of the

species. Jackson (1942) gave a brief account of the

habits of this crayfish and reported having kept an

adult alive in an aquarium for a month.

Hobbs (1942a) assigned pellucidus to the resurrected genus Orconectes, pointing out that Faxonius

of previous authors is a synonym of Orconectes. Balss

(1944 and 1955) contributed no additional information to our knowledge of pellucidus and was unique

in utilizing the combination Cambarus (Cambarus)

pellucidus.

Rhoades (1944) presented a description of his new

subspecies, O. pellucidus packardi, comparing it with

the nominate subspecies and O. p. australis. He summarized their ranges and assigned the three subspecies

of pellucidus and three additional epigean species to

his "Group rafinesquei." Hobbs (1948a) figured the
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

first pleopods of the males of O. pellucidus and related species, questioned Rhoades' division of the

Limosus Section, and expressed the opinion that

inermis represented a species distinct from pellucidus.

Jeannel and Henrot (1949) reported a new locality

for the species, Cave City Cave, Barren County,

Kentucky.

Bott (1950), concerned with Joseph's species reported from Carniole, did not reach a conclusion as

to its identity. Pennak (1953) added no original information about pellucidus, and his figure 286 is an

illustration of O. inermis testii. Villalobos (1953,

1955) considered the existence of Joseph's C. typhlobius as perhaps a relict of the crayfishes "que en

otra epocas poblaron el suelo europeo."

Dearolf (1953) cited dates of observations of this

crayfish in three Kentucky caves.

Eberly (1954) stated that the cave crayfishes (including O. pellucidus) "occupy the highest niche in

the biosystem of the cave." In 1958 and 1960, he

summarized what he considered to be the adaptations

of the crayfishes to a cave environment, agreed with

his predecessors on their antiquity, and gave a broad

generalization regarding their origin. Croizat (1958)

added nothing directly to our knowledge of O.

pellucidus; however, he consistently used Gambarus

instead of Cambarus for the American genus.

Rhoades' (1959) contribution included a history

of the nomenclatural changes among the troglobitic

Orconectes and an interpretation of the relationships

of pellucidus, inermis, and testii, concluding that the

latter is a synonym of inermis and that inermis freely

intergrades with pellucidus in southern Indiana,

thus recognizing O. pellucidus pellucidus and O. p.

inermis.

Cole (1959) and Wells (1959) presented no original information about pellucidus nor did Hobbs and

Barr (1960), Nicholas (1960), or Barr (1961). Hobbs

(1959) included O. (O.) pellucidus in his key to the

crayfishes of the United States, indicating that four

subspecies occurred from Alabama to Indiana.

Brown (1961) reported that a re-examination of

the data of Park et al. (1941) had "indicated a

statistically significant . . . 24-hr, rhythm of activity

to be present . . . with minimum activity about 9

a.m. and maximum about 7:00 p.m." He further

stated that members of pellucidus "have no eyes."

Creaser (1962) discussed the affinities of O. pellucidus, maintaining that only pellucidus and lancifer

should be assigned to the genus Orconectes. Rhoades

(1962) presented a diagnosis of O. p. pellucidus and

stated that it is "widely distributed in the solution

caverns between the Cumberland River and the

counties of southern Indiana" where they "are generally seen in the edges of a clear stygian pools or

hiding under rocks." He further postulated the origin

of the species from a stock which had "its origin in

the Ozarkian Highlands in the late Miocene."

Fitzpatrick (1963), in presenting a background for

elevating the subgenm Faxonella to generic rank, reviewed Creaser's (1962) remarks about O. pellucidus.

Barr (1964) contributed no new data or hypotheses

concerning the species. Holthuis (1964) reviewed the

history of Joseph's three crayfish names and presented a convincing conclusion that C. typhlobius is

a synonym of A. pellucidus Tellkampf, and that Joseph's specimens were simply mislabeled members of

TellkampPs species. C. coecus Joseph was declared

a nomen nudum, and C. stygius Joseph a synonym

of C. typhlobius, becoming thereby a synonym of A.

pellucidus. Vandel (1964) presented an excellent account of the various contributions of others to our

knowledge of the biology of the species. In the same

year, Wolfe and Cornwell reported that although

small amounts of B-carotene and lutein were present

in O. p. pellucidus, "astaxanthin, the principal carotenoid of most Crustacea" is absent, supporting

their conclusion that "pigmentation is dependent on

the amount of carotenoid in the diet rather than on

the presence of light." It is also suggested that this

crayfish lacks, and perhaps never had, "the ability

to oxidize dietary carotenoids."

Mohr (1964) presented a photograph of O. pellucidus and made the statement that it "may live to

twice the age of surface kin." Poulson (1964) recounted the observations of Barr (personal communication) that breeding (ovulation?) in O. pellucidus

"precedes low water by one to two months, and high

water, with maximum organic inwash, by four to

six months. Further, he stated that "the restriction

of .. . young crayfish to springs and backwaters must

be related to current, small food-particle size, clay,

or other factors because these habitats and adult habitats shows no differences in CO2 or carbonate alkalinity." Discussing light, Poulson stated: "Light stimulation of the brain area, reduced eye stalk, or 6th

abdominal ganglion results in negative reaction to

light," and he reported that a "kinetic component
NUMBER 10 5 65

is found . . . [in] O. pellucidus . . . since they turn

away when illuminated on one side."

Frey's (1965) remarks relative to O. pellucidus

were largely condensed from Rhoades' (1944, 1959)

conclusions and postulates. Jegla et al. (1965) compared the maximum lengths of individuals in populations of troglobitic Orconectes (O. i. inermis, O.

pellucidus, and O. australis) and found a difference

between them of 20 mm, the same difference in

length at which they reached sexual maturity, as

well as one "in inflection point for cheliped allometric

growth. These variations may be a result of different

levels of food supply and interspecific competition.

They do not appear to be influenced by two-fold

variation in population size, eight-fold variation in

population density, or ten-fold variation in available

habitat." Predation by fishes was postulated to reduce

population density but seemed to have no effect on

maximum length attained at sexual maturity. Length

was believed to be associated with available food

and perhaps influenced by inter-specific competition.

Hart and Hart (1966) reported that O. pellucidus

is infested with the following entocytherid ostracods:

Dactylocythere ungulata (Hart and Hobbs, 1961),

Sagittocythere barri (Hart and Hobbs), and S. stygia

Hart and Hart. Jegla (1966) reviewed previous

references to molting and reproduction in this species.

Barr (1966) referred to Tellkampfs description of

this species.

Mohr and Poulson (1966) presented a drawing of

O. pellucidus, and stated that "Scientists now believe . . . that the ancestors of our commonest cave

crayfish, Orconectes pellucidus, were isolated in caves

at the beginning of the Pleistocene epoch."

Barr (1967a, 1967b) noted the northern boundary

of the range of O. pellucidus, indicating its occurrence

in "the south Pennyroyal fauna," and included a

photograph of a living first-form male. Neither Fitzpatrick (1967) nor Hobbs (1967a) added to our

knowledge of the species, and Hobbs (1967b) only

proposed that "at least some of the similarities" between Procambarus pecki and O. pellucidus "are due

to convergence."

Thompson (1967) observed that O. pellucidus,

in contrast to epigean crayfish maintained in an

aquarium, "rather than folding its antennae back in

a streamlined position as the epigeal form did while

walking through vegetation, the cave form held them

forward for tactile cues, and consequently often became tangled in the litter. It also lacked cryptic

responses when startled and spent little time hidden

under rocks, compared to the epigeal form." Thompson did not cite the source of his O. pellucidus, and

it is possible that his animal was a member of one

of the subspecies of O. inermis.

Jegla and Poulson (1968), reviewing the work of

Park et al. (1941) and that of Brown (1961), concluded that the data of Park et al. cannot "be used

to distinguish between endogenously and exogenously

controlled rhythms." They found "evidence of a

circadian clock mechanism" with "average periods

of 26, 27, and 34 hours" in O. pellucidus [=0 .

inermis inermis and O. inermis x testii\.

Barr (1968) expressed the opinion that the steepening of stream gradients in the Pennyroyal and

Cumberland plateaus from Alabama to Indiana at

the close of the Pliocene or early Pleistocene was

primarily responsible for the extinction of the ancestors of the troglobitic Orconectes, and that the

quieter streams of caves served as refuges for the

evolving troglobites. He also presented a graph redrawn from the 24-hour activity data of Park et al.

(1941), and criticized Brown (1961) for not utilizing

all of Park's published data.

Barr and Kuehne (1971) discussed the role of this

crayfish in the Mammoth Cave ecosystem.

DIAGNOSIS.—Albinistic; eyes reduced and without

pigment; rostrum with marginal spines or tubercles

delimiting base of acumen, margins subparallel, convex or converging, upper surface concave and without median carina; postorbital ridges usually terminating cephalically in spines or tubercles; hepatic

area with or without two to many spines; at least

one, often several cervical spines or acute tubercles

present; areola 3.7 to 6.0 times longer than broad

and constituting 34.1 to 42.2 percent of entire length

of carapace; chelae not conspicuously setose but with

ciliated tubercles, mesiodorsal surface of palm with

several irregular rows of tubercles; hooks on ischiopodites of third and fourth pereiopods. First pleopod

of first-form male with greatest cephalocaudal diameter of pleopod more than twice that immediately

proximal to base of central projection, always terminating in only two elements; non-corneous mesial

process broad basally, triangular, directed distally

and slightly caudally and extending considerably beyond distal extremity of central projection, frequently

almost obscuring latter in caudal aspect (Figure 14;) ;
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

central projection small, corneous, flattened in cephalocaudal plane, and directed distad; slight swelling

at cephalic base of central projection. Annulus ventralis only slightly broader than long, highest along

median portion, and without shelf-like rim along

caudal margin (Figure 14/).

Topotypic Male, Form I: Body (Figure 14c, h)

subovate, depressed. Abdomen narrower than thorax

(11.0 and 12.6 mm in widest parts, respectively).

Width of carapace greater than depth in region of

caudodorsal margin of cervical groove (12.5 and

10.2 mm). Areola moderately broad (4.5 times

longer than wide), with about 6 widely spaced,

minute punctations across narrowest part. Cephalic

section of carapace 1.6 times as long as areola;

length of areola 38.6 percent of entire length of

carapace. Rostrum approximately twice as broad

as long, excavate, and with length of acumen subequal to width of rostrum at base; cephalic extremity

reaching far beyond antennular peduncle and slightly

beyond antennal peduncle; margins not swollen,

only slightly elevated, and with prominent corneoustipped marginal spines at base of acumen; upper

surface with evenly spaced minute setiferous punctations; subrostral ridges very weak and evident in

dorsal aspect along basal fourth of rostrum.

Postorbital ridges relatively weak, short, with shallow dorsolateral grooves, and terminating cephalically in prominent corneous-tipped spines. Suborbital

angle lacking. Branchiostegal spines acute. Six and

eight cervical spines present on right and left sides

of carapace, respectively. Carapace punctate dorsally

and granulate laterally; hepatic area with about 12

spines of various sizes together with several tubercles.

Abdomen slightly longer than carapace (32.0 and

30.0 mm). Cephalic section of telson with 2 strong

spines in each caudolateral corner, mesial ones

movable.

Epistome (Figure 14g) subtriangular, with almost

straight cephalolateral margins and rounded basal

lateral extremities; ventral surface subplane with

crowded minute setiferous punctations. Eyes much

reduced, completely hidden under rostrum in dorsal

aspect and extending only halfway between margin

of orbit and marginal spines of rostrum. Antennules

of usual form with prominent spine near distoventral

extremity of basal segment. Antennae extending caudally at least to caudal margin of telson. Antennal

scale (Figure 14i) broadest distal to midlength, about

2.2 times longer than broad; outer thickened portion

much narrower than lamellar area and terminating

in prominent corneous-tipped spine. Third maxilliped extending almost to end of basal third of

ultimate podomere of peduncle of antenna.

Chela (Figure 14/) slender and little inflated;

mesial margin of palm 1.3 times longer than width

of palm; palmar area tuberculate, with fine plumose

setae at distal bases of tubercles; ventral surface with

prominent corneous-tipped tubercle opposite base of

articulation of dactyl; tubercles along mesial surface

arranged in sublinear series with 12 tubercles in

mesialmost row; distal two-thirds of lateral margin

of palm and proximal fourth of fixed finger weakly

costate. Fingers not gaping; dorsal and ventral surfaces of both with median longitudinal ridges flanked

by setiferous punctations; opposable margin of immovable finger with row of 14 rounded corneous

tubercles, fourth from base largest; same margin with

large tubercles below row and between eighth and

ninth tubercle; row of minute denticles broken by

8 basal tubercles and continuous distally to base of

corneous tip of finger; opposable margin of dactyl

with row of 18 tubercles, fourth and fifth from base

subequal in size and larger than others in row, row

of minute denticles extending almost entire length

of finger and interrupted by tubercles; mesial surface

of dactyl with tubercles along proximal half and

punctations along distal. Carpus longer than broad

and tuberculate; mesial surface of right with 1 (left

with 2) large spine and more proximal moderately

large acute tubercle; distoventral margin with 2

prominent spines, 1 adjacent to ventrolateral articulation with propodus, 1 situated more mesially. Merus

tuberculate except proximolaterally, upper surface

with 4 or 5 corneous-tipped spines near distal end,

and ventral surface with lateral row of 14 spikelike

tubercles and mesial one of 13, few additional tubercles flanking latter two rows. Ischium with row

of 6 small tubercles on opposable margin, other

smaller ones lateral to row and on outer surface.

Ischia of third and fourth pereiopods (Figure 14&)

with simple hooks, those on third conspicuously

larger than those on fourth but neither extending

proximad of distal extremity of corresponding basis.

Coxae of fourth pereiopods with heavy, caudomesially

projecting prominences; coxae of fifth without prominences except for small mesioventral projection on

ventral border of rim surrounding basal portion of

phallic papillae.

First pleopods (Figure 14a, e, j , k) symmetrical,
NUMBER 10 5 67

FIGURE 14.—Orconectes pellucidus: a, mesial view of first pleopod of topotypic male, form I;

b, mesial view of first pleopod of topotypic male, form II; c, dorsal view of carapace of topotypic male, form I; d, lateral view of first pleopod of topotypic male, form II; e, lateral view

of first pleopod of topotypic male, form I: /, dorsal view of distal podomeres of cheliped of topotypic male, form I; g, epistome of topotypic male, form I; h, lateral view of carapace of topotypic male, form I; i, antennal scale of topotypic male, form I; ;, caudal view of first pleopods

of topotypic male, form I; k, ventral view of caudal thoracic region of topotypic male, form I;

/, annulus ventralis and portion of sternum of topotypic female.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

reaching coxae of third pereiopods when abdomen

is flexed and situated in shallow, broad sternal arch;

tip ending in two parts as described in Diagnosis.

Basal podomere of uropods with 2 prominent

spines, each overhanging lateral portions of corresponding ramus and small but conspicuous "telsonic

condyle" projecting dorsomesially from its mesiodistal

angle. Inner ramus of telson without well-defined

longitudinal median carina and spine but with spine

on distolateral angle; outer ramus with row of spines

on distal margin of proximal section, lateral 1 or 2

longer than others.

Topotypic Female: Differing from holotype in

following respects: Acumen not quite reaching distal

end of antennal peduncle; 5 and 6 cervical spines

on left and right sides of carapace, respectively;

hepatic area with only 7 well-defined spines; projecting portion of epistome shorter with cephalolateral margins slightly convex; third maxillipeds not

quite reaching distal podomere of antennal peduncle;

opposable margin of immovable finger of cheliped

with row of 15 tubercles, fifth from base largest, that

of dactyl with 18, sixth from base largest; upper

surface of merus with 5 and 2 spines on left and right

chelipeds, respectively, and ventrolateral margin with

11 and ventromesial margin with 17 and 12 on right

and left; ischium with 7 irregularly arranged acute

tubercles. First pleopods uniramous and reaching midlength of annulus when abdomen is flexed.

Annulus ventralis (Figure 14/) shallowly situated

in sternum and not firmly fused to sternal plate immediately cephalic to it; outline subovate, only

slightly broader than long, with median portion

elevated ventrally and forming broad arc having its

highest segment distinctly caudal to midlength;

arched portion bearing only faint longitudinal grcove,

ending caudally on vertical wall, plateau-like rim

totally lacking; sinus very short, originating dextral

to caudal end of longitudinal groove, following gentle

arc across median line, passing dorsally and ending

on caudal wall sinistral to median line.

Topotypic Male, Form II: Differing from holotype in following respects: areola with only 4 punctations across narrowest part; only 4 cervical spines

and only 5 hepatic spines on each side of carapace;

opposable margin of immovable finger of cheliped

with row of 10 tubercles, corresponding margin of

dactyl with 12; merus of cheliped with 3 spines on

upper distal surface, 8 in ventrolateral row and 13

in ventromesial row; ischium with row of 5 tubercles.

Hooks on ischia of both third and fourth pereiopods

much reduced, those on fourth to low tubercles;

protuberances on coxae of fourth pereiopods only

slightly reduced in size.

First pleopods (Figure 146, d) markedly similar

to those of holotype; however, central projection

rounded, poorly defined, and non-corneous.

SIZE.—The largest specimen available to us is a

first-form male from Crystal Lake in Mammoth

Cave, Edmonson County, Kentucky, which has a

carapace length of 39.4 mm. The smallest first-form

male, carapace length 22.0 mm, was collected in

Cooks Cave, Logan County, Kentucky.

MEASUREMENT S (in millimeters).—Orconectes

pellucidus:

Topotypic Topotypic Topotypic

male female male

Form I Form II

Carapace :

Height 10.2 12.8 9.8

Width 12.6 16.2 11.0

Length 30.0 36.8 27.2

Rostrum:

Width 4.2 5.2 3.9

Length 8.5 10.0 8.2

Areola:

Width 2.6 3.0 1.9

Length 11.6 14.7 10.2

Chela:

Length of inner margin

of palm 8.4 10.2 7.3

Width of palm 6.3 7.8 4.8

Length of outer margin

of chela 16.6 19.5 25.6

Length of dactyl 9.2 10.6 14.6

TYPES.—Holotype , Zoologisches Museum der

Humboldt-Universitat, Berlin (o*I).

TYPE-LOCALITY. — Mammoth Cave, Edmonson

County, Kentucky.

SPECIMENS EXAMINED.—Specimens from Kentucky

and Tennessee were examined as follows:

KENTUCKY: Barren County: (1) Cave City (=Railroad)

Cave, in town of Cave City, 16" I, T.C.B. and T.G.M.,

VIII/23/65. (2) Diamond Cave, 2 miles N of Park City

on State Rte. 225, 1 $ I, T.C.B., S.B.P., J.R.H., and J. F.

Fitzpatrick, Jr., VI1/25/64. (3) Twyman Cave, 2 miles N

of Hiseville, 2 $ II, 2$ , R.M.N., IX/11/65. Christian

County: (1) Glover's Cave, 4 miles SW of Trenton, 2 $ II,

2 5 , T.C.B., VI/9/57. Edmonson County: (1) Mammoth

Cave, Echo River, 1<JI, 1 $ II, 39 , 7j ,$, 5j$ , W.P.H.,

date?; l^H , 3$ , L.H., VIII/30/39; 49 , Raymond Nelson, VII/28/58. (2) Mammoth Cave, Roaring River, 5 31,
NUMBER 105 69

1 $ II, 5 9 , 2j 3 , 4j 9, W.P.H., XIII/28/01; 1 $ II, C.W.H.,

XI/29/63. (3) Mammoth Cave, Styx River, 131, T.C.B.,

IV/28/62; 531, 13II, 49 , T.C.B., V/13/62. (4) Mammoth Cave, Lake Lethe, 3,51, 13II, T.C.B., X/25/61;

1 3 I, T.C.B. and R.A.K., X/l 1/63; 5 9, T.C.B. and R.A.K.,

XI/25/61. (5) Mammoth Cave, Crystal Lake, 131,

T.C.B., 11/22/64; 13II, T.C.B., IV/F/64. (6) Mammoth Cave, 131, Peter Parker, ?/?/1858; 2j3 , Leonard

Giovannoli, VI1/19/29; 13II, 3 9, 1J9, L.G. VIII/

6/29; 13II, 4 9, V. E. Shelford, VII/4/42; 2j9, ?,

VIII/28/41; 131, Kjell Sandved, VI/?/67; 131, iSU)

2 9 , lj'3, 1J9, W.P.H., date?; 19 , M. E. Mann,

date?; Ij3 , Ellsworth Call, date?; 13H, 39 , coll. and

date ( ?). (7) Blowing Spring Cave in Mammoth Cave

National Park, 19 , L.H., XII/1/56; 19 , O.D. Lee, date?.

(8) Cedar Sink Cave, in Mammoth Cave National Park,

1 3 II, T.C.B., X/29/66. (9) Stream in Long Cave, Mammoth Cave National Park, 13II, 19 , L.H., 1/5/56. (10)

Stillhouse Hollow Cave, Mammoth Cave National Park, 1 9 ,

L.H., XI/17/56. (11) White Cave, 3 9, John E. Younglove, date?. (12) Floyd Collins Crystal Cave, in Mammoth

Cave National Park, 19 , Carl Melton, V/28/60. Hart

County: (1) Bald Knob Cave, 2 miles W of Hardyville,

13II, 19 with eggs, R. M. Norton, IX/11/65, (2) Buckner Hollow Cave, 1.2 miles SE of Hinesdale, 231, 39 ,

J.R.H. and T.C.B., VIII/30/63. (3) Horse (-Hidden

River) Cave, in town of Horse Cave, 2 9, 1 j $, 1 j 9, E. P.

Creaser and M. Becker, VIII/28/29; 19, E.P.C. and B.,

V/13/30; 131, 13II, 19 , Ij3 , L.H., VIII/30/39; 1311,

1 9 , Nat. Spel. Soc, VIII/29/41. (4) Stream in cave, 2

miles SW of Northtown, 131, 19 , L.H., 111/23/57. (5)

Mammoth Onyx Cave on St. Rte. 335, 19 , R.R., VII/

16/39. Logan County: (1) Cooks Cave, 1 mile E of

Adairville, 2 31, 19, T.C.B., VI/25/57. (2) Mud River

Cave, 4 miles E of Russellville, 3 31, 3 3II, 3 9, J.R.H. and

T.C.B., VIII/13/65. Trigg County: (1) Cool Spring Cave,

on Sinking Fork Creek, 2 31, S.B.P. and R.M.N., IV/19/64;

431 , 13II, 5 9, S.B.P., VI/28/65. Warren County: (1)

By-Pass Cave, Bowling Green, 1 9, K. Haas, VI/?/55. (2)

Lost River Cave, 3 miles S of Bowling Green, 15j9, D. H.

Puckett and Herbert Shadowen, VIII/7/64. (3) Pruitt

Saltpeter Cave, 0.8 mile SE of Anna, 2 31, 13II, 39 ,

T.C.B., C.W.H., and D.G.H., XI/30/64.

TENNESSEE: Montgomery County: (1) Sink Hole Cave

on Austin Peay College Farm at Clarksville, 43II, 3 9,

D. A. Etnier, XI/22/68. (2) Bellamy Cave, 3.0 miles S of

Oakwood, 1 3II, 19 , Merlin Tuttle, VIII/29/69.

RANGE.— (Figure 1) The range of Orconectes pellucidus extends southwestward from Hart County to

Trigg County, Kentucky, and Montgomery County,

Tennessee.

VARIATIONS.—Most stable among the features examined in Orconectes pellucidus are perhaps the

secondary sexual characteristics of the adults. The

first pleopods of the male uniformly exhibit a markedly slender (in lateral aspect) distal portion and a

mesial process which extends distally much beyond

the tip of the central projection. The hooks on the

ischia of the third and fourth pereiopods vary in

size, particularly that of the fourth, but are always

clearly evident in the first-form male. Those on the

fourth may be much smaller than those on the third,

and neither greatly swollen; occasionally only the

hook on the third is markedly robust; but in some

males, the hooks on the two pereiopods are subequal

in size, and both swollen. The annulus ventralis,

while showing minor variations in outline and contour of the ventral surface, always has a rather steep,

evenly curved caudal surface and lacks the caudal

plateau-like rim characteristic of the other troglobites

of the genus.

Whereas, there is comparatively little variation in

the secondary sexual characteristics of O. pellucidus,

the carapace and chela show a wide range of differences. Most of those noted in the carapace, including

meristic ones, are associated with the degree of development of the spiny elements. In general, among

those individuals that have a number of strongly

developed cervical and hepatic spines, the marginal

spines of the rostrum, the acumen, the postorbital

spines, and that on the antennal scale are longer

than those in specimens having reduced, and usually

fewer cervical and hepatic spines (see Figure 15M,

v). Inasmuch as, through convention, the length of

the carapace is given as the distance between the tip

of the rostrum and the midcaudodorsal margin of

the carapace, it is obvious that a spiny (hence, having

a longer rostrum) individual of approximately the

same size as one that is relatively aspinous has a

proportionately greater carapace length. The longer

rostrum is also reflected in differences noted in the

relative length of the areola, which is expressed in

the percentage of the total length of the carapace;

that is, an animal having a longer rostrum has a

proportionately shorter areola. Comparing the same

two illustrations mentioned above, the areolae constitute 41.4 and 38.9 percent of the carapace length,

respectively, although there is only 0.1 mm difference

in the actual length of the areolae of the two. Thus,

the relative lengths of the areola are not so much

determined by variation in its length as by the variation in the length of the rostrum, and the latter

seems, for the most part, to be correlated with the

greater development of the acumen (essentially a

spine). Furthermore, it seems at least possible, on
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 10 5 71

a \

^ /

) I

FIGURE 16.—Cephalic portion of carapace of Orconectes pellucidus illustrating variation in

spination (all localities in Kentucky): a, Mud River Cave, Logan Co. ( $1); b, c, Cooks Cave,

Logan Co. ( <J I, ? ); d, Glovers Cave, Christain Co. ($);« , Cool Spring Cave, Trigg Co.

the basis of our limited data, that the aspinous condition may be correlated with more strongly lotic

habitats.

The loss and regeneration of appendages among

troglobitic crayfishes is commonplace, and the regeneration is often so complete that frequently it is

difficult to determine whether or not a given cheliped

is the original one or a replacement. The occasional

occurrence of a comparatively robust chela on a

large male elicits the conclusion that few individuals

reach adulthood without having regenerated at least

one member of this pair of appendages. Even with the

tremendous range of variation that has been observed

in them, an elongate, comparatively narrow, tuberculate palm is characteristic of all of the troglobitic

species of the genus. We have been unable to discover

FIGURE 15.—Cephalic portion of carapace of Orconectes

pellucidus illustrating variation in spination (all localities in

Kentucky): a, Horse Cave, Hart Co. (ill) ; b, Buckner

Hollow Cave, Hart Co. ( 9 ) ; c, Cave 2 mi. W. of Northtown, Hart Co. ( $ I) ; d, Horse Cave, Hart Co. ($1); e-k,

Mammoth Cave, Edmonson Co. (9 , 9 , 6*11, d* II, 6*1,

9 ) ; /, Long Cave, Edmonson Co. ( 9 ) ; m, Cedar Sink,

Edmonson Co. ( 9 ) ; n, White Cave, Edmonson Co. ( 9 ) ; o,

Stillhouse Hollow Cave, Edmonson Co. (9) ; p, By-Pass

Cave, Warren Co. ( 9 ) ; q, Pruitt Saltpeter Cave, Warren

C o

- (<5I); r, s, Lost River Cave, Warren Co. (9 , 9) ; *,

Diamond Cave, Barren Co. (31) ; u, Cave City Cave, Barren Co. ( $ I) ; v, Twyman Cave, Barren Co. ( $ I) ; w, Mud

River Cave, Logan Co. ( 9 ).

any consistent feature of the cheliped which is unique

to one of the species.

Despite the variations in spination, no particular

pattern or degree of development of the spines seems

to be correlated with a major portion of the range

of the species. In some localities, all the specimens

collected have been very spiny, and in others, the

only traces of spines on the carapace are limited to

a single, small cervical spine and one or two adjacent

small tubercles, weak marginal spines on the rostrum,

and a broad, short acumen. Yet in other localities,

while one of the two types seem to predominate, both

are represented in the population. Further, to make

more difficult correlation with a portion of the range

of the species or with ecological factors, there is

considerable variation in the degree of development

of the individual spines, so that two specimens from

the same or different localities may have virtually an

identical number of spines, but in one they are very

prominent and in the other comparatively weak.

From the eastern portion of the range in Barren,

Edmonson, and Hart counties, we have 131 specimens from 15 localities. In Hart County, only in

Horse Cave are there specimens which might be

classified as spiny (Figure 15a), but even there, only

3 of the 15 specimens could be so designated. The

others have very weak spines, or the hepatic spines

are reduced or lacking (Figure \bd). Among them

the areola constitutes from 38.5 to 40.6 percent of

the length of the carapace. A single specimen from
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Floyd Collins Crystal Cave has a large number of

small spines, and its areola constitutes 38.6 percent

of the carapace length. From Buckner Hollow Cave

(Figure \5b) are 5 specimens, of which 3 are almost

smooth, and only 1 moderately spiny; their areolae

range from 39.1 to 41.1 percent. The remaining 5

specimens from Bald Knob Cave, Mammoth Onyx

Cave, and the unnamed cave two miles southwest of

Northtown (Figure 15c) have few poorly developed

spines, and the areola ranges from 39.6 to 41.7

percent of the carapace length.

In Edmonson County, the range of variation in

the 95 specimens from Mammoth Cave is almost as

great as that from all the other caves in the county

combined (Figure \5e-k), and the areola ranges

from 38.0 to 41.6 percent of the total length of the

carapace. Our 3 specimens from White Cave are

comparatively spiny (Figure 15n) and their areolae

constitute 38.9 to 40.4 percent of the carapace

length. In the remaining localities, the 5 available

specimens are relatively aspinous (Figure 15/, m, o),

and their areolae vary from 39.7 to 42.4 percent of

the carapace length.

The collection from Twyman Cave, Barren County,

consists of 3 very spiny individuals (Figure 15y),

and 1 which is almost spineless, resembling the 3

specimens from Diamond Cave and Cave City Cave

(Figure \5t, u). In the specimens from the former

cave the areola ranges from 38.5 to 40.5 percent of

the carapace length, and in the latter two, 41.8 and

41.4 percent, respectively.

In the central portion of the range, in Warren

and Logan counties, a similar range of variation

exists. Among 6 specimens from Pruitt Saltpeter

Cave and 15 specimens from Lost River Cave, both

moderately spiny and aspinous (Figure I5q-s) individuals are present in which the areola comprises

38.9 to 41.3 percent of the length of the carapace.

Our single specimen from By-Pass Cave is aspinous

(Figure 15p), with the areola constituting 40.5 percent.

In Logan County, the 3 specimens from Cooks

Cave are very weakly spined (Figure 16fr, c), with

areolae constituting 38.1 to 39.9 percent of the carapace length. In contrast, the 9 specimens from Mud

River Cave are more spiny (Figure 16a) and, except

for the largest first-form male, which has an areola

comprising 40.6 percent of the carapace length, they

exhibit a range of 37.6 to 39.8 percent.

In the three caves in the southwestern part of the

range there is essentially a repetition of what exists

elsewhere. Our 4 specimens from Glover Cave, Christian County, are relatively smooth (Figure \6d) and

have areola constituting 39.6 to 41.6 percent of the

carapace length. The 12 specimens from Cool Spring

Cave, Trigg County, are among the most spiny specimens (Figure 16<?) we have observed, and their long

acumen is reflected in the relative areola length

which ranges from 37.6 to 38.7 percent of the carapace length. None of the 9 specimens from Montgomery County, Tennessee is so spiny as those from

Cool Spring Cave, and considerable variation exists

among them, as might be assumed by the range of

their relative areola length, 39.3 to 44.2 percent of

the carapace length.

LIFE HISTORY NOTES.—First-form males have been

collected each month of the year except January,

September, and December. Collections made during

these months consist of two, six, and one specimens,

respectively; thus there is little reason to doubt that

breeding males occur throughout the year. The only

female carrying eggs that we have examined was

collected on 11 September 1965 in Bald Knob Cave,

Hart County, Kentucky. No female carrying young

has been reported since Eigenman (1899) collected

a specimen on 23 November 1898.

9

juv. $

juv. $

ovig. $

Totals

SEASONAL COLLECTIONS OF SPECIMENS EXAMINED

Not

Sex Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. known Total

1 3

1

-

-

-

-

4

5

1

6

-

-

-

12

7

3

9

-

-

-

19

1

-

1

2

-

-

4

9

10

15

4

23

-

61

-

3

2

-

-

1

6

4

2

-

-

-

_

6

2

6

12

-

_

_

20

_

_

1

_

_

_

1

6

7

20

9

6

_

48

39

34

68

15

29

1

186
NUMBER 105 73

Literature Cited

Anonymous

1843a. Die Mammuthhohle. Das Ausland, number 13,

pages 19-20.

1843b. [Notice of Eyeless Crayfish and Eyeless Fish from

Mammoth Cave, Kentucky.] Proceedings of the

Academy of Natural Sciences of Philadelphia,

1:175.

Apfelbeck, M. Victor

1895. Sur la faune des cavernes de Bosnie et d'Herzegovine. Spelunca, Bulletin de la Societe de

Speleologie, number 1, pages 23-24.

Balss, Heinrich

1944. Decapoda, III Morphologischer Teil. Pages 321-

480, Textabb. 361-516, in H. G. Bronn's Klassen

und Ordnungen des Tierreichs, Band V. Abteilung

I, Buch 7, Lieferung 3.

1955. Decapoda, VI Okologie. Pages 1283-1367, Textabb. 1043-1069, in H. G. Bronn's Klassen und

Ordnungen des Tierreichs, Band V, Abteilung I,

Buch 7, Lieferung 10.

Banta, Arthur M.

1905. The Fauna of Mayfield's Cave. Science, new

series, 21 (544) =853-854.

1907. The Fauna of Mayfield's Cave. Carnegie Institution of Washington, Publication 67, 114 pages.

Barr, Thomas C, Jr.

1961. Caves of Tennessee. Bulletin of the Tennessee

Department of Conservation and Commerce, number 64, 567 pages, 150 figures.

1964. The Fauna of Mammoth Cave: A Study in Community Composition and Evolution. Bulletin of

the National Speleological Society, 26(2):78-79.

1966. Evolution of Cave Biology in the United States,

1822-1965. Bulletin of the National Speleological

Society, 28(1): 15-21.

1967a. Ecological Studies in the Mammoth Cave System

of Kentucky I: The Biota. International Journal

of Speleology, 3:147-204, plates 37-64.

1967b. Observations on the Ecology of Caves. The

American Naturalist, 101 (922) :475-491, 4 figures.

1968. Cave Ecology and the Evolution of Troglobites.

Pages 35-102, 24 figures, in volume 2 of Evolutionary Biology, by Theodosius Dobzhansky, M. K.

Hecht, and W. C. Steere. New York, AppletonCentury-Crofts.

Barr, Thomas C, Jr., and Robert A. Kuehne

1971. Ecological Studies in the Mammoth Cave System

of Kentucky. II. The Ecosystem. Annales de

Spiliologie, Revue Trimestrielle, 26(l):47-96,

12 figures.

Bell, J. Carlton

1906. The Reactions of Crayfish to Chemical Stimuli.

Journal of Comparative Neurology and Psychology, 16(4):299-326.

Binkerd, A. D.

1869. The Mammoth Cave and Its Denizens: A Complete Descriptive Guide, vi + 95 pages. Cincinnati, Robert Clarke & Co.

Blatchley, W. S.

1897. Indiana Caves and Their Fauna. Twenty-first

Annual Report of the Indiana Department of Geology and Natural Resources, pages 121-212,

plates 4-13.

Bolivar, C, and R. Jeannel

1931. Biospeologica LVI. Campagne speologique dans

l'Amerique du nord en 1928. Archives de Zoologie Experiment ale et Generate, 71:293-499, 212

figures.

Bott, Richard

1950. Die Flusskrebse Europas (Decapoda, Astacidae).

Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, number 483, 36 pages, 25

figures, 6 plates.

Bouvier, E. L.

1940. Fauna de France, 37, Decapodes Marcheurs. 339

pages, 222 figures, 14 plates. Paris, Paul LeChevalier et Fils, 12 Rue de Tournon.

Brown, Frank A.

1961. Diurnal Rhythm in Cave Crayfish. Nature, 191

(4791)=929-930.

Bundy, W. F.

1876. Appendix, pages 24-25, in List of Illinois Crustacea with Descriptions of New Species, by S. A.

Forbes. Bulletin of the Illinois Museum of Natural History, number 1, 25 pages.

Burroughs, W. G.

1923. A Pottsville-filled Channel in the Mississippian.

Kentucky Geological Survey, series 6, 10:115-126.

Call, Richard Ellsworth

1897. The Natural History of the Cave. 116 pages, 24

figures, in Mammoth Cave of Kentucky, by H. C.

Hovey and R. E. Call. Louisville, Kentucky,

John P. Morton and Co.

1901. [The natural history of the cavern; a reprint of

the above.] In Mammoth Cave of Kentucky, an

Illustrated Manual, by H. C. Hovey and R. E.

Call, v + 112 pages. Louisville, Kentucky, John

P. Morton and Company.

Chappuis, P. A.

1927. Die Tierwelt der Unterirdischen Gewasser. Volume 3, 175 pages, 62 figures, 4 plates, in Die

Binnengewasser, by August Thienemann. Stuttgart, E. Schweizerbart'sche Verlagsbuchhandlung.

Cole, Gerald A.

1959. A Summary of Our Knowledge of Kentucky Crustaceans. Transactions of the Kentucky Academy

of Science, 20(3-4): 66-81.

Collett, John

1874. Geology of Lawrence County, Indiana. Fifth Annal Report of the Geological Survey of Indiana,

pages 260-312.

1879. Geological Report on Harrison and Crawford

Counties, Indiana. Eighth, Ninth, and Tenth

Annual Reports of the Geological Survey of Indiana, pages 291-522.
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cooper, John E.

1966. Preliminary Observations on the Biology of Shelta

Cave, Alabama. Bulletin of the National Speleological Society, 28(2):96-97.

1967. Animals in Alabama Caves. Pages 14-16 in The

Caves of Alabama, A Guide for the 1967 Convention of the National Speleological Society at Birmingham and Huntsville, Alabama, June 1967,

by John Veitch and John E. Cooper.

1968. Shelta Cave, a Biological Marvel. NSS News,

26(2):33-35.

Cooper, J. E., and Martha Riser Cooper

1969a. Comments on Alabama Cave Biology. Pages 25-

30 in Section 2 (Cave Science) of Speleo Digest,

William T. Plummer, editor.

1969b. Cave-Associated Herpetozoa II: Salamanders of

the genus Gyrinophilus in Alabama Caves. Bulletin of the National Speleological Society, 30(2):

19-24, 1 figure.

Cooper, J. E., and Thomas L. Poulson

1968. A Guide for Biological Collecting in Caves. NSS

News, 26(9): 127-138, 25 figures.

Cooper, Martha Riser

1969. Sensory Specialization and Allometric Growth in

Cavernicolous Crayfishes. Actes Du IVe Congres

International de Spileologie en Yugoslavie (12-

26 IX 1965), 4-5:203-208, 1 figure.

Cope, Edward Drinker

1871a. Wyandotte Cave and Its Life. The Indianapolis

Journal, 5 Sept. 1871, page 4.

1871b. Life in the Wyandotte Cave. Annals and Magazine of Natural History, series 4, 8(47):368-370

[reprinted from Indianapolis Journal, 5 Sept

1871].

1872a. On the Wyandotte Cave and Its Fauna. American Naturalist, 6 (7): 406-422, figures 109-116.

1872b. Report on the Wyandotte Cave and its Fauna.

Third and Fourth Annual Report of the Geological Survey of Indiana, pages 157-182, figures 109-

116. [Reprinted from American Naturalist,

6(7):406-422.]

1872c. [In minutes of meeting of the Academy, 28 November 1871.] Proceedings of the Academy of

Natural Sciences of Philadelphia, part I, new

series, pages 297-298.

1879. Observations on Wyandotte Cave and its Fauna.

Eighth, Ninth, and Tenth Annual Reports of the

Geological Survey of Indiana, pp. 489-506, figures

109-116. [Reprinted from American Naturalist,

6(7): 406-422.]

1881. Orconectes hamulatus. Pages 881-882, plate 7,

figure 1, in The Fauna of the Nickajack Cave, by

E. D. Cope and A. S. Packard, Jr. American

Naturalist, Volume 15.

Cope, E. D., and A. S. Packard, Jr.

1881. The Fauna of the Nickajack Cave. American

Naturalist, 15:877-882, plate 7.

Creaser, Edwin P.

1931. Three New Crayfishes (Cambarus) from Puebla

and Missouri. Occasional Papers of the Museum

of Zoology, University of Michigan, number 224,

10 pages, 17 figures.

1932. The Decapod Crustacea of Wisconsin. Transactions of the Wisconsin Academy of Sciences, Arts

and Letters, 27:321-338, 13 figures.

1962. Notes on Homologies and Genetic Relationships in

the Cambarinae Crayfishes. 7 pages, 22 figures.

Privately printed.

Crocker, Denton W.

1957. The Crayfishes of New York State (Decapoda,

Astacidae). New York State Museum and Science

Service Bulletin, number 335, 97 pages, 5 plates,

6 figures.

Crocker, D. W., and David W. Barr

1968. Handbook of the Crayfishes of Ontario, ix +

158 pages, 87 figures. University of Toronto Press.

Croizat, Leon

1958. Panbiogeography, or an Introductory Synthesis of

Zoogeography, Phytogeography and Geology; with

Notes on Evolution, Systematics, Ecology, Anthropology, etc. Volume I (The New World), xxxi

+ 1018 pages, 119 figures. Distributed by Wheldon and Wesley, Ltd., Nr. Hitchin, Herts, England.

Cunningham, J. T.

1893. Blind Animals in Caves. Nature, 47:439.

Dana, James D.

1852. Crustacea, Part 1. Volume 13, 685 pages, in

United States Exploring Expedition During the

Years 1838, 1839, 1840, 1841, 1842 Under the

Command of Charles Wilkes, US.N. Philadelphia, C. Sherman.

Darwip, Charles

1859. On the Origin of Species, by Means of Natural

Selection, or the Preservation of Favored Races in

the Struggle for Life, ix + 502 pages. London,

J. Murray.

Dearolf, Kenneth

1942. Report of a Biological Reconnaissance of the New

Discovery in Mammoth Cave, Kentucky, January

7-11, 1941. Bulletin of the National Speleological Society, number 4, pages 48—52.

1953. The Invertebrates of 75 Caves in the United

States. Proceedings of the Pennsylvania Academy

of Science, 27:225-241.

Eberly, William R.

1954. The Use of Crayfish to Illustrate some Ecological

Principles. Turtox News (General Biological Supply House, Inc.), 32(3):58-60.

1955. Summary of the Distribution of Indiana Crayfishes, Including New State and County Records.

Proceedings of the Indiana Academy of Science,

65:281-283.

1958. Competition and Evolution in Cave Crayfishes of

Southern Indiana. Mimeographed, 7 pages.

1960. Competition and Evolution in Cave Crayfishes of

Southern Indiana. Systematic Zoology, 9:29-32.
NUMBER 10 5 75

Eigenmann, Carl H.

1899. Explorations in the Caves of Missouri and Kentucky. Proceedings of the Indiana Academy of

Science, 1898, pages 58-61.

1903. The Eyes of the Blind Vertebrates of North

America. V. The History of the Eye of the

Blind Fish Amblyopsis from its Appearance to its

Disintegration in Old Age. Pages 167-204, 4

plates, in Mark Anniversary Volume, George H.

Parker, editor. New York, Henry Holt and Company. 4 plates.

Erichson, W. F.

1846. Uebersicht der Arten der Gattung Astacus. Archiv fur Naturgeschichte, Berlin, 1:86-103.

Fage, Louis

1931. Crustaces Amphipodes et Decapodes. Pages 361-

374, 21 figures, in Biospeologica LVI. Campagne

speologiquc dans FAmerique du nord en 1928, by

C. Bolivar and R. Jeannel. Archives de Zoologie

Expirimentale et Generate, 71:361-374, 21 figures.

Faxon, Walter

1884. Descriptions of New Species of Cambarus: to

Which is Added a Synonymical List of the Known

Species of Cambarus and Astacus. Proceedings of

the American Academy of Arts and Science,

20:107-158.

1885a. A Revision of the Astacidae. Part I. The Genera

Cambarus and Astacus. Memoirs of the Museum

of Comparative Zoology at Harvard College,

volume 10, number 4, vi+186 pages, 10 plates.

1885b. A List of the Astacidae in the United States National Museum. Proceedings of the United States

National Museum, 8(23): 356-361.

1889. Cambarus setosus. Page 237, figures 1, 2, 3, 7 in

Cave Animals in Southwestern Missouri, by Harrison Garman. Bulletin of the Museum of Comparative Zoology at Harvard College, volume 17,

number 6.

1890. Notes on North American Crayfishes, Family

Astacidae. Proceedings of the United States National Museum, 12(785):619-634.

1898. Observations on the Astacidae in the United States

National Museum and in the Museum of Comparative Zoology, with Descriptions of New Species. Proceedings of the United States National

Museum, 20(1136):643-694, plates 62-70.

1914. Notes on the Crayfishes in the United States National Museum and the Museum of Comparative

Zoology, with Descriptions of New Species and

Subspecies. Memoirs of the Museum of Comparative Zoology at Harvard College, 40(8):351-427,

13 plates.

Fingerman, Milton, and W. C. Mobberly, Jr.

1960. Trophic Substances in a Blind Cave Crayfish.

Science, 132(3418):44-45.

Fitzpatrick, Joseph F., Jr.

1963. Geographic Variation in the Crawfish Faxonella

clypeata (Hay) with the Definition and Defense

of the Genus Faxonella Creaser (Decapoda,

Astacidae). Tulane Studies in Zoology, 10(1):

57-79, 20 figures.

1967. The Propinquus Group of the Crawfish Genus

Orconectes (Decapoda: Astacidae). The Ohio

Journal of Science, 67(3): 129-172, 27 figures.

Fitzpatrick, J. F., Jr., and Rudolph Prins

1965. The First-Form Male of Orconectes bisectus

Rhoades, a Poorly Known Kentucky Crawfish.

American Midland Naturalist, 74(1): 141-147,

12 figures.

Fleming, Robert S.

1939. The Larger Crustacea of the Nashville Region.

Journal of the Tennessee Academy of Science,

13(4):296-314;14(2):261-264; 14(3):299-324;

22 plates.

Frey, David G.

1965. Other Invertebrates—an Essay in Biogeography.

Pages 613-631 in The Quaternary of the United

States, VII Congress of the International Association for Quaternary Research. Princeton University Press.

Garman, Samuel

1889. Cave Animals from Southwestern Missouri. Bulletin of the Museum of Comparative Zoology at

Harvard College, 17(6): 225-259, 2 plates.

1924. A Little Known Cave Crayfish. Transactions of

the Kentucky Academy of Science, Seventh Annual Meeting, 1 (1914-1923): 87-94, 20 figures.

Gibbes, L. R.

1850. On the Carcinological Collections of the United

States, an Enumeration of the Species Contained

in Them, with Notes on the Most Remarkable and

Descriptions of New Species. Proceedings of the

American Association for the Advancement of

Science, 3:167-201.

Giovannoli, Leonard

1933a. Invertebrate Life of Mammoth and Other Neighboring Caves. Pages 600-623, figures 86-90, in

Cave Life of Kentucky, by Vernon Bailey. American Midland Naturalist, volume 14, number 5.

1933b. Invertebrate Life of Mammoth and Other Neighboring Caves. Pages 218-241, 4 figures, in Cave

Life of Kentucky Mainly in the Mammoth Cave

Region, by Vernon Bailey. Notre Dame University Press.

Girard, Charles

1852. A Revision of the North American Astaci, with

Observations on Their Habits and Geographical

Distribution. Proceedings of the Academy of

Natural Sciences of Philadelphia, 6:87-91.

Graeter, E.

1909. Die Zoologische Erforschung der Hohlengewasser

seit dem Jahre 1900 mit Ausschluss der Vertebraten. Internationale Revue der Gesamten

Hydrobiologie und Hydrographie. Dr. Werner

Klinkhardt, Leipzig, 2:457-479.

Graham, Dick

1969. Indian Rock Cave. Alabama Caving 68 '69,

page 4.
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hagen, Hermann A.

1870. Monograph of the North American Astacidae.

Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, number 3,

109 pages, 11 plates.

1872. The Blind Crayfish. American Naturalist, 6:494-

495.

Harris, J. Arthur

1903a. The Habits of Cambarus. American Naturalist,

37:601-608.

1903b. An Ecological Catalogue of the Crayfishes Belonging to the Genus Cambarus. Kansas University

Science Bulletin, 2(3):51-187, 4 plates.

Hart, C. W., Jr., and Dabney G. Hart

1966. Four New Entocytherid Ostracods from Kentucky,

with Notes on the Troglobitic Sagittocythere barri.

Notulae Naturae, number 388, 10 pages, 13

figures.

1969. The Functional Morphology of Entocytherid Ostracod Copulatory Appendages, with a Discussion

of Possible Homologues in other Ostracods. Pages

154-167, 12 figures, in The Taxonomy, Morphology and Ecology of Recent Ostracoda, John W.

Neale, editor. Edinburgh, Oliver and Boyd.

Hart, C. W., Jr., and Horton H. Hobbs, Jr.

1961. Eight New Troglobitic Ostracods of the Genus

Entocythere (Crustacea, Ostracoda) from the

Eastern United States. Proceedings of the Academy of Natural Sciences of Philadelphia, 113(8):

173-185, figures 1-32.

Hay, William Perry

1891. The Crustacea of Indiana. Proceedings of the

Indiana Academy of Science, 1891, pages 147—151.

1893. Observations on the Blind Crayfishes of Indiana,

with a Description of a New Subspecies; Cambarus pellucidus testii. Proceedings of the United

States Na'ional Museum, 16(935): 283-286,

plates 64—65.

1896. The Crawfishes of the State of Indiana. Twentieth Annual Report of the Department of Geology and Natural Resources of Indiana, pages

475-507, 15 figures.

1897. On a Collection of Crustaceans from Indiana

Caves. Pages 206-210 in Indiana Caves and their

Fauna, by W. S. Blatchley. Twenty-first Annual

Report of the Department of Geology and Natural Resources of Indiana.

1899. Synopsis of North American Invertebrates—The

Astacidae of North America. American Naturalist, 33(396):957-966.

1902a. Observations on the Crustacean Fauna of the Region about Mammoth Cave, Kentucky. Proceedings of the United States National Museum, 25

(1285):223-236.

1902b. Observations on the Crustacean Fauna of Nickajack Cave, Tennessee, and Vicinity. Proceedings

of the United States National Museum, 25(1292):

417-439, sketch map, figures I, II, 1-8.

Hobbs, Horton H., Jr.

1940. Seven New Crayfishes of the Genus Cambarus

from Florida, with Notes on Other Species. Proceedings of the United States National Museum,

89(3097):387-423, figures 14-22.

1941. Three New Florida Crayfishes of the Subgenus

Cambarus (Decapoda, Astacidae). American

Midland Naturalist, 26(1) : 110-121, 2 plates.

1942a. A Generic Revision of the Crayfishes of the

Subfamily Cambarinae (Decapoda, Astacidae)

with the Description of a New Genus and Species.

American Midland Naturalist, 28(2):334-357, 3

plates.

1942b. The Crayfishes of Florida. University of Florida

Publications, Biological Science Series, volume 3,

number 2, 179 pages, 24 plates.

1943. Two New Crayfishes of the Genus Procambarus

from Mexico (Decapoda, Astacidae). Lloydia,

6:198-206, 26 figures.

1948a. On the Crayfishes of the Limosus Section of the

Genus Orconectes. Journal of the Washington

Academy of Science, 38(1): 14-21, 28 figures.

1948b. A New Crayfish of the Genus Orconectes from

Southern Tennessee. Proceedings of the Biological Society of Washington, 61:85-92, 9 figures.

1952. A New Albinistic Crayfish of the Genus Cambarus

from Southern Missouri with a Key to the Albinistic Species of the Genus (Decapoda, Astacidae ). American Midland Naturalist, 48 (3) : 689-

693, 8 figures.

1958. The Evolutionary History of the Pictus Group of

the Crayfish Genus Procambarus (Decapoda, Astacidae). Quarterly Journal of the Florida Academy of Sciences, 21(1): 71-91, 20 figures.

1959. [Key to Crayfishes.] Pages 869-901, figures 31.

18-31.41, in Ward and WhippWs Freshwater Biology, ed. 2, W. T. Edmondson, editor. New

York, J. Wiley & Sons, Inc.

1962. Notes on the Affinities of the Members of the

Blandingii Section of the Crayfish Genus Procambarus (Decapoda, Astacidae). Tulane Studies

in Zoology, 9(5): 273-279, 72 figures.

1965. A New Crayfish of the Genus Cambarus from

Tennessee with an Emended Definition of the

Genus (Decapoda, Astacidae). Proceedings of

the Biological Society of Washington, 78:265-273,

12 figures.

1967a. The Current Status of the Crayfishes Listed by

Girard (1852) in His "A Revision of the North

American Astaci . . ." (Decapoda, Astacidae).

Crustaceana, 12(2): 124-132.

1967b. A New Crayfish from Alabama Caves with Notes

on the Origin of the Genera Orconectes and

Cambarus (Decapoda, Astacidae). Proceedings

of the United States National Museum, volume

123, number 3621, 17 pages, 21 figures.

1969. On the Distribution and Phylogeny of the Crayfish Genus Cambarus. Pages 93-178, 20 figures,

in The Distributional History of the Biota of the
NUMBER 10 5 77

Southern Appalachians, Part I, Invertebrates,

Perry C. Holt, Richard L. Hoffman, and C. Willard Hart, Jr., editors. Virginia Polytechnic Institute, Research Division Monograph.

1970. A New Crayfish from the Nashville Basin, Tennessee. Proceedings of the Biological Society of

Washington, 83(14): 161-169, 1 figure.

1971a. The Entocytherid Ostracods of Mexico and Cuba.

Smithsonian Contributions to Zoology, number 81,

55 pages, 31 figures.

1971b. A New Troglobitic Crayfish from Florida. Quarterly Journal of the Florida Academy of Sciences,

34(2): 114-124, 19 figures.

Hobbs, H. H., Jr., and Thomas C. Barr, Jr.

1960. The Origins and Affinities of the Troglobitic

Crayfishes of North America (Decapoda, Astacidae). I. The Genus Cambarus. American Midland Naturalist, 64(1): 12-33, 57 figures.

Hobbs, H. H., Jr., and M. S. Bedinger

1964. A New Troglobitic Crayfish of the Genus Cambarus (Decapoda, Astacidae) from Arkansas with

a Note on the Range of Cambarus cryptodytes

Hobbs. Proceedings of the Biological Society of

Washington, 77:9-15, 11 figures.

Hobbs, H. H., Jr., and J. F. Fitzpatrick, Jr.

1962. A New Crayfish of the Propinquus Group of the

genus Orconectes from the Ohio Drainage System

in West Virginia (Decapoda, Astacidae). Proceedings of the Biological Society of Washington,

75:207-214, 15 figures.

Hobbs, H. H., Jr., and Alejandro Villalobos F.

1964. Los Cambarinos de Cuba. Anales del Instituto

de Biologia de la Universidad Nacional Autonoma

de Mexico, 34(1-2): 307-366, 50 figures.

Holthuis, L. B.

1956. Proposed Addition to the "Official List of Generic

Names in Zoology . . . ." Bulletin of Zoological

Nomenclature, 12(4): 107-119.

1964. On the Status of Two Allegedly European Crayfishes, Cambarus typhlobius Joseph, 1880, and

Austropotamobius pallipes bispinosus Karaman,

1961 (Decapoda, Atacidae). Crustaceana, 7 (1):

42-48, 1 plate.

Hovey, Horace C.

1882. Celebrated American Caverns, Especially Mammoth, Wyandot, and Luray, Together with Historical Scientific and Descriptive Notices of Caves

and Grottoes in other Lands; with Maps and

Illustrations, xii+288 pages, illustrated. Cincinnati, Robert Clarke and Company.

1891. Guide Book to the Mammoth Cave of Kentucky;

Historical, Scientific and Descriptive. 75 pages, illustrated. Cincinnati, Robert Clarke and Company.

1912. Mammoth Cave of Kentucky, with an Account of

Colossal Cavern. 131 pages, illustrated. Louisville, John P. Morton and Company.

Hubbard, H. G.

1880. Two Days' Collecting in the Mammoth Cave, with

Contributions to a Study of its Fauna. American

Entomologist, 3:34-40, 79-86, figures 8-10, 18-

24.

Huheey, James E.

1961. A Cyclical Evolutionary Process. Systematic Zoology, 10(l):43-45.

Jackson, George F.

1942. Animal Life of Caverns. Bulletin of the National

Speleological Society, number 4, pages 3-7.

1953. Wyandotte Cave. 66 pages, illustrated. Narberth, Pennsylvania, Livingston Publishing Company.

1955. Caves of Indiana. Bulletin of the National Speleological Society, 16:55-64.

Jegla, Thomas C.

1964. Reproductive and Molt Periods in a Cave Crayfish. Bulletin of the National Speleological Society, 26(2):81.

1965a. Studies of the Eyestalks of Cave Crayfish. American Zoologist, 5(4) :647.

1965b. Studies of the Eyestalk, Metabolism, and Molting

and Reproductive Cycles in a Cave Crayfish [Orconectes pellacious]. Dissertation Abstracts, 25

(2):1421.

1966. Reproductive and Molting Cycles in Cave Crayfish. Biological Bulletin, 130(3):345-358, 4 figures.

1969. Cave Crayfish: Annual Periods of Molting and

Reproduction. Actes du IVe Congres International de Speleologie en Yugoslavie (12-26 IX

1965), 4-5: 135-137.

Jegla, T. C, and Thomas L. Poulson

1968. Evidence of Circadian Rhythms in a Cave Crayfish. Journal of Experimental Zoology, 168(2):

273-282, 5 figures.

1970. Circannian Rhythms—I. Reproduction in the

Cave Crayfish, Orconectes peUucidus inermis.

Comparative Biochemical Physiology, 33:347-

355, 4 figures.

Jegla, T. C, T. L. Poulson, and M. R. Cooper

1965. Interpopulation Variations in a Cave Crayfish

(Orconectes peUucidus). American Zoologist,

5(4):639.

Jeannel, R., and H. Henrot

1949. Les cole"opteres cavernicoles de la region des

Appalaches. Notes BiospeoUogiques, Musium

National d'Histoire Naturelle, 4(12):7-116.

Jillson, Willard Rouse

1954. Geology of Crystal Cave. 39 pages. Frankfort,

Kentucky, Roberts Printing Co.

Jones, Walter B., and William W. Varnedoe, Jr.

1968. Caves of Madison County, Alabama. Geological

Survey of Alabama, Circular 52, 177 pages, 128

figures, 1 plate.

Joseph, Gustav

1880. Demonstration eines neuen zur Familie der Astaciden gehorenden blinden Grottenkrebses aus

Krain. Jahres-Bericht der Schlesische Gesellschaft fur vaterlandische Kultur, 57:202.
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1881. Erfahrungen im wissenschaft lichen Sammeln und

Beobachten der den Krainer Tropfsteingrotten

eigenen Arthropoden. Berliner Entomologische

Zeitschrift, 25:233-282.

1882. Systematisches Verzeichniss der in der TropfsteinGrotten von Krain einheimischen Arthropoden

nebst Diagnosen der vom Verfasser entdeckten

und bisher noch nicht beschriebenen Arten. Schluss

der Abhandl.: Erfahrungen im wissenschaftlichen

Sammeln und Beobachten der den Krainer Tropfsteingrotten eigenen Arthropoden. Berliner Entomologische Zeitschrift, 26:1-50.

Larimer, James L., Daniel L. Trevino, and Rberts A. Ashby

1966. A Comparison of Spectral Sensitivities of Caudal

Photoreceptors of Epigeal and Cavernicolous Crayfish. Comparative Biochemical Physiology, 19

(2):409-415, 1 figure.

Leydig, Franz

1883. Untersuchungen zur Anatomie und Histologie der

Thiere. 174 pages, 8 plates. Bonn, Verlag von

Emil Strauss.

Lonnberg, Einar

1894. Cambarids from Florida, A New Blind Species.

Zoologischer Anzeiger, 17:125-127.

1895. Cambarids from Florida, a New Blind Species.

Bihang till Koniglische Svenska V etenskaps-Akademiens Handlingar, 22(4): 3-14, 6 figures.

Lucas, H.

1864. [Note on Astacus pellucidus.] Bulletin Sociiti

Entomologique de France, 4:iv.

Meredith, W. G., and F. J. Schwartz

1960. Maryland Crayfishes. Maryland Department of

Research and Education, Educational Series, number 46, 32 pages, 17 figures.

Minckley, W. L.

1963. The Ecology of a Spring Stream, Doe Run, Meade

County, Kentucky. Wildlife Monographs, number 11, 124 pages, 30 figures.

Mohr, Charles E.

1964. Exploring America Underground. National Geographic, 125 (6): 803-837, illustrated.

Mohr, Charles E., and Thomas L. Poulson

1966. The Life of the Cave. Our Living World of Nature series. 232 pages, illustrated. New York,

McGraw Hill Book Company.

Newport, George

1855. On the Ocelli of the Genus Anthophorabia.

Transactions of the Linnean Society of London,

21:161-165, figures 10-15.

Nicholas, Brother G.

1960. Checklist of Macroscopic Troglobitic Organisms

of the United States. American Midland Naturralist, 64(1): 123-160.

1969. Troglobites of Alabama—The Beetles. Pages 18-

25 in Section 2, Cave Science, of Speleo Digest,

William T. Plummer, editor.

1970. Life in the Underground World. National Wildlife, 8(6):21-24, illustrated.

Ortmann, A. E.

1892. Decapod Crustacea in Strassburger Museums.

Zoologische Jahrbiicher abteilung fur Systematik,

Okologie und Geographie der Tiese, 6:1-13.

1902. The Geographical Distribution of Freshwater Decapods and its Bearing Upon Ancient Geography.

Proceedings of the American Philosophical Society, 41 (171): 267-400, 8 figures.

1905a. The Mutual Affinities of the Species of the Genus

Cambarus and Their Dispersal over the United

States. Proceedings of the American Philosophical Society, 44( 180) :91-136, plate 3.

1905b. Procambarus, a New Subgenus of the Genus

Cambarus. Annals of the Carnegie Museum,

3(3):435-442, 3 figures.

1906. The Crayfish of the State of Pennsylvania. Memoirs of the Carnegie Museum, 2(10): 343-534,

plates A, B, 39-43.

1918. Higher Crustaceans (Malacostraca). Pages 828-

850, figures 1303-1318, in Freshwater Biology,

by H. B. Ward and G. C. Whipple, New York,

John Wiley and Sons, Inc.

1931. Crawfishes of the Southern Appalachians and the

Cumberland Plateau. Annals of the Carnegie

Museum, 20(2) :61-160.

Osborn, R. C.

1912. The Crayfish. Zoological Society Bulletin, 16

(54):921-927.

Packard, A. S., Jr.

1871. On the Crustaceans and Insects. In The Mammoth Cave and Its Inhabitants, by A. S. Packard

and F. W. Putnam. American Naturalist, 5(12):

744-761, figures 122-133.

1872a. [Note on Cope 1871b.] American Naturalist, 6:

30.

1872b. In The Mammoth Cave and its Inhabitants; or

Descriptions of the Fishes, Insects and Crustaceans

Found in the Cave; wtih Figures of the Various

Species, and an Account of Allied Forms, Com~

prising Notes upon their Structure, Development

and Habits, with Remarks upon Subterranean

Life in General, by A. S. Packard, Jr., and F. W.

Putnam. [Consists largely of reprints of articles

by Packard (American Naturalist, 1871) and

Putnam (Annual Report of the Peabody Academy,

1871, and American Naturalist, 1872).] 62 pages,

8 figures, 2 plates. Salem, Massachusetts, Salem

Press.

1873. On the Cave Fauna of Indiana. Fifth Annual

Report of the Peabody Academy of Science, July

1873, pages 93-97.

1874. [Remarks] in Minutes of the Section of Entomology Meeting of the Boston Society of Natural History, January 1, 1874. Proceedings of the Boston

Society of Natural History, 16:209.

1879. Zoology for Students and General Readers. vi+

719 pages, 545 figures. New York, Henry Holt

and Company.
NUMBER 10 5 79

1888. The Cave Fauna of North America, with Remarks

on the Anatomy of the Brain and Origin of the

Blind Species. Memoirs of the National Academy

of Sciences, volume 4, 156 pages, 21 figures, 27

plates.

1890. The Effect of Cave Life on Animals, and its

Bearing on the Evolutionary Theory. Popular

Science Monthly, 36(3):389-397.

1894. On the Origin of the Subterranean Fauna of

North America. American Naturalist, 28:727-

751.

Park, Orlando

1938. Studies in Nocturnal Ecology, VII. Preliminary

Observations on Panama Rain Forest Animals.

Ecology, 19(2): 208-223.

Park, O., T. W. Roberts, and S. J. Harris

1941. Preliminary Analysis of Activity of the Cave

Crayfish, Cambarus pellucidus. American Naturalist, 75(757): 154-171, 5 figures.

Parker, G. H.

1890. The Eyes in Blind Crayfishes. Bulletin of the

Museum of Comparative Zoology at Harvard ColItge, 20(5): 153-162, 1 plate.

Pearse, A. S.

1910. The Crawfishes of Michigan. Michigan State

Biological Survey, Publication 1:9-22, 8 plates.

Penn, George H., Jr.

1952. A New Crayfish of the Virilis Section of the

Genus Orconectes. Natural History Miscellanea,

Chicago Academy of Sciences, number 109, 7

pages, 12 figures.

Penn, G. H., Jr., and J. F. Fitzpatrick, Jr.

1963. Interspecific Competition Between Two Sympatric

Species of Dwarf Crawfishes. Ecology, 44(4):

793-797, 1 figure.

Pennak, Robert W.

1953. Freshwater Invertebrates of the United States.

be+ 769 pages, 470 figures. New York, The Ronald Press Company.

Pope, T. E. B.

1926. Blind Fishes and Crayfishes. Yearbook of the

City of Milwaukee, 6:163-171, figures 156, 157.

Poulson, Thomas L.

1964. Animals in Aquatic Environments: Animals in

Caves. Pages 749-771 in Handbook of Physiology, D. B. Dill, editor. Washington, American

Physiological Society.

Poulson, T. L., and Thomas C. Jegla

1966. Circadian Rhythms and Seasonal Cycles in Cave

Fishes and Crayfish. Bulletin of the National

Speleological Society, 28(2):98.

Poulson, T. L., and William B. White

1969. The Cave Environment. Science, 165(3897):

971-981, 3 figures.

Powell, Richard L.

1961. Caves of Indiana. Indiana Department of Conservation, Geological Survey, Circular 8, 127

pages, 58 figures, 4 plates.

Pratt, Henry Sherring

1916. A Manual of the Common Invertebrate Animals

Exclusive of Insects. 737 pages, illustrated. A.

C. McClurg and Company.

Price, F. M.

1900. The Eyes of Cambrus pellucidus [Title, without

text, of a paper presented to the Academy]. Proceedings of the Indiana Academy of Science, 1899,

page 155.

Prins, Rudolph

1968. Comparative Ecology of the Crayfishes Orconectes

rusticus rusticus and Cambarus tenebrosus in Doe

Run, Meade County, Kentucky. Internationale

Revue der gesamten Hydrobiologie, 53(5): 667-

714, 13 figures.

Putnam, F. W.

1872. The Blind Fishes of Mammoth Cave and Their

Allies. American Naturalist, 6:6-30, 2 plates.

1875a. Remarks on Living Fishes and Crayfishes of Mammoth Cave. (In Minutes of the Meeting of the

Boston Society of Natural History, December 2,

1874.) Proceedings of the Boston Society of

Natural History, 17:222-225.

1875b. Remarks on the Mammoth Cave and Some of its

Animals. (In Minutes of the Meeting of the Essex

Institute, Salem, Massachusetts, November 25,

1874.) Bulletin of the Essex Institute, 6(2): 190-

200.

1877. On Some of the Habits of the Blind Crawfish,

Cambarus pellucidus, and the Reproduction of

Lost Parts. Proceedings of the Boston Society

of Natural History, 18:16-19.

Rab£,Fefix

1890. La Question de L'Ecrevisse. 83 pages. Imprimerie de Sezanne, H. Patoux.

Rhoades, Rendell

1941. Notes on Some Crayfishes from Alabama Caves,

with the Description of a New Species and a New

Subspecies. Proceedings of the United States

Notional Museum, 91(3129): 141-148, 2 figures.

1944. The Crayfishes of Kentucky with Notes on Variation, Distribution and Descriptions of New Species

and Subspecies. American Midland Naturalist,

31(1): 111-149, 10 figures.

1959. A Review of the Status of Orconectes inermis Cope

and a Suggested Nomenclatorial Change. American Midland Naturalist, 61(2): 399-402.

1962. The Evolution of Crayfishes of the Genus Orconectes Section Limosus. (Crustacea, Decapod*).

Ohio Journal of Science, 62(2): 65-96, 8 figures.

Semper, Karl

1881. Animal Life as Affected by the Natural Conditions

of Existence, rvi+472 pages, 106 woodcuts.

New York, D. Appleton and Company.

Shaler, N. S.

1875. On the Antiquity of the Caverns and Cavern Life

of the Ohio Valley. Memoirs of the Boston Society of Natural History, 2:355-363. [Repub-
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lished, 1876, in Memoirs of the Kentucky Geological Survey, 1(1): 1-13.]

Silliman, B., Jr.

1851. On the Mammoth Cave of Kentucky. [From a

letter addressed to Prof. Guyot, Cambridge; dated

November 8, 1950, Louisville.] American Journal of Science and Arts, series 2, 11:332-339.

Sloan, John

1888. Letters to Packard Written in 1873. In Packard,

A. S., The Cave Fauna of North American, with

Remarks on the Anatomy of the Brain and Origin

of the Blind Species. Memoirs of the National

Academy of Sciences, 4:15, 16.

Smith, S. I.

1873. The Crustacea of the Freshwaters of the United

States. Report of the United States Commission

of Fish and Fisheries, (1872-1873), pages 637-

665.

1875. The Crustaceans of the Caves of Kentucky and

Indiana. American Journal of Science and Arts,

series 3,9:476-477.

Spandl, Hermann

1926. Die Tierwelt der Unterirdischen Gewasser. xi +

235 pages, 116 figures. Spelaologische Monographien herausgegeben vom Spelaologischen Institut der Bundeshohlen-kommission, Wien, 11.

Spurgeon, C. H.

1915. The Eyes of Cambarus setosus and Cambarus

pellucidus. Biological Bulletin, 28:385-396, 9

figures, 1 plate.

Stammer, H. J.

1932. Die Fauna des Timavo. Ein Beitrag sur Kenntnis

der Hohlen Gewasser, des Suss-und Brackwassers

im Karst. Zoologische Jahrbucher abteilung fur

Systematik, Okologie und Geographic der Tiese,

63:521-656, 16 figures.

Stebbing, Thomas R. R.

1893. A History of Crustacea, Recent Malacostraca.

viii + 466 pages, illustrated. New York, D. Appleton and Company.

Steele, Mary Isabelle

1902. Crayfish of Missouri. Publications of the University of Cincinnati, series 2, volume 2, bulletin 10,

33 pages, 5 plates.

Stenuit, Robert, and Marc Jasinski

1964. Marveilleux monde souterrain. 95 pages, illustrated. Western Publishing-Hachette International.

1966. Caves and the Marvelous World Beneath Us.

[Translated from the French by Harry Pearman.]

95 pages, illustrated. New York, A.S. Barnes and

Company.

Stiles, C. W., and Albert Hassall

1927. Key-Catalogue of the Crustacea and Arachnoids

of Importance in Public Health. Hygienic Laboratory Bulletin, number 148, pages 197-289.

Tarkington, Terry W., William W. Varnedoe, Jr., and John

D. Veitch

1965. Alabama Caves, iv+81 pages and maps.

Huntsville, Alabama, Huntsville Grotto of the National Speleological Society.

Tellkampf, Theodor G.

1844a. Ausflug nach der Mammuthhohle in Kentucky.

Das Ausland, (168) :671-672, (169):675-676,

(170):679-680, (171):683-684, (172):687-688,

(173):691-692, (174):695-696, (175):699-700.

1844b. Ueber den blinden Fisch der Mammuthhohle in

Kentucky, mit Bemerkungen uber einige andere in

dieser Hohle lebende Thiere. Archiv fur Anatomie, Physiologie und wissenschaftliche Medicin,

Heft IV, pages 381-394, Tafel IX.

1845. Memoirs on the Blind Fishes and some other

Animals Living in the Mammoth Cave in Kentucky. New York Journal of Medical and Collateral Science, pages 84-93, 1 plate.

Thompson, David H.

1967. A Comparison of Visual Acuity between Cave and

Epigeal Crayfish. Wisconsin Speleologist, Summer-Fall, 1967, page 45.

Thompson, W. M.

1844. Notice of the Blind Fish, Cray-fish and Insects

from the Mammoth Cave, Kentucky. Annals and

Magazine of Natural History, 13:111-113.

Torode, William

1968. Alabama 1000. Huntsville Grotto Newsletter, 9

(10):152-153.

1969. Alabama 1000. Alabama Caving '68-69, Special

Publication of the Huntsville Grotto of the National Speleological Society, page 16.

Turner, C. L.

1935. The Aberrant Secondary Sex Characters of the

Crayfishes of the Genus Cambarus. American

Midland Naturalist, 16(6): 863-882.

Underwood, Lucien M.

1886. List of the Described Species of Freshwater Crustacea from America, North of Mexico. Bulletin

of the Illinois State Laboratory of Natural History, 2(5):323-386.

Vandel, Albert

1964. Biospeologie. La Biologie des Animaux Cavernicoles. xviii + 619 pages, 80 figures, 11 plates.

Paris, Gauthier-Villars.

Van Straelen, Victor

1942. A propos de la distribution des ecrevisses, des

homards et des crabes d'eau douce. Bulletin

Musee Royal d'Histoire des Sciences Naturelles

de Belgique, 18(56): 1-11.

Villalobos, Alejandro F.

1953. Distribucicn geografica y notas ecologicas de los

cambarinos mexicanos. Memoria del Congresso

Cientifico Mexicano, IV Centario de la Universidad de Mexicano, 7:343-374, 4 plates.

1954. Estudios de los cambarinos mexicanos XII. Parte

1. Revision de las especies afines a Procambarus

mexicanus (Erichson) con Descripcion de formas

nuevas. Anales del Instituto de Biologia, Universidad Autonoma de Mixico, 25(1,2): 299-379,

19 plates.
NUMBER 10 5 81

1955. Cambarinos de la fauna mexicana (Crustacea

Decapoda). Tesis, Facultad de Ciencias, xvi + 290

pages, 62 plates. Universidad Nacional Autonoma de Mexico.

Wells, P. H.

1959. Response to Light by Cave Crayfishes. Occasional Papers of the National Speleological Society,

number 4, 15 pages, 10 figures.

Wolf, Benno

1934-1938. Volume 3, pages 103-105, in Animalium

Cavernarum Catalogus, by Dr. W. Junk. Verlag

fur Naturwissenschaften, 's-Gravenhage.

Wolfe, Douglas A., and David G. Cornwell

1964. Can item >ids of Cavernicolous Crayfish. Science,

144:1467-1469.

Wright, R. Ramsey

1884. Leydig's Researches in Anatomy and Histology.

[A review of Leydig's work (1893) and a comparison of the olfactory organs of O. pellucidus

with those of O. propinquus.] American Naturalist, 10(3):272-273.

Appendix

LIST OF CAVES.—The caves listed below are those

from which troglobitic crayfishes of the genus Orconectes have been reported. An asterisk indicates

those from which we have examined no specimens;

for these the earliest reference to the occurrence of

crayfish in them is cited. More precise locations for

most of the caves may be found under the Specimens

Examined section of the species treatments.

The number in parentheses following most of the

cave names identifies the troglobitic crayfish inhibiting it, as listed below:

1. Orconectes australis australis (Rhoades)

2. Orconectes australis packardi Rhoades

3. Orconectes incomptus, new species

4. Orconectes inermis inermis Cope

5. Orconectes inermis testii (Hay)

6. Orconectes pellucidus (Tellkampf)

Also enclosed in these parentheses are letters referring to the associated commensal entocytherid ostracods listed below (these ostracod records have been

extracted from Hart and Hobbs (1961) and Hart and

Hart (1966), with corrections and emendations in

locality records and in the identities of the crayfish

hosts) :

A. Dactylocythere prionata (Hart and Hobbs)

B. Dactylocythere steevesi (Hart and Hobbs)

C. Dactylocythere ungulata (Hart and Hobbs)

D. Donnaldsoncythere donnaldsonensis (Klie)

E. Donnaldsoncy there hiwasseensis (Hobbs and Walton)

r\ Donnaldsoncy there tuberosa (Hart and Hobbs)

G. Sagittocythere barri (Hart and Hobbs)

H. Sagittocythere stygia Hart and Hart

I. Uncinocythere simondsi (Hobbs and Walton)

J. Uncinocythere zancla Hobbs and Walton

ALABAMA

Jackson County

Bell Spring Cave (1)

Borderline Cave (1)

Canyon Cave (1)

Doodlebug Hole [ = Blowing Cave] (1)

Doug Green Cave (1)

Fern Cave (1 G)

Guess Creek Cave (1)

Indian Rock Cave (Graham, 1969)

Jess Elliott Cave (1)

Kennamer Cave (1)

Larkins Cave (1)

Limrock Blowing Cave (1)

McFarland Cave (1)

Paint Rock Cave (1)

Saltpeter Cave (1)

Salt River Cave (1)

Sauta Cave [ = Blowing Cave ] (1 G)

Madison County

Aladdin Cave (1)

Big Spring [ = Huntsville] Cave (1)

Burwell Cave (1)

Byrd Spring Cave (1)

Cold Spring Cave (1)

Fuqua Spring Cave (1)

•Grayson Spring Cave (Jones and Varnedoe,

1968)

Hering Cave [ = Cave Spring Cave] (1)

Matthews Cave (1)

Sadler Spring Cave (1)

Shelta Cave (1 G)

INDIANA

Bartholomew County: Clifty Caves incorrectly

cited by Hay, 1893; see Washington County.

Rhoades, 1959, mentioned crayfish from the

County but cited no localities.

Brown County: Rhoades, 1959; no specific localities were mentioned.

Crawford County

Archibald Cave (4)

•Binkleys Cave (Sight record, T.C.B.)

•Carter Byrnes Cave (Sight record, T.C.B.)

Crawfish Spring (4)
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

*"Evaston" [ = Everton] Cave (Blatchley, 1897)

Marengo Cave (4)

Sibert's Well Cave (4 G)

•Wild Cat Cave (Hay, 1893)

Wyandotte Cave (4)

Harrison County

Bradford Cave (4 G)

*Cave near Mauckport (4)

King's Cave (4)

Rhodes Cave (Collett, 1879)

Jefferson County

•Caves near Madison (Hay, 1893)

Lawrence County

•Bedford Cave (Packard, 1890)

Blue Spring Cave (4)

•"Connelly's" [ = Connerly's] Cave (Hay, 1893)

Donaldson's [Donaldson's-Bronson's] Cave (4 D)

Mitchell Cave (Eigenmann 1903, Banta 1907)

Shawnee Cave (Spurgeon, 1915)

•"Donnihue's" [=Donnehue's] Cave (Hay, 1893)

•Down's Cave (Hay, 1893)

•Hamer's Cave (Collett, 1874)

Harrison Cave (4)

Pless Cave (4)

Shiloh Cave (4)

Sullivan Cave (4)

Wagoner Cave (4)

Monroe County

Carmichael Cave (5)

Eller's Cave (5 G)

•Goode's Cave (Powell, 1961)

Mayfield's Cave (5)

May's Cave (5 G)

•Ranard School Cave (Powell, 1961)

Reeve's Cave (5)

Salamander Cave (5)

Shaft Cave (5)

•"Truett's" [ = Truitt's] Cave (Hay, 1891)

Orange County

•Cave in Orleans (Packard, 1888)

•Cave near Paoli (Hay, 1893)

•Lost River Cave (Packard, 1888)

Murray Spring Cave (4)

Washington County

Endless Cavern [ = Clifty Cave] (4)

Fredericksburg Cave (4)

River [ = Clifty] Cave (4)

KENTUCKY

Barren County

Cave City [ = Railroad] Cave (6)

Diamond Cave (6 G)

Twyman Cave (6)

Breckenridge County

Bandy Cave (4)

Bat Cave (?Packard, 1888) (4)

Cave near Big Spring (4)

Lockard Cave (4 G)

•McCubbin's Cave (Jegla and Poulson, 1968)

Thornhill Cave (4)

Christian County

Glover's Cave (6)

Edmonson County

Blowing Spring Cave (6)

Cedar Sink Cave (6)

Floyd Collins Crystal Cave (6)

•?Ganter's Cave [ = "cave . . . down the Green

River from Mammoth Cave"] (Putnam, 1875b)

Long Cave (6)

Mammoth Cave (6 G, H)

•Martin Cave (Sight record, T.C.B.)

Stillhouse Hollow Cave (6)

White Cave (6)

Green County

Brush Creek Cave (4)

Scott Cave (4)

Hardin County

Bland Cave (4 G, J)

Nelson Cave (4)

Hart County

Bald Knob Cave (6)

Buckner Hollow Cave (6 C, G)

Cave near Northtown (6)

Cooch Webb Cave (4)

Cub Run Cave (4)

Horse [ = Hidden River] Cave (6)

Mammoth Onyx Cave (6)

Riders Mill Cave (4 G, J)

Turner Cave (4)

Logan County

Cooks Cave (6)

Mud River Cave (6)

McCreary County

Eureka Cave (2)

Steele Hollow Cave (2)

Meade County

Joe Jones Cave (4)

Lime Kiln Cave (4)

Rockhaven Cave (4 E, G)
NUMBER 10 5 83

Shackletts Cave (4)

Pulaski County

Baker Cave (2)

Cumberland Crystal [ = Sloans Valley] Cave (2)

Hydens Cave (2 A, G, I)

Old Kentucky Cave (2)

Pourover Cave (2)

Wind Cave (2 A)

Rockcastle County

Duvalt Cave (2)

Fletchers Spring Cave (2)

Pine Hill Cave (2)

Teamers Cave (2 G)

Trigg County

Cool Spring Cave (6 G)

Warren County

By-Pass Cave (6)

Lost River Cave (6)

Pruitt Saltpeter Cave (6)

Wayne County

Blowing Cave (2)

Horse Hollow Cave (2 B, C, F)

Johnson Fork Cave (2)

Kogers Cave (2)

TENNESSEE

Fentress County

Buffalo Cave (1)

Sells Cave (1)

Wolf River Cave (1)

Franklin County

Caroline Cove Cave (1)

Putman Spring Cave (1)

Walker Spring Cave (1)

Wet Cave (1)

Grundy County

Bear Cave (1)

Big Mouth Cave (1 C)

Crystal Cave (1)

Wonder Cave (1)

Jackson County

Carter Cave (3 G)

Cherry Cave (3)

Haile Cave (3)

Montgomery County

Bellamy Cave (6)

Sinkhole Cave (6)

Overton County

•East Water Supply Cave (Sight record, T.C.B.)

Raven Bluff Cave (1)

Sheep Cave (1)

Putnam County

Blind Fish Cave (1 B, C, F, G)

Bridge Creek Cave (1 G)

Johnson Saltpeter Cave (1)

Warren County

•Blowing Cave (Sight record, T.C.B.)

Cumberland Caverns (1)

Turkeyscratch Cave (1)

White County

•Haskell Sims Cave (Sight record, T.C.B.)

Indian Cave (1)

Ross Cave (1)

Ward [ = Dairy House] Cave (1)

Wildcat Cove Cave (1)

COLLECTORS.—Listed below, alphabetically by the

initials used to identify them in the species treatments,

are the collectors of the specimens studied:

A.F. A. Fiske

A.R.B Andrew R. Boone

ARC. A. R. Cahn

A.W.D. Arthur W. Dobson

B. J. C. Beakley

B.B. Bret Blosser

B.D. Bert Denton

B.D.V. B. D. Valentine

C.E.B. C. E. Bush

C.E.M. Charles E. Mohr

C.E.Mc. C. E. McCary

C.H. C. Harrel

CM. Carl Melton

C.W.H. C. W. Hart, Jr.

D.A.E. D. A. Etnier

D.B. Donald Blair

D.E. D. Egbert

D.G.H. Dabney G. Hart

D.H.P. D. Hugh Puckett

D.P.B. David P. Beiter

E.C. Ellsworth Call

E.P.C. E. P. Creaser

F.N.B. F. N. Blanchard

G.H.E. G. H. Ehlers

H.H. H. Henrot

H.H.H. III H. H. Hobb« III

H.R.S. H. R. Steeves, Jr.

H.R.S. III H. R. Steeves HI

H.S. Herbert Shadowen

H.T. Hugh Thomas

J.E.C. J. E. Cooper

J.E.Cr. J. E. Crouch

J.E.Y. John E. Younglove

J.F.F. J. F. Fitzpatrick, Jr.

J.H. J. Harris

J.H.C. Jerry H. Carpenter
8 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

J.M.V. J. M. Valentine

J.N.D. J. N. Dent

J.P. J. Purcell

J.P.V. J. P. Voigt

J.R. J. Rhines

J.R.H. J. R. Holsinger

J.R.R. James R. Reddell

J.S. Jerry Stade

K.B. Kenneth Burdsall

K.H. K. Haas

K.S. Kjell Sandved

L.G. Leonard Giovannoli

L.G.C. Lloyd G. Carr

L.G.Co. Lyle G. Conrad

L.H. Leslie Hubricht

L.M. L. Merkle

M.B. Mark Becker

M.E.M. M. E. Mann

M.R.C. Martha R. Cooper

M.T. Merlin Tuttle

O.D.L. O. D. Lee

P.P. Peter Parker

R.A.K. R. A. Kuehne

R.B. R. Baroody

R.B.C.

R.G.

R.M.

R.M.N

R.N.

R.P.

R.R.

R.T.

R.W.

S.B.P.

S.M.

S.R.G.

T.C.B.

T.C.B.

T.C.J.

T.E.S.

T.G.M

V.E.S.

W.B.J.

W.L.M

R. B. Cumming

Richard Graham

R. McAdams

. R. M. Norton

Raymond Nelson

Rudolph Prins

Rendell Rhoades

R. Taylor

R. Walker

Stewart B. Peck

Sherman Minton

S. R. Gorin

T. C. Barr, Jr.

Ill T. C. Barr III

Thomas C. Jegla

T. E. Simpson

Terrence G. Marsh

V. E. Shelford

Walter B. Jones

[. W. L. Minckley

W.M.A. W. M. Andrews

W.P.H

W.R.E

W.T.

. W. P. Hay

. W. R. Eberly

William Torode

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