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number 20

June 1994

N.C. DOCUMENTS
CLEARINGHOUSE

AUG 1 5 1994 EDITORIAL STAFF

N.C. STATE L!BF^ARYR^CHARD a. Lancia, Editor

RALEIGH ^ ^ ^ . ..^.

Suzanne A. Fischer, Assistant Editor
Eloise F. Potter, Production Manager

EDITORIAL BOARD

James W. Hardin Rowland M. Shelley

Professor of Botany Curator of Invertebrates

North Carolina State University North Carolina State Museum

of Natural Sciences

William M. Palmer Robert G. Wolk

Director of Research and Collections Director of Programs

North Carolina State Museum North Carolina State Museum
of Natural Sciences of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State
Museum of Natural Sciences, appears twice yearly in consecutively
numbered issues. Subject matter focuses on systematics, evolution,
zoogeography, ecology, behavior, and paleozoology in the southeastern
United States. Papers stress the results of original empirical field
studies, but synthesizing reviews and papers of significant historical
interest to southeastern zoology are also included. Brief communications
are accepted.

All manuscripts are peer reviewed by specialists in the Southeast
and elsewhere; final acceptability is determined by the Editor. Address
manuscripts and related correspondence to Editor, Brimleyana, North
Carolina State Museum of Natural Sciences, P.O. Box 29555, Raleigh,
NC 27626. Information for contributors appears in the inside back
cover.

Address correspondence pertaining to subscriptions, back issues,
and exchanges to Brimleyana Secretary, North Carolina State Museum
of Natural Sciences, P.O. Box 29555, Raleigh, NC 27626.

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Environment,

Health, and Natural Resources

James B. Hunt Jr., Governor
Jonathan B. Howes, Secretary

CODN BRIMD 7
ISSN 0193-4406

Review of Biologically Significant Caves and their
Faunas in Florida and South Georgia

Richard Franz

Florida Museum of Natural History

University of Florida

Gainesville, Florida 32611

Judy Bauer

National Cave Diving Association

P.O. Box 14492

Gainesville, Florida 32604

AND

Tom Morris

2629 NW 12th Avenue
Gainesville, Florida 32605

ABSTRACT — At least 267 biologically significant caves have been
identified in Florida and south Georgia. Alachua, Jackson, Marion,
and Suwannee counties, Florida, contain over half of these
localities. The macroscopic troglobitic faunas in these caves
include 27 invertebrates and one vertebrate. The terrestrial component
consists of an insect and a spider; the rest are aquatic. One
branchiobdellid annelid and three entocytherid ostracod crusta-
ceans are obligate symbionts on troglobitic crayfishes and probably
should be considered troglobites. The rest of the region's re-
ported cave fauna is composed of 23 troglophiles, 47 trogloxenes,
and 37 accidentals. At least one bat {Myotis grisescens) is
regularly dependent on Florida caves for certain parts of its
life cycle, but leaves the cave environment to feed and mi-
grate. Two other cave-dependent bats {Myotis keeni and Myotis
sodalis) have been recorded only rarely from the region. Two
bats routinely use Florida caves, but also roost in other habi-
tats. The region's obligate cave species are grouped into six
cave faunas: Econfina Creek, Apalachicola, Woodville, Ocala,
St. Johns River, and Miami. Each fauna is restricted to a
particular karst region and is characterized by precinctive taxa.
The Ocala Fauna has the largest number of taxa (12) and
inhabits the largest geographic area, whereas the Econfina Creek
Fauna has the smallest number (2). The latter also is the only
fauna in the region that does not include at least one troglobitic
crayfish and amphipod. The two most complex faunas (Ocala
and St. Johns River) are subdivided into smaller assemblages.
Most Florida troglobitic taxa have been identified by the Florida
Committee on Rare and Endangered Plants and Animals as

Brimleyana 20:1-109, June 1994 1

Richard Franz, Judy Bauer and Tom Morris

deserving state and/or federal protection; currently one cave
crayfish is considered a Species of Special Concern by the
State of Florida; and the Squirrel Chimney Cave Shrimp is
listed as Threatened under provisions of the U. S. Endangered
Species Act of 1973, as amended.

Since 1894, naturalists have identified Florida springs, sinkholes,
and caves as habitats of plants and animals (Hubbard 1901; Lonn-
berg 1894, 1895; Faxon 1898; Hubbell 1936; Carr 1939; Hobbs 1940^,
1941, 1942fl, 19426; Young 1942, etc.). H. H. Hobbs, Jr. and H. B.
Sherman apparently were the first to attempt a systematic survey of
Florida's caves in order to enhance their studies of cave species (Hobbs
19426; Hobbs et al. 1977). Ferguson et al. (1947) followed with the
first catalogue of the springs of Florida, a work that was later updated
by Rosenau et al. (1977). In 1962, the Florida Speleological Society
published the first extensive inventory of caves of Florida and south
Georgia (Hippenmeier et al. 1962). This list included 188 caves in
Florida and south Georgia that could be entered using traditional caving
methods. The pioneering survey efforts of the Florida Speleological
Society also led to papers on the state's cave-associated vertebrates
(Pylka 1957) and aquatic troglobites (Pylka and Warren 1958, Warren
1961). The crayfish portion of Warren's 1961 list was updated by
Hobbs et al. (1977), Franz (1982), Franz and Lee (1982), Hobbs (1989),
and Franz and Franz (1990). Peck (1970) provided the first comprehensive
list of terrestrial arthropods from Florida, including the first records
of terrestrial troglobites from the state. Franz et al. (1971) presented
a list of gastropods from caves in the Marianna area of west Florida.

Since 1968, many important biological discoveries in Florida
caves have led to the description of 11 new troglobitic taxa. Most of
these discoveries resulted directly from efforts of the cave diving
community, particularly divers associated with the Cave Diving Section
of the National Speleological Society and National Association of
Cave Divers. Aided by advances in SCUBA equipment, divers penetrated
many previously inaccessible underwater cave systems. These discoveries
have enhanced our ability to interpret better the factors that have led
to the development of the region's complex aquatic troglobitic faunas.

This paper provides an updated list of the troglobitic species of
Florida and south Georgia. It summarizes distributional records for
both troglobitic and other cave-inhabiting species and comments on
the zoogeographic patterns exhibited by troglobitic species in the
region. The distributional patterns for cave crayfishes originally proposed
by Franz and Lee (1982) for the most part remain intact, i.e., (1)

Caves and their Faunas in Florida and South Georgia

faunas of the Apalachicola, Woodville, Ocala, and St. Johns River
karst areas are found in groundwater habitats associated with the Floridan
aquifer, primarily the upper Eocene Ocala Group limestones; (2) limestones
covered with deep sand and clay overburdens tend to lack troglobites;
and (3) the distribution of certain troglobites are correlated with the
relative levels of organic materials that collect in subterranean systems.

METHODS

We obtained information on caves and their faunas from many
sources. Over the years, dozens of cavers and cave divers provided
critical specimens of cave species and freely contributed information
on cave locations, maps, and descriptions, which have given us a
better appreciation of this region's complex cave faunas.

Precise locations of caves have been omitted from our paper
given the sensitivity of caves and their faunas to human disturbance.
If someone needs specific information concerning particular sites
they should contact the authors, members of the Florida Speleological
Society, Cave Diving Section of the National Speleological Society,
National Association of Cave Divers, or the staff of the Florida Natural
Areas Inventory (FNAI) (1018 Thomasville Road, Tallahassee, FL
32303).

We obtained voucher specimens from previously unsampled caves
in a variety of ways. Many samples were either caught by hand or
with dip nets. When possible, cave crayfishes were caught in minnow
traps that were baited with canned cat food. Cans were punctured and
placed inside the closed traps. This technique proved very successful,
e.g., two traps left for 24 hours in the main pool at Sunday Sink
(Marion County) attracted 125 crayfish (Procambarus lucifugus); however,
this technique was not effective in catching spider cave crayfishes
(Troglocambarus) in more than a hundred trapping attempts.

Terms — Common names for decapods follow Williams et al. (1989),
except when new distributional information or taxonomic interpretations
beg for the use of another name. We also propose common names for
recently described taxa that were not included in Williams et al.
(1989). Additional common names have been applied over the years
to a number of the taxa and are listed in the Other Common Names
section of appropriate species accounts.

We use an ecological classification for cavernicoles modified
from Peck (1970), Hobbs III (1992), and Holsinger (personal communication,
Old Dominion University). Troglobites (or stygobionts) are restricted
to caves or groundwater habitats and have obvious morphological
adaptations for subterranean habitats (e.g., reductions of the eye structure

Richard Franz, Judy Bauer and Tom Morris

and loss of pigmentation); troglophiles may complete their life cycles
in caves, as well as in certain non-cave habitats, and do not show the
extreme morphological adaptations that are usually reserved for troglobites;
trogloxenes regularly are encountered in caves but can not complete
their life cycles in them; accidentals are species that do not normally
inhabit caves but for one reason or another had taken refuge in specific
caves at the time of visitation.

We follow Frank and McCoy (1989) in their uses of the terms
indigenous and precinctive, rather than native and endemic. Based on
their interpretations, "indigenous" is preferred to "native" because
the latter has "subsidiary meanings in English;" "precinctive" is restricted
to taxa that are indigenous and "known from no other area."

HISTORY OF FLORIDA BIOSPELEOLOGY
Early Period (1893-1897)— Florida biospeleology began in 1893
with the discovery of white crayfishes in a hand-dug well at Lake
Brantley, near Orlando, Seminole County. This crustacean was named
Cambarus acherontis by its collector, the Swedish naturalist Einar
Lonnberg (Lonnberg 1894, 1895).

The next discoveries occurred in 1894 while H. G. Hubbard, the
noted entomologist who worked as a Special Agent of the U.S. Entomological
Commission (United States Department of Agriculture), was visiting
colleagues, W. T. Webber and H. J. Swingle at Eustis, Florida. According
to Hubbard (1901:395-396), after receiving word of caves in Hernando
and Citrus counties, they traveled the 30 or 40 miles (48-64 km) from
Eustis to explore caves at "Istachatta on the Withlacoochie River"
and "Double Hammock country, in Citrus County." At Double Hammock,
they encountered a large "cavern, 75 to 100 feet deep, in a hillside of
open pine woods." They noted "white crawfish very much like those
in the Mammoth Cave," bats, streblid flies (Trichobius major), mites,
"hairy Muscid" (fly), minute black gnats, spiders, "Hemipteron," molds,
and cave fungi. They named this cave Gum Tree Cavern (later called
Gum Cave or Sweet Gum Cave) because of a large sweet gum tree
that grew on the verge of the sinkhole entrance. Their collection of
white crayfishes from this cave currently resides in the crustacean
collection at the U.S. National Museum of Natural History. Faxon
(1898) examined these specimens, but failed to recognize their unique-
ness. He assigned them to Lonnberg's Cambarus acherontis, an
interpretation that was followed by Harris (1903) and Ortmann (1902,
1905). The Gum Tree Cavern specimens were eventually described as
Cambarus (=Procambarus) lucifugus lucifugus by Horton H. Hobbs,
Jr. (1940fl). In addition, there was a single specimen of Troglocambarus

Caves and their Faunas in Florida and South Georgia

maclanei in the original Gum Tree Cavern collection that escaped the
attention of Faxon and other workers and remained unnoticed until
Martha R. Cooper rediscovered the specimen mixed with the others
over 80 years later! Webber and Swingle also provided additional
specimens of white crayfishes to the U.S. National Museum of Natural
History, but there is some question as to their origins. The collection
data stated only that they were received from Swingle and Webber
from Eustis, Lake County, Florida, in November 1897. It is not known
whether the specimens originated in the Eustis area or were collected
elsewhere and shipped from Eustis. Hobbs (1940^) considered these
specimens similar to his Cambarus lucifugus from Gum Cave but
indicated that they possessed certain unique features. This led him to
suggest that they probably represented an unidentified subspecies of
lucifugus. As far as we are aware, there have been no further collections
of this unique crayfish.

Hobbs Period (1935-1946) — Little happened in Florida speleology
between 1897 and 1935. In 1931, Horton H. Hobbs, Jr. began his long
and productive scientific career as an undergraduate student at the
University of Florida (Hobbs 1986). Upon graduation in 1935, he
enrolled in the master's program at Florida where he began his studies
on Florida crayfishes under the direction of Dr. J. Speed Rogers.
Hobbs was encouraged first to survey the crayfishes of Alachua County
for his master's degree (Hobbs 1936), before launching into his definitive
survey of the state's crayfish fauna for his Ph.D., which he completed
in May 1940 (Hobbs 1940/?). As part of this scholastic program, he
became one of the leaders in the natural history revolution that occurred
at the University of Florida during the pre-World War II years. Hobbs
was the third person to earn a Ph.D in biology from this institution,
following Archie Carr and H. K. Wallace.

In March 1935, Hobbs found his first cave crayfishes
(Procambarus pallidus) in "... a small area of subterranean water
exposed in the bottom of a cavelike lime sink in the southern part of
Columbia County" (=Riverbed Cave) (Hobbs 1940^, 1986). This site
was located in an abandoned stream valley at the base of the railroad
embankment along U.S. Highway 27 west of High Springs. We have
searched the immediate area around the railroad embankment but were
not successful in relocating an exposure of subterranean water at this
site; however, we have found troglobites in other sinks north of the
embankment further up the valley.

Between 1936 and 1942, Hobbs collected additional material of
Procambarus lucifugus lucifugus at Sweet Gum Cave in Citrus County,

Richard Franz, Judy Bauer and Tom Morris

the site of Hubbard's original collection in 1894, with the assistance of
H. B. Sherman and other colleagues. Hobbs (1940«) also reported a
second collection of this crayfish from an unidentified cave, 23.3 km
(14 miles) north of Weekiwachee Springs in Hernando County, obtained
by Albert Greenburg in 1937.

The first specimens of Procambarus lucifugus alachua were taken
in Alachua County at Hog Sink in November 1937 (Hobbs 1940(2) and
at Goat Sink in January 1938. During the following November, Hobbs
visited Palm Springs in Seminole County and for the first time since
the original collection by Lonnberg he obtained additional material of
Procambarus acherontis in the spring pool (Hobbs 19426, 1986). Following
these successes, Hobbs and Dr. Sherman used aerial photographs to
locate sinkholes and other depressions, then visited the sites to determine
whether they led to subterranean water. This systematic survey in the
vicinity of Gainesville, Alachua County, led to the discovery of sites
such as Squirrel Chimney (Hobbs et al. 1977). Between 1937-1942,
students and friends, particularly William M. McLane, Lewis J. Marchand,
and Alphonse C. Chable, accompanied Hobbs during his explorations.
Other companions included J. Adams, W. Beck, L. Berner, A. Carr,
T. Carr, J. C. Dickinson, C. S. Goodnight, M. L. Goodnight, B. J.
Kaston, J. Kilby, J. M. Martin, C. Mohr, G. Pournelle, H. B. Sherman,
K. Spurr, H. K. Wallace, and F. N. Young. McLane discovered the
first specimens of the Northern Spider Cave Crayfish at Squirrel Chimney
in March 1941, which Hobbs named Troglocambarus maclanei in
McLane's honor (see Hobbs et al. 1977 for a discussion of this discovery).

In 1939, Archie Carr obtained the original specimen of the cave
salamander, Haideotriton wallacei. This specimen was retrieved by
Mr. Hummel, sanitary engineer with Dougherty County (Georgia),
from a "200-foot well" in Albany, Georgia, after it was air-lifted to
the surface (Carr 1939). The animal was named in honor of H. K.
Wallace, the spider expert, Carr's colleague and friend at Florida,
and Hobbs' brother-in-law. The holotype was sent alive through the
mail in a Mason jar to H. K. Wallace who brought it to the attention
of Archie Carr (H. K. Wallace, personal communication, [retired]
University of Florida). Wallace had met Mr. Hummel when he and A.
P. Black were consulting with city engineers in Albany, Georgia, in
early May 1939 (H. K. Wallace, personal communication).

World War II interrupted the flow of natural history research at
Florida, although several students returned after the war to finish
their studies. Hobbs continued to teach at Florida until 1946 when he
moved to the Biology Department at the University of Virginia. He
served as the Director of the Mountain Lake Research Station between

Caves and their Faunas in Florida and South Georgia

1956-1960, and then moved on to the Department of Zoology at the
U.S. National Museum of Natural History in 1962. Although retired
from the Smithsonian since 1984, he retains his emeritus status there
and continues to pursue his studies on the systematics of cray-
fishes, ostracods, and other crustaceans. Between 1940-1942, Hobbs
described Cambarus (=Procambarus) lucifugus lucifugus, Cambarus
{-Procambarus) lucifugus alachua, Cambarus (=Procambarus)
pallidus, Troglocambarus maclanei, and Cambarus cryptodytes
(Hobbs 1940fl, 1941, 1942a). His collections of other cave organisms
from Alachua County caves led to the descriptions of the isopod Asellus
(=Caecidotea) hobbsi (Maloney 1939) and the amphipod Crangonyx
hobbsi (Shoemaker 1941). Hobbs' early crayfish studies culminated
with the publication of his treatise Crayfishes of Florida (Hobbs \942b).
This book continues to be the single most important reference on the
state's crayfish fauna.

Post-Hobbs Period and the Florida Speleological Society (FSS)
(1946-1969) — Little activity occurred in Florida biospeleology im-
mediately following Hobbs' departure from the state. In 1954, Pirkle
and Babb obtained a specimen of Procambarus pallidus from a well
at Four O'Clock Church (Fort Clark) in Alachua County. Between
1952-1954, Robert B. Cumming, while an undergraduate student at
the University of Florida, collected troglobitic crustaceans. His Florida
cave collections included Troglocambarus maclanei from Squirrel Chimney
and Sweet Gum caves in 1953 and 1954, respectively; Procambarus
lucifugus and Procambarus pallidus from Eichelberger Cave in 1952;
and the first specimen (a female) of Palaemonetes cummingi from
Squirrel Chimney in Alachua County in 1953 (Chace 1954). The presence
of Procambarus pallidus in Eichelberger Cave later was questioned
by Franz and Lee (1982) because this locality is 40 km from the next
closest documented site, and no other specimens of this crayfish have
been collected from caves in the vicinity of Eichelberger Cave before
or since Cumming's specimens were obtained. Eichelberger Cave has
been destroyed by quarrying activities. Franz and Lee (1982) suggested
that the collection had been mixed accidentally, and the Procambarus
pallidus specimens actually were collected elsewhere. Search should
be continued for the species in Marion County.

A second inventory of Florida and south Georgia caves began in
1949 with the establishment of a caving club in Gainesville. The survey
was expanded to include biological and paleontological surveys when
the club became a University of Florida student grotto of the National
Speleological Society in 1952.

Richard Franz, Judy Bauer and Tom Morris

In 1956, Peter Drummond, an early member of the Florida Speleological
Society, collected the first specimens of Cambarus cryptodytes from
Climax Cave, Decatur County, Georgia, and Horst R. H. Heineman
obtained specimens of what Hobbs initially identified as Procambarus
pallidus {=Procambarus orcinus) at Clay Sink (probably Gopher Sink)
in Leon County, Florida. Richard D. Warren was active with the
Florida Speleological Society between 1956-1965. He contributed specimens
of Procambarus lucifugus X alachua from Roosevelt Cave in 1960;
Procambarus erythrops from Hildreth Cave in 1962; and Troglocambarus
maclanei from Indian Cave in 1962. He also collected specimens of
Cambarus cryptodytes in 1961 and 1963 at Climax Cave, the type
series of Asellus {=Remasellus) parvus from Ten Inch Cave in 1962,
and with Bousfield the type series of Crangonyx grandimanus from
Indian Cave (Marion County) in 1962. Gerard M. Miller and Warren
were the first to discover Haideotriton wallacei in Florida. They initially
recovered specimens from Gerard's Cave in Jackson County in 1957
(Pylka and Warren 1958), and later found this salamander in Judge
Cave and Washed-out Cave in Florida and at Climax Cave in Georgia
(Warren 1961). Additional specimens from Climax Cave were secured
by Alberta Etters (Smith) and Vernal Harkness in 1960. A brief list of
cave-associated vertebrates (without specific localities) was provided
by Pylka (1957). At the same time. Dale W. Rice and William L.
Jennings were investigating the distribution and ecology of cave bats,
particularly in the Marianna area (Jennings and Layne 1957; Jennings
1958; Rice 1955fl, 1955Z), 1957).

Biospeleologists John E. and Martha R. Cooper visited Richard
D. Warren in 1964-1965. The Coopers obtained specimens of Procambarus
acherontis in 1964 and 1965 at Palm Springs. The initial specimens
were netted from aquatic vegetation in the spring pool; a second
series was acquired from the pool by the property manager of the
spring property and given to the Coopers when they returned to Palm
Springs in 1965 (Cooper 1965^). With Warren, they also visited Goat
Sink, Protheroe Sink, and Still Sink where they obtained samples of
troglobitic crayfishes (Cooper 19656).

Frank Hurt retrieved the first record of Cambarus cryptodytes
from the cave at Waddell's Mill Pond in 1965. S. B. Peck made a
collection of Procambarus pallidus at Warren Cave in 1965. This
appears to be the first collection of the species from Warren Cave
since Hobbs' original specimens. Notes associated with the collection
indicated that the cave had been dry for 4 years prior to his finding
them. Peck also obtained specimens of Cambarus cryptodytes at Climax
Cave (Hobbs 1981) and two new staphylinid beetles at Miller's Cave

Caves and their Faunas in Florida and South Georgia

(Klimaszewski and Peck 1986). Peck's inventory of Florida caves
included three collembolans, three orthopterans, six beetles, two opilionids,
five spiders, one millipede, and one centipede (Peck 1970).

David S. Lee, then a student at Florida Southern College, started
to visit caves in Alachua, Citrus, and Jackson counties, Florida, in
1965 (D. S. Lee, personal communication, North Carolina State Museum
of Natural Science). Between 1965 and 1970, he discovered many
new biologically significant sites, particularly in northwest Florida,
and made several important collections, one of which was the first
and only ovigerous female Palaemonetes cummingi from Squirrel Chimney.
He conveyed this specimen alive to Sheldon Dobkin at Florida Atlantic
University who successfully reared the developing larvae and published
the first description of the larval development for this species (Dobkin
1971). Merlin Tuttle also visited caves in Jackson County during this
same period in pursuit of gray bats (Myotis grisescens). Lee and Tuttle
combined their expertise and encouraged the Florida Park Service to
gate Old Indian Cave at Florida Caverns State Park to protect its
important bat colony (Lee and Tuttle 1970), and Lee later encouraged
them to limit access to other biologically sensitive caves in the Park.
Lee introduced Richard Franz to Florida biospeleology in 1967. On
one of the trips to Florida from Maryland, Lee and Franz visited
Squirrel Chimney in hopes of obtaining additional specimens of the
Squirrel Chimney Cave Shrimp, but instead, rescued a large eastern
diamondback rattlesnake {Crotalus adamanteus) from the bottom of
the vertical entry shaft (Franz 1968). Lee and Franz continued to
travel to west Florida between 1968 and 1970 to explore caves in
Jackson and Washington counties. Data from this period provided the
basis for studies on the predatory snail Euglandina rosea (Franz et al.
1971) and on the cave salamander, Haideotriton wallacei, and other
vertebrates in Jackson County caves (Lee \969a, \969b, 1969c, \969d,
1976). In 1970, they collected the first specimens of Cambarus cryptodytes
and Haideotriton wallacei from Pool Cave in Florida Caverns State
Park. The next year, Franz took Archie Carr to this cave and showed
him his first live Haideotriton since he had described the salamander
in 1939. In 1972, Franz joined the faculty of the Florida State Museum
(recent name change to Florida Museum of Natural History) at the
University of Florida where he currently continues his cave studies.

Modern Period (1970-1992) — The discovery of a new troglobitic
crayfish in a Miami well in 1968 rekindled interest in the state's cave
crayfish fauna. This species, eventually described as Procambarus
milleri (Hobbs 1971), was the first new cave crayfish found in Florida

10 Richard Franz, Judy Bauer and Tom Morris

since 1942 and the first new troglobite since the descriptions of Crangonyx
grandimanus (Bousfield 1963) and Asellus parvus (Steeves 1964).
The discovery of the Miami Cave Crayfish set the stage for the disclosure
of four other crayfish discoveries in the 1970s.

D. Bruce Means, who began visiting northwest Florida caves in
the mid-1960s, made numerous important collections of cave crayfishes
and vertebrates from caves and sinks in the Tallahassee and Marianna
areas. Means made several collections of a new crayfish at Gopher
Sink and Culley's Cave in 1970 and 1971, which he and Hobbs later
named Procambarus orcinus (Hobbs and Means 1972). Other collectors
who contributed specimens used in the type description of this crayfish
included H. R. H. Heineman (Clay Sink, 1956), J. Halusky (Gopher
Sink, 1970), J. Bishop (Osgood Sink, 1968), J. Couch (cave 3 miles
(4.8 km) south of Woodville, 1962), and L. B. Trott (Wakulla Springs,
1957). Specimens of a second new crayfish, collected first by Michael
N. Horst at Big Blue Spring on the Wacissa River in Jefferson County
in 1970, were described as Procambarus horsti in the same paper
with Procambarus orcinus (Hobbs and Means 1972).

Barry Mansell and Frank Hurt collected the first specimens of a
new cave crayfish at Sim's Sink {Procambarus erythrops Relyea and
Sutton, 1975) in Suwannee County in 1971 (B. Mansell, personal
communication, Jacksonville, Florida); they also took specimens of
Troglocambarus maclanei from Sim's Sink, Procambarus lucifugus
from Bat Cave (Alachua County), Sweet Gum Cave, and Indian Cave
(Marion County), and Procambarus pallidus from Squirrel Chimney
in 1971-1972. Mansell and Bruce Sutton collected specimens of Haideo-
triton wallacei from Gerard's cave in 1969. Kenneth Relyea and Sutton's
explorations of north Florida caves were contemporary with those of
Means, Mansell, and Hurt. They collected additional material of Pro-
cambarus erythrops from Sim's Sink in 1971-1972; Procambarus pallidus
from Pallidus Sink, Squirrel Chimney, and Martin Cave in 1972; and
Troglocambarus maclanei from Sim's Sink in 1975. The new material,
plus those collected by Mansell and Hurt, from Sim's Sink allowed
Relyea and Sutton to describe Procambarus erythrops (Relyea and
Sutton 1975). Relyea also obtained two specimens of a unique crayfish
from Alexander Springs in Lake County in 1973 and 1974 (Relyea et
al. 1976) that later was described as Procambarus delicatus following
the collection of a third specimen in 1985 by J. B. Smith and D.
Haren H (Hobbs and Franz 1986).

In 1973, Stephen R. Humphrey and Franz visited Orange Lake
Cave in Marion County for the first time after cavers reported large
numbers of bats present in the cave. Lee and Franz returned to this

Caves and their Faunas in Florida and South Georgia 11

site in 1974-1975 and collected several series of crayfishes. They
were later described as Procambarus franzi (Hobbs and Lee 1976).
Subsequent collections at Hell Hole and Trade Wind Farm's Sink
extended the known range of this species several kilometers south
toward the Reddick area. Chert Cave and Sunday Sink, south of Ocala,
were found to have large populations of Procambarus lucifugus intergrades
and Troglocambarus maclanei. The latter was located as a result of
surveys associated with the Cross-Florida Barge Canal study. Ray
Ashton and Pat Sawyer Ashton collected the first specimens of Procambarus
lucifugus intergrades from Ocala Caverns in 1976.

The momentum established in the 1970s continued into the 1980s
and 1990s. Additional discoveries led to the descriptions of Procambarus
leitheuseri in Hernando and Pasco counties (Franz and Hobbs 1983),
Procambarus delicatus in Lake County (Hobbs and Franz 1986), Procambarus
morrisi in Putnam County (Hobbs and Franz 1990), Procambarus attiguus
in Marion County (Hobbs and Franz 1992), and Dasyscias franzi in
Washington County (Thompson and Hershler 1991). During this period,
there were also collections of several crustaceans that still remain
unstudied (e.g., two Caecidotea from Orange and Washington counties
and a Troglocambarus from Orange County). Recent collections of
Steeves' Asellus parvus at Split Sink and Peacock Springs allowed
Bowman and Sket to recognize the uniqueness of this isopod for
which they erected the new genus Remasellus (Bowman and Sket
1985).

Throughout this period, cavers and divers continued to provide
many other important specimens that added tremendously to our knowledge
of the distributions of Crangonyx grandimanus, C. hobbsi, Procambarus
acherontis, P. horsti, P. lucifugus, P. orcinus, P. pallidus, and Troglocambarus
maclanei. Most recently, Buford Pruitt has taken many important voucher
specimens of Procambarus lucifugus, P. pallidus and Troglocambarus
maclanei in underwater caves of Levy and Hamilton counties. Pruitt
also generously purchased and donated Sim's Sink to The Nature
Conservancy as a cave crayfish preserve in 1987.

BIOLOGICALLY IMPORTANT CAVES

We recorded a total of 267 caves in Florida and south Georgia
where biological materials have been recovered (Appendix 1). More
than half are concentrated in Alachua (47 caves), Jackson (34 caves),
Marion (27 caves), and Suwannee (43 caves) counties (Fig.l). Other
biologically significant caves are found in Columbia (15), Levy (11),
Lafayette (11), Leon (11), Hamilton (11), Gilchrist (9), Wakulla (8),
Citrus (7), Hernando (6), Madison (5), Orange (4), Pasco (4), Seminole

12

Richard Franz, Judy Bauer and Tom Morris

Fig. 1. Map of Florida showing county locations. Numbers of caves
are shown for Alachua, Jackson, Marion, and Suwannee counties. Dots
indicated other important cavernous counties.

(3), Lake (2), Washington (2), Dade (2), Holmes (1), Jefferson (1),
Pinellas (1), and Putnam (1) counties, Florida, and Decatur (1) and
Doughtery (1) counties, Georgia. Caves theoretically could exist anywhere
in Florida and south Georgia since most of this region is underlain
with extensive beds of Oligocene and Eocene limestones. These limestones
are components of the Floridan aquifer, while limestones in Dade
County are part of the Biscayne aquifer.

Caves and their Faunas in Florida and South Georgia 13

There are no reports of troglobites being recovered from especially
deep wells such as those reported in the San Antonio Pool of the
Edwards aquifer, Texas (blind catfishes, genera Satan and Trogloglanis,
at groundwater depths between 305-582 m) (Cooper and Longley 1919a,
19796). The deepest record of a Florida troglobite is for Procambarus
orcinus (and possibly Procambarus horsti) at Wakulla Springs at about
100-m water depth. Other depth records include 67.6 m (231 ft) for
Procambarus leitheuseri at Eagle's Nest Sink (Hernando County);
9 m (28.8 ft) for Procambarus milleri at the well northeast of Home-
stead; 44 m (150 ft) and 36.7 m (125 ft) for Procambarus pallidus at
Fort Clark Church well and Hornsby Sink, respectively (Alachua
County); 67.6 m (231 ft) for Troglocambarus maclanei at Eagle's Nest
Sink (Hernando County); 26.4 m (80 ft) for Cambarus cryptodytes at
Hole-in-Wall (Jackson County); and 60.8 m (200 feet) for Haideotriton
wallacei at the Albany, Georgia, well.

Franz and Lee (1982) reasoned that limestones covered with
thick layers of unconsolidated marine or aeolian sediments lack sufficient
food imput to sponsor colonizing organisms. However, recent discoveries
of certain colonial bacteria in a few underwater caves might provide
limited grazing for troglobitic crustaceans (Hobbs and Franz 1992).

The most cavernous parts within the study region are the mature
karst areas in western Alachua and Marion counties, eastern Citrus,
and northcentral Jackson County, Florida. The current list maintained
by the Florida Speleological Society for this four-county area includes
430 of 630 caves reported from Florida (Al Krause, personal communica-
tion, Florida Speleological Society). Caves in mature karst areas tend
to have portions of their passages above the local water table. As a
result many of the larger caves are inhabited by cave-dependent bats
which supply large quantities of guano to fuel the biotic system.
Above-water caves in the Citrus, Marion, and Jackson counties tend
to be shallow and have meandering passages and multiple entrances,
whereas many in Alachua County are deeper and have vertical shafts
as entrances that require technical climbing equipment to negotiate.
Some of the Alachua entrance holes are over 17.6 m (50 ft) deep.
Speleothem development is rare in most Florida caves, although some
caves in Citrus, Marion, and Jackson counties have interesting displays.

Saturated riverine karsts are developed along portions of the
Choctawhatchee, (western) Econfina, Chipola, Flint (Georgia), St. Marks-
Wakulla, Wacissa, Suwannee, and St. Johns rivers and their tributaries.
Caves within these karsts have little or no dry passage. Access to
them is usually through spring outlets, spring-siphons, or water-filled
sinkholes.

14 Richard Franz, Judy Bauer and Tom Morris

Rosenau et al. (1977) listed over 300 springs in 42 Florida counties,
most of which have their water sources in the Floridan aquifer, particularly
the Ocala Group of Eocene limestones. Twenty-seven of these springs
have recorded water flows that average six billion gal/day each and
are considered first magnitude springs (Rosenau et al. 1977). Counties
with saturated riverine karsts where biological specimens have been
retrieved include Columbia, Dade, Gilchrist, Hamilton, Hernando, Jefferson,
Lafayette, Lake, Leon, Levy, Madison, Orange, Pasco, Pinellas, Seminole,
Suwannee, Wakulla, and Washington. Extensive underwater cave systems
found in these counties form important conduits for local groundwater
circulation. Knab (1991) noted that six of the ten longest known
underwater caves in the world are located in the saturated karsts of
the Suwannee and Wakulla drainage basins in Florida — Falmouth Spring
(or Cathedral Sink), 3,291.8 m (10,828.3 ft); Chip's Hole, 3,169.9 m
(10,427.3 ft); Sullivan's Sink-Cherry Sink, 2,590.8 m (8,522.4 ft)
(Leon County, Wakulla River); Manatee Springs, 2,342.4 m (7,705.3
ft) (Levy County, Suwannee River); Luraville-Telford Spring, 2,194.5
m (7,218.7 ft) (Suwannee County, Suwannee River); Hornsby Sink,
2,055.3 m (6,760.8 ft) (Alachua County, Suwannee River). Recent
explorations of the Leon Sinks complex under Leon and Wakulla
counties have revealed over 14,288 m (47,000 ft) of continuous mapped
passage (Gary Knecht, personal communication, Tallahassee, Florida).

SPECIES ACCOUNTS
FOR FLORIDA AND SOUTH GEORGIA TROGLOBITES

Phylum MOLLUSCA

Class GASTROPODA

Order PROSOBRANCHIA

Family Hydrobiidae

Dasyscias franzi Thompson and Hershler

SHAGGY GHOSTSNAIL

Dasyscias franzi Thompson and Hershler, 1991. Malacological

Review 24:57-61. TYPE LOCALITY: Blue Spring Cave System, Econfina

River, Washington County, Florida. Holotype (UF 93964), 27 paratypes

(UF).

DISTRIBUTION: ECONFINA CREEK FAUNA. Known only from
the type locality.

ETYMOLOGY: Named for Richard Franz, discover of the snail.
REFERENCES: Thompson and Hershler 1991 (original description,
SEM photograph).

Caves and their Faunas in Florida and South Georgia 15

Phylum ARTHROPODA

Class MALACOSTRACA

Order ISOPODA

Family Asellidae

Caecidotea hobbsi (Maloney)

HOBBS' CAVE ISOPOD

Asellus hobbsi Maloney, 1939. Proceedings of U.S. National Museum
68(3057):457. TYPE LOCALITY: Dudley Cave, Alachua County, Florida.
Holotype (USNM 76434), Horton H. Hobbs, Jr. (coll.), 31 October
1937. Paratypes from Dudley Cave and from crayfish burrows at Blountstown,
Calhoun County, Florida.

Caecidotea hobbsi. — Bowman, 1975:339-340.

DISTRIBUTION: APALACHICOLA (Marianna Lowlands) and
OCALA FAUNAS (Upper Suwannee, Marion); also northcentral Georgia.
Known from groundwater habitats in Alachua, Calhoun, Jackson and
Marion counties, Florida, and in DeKalb County, Georgia. This species
is not restricted to limestone areas and may be more widely-distributed
than records indicate. It probably lives in interstices saturated with
groundwater in unconsolidated sediments, as well as in cave pools,
similar to other members of the Hobbsi group (Lewis 1982, Lewis
and Holsinger 1985). The isopod was listed as a Species of Special
Concern by the Florida Committee on Rare and Endangered Plants
and Animals because of its restriction to groundwater habitats and its
apparent rarity (Franz 1982).

SPECIFIC LOCALITIES: FLORIDA, A/ac/^wa County. Aulsbrook
Cave (RF), Bat Cave (USNM), cave 21.7 km (13 mi) west of Gainesville
(probably Dudleys Cave) (USNM), Dudley Cave-type locality (USNM,
Maloney 1939, Hobbs 1942^), well near Micanopy (USNM). Calhoun
County: 3.6 km (2 mi) south of Altha, near Blountstown (burrow of
the Apalachicola Burrowing Crayfish, Procambarus rogersi) (USNM).
Jackson County. Gerard's Cave (USNM). Marion County. Hollowed
Ground Cave? (J. Lewis), Rainbow Acre's Cave (USNM), Roosevelt
Cave (USNM). GEORGIA, DeKalb County, spring on Walter Chandler
Estate at Emory University (USNM).

ETYMOLOGY: Named in honor of Horton H. Hobbs, Jr., the
collector of the type series.

REMARKS: The Georgia record constitutes a major range extension
for this species. Their identification was confirmed by T. E. Bowman
(personal communication, U.S. National Museum of Natural History).

REFERENCES: Franz 1982 (conservation status), Harris 1968
(as prey); Hobbs 1942Z? (records); Lee 1969^ (as prey); Maloney 1939

16 Richard Franz, Judy Bauer and Tom Morris

(type description), Steeves 1964 (records), 1966 (taxonomy), Warren
1961 (records, photograph).

Caecidotea sp. 1
REMARKS: Specimens of asellid isopods were collected on floating
wood in the cave stream of the Econfina Blue Spring Cave System in
Washington County, Florida (ECONFINA CREEK FAUNA). According
to J. Lewis (personal communication, Louisville, Kentucky), this species
is closely related to Caecidotea alabamensis and C. nickajackensis,
which are found at cave sites in northern Alabama, and in all probability
it represents an undescribed taxon. Additional collecting is necessary
in order to resolve this isopod's taxonomic assignment (J. Lewis,
personal communication).

Caecidotea sp. 2
REMARKS: An undescribed asellid isopod was collected by Roger
Werner in Rock Springs Cave, Orange County, Florida (USNM) (ST.
JOHNS RIVER FAUNA, Wekiva). Its relationships with other Caecidotea
are unstudied (T. E. Bowman, personal communication).

Remasellus parvus (Steeves)
SWIMMING FLORIDA CAVE ISOPOD

Asellus parvus Steeves, 1964. American Midland Naturalist 71(2):450-
451. TYPE LOCALITY: Ten Inch Cave, 8.4 km (5 mi) of Newberry,
Alachua County, Florida. Holotype and allotype (USNM 111142), a
paratypic female (USNM 111140) and paratypic male (USNM 111141),
R. D. Warren (coll.), 9 June 1961.

Remasellus parvus. — Bowman and Sket, 1985:554.

DISTRIBUTION: WOODVILLE and OCALA (Upper Suwannee)
FAUNAS. Known from groundwater habitats in karst areas of the
Wakulla and upper Suwannee rivers. It may be more common than
collections indicate.

SPECIFIC LOCALITIES: Alachua County. Ten Inch Cave-type
locality (USNM). Madison County: Thunderhole Sink (USNM). Suwannee
County: Peacock Springs Cave System (USNM). Wakulla County: Split
Sink (USNM).

ETYMOLOGY: Remasellus from remus=oar (Latin) + Asellus,
referring to the shape and function of pereopod 2-7 (Bowman and
Sket 1985); parvus refers to small; so named because of its relatively
small size (Steeves 1964).

Caves and their Faunas in Florida and South Georgia 17

REMARKS: Unlike Caecidotea hobbsi, this isopod may be restricted
to cave habitats in light of its unusual morphology and swimming
mode of locomotion.

REFERENCES: Bowman and Sket 1985 (definition of Remasellus,
records); Steeves 1964 (original description).

Order AMPHIPODA

Family Crangonyctidae

Crangonyx grandimanus Bousfield

FLORIDA CAVE AMPHIPOD

Crangonyx grandimanus Bousfield, 1963. National Museum of
Canada, Natural History Paper No. 18:1-9. TYPE LOCALITY: Indian
Cave, 11.7 km (7 mi) southwest of Ocala, Marion County, Florida.
Holotype (NMC 5002), R. D. Warren (coll.), 18 February 1961. Paratype
(NMC 5003) from Huggin's Cave, Alachua County, R. D. Warren
(coll.), December 1961.

DISTRIBUTION: WOODVILLE, OCALA (Upper Suwannee, Lower
Suwannee, Marion, Gulf Coastal Lowlands) and MIAMI FAUNAS.
This amphipod was listed as a Species of Special Concern by the
Florida Committee on Rare and Endangered Plants and Animals (Franz
1982) because of its subterranean habits and limited distribution, and
as a candidate for federal protection (Wood 1992). It appears to be
widespread in groundwater habitats associated with limestone areas
on the western slope of the old Northern Highlands in the Florida
peninsula. This amphipod is known from the Woodville Karst in the
St. Marks-Wakulla River basin, northern peninsula karsts from the
Suwannee River south to Pasco County and from the Miami karst;
however, it is rare at most sites. This amphipod has not been collected
on the eastern slope in the St. Johns River valley.

SPECIFIC LOCALITIES: Alachua County: Dudley Cave (JRH),
Goat Sink (JRH), Hertzog Cave (JRH), High Springs Cave (JRH),
Huggins Cave (JRH), well (Archie Carr Farm) near Micanopy (JRH).
Citrus County. Sweet Gum Cave (JRH). Dade County: well. Little
Bird Nursery and Garden Store, Miami (JRH). Gilchrist County: Devil's
Eye and Ear Springs (JRH). Hernando County: Eagle's Nest Sink
(JRH). Leon County: Little Dismal Sink (JRH). Levy County: Archer
Cave (JRH), well at Chiefland (JRH). Madison County: Madison Blue
Spring (JRH). Marion County: Indian Cave-type locality (Bousfield
1963), well 3.6 km (2 mi) northeast of Anthony (JRH). Pasco County:
Nexus Sink (JRH). Suwannee County: Cisteen Sink (JRH), Orange
Grove Sink (JRH), Peacock Springs Cave System (JRH). Wakulla County:

18 Richard Franz, Judy Bauer and Tom Morris

Emerald Sink (JRH), McBride Slough (JRH), River Sinks (JRH), Sally
Ward Spring (JRH), Shepard Blue Spring (JRH).

ETYMOLOGY: Although Bousfield (1963) did not indicate the
origin of the name "grandimanus" in the original description, we
presume that he was making reference to the "very large gnathopods"
that characterize this species.

REMARKS: Holsinger (1972) listed this amphipod as a member
of the obliquus-richmondensis group.

REFERENCES: Bousfield 1963 (type description); Franz 1982
(conservation status); Holsinger 1972 (records, key), 1977 (taxonomy).

Crangonyx hobbsi Shoemaker
HOBBS' CAVE AMPHIPOD

Crangonyx hobbsi Shoemaker, 1941. Charleston Museum Leaflet
16:457. TYPE LOCALITY: Huggins Cave, Alachua County, Florida.
Holotypic male (USNM 79362), paratypic female (USNM 109623),
Horton H. Hobbs, Jr. (coll.).

DISTRIBUTION: WOODVILLE, OCALA (Upper Suwannee, Lower
Suwannee, Orange Lake, Marion, Withlacoochee, Gulf Coastal Lowlands),
and MIAMI FAUNAS. This amphipod is widespread in limestone
areas of the Florida peninsula, east of the Apalachicola River, and on
occasions can be abundant at certain sites. Its distribution coincides
with that of Crangonyx grandimanus. The species was listed as a
Species of Special Concern by the Florida Committee on Rare and
Endangered Plants and Animals (Franz 1982) because of its dependence
on specialized subterranean habitats and its limited distribution; it is
a candidate for federal listing (Wood 1992).

SPECIFIC LOCALITIES: Alachua County. Cave in riverbed at
High Springs (=River Bed Cave?) (JRH), Crumbly Sink (JRH), Devils
Hole (JRH), Dudleys Cave (JRH), Goat Sink (JRH, Hobbs 19426),
High Springs Cave (JRH), Huggins Cave-type locality (USNM, Shoemaker
1941, Bousfield 1963), Still Sink (Warren 1961, not seen by JRH)),
well (Archie Carr Farm) near Micanopy (JRH). Citrus County. Sweet
Gum Cave (JRH, Hobbs 1942/?). Columbia County. Bussey's Sink
(JRH), River Bed Cave (Hobbs 19426, Warren 1961). Dade County.
well, Little Bird Nursery and Garden Store (JRH). Gilchrist County.
Devil's Eye and Ear Spring (JRH). Hernando County. Eagle's Nest
Sink (JRH). Leon County. Sullivan's Tunnel (JRH). Levy County.
Friedman's Sink (JRH), Manatee Springs (JRH), well at Chiefland
(JRH). Madison County. Madison Blue Spring (JRH). Marion County.
Chert Cave (JRH), Hell Hole (JRH), Indian Cave (Bousfield 1963,
JRH), Orange Lake Cave (JRH), Roosevelt Cave (Warren 1961), Sunday

Caves and their Faunas in Florida and South Georgia 19

Sink (JRH). Pasco County. Nexus Sink (JRH), 38 m (125 ft) well at
Lacoochee (JRH). Suwannee County: Challenge Sink (JRH), Cisteen
Sink (JRH), Orange Grove Sink (JRH), Peacock Springs Cave System
(JRH), Sim's Sink (JRH). Wakulla County: McBride Slough (JRH),
River Sinks (JRH), Sally Ward Spring (JRH), Shepard Blue Spring
(JRH).

ETYMOLOGY: Named for Horton H. Hobbs, Jr., the collector
of the type series (Shoemaker 1941).

REMARKS: Holsinger (1972) placed this unique species in the
monotypic hobbsi group.

REFERENCES: Franz 1982 (conservation status); Holsinger 1972
(records, key), 1977 (taxonomy); Shoemaker 1941 (type description);
Warren 1961 (records).

Order DECAPODA

Family PALAEMONIDAE

Palaemonetes cummingi Chace

SQUIRREL CHIMNEY CAVE SHRIMP

Palaemonetes (Palaemonetes) cummingi Chace, 1954. Journal

of Washington Academy of Science 44(10):319-323. TYPE LOCALITY:

Squirrel Chimney, Alachua County, Florida. Holotype, female (USNM

95795), Robert B. Gumming (coll.), 11 July 1953.

Palaemonetes cummingi. — Warren, 1961:6.

OTHER COMMON NAMES: Florida Cave Shrimp.

DISTRIBUTION: OCALA FAUNA (Upper Suwannee). Known
only from the type locality. This cave shrimp was proposed as Threatened
by the Florida Committee on Rare and Endangered Plants and Animals
(Franz 1982) and was listed as Threatened on 21 June 1990 under
provisions of the U. S. Endangered Species Act of 1973, as amended
(Anonymous 1990). Because it is known from only one site in a
rapidly developing urban area, the shrimp is vulnerable to extinction
from groundwater contamination and deterioration of the surface habitat
around the Squirrel Chimney sinkhole. Divers failed to find shrimps
after an extensive search in this underwater cave in October 1992
(Morris and Butt 1992).

ETYMOLOGY: Named in honor of Robert B. Gumming, the
collector of the first specimen (Chace 1954).

REFERENCES: Chace 1954 (type description); Dobkin 1971 (larval
development); Franz 1982 (conservation status); Hobbs et al. 1977
(description); Strenth 1976 (taxonomy).

20 Richard Franz, Judy Bauer and Tom Morris

Family CAMBARIDAE .

Procambarus (Leconticambarus) milleri Hobbs
MIAMI CAVE CRAYFISH

Procambarus milleri Hobbs, 1971. Quarterly Journal of Florida
Academy of Science 34(2):115. TYPE LOCALITY: well at the Little
Bird Nursery and Garden Store, Miami, Dade County, Florida. Holotype
(USNM 131257), Billy R. Drummond, George C. Miller, and others
(colls.), 2 May 1968. Morphotype (USNM 131258), 16 paratypes.

Procambarus (Leconticambarus) milleri. — Hobbs, 1972:7 (by
implication).

DISTRIBUTION: MIAMI FAUNA. Known from two localities
in the Miami area. This species was recently collected at another site
in the Miami area by W. F. Loftus and P. Radice. The new specimens
are currently under study, and more information will become available
at a future date (W. F. Loftus, personal communication. Everglades
National Park). This crayfish is potentially vulnerable to extinction
from groundwater pollution and salt water intrusion, because its entire
range lies within a major urban area. The crayfish is listed as a
Species of Special Concern by the Florida Committee on Rare and
Endangered Plants and Animals (Franz 1982).

SPECIFIC LOCALITIES: Dade County: well at Little Bird Nursery
and Garden Store-type locality (USNM), 9-m deep well northeast of
Homestead (USNM).

ETYMOLOGY: Named in honor of George C. Miller who together
with Billy R. Drummond forwarded the type specimens to Horton H.
Hobbs, Jr. Mr. Miller was a long-time friend of Hobbs and fellow
student of crayfish (Hobbs 1971).

REFERENCES: Caine 1974 (evolution); Franz 1982 (conservation
status); Franz and Lee 1982 (distribution, evolution, records); Hobbs
1971 (original description); Hobbs et al. 1977 (description, records);
Hobbs and Hobbs 1991 (key).

Procambarus (Lonnbergius) acherontis (Lonnberg)

ORLANDO CAVE CRAYFISH .

Cambarus acherontis Lonnberg, 1895. Bihang till Koniglische
Svenska Veterskaps-Akademiens Handlingar 22:6. TYPE LOCALITY:
subterranean rivulet about 12.8 m (42 ft) from the surface in a hand-
dug well. Lake Brantley, Seminole County, Florida. Syntypes (ZIAS
1/4412) (two Form II males), one of which is still extant (Hobbs
1989).

Caves and their Faunas in Florida and South Georgia 21

Cambarus (Cambarus) acherontis. — Ortmann, 1905:102.

Cambarus (Ortmannicus) acherontis. — Fowler, 1912:341 (by implication).

Procambarus acherontis. — Hobbs, 1942^:342 (by implication).

Procambarus (Lonnbergius) acherontis. — Hobbs, 1972:8

OTHER COMMON NAMES: Orange-Seminole Cave Crayfish.
We prefer the use of the name Orlando Cave Crayfish because the
entire world distribution occurs within the Orlando metropolitan area.
It brings into focus the precarious conservation status of this species
in one of the most rapidly expanding urban areas in Florida.

DISTRIBUTION: ST. JOHNS RIVER FAUNA (Wekiva). This
crayfish is found in groundwater habitats in a limestone area along
the Wekiva River in Orange and Seminole counties, Florida. It was
proposed as Threatened by the Florida Committee on Rare and Endangered
Plants and Animals (Franz 1982).

SPECIFIC LOCALITIES: Orange County: Apopka Blue Hole
(USNM), Wekiwa Springs (USNM), well at Long Lake (RF). Seminole
County: Palm Springs (USNM), well at Altamonte Springs (RF), well
at Lake Brantley-type locality (ZIAS, Lonnberg 1894, 1895).

ETYMOLOGY: The name acherontis apparently refers to "Acheron,"
the name of the river of woe in Greek and Roman mythology, one of
the five rivers that surrounds Hades, across which Charon ferried the
dead.

REFERENCES: Cooper 1965a (records); Franz 1982 (conservation
status, records); Franz and Lee 1982 (distribution, evolution); Hobbs
19426 (description, records); Hobbs et al. 1977 (description, records);
Hobbs and Hobbs 1991 (key); Lonnberg 1894, 1895 (original description);
Walton and Hobbs 1959 (Ul as commensal).

Procambarus (Lonnbergius) morrisi Hobbs and Franz
PUTNAM COUNTY CAVE CRAYFISH

Procambarus (Lonnbergius) morrisi Hobbs and Franz, 1990.
Proceedings of Biological Society of Washington 104(l):56-62. TYPE
LOCALITY: Devil's Sink, 7.1 km west of Interlachen, Putnam County,
Florida. Holotype, allotype, morphotype (USNM 220374, 220375, 220376,
respectively), 12 paratypes (USNM), Tom Morris (coll.), 9 May 1989,
Tom Morris and Paul Smith (colls.), 6-8 March 1990.

DISTRIBUTION: ST. JOHNS RIVER FAUNA (Lake George).
Known only from the type locality.

ETYMOLOGY: This crayfish is named in honor of Tom Morris,
biologist and cave diver, who with Paul Smith collected the type

22 Richard Franz, Judy Bauer and Tom Morris

series and provided detailed information on Devils Sink (Hobbs and
Franz 1990).

REFERENCES: Franz and Franz 1990 (listed as a new cavernicolous
crayfish from Putnam County); Hobbs and Franz 1990 (original
description).

Procambarus (Ortmannicus) attiguus Hobbs and Franz
SILVER GLEN SPRINGS CAVE CRAYFISH

Procambarus {Ortmannicus) attiguus Hobbs and Franz 1992.
Proceedings of Biological Society of Washington 105(2):359-365. TYPE
LOCALITY: Silver Glen Springs, 14.4 km northwest of Astor Park,
Marion County, Florida. Holotype, allotype, and juvenile female paratype
(USNM 220683, 220684, and 220685, respectively), Tom Morris (coll.),
16 August 1990 (holotype only).

DISTRIBUTION: ST. JOHNS RIVER FAUNA (Lake George).
Known only from the type locality. One specimen (USNM) was taken
by Mike Spelman and Mike Hill on 5 June 1991 from Silver Glen
Well Cave that lies on the west side of the main spring pool and
represents a secondary outflow of the main spring cave.

ETYMOLOGY: Attiguus from Latin meaning neighboring; alluding
to the geographic proximity of the type, and only known locality, of
this species to Procambarus delicatus at Alexander Springs, as well
as to the close kinship of these two crayfishes (Hobbs and Franz
1992).

REFERENCES: Hobbs and Franz 1992 (original description).

Procambarus {Ortmannicus) delicatus Hobbs and Franz
BIG CHEEKED CAVE CRAYFISH

Procambarus {Ortmannicus) delicatus Hobbs and Franz, 1986.
Journal of Crustacean Biology 6(3):509. TYPE LOCALITY: Alexander
Springs, 9 km south of Astor Park, Lake County, Florida. Holotype,
allotype, paratype (USNM 218528, 144848, 145578, respectively);
holotype, Jeffrey B. Smith and Don Haren II (colls.).

OTHER COMMON NAMES: Alexander Springs Cave Crayfish.

DISTRIBUTION: ST. JOHNS RIVER FAUNA (Lake George).
Known only from the type locality.

ETYMOLOGY: The name delicatus (Latin= dainty) refers to the
delicate mien of this species.

REFERENCES: Franz and Lee 1982 (distribution); Hobbs and
Franz 1986 (original description), 1992 (comparison with attiguus);
Hobbs and Hobbs 1991 (key); Relyea et al. 1976 (record).

Caves and their Faunas in Florida and South Georgia 23

Procambarus {Ortmannicus) erythrops Relyea and Sutton
SANTA FE CAVE CRAYFISH

Procambarus {Ortmannicus) erythrops Relyea and Sutton, 1975.
Tulane Studies in Zoology and Botany 19(l-2):8. TYPE LOCALITY:
Sim's Sink, \.l km (1 mi) west of junction of U.S. Highways 27 and
129, 0.17 km (0.1 mi) south of U.S. Highway 27, Suwannee County,
Florida. Holotype, allotype, morphotype (USNM 133471, 133472, 133473,
respectively), 14 paratypes (USNM, RNHL, BMNH).

OTHER COMMON NAMES: Red-eyed Cave Crayfish, Sim Sink
Cave Crayfish.

DISTRIBUTION: OCALA FAUNA (Upper Suwannee). Restricted
to groundwater habitats in southern Suwannee County, Florida.

SPECIFIC LOCALITIES: Suwannee County. Azure Blue Sink
(USNM), Bufo Sink (Relyea and Sutton 1975), Hildreth Cave (USNM),
Quarry Sink (B. Sutton), Sim's Sink-type locality (USNM, Relyea
and Sutton 1975).

ETYMOLOGY: The species name erythrops from "Erythros" (Gr.)
for the color red, and "Ops" (Gr.) for eye; alluding to the red pigment
spot in the eye of this crayfish (Relyea and Sutton 1975).

REMARKS: Bufo Sink, 0.33 km (0.2 mi) south of (Sims Sink),
was listed as an unnamed sink in the original description (B. Sutton,
personal communication, Gainesville, Florida).

REFERENCES: Franz 1982 (conservation status); Franz and Lee
1982 (distribution, evolution, records); Hobbs et al. 1977 (description,
records); Hobbs and Hobbs 1991 (key); Mellon 1977 (eye structure);
Mellon and Lnenicka 1980 (eye structure); Relyea and Sutton 1975
(original description).

Procambarus {Ortmannicus) franzi Hobbs and Lee
ORANGE LAKE CAVE CRAYFISH
Procambarus {Ortmannicus) franzi Hobbs and Lee, 1976.
Proceedings of Biological Society of Washington 89(32):384. TYPE
LOCALITY: Orange Lake Cave, 0.64 km (0.4 mi) south of junction
of U.S. Highway 441 and State Route 318, Marion County, Florida.
Holotype, allotype, morphotype, (USNM 146992, 146993, 146994,
respectively), seven paratypes (USNM).

DISTRIBUTION: OCALA FAUNA (Orange Lake). Restricted to
groundwater habitats in the vicinity of Orange Lake in northern Marion
County, Florida.

SPECIFIC LOCALITIES: Marion County: Hell Hole (USNM),
Orange Lake Cave-type locality (USNM, Hobbs and Lee 1976), Orange

24 Richard Franz, Judy Bauer and Tom Morris

Lake Quarry solution pipes (DSL), Trade Winds Farm Cave (USNM).

ETYMOLOGY: Named for Richard Franz who with D. S. Lee
collected the type series at Orange Lake Cave (Hobbs and Lee 1976).

REFERENCES: Dickson and Franz 1980 (gill respiration); Franz
1982 (conservation status); Franz and Lee 1982 (evolution, distribution,
records); Hobbs and Lee 1976 (original description); Hobbs et al.
1977 (description, records); Hobbs and Hobbs 1991 (key).

Procambarus (Ortmannicus) horsti Hobbs and Means
BIG BLUE SPRINGS CAVE CRAYFISH

Procambarus horsti Hobbs and Means, 1972. Proceedings of
Biological Society of Washington 84(46): 401. TYPE LOCALITY: Big
Blue Spring (tributary to the Wacissa River), 3.7 km (2.2 mi) south of
the crossroads in the town of Wacissa, Jefferson County, Florida.
Holotype, allotype, morphotype (USNM 132043, 132044, 132045,
respectively), four paratypes (USNM), Michael N. Horst (coll.), October
1970.

Procambarus {Ortmannicus) horsti. — Holt, 1973Z?:246.

OTHER COMMON NAMES: Horst's Cave Crayfish.

DISTRIBUTION: WOODVILLE FAUNA. Confined to subterranean
habitats in limestone areas of the Woodville Karst Plain in Jefferson,
Leon, and Wakulla counties, Florida.

SPECIFIC LOCALITIES: Jefferson County: Big Blue Spring (Wacissa
River)-type locality (USNM). Leon County: well 7.5 km (4.5 mi) east
of Tallahassee (USNM). Wakulla County: Shepards Blue Spring (USNM).
Questionable record from Wakulla Spring (see Morris 1989).

ETYMOLOGY: Named for Michael N. Horst who donated the
type series to the U.S. National Museum of Natural History (Hobbs
and Means 1972).

REMARKS: This crayfish is a member of the pallidus complex
(Franz and Lee 1982).

REFERENCES: Franz 1982 (conservation status); Franz and Lee
1982 (evolution, distribution, records); Hobbs and Means 1972 (original
description); Hobbs et al. 1977 (description, records); Hobbs and Hobbs
1991 (key); Morris 1989 (records). ,^

Procambarus {Ortmannicus) leitheuseri Franz and Hobbs

COASTAL LOWLANDS CAVE CRAYFISH

Procambarus {Ortmannicus) leitheuseri Franz and Hobbs, 1983.

Proceedings of Biological Society of Washington 96(2): 323. TYPE

LOCALITY: Eagle's Nest Sink, 6.6 km (4.0 mi) northwest of junction

Caves and their Faunas in Florida and South Georgia 25

of U.S. Highway 19 and State Road 50, Hernando County, Florida.
Holotype, allotype, and morphotype (USNM 178361, 178582, 178585,
respectively), three paratypes (UF specimens transferred to USNM),
A. T. Leitheuser and L. F. Collins (colls.).

DISTRIBUTION: OCALA FAUNA (Gulf Coastal Lowlands). Restricted
to deep groundwater habitats in western Hernando and Pasco counties,
Florida.

SPECIFIC LOCALITIES: Hernando County: Die Polders 2 Sink
(USNM), Die Polders 3 Sink (USNM), Eagle's Nest Sink-type locality
(USNM), Little Salt Springs (ATL), Little Springs (ATL). Pasco County:
Arch Sink (USNM), Black Hole (USNM), Nexus Sink (USNM).

ETYMOLOGY: Named for its discoverer, Arthur T. Leitheuser,
who has added much to our knowledge of the distribution of troglobitic
crayfishes of Florida (Franz and Hobbs 1983).

REMARKS: This crayfish is a member of the lucifugus complex.

REFERENCES: Franz and Hobbs 1983 (original description); Hobbs
and Hobbs 1991 (key).

Procambarus (Ortmannicus) lucifugus lucifugus (Hobbs)
WITHLACOOCHEE LIGHT-FLEEING CAVE CRAYFISH

Cambarus acherontis. — Faxon, 1898:645.

Cambarus lucifugus lucifugus Hobbs, 1940fl. Proceedings of U.S.
National Museum 89(3097):398. TYPE LOCALITY: Gum Cave (=Sweet
Gum Cave), 11.2 km (7 mi) southwest of Floral City, Citrus County,
Florida. Holotype, allotype, "morphotype" (USNM 77916, 77917, 77918,
respectively), 32 paratypes (MCZ, USNM, FSBC, OSM), Kilby, Sherman,
Hobbs Jr. (colls., holotype), 1 May 1936.

Procambarus lucifugus lucifugus. — Hobbs, 1942^:343 (by implication).

Procambarus {Ortmannicus) lucifugus lucifugus. — Hobbs, 1972:9.

OTHER COMMON NAMES: Florida Cave Crayfish (in part).

DISTRIBUTION: OCALA FAUNA (Withlacoochee). Restricted
to groundwater habitats in Citrus and northern Hernando counties,
Florida.

SPECIFIC LOCALITIES: Citrus County: Sweet Gum Cave-type
locality (USNM, Hobbs 1940^). Hernando County: cave 23.3 km (14
mi) north of Weekiwachee Springs (USNM).

ETYMOLOGY: Lucifugus meaning light-fleeing, referring to its
habitation of unlighted cave environments.

REMARKS: This crayfish is a member of the lucifugus complex
(Franz and Lee 1982).

26 Richard Franz, Judy Bauer and Tom Morris

REFERENCES: Franz 1982 (conservation status); Franz and Lee
1982 (evolution, distribution, records); Hobbs 1940a (original description),
19A2b (description, records); Hobbs et. al. 1977 (description, records);
Hobbs and Hobbs 1991 (key); Hobbs III 1992 (photograph).

Procambarus (Ortmannicus) lucifugus alachua (Hobbs)
ALACHUA LIGHT-FLEEING CAVE CRAYFISH

Cambarus lucifugus alachua Hobbs, 1940fl. Proceedings of U.S.
National Museum 89(3097):402. TYPE LOCALITY: a small cave.
Hog Sink, about 16.7 km (10 mi) west of Gainesville, Alachua County,
Florida. Holotype, allotype, "morphotype" (USNM 76592), 50 paratypes
(USNM, FSBC), H. H. Hobbs Jr. (coll., holotype), 30 November 1937.

Procambarus lucifugus alachua. — Hobbs, 1942^:343 (by implication).

Procambarus {Ortmannicus) alachua. — Hobbs, 1972:10.

OTHER COMMON NAMES: Florida Cave Crayfish (in part).

DISTRIBUTION: OCALA FAUNA (Upper Suwannee). Restricted
to groundwater habitats in the Western Valley (Newberry Karst Plain)
of western Alachua and northeastern Levy counties, Florida.

SPECIFIC LOCALITIES: Alachua County: Bat Cave (USNM),
Crumbly Sink (USNM), Cueva Fria (USNM), Dudley Cave (USNM),
Goat Sink (USNM), Hog Sink-type locality (USNM), Martin's Cave
(Relyea and Sutton 1974), Protheroe Sink (Warren 1961), Seven Chimneys
Sink (RF), Squirrel Chimney (USNM), Tusk Cave (USNM). Levy County:
Gunpowder Cave (USNM), Williston Blue Sink (USNM).

ETYMOLOGY: The subspecific name refers to Alachua County,
Florida, the area where most of the specimens of this race have been
collected (RF).

REMARKS: This crayfish is a member of the lucifugus complex
(Franz and Lee 1982).

REFERENCES: Franz 1982 (conservation status); Franz and Lee
1982 (evolution, distribution, records, photograph); Hobbs 1940^ (original
description), \9A2b (description, records); Hobbs et al. 1977 (description,
records); Hobbs and Hobbs 1991 (key); Holt 19736 (CI as commensal);
Lee 1969c (as prey).

Procambarus I. lucifugus X /. alachua
Intergrade populations of LIGHT-FLEEING CAVE CRAYFISHES

DISTRIBUTION: OCALA FAUNA (Lower Suwannee and Marion).
The two intergrade populations appear isolated from one another and
from the P. I. lucifugus population on the western side of the Withlacoochee

Caves and their Faunas in Florida and South Georgia 27

River and P. L alachua population in Alachua and northeast Levy
counties. The taxonomic relationships between the various populations
are in need of further study.

SPECIFIC LOCALITIES: Lower Suwannee population in Gilchrist
County. Kelley Sinks (RF), Old Walker Farm Sink (USNM), Robert's
Cave (Warren 1961). Levy County. Friedman's Sink (USNM), Manatee
Springs (USNM). Marion population in Marion County. Briar Cave
(USNM), Chert Cave (USNM), Eickelberger Cave (USNM), Indian
Cave (USNM), Ocala Caverns (USNM), Redding Catacombs (RF),
Roosevelt Cave (USNM), Silver Springs (USNM), Steeple Cave (RF),
Sunday Sink (USNM), Waldo Cave (Warren 1961).

REFERENCES: Cooper 19656 (records); Franz 1982 (conservation
status); Franz and Lee 1982 (evolution, distribution, records); Hobbs
19426 (description, records); Hobbs et al. 1977 (description, records);
Holt 19736 (CI as commensal).

Procambarus (Ortmannicus) orcinus Hobbs and Means
WOODVILLE KARST CAVE CRAYFISH

Procambarus pallidus. — Hobbs 1958:81 (part).

Procambarus orcinus Hobbs and Means, 1972. Proceedings of
Biological Society of Washington 84(46):394. TYPE LOCALITY: Gopher
Sink, 5 km (3.1 mi) southwest of State Road 61 and 0.3 km (0.2 mi)
east of State Route 369, Leon County, Florida. Holotype, allotype,
morphotype (USNM 132031, 132032, 132033, respectively), 21 paratypes
(USNM).

Procambarus {Ortmannicus) orcinus. — Hobbs, 1972:58.

DISTRIBUTION: WOODVILLE FAUNA. Apparently restricted
to groundwater habitats in the limestone areas of the western Woodville
Karst Plain in Leon and Wakulla counties, Florida.

SPECIFIC LOCALITIES: Leon County: Bird Sink Swallet (USNM),
cave 5 km (3 mi) north of Woodville (USNM), Clay Sink (USNM,
Warren 1961), Culley's Cave (USNM), Falcon's Nest (USNM), Gopher
Sink-type locality (USNM, Hobbs and Means 1972), Little Dismal
Sink (USNM), Osgood Sink (USNM), Sullivan's Tunnel (USNM). Wakulla
County. Emerald Sink (USNM), Indian Springs (USNM), McBride
Spring (USNM), Sally Ward Spring (Morris 1989), River Sinks (Caine
1978), Wakulla Springs (USNM).

ETYMOLOGY: The species name orcinus (L.) for the nether
world, referring to the spelean habitat of this crayfish (Hobbs and
Means 1972).

REMARKS: Franz and Lee (1982) listed this crayfish as a member
of the pallidus complex.

28 Richard Franz, Judy Bauer and Tom Morris

REFERENCES: Caine 1978 (ecology, records); Franz 1982
(conservation status, records); Franz and Lee 1982 (evolution, distri-
bution, records); Hobbs and Means 1972 (original description); Hobbs
et al. 1977 (description, records); Hobbs and Hobbs 1991 (key); Holt
19736 (CI as commensal).

Procambarus {Ortmannicus) pallidus (Hobbs)
PALLID CAVE CRAYFISH

Cambarus acherontis. — Hobbs, 1937:154.

Cambarus acherontis pallidus Hobbs, 1938:90-91. Nomen nudum.

Cambarus pallidus Hobbs, 1940fl. Proceedings of U.S. National
Museum 89(3097):394. TYPE LOCALITY: Warren Cave, 17.6 km
(11 mi) northwest of Gainesville, Alachua County, Florida. Holotype,
allotype (both listed as USNM 76591), 13 paratypes (USNM, MCZ,
FSBC), H. H. Hobbs, Jr. et al. (colls., holotype).

Procambarus pallidus. — Hobbs, 1942^:343 (by implication).

Procambarus {Ortmannicus) pallidus. — Hobbs, 1972:10.

DISTRIBUTION: OCALA FAUNA (Upper Suwannee). Groundwater
habitats in limestone areas of the (northern) Withlacoochee River,
upper Suwannee River, lower Santa Fe River, and the Newberry Karst
Plain (western Alachua and northeast Levy counties), Florida. The
species may eventually be discovered in karst areas along the Withlacoochee
River in southern Georgia because the M2 Cave site in Madison County
(Florida) lies 9 km (6 mi) south of the Florida-Georgia border.

SPECIFIC LOCALITIES: A/flc/zMfl County. Alachua Sink (USNM),
Chimney Sink (USNM), Cueva Fria (USNM), Devil's Hole (USNM),
Goat Sink (USNM), Hertzog Cave (USNM), High Springs Cave (USNM),
Hog Sink (USNM), Hornsby Sink (USNM), Hornsby Spring (ATL),
McGeehee Blue Hole Cave (ATL), Pallidus Sink (USNM), Protheroe
Sink (Warren 1961), Squirrel Chimney (USNM), Still Sink (USNM),
Warren Cave-type locality (USNM, Hobbs 1940fl), well at Fort Clark
(USNM), 32-Foot Cave (RF). Columbia County. Big Grungy Swallet
(USNM), Big Room Cave (TM), Bussey's Sink (USNM), Columbia
Spring (TM), Fossil Cave Sink (TM), Riverbed Cave (USNM), Rose
Creek Swallet (USNM), Rose Creek Overflow Cave (TM), Russell's
Rub (TM), Shiloh Cave (USNM). Gilchrist County. Devil's Eye and
Ear Spring (USNM), Ginnie Springs (RF), Rocky Bluff Spring (USNM).
Hamilton County: Adams Spring (TM), Corbett Spring Cave (USNM),
Firecracker Cave (USNM), Overflow Cave (USNM), Pott Spring (TM),
Rossiter Spring (TM), Shallow Spring (ATL), Underhung Sink (USNM).
Lafayette County: Alligator Rescue Spring (TM), Aliens Mill Pond
Spring (USNM), Kassermans Sink (ATL), Lafayette Blue Spring (ATL),

Caves and their Faunas in Florida and South Georgia 29

Main Sink (ATL), Owens Spring (TM), Perry Spring (ATL), Ruth Spring
(USNM), Troy Spring (USNM). Levy County. Archer Caves (RF), Devils
Den (USNM). Madison County. Baseline Cave (USNM), Madison Blue
Spring (ATL), M2 Blue Cave (USNM), Suwannacoochee Spring (USNM),
Thunderhole Sink (USNM). Suwannee County. Anderson Spring (TM),
Bonnett Spring (TM), Challenge Sink (TM), Charles Spring (TM), Cisteen
Sink (USNM), Cow Spring Cave (USNM), Crazy Horse Sink (USNM),
Double Sink (ATL), Edwards Spring (USNM), Falmouth Spring (JB),
Ghoul Sink (USNM), Irvine Slough Spring (TM), Lineater Spring (TM),
Little River Spring (USNM), Mirkwood Sink (USNM), Olsen Sink (TM),
Orange Grove Sink (USNM), Peacock Spring (USNM), unnamed sink
in Peacock System (USNM), Peacock Sink 3 (JB), Pot Hole (JB), Register
Sink (ATL), Sandbag Spring (TM), Smith Sink (ATL), Stick Sink (ATL),
Telford Springs (USNM), Ten Mile Hollow Cave (TM), Water Hole 3
Sink (JB).

ETYMOLOGY: The name pallidus refers to the pallid appearance
of this cave species (RF).

REMARKS: The record from Eichelberger Cave in Marion County
(Hobbs et al. 1977) was questioned by Franz and Lee (1982). Crayfishes
from the Big Grungy population are less tuberculate and spiniform
than more typical P. pallidus (HHH). We encourage the collections
of more specimens from this site in order to resolve its taxonomic
status. P. pallidus is a member of the pallidus complex (Franz and
Lee 1982).

REFERENCES: Cooper \965b (records); Dickson and Franz 1980
(gill respiration); Franz 1982 (conservation status, records); Franz
and Lee 1982 (evolution, distribution, records); Hobbs 1940a (original
description), 1942^, 1942^ (descriptions, records); Hobbs et al. 1977
(descriptions, records); Hobbs and Hobbs 1991 (key); Relyea and Sutton
1913a (egg-bearing); Streever 1992Z? (crayfish kill at Peacock Springs);
Walton and Hobbs 1959 (CI as commensal).

Troglocambarus maclanei Hobbs
NORTHERN SPIDER CAVE CRAYFISH

Troglocambarus maclanei Hobbs, 19A2a. American Midland
Naturalist 28(2):345. TYPE LOCALITY: Squirrel Chimney, 17.6 km
(11 mi) northwest of Gainesville, Alachua County, Florida. Holotype,
allotype, paratypic male (form II) (USNM 79385, 79386, 79387,
respectively), other paratypes (MCZ, USNM). H. H. Hobbs (coll., holotype),
25 March 1941.

OTHER COMMON NAMES: Spider Cave Crayfish (in part),
McLane's Cave Crayfish.

30 Richard Franz, Judy Bauer and Tom Morris

DISTRIBUTION: OCALA FAUNA (Upper Suwannee, Lower
Suwannee, Orange Lake, Marion, Withlacoochee, and Gulf Coastal Lowlands).
Troglocambarus maclanei is the most widely distributed troglobitic
crayfish in Florida. It ranges from southern Suwannee County, southwest-
ward to Pasco County. Based on the sighting of a shrimp-like crayfish
in Knight Sink, near Tarpon Springs, this species may range south into
Pinellas County (P. Heinerth, personal communication, Hudson, Florida).

SPECIFIC LOCALITIES: Alachua County. Goat Sink (USNM),
Hertzog Cave (USNM), Squirrel Chimney-type locality (USNM, Hobbs
1942fl). Columbia County: Columbia Spring (TM), Fossil Cave Sink
(TM). Gilchrist County. Devil's Eye and Ear Spring (USNM). Hernando
County: Eagle's Nest Sink (USNM). Levy County: Manatee Springs
(USNM), Peanut Cave (USNM). Marion County: Chert Cave (USNM),
Indian Cave (USNM), Orange Lake Cave (USNM), Sunday Sink (RF),
Trade Winds Farm Sink (USNM). Suwannee County: Azure Blue Sink
(USNM), Sim's Sink (USNM).

ETYMOLOGY: Named for William A. McLane, collector of the
original specimens.

REFERENCES: Cooper 19656 (records); Franz 1982 (conservation
status, records, photograph); Franz and Lee 1982 (evolution, distribution,
records); Franz and Franz 1990 (distribution); Hobbs 1942a (original
description, photograph), 19426 (description, records, photograph); Hobbs
et al. 1977 (description, records); Hobbs and Hobbs 1991 (key); Holt
19736 (CI as commensal); Mohr and Poulson 1966 (photo).

Troglocambarus sp.
ORLANDO SPIDER CAVE CRAYFISH

DISTRIBUTION: ST. JOHNS RIVER FAUNA (Wekiva). Known
only from Apopka Blue Sink, Orange County, Florida (USNM). This
cave may represent an upstream part of the Rock Spring Cave System.
REMARKS: Specific identification of this crayfish awaits the collection
of Form I males (HHH).

REFERENCES: Hobbs III 1992 (photograph).

Cambarus (Jugicambarus) cryptodytes Hobbs
APALACHICOLA CAVE CRAYFISH

Cambarus {Cambarus) cryptodytes Hobbs. 1941. American
Midland Naturalist 26(1): 110. TYPE LOCALITY: well on the R. W.
Williams farm, 3.2 km (2 mi) south of Graceville, Jackson County,

Caves and their Faunas in Florida and South Georgia 31

Florida. Holotype, allotype, morphotype (USNM 79339, 79340, 79343,
respectively), five paratypes (MCZ, USNM).

Cambarus cryptodytes . — Hobbs, 1942^:354.

Cambarus {Jugicambarus) cryptodytes. — Hobbs, 1969:107.

OTHER COMMON NAMES: Marianna Lowlands Cave Crayfish,
Dougherty Plain Cave Crayfish. We propose the new common name,
Apalachicola Cave Crayfish, which we believe better reflects this
species and its distribution.

DISTRIBUTION: APALACHICOLA FAUNA (Marianna and
Southwest Georgia). Groundwater areas of northcentral Jackson Coun-
ty, Florida, and Decatur County, Georgia.

SPECIFIC LOCALITIES: Jackson County. Cave-in-Woods (RF),
Ellis Cave (USNM), Gerard Cave (USNM), Gerome's Cave (Hobbs et
al. 1977), Hole-in Wall (USNM), Jackson Blue Spring (USNM), Judges
Cave (Warren 1961), Miller's Cave (DSL), Milton's Well Cave (DSL),
Pool Cave (Hobbs et al. 1977), Pottery Cave (Warren 1961), Ray's
Cave (FNAI), Rockwell Cave (Hobbs et al. 1977), Soda Straw Cave
(Warren 1961), Twin Cave (USNM), Vetter's Cave (Hobbs et al. 1977),
Waddell's Mill Cave (USNM), Washed-out Cave (Warren 1961), well
3.3 km (2 mi) south of Graceville-type locality (USNM, Hobbs 1941).
Decatur County (Georgia): Climax Cave (USNM).

REMARKS: The type locality was an 18.2-m (60 ft) deep well
on the farm of Robert W. Williams. During a 1983 visit with Mr.
Robert Williams, who obtained the original series for Hobbs, we learned
that the well had been filled many years before. The Washington
County specimen mentioned by Hobbs (1989), we believe, must have
come from Jackson County (probably Gerards Cave), based on road
mileage listed with the specimen. Unusual specimens of Cambarus
were recently collected at Blue Hole in Florida Caverns State Park by
park rangers and at Vortex Spring in Holmes County that may represent
one or more distinct taxon (HHH). Adult material is necessary before
this taxonomic problem can be resolved.

ETYMOLOGY: Crypto from the Greek=hidden, dytes=to dwell,
referring to the crayfish dwelling in the cave environment.

REFERENCES: Franz 1982 (conservation status); Franz and Lee
1982 (evolution, distribution, records); Harris 1968 (associate of Haideo-
triton); Hobbs 1941 (original description), 1942^, 19426 (description,
records), 1969 (taxonomy), 1981 (description, records in Georgia);
Hobbs et al. 1977 (description, records); Hobbs and Hobbs 1991 (key);
Hobbs and Walton 1968 (Uw as commensal); Pylka and Warren 1958
(records); Warren 1961 (records).

32 Richard Franz, Judy Bauer and Tom Morris

Class INSECTA

Order COLLEMBOLA

Family Entomobryidae

Pseudosinella pecki Christiansen and Bellinger

MARIANNA CAVE SPRINGTAIL

Pseudosinella pecki Christiansen and Bellinger, 1980. Grinnell
College Special Paper: 988-989pp. TYPE LOCALITY: Miller's Cave,
(Florida) Caverns State Park, Jackson County, Florida. S. B. Peck
(coll.), 28 December 1965.

DISTRIBUTION: APALACHICOLA FAUNA (Marianna Lowlands
and Southwest Georgia). Known from caves in Jackson County (Florida),
Jackson County (Alabama), and Decatur, Randolph, and Stewart coun-
ties, Georgia (Christiansen and Bellinger 1980).

REMARKS: According to Christiansen and Bellinger (1980), this
species is probably a troglobitic derivative of Pseudosinella argentea.

ETYMOLOGY: Named for Stewart B. Peck, collector of the
collembolan.

REFERENCES: Peck 1970 (record); Christiansen and Bellinger
1980 (original description).

Class ARCHNIDA

Order ARANEA

Family Linyphiidae

Islandiana sp.

MARIANNA CAVE SHEETWEB WEAVER SPIDER

DISTRIBUTION: APALACHICOLA FAUNA (Marianna Lowlands).
This undescribed spider is known only from Miller's Cave, Florida
Caverns State Park, Jackson County, Florida. REMARKS: This spider
is currently under study by W. T. Gertsch (S. B. Peck, personal com-
munication, Carlton University).

REFERENCES: Peck 1970 (record).

Phylum CHORDATA
Class AMPHIBIA
Order CAUDATA
Family Plethodontidae '

Haideotriton wallacei CsLTT •:

GEORGIA BLIND SALAMANDER
Haideotriton wallacei Carr, 1939. Occasional Papers of Boston
Society of Natural History 8:335-336. TYPE LOCALITY: 60.8-m (200

Caves and their Faunas in Florida and South Georgia 33

ft) deep well in Albany, Dougherty County, Georgia. Holotype (MCZ
19875), Mr. Hummel (Dougherty County Sanitary Engineer) (coll.), 19
May 1939.

DISTRIBUTION: APALACHICOLA FAUNA (Marianna
Lowlands and Southwest Georgia). Restricted to groundwater
habitats in Jackson County, Florida, and Decatur and Dougherty
counties, Georgia. This salamander was listed as Rare by the Florida
Committee on Rare and Endangered Plants and Animals (Means 1978,
1992).

SPECIFIC LOCALITIES: FLORIDA, Jackson County. Cave-in-
Woods (RF), Ellis Cave (DSL), Gerard's Cave (UF, USNM, NCSM,
MCZ, Pylka and Warren 1958), Hole-in-Wall Cave (UF), Jackson
Blue Spring (RF), Judges Cave (DSL), Miller's Cave (DSL), Milton's
Well Cave (DSL), Twin Cave (RF), Washed-out Cave (Warren 1961).
GEORGIA, Decatur County: Climax Cave (UF). Dougherty County.
well in Albany (MCZ).

ETYMOLOGY: Named for H. K. Wallace, spider expert and
Carr's colleague at the University of Florida, Gainesville.

REFERENCES: Carr 1939 (original description, photo); Bishop
1947 (description, photo); Brandon 1967 (description, literature);
Dundee 1962 (response to metamorphic agent); Franz and Lee 1982
(crayfish associate); Harris 1968 (ecology); Hilton 1945 (skeleton);
Hobbs III 1992 (photo); Lee 1969a (food habits), 1969^?; Means 1977
(distribution), 1978, 1992 (conservation); Mohr and Poulson 1966
(photo); Peck 1973 (feeding efficiency); Valentine 1964 (morphology);
Vandel 1965^ (records); Wake 1966 (taxonomy); Pylka and Warren
1961 (record).

SUMMARY — The troglobitic fauna is composed of three isopods,
two amphipods, one shrimp, 18 crayfishes (including two subspecies
and two intergrade populations), one snail, one spider, one spring-
tail, and one salamander. At least one species (Caecidotea hobbsi)
occurs in groundwater habitats outside of caves, per se; the amphipods
(Crangonyx hobbsi and Crangonyx grandimanus) also may use
similar habitats, particularly in light of their wide geographic dis-
tributions. Several taxa in this list remain undescribed, due to the
lack of critical material in collections. Faunas and assemblages are
described in the section entitled Obligate Cave Faunas and Karst
Regions.

34 Richard Franz, Judy Bauer and Tom Morris

ANNOTATED LIST
OF OTHER CAVE-ASSOCIATED SPECIES

Phylum ANNILIDA

Class OLIGOCHAETA

Family Branchiobdellidae

Cambarincola leoni Holt (commensal on troglobites). Alachua
County: on Troglocambarus maclanei from Squirrel Chimney; on
Procambarus lucifugus alachua from Goat Sink. Leon County:
on Procambarus orcinus from Gopher Sink. Marion County: on
Procambarus lucifugus X alachua from Indian Cave. REMARKS:
Holt (19736) lists questionable records of this species on crayfishes
from Squirrel Chimney (Alachua County), The Bat Hole (=Roberts
Cave) (Gilchrist County), and Sim's Sink (Suwannee County).

Cambarincola manni Holt (accidental?). Seminole County:
on Procambarus acherontis from Palm Spring (Holt 19736). REMARKS:
This annelid also has been retrieved from epigean crayfishes {Pro-
cambarus alleni, P. fallax, P. paeninsulanus) (Holt 1973a). Its
presence on Procambarus acherontis may have been fortuitous
because the specimens were acquired from crayfishes found in
the spring pool at Palm Springs rather than from the adjacent
cave environment. Infestations may have originated with surface
crayfishes coinhabiting the spring pool.

Other Branchiobdellid Annelids. Unidentifiable branchiobdellid
materials were collected on Procambarus erythrops (listed as Pro-
cambarus lucifugus) from Sim's Sink (Suwannee County), on Procam-
barus horsti from Big Blue Spring (Jefferson County), and on
Cambarus cryptodytes from Gerard's Cave (Jackson County) and
Climax Cave (Decatur County, GA) (Holt 19736).

Other Annelids. Aeolosomatid oligochaetes were reported as
part of the benthos from the cave tunnel and from the shallower
regions of Vortex Blue Spring in Holmes County (Helfman 1986).
Other caves with populations of aquatic annelids include: Columbia
County: Rose Creek Swallet and Overflow (TM), Blue Sink (TM).
Hamilton County: Shallow Spring (TM), Rossiter Spring (TM).
Madison County: Thunderhole (TM). Suwannee County: Alligator
Rescue Spring (TM), Bonnett Spring (TM), Charles Spring (TM),
Irvine Slough Spring (TM), Peacock Springs (TM, Streever 19926),
Lineater Spring (TM), Telford Spring (TM), Water Hole Cave
(TM).

Caves and their Faunas in Florida and South Georgia 35

Phylum MOLLUSCA
Class BIVALVIA
Family Cyrenidae
Corbicula fluminea (O.F.M.) (troglophile?). Columbia County.
Siphon Creek Cave (TM); Lafayette County: Green Sink (TM); Suwan-
nee County. Peacock Springs Cave (Streever \992a).

Family Unionidae
Uniomerus obesus (Lee) (trogloxene?). Columbia County. Rose
Creek Swallet (James D. Williams, personal communication, National
Biological Survey, Gainesville, Florida). REMARKS: Mussels were
collected alive buried in hard sand on the floor of this spring cave
between the entrance and 60 meters penetration at water depths up to
12 meters. Their distribution in the cave probably is limited by the
availability of filterable foods and the abilities of the host fish that
carry the parasitic glochidia to penetrate subterranean habitats. The
food supply may include the thin veneer of organic silt that covers
the sand at this site.

Class GASTROPODA

Subclass PROSOBRANCHIA

Family Pleuroceridae

Elimia clenchi (Goodrich), Slackwater Elimia (trogloxene). Holmes

County. Vortex Blue Spring (Helfman 1986). REMARKS: Reported

as benthos from the cavern portion of Vortex Spring, but not from the

deeper portions of the cave.

Elimia curvicostata (Reeve), Graphite Elimia (trogloxene).
Washington County: Econfina Blue Springs Cave (RF, Thompson and
Hershler 1991). REMARKS. This species was collected with the troglobitic
snail, Dasyscias franzi, in the cave. Elimia curvicostata also occurred
in the outflow and spring pool of Econfina Blue Spring; other snail
species, including Elimia athearni, were found in the spring pool but
not in the cave stream.

Family Hydrobiidae
Amnicola retromargo Thompson (troglophile). Hamilton County:
Shallow Spring (FGT). REMARKS: These snails were collected approxi-
mately 60-600 meters inside the cave (water depth 12 meters) on
boulders on the floor and on the cave walls in strong water currents.
These specimens were lighter in color than individuals found in surface
streams (F. G. Thompson personal communication, Florida Museum of
Natural History).

36 Richard Franz, Judy Bauer and Tom Morris

Family Thiaridae

Melanoides tuberculatus (Mueller) (trogloxene?) Citrus County.
Restinghouse Siphon (FGT). REMARKS: This snail is introduced from
southeast Asia. It was found on the floor of this flooded cave between
0-60 meters penetration and at water depths up to 12 meters.

Tarebia granifera (Lamarck) (trogloxene?). Citrus County.
Restinghouse Siphon (FGT). REMARKS: This snail is introduced from
southeast Asia. This snail was found in association with Melanoides
tuberculatus on the cave floor between 0-60 meters penetration and at
water depths up to 12 meters.

Subclass PULMONATA
Family Endodontidae

Anguispira alternata (Say) (trogloxene). Jackson County, caves
(Franz et al. 1971).

Family Haplotrematidae

Haplotrema concavum (Say) (trogloxene). Jackson County, caves
(Franz et al. 1971).

Family Oleacinidae

Euglandina rosea (Ferussac) {iio^oxtnt). Jackson County: Gerome's
Cave, Kramers cave, Milton's Well Cave, Pottery Cave, River Cave,
Vetter's Cave (Franz et al. 1971). Washington County. Falling Waters
Trail Cave (Franz et al. 1971). REMARKS: Nests with eggs of this
snail were reported from several caves in Jackson County (Franz et al.
1971).

Family Polygyridae

Mesodon inflectus (Say) (trogloxene). Jackson County, caves (Franz
et al. 1971).

Mesodon thyroidus (Say) (trogloxene). Jackson County, caves (Franz
et al. 1971).

Triodopsis hopetonensis (Shuttle worth) (trogloxene). Alachua County.
Bat Cave (FGT).

Family Zonitidae
Vintridens demissus (Binney) (trogloxene). Jackson County: caves
(Franz et al. 1971). ^.

Caves and their Faunas in Florida and South Georgia 37

Family Planorbidae

Gyraulus parvus (Say), Ash Gyro (trogloxene?). Alachua County:
Bat Cave (FGT). REMARKS: The presence of this snail and Promenetus
in Bat Cave is puzzling. The cave is located in the Newberry Karst
Plain and is not close to any standing or flowing bodies of surface
water (RF).

Promenetus sp. (trogloxene?). Alachua County. Bat Cave (FGT).

Unidentified snails. Madison County. Madison Blue Spring (TM).

Phylum ARTHROPODA

Class CRUSTACEA
Subclass OSTRACODA
Family Entocytheridae

Uncinocythere ambophora (Walton and Hobbs) (commensal on
troglobites). Putnam County, on Procambarus morrisi at Devil's Sink
(Hobbs and Franz 1990). Seminole County, on Procambarus acherontis
at Palm Springs (Walton and Hobbs 1959).

Uncinocythere equicurva (Hoff) (trogloxene?). Reported by Hoff
(1944) on Procambarus lucifugus alachua (specific data not mentioned);
also known on surface crayfishes (Hoff 1944); identity questioned by
Walton and Hobbs (1959).

Uncinocythere lucifuga (Walton and Hobbs) (commensal on
troglobites). Alachua County: on Procambarus lucifugus alachua
from Hog Sink; on Procambarus pallidus from Squirrel Chimney (Walton
and Hobbs 1959). Marion County: on Procambarus franzi from Orange
Lake Cave (A. Norden, personal communication, Maryland Natural
History Society, Baltimore).

Uncinocythere warreni Hobbs and Walton (commensal on
troglobites). Decatur County, Georgia: on Cambarus cryptodytes from
Climax Cave (Hobbs and Walton 1968). REMARKS: This ostracod
also should occur on Cambarus cryptodytes in caves of Jackson County,
Florida.

Other Ostracods. REMARKS: Lee (1969^:175) reported "Six
species of ostracods of the families Canoninae and Cypridacea...
including one specimen identified to the genus Darwinula" from the
gut of Haideotriton wallacei from Gerards Cave in Jackson County.

Subclass EUCOPEPODA
Unidentified copepods. Lee (1969^) reported finding copepods in
the gut of Haideotriton wallacei from Gerards Cave. Hobbs (1942Z?)

38 Richard Franz, Judy Bauer and Tom Morris

also mentioned copepods from Gum Cave. Other copepod records include
Columbia County: Jug Spring (TM); Gilchrist County: Devil's Eye and
Ear Spring (TM). Leon County: Little Dismal Sink (TM). Levy County:
Gunpowder Sink (TM). Marion County: Silver Springs (TM); Suwannee
County: Charles Spring (TM), Irvine Slough Spring (TM). REMARKS:
Copepods frequently were found in association with catfish feces (TM).

Subclass MALACOSTRACA
Order ISOPODA
Family Trichoniscidae
Miktoniscus alabamensis Muchmore (troglophile). Jackson County:
Florida Caverns, Miller's Cave (Vandel 1965^, 1965^; Peck 1970).

Order AMPHIPODA
Family Crangonyctidae
Crangonyx floridanus Bousfield (troglophile). Dade County: well
at Little Bird Nusery and Garden Store in Miami (JRH). Jackson
County: China Cave (JRH), Gerard's Cave (Lee 1969«), Geromes Cave
(JRH), Judges Cave (JRH), Millers Cave (JRH), Pool Cave (JRH).
Suwannee County: Azure Blue Sink (JRH). REMARKS: The type
series of this amphipod was collected in a swamp at Highlands Ham-
mock State Park, Florida (Bousfield 1963). It also has been taken from
a swamp in St. Tammany Parish, Louisiana (Holsinger 1972). Specimens
from the well in Miami do not differ from those in northern Florida
and do not represent an undescribed subspecies as suggested earlier by
Holsinger (1972; personal communication, Old Dominion University).

Family Talitridae
Hyallela azteca (Saussure) (trogloxene). Holmes County: Vortex
Blue Spring (Helfman 1986). REMARKS: Reported as part of the
benthos from the cave tunnel and from the shallower cavern region at
Vortex Spring.

Order DECAPOD A v

Family Cambaridae
Procambarus alleni (Faxon) (trogloxene). Monroe County: shallow
solution channels on Big Pine Key (DSL).

Procambarus paeninsulanus (Faxon) (irogloxenQ). Jackson County:
Gerome's Cave (DSL). REMARKS: This crayfish was abundant in the

Caves and their Faunas in Florida and South Georgia 39

main cave pool at Geromes Cave; ovigerous females were found in
burrows constructed in bat guano along its bank (DSL).

Procambarus fallax (Hagen) (trogloxene). Alachua County: Alachua
Sink (Hobbs et al. 1977).

Family Palaemonidae

Marcobrachium carinus (L.) (accidental?). Marion County: Silver
Glen Spring (Hobbs and Franz 1992), Silver Springs (Tom Allen, personal
communication, formerly Gainesville, Florida). REMARKS: This shrimp
has been encountered in the entrances of spring caves in the lower St.
Johns River system. Prior to the building of Rodman Dam, the species
once was common in Silver Springs; it apparently has been extirpated
from this site (Tom Allen, personal communication). Specimens have
been retrieved from the stream below the dam as recently as 1990.

Palaemonetes paludosa (Gibbes) (accidental?). Columbia County.
Siphon Creek Cave (TM). Gilchrist County: Devils Eye and Ear Spring
(TM). Hernando County: Die Polder 3 (PH), Eagles Nest (TM). Leon
County: Gopher Sink (Hobbs and Means 1972). Wakulla County: Sally
Ward Spring (TM), Wakulla Springs (TM). REMARKS: Specimens
have been seen at water depths near 60 m in Die Polders 3 (TM). An
unusual specimen of Palaemonetes was collected in Silver Springs
that warrants additional study (HHH). This specimen may represent
an undescribed taxon.

Palaemonetes kadiakensis Rathbun (accidental?). Leon County:
Bird Sink (USNM). REMARKS: Two females were collected in the
dark zone, 60 m inside the cave in September 1993 (B. Pruitt, pers.
comm.).

Family Portunidae
Callinectes sapidus Rathbun (accidental?). Marion County: Salt
Springs (RF), Silver Glen Springs (Hobbs and Franz 1992). REMARKS:
Female blue crabs commonly invade the mouths of springs around
Lake George on the St. Johns River during their annual migrations.

Other Crustaceans. An unstudied crab (trogloxene?) was collected
in the cave at Silver Glen Springs, Marion County (HHH).

Class INSECTA
Order COLLEMBOLA
Family Isotomidae
Isotoma notabilus Schaffer (troglophile). Alachua County: Warren
Cave (Peck 1970).

40 Richard Franz, Judy Bauer and Tom Morris

Family Tomoceridae
Tomocerus dubius Christiansen (troglophile). Alachua County: Warren
Cave (Peck 1970). Jackson County. Miller's Cave (Peck 1970).

Order ORTHOPTERA
Family Gryllacrididae

Ceuthophilus gracilipes (Haldeman) (trogloxene). Jackson County:
Bat Cave, Blue Spring Cave, Florida Caverns, Gerard's Cave, Miller's
Cave, small cave near Judges Cave (Hubbell 1936, Peck 1970).

Ceuthophilus latibuli Scudder (trogloxene). Alachua County: Bat
Cave, Dudley Cave, Grant's Cave, Jook Cave, O'Steen's Cave, Warren
Cave (Hubbell 1936, Peck 1970). Citrus County: Blowing Hole Cave,
Dr. Doans (Dames) Cave (Hubbell 1936, Peck 1970). Marion County:
Belleview Cave, Jennings Cave, Waldo Cave (Hubbell 1936, Peck
1970).

Ceuthophilus virgatipes Rehn & Hebard (trogloxene). Alachua
County: Dudley Cave (Hubbell 1936, Peck 1970). Marion County:
Villa Height's Cave (Hubbell 1936, Peck 1970).

Order COLEOPTERA
Family Leiodidae

Nemadus sp. (troglophile). Alachua County: Warren Cave (Peck
1970). Jackson County: Gerard's Cave (Peck 1970).

Prionchaeta opaca (Say) (troglophile). Jackson County: Gerard's
Cave, Miller's Cave (Peck 1970).

Ptomaphagus cavernicola Schwarz (troglophile). A/ac/zM^ County:
Warren Cave (Peck 1970). Jackson County: Gerard's Cave (Lee 1969<2).
REMARKS: Lee (1969«) found this beetle in the stomaches oi Haideo-
triton wallacei from Gerard's Cave. Although it is found regularly in
caves from Mexico to Florida, this beetle recently was recorded away
from caves in mixed pine and deciduous forests of floodplain loam or
sandy soils in Jackson, Leon, and Suwannee counties, Florida, and in
Macon County, Georgia (Peck 1982). Peck (1982) concluded that P.
cavernicola is a contemporary inhabitant of forests of the southeastern
United States, is not cave-limited or a Recent climatic relict, and
probably can colonize suitable cave sites.

Family Dytiscidae
Hydroporus clypealis Sharp (accidental?). A/ac/ima County: Devil's
Hole (Hobbs 19426, Young 1942).

Caves and their Faunas in Florida and South Georgia 41

Family Staphylinidae

Atheta (Atheta) annexa Casey (trogloxene). Jackson County: Miller's
Cave (Klimaszewski and Peck 1986).

Atheta (Dimetrota) lucifuga Klimaszewski and Peck (trogloxene).
Jackson County: Miller's Cave (Klimaszewski and Peck 1986).

Atheta (Dimetrota) troglophila Klimaszewski and Peck (troglophile).
Jackson County: Miller's Cave (Klimaszewski and Peck 1986).

Family Catopidae
\ Unidentified catopid beetle. Lee {\969a) reported these beetles

in the gut of Haideotriton wallacei from Gerards Cave.

Family Carabidae
Rhadine larvalis LeConte (troglophile). Jackson County: "from
at least one Marianna area cave" (T. C. Barr, Jr., personal communica-
tion, University of Kentucky). REMARKS: This beetle also was found
in Turk's Cave, near Evergreen, Conecuh County, Alabama, and could
occur in any Florida cave (T. C. Barr, Jr., personal communication).

Order DIPTERA
Family Nycteribiidae
Basilia boardmanii Rozeboom (trogloxene). Marion County: on
Myotis austroriparious from Orange Lake Cave. (James Bain, personal
communication, Flagstaff, Arizona).

Family Streblidae
Trichobius major Coquillet (trogloxene). Citrus County: Sweet
Gum Cave (Hubbard 1901). Marion County: on Myotis austroriparious
from Orange Lake Cave (James Bain, personal communication).

Class ARACHNIDA

Order UROPYGI

Family Thelyphonidae

Mastigoproctus giganteus (accidental). Citrus County: Sweet Gum

Cave (Hobbs 19426). REMARKS: Listed as Thelyphonus giganteus

by Hobbs (1942b).

Order OPILIONES
Family Phalangodidae
Phalangodes (Bishopella) laciniosa Crosby & Bishop (troglophile).
Jackson County: Florida Caverns (listed as P. marianna by Goodnight
and Goodnight 1942, 1953).

42 Richard Franz, Judy Bauer and Tom Morris

Phalangodes (Crosbyella) spinturnix Crosby and Bishop (troglophile).
Jackson County: Gerard's Cave, Milton Cave, Miller's Cave (Peck
1970). . J,

Order ARANEAE

Family Argiopidae

Azilla affinis (Simon) (troglophile). Jackson County: "Spring Cave"

and "Two Entrance Cave" in Florida Caverns State Park (Peck 1970).

REMARKS: Listed by Peck (1970) as Azilla vagepicta Simon. Levi

(1980) suggested the use of A. affinis instead of A. vagepicta.

Family Linyphiidae
Centromerus latidens (Emerton) (troglophile). Jackson County:
(Old) Indian Cave (Peck 1970). REMARKS: An undetermined female
Centromerus also was collected in Dr. Dames Cave, Citrus County
(Peck 1970).

Family Nesticidae

Gaucelmus augustinus Keyserling (troglophile). Alachua County:
Bat Cave, Dudley Cave, Squirrel Chimney, Warren Cave (Peck 1970,
Gertsch 1984). Citrus County: "caves," small cave 8 km (5 mi) north
of Dr. Dames Cave, Dr. Dames Cave (Peck 1970, Gertsch 1984)).
Jackson County: Florida Caverns, Milton's Cave, (Waddells) Mill Pond
Cave, Miller's Cave (Peck 1970, Gertsch 1984). Marion County: Mefford
Cave (Gertsch 1984, Peck 1970)

Eidmannella pallida (Emerton) (troglophile). Alachua County: Bat
Cave, Grant's Cave, Dudley Cave (Peck 1970, Gertsch 1984). Citrus
County: Blowing Hole Cave, Dr. Dames Cave (Peck 1970, Gertsch
1984). Jackson County: cave near Gerards Cave, Florida Caverns,
Gerard's Cave, Judges Cave, Miller's Cave, (Miltons) Well Cave No.l,
(Waddells) Mill Pond Cave, Old Indian Cave (Peck 1970, Gertsch 1984)
REMARKS: New name combination was suggested by Gertsch (1984).
Formerly called Nesticus pallidus.

Family Cteniidae
Ctenus captiosus (troglophile?). Probably in Florida caves (Edwards
1989).

Order ACARI
Unidentified mites. Lee (1969«) reported a single mite in the gut
of a Haideotriton wallacei from Gerard's Cave.

Caves and their Faunas in Florida and South Georgia 43

Class DIPLOPODA
Family Cambalidae

Cambala annulata (Say) (troglophile). Jackson County: Gerard's
Cave, Indian Cave, Judges Cave, Miller's Cave, Milton's Cave (Peck
1970).

Class CHILOPODA
' Family Lithiobiidae

Lithobius atkinsoni Bollman (troglophile). Jackson County: Gerard's
Cave, Milton's Cave (Peck 1970).

Other Invertebrates. P. Heinerth (personal communication)
found sponges on divers' guide lines that extended into Black Hole,
Pasco County. This cave lies in a tidal marsh near the Gulf of Mexico.
The water in the sink is usually heavily stained by tannins, which
accounts for the cave's name. In addition, colonies of freshwater
colonial cnidarians, Cordylophora lacustris, and several types of
zooplankton were reported from Little River Spring Cave in Suwannee
County (Streever 1993).

Phylum CHORDATA

Class OSTEICHTHYES

Family Anguillidae

Anguilla rostrata (Lesueur), American Eel (trogloxene). Holmes
County: Vortex Blue Spring (Helfman 1986). Jackson County: Gerard's
Cave (Pylka and Warren 1958, DSL), Hole-in-the-Wall (TM), Jackson
Blue Spring (TM), Milton's Well Cave (DSL), Twin Cave (PS, TM).
Leon County: Bird Sink (TM), Gopher Sink (Hobbs and Means 1972),
Little Dismal (TM), Munson Slough Blue (TM), Sullivan's Sink (TM).
Madison County: Thunderhole (TM). Marion County: Silver Glen Spring
(Hobbs and Franz 1992). Suwannee County: Bonnett Spring (TM), Charles
Spring (TM). Wakulla County: Wakulla Springs (TM).

Family Aphredoderidae
Aphredoderus sayanus (Gilliams), Pirate Perch (trogloxene). Jackson
County: Ellis Cave (Brockman and Bortone 1977). Suwannee County:
Irvine Slough Spring (TM), Orange Grove Sink (TM), Peacock Springs
(TM). REMARKS: This fish has been seen up 200 m inside the cave at
Peacock Springs and 16 m inside Irvine Slough Cave (TM).

44 Richard Franz, Judy Bauer and Tom Morris

Family Poeciliidae
Gambusia holbrooki Girard, Mosquitofish (trogloxene?). Alachua
County. Twin Chimneys Sink (Marshall 1947). Jackson County: Pool
Cave (DSL). Leon County. Gopher Sink (Hobbs and Means 1972).
Levy County. Half-Moon Cave (Marshall 1947). REMARKS: Recent
name change from Gambusia affinis holbrooki.

Family Moronidae
Morone saxitilus (Walbaum), Striped Bass (accidental). Marion
County. Silver Glen Spring (Hobbs and Franz 1992). REMARKS: Specimens
of this fish followed divers into the cave system at Silver Glen Springs
(Hobbs and Franz 1992).

Family Cyrinidae
Notropis harperi Fowler, Redeye Chub (trogloxene). Alachua
County. Bat Cave (Relyea and Sutton 19736), Cow Sink (Marshall
1947), Fern Cave (TM), Hog Sink (Hobbs 19426, Marshall 1947), Jerome
Sink (Marshall 1947), Martin's Cave (Relyea and Sutton 19736), Squirrel
Chimney (TM), Zamia Sink (Marshall 1947). Citrus County, unidentified
caves in Withlacoochee State Forest (DSL). Gilchrist County. Devil's
Eye and Ear Spring (TM), Hart Spring (TM), Little Devil's Spring
(TM), Otter Springs (TM). Holmes County. Vortex Spring (TM). Jackson
County. Jackson Blue Spring (TM), Twin Cave (TM). Leon County.
Gopher Sink (Hobbs and Means 1972), Half-Moon Cave (Marshall
1947), Pool Cave (O. G. Brock, personal communication, Florida Park
Service). Levy County. Gunpowder Sink (TM). Madison County. Baseline
Cave (TM), Thunderhole (TM). Marion County. Silver Glen Spring
(Hobbs and Franz 1992). Suwannee County. Bonnett Spring (TM), Irvine
Slough Cave (TM), Peacock Springs (TM), Wingate Well (TM). Wakulla
County. Wakulla Springs (TM). Washington County. Econfina Blue
Spring Cave (RF).

Family Ictaluridae
Ameiurus natalis (Lesueur), Yellow Bullhead {ixogXoxtnt). Alachua
County. Bat Cave (RF, Relyea and Sutton 19736), Chimney Sink (USNM),
Crumbly Sink (TM), Hornsby Spring (TM), Martin Cave (Relyea and
Sutton 19736), Still Sink (Cooper 19656). Columbia County. Jug Spring
on Ichetucknee River (RF), Rose Creek Swallet and Overflow (TM).
Gilchrist County. Devil's Eye and Ear Spring (TM), Hart Spring (TM),
Otter Spring (TM), Siphon Creek Cave (TM), Rock Bluff Spring (TM).
Hamilton County. Firecracker Cave (Pruitt 1991^, 1992), Pott Spring
(TM), Rossiter Spring (TM), White Springs (all dead, TM). Hernando

Caves and their Faunas in Florida and South Georgia 45

County: Die Polder 3 (TM). Holmes County: Vortex Spring (TM). Jackson
County: Hole in Wall (TM), Jackson Blue Spring (TM), Twin Cave
(PS). Lafayette County: Alligator Rescue Spring (TM), Aliens Mill
Pond Spring (TM), Lafayette Blue Spring (Pruitt 1992), Perry Spring
(TM). Leon County: Bird Sink Swallett (TM), Gopher Sink (TM), Munson
Slough Blue (TM). Levy County: Manatee Springs (TM), Octopus Cave
(Pruitt 1991fl). Madison County: M2 Blue Cave (Pruitt 1991J, 1992).
Thunderhole Sink (PH). Suwannee County: Anderson Spring (TM),
Azure Blue (TM), Bonnett Springs (TM), Irvine Slough (TM), Peacock
Springs (TM), Stick Sink (TM), Water Hole Cave (TM). Wakulla County:
McBride Slough (TM), Wakulla Springs (TM). REMARKS. Procambarus
pallidus was found in the stomach of a Chimney Sink catfish (notes
associated with USNM crayfish specimen); Procambarus lucifugus in
the stomach of Bat Cave catfish (Relyea and Sutton 19736). Pruitt
(1992) noted over a hundred individuals of this fish in Firecracker
Cave in July.

Ameiurus nebulosus (Lesueur) Brown Bullhead (trogloxene?).
Hamilton County: Firecracker Cave (Pruitt 1991c, 1992). Holmes County:
Vortex Spring (TM).

Family Centrachidae

Lepomis macrochirus Rafinesque, Blue Gill (trogloxene). Jackson
County: cave in Florida Caverns State Park (DSL), cave near dump
south of Marianna (DSL).

Pomoxis nigromaculatus (Lesueur), Black Crappie (accidental).
Suwannee County: Peacock III Spring (TM).

Class AMPHIBIA
Order CAUDATA
Family Ambystomatidae
Ambystoma opacum (Gravenhurst), Marbled Salamander
(accidental). Jackson County: unidentified cave on Milton property (DSL).
Ambystoma tigrinum tigrinum (Green), Eastern Tiger Salamander
(accidental). Jackson County: Milton's Well Cave (DSL).

Family Plethodontidae

Eurycea cirrigera (Green), Southern Two-lined Salamander
(accidental). Jackson County: Ellis Cave (DSL).

Eurycea longicauda guttolineata (Holbrook), Three-lined
Salamander (troglophile). /flc/:5(9« County: Ellis Cave (DSL), Gerome's
Cave (DSL), Miller's Cave (RF), Milton's Well Cave (DSL), Pottery

46 Richard Franz, Judy Bauer and Tom Morris

Cave (RF), Gerard's Cave (Lee 1969c). REMARKS: Larvae, trans-
formed juveniles, and adults were present in several caves.

Plethodon grobmani Neill (trogloxene). Jackson County: Ellis Cave
(DSL), Miller's Cave (RF), Milton's Well Cave (DSL), Pottery Cave
(RF).

Order ANURA ;

Family Leptodactylidae

Eleutherodactylus planirostris planirostris (Cope), Greenhouse

Frog (trogloxene). Alachua County: Bat Cave (RF). Citrus County:

Sweet Gum Cave (RF). Levy County: Octopus Cave (Pruitt 1991a).

Marion County: Orange Lake Cave (RF, DSL).

Family Bufonidae
Bufo terrestris Bonnaterre, Southern Toad (trogloxene). Jackson
Cow«/y: Milton's Well Cave (DSL).

Family Pelobatidae
Scaphiopus holbrooki holbrooki (Harlan), Eastern Spadefoot
(accidental). Alachua County: Warren Cave (RF).

Family Hylidae

Acris gryllus dorsalis (Harlan), Florida Cricket Frog (accidental).
Jackson County: Milton's Well Cave (DSL).

Pseudacris crucifer crucifer Weid, Northern Spring Peeper
(accidental). Jflc^50Az County: Milton's Well Cave (DSL).

Hyla gratiosa Leconte, Barking Tree Frog (accidental). Jackson
County: Milton's Well Cave (DSL), Miller's Cave (RF).

Hyla squirella Sonnini and Latreille, Squirrel Treefrog
(accidental). Jackson County: Milton's Well Cave (DSL).

Family Ranidae

Rana catesbeiana Shaw, Bullfrog (accidental?). Alachua County:
Bat Cave (RF), Goat Sink (Lee 1969c). Jackson County: Gerome's
Cave (DSL), Ellis Cave (RF), Gerard's Cave (Lee 1969c). Suwannee
County: Sim's Sink (RF). REMARKS: Large female bullfrog (173 mm
SVL) from Gerome's Cave contained an unidentified bat in its stomach
(DSL).

Rana clamitans Latreille, Bronze Frog (accidental). Jackson County:
Milton's Well Cave (DSL).

Rana grylio, Pig Frog (accidental). Levy County: Octopus Cave
(Pruitt 1991fl).

Caves and their Faunas in Florida and South Georgia 47

Rana utricularia Cope, Southern Leopard Frog (accidental?). Alachua
County: Bat Cave (RF), Warren Cave (Hobbs 1942b). Columbia County:
Riverbed Cave (Hobbs \9A2b). Jackson County: Gerard's Cave (Lee
1969c), Waddell's Mill Spring Cave (DSL). Levy County: Octopus Cave
(Pruitt 199\a). Marion County: Orange Lake Cave (RF, DSL), Sunday
Sink (RF).

Class REPTILIA

Order CROCODYLIA

Family Alligatoridae

Alligator mississipiensis (Daudin), American Alligator (accidental).

Jackson County: Small cave along the Chipola River floodplain (RF).

Lafayette County: Alligator Rescue Spring (TM). Levy County: Maddox

Cave (BP, BH, AK). REMARKS: A small alligator (approx. 1 m in

length) was wedged into the passage of a small cave at the base of a

limestone bluff along the floodplain of the Chipola River. A 2-m individual

repeatedly was found in Maddox Cave about 15 m inside the entrance.

The Lafayette animal was rescued from the bottom of a collapse sink

where it had apparently fallen and became entrapped. Other alligators

are reported from caves in the vicinity of Sawgrass Lake, southeast of

Archer, Levy County, Florida (A. Krause, personal communication).

Order TESTUDINES
Family Chelydridae
Chelydra serpentina osceola Stejneger, Florida Snapping Turtle
(accidental). Marion County Nickelberger Cave (PS).

Family Emydidae
Pseudemys floridana peninsularis Carr, Peninsula Cooter
(accidental). Citrus County: Hall's Bat Cave (DSL). REMARKS: Uniden-
tified Pseudemys also are reported from Octopus Cave, Levy County
(Pruitt \99\a).

Order SQUAMATA
Suborder LACERTIDAE
Family Anguidae
Ophisaurus ventralis (Linnaeus), Eastern Glass Lizard (accidental).
Jackson County: Milton's Well Cave (RF).

Suborder SERPENTES
Family Colubridae
Nerodia fasciata fasciata (Linnaeus), Banded Watersnake
(accidental). Jackson County: Pool Cave (RF).

48 Richard Franz, Judy Bauer and Tom Morris

Elaphe guttata guttata (Linnaeus), Corn Snake (trogloxene). Listed
only as Florida caves by Pylka (1957).

Elaphe obsoleta quadrivittata (Holbrook), Yellow Ratsnake
(trogloxene). Alachua County. Bat Cave (RF). Citrus County. Sweet
Gum Cave (Hobbs 19426, RF).

Elaphe obsoleta spiloides (Dumeril, Bibron, Dumeril), Gray Ratsnake
(trogloxene). Jackson County. Geromes Cave (RF, DSL), Gerard's Cave
(Lee 1969c).

Family Viperidae

Agkistrodon piscivorous conanti Gloyd, Florida Cottonmouth
(accidental?). Jackson County. Ellis Cave (DSL), Judges Cave (DSL).

Crotalus adamanteus Beauvois, Eastern Diamondback Rattlesnake
(accidental). Alachua County. Squirrel Chimney (Franz 1968).

Class AVES
Family Strigidae
Strix varia alleni Ridgway, Barred Owl (accidental?). Levy County.
Octopus Cave (Pruitt 1991^).

Family Cathartidae
Coragyps atratus (Bechstein), Black Vulture (accidental?, nesting
in cave). Jackson County. Unnamed cave on Merritt's Mill Pond.
REMARKS: Adult and two young were found 7 m inside the cave
entrance (TM).

Class MAMMALIA
Family Didelphidae
Didelphus virginana pigra Bangs, Virginia Opossum (accidental).
Listed only as Florida caves by Pylka (1957).

Family Soricidae
Blarina carolinensis carolinensis (Bachman), Short-tailed Shrew
(accidental). Levy County. Octopus Cave (Pruitt 1991^).

Family Molossidae
Tadarida brasiliensis cynocephala (LeConte), Freetail Bat
(accidental). Marion County, "several caves" (Morgan 1985). REMARKS:
Record based on single observations at Orange Lake Cave and Sunday
Sink in September 1974 and 1975, respectively. In each case, single
bats (males?) were found on the walls at night near the cave entrances.
The bats immediately took flight and left the cave when they were
disturbed (RF field notes).

Caves and their Faunas in Florida and South Georgia 49

Family Vespertilionidae

Pipistrellis subflavus floridanus Davis, Least Bat (trogloxene).
Alachua County: Goat Sink (RF). Gilchrist County. Roberts Cave (RF).
Levy County: Octopus Cave (Pruitt \99\a). Jackson County: Gerome's
Cave (DSL), Miller's Cave (Brock, personal communication), Old Indian
Cave (Rice 1955^, 19556, Jennings and Layne 1957), Gerard's Cave
(Lee 1969c). Marion County: Orange Lake Cave (RF), Sunday Sink
(RF).

Myotis austroriparious (Rhoads), Southeastern Bat (trogloxene).
Alachua County: Bat Cave (RF), Grant's Cave (RF, Rice 1957), Hog
Sink (Rice 1957), Jones Cave (McNab 1974), Seven Chimneys Sink
(McNab 1974), Warren Cave (McNab 1974). Citrus County: Sweet
Gum Cave (RF). Gilchrist County: Roberts Cave (RF, Rice 1957). Jack-
son County: Gerard's Cave (DSL), Old Indian Cave (Jennings and
Layne 1957), Mud Cave (Rice 19556). Levy County: Devil's Den (Pruitt
19916), Octopus Cave (Pruitt I99\a). Marion County: Hell Hole (BH),
Orange Lake Cave (RF), Sunday Sink (RF). Suwannee County: Devil's
Head and Horns, Mulky Road Sink (RF). (Also see Humphrey and
Gore 1992). REMARKS: This bat is proposed a candidate for federal
listing (Wood 1993)

Myotis grisescens Howell, Gray Bat (trogloxene). Jackson County:
Gerome's Cave (DSL), Gerard's Cave (DSL), Milton's Cave (DSL),
Old Indian Cave (Rice 19556, Jennings and Layne 1957, Lee and Tuttle
1970, Humphrey and Tuttle 1978), ONS Cave (DSL). REMARKS: The
Florida Committee on Rare and Endangered Plants and Animals listed
this bat as Endangered (Humphrey and Tuttle 1978; Gore 1992). This
bat also is listed by both the State of Florida and by U. S. Fish and
Wildlife Service as Endangered (Wood 1993).

Myotis keenii septentrionalis (Trouessart), Keen's Bat (trogloxene).
Jackson County: Old Indian Cave (Rice 19556, Scudder and Humphrey
1978). REMARKS: Only two specimens of this bat are known from
Florida. This bat was considered Endangered in the 1982 Florida Rare
and Endangered Biota volume on mammals (Scudder and Humphrey
1978) but not listed in the 1992 revised volume (Humphrey 1992).

Myotis sodalis Miller and Allen, Social Bat (trogloxene). Jackson
County: Old Indian Cave (Jennings and Layne 1957, Humphrey and
Scudder 1978). Only one specimen is known from Florida. The Florida
Committee on Rare and Endangered Plants and Animals listed this bat
as Endangered (Humphrey and Scudder 1978; Humphrey 1992). This
bat also is listed by the State of Florida and the U.S. Fish and Wildlife
Service as Endangered (Wood 1993).

50 Richard Franz, Judy Bauer and Tom Morris

Family Castoridae
Castor canadensis carolinensis Rhoads, Beaver (accidental).
Jackson County: Gerome's Cave (DSL).

Family Sciuridae
Sciurus carolinensis carolinensis Gmelin, Gray Squirrel (accidental).
Listed only as Florida caves by Pylka (1957).

Family Cricetidae

Peromyscus gossypinus gossypinus (Leconte), Cotton Mouse
(accidental). Levy County. Octopus Cave. Jackson County: Gerome's
Cave (DSL), Gerard's Cave (Lee 1969c).

Neotoma floridana floridana (Ord), Eastern Woodrat (trogloxene).
Columbia County: Bussey's Sink (RF). Jackson County: Gerome's
Cave (DSL), Old Indian Cave (DSL), Pool Cave (DSL), Waddell's
Mill Pond Cave (DSL), Gerard's Cave (Lee 1969c). Marion County:
Sunday Sink (RF). Suwannee County: Mulky Road Sink (RF).

Family Procyonidae
Procyon lotor elucus Bangs, Raccoon (accidental?). Levy County:
Octopus Cave (Pruitt \99la).

SUMMARY — The less specialized members of the Florida and
south Georgia cave faunas include: unidentified zooplankton, sponges,
one colonial cnidarian, two branchiobdellid annelids, unidentified
aeolosomatid oligocheates, two bivalves, five prosobranch and nine
pulmonata gastropods, four entocytherid and several other ostracods,
unidentified copepods, one isopod, two amphipods, eight decapods
(shrimps, crayfishes, and crab), two springtails, three crickets, nine
beetles, two parasitic flies, one vinegaroon, two harvestmen, five
spiders, unidentified mites, one millipede, one centipede, nine fishes,
five salamanders, 11 frogs, one crocodilian, two turtles, one lizard,
six snakes, two birds, and 13 mammals. The list consists of 37 acciden-
tals, 47 trogloxenes, 23 troglophiles, and four obligate commensals
on troglobite hosts. Of the vertebrates, a few fishes {Anguilla rostrata,
Notropis harperi, Ameiurus natalis), one salamander (Eurycea
longicauda), two frogs (Rana catesbeiana, R. utricularia, and five bats
{Pipistrellis subflavus, Myotis austroriparius, M. grisescens, M. keeni,
and M. sodalis) appear to have more than a casual relationship with
Florida caves. Myotis grisescens, M. keeni, and M. sodalis are the only

Caves and their Faunas in Florida and South Georgia 51

bat species recorded from Florida that are dependent on caves; the
latter two are reported only rarely and probably are not regular members
of Florida and south Georgia's cave faunas.

OBLIGATE CAVE FAUNAS AND KARST REGIONS

The biogeography of Florida and south Georgia's troglobites have
been discussed by Hobbs (1958), Caine (1974), Relyea et al. (1976),
Hobbs et al. 1977, Means (1977), Franz and Lee (1982), and others.
Each new discovery invites interpretative changes. The new distri-
butional records accumulated since 1982 necessitate further
comments concerning distributional patterns exhibited by this unique
group of species.

Six distinct cave faunas are suggested in the Florida and south
Georgia region (Fig. 2). Other unidentified faunas may emerge when
limestone areas outside of the geographic ranges of the six are better
surveyed. Each of the six faunas occupies a specific geographic range,
has precinctive taxa, and is characterized by specific geologic and
hydrologic characteristics. The two largest faunas (Ocala, St. Johns)
are broken into smaller assemblages (Table 1, Fig. 3). An assemblage
is defined as an isolated segment of a fauna that possesses distinctive
taxa.

Taxa are listed for each fauna in the accompanying faunal
descriptions; taxa associated with a specific assemblage are shown on
Table 1. In these faunal descriptions, one asterisk (*) preceding a
name indicates a precinctive species; double asterisks (**) identify
taxa that occur in three or more faunas.

EcoNFiNA Creek Fauna

Species List — *Dasyscias franzi, *Caecidotea sp.l. The fauna is
known from a single cave in the Econfina Creek basin. The karst area
associated with the Econfina Creek Fauna is located in southern Washing-
ton and northern Bay counties. It remains largely unexplored, although
there are numerous spring outlets along the mid-portions of Econfina
Creek where elements of the fauna may eventually be found. Additional
spring water also emerges from the Floridan aquifer directly into
Econfina Creek through fissures in the stream bed and from the base
of bluffs at points where the stream breaches overlying terrace deposits
(Vernon 1942, Musgrove et al. 1965). Econfina Blue Springs are composed
of several spring outlets along the edge of a low bluff that borders a
large spring pool on the east side of Econfina Creek. Combined flow
rates for these springs ranged from 32-51 cu. ft/sec. (1941-1972) (Rosenau
et al. 1977). The springs are developed at the contact between an upper

52

Richard Franz, Judy Bauer and Tom Morris

Fig. 2. Distributions of the six regional faunas. Dots indicate the
position of biologically significant caves in Florida (see Appendix 1).

shell bed and a fine- to medium-grained dolomitic calcarenite that
appears to be related to the Jackson Bluff group of marine sedi-
ments, probably Miocene in age (Muriel Hunter, personal communi-
cation, Tallahassee, Florida). Cooke (1945) considered the shell layer
to represent the "Cancellaria Zone" in the Duplin marl which overlies
a "cavernous limestone" in the Shoal River Formation. These spring
caves lie close enough to the surface that they are breached by
numerous small conical sinks in the adjoining upland. Available
samples of the Econfina Creek Fauna were obtained from a small,
permanently flowing cave stream at the base of one of these sink-
holes behind the most westerly spring outlet.

Caves and their Faunas in Florida and South Georgia 53

GULF COASTAL |^.-.r^>"

UPPER SUWANNEE

ORANGE LAKE
': MARION . I \

t #'{WITHLACOOCHEE ^

;:^^-

Fig. 3. Distribution of the assemblages associated with the Ocala
Fauna.

Apalachicola Fauna.

Species List — **Caecidotea hobbsi, *Cambarus cryptodytes,
Pseudosinella pecki, '^Islandiana sp., *Haideotriton wallacei. Three
of five taxa associated with the Apalachicola Fauna are precinctive.
This faunal region contains the only known terrestrial troglobites in
the state. The Apalachicola Fauna occurs in two segments, one in the
Marianna Lowlands of Jackson County, Florida, and the other in the
Dougherty Plain along the Flint River in Decatur and Dougherty coun-
ties, Georgia (Fig. 4) (Beck and Arden 1984, Lane 1989). Franz and
Lee (1982) suggested that the Marianna species were associated with
caves that were developed in the Ocala Group limestones at or near the

54 Richard Franz, Judy Bauer and Tom Morris

Table 1. Species composition of the assemblages associated with the Ocala
and St. Johns River cave faunas. Asterisk (*) identifies unique taxa in each
assemblage.

OCALA FAUNA

Upper Suwannee Assemblage.

Caecidotea hobhsi

Remasellus parvus

Crangonyx grandimanus

Crangonyx hobbsi
*Palaemonetes cummingi
*Procambarus erythrops
*Procambarus lucifugus alachua
*Procambarus pallidus

Troglocambarus maclanei
Lower Suwannee Assemblage

Crangonyx grandimanus

Crangonyx hobbsi

*Procambarus lucifugus X alachua

Troglocambarus maclanei
Orange Lake Assemblage

Crangonyx hobbsi
*Procambarus franzi

Troglocambarus maclanei
Marion Assemblage

Caecidotea hobbsi '

Crangonyx grandimanus

Crangonyx hobbsi
*Procambarus lucifugus X alachua

Troglocambarus maclanei
Withlacoochee Assemblage

Crangonyx hobbsi
*Procambarus lucifugus lucifugus

Troglocambarus maclanei
Gulf Coastal Lowlands Assemblage

Crangonyx grandimanus

Crangonyx hobbsi
*Procambarus leitheuseri

Troglocambarus maclanei

ST. JOHNS RIVER FAUNA
Wekiwa Assemblage
*Caecidotea sp. 2

*Procambarus acherontis d

*Troglocambarus sp.
Lake George Assemblage

*Procambarus attiguus .

*Procambarus delicatus
*Procambarus morrisi

Caves and their Faunas in Florida and South Georgia 55

Fig. 4. Caves associated with the (A) Marianna Lowlands and South-
west Georgia segments of the Apalachicola Fauna. Georgia sites — (B)
Climax Cave and (C) Albany well.

contact with the younger Suwannee and Marianna limestones (Floridan
aquifer). The troglobite records from southwest Georgia were taken
from a well at Albany in the heart of the Dougherty Plain and in cave
pools at Climax Cave near Climax, Georgia. Climax Cave lies at base
of the Pelham Escarpment of the Tipton Upland on the eastern edge of
the Dougherty Plain (Beck and Arden 1984). Like caves in the Marianna
area, Climax Cave is developed near the contact between the Suwannee
Limestone and the Ocala Group limestones (Floridan aquifer) (See Fig.
12b in Beck and Arden 1984).

Means (1977) felt that the two segments were continuous and
that the aquatic troglobites could disperse between the two areas wherever

56 Richard Franz, Judy Bauer and Tom Morris

there were solution channels in the limestone large enough to accommodate
them. Unfortunately, there are no records of these species from intermediate
areas, although Means (1977) noted a cave along the Apalachicola
River that should be sampled. No troglobites were located during pre-
liminary surveys of caves in Dothan County, Alabama (John E. Cooper,
personal communication, Raleigh, North Carolina). Slight or no morpho-
logical differences have been noted between the populations of Cambarus
cryptodytes and Haideotriton wallacei from the two areas (Pylka and
Warren 1958, Means 1977, Hobbs 1981), which further supports the
continuous population hypothesis.

WooDviLLE Fauna

Species List — Remasellus parvus, **Crangonyx hobbsi, **Crangonyx
grandimanus, *Procambarus horsti, *P. orcinus.The Woodville Fauna
is associated with the Ocala Group limestones (Floridan aquifer) in
the eroded portions of the Tallahassee Hills and the Woodville Karst
Plain, along and below the Cody Scarp, respectively. It is bounded on
the west by the Apalachicola Coastal Lowlands (see Hendry and Sproul
1966). The eastern limits remain to be defined, although none of the
fauna is known to occur east of the Aucilla drainage. Hendry and
Sproul (1966) and Yon (1966) show a more or less continuous limestone
shelf below the Cody Scarp (Woodville Karst Plain) across southern
Leon and Jefferson counties. Lane (1986:32) describes this area as a
"flat to gently undulating surface of sand that overlies carbonate rock.
The carbonates, which lie at shallow depths of 30 feet or less, have
undergone extensive solution by groundwater. This plain exhibits karst
features that are still evolving, for example: many old, well developed
sinkholes that are either permanently or intermittently flooded (Big
Dismal Sink), disappearing streams and natural bridges (Natural Bridge),
Wakulla Springs, and new sinkholes reported periodically."

The fauna appears to follow the riverine karsts associated with
the Wakulla-St. Marks rivers and the Wacissa River. In this way, the
endemic crayfishes parallel the distribution of the closely-related Pro-
cambarus pallidus that tracks the riverine karsts of the upper Suwannee
River and its tributaries. P. orcinus may be restricted to the Wakulla
drainage, whereas P. horsti may be more closely-tied to the St. Marks
and Wacissa drainages. The ecological relationships between them
remain unclear, particularly in lieu of an account that both crayfishes
co-exist in the Wakulla Springs Cave System (Morris 1989). More
collecting is necessary in caves of the Woodville Karst Plain to deter-
mine the actual geographic extent of this fauna and its ecological
specializations.

Caves and their Faunas in Florida and South Georgia 57

OcALA Fauna

Species List — **Caecidotea hobbsi, Remasellus parvus, **Crangonyx
grandimanus, **C. hobbsi, "^Palaemonetes cummingi, *Procambarus
erythrops, *P. franzi, *P. leitheuseri, *P. lucifugus lucifugus, *P. lucifugus
alachua, *P. lucifugus X alachua, *P.pallidus, *Troglocambarus maclanei.
The Ocala Fauna occurs in mature and riverine karst areas associated
with Ocala Group limestones (Floridan aquifer), from the Suwannee
River drainage, southwest through Alachua, Marion, Levy, Citrus, Hernando,
Pasco, and possibly Pinellas counties. Within this region, the fauna
appears fragmented into a series of six geographically distinct assemblages,
each characterized by endemic taxa (Table 1). The factor that consolidates
the Ocala Fauna is the presence of the crayfishes Procambarus lucifugus
(and allied species) and Troglocambarus maclanei which are represented
in every Ocala assemblage.

The Upper Suwannee appears to be the most distinctive of the
six assemblages. The Upper Suwannee incorporates the riverine karsts
along the upper Suwannee, (northern) Withlacoochee, and lower Santa
Fe rivers. It also spills over through the High Springs Gap onto the
Western Valley (=Newberry Karst Plain) between the Cody Scarp and
the (northern) Brooksville Ridge in western Alachua and northeastern
Levy counties (see Fig. 4 and discussions by White 1970, Hoenstine
and Lane 1991). In the Western Valley, the assemblage tracks what
appears to be an ancient stream channel that may have been a former
surface tributary of the Santa Fe River. The Lower Suwannee Assem-
blage is centered in a small karst area between the town of Bell in
Gilchrist County and the Chiefland-Manatee Springs area in Levy
County, west of Bell Ridge and the Waccasassa Flats. The Upper and
Lower Suwannee assemblages are each distinctive: Palaemonetes
cummingi, Procambarus lucifugus alachua, and P. pallidus in the
Upper Suwannee; Procambarus lucifugus X alachua, in the Lower
Suwannee. The Upper Suwannee includes populations of the isopod
Remasellus parvus that also occurs in the Wakulla area south of Talla-
hassee, but yet unrecorded from the Lower Suwannee. The Upper
Suwannee Assemblage (notably Procambarus pallidus) ranges down
the Suwannee River as far south as Rock Bluff Spring, but apparently
does not occur in the Bell karst. For unknown reasons there are
no troglobites known from limestone areas on the other bank of the
Suwannee River in Dixie County. Barriers to dispersal, if any exist,
that separate the Upper and Lower Suwannee assemblages have not
been identified.

The Orange Lake and Marion assemblages lie in karsts of Marion
County. The Orange Lake Assemblage, which includes the endemic

58

Richard Franz, Judy Bauer and Tom Morris

Sumter Upland

Fig. 5. Map showing important physiographic units that might influ-
ence the distributions of species of the Upper Suwannee, Orange Lake,
and Marion assemblages in Alachua and Marion counties.

Procambarus franzi, is associated with the eastern Fairfield Hills, south
of Orange Lake, whereas the Marion Assemblage occurs in a southern
extension of the Western Valley, the Sumter Upland, and the Oklawaha
portion of the Central Valley (at Silver Springs) (Fig. 5). Like the
Lower Suwannee, the Marion Assemblage includes an intergrade
population of Procambarus lucifugus.

Hydrologically, the Orange Lake and Marion assemblages fall
within the Oklawaha River Drainage Basin as defined by Faulkner
(1973) and Lane and Hoenstine (1991). The Fairfield Hills area repre-
sents the extreme northwestern corner of the Silver Springs Drainage
Area, which has its outlet at Silver Springs on the Oklawaha River in
the Central Valley. The Marion Assemblage occurs both in the Silver
Springs Drainage Area (at Silver Springs on the Oklawaha River) and
the Rainbow Springs Drainage Area with its primary outlet through
Rainbow Springs on the (southern) Withlacoochee River (Lane and
Hoenstine 1991).

Caves and their Faunas in Florida and South Georgia 59

The Withlacoochee Assemblage definitely is known from only
one locality (Sweet Gum Cave) on the east slope of the (southern)
Brooksville Ridge. A second record (cave 23.3 km north of Weekiwachee)
was listed in Hobbs (1942Z?) from Hernando County, but its specific
location remains unknown. The unique Procambarus lucifugus lucifugus
is reported from both sites. The directions listed with the Hernando
specimens, if taken literally, places the locality near the Citrus-Hernando
county line, probably near Chassahowitzka in Citrus County, on the
west side of the (southern) Brooksville Ridge. This area has low
relief, and any cave would be flooded and would likely function as a
spring, such as the Chassahowitzka Springs group near the town with
the same name (see Rosenau et al. 1977). However, if the collector's
directions were slightly off and the cave was actually northeast of
Weekiwachee, then the locality would fall on the eastern side of the
Brooksville Ridge, within 6 km of Sweet Gum Cave. More effort is
needed to locate appropriate caves along the (southern) Brooksville
Ridge, particularly in the vicinity of the Citrus-Hernando county line,
to resolve the Withlacoochee Assemblage's actual geographic extent.

The final assemblage occurs in the Gulf Coastal Lowlands in
southwestern Hernando and northwestern Pasco counties. The area
lies on the west side of the (southern) Brooksville Ridge, between the
towns of Weekiwachee and Hudson. A sighting of a small Troglo-
cambarus-likt crayfish in Knights Sink at Lake Tarpon suggests that
this assemblage's geographic range actually might extend as far south
as northern Pinellas County. It includes the unique Procambarus
leitheuseri. All of the known localities occur in a small coastal limestone
strip subdued in part by marine terrace deposits, between the 0.6-m (2
ft) and 9.1-m (30 ft) contour lines (Wetterhall 1965). Troglobites in
this assemblage have been recovered from several deep sinks and
springs associated with the ridge terrace (Die Polders) and from a
spring (Black Hole) in a coastal salt marsh.

St. Johns River Fauna

Species List — *Caecidotea sp. 2, '^Procambarus acherontis, *P.
attiguus, *P. delicatus, *P. morrisi, ^Troglocambarus sp. The St. Johns
River Fauna is still incompletely known, and more new taxa are expected
as divers penetrate unexplored karst windows and springs along the St.
Johns and lower Oklawaha rivers. There are two related assemblages
of troglobites within this fauna, one focusing on a small karst area
along the Wekiva River in the Orlando metropolitan area and the other
occurring along the west shore of Lake George, Alexander Spring Run,
and the lower Oklawaha River. No species have been found in the

60 Richard Franz, Judy Bauer and Tom Morris

30-lcm stretch between the two areas. All of the known occurrences of
this fauna are in flooded caves associated with springs, sinkholes, and
wells. These cave systems are developed near the contact between the
basal units of the Hawthorne Formation and the underlying Ocala Group
limestones (Floridan aquifer) (Hobbs and Franz 1986, 1990, 1992).

The Wekiva Assemblage includes an undescribed isopod (Caecidotea
sp. 2), Procambarus acherontis, and an unstudied population of Troglo-
cambarus. It occurs in a small limestone plain that lies along the
Wekiva River, and east of the Lake Apopka basin, in western Orange
and Seminole counties. Groundwater originates in a sand hill region
south of the sinkhole plain. A north-directed branch of this water
flows northeast towards the Wekiva valley, but an eastern branch
intersects a "bad water zone" in eastern Orange County that contains
dissolved solids in excess of 1,800 ppm (Barraclough 1962, Lichtler
et al. 1968). The north branch emerges as springs along the south
escarpment of the river (Rock, Wekiwa, Palm, Sanlando, Witherington,
and Barrel springs). Analysis of water from these springs shows less
than 150 ppm of dissolved solids, and several, such as Palm Springs,
have the odor and white slimy bacteria that are associated with sulphur
springs.

The Lake George Assemblage remains poorly known. Three
crayfishes are known: Procambarus attiguus, P. delicatus, and P.
morrisi. The first two belong to the seminolae group and probably are
related to Troglocambarus, whereas Procambarus morrisi is related
to P. acherontis. Riverine karsts extend north of the Lake George
springs area to at least Green Cove Springs in Clay County. Divers
that penetrated the spring at Green Cove reported no troglobites (WS),
although more effort is necessary, possibly using baited traps, to verify
their observations. More exploration in the riverine karsts along the
Oklawaha River should be encouraged. The presence of Procambarus
attiguus at Devils Sink in Putnam County and Procambarus lucifugus
and sightings of isopods and amphipods in Silver Springs suggest a
region with a complex biota. ^

Miami Fauna

Species List — **Crangonyx grandimanus, **C. hobbsi, "^Procambarus
milleri. The Miami Fauna was collected from shallow wells at the
Little Bird Nursery and Garden Store in Miami (Hobbs 1971) and in
northeast Homestead (HHH, personal communication). The latter site
is approximately 22 km southwest of the type locality (W. Loftus,
personal communication). According to Hobbs (1971), the original specimens
from the type locality were obtained from a trap at the outlet of a
motorized pump associated with the Little Bird well in 1968. Both

Caves and their Faunas in Florida and South Georgia 61

sites are associated with the Miami oolite. Parker and Cook (1944)
indicated that since so much of the Miami oolite is occupied by solution
holes it is highly permeable in a vertical direction and is a good aquifer
that furnishes supplies to many small wells. However, they report it to
have a low horizontal permeability which theoretically might restrict
the dispersal of larger troglobites. This fauna is currently known from
only two sites, and it should be searched for elsewhere in suitable
groundwater habitats in Dade County. The Miami Fauna is associated
with the Biscayne aquifer, unlike cave faunas in north Florida and
south Georgia that occur in the Floridan aquifer.

FAUNAL RELATIONSHIPS

Cave crayfishes are the best indicators of faunal relationships
because the other Florida troglobites either have very limited distributions
or range over at least three faunal areas. Troglobitic crayfishes are
found in all but the Econfina Creek Fauna.

Six crayfish lineages are involved: (1) Cambarus (subgenus
Jugicambarus) represented by Cambarus cryptodytes restricted to the
Apalachicola Fauna; (2) Procambarus (subgenus Leconticambarus),
by Procambarus milleri, Miami Fauna; (3) Procambarus (subgenus
Lonnbergius), by Procambarus acherontis and P. morrisi, St. Johns
River Fauna; (4) Procambarus (subgenus Ortmannicus, Pictus Group,
lucifugus complex), by Procambarus erythrops, P. franzi, P. leitheuseri,
P. lucifugus lucifugus, P. lucifugus alachua, P. lucifugus intergrade
populations, Ocala Fauna; (5) Procambarus (subgenus Ortmannicus,
Pictus Group, pallidus complex), by Procambarus horsti, P. orcinus,
P. pallidus, Woodville and Ocala faunas; and (6) Procambarus (sub-
genus Ortmannicus, Seminolae Group) and Troglocambarus, by Pro-
cambarus attiguus, P. delicatus, Troglocambarus maclanei, and
Troglocambarus sp., St. Johns River and Ocala faunas.

Lineages 1-4 — The first four lineages have restricted distributions
and therefore provide little information concerning the relationships
between faunal groups.

Lineage 5 — The pallidus complex occurs in both the Woodville
Fauna and the Upper Suwannee Assemblage of the Ocala Fauna. As a
group, they are thought to have had a common ancestor, one similar
in morphology to the extant Procambarus leptodactylus, which now
occurs in streams north of Florida (Hobbs 1958, Franz and Lee 1982,
Hobbs and Franz 1986). The distribution of the pallidus complex cor-
responds to the western slope of the old Northern Highlands described
by White (1970) and is associated with Eocene limestone areas along
the Cody Scarp (Fig. 6). Troglobites have not been found in the intervening

62

Richard Franz, Judy Bauer and Tom Morris

Fig. 6. Distribution of Procambarus horsti, Procambarus orcinus, and
Procambarus pallidus.

area that separates the two faunas, although more exploration in the
50-km-wide hiatus may change this view. We suggest that the evolutionary
history of the pallidus complex involved independent invasions of
ancestral leptodactylus-like stocks into geographically isolated ground-
water reservoirs associated with the Cody Scarp in the Woodville and
Upper Suwannee areas. Subsequent evolution led to the differentiation
of the horstiorcinus branch in the St. Marks-WakuUa-Wacissa drainages
and the pallidus branch in the Suwannee basin.

Lineage 6 — Hobbs (1942) and Hobbs and Franz (1986) pointed
out the similarities between Troglocambarus maclanei and members
of the Seminolae Group in the genus Procambarus. They felt that
these features indicated a evolutionary relationship between the two

Caves and their Faunas in Florida and South Georgia 63

■ATTIGUUS

•i ^'^<''' ■ ^
TROGLOCAMBARUS, ,^ N^. 'r'^CDELICATUS

^TROGLOCAMBARUS
sp.

Fig. 7. Distribution of Procambarus attiguus, Procambarus delicatus,
Troglocambarus sp., and Troglocambarus maclanei. Arrow shows the
suggested direction for the dispersal of Troglocambarus lineage.

groups of crayfishes. The recent discoveries of two troglobitic members
of the Seminole Group {Procambarus attiguus and P. delicatus) and an
unstudied population of Troglocambarus in the St. Johns River basin
(Fig. 7) allows for further speculation concerning the evolution of this
lineage. We propose that the initial differentiation of Troglocambarus
from ancestral Procambarus stocks occurred in karsts associated with
the St. Johns River. Then, one branch of Troglocambarus dispersed
from the St. Johns River karst area, via subterranean routes, into cave
systems on the western slope of the Northern Highland (Fig. 7). They
probably arrived on the western slope already preadapted for co-
existence with larger, less specialized cave Procambarus (lineages 4
and 5). Unfortunately, the absence of Form I males for the Orlando

64 Richard Franz, Judy Bauer and Tom Morris

Troglocambarus prevents a more scholarly interpretation of the relation-
ships between the St. Johns and Ocala populations of Troglocambarus
and between the Orlando Troglocambarus and Procambarus attiguus
and P. delicatus.

CONSERVATION

Troglobitic populations of most Florida and south Georgia faunas
are potentially susceptible to human disturbance. Because these cave
faunas are composed of primarily aquatic species, they could be threat-
ened by (1) groundwater contamination that results from saltwater
intrusion due to excessive pumping of groundwater and from the
introduction of agricultural and industrial chemicals as well as human
and domesticated animal wastes into groundwater reservoirs. Ground-
water contamination may also result from vandalism and other direct
human-related disturbances.

Cave fauna kills have been documented on at least four occasions
in caves associated with the Suwannee and Wakulla drainages. On
one occasions, Streever (1992^) reported a complete faunal kill that
included, not only cave crayfishes, but also yellow bullheads, American
eels, and Corbicula clams. This event followed the introduction of
cold flood waters into the cave from the Suwannee River which forced
the spring to reverse. The kill may have resulted from sudden changes
in water temperature, oxygen deprivation, or the introductions of toxic
materials. Another catfish kill was noted at White Springs, but no
other information is available about the conditions that may have
sponsored this event (TM). Crustacean kills were also noted at Edwards
Spring and Falmouth Spring-Siphon in May 1989. Unlike Streever's
account, these springs had not reversed, and there was no injection of
cold water. Over 50 cave crayfish (probably Procambarus pallidus)
and large numbers of isopods and amphipods were found dead at
Edwards Spring. Divers reported that they found blue pellets spread
on the ground in a managed pine forest in the vicinity of several large
sinks that formed the upstream portion of the Edwards Spring cave.
They suggested that the pellets were possibly a herbicide used to
control hardwoods in the plantation.

The Squirrel Chimney Cave Shrimp was federally listed as
Threatened (Anonymous 1990), and the Florida Cave Amphipod,
Hobbs' Cave Amphipod, and Orlando Cave Crayfish are being con-
sidered as candidates for listing by the U.S. Fish and Wildlife Service
(Wood 1992). The Santa Fe Cave Crayfish is considered a Species of
Special Concern by the State of Florida (Wood 1992). Most of the
other Florida troglobitic fauna has been proposed by the Florida

Caves and their Faunas in Florida and South Georgia 65

Committee on Rare and Endangered Plants and Animals for listing by
the state (Franz 1982).

Populations of the following troglobitic taxa have been found in
caves that occur on federal, state, and private conservation lands where
they receive some protection: Dasyscias franzi (Econfina Blue Spring
Preserve [Northwest Water Management District]), Caecidotea hobbsi
(Dudley Farm State Historical Site [Florida Park Service]), Caecidotea
sp. 1 (Econfina Blue Spring Preserve), Caecidotea sp. 2 (Rock Springs
County Park), Remasellus parvus (Peacock Springs State Preserve [Florida
Park Service]), Crangonyx grandimanus (Peacock Springs State Preserve,
Leon Sinks Recreation Area [U.S. Forest Service], Wakulla Springs
State Park [Florida Park Service]), C. hobbsi (Dudley Farm State
Historical Site, Peacock Springs State Preserve, Wakulla Springs State
Park), Procambarus acherontis (Wekiwa Springs State Park [Florida
Park Service]), P. attiguus (Silver Glen Springs Recreation Area [U.
S. Forest Service]), P. delicatus (Alexander Springs Recreation Area
[U.S. Forest Service]), P. erythrops (Sims Sink Preserve [The Nature
Conservancy]), P. lucifugus alachua (Dudley Farm State Historical
Site), P. lucifugus X alachua (Manatee Springs State Park [Florida
Park Service]), P. orcinus (Wakulla Springs State Park, Leon Sinks
Recreation Area), P. pallidus (Suwannee River State Park [Florida
Park Service], Peacock Springs State Preserve), Troglocambarus maclanei
(Manatee Springs State Park), Cambarus cryptodytes (Florida Caverns
State Park [Florida Park Service]), Pseudosinella pecki (Florida Caverns
State Park), Islandiana sp. (Florida Caverns State Park), and Haideotriton
wallacei (Florida Caverns State Park). There is no such protection for
six taxa {Procambarus franzi, P. horsti, P. leitheuseri, Marion population
of Procambarus lucifugus X alachua, P. morrisi, and Troglocambarus
sp.). Important populations of these invertebrates should be incorporated
into future land acquisition projects in order to reduce their chances
of extinction.

CONCLUSIONS

More survey work is needed in Florida and south Georgia to
understand the complex relationships between the regional faunas.
New troglobites undoubtedly await discovery when cavers and cave
divers explore karst windows outside normally visited areas. Large
areas of the state still remain uncharted. We recommend that divers
wishing to assist in future biological discoveries look in the Choc-
tawhatchee River basin, Econfina Creek karst area in Bay and Wash-
ington counties, scarp areas between Tallahassee and the Suwannee

66 Richard Franz, Judy Bauer and Tom Morris

River, Georgia portions of tlie (northern) Withlacoochee River and the
Flint River, St. Johns River basin, and southern Florida.

We encourage the collections of specimens of both macro- and
micro-invertebrates that represent new records; specimens should be
preserved in alcohol (70% ethanol) at the time of collection. All specimens
should be donated to appropriate collections where they can be properly
studied and curated. If maintained alive, specimens need to be brought
to a specialist immediately, for upon death these fragile organisms
decay rapidly and are useless. . ^,

ACKNOWLEDGEMENTS. —Since this project spans 25 years, it
is difficult to list everyone who has contributed information to the data
base. However, we particularly are indebted to David S. Lee (DSL),
who introduced Richard Franz (RF) to Florida caves in the late 1960s,
the many cavers and cave divers who graciously contributed specimens
and observations, and especially the land owners who have allowed
access to their caves over the years. We wish to thank Thomas E.
Bowman (TB), Horton H. Hobbs, Jr. (HHH), Jerry Lewis (JL), John R.
Holsinger (JRH), Arnold Norden (AN), and Fred G. Thompson (FGT)
for aid in identification of crustacean and molluscan materials; Jim
Stevenson (Florida Department of Natural Resources) and Joe Fredericks
(former park manager at Florida Caverns State Park) who allowed Lee
and Franz to work in caves of Florida Caverns and Falling Waters state
parks in the late 1960s; Bill and Sandy Fehring, Steve Gerrard, Paul
Heinerth (PH), Brian Houha (BH), the late Bill Hurst (BHT), David S.
Lee (DSL), A. Terry Leitheuser (ATL), Buford Pruitt (BP), Wes Skiles
(WS), Paul Smith (PS), and Roger Werner who have continued to
contribute important specimens through the years often from difficult
sites; the Florida Speleological Society (FSS) and Al Krause (AL) for
sharing information in their files; Walter Auffenberg, Peter Drummond,
Horton H. Hobbs Jr., David S. Lee, Barry Mansell, Bill and Shirley
Oldacre, Joseph M. Pylka, Richard D. Warren, and H. K. Wallace for
help in reconstructing the history of biospeleology in Florida; Thomas
C. Barr, Jr., C. Kenneth Dodd, Jr., Howard Frank, Horton H. Hobbs,
Jr., Al and Martha Krause, and David S. Lee who carefully reviewed
parts or all of the manuscript; and the staff of the Florida Museum of
Natural History for their continued support of this project. The following
museum collections include important specimens from Florida caves:
British Museum of Natural History (BMNH); Florida Department of
Natural Resources (FSBC=FDNR); Florida Museum of Natural History
(UF); National Museum of Natural History (Smithsonian Institution)
(USNM); North Carolina State Museum of Natural Science (NCSM);

Caves and their Faunas in Florida and South Georgia 67

Museum of Comparative Zoology (MCZ); National Museum of Canada
(NMC); Ohio State Museum (OSM); Zoology Institute, Academy of
Science, Russia, St. Petersburg (ZIAS).

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Received 12 October 1992
Accepted 1 August 1993

Caves and their Faunas in Florida and South Georgia 79

APPENDIX 1. List of biologically significant caves and
springs in Florida and South Georgia. Sites are listed by county.
Pertinent data for each site include an identification code, accepted
site name (and other names when appropriate), location (section,
township, range, and name of the appropriate U. S. Geological
Survey quadrangle map), regional faunas and assemblages, list
of species (collection, authority, or reference), ownership, and
relevant references with annotations (abbreviations represent troglo-
bitic taxa). Abbreviations used in this section are as follows:
Cl=Cambarincola leoni, \52i-Uncinocy there ambophora, \J\=U. lucifuga,
Uw=U. warreni, C2ih=Caecidotea hobbsi, Rp=Remasellus parvus,
Ch=Crangonyx hobbsi, Cg=C. grandimanus, Vc=Palaemonetes cum-
mingi, Fa=Procambarus acherontis, Pat=P. attiguus, Pd=P. delicatus,
Pe=P. erythrops, Ph=P. horsti, Ff=P. franzi, Ple=P. leitheuseri,
Pla=P. lucifugus alachua, FU=Procambarus lucifugus lucifugus,
PlXa=P. lucifugus intergrade, Pmi=P. milleri, Pm=F. morrisi, Po=P.
orcinus, Pp=P. pallidus, Cc=Cambarus cryptodytes, Tm=Troglocam-
barus maclanei, Tsp=Troglocambarus sp., Vsp=Pseudosinella pecki,
lp=Islandiana sp., Df=Dasyscias franzi, Hw=Haideotriton wallacei.
Other abbreviations are listed in the Methods and Acknowledgments
sections.

ALACHUA COUNTY

ALA-1. ALACHUA SINK (or Lime Sink, Alachua Green Sink)
(Sec.lO, T.8S, R.18E, Alachua Quad.). OCALA FAUNA (Upper Suwan-
nee), Procambarus pallidus (USNM), unidentified cave amphipods
(TM). Private, susceptible to groundwater pollution from parking lot
run-off. REFERENCES: Franz and Lee 1982 (Pp).

ALA-2. AULSBROOK CAVE (Sec.35, T.9S, R.17E, Newberry
Quad.). OCALA FAUNA (Upper Suwannee), Caecidotea hobbsi (RF).
Private. REFERENCES: Franz 1982 (Cah).

ALA-3. BARBIES CAVE (Sec.l4, T.IOS, R.17E, Archer Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes
(FSS files). Private.

ALA-4. BAT CAVE (Sec.l8, T.9S, R.17E, Waters Lake Quad.).
OCALA FAUNA (Upper Suwannee), Caecidotea hobbsi (USNM),
Procambarus lucifugus alachua (USNM), and other cave associated
species. Private. REFERENCES: Franz 1982 (Pla); Hobbs et al. 1977
(Pla); NcNab 1974 (bats); Relyea and Sutton 1973/? (Pla, fish); Rice
1957 (bats).

80 Richard Franz, Judy Bauer and Tom Morris

ALA-5. BUZZARD'S ROOST (Sec.28, T.8S, R.18E, Alachua Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes. Private.

ALA-6. CHIMNEY SINK (unidentified site). OCALA FAUNA
(Upper Suwannee), Procambarus pallidus (USNM), other cave-
associated species. Private. REFERENCES: Franz 1982 (Pp); Hobbs et
al. 1977 (Pp).

ALA-7. CRUMBLY SINK (Sec.22, T.9S, R.17E, Newberry Quad.).
OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi (JRH), Pro-
cambarus lucifugus alachua (USNM). Private.

ALA-8. CUEVA FRIA (Sec.24, T.IOS, R.18E, Arrendondo Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus lucifugus alachua
(USNM), Procambarus pallidus (USNM). Private. REFERENCES:
Franz 1982 (Pla, Pp); Hobbs l9A2b (Pla).

ALA-9. COW SINK (Sec.21, T.9S, R.17E, Newberry Quad.)
OCALA FAUNA (Upper Suwannee), cave-associated species.
REFERENCES: Marshall 1947 (fish).

ALA-10. DEVIL'S HOLE (or Glover Sink) (Sec.l8, T.IOS, R.18E,
Archer Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi
(Hobbs 1942Z?), Procambarus pallidus (USNM), other cave-associated
species. Private. REFERENCES: Franz 1982 (Ch, Pp); Hobbs 1942Z?
(Ch, Pp); Hobbs et al. 1977 (Pp).

ALA-11. DOUBLE-BARREL SINK (or Double Sink Cave) (Sec.
24, T.IOS, R.17E, Archer Quad.). OCALA FAUNA (Upper Suwannee),
unidentified cave crayfishes (BHT, pers. comm., 1975). Private.

ALA-12. DUDLEY CAVE (Sec.32, T.9S, R.18E, Newberry Quad.).
OCALA FAUNA (Upper Suwannee), Caecidotea hobbsi-type locality
(USNM), Crangonyx hobbsi (JRH), Procambarus lucifugus alachua
(USNM), other cave-associated species. Public, Dudley Farm State Historical
Site, Florida Dept. Natural Resources. REFERENCES:
Franz 1982 (Cah, Ch, Pla); Hobbs 1942^ (Cah, Ch, Pla); Krause 1990fl
(cave description, map); Maloney 1939 (Cah-type description); Peck
1970 (terrestrial arthropods); Warren 1961 (Cah, Ch, Pla).

ALA-13. DUDLEY TUNNEL (or Dudleys Cave II) (Sec.32, T.9S,
R.18E, Newberry Quad.). OCALA FAUNA (Upper Suwannee),
Caecidotea hobbsi (USNM). Public, Dudley's Farm State Historical
Site, Florida Dept. Natural Resources. REFERENCES: Franz 1982 (Cah);
Hobbs 1942 (Cah); Krause 1990^ (cave description, map); Warren 1961
(Cah).

ALA-14. DUFF'S CAVE (Sec.l4, T.IOS, R.17E, Archer Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes
(FSS files). Private.

Caves and their Faunas in Florida and South Georgia 81

ALA-15. FELLOE CAVE (Sec.2, T.IOS, R.18E, Gainesville West
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave cray-
fishes (FSS files). Private.

ALA-16. FERN CAVE (or Fern Sink) (Sec.34, T.8S, R.17E, Newberry
Quad.). OCALA FAUNA (Upper Suwannee), unidenti-
fied cave crayfish (FSS files), other cave-associated species. Private.

ALA-17. GOAT SINK (Sec.20, T.9S, R.18E, Newberry Quad.).
OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi, Procam-
barus lucifugus alachua (USNM), Procambarus pallidus, Troglo-
cambarus maclanei (USNM), other cave-associated species. Private.
REFERENCES: Cooper 1965/? (Pla, cave description); Hobbs 19426
(Pla); Hobbs et al. 1977 (Pla, Pp); Holt 19736 (CI as commensal on
Pla); Lee 1969c (bullfrogs); Warren 1961 (Pla).

ALA-18. GRANT'S CAVE (Sec.3, T.IOS, R.18E, Newberry Quad.).
OCALA FAUNA, cave-associated species. Private. REFERENCES: Peck
1970 (terrestrial arthropods); Rice 1957 (bats).

ALA-19. HAGUE CAVE (Sec.9, T.8S, R.18E, High Springs Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes
(FSS files). Private.

ALA-20. HERTZOG CAVE (or Herzog's Cave, Wagon Wheel
Cave) (Sec. line 17/18, T.IOS, R.19E, Arrendondo Quad.). OCALA
FAUNA (Upper Suwannee), Crangonyx grandimanus (JRH), Procam-
barus pallidus (USNM), Troglocambarus maclanei (USNM). Private.

ALA-21. HIGH SPRINGS CAVE (Sec.2, T.8S, R.17E, High Springs
Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi, Pro-
cambarus pallidus (USNM). Private. REFERENCES: Franz 1982 (Ch,
Pp); Hobbs 19426 (Ch, Pp); Hobbs et al. 1977 (Pp); Warren 1961 (Ch,
Pp).

ALA-22. HOG SINK (Sec.24, T.IOS, R.18E, Arrendondo Quad.).
OCALA FAUNA (Upper Suwannee), Uncinocythere lucifuga (Walton
and Hobbs 1959), Procambarus lucifugus alachua-type locality
(USNM), Procambarus pallidus (USNM), other cave-associated species.
Private. REFERENCES: Franz 1982 (Ch, Pla); Franz and Lee 1982
(Pla); Hobbs 19426 (Pla); Hobbs et al. 1977; (Pla) Marshall 1947 (fish);
Rice 1957 (bats); Walton and Hobbs 1959 (Ul); Warren 1961 (Ch, Pla,
Pp).

ALA-23. HORNSBY SINK (Sec.26, T.7S, R.17E, High Springs
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM). Private, children's camp. REFERENCES: Franz 1982 (Pp);
Hobbs et al. 1977 (Pp).

ALA-24. HORNSBY SPRING (Sec.27, T.7S, R.17E, High Springs
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus

82 Richard Franz, Judy Bauer and Tom Morris

(ATL). Private, children's camp. REFERENCES: Auffenberg 1957^
(fossils); Dolan and Allen 1961 (archeology). Lane 1986 (geology),
Rosenau et al. 1977 (spring description); Webb 1974 (fossils).

ALA-25. HUGGINS CAVE (Sec.35, T.8S, R.17E, Newberry Quad.).
OCALA FAUNA (Upper Suwannee), Crangonyx grandimanus,
Crangonyx hobbsi-type locality (USNM). Private. REFERENCES:
Franz 1982 (Cg, Ch); Shoemaker 1941 (Ch-type description); Warren
1961 (Cg, Ch).

ALA-26. JEROME SINK (Sec.22, T.9S, R.17E, Newberry Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes (FSS
files), other cave-associated species. Private. REFERNCES: Marshall
1947 (fish).

ALA-27. JONES CAVE (or Witches Den) (Sec.l7, T.9S, R.17E,
Waters Lake Quad.). OCALA FAUNA (Upper Suwannee), cave-
associated species. REFERENCES: McNab 1974 (bats).

ALA-28. JOOK'S CAVE (unidentified site). OCALA FAUNA
(Upper Suwannee), cave-associated species. REFERENCES: Hubbell
1936 (crickets); Peck 1970 (terrestrial arthropods).

ALA-29. MARTIN'S CAVE (possibly Seven Chimneys) (Sec.l8,
T.9S, R.17E, Waters Lake Quad.). OCALA FAUNA (Upper Suwannee),
Procambarus lucifugus alachua, other cave-associated species. Private.
REFERENCES: Franz 1982 (Pla); Relyea and Sutton 1973Z> (Pla,
catfish).

ALA-30. MCGEEHEE BLUE HOLE (Sec. 15, T.9S, R.17E,
Newberry Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidas (ATL). Private.

ALA-31. MCGEEHEE CHIMNEY (Sec. 16, T.9S, R.17E,
Newberry Quad.). OCALA FAUNA (Upper Suwannee), unidentified
cave crayfishes (FSS files). Private.

ALA-32. O'STEEN'S CAVE (Sec.26, T.8S, R.17E, High Springs
Quad.). OCALA FAUNA (Upper Suwannee), cave-associated species.
REFERENCES: Hubbell 1936 (crickets); Peck 1970 (terrestrial
arthropods).

ALA-33. PALLIDUS SINK (Sec.l5, T.8S, R.17E, High Springs
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM). Private. REFERENCES: Franz 1982 (Pp); Hobbs l9A2b (Pp);
Hobbs et al. 1977 (Pp); Warren 1961 (Pp).

ALA-34. POWERHOUSE SINK (Sec.22, T.8S, R.18E, Alachua
Quad.). OCALA FAUNA (Upper Suwannee), unidentified crayfishes
(FSS files). Private.

ALA-35. PROTHEROE SINK (Sec.24, T.IOS, R.18, Arrendondo
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus lucifugus

Caves and their Faunas in Florida and South Georgia 83

alachua (Cooper 1965^?), Procambarus pallidus (Cooper 19656). Pri-
vate. REFERENCES: Cooper 1965b (Pla, Pp, cave description); Franz
1982 (Pla. Pp); Hobbs et al. 1977 (Pla, Pp); Warren 1961 (Pla, Pp).

ALA-36. SCHOUTEN CAVE (Sec.5, T.IOS, R.18E, Newberry
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave cray-
fishes (FSS files). Private.

ALA-37. SEVEN CHIMNEYS SINK (Sec.l7, T.9S, R.17E, Waters
Lake Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
lucifugus alachua. REFERENCES: Franz 1982 (Pla); Hobbs et al. 1977
(Pla); McNab 1974 (bats).

ALA-38. SQUIRREL CHIMNEY (Sec.21, T.9S, R.18E, Newberry
Quad.). OCALA FAUNA (Upper Suwannee), Uncinocythere lucifuga
(Walton and Hobbs 1959), Palaemonetes cummingi-typt locality
(USNM), Procambarus lucifugus alachua, Procambarus pallidus
(USNM), Troglocambarus maclanei-type locality (USNM), other cave-
associated species. Private. REFERENCES: Anonymous 1990 (Pc,
federal conservation status); Dickson and Franz 1980 (Pp, gill respira-
tion); Dobkin 1971 (Pc, larval development); Franz 1982 (Ch, Pc, Pla,
Pp, Tm); Franz and Lee 1982 (Pla, Pp, Tm, ecology); Hobbs 1942«
(Tm-type description), 1942/? (Pp, Tm); Hobbs et al. 1977 (Pla, Pp,
Tm); Holt 1973/? (CI as commensal of TM); Mohr and Poulson 1966
(Pla, Tm, cave description, photo); Morris and Butt 1992 (Pc, cave
description); Peck 1970 (terrestrial arthropods); Relyea and Sutton 1973^
(Pp, egg-bearing); Walton and Hobbs 1959 (Ul); Warren 1961 (Pp,
Tm).

ALA-39. STILL SINK (Sec.29, T.9S, R.18E, Newberry Quad.).
OCALA FAUNA (Upper Suwanneee), Crangonyx hobbsi (JRH), Pro-
cambarus pallidus (USNM). Private. REFERENCES: Cooper 1965/?
(Pp, cave description); Franz 1982 (Pp); Hobbs, et al. 1977 (Pp).

ALA-40. TEN INCH CAVE (Sec. 3, T.9S, R.17E, Newberry
Quad.). OCALA FAUNA (Upper Suwannee), Remasellus parvus-typc
locality (USNM). Private. REFERENCES: Steeves 1964 (Rp, type
description).

ALA-41. TUSK CAVE (Sec.34, T.9S, R.18E, Gainesville West
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus lucifugus
alachua (USNM). Private. REFERENCES: Franz and Lee 1982 (Pla).

ALA-42. WARREN CAVE (Sec.l3, T.9S, R.18E, Gainesville West
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus-iypQ locality (USNM), other cave-associated species. Private,
National Speleological Society, cave preserve. REFERENCES: Franz
1982 (Pp); Franz and Lee 1982 (Pp); Hobbs 1942/? (Pp, type description);

84 Richard Franz, Judy Bauer and Tom Morris

Hobbs et al. 1977 (Pp); Krause 1992 (cave description, map); McNab
1974 (bats); Peck 1970 (terrestrial arthropods); Warren 1961 (Pp).

ALA-43. WELL, FORT CLARK CHURCH. (Sec.31-32, T.9S. R.19E,
Gainesville West. Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidas (USNM), Private. REFERENCES: Franz 1982 (Pp); Hobbs et
al. 1977 (Pp).

ALA-44. WELL, MICANOPY (Unidentified site). OCALA FAUNA
(Orange Lake), Caecidotea hobbsi (USNM). Private. REFERENCES:
Steeves 1964 (Cah).

ALA-45. WELL, MICANOPY (Archie Carr Farm) (Sec. 34,
T.llS, R.20E, Flemington Quad.). OCALA FAUNA (Orange Lake),
Crangonyx grandimanus (JRH), Crangonyx hobbsi (JRH). Private.

ALA-46. ZAMIA SINK (Sec.8, T.IOS, R.18E, Newberry Quad.).
OCALA (Upper Suwannee), cave-associated species. REFERENCES:
Marshall 1947 (fish).

ALA-47. 32-FOOT CAVE (Sec.l8, T.IOS, R.19E, Gainesville West
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM). Private. REFERENCES: Franz and Lee 1982 (Pp).

CITRUS COUNTY

CIT-1. CAVE (6 mi N of Lecanto). OCALA FAUNA (Withlacoochee),
cave-associated species. REFERENCES: McNab 1974 (bats).

CIT-2. BLOWING HOLE CAVE (Sec.21, T.20S, R.19E, Brooksville
NW Quad.). OCALA FAUNA (Withlacoochee), cave-associated species.
Public, Withlacoochee State Forest, Florida Dept. Agriculture, gated
cave. REFERENCES: Hubbell 1936 (crickets); Peck 1970 (terrestrial
arthropods).

CIT-3. DR. DAMES CAVE (or Dr. Doan's Cave) (Sec.30, T.20S,
R.19E, Brooksville NW Quad.). OCALA FAUNA (Withlacoochee),
cave-associated species. Public, Withlacoochee State Forest, Florida
Dept. Agriculture. REFERENCES: Hubbell 1936 (crickets); Peck 1970
(terrestrial arthropods).

CIT-4. HALL'S BAT CAVE (or Rock Pile Cave, also known as
Trail 10 Cave) (Sec.30, T.19S, R.18E, Brooksville NW Quad.). OCALA
FAUNA (Withlacoochee), cave-associated species. REFERENCES: Lee
field notes.

CIT-5. HOMOSASSA SPRINGS. (Sec. 28, T.19S., R.17E.,
Homosassa Quad.). OCALA FAUNA (?), unidentified cave amphipods
and isopods. Public, Homosassa Springs State Park, Florida Dept. Natural
Resources. REFERENCES: Rosenau et al. 1977; Karst Environ-mental
Services (cave map).

Caves and their Faunas in Florida and South Georgia 85

CIT-6. RESTINGHOUSE SIPHON (Sec. 28, T.19S., R.17E,
Homosassa Quad.). OCALA FAUNA, cave-associated species (snails).
Private.

CIT-7. SWEET GUM CAVE (Sec.36, T.20S, R.19E, Nobleton
Quad.). OCALA FAUNA (Withlacoochee), Crangonyx hobbsi
(JRH), Procambarus lucifugus lucifugus-type locality (USNM), Troglo-
cambarus maclanei (USNM), other cave-associated species. Private.
REFERENCES: Faxon 1898 (PU as Pa); Franz 1982 (Pll); Hobbs 1940«
(Pll, type description); 19426 (Pll); Hobbs et al. 1977 (Pll); Hobbs III
1992 (photo of Pll); Hubbard 1901 (terrestrial arthropods); Mohr and
Poulson 1966 (Pll); Warren 1961 (Pll).

COLUMBIA COUNTY

COL-1. BIG GRUNGY SWALLET (Sec.22, T.7S, R.17E, High
Springs Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (USNM), unidentified cave amphipods (TM). Public, O'Leno
State Park, Florida Dept. Natural Resources. REFERENCES: TM (map).

COL-2. BIG ROOM CAVE SINK (Sec.l8, T.7S, R.17E, Fort
White Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (TM), unidentified cave amphipods (TM). Private.

COL-3. BUSSEY SINK (Sec.l8, T.7S, R.17E, Fort White Quad.).
OCALA FAUNA (upper Suwannee), Crangonyx hobbsi (JRH), Pro-
cambarus pallidus (USNM), other cave-associated species. Private.

COL-4. COLUMBIA SPRINGS (or Olustee Creek Spring) (Sec.29,
T.6S, R.18E, Mikesville Quad.). OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (TM). Private. REFERENCES: TM (map).

COL-5. FOSSIL CAVE SINK (or Jebs Hole) (sensitive, T.7S,
R.17E, High Springs SW Quad.). Private. OCALA FAUNA (Upper
Suwannee), Procambarus pallidus (TM), Troglocambarus maclanei
(TM), unidentified cave amphipods (TM). REFERENCES: TM (map).

COL-6. JUG SPRING (or Blue Hole Spring) (Sec.7, T.6S, R.16E,
Hildreth Quad.). OCALA FAUNA (Upper Suwannee), unidentified
cave amphipods and isopods (TM), other cave-associated species.
Public, Ichetucknee River State Park, Florida Dept. Natural Resources.
REFERENCES: Rosenau, et. al. 1977 (spring description); Auffenberg
19576 (fossils, cave map).

COL-7. RIVERBED CAVE (Sec.l9, T.7S, R.17E, High Springs
SW Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi
(Hobbs 1942b), Procambarus pallidus (USNM), other cave-associated
species. Private, railroad right-of-way. REFERENCES: Franz 1982 (Pp);
Hobbs 1940fl (Pp), 19426 (Ch, Pp, cave description); Hobbs et al.
1977 (Pp); Warren 1961 (Ch, Pp).

86 Richard Franz, Judy Bauer and Tom Morris

COL-8. ROBINS NEST SPRING/SIPHON. (Sec.35, T.6S, R.16E,
Fort White Quad.). OCALA FAUNA (Upper Suwannee), unidentified
cave amphipods and crayfishes (TM). Private.

COL-9. ROSE CREEK SWALLET I (or Duckweed I) (Sec. 10,
T.5S, R.16E, Columbia Quad.). OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (USNM), unidentified cave amphipods and
isopods (TM), other cave-associated species. Private. REFERENCES:
TM (map).

COL-10. ROSE CREEK OVERFLOW SWALLET (or Duckweed
II) (Sec. 15, T.5S, R.16E, Columbia Quad.). OCALA FAUNA (Upper
Suwannee), Procambarus pallidus (TM), unidentified cave ampi-
pods and isopods (TM), other cave-associated species. Private.
REFERENCES: TM (map).

COL-11. RUSSELL'S RUB (Seel, T.6S, R.15E, Hildreth Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (TM).
Private.

COL-12. SHILOH CAVE (or Railroad Cave) (Sec.l3, T.7S, R.16E,
Fort White Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (USNM). Private, railroad right-of-way.

COL-13. SIPHON CREEK CAVE (Sec.28, T.7S, R.16E, High
Springs SW Quad.). OCALA FAUNA (Upper Suwannee), cave-associated
species. Public, state waters.

COL-14. TROOP SINK (sensitive, T.5S, R.14E, O'Brien SE Quad.).
OCALA FAUNA (Upper Suwannee), unidentified crayfishes {Procam-
barus lucifugus complex ?) (TM). Private. REFERENCES: TM (map).

COL-15. WHITE SPRINGS (Sec.7, T.2S., R.16E, White Springs
West Quad.). OCALA FAUNA (Upper Suwannee), cave-associated species.
REMARKS: Catfish kill in cave (TM).

DADE COUNTY

DAD-1. WELL, LITTLE BIRD NURSERY AND GARDEN
STORE (Sec.l5, T.54S, R.40E, South Miami Quad.). MIAMI FAUNA,
Crangonyx grandimanus (JRH), Crangonyx hobbsi (JRH), Procam-barus
milleri-typQ locality (USNM), other cave-associated species. Private,
commercial site. REFERENCES: Franz 1982 (Pmi); Franz and Lee
1982 (Pmi); Hobbs 1971 (Pmi-type description); Hobbs et al. 1977
(Pmi); Holsinger 1972 (Ch, Cg).

DAD-2. WELL 4.5 KM NORTHEAST OF HOMESTEAD (Sec.29,
T.56S, R.39E, Goulds Quad.). MIAMI FAUNA, Procambarus milleri
(USNM). Private. REMARKS: Five specimens were taken from a nine
meter deep well in the Biscayne aquifer on 1 June 1992 and 22 August

Caves and their Faunas in Florida and South Georgia 87

1992 by W. F. Loftus and P. Radice (W. F. Loftus and HHH, pers.
comm.).

GILCHRIST COUNTY

GIL-1. DEVIL'S EYE AND EAR SPRINGS (Sec. 34, T.7S,
R.16E, Hildreth Quad.). Ginnie Springs Cave System. OCALA FAUNA
(Upper Suwannee), Crangonyx grandimanus (JRH), Crangonyx
hobbsi (JRH), Procambarus pallidus (USNM), Troglocambarus
maclanei (USNM). Public, state waters, proximity of commercial
recreation area. REFERENCES: Franz and Lee 1982 (Pp); Rosenau et
al. 1977 (spring description).

GIL-2. GINNIE SPRINGS (Sec.34, T.7S, R.16E, High Springs
Quad.). Ginnie Springs Cave System. OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (Pp). Private, commercial recreation area.
REFERENCES: Franz and Lee 1982 (Pp); Rosenau et al. 1977 (spring
description).

GIL-3. HART SPRINGS (Sec. 30, T.9S, R.14E, Wannee Quad.).
OCALA FAUNA (Lower Suwannee), unidentified cave crayfishes and
amphipods (TM). Public, Hart Springs County Park, Gilchrist County.

GIL-4. KELLEY'S SINKS (Sec.34, T.8S, R.14E, Wannee Quad.).
OCALA FAUNA (Lower Suwannee), Procambarus lucifugus X
alachua (RF). Private. REFERENCES: Franz and Lee 1982 (PlXa);
FSS files.

GIL-5. LITTLE DEVIL'S SPRING (Sec.34, T.7S, R.16E, High
Springs SW Quad.). OCALA FAUNA (Upper Suwannee), unidentified
cave amphipods (TM). REFERENCES: WS (map).

GIL-6. OLD WALKER FARM SINKS (Sec.3, T.9S, R.14E, Wannee
Quad.). OCALA FAUNA (Lower Suwannee), Procambarus lucifugus
X alachua (USNM). Private. REFERENCES: Franz 1982 (PlXa); Franz
and Lee 1982 (PlXa).

GIL-7. OTTER SPRINGS (Sec.6, T.IOS, R.14E, Wannee Quad.).
OCALA FAUNA (Lower Suwannee), unidentified cave crayfishes and
amphipods (TM), other cave-associated species (TM). Private, com-
mercial recreation area. REFERENCES: Rosenau et. al. 1977 (spring
description).

GIL-8. ROBERT'S CAVE (or Bells Bat Cave, Bat Hole) (Sec.ll,
T.9S, R.14E, Wannee Quad.). OCALA FAUNA (Lower Suwannee),
Procambarus lucifugus X alachua (Warren 1961), other cave-associated
species. Private. REFERENCES: Franz 1982 (PlXa); Hobbs et al. 1977
(PlXa); Holt 19736 (CI as commensal on Pla); Rice 1957 (bats); Warren
1961 (PlXa).

88 Richard Franz, Judy Bauer and Tom Morris

GIL-9. ROCK BLUFF SPRING (Sec.9, T.8S, R.14E., Hatchbend
Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx sp.? (JRH),
Procambarus pallidas (USNM). Private. REFERENCES: Rosenau et
al. 1977 (spring description).

HAMILTON COUNTY

HAM-1. ADAMS SPRING/SIPHON (Sec. 8, T.IS, R.12E,
Ellaville Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (TM). Private.

HAM-2. CORBET SPRING CAVE (Sec. 10, T.IN., R.llE.,
Octahatchee Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (USNM). Private.

HAM-3. FIRECRACKER CAVE (Sec. 5, T.IN, R.llE.,
Octahatchee Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (USNM), unidentified cave amphipods and isopods, other associated-
cave species. Private. REFERENCES: Pruitt 1991c, 1992 (Pp, cave
description, map, biota).

HAM-4. HYDRANT SPRING (sensitive, T.IS, R.12E, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave cray-
fishes (TM). Public, state waters. REFERENCES: TM (map).

HAM-5. NATURAL BRIDGE SPRING (sensitive, T.IS, R.12E,
Ellaville Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave
crayfishes. Public, Suwannee River State Park, Florida Dept. Natural
Resources. REFERENCES: TM (map).

HAM-6. OVERFLOW SPRING CAVE (Sec.l3, T.IS, R.llE, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), unidentified cave amphipods (TM). Public, Suwannee River
State Park, Florida Dept. Natural Resources.

HAM-7. POTT SPRING (sensitive, T.IN, R.llE, Ellaville Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (TM), un-
identified cave amphipods (TM). Private.

HAM-8. RAVINE INTERMITTENT SPRING (sensitive, T.IS, R.12E,
Ellaville Quad.), unidentified cave crayfishes (TM). Private.

HAM-9. ROSSITER SPRING (sensitive, T.IN, R.llE, Octahatchee
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(TM), unidentified cave amphipods (TM). Private.

HAM-10. SHALLOW SPRING (Sec.lO, T.IN, R.llE, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), unidentified cave amphipods (TM), cave-associated species
(snails). Private.

Caves and their Faunas in Florida and South Georgia 89

HAM-11. UNDERHUNG SINK (Sec.23, T.IN, R.llE, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM). Private.

HERNANDO COUNTY

HER-1. CAVE (unidentified site 23.3 km [14 mi] N of Weekiwachee
Springs). OCALA FAUNA (Withlacoochee), Procambarus lucifugus
lucifugus (USNM). REMARKS: Two specimens (Form I male and female)
collected by Albert Greenburg in 1937 (data with USNM specimens).
Franz 1982 (Pll); Hobbs 1940« (Pll), 1942Z? (Pll); Hobbs et al. 1977
(Pll); Warren 1961 (Pll).

HER-2. DIE POLDER 2 SINK (Sec.5, T.23S, R.18E, Weekiwachee
Springs Quad.). OCALA FAUNA (Gulf Coastal Lowlands), Procam-
barus leitheuseri (USNM). Private, boy scout camp. REFERENCES:
Franz and Hobbs 1983 (Pie).

HER-3. DIE POLDER 3 SINK (Sec.5, T.23S, R.18E, Weekiwachee
Springs Quad.). OCALA FAUNA (Gulf Coastal Lowlands), Procambarus
leitheuseri (USNM), Troglocambarus maclanei (TM), other cave-
associated species. Private, boy scout camp. REFERENCES: Franz and
Hobbs 1983 (Pie).

HER-4. EAGLE'S NEST SINK (or Lost Sink) (Sec.21, T.22S,
R.17E, Weekiwachee Springs Quad.). OCALA FAUNA (Gulf Coastal
Lowlands), Crangonyx grandimanus (JRH), Crangonyx hobbsi (JRH),
Procambarus leitheuseri-typQ locality (USNM), Troglocambarus maclanei
(USNM), other cave-associated species. Private. REFERENCES: Franz
and Hobbs 1983 (Pie-type description).

HER-5. LITTLE SALT SPRINGS (Sec. 29, T.22S, R.17E,
Weekiwachee Springs Quad.). OCALA FAUNA (Gulf Coastal Lowlands),
Procambarus leitheuseri (ATL). Private. REFERENCES: Franz and
Hobbs 1983 (Pie); Rosenau et al. 1977 (spring description).

HER-6. LITTLE SPRINGS (or Twin D's) (Sec.2, T.22S, R.17E,
Weekiwachee Springs Quad.). OCALA FAUNA (Gulf Coastal Lowlands),
Procambarus leitheuseri (ATL). Private, commercial attraction (Weeki-
wachee Springs). REFERENCES: Franz and Hobbs 1983 (Pie); Rosenau
et al. 1977 (spring description), Wetterhall 1965 (hydrology).

HOLMES COUNTY

HOL-1. VORTEX BLUE SPRING (Sec.9, T.4N, R.17W, Prosperity

Quad.). Unstudied cave crayfish (USNM), other cave-associated species.

Private, commercial recreation site. REMARKS: This spring occurs in

the Choctawhatchee drainage, west of the Marianna Lowlands of Jackson

90 Richard Franz, Judy Bauer and Tom Morris

County. At least three other large springs occur in this county (Rosenau
et al. 1977). Recent collections of cave crustaceans in the Vortex system
suggests that this spring area is in need of more explora-tions. REFERENCES:
Helfman 1986 (fish); Rosenau et al. 1977 (spring description).

JACKSON COUNTY

JAC-1. BAT CAVE (unidentified site). APALACHICOLA FAUNA
(Marianna Lowlands), cave-associated species. REMARKS: Hubbell
(1936) indicated the cave was located on the bank of the Chipola River
near Marianna. REFERENCES: Hubbell 1936 (crickets); Peck 1970
(terrestrial arthropods).

JAC-2. BLUE HOLE SPRING (Sec.21, T.5N, R.lOW, Marianna
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cambarus
cf. cryptodytes (USNM). Public, Florida Caverns State Park, Florida
Dept. Natural Resources. REMARKS: Unusual crayfish specimen;
more material is needed from this site in order to determine this popula-
tion's specific identity (HHH).

JAC-3 BLUE SINK (Sec. 2, T.3N, R.llN, Kynesville Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), Haideotriton wallacei
(TM). Private.

JAC-4. CAVE-IN-WOODS (Sec. 26, T.5N, R.llW, Cottondale
East Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cam-
barus cryptodytes (RF), Haideotriton wallacei (RF). Private.
REFERENCES: Franz and Lee 1982 (Cc).

JAC-5. CHINA CAVE (Sec.27, T.5N, R.lOW, Marianna Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), Haideotriton wallacei,
other cave-associated species. Public, Florida Caverns State Park,
Florida Dept. Natural Resources. REFERENCES: Hobbs III 1992 (Hw,
photo).

JAC-6. COFFIN SPRING (Sec.35, T.4N, R.llW, Kynesville Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), unidentified cray-
fishes (TM). REFERENCES: map (TM, WS).

JAC-7. ELLIS CAVE (or Honey Comb Hill Cave) (Sec. 28, T.5N,
R.lOW, Marianna Quad.). APALACHICOLA FAUNA (Marianna Low-
lands), Cambarus cryptodytes (USNM), Haideotriton wallacei (RF),
other cave-associated species. Private. REFERENCES: Brockman and
Bortone 1977 (fish); Franz 1982 (Cc); Hobbs et al. 1977 (Cc).

JAC-8. FLORIDA CAVERNS (commercial cave?) (Sec.27, T.5N,
R.llE, Marianna Quad.). APALACHICOLA FAUNA (Marianna Low-
lands), cave-associated species. Public, Florida Caverns State Park,
Florida Dept. Natural Resources. REFERENCES: Lane 1986 (cave
description); Peck 1970 (terrestrial arthropods); Vandel 1965a (biota).

Caves and their Faunas in Florida and South Georgia 91

JAC-9. GERARD'S CAVE (or Sam Smith Cave) (Sec.23, T.5N,
R.llE, Cottondale East Quad.). APALACHICOLA FAUNA (Marianna
Lowlands), Caecidotea hobhsi (USNM), Cambarus cryptodytes (USNM),
Haideotriton wallacei (UF, USNM, NCSM, MCZ), other cave-associated
species. Private. REFERENCES: Caine 1978 (Cc, ecology); Franz
1982 (Cc); Franz et al. 1971 (snails); Hobbs et al. 1977 (Cc); Lee
1969fl, 19696, 1969c, 1969J (cave-associated species); Pylka and
Warren 1958 (Hw); Warren 1961 (Cah, Cc, Hw).

JAC-10. GEROME'S CAVE (or Bumpnose Cave) (Sec.l8, T.5N,
R.lOW, Cottondale East Quad.). APALACHICOLA FAUNA (Mari-anna
Lowlands). Cambarus cryptodytes (DSL), other cave-associated species.

JAC-11. HOLE-IN-WALL SPRING (Sec.5, T.4N, R.9W, Marianna
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cambarus
cryptodytes (USNM), Haideotriton wallacei (UF), unidentified amphi-
pods (TM). Private. REFERENCES: Exley 1978 (cave description,
map).

JAC-12. JACKSON BLUE SPRING (Sec. 33, T.5N, R.9W,
Marianna Quad.). APALACHICOLA FAUNA (Marianna Lowlands),
Cambarus cryptodytes (USNM), Haideotriton wallacei (JB). Public,
state waters.

JAC-13. JUDGES CAVE (Sec.35, T.5N, R.lOW, Marianna Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), Cambarus crypto-
dytes (Warren 1961), Haideotriton wallacei (Warren 1961). Public,
Judges Cave Bat Preserve, Florida Game and Fresh Water Fish Com-
mission. REFERENCES: Franz 1982 (Cc); Hobbs et al. 1977 (Cc);
Warren 1961 (Cc, Hw).

JAC-14. LIMESTONE CAVE AT BLUE SPRING (unidentified
site). APALACHICOLA FAUNA (Marianna Lowlands), cave-associated
species. REMARKS: Hubbell (1936) listed the site as "...small limestone
caves at Blue Spring."REFERENCES: Hubbell 1936 (crickets); Peck
1970 (terrestrial arthropods).

JAC-15. KRAMER'S CAVE (unidentified cave in Florida Caverns
State Park). APALACHICOLA FAUNA (Marianna Lowlands), cave-
associated species. Public, Florida Caverns State Park, Florida Dept.
Natural Resources. REFERENCES: Franz et al. 1971 (snails).

JAC-16. MILLER'S CAVE (Sec.28, T.5N, R.lOW, Marianna
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cambarus
cryptodytes (RF), Haideotriton wallacei (RF), Pseudosinella pecki
(Christiansen and Bellinger 1980), other cave-associated species.
Public, Florida Caverns State Park, Florida Dept. Natural Resources.
REFERENCES: Christiansen and Bellinger 1980 (Psp); Peck 1970
(terrestrial arthropods); Klimaszewski and Peck 1986 (beetles).

92 Richard Franz, Judy Bauer and Tom Morris

JAC-17. MILTON'S CAVE (Sec. 13, T.5N, R.llW, Cottondale
East Quad.). APALACHICOLA FAUNA (Marianna Lowlands), cave-
associated fauna.

JAC-18. MILTON'S WELL CAVE (Sec. 13, T.5N, RllW,
Cottondale East Quad.). APALACHICOLA FAUNA (Marianna Low-
lands), Cambarus cryptodytes (DSL), Haideotriton wallacei (DSL),
other cave-associated species. Private. REFERENCES: Franz 1982
(Cc); Franz et al. 1971 (snails); Hobbs et al. 1977 (Cc).

JAC-19. MUD CAVE (Sec.3, T.4N, R.IOE, Marianna Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), cave-associated
species. Private. REFERENCES: Rice 1957 (bats).

JAC-20. OLD INDIAN CAVE (Sec.21, T.5N, R.lOW, Marianna
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), cave-
associated species. Public, Florida Caverns State Park, Florida Dept.
Natural Resources. REFERENCES: Lee and Tuttle 1970 (bat protection);
McNab 1974 (bats); Peck 1970 (terrestrial arthropods); Rice 1955^,
19556 (bats).

JAC-21. POOL CAVE (or Pond Cave, Salamander Cave)
(Sec.27, T.5N, R.lOW, Marianna Quad.). APALACHICOLA FAUNA
(Marianna Lowlands), Cambarus cryptodytes (RF), Haideotriton
zwallacei (UF), other cave-associated species. Public, Florida Caverns
State Park, Florida Dept. Natural Resources. REFERENCES: Franz
1982 (Cc); Hobbs et al. 1977 (Cc).

JAC-22. POTTERY CAVE (Sec.27/28, T.5N, R.lOW, Marianna
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cambarus
cryptodytes (Warren 1961), other cave-associated species. Public,
Florida Caverns State Park, Florida Dept. Natural Resources.
REFERENCES: Franz 1982 (Cc); Franz et al. 1971 (snails); Hobbs et
al. 1977 (Cc).

JAC-23. RAY'S CAVE (Sec.22, T.5N, R.llW, Cottondale East
Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cambarus
cryptodytes. Private. REFERENCES: FNAI record.

JAC-24. RIVER CAVE (unidentified cave in Florida Caverns
State Park). APALACHICOLA FAUNA (Marianna Lowlands), cave-
associated species. Public, Florida Caverns State Park, Florida
Dept. Natural Resources. REFERENCES: Peck 1970 (terrestrial
arthropods).

JAC-25. ROCKWELL CAVE (unidentified site). APALACHICOLA
FAUNA (Marianna Lowlands), Cambarus cryptodytes (Franz 1982).
REFERENCES: Franz 1982 (Cc); Hobbs et al. 1977 (Cc).

JAC-26. SODA STRAW CAVE (or Walt's Misery) (Sec.2, T.4N,
R.lOW, Marianna Quad.) APALACHICOLA FAUNA (Marianna Low-

Caves and their Faunas in Florida and South Georgia 93

lands), Cambarus cryptodytes (Warren 1961). Public, Florida Caverns
State Park, Florida Dept. Natural Resources. REFERENCES: Franz
1982 (Cc); Hobbs et al. 1977 (Cc); Warren 1961.

JAC-27. "SPRING CAVE" (unidentified cave in Florida Caverns
State Park). APALACHICOLA FAUNA (Marianna Lowlands),
cave-associated species. Public, Florida Caverns State Park, Florida
Dept. Natural Resources. REFERENCES: Peck 1970 (terrestrial
arthropods).

JAC-28. SWEETWATER SPRING (=Bozell Spring) (Sec.l6, T.7N,
R.lOW, Marianna Quad.). APALACHICOLA FAUNA (Marianna Low-
lands), unidentifed cave crayfishes (TM). Private.

JAC-29. TWIN CAVE (Sec.6, T.4N, R.9W, Marianna Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), Cambarus crypto-
dytes (RF), Haideotriton wallacei (RF), unidentified cave amphipods
and isopods (TM). Private. REFERENCES: Exley 1978 (cave descrip-
tion, map); Franz and Lee 1982 (Cc).

JAC-30. "TWO ENTRANCE CAVE" (unidentified cave in Florida
Caverns State Park, possibly Millers Cave). APALACHICOLA FAUNA
(Marianna Lowlands), cave-associated species. Public, Florida Caverns
State Park, Florida Dept. Natural Resources. REFERENCES: Peck 1970
(terrestrial arthropods).

JAC-31. VETTER'S CAVE (Sec.27, T.5N, R.lOW, Marianna Quad.).
APALACHICOLA FAUNA (Marianna Lowlands), Cambarus cryptodytes
(RF), other cave-associated species. Public, Florida Caverns State Park,
Florida Dept. Natural Resources. REFERENCES: Franz 1982 (Cc); Franz
et al. 1971 (snails); Hobbs et al. 1977 (Cc).

JAC-32. WADDELL'S MILL POND CAVE (Sec.33, T.6N, R.llW,
Sills Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cam-
barus cryptodytes (USNM), other cave-associated species. Private.
REFERENCES: Hobbs et al. 1977 (Cc).

JAC-33. WASHED-OUT CAVE (Sec.23, T.5N, R.llW, Cottondale
East Quad.). APALACHICOLA FAUNA (Marianna Lowlands), Cam-
barus cryptodytes (Warren 1961), Haideotriton wallacei (Warren 1961).
Private. REFERENCES: Franz 1982 (Cc); Hobbs et al. 1977 (Cc); Warren
1961 (Cc).

JAC-34. WELL, 2 mi south of Graceville (Sec.15, T.6N, R.13W,
Graceville Quad.). APALACHICOLA FAUNA (Marianna Lowlands),
Cambarus cryptodytes-typt locality (USNM). Private. REMARKS: well
filled (R. Williams, personal communication, Graceville, Florida,
1983). REFERENCES: Hobbs 1941 (Cc-type description), 19426 (Cc);
Warren 1961 (Cc).

94 Richard Franz, Judy Bauer and Tom Morris

JEFFERSON COUNTY
JEF-1. WACISSA BIG BLUE SPRING (Sec.l2, T.2S., R.3N, Wacissa
Quad.). WOODVILLE FAUNA, Procambarus horsti-type locality (USNM),
unidentified cave amphipods and isopods (TM). Public, state waters.
REFERENCES: Hobbs and Means 1972 (Ph-type description); Hobbs
et al. 1977 (Ph); Franz 1982 (Ph); Franz and Lee 1982 (Ph); Rosenau
et al. 1977 (spring description).

LAFAYETTE COUNTY

LAF-1. ALLENS MILL POND SPRING (Sec.5, T.4S, R.llE, Dowling
Park Quad). OCALA FAUNA (Upper Suwannee), Procam-barus pallidus
(USNM), unidentified cave amphipods and isopods (TM). Public, Suwannee
River Water Management District, recreation area. REFERENCES: Rosenau
et al. 1977 (spring description).

LAF-2. ALLIGATOR RESCUE SPRING (Sec.25, T.4S, R.llE,
Mayo Quad.). OCALA FAUNA (Upper Suwannee). Procambarus pallidus
(TM). Public, state waters. REFERENES: map (TM).

LAF-3. BOBCAT SINK (Sec.21, T.4S, R.llE, Mayo Quad.). Lafayette
Blue Spring System. OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (TM), unidentified cave amphipods (TM). Private.

LAF-4. GREEN SINK (Sec.21, T.4S, R.llE, Mayo Quad.).
Lafayette Blue Spring System. OCALA FAUNA (Upper Suwannee),
unidentified cave crayfishes, other cave-associated species (TM).
Public, Blue Springs County Park, Lafayette County.

LAF-5. KASSERMAN SINK (Seel, T.6S, R.13E, Branford Quad.).
Ruth Spring Cave System. OCALA FAUNA (Upper Suwannee), Procam-
barus pallidus (ATL). Private.

LAF-6. LAFAYETTE BLUE SPRING (Sec.21, T.4S, R.llE, Dowling
Park Quad.). Lafayette Blue Spring Cave System. OCALA FAUNA
(Upper Suwannee), Procambarus pallidus (ATL). Public, Blue Springs
County Park, Lafayette County recreation area. REFERENCES:
Rosenau et al. 1977 (spring description).

LAF-7. MAIN SINK (Seel, T.6S, R.13E, Branford Quad.). Ruth
Spring Cave System. OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (ATL). Private.

LAF-8. OWENS SPRING (Sec.21, T.5S, R.13E, Mayo SE Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (TM).
Private.

LAF-9. PERRY SPRING (Sec.35, T.4S, R.llE, Mayo Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL), unidentified
cave amphipods and isopods (TM). Private. REFERENCES: Rosenau
et al. 1977 (spring description).

Caves and their Faunas in Florida and South Georgia 95

LAF-10. RUTH SPRING (Seel, T.6S, R.13E, Branford Quad.).
Ruth Spring Cave System. OCALA FAUNA (Upper Suwannee), Procam-
barus pallidus (USNM). Private. REFERENCES: Rosenau et al. 1977
(spring description).

LAF-11. TROY SPRING (Sec.34, T.5S, R.13E, O'Brien Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (USNM).
Private. REFERENCES: Franz and Lee 1982 (Pp); Rosenau et al.
1977 (spring description).

LAKE COUNTY

LAK-1. ALEXANDER SPRINGS (Sec.39, T.16S, R.27E, Alexander
Spring Quad.). ST. JOHNS RIVER FAUNA (Lake George), Procam-
barus delicatus-type locality (USNM), unidentified cave isopods (TM).
Public, Ocala National Forest, U.S. Forest Service, recreation area.
REFERENCES: Franz and Lee 1982 (unidentified crayfish); Hobbs
and Franz 1986 (Pd-type description); Relyea et al. 1976 (unidentified
crayfish); Rosenau et al. 1977 (spring description).

LAK-2. EUSTIS (unidentified site). OCALA FAUNA (? area),
Procambarus lucifugus subspecies? (USNM). REFERENCES: Hobbs
1940« (PIX?); Hobbs 1942^ (PIX?).

ti -

LEON COUNTY

LEO-1. BIRD SINK SWALLET (Sec.l7, T.IN, R.3E, Lloyd Quad.).
WOODVILLE FAUNA, unidentified cave crayfishes (possibly Procam-
barus horsti) (TM). Private.

LEO-2. CAVE, 3 mi. north of Woodville (unidentified site, possibly
Gopher Sink). WOODVILLE FAUNA, Procambarus orcinus (USNM).

LEO-3. CLAY SINK (unidentified site, possibly Gopher Sink).
WOODVILLE FAUNA, Procambarus orcinus (USNM) (listed in Warren
[1961] 2iS, Procambarus pallidus). REFERENCES: Warren 1961 (Po).

LEO-4. CULLEY'S CAVE (Sec.l7, T.2S, R.IW, Lake Munson
Quad.). WOODVILLE FAUNA, Procambarus orcinus (USNM). Leon
Sinks Recreation Area, Apalachicola National Forest, U.S. Forest Ser-
vice. REFERENCES: Hobbs and Means 1972 (Po), Hobbs III 1992
(photo of sink).

LEO-5. GOPHER SINK (Sec. 16, T.2S, R.IW, Lake Munson
Quad.). WOODVILLE FAUNA, Procambarus orcinus-type locality
(USNM), unidentified cave amphipods, other cave-associated species.
Private. REFERENCES: Franz 1982 (Po); Hobbs et al. 1977 (Po);
Hobbs and Means 1972 (Po, type description); Holt 1973b (CI as commensal
of Po).

96 Richard Franz, Judy Bauer and Tom Morris

LEO-6. LITTLE DISMAL SINK (Sec.17, T.2S, R.IW, Lake Munsion
Quad.). WOODVILLE FAUNA, Crangonyx grandimanus (JRH), Pro-
cambarus orcinus (USNM), unidentifed cave amphipods and isopods
(TM). Public, Leon Sinks Recreation Area, Appalachicola National Forest,
U.S. Forest Service.

LEO-7. MUNSON SLOUGH BLUE (sensitive, T.2S, R.IW, Lake
Munson Quad.). WOODVILLE FAUNA, unidentified cave crayfishes
(TM). Private. REFERENCES: TM (map).

LEO-8. NATURAL BRIDGE SPRING (Sec.29, T.2S, R.2E, Wood-
ville Quad.). WOODVILLE FAUNA, unidentified cave amphipods and
crayfishes (TM). Private. REFERENCES: Lane 1986 (description).

LEO-9. OSGOOD SINK (Sec.ll, T.2S, R.IE, Woodville Quad.).
WOODVILLE FAUNA, Procambarus orcinus (USNM). Private.
REFERENCES: Franz 1982 (Ph); Hobbs et al. 1977 (Ph); Hobbs and
Means 1972 (Ph).

LEO-10. SULLIVAN'S TUNNEL (or Hole-in-the-Ground) (Sec.l3,
T.2S, R.2W, Hilliardville Quad.). WOODVILLE FAUNA, Crangonyx
hobbsi (JRH), Procambarus orcinus (USNM), unidentified cave isopods
(TM). Public, Apalachicola National Forest, U.S. Forest Service.
REFERENCES: Knab 1991 (cave length).

LEO-11. WELL, 4.5 mi east of Tallahassee (unidentified site.).
WOODVILLE FAUNA, Procambarus horsti (USNM). Private.
REFERENCES: Franz 1982 (Ph); Hobbs et al. 1977 (Ph); Hobbs and
Means 1972 (Ph).

LEVY COUNTY

LEV-1. ARCHER CAVES (Sec. 4, T.12S, R.18E, Bronson NE
Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx grandimanus
(JRH), Procambarus pallidus (USNM), other cave-associated species.
Private. REFERENCES: Franz and Lee 1982 (Pp).

LEV-2. BLUE GROTTO (or Williston Blue Sink) (Sec.2, T.13S,
R.18E, Williston Quad.). OCALA FAUNA (Lower Suwannee), Procambarus
lucifugus X alachua (USNM). Private, commercial recreation area.

LEV-3. DEVIL'S DEN (Sec.26, T.12S, R.18E, Williston Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (USNM).
Private. REFERENCES: Kurten 1966 (fossils); Martin 1974 (fossils);
Pruitt 19916 (cave description, map); Rice 1957 (bats); Webb 1974
(fossils).

LEV-4. FOUR CAVE (Sec.28, T.12S, R.18E, Bronson NE Quad.).
OCALA FAUNA (Upper Suwannee), cave-associated species. Private.
REFERENCES: Pruitt 1990 (cave description, fish).

Caves and their Faunas in Florida and South Georgia 97

LEV-5. FRIEDMAN'S SINK (Sec. 35, T.US, R.13E., Manatee
Springs Quad.). Manatee Springs Cave System. OCALA FAUNA
(Lower Suwannee), Crangonyx hobbsi (JRH), Procambarus lucifugus
X alachua (USNM). Public, Manatee Springs State Park, Florida
Dept. Natural Resources. REFERENCES: Exley 1984 (description,
map, photo).

LEV-6. GUNPOWDER CAVE (Sec. 26, T.12S, R.14E, Williston
Quad.). OCALA FAUNA (Upper Suwannee), Troglocambarus
maclanei (USNM), unidentified cave crayfishes {Procambarus
lucifugus complex?). Private.

LEV-7. HALF MOON CAVE (Sec. 29, T.12S, R.18E, Williston
Quad.). OCALA FAUNA, cave-associated species. Private. REFER-
ENCES: Marshall 1947 (fish).

LEV-8. MANATEE SPRINGS (Sec.26, T.llS, R.13E, Manatee
Springs Quad.). Manatee Springs Cave System. OCALA FAUNA (Lower
Suwannee). Crangonyx hobbsi (JRH), Procambarus lucifugus X alachua
(USNM), Troglocambarus maclanei (USNM). Public, Manatee Springs
State Park, Florida Dept. Natural Resources. REFERENCES: Exley
1984 (exploration, map, photos); Franz 1982 (PlXa); Franz and Lee
1982 (PlXa); Knab 1991 (cave length); Rosenau et al. 1977 (spring
description).

LEV-9. OCTOPUS CAVE (Sec.4, T.12S, R.18E, Bronson NE
Quad.). OCALA FAUNA (Upper Suwannee), associated cave fauna.
Private. REFERENCES: Pruitt \991a (description, map, fauna).

LEV-10. PEANUT CAVE (Sec.24, T.13S, R.18E, Morristown
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus lucifugus
alachua (USNM). Private.

LEV-11. WELL, CHIEFLAND (unidentified site). OCALA
FAUNA (Lower Suwannee), Crangonyx grandimanus (JRH), Crangonyx
hobbsi (JRH). Private. REFERENCES: Franz 1982 (Ch).

MADISON COUNTY

MAD-1. BASELINE CAVE (Sec.33, T.IS, R.llE, Falmouth
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), unidentified cave amphipods and isopods (TM). Private.
REFERENCES: map (TM, BP).

MAD-2. MADISON BLUE SPRING (Sec. 17, T.IN, R.llE,
Ellaville Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx
grandimanus (JRH), Crangonyx hobbsi (JRH), Procambarus pallidus
(ATL), other cave-associated species. Private. REFERENCES: Knab
1991 (cave length); Martin and Harris 1993 (mineralogy); Rosenau et
al. 1977 (spring description).

98 Richard Franz, Judy Bauer and Tom Morris

MAD-3. M2 BLUE SPRING (Sec. 32, T.2N, R.llE, Octahatchee
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), other cave-associated species. Public, Suwannee River Water
Management District. REFERENCES: Pruitt 1991(i, 1992 (cave descrip-
tion, map).

MAD-4. SUWANNACOOCHEE SPRING (Sec.24, T.IS, R.llE.,
Ellaville Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (USNM). Public, Suwannee River State Park, Florida Dept.
Natural Resources. REFERENCES: Franz 1982 (Pp); Hobbs et al.
1977 (Pp); Rosenau et al. 1977 (spring description).

MAD-5. THUNDERHOLE SINK (Sec. 10, T.IS, R.llE, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Remasellus parvus
(USNM), Procambarus pallidus (USNM), unidentified cave amphipods
(TM), other cave-associated species. Private. REFERENCES: Franz
and Lee 1982 (Pp).

MARION COUNTY

MAR-1. BELLEVIEW CAVE (unidentified site, possibly Ocala
Caverns). OCALA FAUNA (Marion), cave-associated species. REFER-
ENCES: Peck 1970 (terrestrial arthropods).

MAR-2. BRIAR CAVE (Sec.35, T.15S, R.21E, Ocala West Quad.).
OCALA FAUNA (Marion), Procambarus lucifugus X alachua (USNM).
Private. REFERENCES: Johnson 1990; A. Krause 19906; M. Krause
1990 (cave description, geology).

MAR-3. CHERT CAVE (Sec.lO, T.16S, R.22E, Belleview Quad.).
OCALA FAUNA (Marion), Crangonyx hobbsi (JRH), Procambarus
lucifugus X alachua (USNM), Troglocambarus maclanei (USNM).
Private.

MAR-4. COON CAVE (Sec. 30, T.14S, R.22E, Anthony Quad.).
OCALA FAUNA (Marion), Procambarus lucifugus X alachua (USNM).
Private.

MAR-5. EICHELBERGER CAVE (Sec.2, T.17S, R.22E, Belleview
Quad.). OCALA FAUNA (Marion), Procambarus lucifugus X alachua
(USNM), Procambarus pallidus"} (USNM). Private. REMARKS: Franz
and Lee (1982) questioned the validity the Procambarus pallidus
record. Unfortunately this cave was destroyed by limestone mining
activities. REFERENCES: Auffenberg 1958, 1963 (fossils); Brodkorb
1956 (fossils); Franz 1982 (PlXa, Pp); Franz and Lee 1982 (PlXa,
Pp); Hobbs et al. 1977 (PlXa, Pp).

MAR-6. FROG CRAWL CAVE (Sec. 15, T.14S, R.20E, Fairfield
Quad.). OCALA FAUNA (Marion), unidentified cave crayfishes (FSS
files). Private.

Caves and their Faunas in Florida and South Georgia 99

MAR-7. HELL HOLE (Sec. 6, T.14S, R.21E, Reddick Quad.).
OCALA FAUNA (Orange Lake), Crangonyx hobbsi (JRH), Procam-
harus franzi (USNM). Private. REFERENCES: Franz and Lee 1982
(Pf); Krause 1991 (cave description, map).

MAR-8. HOLLOWED GROUND CAVE. (Sec. 36, T.15S, R.20E,
Cotton Plant Quad.). OCALA FAUNA (Marion), Caecidotea hobbsH
(J. Lewis, personal communication). Private.

MAR-9. INDIAN CAVE (or Last Resort Cave) (Sec. 36, T.15S,
R.20E, Cotton Plant Quad.). OCALA FAUNA (Marion), Crangonyx
grandimanus-type locality (NMC), Crangonyx hobbsi (USNM), Pro-
cambarus lucifugus X alachua (USNM), Troglocambarus maclanei
(USNM). Private. REFERENCES: Bousfield 1963 (Cg, type descrip-
tion); Cooper l96Sa (PlXa); Franz 1982 (Cg, PlXa, Tm); Hobbs
19426 (PlXa); Hobbs et al. 1977 (PlXa); Warren 1961 (PlXa).

MAR-10. JENNING'S CAVE (or Confederate Cave) (Sec.26, T.15S,
R.19E, Romeo Quad.). OCALA FAUNA (Marion), cave-associated
species. Private. REFERENCES: Peck 1970 (terrestrial arthropods).

MAR-11. MEFFORD CAVE (Sec.l5, T.13S, R.21E, Reddick
Quad.). OCALA FAUNA (Marion), cave-associated species. Private.
REFERENCES: Auffenberg 1951a, 1958, 1963 (fossils); Gertsch 1984
(spiders); McNab 1974 (bats); Peck 1970 (terrestrial arthropods).

MAR-12. NICKELBERGER CAVE (Sec.2, T.17S, R.22E, Belleview
Quad.). OCALA FAUNA (Marion), unidentified cave crayfishes, other
cave-associated species (PS, BP, FSS files). Private.

MAR-13. OCALA CAVERNS (Sec.23, T.16S, R.22E, Belleview
Quad.). OCALA FAUNA (Marion), Procambarus lucifugus X alachua
(USNM). Private, previously commercialized. REFERENCES: Franz
and Lee 1982 (PlXa).

MAR-14. ORANGE LAKE CAVE (Sec.34, T.12S, R.21E, Mcintosh
Quad.). OCALA FAUNA (Orange Lake), Uncinocythere lucifuga (AN),
Crangonyx hobbsi (JRH), Procambarus franzi-typQ locality (USNM),
Troglocambarus maclanei (USNM). Private. REFERENCES: Dickson
and Franz 1980 (Pf, gill respiration); Davis and Rand 1982 (lime-
encrusting algae); Franz 1982 (Ch, Pf, Tm); Franz and Lee 1982 (Tm);
Hobbs and Lee 1976 (Pf, type description).

MAR-15. ORANGE LAKE QUARRY SOLUTION PITS (or
Quarry Crevice Caves) (Sec.34, T.12S, R.21E, Mcintosh Quad.). OCALA
FAUNA (Orange Lake), Procambarus franzi (DSL). Private.

MAR-16. REDDING CATACOMBS (Sec.20, T.16S, R.22E, Shady
Quad.). OCALA FAUNA (Marion), Procambarus lucifugus X alachua
(RF). Private. REFERENCES: Franz and Lee 1982 (PlXa).

100 Richard Franz, Judy Bauer and Tom Morris

MAR-17. RAINBOW ACRES CAVE (unidentified site, possibly
Jennings Cave). OCALA FAUNA (Marion), Caecidotea hobbsi (USNM).

MAR-18. ROOSEVELT CAVE (or Tillman's Cave) (Sec.32, T.15S,
R.22E, Ocala East Quad.). OCALA FAUNA (Marion), Caecidotea hobbsi
(USNM), Crangonyx hobbsi (Steeves 1964), Procambarus
lucifugus X alachua (USNM). Private. REFERENCES: Franz 1982
(Ch, CaH, PlXa); Hobbs et al. 1977 (PlXa); Steeves 1964 (Cah); Warren
1961 (Cah, PlXa).

MAR-19. SILVER GLEN SPRINGS (Sec.25, T.14S, R.26E, Juniper
Springs Quad.). ST. JOHNS RIVER FAUNA (Lake George), Pro-
cambarus attiguus-type locality (USNM), unidentified cave amphipods
(TM), other cave-associated species. Public, Silver Glen Springs
Recreation Area, U.S. Forest Service. REFERENCES: Hobbs and Franz
1992 (Pat); Rosenau et al. 1977 (spring description).

MAR-20. SILVER SPRINGS (Sec.6, T.15S, R.23E, Ocala East
Quad.). ST. JOHNS RIVER FAUNA (Oklawaha), Procambarus lucifugus
X alachua (USNM), unidentified cave amphipods and isopods (TM),
other cave-associated species. Private, commercial recreation area.
REFERENCES: Rosenau et al. 1977 (spring description).

MAR-21. STEEPLE CAVE (Sec.8, T.16S, R.22E, Shady Quad.).
OCALA FAUNA (Marion), Procambarus lucifugus X alachua (Franz
1982). Private. REFERENCES: Franz 1982 (PlXa); Hobbs et al. 1977
(PlXa).

MAR-22. SUNDAY SINK (Sec.8, T.16S, R.22E, Shady Quad.).
OCALA FAUNA (Marion), Crangonyx hobbsi (JRH), Procambarus
lucifugus X alachua (USNM), Troglocambarus maclanei (USNM), other
cave-associated species. Private. REFERENCES: Franz 1982
(PlXa, Tm); Franz and Lee 1982 (Tm); Hobbs et al. 1977 (PlXa, Tm).

MAR-23. TRADE WINDS FARM CAVE (unidentified site). OCALA
FAUNA (Orange Lake), Procambarus franzi (USNM), Troglocambarus
maclanei (USNM). Private.

MAR-24. VILLA HEIGHTS CAVE (unidentified site). OCALA
FAUNA (Marion), cave-associated species. REMARKS: Hubbell (1936)
noted that the cave was located 51.6 km (31 miles) south of Gainesville
on State Highway 2. REFERENCES: Hubbell 1936 (crickets); Peck
1970 (terrestrial arthropods).

MAR-25. WALDO CAVE (Sec.35, T.15S, R.21E, Ocala West
Quad.). OCALA FAUNA (Marion), Procambarus lucifugus X alachua
(Hobbs 1942), other cave-associated species. Private. REFERENCES:
Franz 1982 (PlXa); Hobbs 1942/? (PlXa); Hobbs et al. 1977 (PlXa);
Peck 1970 (terrestrial arthropods); Warren 1961 (PlXa).

Caves and their Faunas in Florida and South Georgia 101

MAR-26. WELL (2 mi NE of Anthony) (Sec.9?, T.14S, R.22E,
Anthony Quad.). OCALA FAUNA (Orange Lake?), Crangonyx grandi-
manus (JRH). Private.

MAR-27. WOODS CAVE (Sec.l3, T.16S, R.21E, Shady Quad.).
OCALA FAUNA (Marion), unidentified cave crayfishes. Private.
REFERENCES: FSS files.

MAR-28. ZUBER SINK (Sec.9, T.14S, R.20E, Fairfield Quad.).
OCALA FAUNA (Orange Lake?), unidentified cave crayfishes (JB).
Private.

ORANGE COUNTY

ORA-1. APOPKA BLUE HOLE (Sec.l6, T.20S, R.28E, Sorrento
Quad.). ST. JOHNS RIVER FAUNA (Wekiva), Procambarus acherontis
(USNM), Troglocambarus sp. (USNM). Private, vulnerable to urban
development. REFERENCES: Hobbs III 1992 (Tsp, photo).

ORA-2. ROCK SPRINGS Sec.l5, T.20S, R.28E, Sorrento Quad.).
ST. JOHNS RIVER FAUNA (Wekiva), Caecidotea sp.2 (USNM). Public,
Rock Springs County Park. REFERENCES: Auffenberg 1963 (fossils);
Rosenau et al. 1977 (spring description); Webb 1974 (fossils).

ORA-3. WEKIWA SPRINGS (Sec.36, T.20S, R.28E, Forest City
Quad.). ST. JOHNS RIVER FAUNA (Wekiva), Procambarus acherontis
(USNM). Public, Wekiva Springs State Park, Florida Dept. Natural
Resources. REMARKS: Cooper (1965a) noted that "white crayfish had
been seen around 1890." The USNM specimen was collected in moss
on the outside of one of the cracks in the main spring in about 1.5
meters water depth by David A. Sukkert on 29 September 1990 (Rosi
Mulholland, personal communication, Florida Park Service).
REFERENCES: Cooper 1965^ (description); Rosenau et al. 1977 (spring
description).

ORA-4. WELL, LONG LAKE (Sec.36, T.21S, R.28E, Orlando
West Quad.). ST. JOHNS RIVER FAUNA (Wekiva), Procambarus
acherontis (Franz and Lee 1982). Public, county water well.
REFERENCES: Franz 1982 (Pa); Franz and Lee 1982 (Pa).

PASCO COUNTY

PAS-1. ARCH SINK (or Arch-Way Sink) (Sec.2, T.24S, R.17E,
Port Richey NE Quad.). OCALA FAUNA (Gulf Coastal Lowlands),
Procambarus leitheuseri (USNM). Private. REFERENCES: Franz and
Hobbs 1983 (Pie).

PAS-2. BLACK HOLE (Sec.l4, T.24S, R.16E, Aripeka Quad.).
OCALA FAUNA (Gulf Coastal Lowlands), Procambarus leitheuseri

102 Richard Franz, Judy Bauer and Tom Morris

(USNM), other cave-associated species. Private. REFERENCES:
Franz and Hobbs 1983 (Pie).

PAS-3. NEXUS SINK (Sec.3, T.25S, R.16E, Port Richey Quad.).
OCALA FAUNA (Gulf Coastal Lowlands), Crangonyx grandimanus
(JRH), Crangonyx hobbsi (JRH), Procamharus leitheuseri (USNM).
Private. REFERENCES: Franz and Hobbs 1983 (Pie).

PAS-4. WELL, LACOOCHEE (unidentified site). OCALA
FAUNA (Gulf Coastal Lowlands), Crangonyx hobbsi (JRH).

PINELLAS COUNTY
PIN-1. KNIGHT'S SINK (Sec.l9, T.27S, R.16E, Elfers Quad.)
OCALA FAUNA (Gulf Coastal Lowlands), unidentified cave crayfish
(P. Heinerth, personal communication, 1984). Public, Anderson County
Park, next to Tarpon Sink on west side of Lake Tarpon. REFERENCES:
Wetterhall 1965 (hydrology).

PUTNAM COUNTY
PUT-1. DEVIL'S SINK (Sec.l3, T.IOS, R.23E, Interlachen Quad.).
ST.JOHNS RIVER FAUNA (Oklawaha), Uncinocythere ambophora
(Walton and Hobbs 1959), Procambarus morrisi-iype locality (USNM),
unidentified amphipods. Private, vulnerable to groundwater pollution
from unauthorized dumping. REFERENCES: Hobbs and Franz 1990
(Pm, type description, Ua as commensal of Pm).

SEMINOLE COUNTY

SEM-1. PALM SPRING (Sec.2, T.21S, R.29E, Forrest City Quad.).
ST. JOHNS RIVER FAUNA (Wekiva), Uncinocythere ambophora-
type locality (USNM), Procambarus acherontis (USNM), other cave-
associated species. Private, urban development. REMARKS: The Palm
Springs basin (surface drainage area, 1.77 square miles) consists of
Lake Marion, Eleven Hole Pond, and several small unconnected sinks
(Anderson and Hughes 1975). The drainage area includes a golf course
which is the probable location of the type locality of Procambarus
acherontis. REFERENCES: Anderson and Hughes 1975 (hydrology);
Cooper 1965fl (spring description. Pa collection information); Franz
1982 (Pa); Hobbs 1940« (Pa redescription); Hobbs 1942ft (Pa account);
Hobbs et al. 1977 (Pa); Rosenau et al. 1977 (spring description);
Walton and Hobbs 1959 (Ua, type locality); Warren 1961 (Pa).

SEM-2. WELL, ALTAMONTE SPRINGS (Sec. 13, T.21S, R.29E,
Casselberry Quad.). ST. JOHN RIVER FAUNA (Wekiva), Procambarus
acherontis (RF). Private, urban. REFERENCES: Franz 1982 (Pa); Franz
and Lee 1982 (Pa).

Caves and their Faunas in Florida and South Georgia 103

SEM-3. WELL, LAKE BRANTLEY (Sec.3, T.21S, R.29E, Forrest
City Quad.). ST. JOHNS RIVER FAUNA (Wekiva), Procambarus
acherontis-iype locality (ZIAS). Private, urban. REMARKS: The
acherontis specimens described by Einar Lonnberg were taken from a
hand dug well on the farm of A. E. Sjoblom near Lake Brantley in
1893 (Lonnberg 1894). From land records in the Orange County Court
House, we found that A. E. Sjoblom paid taxes on the following
tracts in 1894. The land descriptions as found in the tax records are
as follows: Plot 1. "Begins 573 feet north, 712 feet east of the SW
corner of NW 1/4 (of Section 3), runs east 300 feet, north578 feet,
west 300 feet, and south 578 feet." Plot 2. "Begins 4.5 chains west of
SE corner of SW 1/4, of the NW 1/4, north 4.13 chains, west 1.21
chains, south 4.13 chains." (chain= 66 feet). This site is now located
on a portion of the Sabal Point Golf Course. The owners and grounds
people at the golf course knew of no open wells on the property, and
we presume the well has been filled. RFERENCES: Faxon 1898; Franz
1982 (Pa); Hobbs 1940« (Pa, description), 1942Z? (Pa); Lonnberg 1894,
1895 (Pa, type description); Warren 1961 (Pa).

SUWANNEE COUNTY

SUW-1. ANDERSON SPRING (Sec.35, T.IS, R.llE, Falmouth
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave cray-
fishes and amphipods (TM). Public, Florida state waters.

SUW-2. AZURE BLUE SINK (or Collins Farm Sink) (Sec.9, T.6S,
R.15E, Hildreth Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
erythrops (USNM), Troglocambarus maclanei (USNM), unidentified
cave amphipods (TM), other cave-associated species. Private.
REFERENCES: Franz and Lee 1982 (Pe).

SUW-3. BLUE SINK (Sec.lO, T.2S, R.15E, White Springs West
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave amphi-
pods (TM). Private.

SUW-4. BONNETT SPRING (Sec.20, T.4S, R.12E, Mayo Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL),
unidentified cave amphipods (TM). Private. REFERENCES: Rosenau
et al. 1977 (spring description).

SUW-5. BUFO SINK (Sec.24, T.6S, R.14E, Branford Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus erythrops (B.
Sutton). Private. REMARKS: One of the sites mentioned by Releya
and Sutton in the description of Procambarus erythrops (B. Sutton,
personal communication, Gainesville, Florida). REFERENCES: Franz
1982 (Pe); Hobbs et al. 1977 (Pe); Relyea and Sutton 1975 (Pe).

104 Richard Franz, Judy Bauer and Tom Morris

SUW-6. CHALLENGE SINK (Sec.20, T.4S, R.12E, Bowling Park
Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi (JRH),
Procambarus pallidus (ATL). Public, Peacock Springs State Park, Florida
Dept. Natural Resources.

SUW-7. CHARLES SPRING (Sec.4, T.4S, R.llE, Dowling Park
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus,
unidentified cave amphipods (TM). Public, Suwannee County park.

SUW-8. CISTEEN SINK (Sec.21, T.4S, R.llE, Dowling Park
Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx grandimanus
(JRH), Crangonyx hobbsi (JRH), Procambarus pallidus (USNM).
Public, Peacock Springs State Park, Florida Dept. Natural Resources.

SUW-9. COW SPRING CAVE (Sec. 28, T.4S, R.12E, Dowling
Park Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), unidentified cave amphipods (TM). Private.

SUW-10. CRAZY HORSE SINK (Sec.21, T.4S, R.12E, Mayo
Quad.). Mirkwood Sink Cave System. OCALA FAUNA (Upper Suwan-
nee), Procambarus pallidus (USNM), unidentified cave amphipods (TM).
Private.

SUW-11. DEBRIS CONE SPRING (Sec.9, T.4S, R.llE, Dowling
Park Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave
crayfishes (TM). Private. REFERENCES: map (TM, J. Brown).

SUW-12. DEVIL'S HEAD AND HORNS (Sec.l2, T.5S, R.13E,
O'Brien SE Quad.). OCALA FAUNA (Upper Suwannee), cave-associated
species. Private. REMARKS: Reportedly filled in (Julie Hovis, personal
communication, Florida Game and Fresh Water Fish Commission, 1991).
REFERENCES: Rice 1957 (bats).

SUW-13. DOUBLE SINK (Sec.l6, T.5S, R.14E, O'Brien Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL). Private.

SUW-14. EDWARDS SPRING (Sec.24, T.IS, R.llE, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(USNM), unidentified amphipods and isopods (TM). Private.
REFERENCES: S. Exley (map)

SUW-15. FALMOUTH SPRING/SIPHON (or Cathedral Cave) (Sec.32,
T.IS, R.12E, Falmouth Quad.). Falmouth Cave System. OCALA FAUNA
(Upper Suwannee), Procambarus pallidus (ATL), unidentified amphipods
(TM). Private. REFERENCES: Rosenau et al. 1977 (spring description),
S. Exley (map).

SUW-16. GHOUL SINK (Sec.32, T.IS, R.12E, Falmouth Quad.).
Falmouth Spring Cave System. OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (USNM). Private.

SUW-17. HILDRETH CAVE (Sec.l6, T.6S, R.15E, Hildreth Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus erythrops (USNM).
Private. REFERENCES: Relyea and Sutton 1975 (Pe).

Caves and their Faunas in Florida and South Georgia 105

SUW-18. IRVINE SLOUGH SPRING (Lauraville Spring) (Sec.24,
T.4S, R.llE, Mayo Quad.). OCALA FAUNA (Upper Suwannee), Pro-
cambarus pallidas (ATL), unidentified cave amphipods and isopods.
Private. REFERENCES: Rosenau et al. 1977 (spring description), TM
(map).

SUW-19. LINEATER SPRING (Sec. 7, T.IS, R.12E, Ellaville
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidas
(TM), unidentified cave amphipods (TM). Private.

SUW-20. LITTLE RIVER SPRING (Seel, T.6S, R.13E,
Branford Quad.). OCALA FAUNA (Upper Suwannee), Procambaras
pallidas (USNM), unidentified cave amphipods and isopods (TM), other
cave-associated species. Public, Little River Springs County Park.
REFERENCES: Franz and Lee 1982 (Pp); Rosenau et al. 1977 (spring
description); Streever 1993 (invertebrates).

SUW-21. MIRKWOOD SINK (Sec.21, T.4S, R.llE, Mayo Quad.).
OCALA FAUNA (Upper Suwannee), Procambaras pallidas (USNM).
Private.

SUW-22. MULKY ROAD SINK (Sec.29, T.6S, R.15E, Hildreth
Quad.). OCALA FAUNA (Upper Suwannee), cave-associated species
(RF). Private.

SUW-23. O'HARA CAVE (Sec.24, T.IS, R.llE, Ellaville Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave crayfishes (PS,
1977). Private.

SUW-24. OLSEN SINK (Sec.20, T.4S, R.llE, Dowling Park
Quad.). Peacock Slough Cave System. OCALA FAUNA (Upper Suwannee),
Procambaras pallidas (JB). Public, Peacock Springs State Park,
Florida Dept. Natural Resources.

SUW-25. ORANGE GROVE SINK (Sec.20, T.4S, R.12E, Dowling
Park Quad.). Peacock Slough Cave System. OCALA FAUNA (Upper
Suwannee), Crangonyx grandimanus (JRH), Crangonyx hobbsi (JRH),
Procambaras pallidas (USNM), other cave-associated species. Public,
Peacock Springs State Park, Florida Dept. Natural Resources.
REFERENCES: DeLoach and Arteaga 1972 (cave description); Rosenau
et al. 1977 (spring description).

SUW-26. OSTEEN SINK (unidentified site). OCALA FAUNA
(Upper Suwannee), Procambaras pallidas (USNM).

SUW-27. PEACOCK SLOUGH (unnamed cave in Peacock Slough
Cave system) (Sec.20, T.4S, R.12E, Dowling Park Quad.). OCALA
FAUNA (Upper Suwannee), Procambaras pallidas (USNM). Public,
Peacock Springs State Park, Florida Dept. Natural Resources.
REFERENCES: Hobbs 1971 (Pp); Hobbs et al. 1977 (Pp); Martin and
Harris 1993 (mineralogy).

106 Richard Franz, Judy Bauer and Tom Morris

SUW-28. PEACOCK SPRING CAVE (Sec.20, T.4S, R.12E,
Dowling Park Quad.). Peacock Slough Cave System. OCALA FAUNA
(Upper Suwannee), Remasellus parvus (USNM), Crangonyx grandi-
manus (JRH), Crangonyx hobbsi (JRH), Procambarus pallidus (USNM),
other cave-associated species. Public, Peacock Springs State Park, Florida
Dept. Natural Resources. REFERENCES: Bowman and Sket 1985 (Rp);
Exley and DeLoach 1981 (cave description); Exley and Fisk 1978 (cave
description); Franz 1982 (Pp); Fisk and Exley 1977 (description); Rosenau
et al. 1977 (spring description), Streever 19926 (crayfish kill).

SUW-29. PEACOCK SINK 3 (Sec.20, T.4S, R.12E, Dowling Park
Quad.). Peacock Slough Cave System. OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (JB). Public, Peacock Springs State Park, Florida
Dept. Natural Resources.

SUW-30. POT HOLE SINK (Sec.20, T.4S, R.12E, Mayo Quad.).
Peacock Slough Cave System. OCALA FAUNA (Upper Suwannee),
Procambarus pallidus (JB). Public, Peacock Springs State Park,
Florida Dept. Natural Resources. REFERENCES: Rosenau et al. 1977
(spring description).

SUW-31. QUARRY SINK (Sec.23, T.6S, R.14E, Branford Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus erythrops (Franz
1982). Private. REMARKS: According to B. Sutton (personal communi-
cation), this was one of the unidentified localities where Relyea and
Sutton found crayfishes during their surveys in preparation for the
description oi Procambarus erythrops. REFERENCES: Franz 1982 (Pe);
Relyea and Sutton 1975 (Pe).

SUW-32. REGISTER SINK (Sec.l7, T.5S, R.14E, O'Brien Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL). Private.

SUW-33. SANDBAG SPRING (Sec. 34, T.4S, R.12E, Mayo SE
Quad.). OCALA FAUNA (Upper Suwannee), Procambarus pallidus
(TM), unidentified cave amphipods (TM). Private.

SUW-34. SIM'S SINK (also Simm's Sink) (Sec.24, T.6S, R.14E,
Branford Quad.). OCALA FAUNA (Upper Suwannee), Crangonyx hobbsi
(JRH), Procambarus erythrops-iype locality (USNM), Troglocambarus
maclanei (USNM), unidentified cave isopods (TM). Private, The Nature
Conservancy, cave crayfish preserve. REFERENCES: Franz 1982 (Ch,
Pe, Tm); Franz and Lee 1982 (Pe); Hobbs et al. 1977 (Pe, Tm); Holt
19736 (CI as commensal on Pe); Mellon 1977 (Pe, ocular response);
Mellon and Lnenicka 1980 (Pe, ocular response); Relyea and Sutton
1975 (Pe, type description).

SUW-35. SMITH SINK (Sec. 20, T.5S, T.14E, O'Brien Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL). Private.

Caves and their Faunas in Florida and South Georgia 107

SUW-36. STICK SINK (Sec. 34, T.4S, R.14E, O'Brien Quad.).
OCALA FAUNA (Upper Suwannee), Procambarus pallidus (ATL). Private.

SUW-37. SUWANNEE BLUE SPRING (Seel, T.4S, R.llE, Dowling
Park Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave
amphipods (TM). Private.

SUW-38. TELFORD SPRING (or Tilford Spring) (Sec.25, T.4S,
R.llE, Mayo Quad.). Luraville-Telford Spring System. OCALA FAUNA
(Upper Suwannee), Procambarus pallidus (USNM). Private.
REFERENCES: Knab 1991 (cave length); Martin and Harris 1993
(mineralogy); Rosenau et al. 1977 (spring description), Streever et al.
1993 (sediment deposition).

SUW-39. TEN MILE HOLLOW CAVE (Sec.35, T.4S, R.12E,
Mayo SE Quad.). OCALA FAUNA (Upper Suwannee), Procambarus
pallidus (TM). Private.

SUW-40. WATER HOLE 3 SINK (Sec.20, T.4S, R.12E, Mayo
Quad.). Peacock Slough Cave System. OCALA FAUNA (Upper Suwan-
nee), Procambarus pallidus (JB). Public, Peacock Springs State Park,
Florida Dept. Natural Resources.

SUW-41. WATERING HOLE SPRING (Sec.9, T.4S, R.llE, Dowling
Park Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave
crayfishes and amphipods (TM). Private.

SUW-42. WINGATE WELL (Sec.23, T.5S, R.13E, Mayo SE Quad.).
OCALA FAUNA (Upper Suwannee), unidentified cave cray-fishes, amphipods,
and isopods (TM). Private.

SUW-43. YUCK SPRING (Sec. 22, T.4S, R.llE, Dowling Park
Quad.). OCALA FAUNA (Upper Suwannee), unidentified cave cray-
fishes, amphipods, and isopods (TM). Private.

WAKULLA COUNTY

WAK-1. EMERALD SINK (Sec.20, T.2S, R.IW, Lake Munson
Quad.). Emerald Sink Cave System. WOODVILLE FAUNA, Crangonyx
hobbsi (JRH), Crangonyx grandimanus (JRH), Procambarus orcinus
(USNM). Public, Apalachicola National Forest, U.S. Forest Service.
REFERENCES: Exley and Goodman 1981 (cave description, map).

WAK-2. INDIAN SPRINGS (Sec.3, T.3S, R.IW, Crawfordville
Quad.). WOODVILLE FAUNA, Procambarus orcinus (USNM). Private.
REFERENCES: Rosenau et al. 1977 (spring description); Exley and
Goodman 1981 (cave description).

WAK-3. MCBRIDE SLOUGH SPRING (Sec. 7, T.3S,. R.IE,
Crawfordville Quad.). Wakulla Springs Cave System. WOODVILLE
FAUNA, Crangonyx grandimanus (JRH), Crangonyx hobbsi (JRH),

108 Richard Franz, Judy Bauer and Tom Morris

Procambarus orcinus (USNM), unidentified cave amphipods and
isopods (Morris 1989). Private. REFERENCES: Morris 1989 (Po).

WAK-4. RIVER SINKS (Sec.28, T.2S, R.IW, Lake Munson
Quad.). Emerald Sink Cave System. WOODVILLE FAUNA, Crangonyx
sp. (JRH), Crangonyx hobbsi (JRH), Procambarus orcinus (Caine
1978). Private. REFERENCES: Caine 1978 (Po, ecology); Rosenau et
al. 1977 (spring description); Exley and Goodman 1981 (map).

WAK-5. SALLY WARD SPRING (or Numero Uno Spring) (Sec. 11,
T.3S, R.IW, Crawfordville Quad.). Wakulla Springs Cave System.
WOODVILLE FAUNA, Crangonyx sp. (JRH), Crangonyx hobbsi (JRH),
Procambarus orcinus (Morris 1989). Public, Wakulla Springs State
Park, Florida Dept. Natural Resources. REFERENCES: Morris 1989
(Po, photo); Wilson and Sparks 1992 (hydrology).

WAK-6. SHEPARD BLUE SPRING (Land Grant Sec. 99, Hartsfield
Survey, Spring Creek Quad.). WOODVILLE FAUNA, Crangonyx
grandimanus (JRH), Crangonyx hobbsi (JRH), Procambarus horsti
(USNM). Public, state waters. REFERENCES: Rosenau et al. 1977
(spring description).

WAK-7. SPLIT SINK (Sec.20, T.2S, R.IW, Lake Munson Quad.).
Emerald Sink Cave System. WOODVILLE FAUNA, Remasellus parvus
(USNM). Public, Apalachicola National Forest, U.S. Forest Service.
REFERENCES: Exley and DeLoach 1981 (cave description, map);
Bowman and Sket 1985 (Rp, generic description); Exley and Goodman
(map).

WAK-8. WAKULLA SPRINGS (Sec.ll, T.3S, R.IW, Crawfordville
Quad.). Wakulla Springs Cave System. WOODVILLE FAUNA, Pro-
cambarus orcinus (USNM), Procambarus horsti{l) (Morris 1989),
unidentified cave amphipods and isopods (Morris 1989). Public, Wakul-
la Springs State Park, Florida Dept. Natural Resources. REFERENCES:
DeLoach et al. 1989 (description); Franz 1982 (Po); Hobbs et al.
1977 (Po); Hobbs and Means 1972 (Po); Lane 1986 (cave description);
Mohr 1964 (fossils); Morris 1989 (Po); Olsen 1958; Rosenau et al.
1977 (spring description); Rupert 1991 (geology); Rupert and Wilson
1989 (geology and hydrology); Skiles 1989 (cave description); Webb
1974 (fossils).

WASHINGTON COUNTY
WAS-1. ECONFINA BLUE SPRING CAVE (Sec.27, T.IN, R.13W,
Bennett Quad). ECONFINA CREEK FAUNA, Caecidotea sp.l (JL),
Dasyscias franzi-typc locality (UF), other cave-associated species.
Private. REFERENCES: Rosenau et al. 1977 (spring description);
Thompson and Hershler 1991 (Df, type description).

Caves and their Faunas in Florida and South Georgia 109

WAS-2. FALLING WATERS TRAIL CAVE (Sec.27, T.4N, R.13W,
Wausau Quad.). APALACHICOLA FAUNA (?), cave-associated species.
Public, Falling Waters State Park, Florida Dept. Natural Resources.
REFERNCES: Franz et al. 1971 (snails); Lane 1986 (map).

DECATUR COUNTY, GEORGIA
DEC-1. CLIMAX CAVE (3 mi N of Climax). APALACHICOLA
FAUNA (SW Georgia), Uncinocy there warreni-type locality (USNM),
Cambarus cryptodytes (USNM), Haideotriton wallacei (UF). Private.
REFERENCES: Beck and Arden 1984 (geology, cave map); Hobbs
1981 (Cc account); Hobbs and Walton 1968 (Uw, type description);
Hobbs et al. 1977 (Cc); Maddox 1992 (radon concentrations); Warren
1961 (Cc, Hw).

DOUGHERTY COUNTY, GEORGIA
DOU-1. WELL, ALBANY (unidentified site). APALACHICOLA
FAUNA (SW Georgia), Haideotriton wallacei-typQ locality (MCZ).
Private. REFERENCES: Carr 1939 (Hw, type description).

The Chonaphini, a Biogeographically Significant

Milliped Tribe in Eastern and Western North America

(Polydesmida: Xystodesmidae)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences

P.O. Box 29555

Raleigh, North Carolina 27626-0555

ABSTRACT— The Chonaphini, the only Nearctic xystodesmid tribe
represented in both the eastern and western faunal regions, is
the only tribe in the family in which the prefemoral process
is typically more complex and of greater taxonomic utility than
the acropodite. The latter structure varies from narrowly blade-
like to acicular, and excepting Montaphe paraphoena, n.sp.,
lacks secondary projections. The prefemoral process, however,
is often elaborate with secondary structures arising from the
stem. Six genera, three monotypic, and twelve species com-
prise the tribe, with Semionellus Chamberlin and S. placidus
(Wood) inhabiting four areas in the eastern United States from
southeastern Minnesota to westcentral Virginia. The other taxa
occur west of the Continental Divide from Montana to north-
central California and Vancouver Island, Canada. Chonaphe Cook
is represented by two new and two established species in
the United States, C. evexa and schizoterminalis, and C.
remissa Chamberlin and armata (Harger). Chonaphe cygneia
and patriotica, both authored by Chamberlin, and C. serratus
Loomis and Schmitt are placed in synonymy under C armata.
Montaphe elrodi (Chamberlin), the dominant xystodesmid from
eastern Washington to western Montana, is projected to occur
in the southern extremity of central British Columbia adjacent
to Idaho and northeastern Washington. Metaxycheir Buckett and
Gardner and Tubaphe Causey are monotypic, M. prolata Buckett
and Gardner occurring in eastern Washington and the adjoin-
ing part of northern Idaho, and the subcylindrical T. levii Causey
occurring in wet rainforests of the Olympic Mountains and the
southwestern corner of Vancouver Island. Selenocheir n. gen.,
characterized by a short prefemoral process less than half as
long as the acropodite, consists of three new species ranging
from southwestern Oregon to the northern California coast and
the northern Sierra Nevada Mountains. Modern descriptions and
illustrations are presented for all tribal taxa along with keys
to genera and to the species of Chonaphe and Selenocheir.

Brimleyana 20:111-200, June 1994 111

112

Rowland M. Shelley

The milliped family Xystodesmidae, the dominant Nearctic
polydesmoid family, occurs in three general regions of the continent:
the eastern United States and southern Ontario and Quebec, Canada,
east of the Central Plains; from southern Texas and New Mexico to
El Salvador; and along the Pacific Coast west of the Sierra Nevada
and Cascade Mountains from Los Angeles to southern Alaska, with
an eastward extension into western Montana (Shelley 1987). The family
is also well represented in east Asia — Japan, the Riu Kiu Archipelago,
Korea, the vicinity of Vladivostok, Russia, and an unknown area in
central China (Hoffman 1978, 1979). Tribal continuity exists between
Meso-America and the eastern Nearctic, as the Rhysodesmini, the
only Meso-American tribe, is represented in eastern North America
by 9 genera and over 20 species. Similarly, the Orophini and Harpaphini
are represented in both the western Nearctic and Asiatic regions. There

Fig. 1. Distribution of the Chonaphini. A smooth curve is drawn
around range extremes in all directions. Though no records are avail-
able from the interior of British Columbia, the western interior region
shows the projected occurrence of M. elrodi near the border with Idaho
and northeastern Washington.

The Chonaphini 113

are no faunal linkages between the western Nearctic and Meso-America,
and the Chonaphini is the only common tribe between the West and
East.

The Chonaphini occurs primarily in the western United States
and Vancouver Island, Canada (Fig. 1), where there are currently 5
genera and 11 nominal species (Shelley 1990, 1993^). West of the
Columbia Plateau, Tubaphe levii Causey inhabits rainforests of the
Olympic Mountains and Vancouver Island, and Chonaphe armata
(Harger) and C. remissa Chamberlin occur sporadically in western
Oregon and Washington, respectively (Shelley 1990, 19936). In the
western interior, the tribe is represented by Montaphe elrodi (Chamberlin),
the dominant xystodesmid from eastern Washington to western Montana,
C. armata, also occurring sporadically in western Montana, northern
Idaho, and northeastern Oregon and Washington, and Metaxycheir
prolata Buckett and Gardner, localized in Latah and Benewah counties,
Idaho, and the adjacent eastern periphery of Whitman County, Washington
(Buckett and Gardner 1969, Shelley 1990). I place three additional
nominal species — C. cygneia and patriotica, both authored by Chamberlin,
and C. serratus Loomis and Schmitt — in synonymy under C. armata.
East of the Central Plains, the Chonaphini is represented by Semionellus
placidus (Wood), which occurs primarily in three general areas: from
southeastern Minnesota to eastern Wisconsin, from central Michigan
to western Indiana, and from western Maryland to westcentral Virginia
(Shelley 1990).

In addition to its biogeographic significance as a trans-Nearctic
taxon, the Chonaphini is unique in that it reverses the basic anatomical
pattern of the family. In other xystodesmid tribes, the gonopodal acropo-
dite possesses most diagnostic features and is often elaborate, whereas
the prefemoral process is variable but generally simple and with little
taxonomic utility. In the Chonaphini, however, the acropodite is typical-
ly the simple structure, while the prefemoral process is usually larger
and often elaborate, displaying flanges and other projections from its
stem. The prefemoral process is thus the dominant feature of most
chonaphine gonopods and exhibits most taxonomic characters. Indeed,
the acropodite, usually a curved acicular to narrowly blade-like projection,
is so thin and fragile in most species that it is easily broken during
dissection, so care is needed to remove a gonopod with this structure
intact.

With these basic attributes of the prefemoral process and acropodite,
the Chonaphini seems at first to be a coherent assemblage whose
study is comparatively straightforward, but there are undescribed forms
that depart from this pattern, cannot be accommodated by other western

114 Rowland M. Shelley

tribes, and appear to relate to known chonaphines. One species, Montaphe
paraphoena, n. sp., occurring on the western periphery of the Columbia
Plateau in central Washington, has a modified acropodite that expands
distad and possesses a secondary projection. These features lend taxonomic
utility to the acropodite and suggest that additional forms with expanded,
modified acropodites, await discovery in central Washington. Because
of similarities with M. elrodi in coloration and the curvature patterns
of the telopodal elements, I provisionally assign this species to Montaphe
instead of erecting a fifth monotypic genus, although this action necessi-
tates a broad generic diagnosis resulting in a heterogeneous taxon.
Furthermore, Selenocheir, n. gen., with three species in southern Oregon
and northern/central California, is assigned to the Chonaphini even
though it has a short prefemoral process that is less than half the
length of the acropodite. The latter structure conforms to the tribal
diagnosis in being narrowly blade-like to acicular, but differences in
its orientation on the prefemur, in the broadness of its arc, and in the
distal configuration again lend it taxonomic utility. This genus is
more compatible with the Chonaphini than with any other western
xystodesmid tribe, and traits of the prefemoral process and acropodite
appear to represent plesiomorphic conditions that bridge anatomical
gaps with the Harpaphini.

Thus the Chonaphini, which appears to be a homogeneous
assemblage characterized by a narrow, unmodified acropodite and a
long, elaborate prefemoral process, is really a highly variable ensemble
with few consistent, unifying features; even the structures of the cypho-
pods and the configurations of the gonopodal apertures vary widely.
Tubaphe and Selenocheir also lack a gonosternum, the coxae being
attached by membrane only, and those taxa with a sclerotized band
differ in the position of the sternal lobes. For example in M. elrodi,
the sternum is short, resulting in narrowly segregated gonopods, and
there are small lateral lobes subtending the coxae; in M. paraphoena
the sternum is also short and the gonopods are narrowly separated,
but there is an elongated medial lobe instead of two lateral ones. The
Chonaphini, therefore, is not united by a few features shared by all
included taxa but by a number of traits that are shared unevenly
among the components such that only one higher category can be
defined to encompass the scope of this variation. Efforts to divide the
group and render it more homogeneous by splitting off dissimilar
forms like Tubaphe and Selenocheir, which lack a gonosternum, or
just the latter, which has the short prefemoral process, or M. paraphoena,
with the medial sternal lobe and an expanded, modified acropodite,
result in undefinable categories that cannot be contrasted with a recon-

The Chonaphini 115

stituted tribe Chonaphini. The chonaphine taxa therefore relate through
differentially shared aspects of all the principal features; two forms
that do not share one attribute share others, or relate through a third
form with which they share still other features.

Separate taxonomic status for this assemblage was first proposed
by Verhoeff (1941), who established the Chonaphinae as a subfamily
in the "Leptodesmidae." Hoffman (1979) reduced the taxon to tribal
status and listed Chonaphe Cook, Montaphe Chamberlin, Semionellus
Chamberlin, and Metaxycheir Buckett and Gardner as component genera.
He and I (Shelley 1990) placed Tubaphe Causey in the Harpaphini, an
understandable misassignment because one gonopod of the holotype
of T. levii Causey is lost, the other is broken, and the descriptive
accounts and illustrations (Causey 1954^) do not begin to portray the
taxonomically important features. I (Shelley 1990) proposed Metaxycheir
pacifica for a chonaphine on Vancouver Island, British Columbia,
that I subsequently (Shelley 1993^) had to place in synonymy under
T. levii after recollecting the latter in the Olympic Mountains. Despite
the unfortunate proposal of a synonym, my description and illustrations
(Shelley 1990) make T. levii the only chonaphine taxon that has been
characterized in accordance with modern standards. Most were proposed
with terse accounts that merely validated the names, and one new
genus and six new species await description. The purpose of this
contribution, therefore, is to present modern descriptions and illustrations
for all chonaphine taxa except T. levii, where such is provided in the
account of M. pacifica (Shelley 1990). Acronyms of sources of preserved
study material are as follows:

AMNH — American Museum of Natural History, New York,

New York.
BYU — Monte L. Bean Life Science Museum, Brigham Young

University, Provo, Utah.
CAS — California Academy of Science, San Francisco.
CMN — Canadian Museum of Nature, Ottawa, Ontario.
EIL — Zoology Department, Eastern Illinois University, Charleston.
FMNH — Field Museum of Natural History, Chicago, Illinois.
FSCA — Florida State Collection of Arthropods, Gainesville.
MCZ — Museum of Comparative Zoology, Harvard University,

Cambridge, Massachusetts.
MPM — Milwaukee Public Museum, Milwaukee, Wisconsin.
NCSM — North Carolina State Museum of Natural Sciences, Raleigh.
NMNH — National Museum of Natural History, Smithsonian

Institution, Washington, D. C.

116 Rowland M. Shelley

PMNH — Peabody Museum of Natural History, Yale University,

New Haven, Connecticut.
RBCM — Royal British Columbia Museum, Victoria, British

Columbia, Canada.
ROM — Royal Ontario Museum, Toronto, Ontario, Canada.
SDMNH — San Diego Museum of Natural History, San Diego,

California.
TMM — Texas Memorial Museum, University of Texas, Austin.
UBC — Zoology Department, University of British Columbia,

Vancouver, British Columbia, Canada.
UCD — Bohart Entomological Museum, University of California

at Davis.
UMN — Entomology Department, University of Minnesota,

St. Paul.
UWBM — Thomas Burke Washington State Museum, University

of Washington, Seattle.
UWY — University of Wyoming Insect Museum, Laramie.
VMNH — Virginia Museum of Natural History, Martinsville.
WAS — Private collection of William A. Shear, Hampden-Sydney,

Virginia.
WSU — James Entomological Museum, Washington State University,

Pullman.
WU — Biology Department, Willamette University, Salem, Oregon.

LITERATURE REVIEW

The subfamily Chonaphinae was proposed by Verhoeff (1941)
for Chonaphe, which was erected by Cook (1904) for Polydesmus
armatus. This species was one of four myriapods that Harger (1872)
described from an unspecified site along the John Day River, Oregon,
which I (Shelley 1993/?) inferred was near Canyon City, Grant County,
on the western slope of the Blue Mountains. The third xystodesmid to
be named from western North America, after P. {Leptodesmus) hay-
denianus Wood (1864), now in Harpaphe, and P. dissectus Wood
(1867), now in Motyxia, C. armata is also the second oldest chonaphine
species behind P. (L.) placidus, described by Wood (1864) from an
unspecified site in Michigan. Immediately following the latter's description.
Wood (1864) proposed the synonym, "P. (L.) floridus var.?," and
Wood (1865) repeated the accounts of both species, providing a gonopod
illustration of the former.

Sixteen years after Harger's work, Bollman (1888) recorded
L. placidus from Boswell, Benton County, Indiana. In a listing of
North American myriapods, Bollman (1893) included L. armatus and

The Chonaphini 117

placidus, placed P. floridus var.? in synonymy under the latter, and
described L. borealis from Winona, Winona County, Minnesota. Ken-
yon (1893«, b) reported L. floridus, misspelled as ''floriaius'' in the
first work, from four towns in Nebraska, as did Gunthorp (1913)
from Jefferson County, Kansas, which Cragin (1885) previously re-
corded as P. floridus. Because these sites are well removed from the
range of S. placidus, Shelley (1989) concluded that the usages refer
to eurymerodesmids and placed the names in synonymy under species
of Eurymerodesmus.

The first reference to a chonaphine in the present century was
by Carl (1903), who redescribed L. placidus. Cook (1904) erected
Chonaphe and assigned the new combination, C. armata, to a specimen
from an unspecified site in Washington. Chamberlin (1911) recorded
L. armatus from Madison, Washington, probably a misspelling of
Madson, Whitman County, a now non-existent community not far
from Viola, Latah County, Idaho. Chamberlin (1913) described L.
(C.) elrodi from Flathead Lake, Montana, and in a statement in an
introductory paragraph of a paper on an unrelated polydesmoid, he
(Chamberlin 1920) proposed Semionellus and designated L. placidus
as the type species. Apparently unaware of this action, Williams and
Hefner (1928) recorded L. placidus from Allen, Hardin, Wood, and
Seneca counties, Ohio. Attems (1938) published anatomical treat-
ments and illustrations of C. armata, ''Trichomorpha" placida, and
''Amphelictogon" elrodi, but his (Attems 1931) account and illustrations
of C. armata from a farm near Olympia, Thurston County, Washington,
clearly refer to C. remissa, described by Chamberlin (1949). Consequent-
ly, his 1938 treatment of C. armata is really of both congeners.

After Verhoeff (1941) erected the Chonaphinae, Chamberlin (1946)
described C. michigana, from Midland County, Michigan, and two
years later (Chamberlin 1948), transferred this species to Semionellus.
In the latter paper, he also described and illustrated S. tertius, from
Kerrville, Kerr County, Texas, and in the previous year, he (Chamberlin
1947) recorded S. placidus from Garrett County, Maryland. Chamberlin
(1949) transferred L. (C.) elrodi into the new genus, Montaphe, and
described three new species of Chonaphe: C. cygneia, from White
Swan, Yakima County, Washington; C. patriotica, from Fourth of
July Canyon, Kootenai County, Idaho; and C. remissa, from Puyallup,
Pierce County, Washington. Chamberlin (1951) recorded S. placidus
from Fort Benning, Chattahoochee/Muscogee counties, Georgia, a mis-
identification of the introduced paradoxosomatid Oxidus gracilis (Koch),
as noted by Shelley (1990). Causey (1952) recorded Trichomorpha
placida from Dane and Milwaukee counties, Wisconsin, and a cave in

118 Rowland M. Shelley

Union County, Illinois. Causey (1954^) erected Tubaphe for T. levii,
from the Olympic rain forest, Jefferson County, Washington, and
(Causey 19546) recorded M. elrodi from Evans, Stevens County, Washing-
ton. Johnson (1954) included S. placidus among his list of Michigan
millipeds but gave no specific localities. Chamberlin and Hoffman
(1958) listed all species recognized at that time, added New York and
Virginia to the range oiS. placidus, placed L. borealis and S. michigana
in synonymy, and transferred S. tertius into the chelodesmid genus
Aphelidesmus, where it clearly belongs. Its purported occurrence in
Texas can only reflect a labelling error, as the genus otherwise occurs
from Costa Rica southward (Hoffman 1979).

Little has appeared on the Chonaphini in recent years. Hoffman
(1969) noted that Semionellus is monotypic, related to Chonaphe and
Montaphe, and cited its range as being from Minnesota to Illinois and
Ohio, with isolated populations in western Maryland, the adjacent
part of West Virginia, and the Virginia Blue Ridge. Buckett and Gardner
(1969) erected Metaxycheir for M. prolata, occurring near Moscow,
Latah County, Idaho, which they erroneously recorded as Nez Perce
County. They correctly related Metaxycheir to Chonaphe, but apparent-
ly unaware of Verhoeff's subfamily, placed these genera, along with
Harpaphe Cook and Hybaphe Cook, in the Orophinae, erected by
Hoffman (1964) to accommodate Orophe Chamberlin and Pamelaphe
Hoffman. This heterogeneous taxon was disassembled by Hoffman
(1979), who transferred Harpaphe and Hybaphe to the new tribe Harpa-
phini and reduced the Chonaphinae and Orophinae to tribal status.
Loomis and Schmitt (1971) described the final nominal species, C.
serratus, from Sanders and Lake counties, Montana, alluded to the
occurrence of C. armata along Puget Sound, Washington, where the
only congener is C. remissa, and reported numerous new records of
M. elrodi from Lake, Missoula, and Sanders counties, Montana, and
Idaho County, Idaho.

The final relevant works concern Canada. Kevan (1983) listed
all the taxa as potential Canadian inhabitants, and Shelley (1990)
described Metaxycheir pacifica from Vancouver Island and summa-
rized the distribution of the tribe and its generic and specific composi-
tion, omitting Tubaphe and T. levii. Shelley (1993a) synonymized
M. pacifica under T. levii after recollecting the latter in the Olympic
Mountains.

Thus, at this writing, five genera and nine species comprise the
Chonaphini. They are listed chronologically below along with type
localities and other reported occurrences.

The Chonaphini 119

Chonaphe Cook, 1904.

C. armata (Harger 1872). Vicinity of Canyon City, Grant County,
Oregon, as deduced by Shelley (19936). Also recorded from Washington
in general (Cook 1904, Attems 1938); and the Puget Sound area (Loomis
and Schmitt 1971); Oregon (Attems 1938, Chamberlin and Hoffman
1958); Idaho (Attems 1938); Madson (misspelled as Madison), Whitman
County, Washington (Chamberlin 1911); near Olympia, Thurston County,
Washington (Attems 1931); and Benton, Clackamas, and Multnomah
counties, Oregon; King, Chelan, Kittitas, and Stevens counties, Washing-
ton; Latah and Clearwater counties, Idaho; and Lincoln and Missoula
counties, Montana (Shelley 1990).

C cygneia Chamberlin 1949. White Swan, Yakima County,
Washington.

C. patriotica Chamberlin 1949. Fourth of July Canyon, Kootenai
County, Idaho.

C. remissa Chamberlin 1949. Puyallup, Pierce County, Washington.

C. serratus Loomis and Schmitt 1971. 1.0 mi (1.6 km) west of
Noxon, Sanders County, Montana. Also recorded from 4.0 mi (6.4
km) west of Noxon and 2.0 mi (3.2 km) north of Yellow Bay, Lake
County, Montana.

Semionellus Chamberlin, 1920.

S. placidus (Wood 1864). Michigan without further specification.
Also recorded from Minnesota, Illinois, and Ohio (Hoffman 1969);
Michigan (Wood 1865, Attems 1938, Johnson 1954); New York and
Virginia (Chamberlin and Hoffman 1958); western Maryland and adjacent
West Virginia (Hoffman 1969); the Virginia Blue Ridge (Hoffman
1969); Winona, Winona County, Minnesota (Bollman 1893); Dane
and Milwaukee counties, Wisconsin (Causey 1952); Union County,
Illinois (Causey 1952); Midland County, Michigan (Chamberlin 1946);
Boswell, Benton County, Indiana (Bollman 1888); and Allen, Hardin,
Wood, and Seneca counties, Ohio (Williams and Hefner 1928).
Montaphe Chamberlin, 1949.

M. elrodi (Chamberlin 1913). Flathead Lake, Lake/Flathead counties,
Montana. Also recorded from Evans, Stevens County, Washington
(Causey 19546, Shelley 1990); Spokane and Whitman counties, Wash-
ington (Shelley 1990); Lowell, Idaho County, Idaho (Shelley 1990)
and 3.1 mi (5.0 km) west of Surveyor Creek, Idaho County (Loomis
and Schmitt 1971); Clearwater and Shoshone counties, Idaho (Shelley
1990); and numerous localities in Lake, Missoula, and Sanders coun-
ties, Montana (Loomis and Schmitt 1971).
Tubaphe Causey, 1954«.

T. levii Causey (= Metaxycheir pacifica Shelley). Graves Creek

120 Rowland M. Shelley

Gampground, Olympic National Park, Jefferson County, Washington.
Also known from Port Renfrew and seven other sites along the Pacific
Coast in the southwestern corner of Vancouver Island, British Columbia,
Canada (Shelley 1990).

Metaxycheir Buckett and Gardner, 1969.

M. prolata Buckett and Gardner. 7 mi (11.2 km) northeast of
Moscow, Latah County, Idaho. Also known from Laird Park, cited as
Laird "Peak," 3.0 mi (4.8 km) southeast of Harvard, Latah County
(Shelley 1990).

ANATOMY AND TAXONOMIC CHARACTERS

Color — The overall "rust" colored appearance distinguishes species
of Montaphe from sympatric xystodesmids of all tribes — Orophini,
Harpaphini, and Chonaphini — and I have even noticed a tinge of this
pigmentation in dried specimens that have been out of alcohol for
years. I characterize the color as "rust" because it is a muted or
dampened red, as if mixed with a touch of brown, as opposed to the
bright red of Appalachian species of Sigmoria (Shelley and Whitehead
1986). The rust color seems characteristic of the genus, as the pigment
was clearly evident in the type series of M. paraphoena after 1 1/2
years in preservative. The color is actually restricted to the paranota
and concolorous bands along the caudal margins of each metatergite,
but the latter are so broad as to cover most of the metazona and
dominate the organism. Although one notices the dark gray to black
base color, the rust pigmentation overwhelms one's senses and identifies
the milliped as a representative of Montaphe.

The red banded pattern of Semionellus placidus is less distinctive
than the color of Montaphe spp., but in its range, S. placidus is about
the only xystodesmid displaying this color. An occasional specimen
of Sigmoria (Rudiloria) spp. may show reddish stripes, but this pattern
is exhibited primarily by congeners to the south in the Carolinas,
Tennessee, and Georgia (Shelley and Whitehead 1986). Yellow is the
dominant color in the forms of Sigmoria, Apheloria, and Brachoria
that are sympatric with S. placidus.

Body Form and Somatic Features — Tubaphe levii and S. placidus
are readily identified by their general body forms. As noted by Shelley
(1990), the unmistakable, nearly julidan appearance of the caudal 2/3
of its body, caused by the absence of paranota caudal to segment 4
(Fig. 45), instantly distinguishes T. levii from sympatric specimens of
Harpaphe h. haydeniana. Though less distinctive, the generally sub-
cylindrical body form, caused by reduced and declined paranota, also
distinguishes S. placidus from the much broader and more robust

The Chonaphini 121

sympatric representatives of the Apheloriini and Rhysodesmini.
Combining body form with color pattern allows one to unfailingly
recognize S. placidus in the field.

Aside from the size and degree of decline of the paranota, there
are few noteworthy aspects of the somatic details. The epiproct of T.
levii displays a narrow constriction (see Causey 1954«, fig. 2), whereas
those of the other taxa are smoothly subtriangular. Low lateral ridges
subtend the anterior coxae, and the glaborous pregonopodal sterna,
depressed on segment 6 and the caudal half of segment 5 to accommodate
the gonopodal telopodites, generally lack modifications aside from
long, diverging projections on the 4th sternite of S. placidus (Fig.
27). The postgonopodal sterna also are glabrous and unmodified, with
only shallow transverse grooves between the leg pairs and gently
curved caudal margins. The pregonopodal coxae oi Metaxycheir prolata
are swollen with slight anteriorly directed lobes but are unmodified
in other species. Prefemoral spines are present in all species, but only
on postgonopodal legs and often only in the caudal half of the body.

Aperture — The aperture in the Chonaphini usually extends caudad
to at least a small degree in the midline. The only exceptions are
T. levii and Chonaphe evexa, which resemble other northwestern
xystodesmids in lacking even a trace of a caudal extension (Figs. 15,
46). This feature of the aperture is particularly noticeable in C. armata
and S. placidus (Figs. 2, 28), which have long, broad extensions, and
is another trait that distinguishes the latter from sympatric representa-
tives of other eastern tribes. Within these extensions is a sclerotized
ledge or shelf that I believe represents the sternum between the 9th
legs. Thus, it is really only the elevated caudal margin that extends or
"peels" caudad, analogous to "peeling" the backing off a gummed
label; the opening itself is ovoid to elliptical. However, in C. armata
and S. placidus, the extensions are so obvious as to appear a functional
part of the opening even though they are not.

Gonopods — As stated previously, the Chonaphini is unique in
that the prefemoral processes are highly variable and possess most
taxonomically important features, whereas the acropodites are simple
and with little taxonomic utility, as opposed to the reverse in all
other xystodesmid tribes. All genera and most species can be diagnosed
from aspects of the prefemoral process, which is the dominant gono-
podal feature, overshadowing the acropodite in most forms. The arrange-
ments of the telopodal elements relative to each other are also important,
the prefemoral process and acropodite being subparallel in Semionellus
and Montaphe, and not parallel in the other genera. It is this shared

122 Rowland M. Shelley

parallel arrangement coupled with the shared rust color that support
placement of paraphoena in Montaphe along with elrodi.

Prefemoral Process — There are three basic types of prefemoral
processes: the short projections of Selenocheir that are less than half
as long as the acropodites (Figs. 54-55, 60-61, 65-66); long ones, as
long or longer than the acropodite, that are narrowly blade-like to
subacicular (Figs. 35-38, 47-48, 52-53); and long ones that are expand-
ed, broad, and laminate (Figs. 3-7, 10-13, 16-19, 22-25, 30-31). The
distally expanded, trifurcate projection oi Montaphe paraphoena (Figs.
42-43), shorter than the acropodite, is an exception. The massive
structures of Chonaphe and Semionellus are especially lamellate distad.
In Chonaphe, the projection is upright and expands at about 1/3 length
into a narrow shelf or ledge on the dorsal side. The outer margin of
the ledge extends distad a short distance as a translucent shield; in C.
schizoterminalis a distal angular flap on the stem of the prefemoral
process overlaps the shield to form a tube; and in C. remissa the
shield connects to the inturned lateral margin of the stem to form a
continuous lamina that extends to the tip of the process. The distal
part of the acropodite inserts onto this shield or through the tube and
extends for varying lengths, emerging from behind the shield and
becoming visible in medial view in C. armata and evexa (Fig. 5),
while being obscured by the continuation of the shield and its linkage
with the inturned lateral margin in C. remissa (Fig. 11). Distal to the
shelf, there is the flap of C. schizoterminalis (Fig. 23), a low angular
ridge in C. evexa (Fig. 18), and a thickened, convoluted projection in
C. armata and remissa about halfway between the ledge and the tip
(Figs. 5-6, 11-12). Taxonomically, the most important aspect of the
prefemoral process of Chonaphe is the apical configuration, which is
a modification of a basically subdivided structure. In C schizoterminalis,
the tip is deeply divided lateral to the midline into two apical projections,
a narrow, lateral, dactyliform branch that is subequal in length to the
larger, medial branch (Figs. 22-24). The process is apically entire in
C. evexa, but the subterminal lateral margin turns inward for a short
distance to form a narrowly rounded lobe, suggesting an apical division
that has become subapical as the apical-medial margin has enlarged
and expanded (Figs. 18-19). The impression that I receive from C.
armata and remissa is of formerly divided apices that have rejoined,
with the suture line still being evident in the latter, particularly when
viewed in ventral perspective (Fig. 13). In both species, it is as if a
previous division has disappeared, leaving a vertical, coaxial flap or
sclerotized, marginally serrate lamella in C. remissa (Figs. 10-12),
and an inturned, subacuminate, transverse medial corner in C. armata

The Chonaphini 123

(Figs. 4-6). Thus, although taxonomically important features occur
on several parts of the prefemoral process, all four species of Chonaphe
can be diagnosed solely from the apical configuration. It seems ironic
that such significance can accrue to such a minute part of this massive
structure.

Though large, expanded, and laminate, the prefemoral process
of Semionellus (Figs. 30-31) has a different configuration from that
of Chonaphe. Instead of being upright, it bends strongly, at about a
right angle, near 1/3 length and exhibits cupulate flanges on the medial
surface proximal and distal to this bend. There is no ledge or shield
as in Chonaphe, but occasionally the acropodite extends through the
concavity of the proximal flange. The margins of these flanges, particularly
the proximal one, are highly irregular and sometimes jagged, with
minute serrations, larger teeth, and in some individuals, a secondary
proximal flange, also marginally irregular, arising from the basal lamella;
the distal flange is further ornamented by a strong, distally directed
spine from the caudal margin. The distal 1/3 of the prefemoral process
expands to a subacuminate tip on the inner distal corner, but its
principal characteristic is a dense pilosity that arises from the inner
and apical margins and nearly obliterates the tip. There are two kinds
of hairs — long, relatively straight ones arising marginally and submargin-
ally, and short, curved ones that arise from an overhanging ledge on
both the medial and lateral sides and extend at most only to the level
of the first hairs. These distal hairs, apomorphic for Semionellus, are
unique not only in the Chonaphini but also in the family Xystodemidae.
I know of no other xystodesmid genus with hairs on either the prefemoral
process or acropodite.

The other type of prefemoral process is the narrow projection,
which is blade-like, at least basally, in Montaphe elrodi and Metaxycheir,
and subacicular in Tubaphe. It curves generally bisinuately, lacks
secondary structures in Metaxycheir (Figs. 52-53), and subtends a
variable arc and possesses modifications in Montaphe and Tubaphe
(Figs. 35-38, 47-48). The projection curves gently and displays minute
apical barbules from the inner margin in Tubaphe (Figs. 47-48), the
latter being shared with Montaphe elrodi, in which the barbs are
strongly pronounced and extend proximad on the stem of the projection,
though still clustered apically (Figs. 35-38). In M. elrodi the prefemoral
process bends or curves dorsad near midlength, extending well beyond
the acropodal curvature, excepting individuals in which the latter is
distended (Fig. 38), and tapering smoothly and continuously to a sub-
acuminate tip. The barbs are located distad, and proximal to the bend

124 Rowland M. Shelley

is a large, variably cupulate projection from the medial surface with a
variably serrate to strongly toothed and jagged distal margin. This
structure further varies from a simple curved lamella, with a variably
irregular distal margin, to one with two or three marginal lobes or
folds, also with variably irregular distal margins.

Acropodite — Except for Montaphe paraphoena, chonaphine
acropodites vary from narrowly blade-like to acicular, and though
devoid of secondary structures, hold taxonomic utility in the general
form of their curvatures and in their positions relative to the prefemoral
process. Disposing first of M. paraphoena, its acropodite expands
basally into a thickened, flange-like overhang on the outer medial
surface and a strong lateral spine, which is positioned opposite the
terminal expansion of the prefemoral process and is obscured by the
latter in medial view (Figs. 42-43). Distally, the acropodite narrows
somewhat and curls in a broad, open loop that encompasses the entire
prefemoral process. By contrast in M. elrodi, the acropodite is typical
for the tribe, being narrow basally and becoming still narrower and
subacicular distad. It usually curls over the prefemoral process at the
level of the latter's bend, but occasionally it is distended and lies
subparallel to the prefemoral process (Figs. 35-38).

In both Metaxycheir and Tubaphe, the acropodite curves in the
form of a broad, open loop, through essentially a single vertical plane
in the latter and through more than one plane in the former. The
structure is broader and blade-like in Metaxycheir, and its loop extends
beyond the distalmost point of the prefemoral process (Figs. 52-53).
In Tubaphe the acropodite is narrower, becoming acicular distad, and
the prefemoral process extends either through the loop or below it
(Figs. 47-48).

Chonaphe and Semionellus are the only two genera with truly
acicular acropodites. In Chonaphe it curls around the prefemoral process,
inserts onto the shelf, being obscured by the shield, and curves distad
along the projection's medial face (Figs. 4-7, 10-13, 22-25). Occasionally
the acropodite is displaced and curls above the shelf, thus being clearly
visible for its entire length (Figs. 16-19). In Semionellus the structure
bends strongly basally, is sublinear for most of its length, and curves
broadly distad (Figs. 30-31). It typically lies below, and runs generally
parallel to, the prefemoral process, but in a few males, it passes
through the curvature of the proximal flange. In Chonaphe, the acropodite
is somewhat coiled and passes through numerous vertical planes, whereas
in Semionellus, the structure is nearly uniplanar.

Cyphopods — The female genitalia are positioned transversely in
the cyphopodal aperture, which encircles the 2nd legs on segment 3;

The Chonaphini 125

the caudal margin of the aperture is strongly elevated above the metazonal
surface and rises to a peak in the midline. The common valvular
surface is visible in situ; the receptacle lies beneath the medial corners
of the valves; and the operculum is closely appressed to their lateral
surfaces. All these structures are hirsute, the receptacles having long
hairs arising from the ventral margins and extending beyond the ventral
margins of the valves. As the receptacles in other eastern tribes are
glabrous, without even a trace of hairs, the hirsute ones enable females
of S. placidus to be distinguished from sympatric females of other
genera. In most chonaphines, the medial valvular margins project
distinctly ventrad, thereby creating a central depression or cavity on
the common, ventral surface (Figs. 8, 14, 20, 39, 49). Semionellus is
an exception in that the ventral surface is flat, without a trace of a
cavity or prolongation (Fig. 32), as is C. schizoterminalis, which has
central valvular lobes (Fig. 26). In keeping with its structurally different
gonopods, the cyphopods of Selenocheir have only slight suggestions
of medial lobes and central impressions (Figs. 58, 63, 67). Likewise,
the receptacle is alate in most species, being cupped below the medial
corners of the valves, narrowing distinctly in the midline, and extending
for varying lengths along the anterior and posterior sides of the valves.
For the most part, the operculum is relatively large, a distinct sclerite
lateral to the valves, instead of an indistinct structure as in most
xystodesmids.

^ Tribe Chonaphini Verhoeff

Chonaphinae Verhoeff, 1941:403.

Chonaphini: Hoffman, 1979:157. Shelley, 1990:2313-2315.

Components — Chonaphe Cook, 1904; Semionellus Chamberlin,
1920; Montaphe Chamberlin, 1949; Tubaphe Causey, 1954^; Metaxycheir
Buckett and Gardner, 1969; Selenocheir, new genus.

Diagnosis — A tribe of moderate-size Xystodesminae with the
following characteristics: gonopodal aperture with caudal margin ele-
vated, ovoid or extending caudad to varying degrees between 9th
legs, sternum between latter present as "shelf" in caudal extension;
gonocoxae joined by membrane, with or without sclerotized sternal
band, latter usually with lobes subtending coxae, occasionally with
medial lobe; telopodal elements parallel or not parallel; prefemoral
process variable, often elaborately ornamented, short and less than
half as long as acropodite or as long or longer than latter, ranging
from acicular to narrowly blade-like to expanded and laminate, usually
with projections arising from stem or with shallow or deep apical

126 Rowland M. Shelley

cleft; acropodite usually simple and unmodified, acicular to narrowly
blade-like, either circumscribing variably broad arc, with or without
abrupt distal curvature change, bending anteriad at right angle proximally
and curving broadly distad, or looping around prefemoral process and
inserting on shelf on latter; cyphopods oriented transversely in aperture,
with or without prolongations of medial valvular corners and variable
central cavities, receptacles alate or flattened, hirsute with variable
numbers of long hairs arising primarily from ventral margins.

Distribution — Occurring in parts of western North America and
the eastern United States east of the Central Plains from southeastern
Minnesota to the Blue Ridge Province of northern Virginia. Mapping
the available museum records reveals that the range is divided into 7
separate areas (Fig. 1): along the Pacific Coast in the southwestern
corner of Vancouver Island, British Columbia; from the Olympic Penin-
sula of Washington to the northern coast of California and the central
Sierra Nevada Mountains, extending eastward across the Cascade Moun-
tains in Washington to the western fringe of the Columbia Plateau
Physiographic Province; from northcentral Oregon and eastern Washing-
ton to western Montana and Idaho north of the Salmon River, probably
extending just across the International Border into southern British
Columbia between Rossland and Creston; from southeastern Minnesota
to southeastern Wisconsin; from the central lower peninsula of Michigan
through western Ohio to western Indiana; a single site in southeastern
Ohio near the Ohio River; and from western Maryland through eastern
West Virginia to the Blue Ridge Province of westcentral Virginia,
terminating in Shenandoah National Park.

Remarks — The anatomical diversity in this diplopod assemblage
presents formidable obstacles to formulating a tribal diagnosis, as
several key features have exceptions. The gonopodal apertures are
large and expanded caudad in all chonaphines except C. evexa, T.
levii, and Selenocheir sinuata; T. levii and the species of Selenocheir
are the only representatives lacking a sternal remnant; Montaphe para-
phoena is the only chonaphine lacking an acicular to narrowly blade-
like acropodite and the only one with the gonosternum lacking lateral
lobes; the species of Selenocheir are the only ones with short prefemoral
processes; and Metaxycheir prolata is the only component with a
long prefemoral process that lacks modifications. To my knowledge
the Chonaphini is the only xystodesmid tribe in which the presence or
absence of a sternum is not constant, but the narrow aperture of
Tubaphe is shared with C. evexa and S. sinuata; the caudally extended
apertures of S. arcuata and S. directa are shared with other chonaphines;
and the barbules on the prefemoral process of T. levii are shared with

The Chonaphini 127

Montaphe elrodi. After evaluating all the specimens cited herein, it is
apparent that all these genera are related and comprise a distinct
faunal assemblage despite the absence of a unifying anatomical feature.
The closest such trait is the narrowly blade-like to acicular acropodite,
the only exception being Montaphe paraphoena.

The previous range description (Shelley 1990) was published
before field expeditions to Washington, Idaho, and Montana. The
specimens collected on these trips plus additional museum samples
reveal that the distribution in the western United States is not contiguous
and that a separate "finger" extends southward along the Blue Mountains
into Oregon from eastern Washington. Foremost among the new speci-
mens are the syntypes of P. armatus at the PMNH (Shelley 19936),
and three additional samples from Grant and Umatilla counties, Oregon
(WAS), which confirm the species in this range. An additional record
of S. placidus from eastern Ohio, shown as a solitary spot in figures 1
and 69, suggests that the central and eastern areas of this species may
eventually be connected.

Key to Genera of the Chonaphini, based primarily on adult males.

1. Prefemoral process short, less than half as long as acropodite

(Figs. 54-55, 60-61, 65-66); Jackson and Curry counties,
Oregon, to Mendocino and El Dorado counties, California ....

Selenocheir, new genus

Prefemoral process longer, nearly as long as to longer than acropodite
2

2. Prefemoral process generally long and slender, acicular or blade-

like, expanding only distad if at all, with or without variable

processes 3

Prefemoral process broadly expanded and laminate throughout length
5

3. Prefemoral process blade-like and unadorned, without projections,

(Figs. 52-53); Latah and Benewah counties, Idaho

Metaxycheir Buckett and Gardner

Prefemoral process either with variably long barbules or distally
expanded 4

4. Paranota variably broad and distinct throughout body; gonoster-

num present (Figs. 34, 41); Douglas County, Washington, to

Lake County, Montana Montaphe Chamberlin

Paranota present on segments 1-5 only, remaining segments
appearing nearly julidan, with at most only ozopore swell-
ings (Fig. 45); gonosternum absent, coxae connected by
membrane only; Vancouver Island, British Columbia, to

128 Rowland M. Shelley
Jefferson County, Washington

Tubaphe Causey

5. Prefemoral process upright, without distal hairs; distal part of
acropodite inserting onto narrow shielded ledge on pre-
femoral process (Fig. 47); King and Chelan counties,
Washington, to Grant and Coos counties, Oregon, and

Missoula County, Montana

Chonaphe Cook

Prefemoral process bent abruptly dorsad at 1/3 length, with dense
distal hairs on inner and apical margins, without ledge; acro-
podite usually lying parallel to prefemoral process (Figs.

30-31); southeastern Minnesota to westcentral Virginia

Semionellus Chamberlin

Genus Chonaphe Cook
Chonaphe Cook, 1904:56. Attems, 1931:65; 1938:155-156. Chamberlin

and Hoffman, 1958:27. Jeekel, 1971:253. Hoffman, 1979:157.

Kevan, 1983:2968.

Type species — Polydesmus armatus Harger, 1872, by original
designation.

Diagnosis — Paranota present and variably distinct on all segments;
epiproct without constriction; gonocoxae widely segregated by narrow
sternal band, latter with lateral lobes; telopodal elements not parallel,
arising proximad on prefemur; prefemoral process long, expanded and
laminate, elaborately ornamented, with shielded ledge or shelf near
midlength on dorsal surface; acropodite acicular, passing through numer-
ous vertical planes, looping around prefemoral process and curling
onto shelf; cyphopod valves with medial corners usually prolonged
ventrad, subtending central cavity.

Description — A genus of moderately large to large chonaphine
Xystodesminae with the following characteristics:

Body composed of head and 20 segments in both sexes, essentially
parallel sided, tapering at both ends.

Head of normal appearance, smooth, polished. Epicranial suture
faint or distinct, terminating above interantennal region. Antennae
relatively short and broad, becoming progressively more hirsute distad,
with 4 conical sensory cones on ultimate article, no other sensory
structures apparent. Genae not margined laterally, with faint or distinct
central impressions, ends narrowly rounded and extending just beyond
adjacent cranial margins. Facial setae with epicranial, interantennal,
frontal, genal, clypeal, and labral series present, with or without supra-
antennal and subantennal series.

The Chonaphini 129

Terga smooth, polished. Collum relatively broad, ends terminating
above or at same level as those of following tergite. Paranota moderate-
size to well developed, broadest on segments 1-5/6, moderately declined,
angling ventrolaterad and interrupting or continuing slope of dorsum,
anterior corners rounded throughout or blunt on segments 2-3,
caudolateral corners variable, either blunt or with suggestion of tooth
on anteriormost segments and round on remaining tergites, or squared
on anteriormost segments, becoming blunt in midbody region and
blunt caudal to midlength. Peritremata broad, distinct to moderately
distinct, moderately to strongly elevated above paranotal surface; ozopores
located caudal to midlength, opening laterad to sublaterad. Caudal
segments normal for family.

Sides of metazonites variable, with or without variable grooves,
impressions, or ridges above pregonopodal coxae. Strictures distinct.
Gonapophyses moderately long and stout. Pregonopodal sterna glabrous;
4th unmodified or with variable lobes, medially coalesced or widely
segregated; 5th sternum with flattened ridges or low, widely segregated
projections between anterior (4th) legs and depression between 5th
legs; 6th sternum with strong depression between both legs to accom-
modate stems of telopodites, 7th legs set slightly farther apart than
6th. Postgonopodal sterna glabrous, with broad, shallow, central im-
pressions but otherwise without modifications, caudal margins smooth,
gently curved. Coxae without projections; prefemora either without
ventrodistal spines or with variably short spines on legs in caudal half
of body; tarsal claws gently curved. Hypoproct broadly rounded, para-
procts with margins strongly thickened.

Gonopodal aperture generally ovoid, either extending caudad
between 9th legs or terminating anterior to latter, with or without
slight anteriolateral indentations, anterior margin and anterior halves
of sides flush with metazonal surface, latter elevating caudad and
continuing onto caudal margin, either descending in midline or con-
tinuing at level elevation. Gonopods in situ with telopodites extending
anteriad from aperture and lying parallel to each other, overhanging
6th and caudal half of 5th sterna. Coxae variable in size, either with
2, or linear field of 4-8, macrosetae, well separated from each other
by narrow sternal band, latter with short to moderate lobes subtending
coxae. Telopodal elements not parallel, arising proximad on prefemur;
latter moderately long and slender; prefemoral process large, arising
from anteromedial or anterior margins, narrow basally, expanding great-
ly at 1/3 to 1/4 length into ledge or shelf on dorsal side, outer margin
extending beyond latter to form translucent shield, stem of prefemoral
process continuing distad with variably irregular margins, either with

130 Rowland M. Shelley

thickened irregular, convoluted projection on dorsal side at 3/4 length
or with angular ridge or flange, stem either broadly rounded apically
and with inturned lateral subapical lobe, or deeply divided with narrow
lateral and broad medial terminations of subequal length, or with
suggestion of division and with lateral side either bent strongly dorsad
and transversely to stem, or not bent and with laminate, coaxial projection
on dorsal side, or expanding distad and leaning over stem. Acropodite
acicular, arising on caudal or caudolateral sides of prefemur, looping
around prefemoral process and curling onto shelf, apically acuminate.
Prostatic groove arising in pit in prefemur, extending along medial
face of latter onto acropodite, running generally along inner surface
of loop to apical opening.

Cyphopodal aperture broad, encircling 2nd legs, sides and caudal
margin elevated above metazonal surface. Cyphopods in situ with
valves oriented transversely, common surface visible in aperture. Valves
variable in size, subequal, lightly to moderately hirsute, medial corners
extending ventrad into variable lobes, subtending central depressions.
Receptacle small to moderately large, hirsute, or with variable numbers
of hairs arising from ventral margins, either cupped around valves or
flat and inconspicuous. Operculum relatively large, located laterad to
valves, with numerous long hairs.

Distribution — Occurring in five segregated areas in the northwestern
United States as follows: Coos County, Oregon; Grant and Umatilla
counties, Oregon; Benton County, Oregon, to Mason and Chelan counties,
Washington; and Stevens and Whitman counties, Washington, to Missoula
County, Montana. This total area covers about 468 mi (753 km) in
the east-west dimension and 351 mi (565 km) in the north-south, and
encompasses parts of the Northern Rocky Mountains, Columbia Plateau,
Sierra Cascade, and Pacific Border Physiographic Provinces.

Species — Four are known; more may occur in isolated pockets
within the generic range.

Key to Species of Chonaphe (based on adult males)

1. Prefemoral process with broad, thickened, convoluted projection

from dorsal surface distal to shelf (Figs. 5-6, 11-12) 2

Prefemoral process with angular ridge or expanded flap on dorsal
surface distal to shelf (Figs. 16, 18, 22-23) 3

2. Apical margin of prefemoral process with lateral side acuminate,

bent abruptly dorsad, lying transverse to axis (Figs. 4-6);
Stevens and Chelan to Yakima counties, Washington;
Multnomah to Benton and Umatilla to Grant counties, Oregon;
and Sanders County, Montana, to Kootenai and Idaho

The Chonaphini 131

counties, Idaho

armata (Harger)

Distal extremity of prefemoral process with coaxial, marginally
serrate projection on dorsal surface (Figs. 10-12); King and
Mason to Skamania counties, Washington

remissa Chamberlin

3. Apical margin of prefemoral process broadly rounded, entire,
distolateral margin inturned dorsad, forming short sub-
terminal lobe; ridge on anterior surface only slightly elevated
above stem of projection, not laminate (Figs. 16-19); Coos
County, Oregon

evexa, new species

Apical margin of prefemoral process gently curved, deeply divided,
lateral side forming long, slender dactyliform projection, sub-
equal in length to broader medial projection; ridge on anterior
surface elevated and expanded into laminate flange or flap,
overlaying shield (Figs. 22-25); Stevens County, Washington

schizoterminalis, new species

Chonaphe armata (Harger)

Figs. 2-8

Polydesmus armatus Harger, 1872:119-120. pi. II, fig. 8.
Leptodesmus armatus: Bollman, 1893:122. Chamberlin, 1911:264.
Chonaphe armata: Cook, 1904:56-57, pi. Ill, figs. 2a-c. Attems, 1938

(in part): 156, fig. 177. Chamberlin, 1949:125. Chamberlin and

Hoffman, 1958:27. Kevan, 1983:2968. Shelley, 1990:2314;

19936:9-10, figs. 4-5.
Chonaphe cygneia Chamberlin, 1949:125, fig. 1. Chamberlin and

Hoffman, 1958:27. Kevan, 1983:2968. NEW SYNONYMY.
Chonaphe patriotica Chamberlin, 1949:127, figs. 2-3. Chamberlin

and Hoffman, 1958, 1958:27. Kevan, 1983:2968. NEW SYNONYMY.
Chonaphe serratus Loomis and Schmitt, 1971:111-112, figs. 1-2. NEW

SYNONYMY.
Chonaphe serrata: Kevan, 1983:2968.

Type specimens — One male and two female syntypes (PMNH)
collected by O. Harger and G. H. Collier in October 1871 from the
vicinity of Canyon City, Grant County, Oregon.

Diagnosis — Dorsal surface of prefemoral process with broad,
thickened, irregular projection distal to shelf; apical margin sublinear
to gently indented, lateral half curving anteriad and lying transversely
to axis, apically acuminate; shield not connecting with lateral margin.

132

Rowland M. Shelley

Fig. 2-8. Chonaphe armata. 2, gonopods in situ, ventral view of
male from Latah County, Idaho. 3, gonocoxae and sternum, dorsal view
of specimen from Chelan County, Washington. 4, left gonopod of male
from Umatilla County, Oregon, medial view. 5, telopodite of the same,
dorsal view. 6, the same, lateral view. 7, the same, ventral view. 8,
left cyphopod of female from Latah County, Idaho, caudal view, a,
acropodite; c, coxa; o, operculum; pfp, prefemoral process; r, recep-
tacle; V, caudal valve. Scale line for Figure 2 = 1.00 mm; line for
other Figures = 2.78 mm for 3, 2.00 mm for 4-7, 1.00 mm for 8.

The Chonaphini 133

Color in life — Metaterga black, without spots and usually without
stripes, paranota white, cream-colored, yellow, or light reddish orange;
collum without a stripe along anterior margin.

Male syntype — Length 23.3 mm, maximum width 4.3 mm, W/L
ratio 18.5%, D/W ratio 65.1%. Body parallel sided for most of length,
tapering at both ends.

Head capsule smooth, polished; epicranial suture shallow, indistinct,
terminating above interantennal region. Width across genal apices 2.3
mm, interantennal isthmus 0.8 mm. Antennae appearing relatively
short and broad, reaching back to caudal margin of 3rd tergite; relative
lengths of antennomeres 2>3>6>4=5>1>7. Genae with moderately dis-
tinct central impressions, ends broadly rounded and extending slightly
beyond adjacent cranial margins. Facial setae as follows: epicranial
1-1, interantennal absent, frontal 1-1, genal 3-3, clypeal about 8-8,
labral about 12-12.

Collum broad, ends terminating at same level as those of adjacent
tergite. Paranota well developed throughout body, broadest on segments
1-6, angling ventrolaterad but interrupting slope of dorsum, anterior
corners rounded, caudolateral corners squared on segments 2-5, slightly
acuminate on 6-8 and 11, blunt on 9-10 and 12-13, angling caudad
on 14-17. Peritremata distinct, strongly elevated above paranotal surface;
ozopores located caudal to midlength, opening sublaterad.

Sides of metazonites generally smooth, polished, with variable
curved, shallow impressions. Pregonopodal sterna glabrous; 4th sternum
unmodified; 5th sternum with two low, rounded, widely separated
elevations between anterior (4th) legs and concave depression between
5th legs to accommodate apices of telopodites. Prefemora of legs on
segments 11-18 with short but distinct, ventrodistal spines.

Gonopodal aperture extending caudad between 9th legs, 1.8-mm
wide and 1.0-mm long at midpoint, without indentations, sides elevating
into broadly rounded, caudolateral lobes, lower in midline. Gonopods
in situ (Fig. 2, not this specimen) with telopodites situated in opposite
sides of aperture, well separated from each other, extending anteriad
in parallel arrangement over coxae of legs 6-7 (on segment 6), termi-
nating over 5th legs and sternum of segment 5. Gonopod structure as
follows (Figs. 3-7): Coxae of normal size and appearance, with linear
fields of 4-6 macrosetae, well segregated by broad sternum, latter
with moderate-size lateral lobes. Prefemur moderately long, with large
prefemoral process arising from anteromedial surface, latter narrow
basally and leaning slightly laterad, expanding at 1/3 length into ledge
or shelf on dorsal side, outer margin extending beyond surface of
ledge as translucent shield, stem of prefemoral process continuing distad

134 Rowland M. Shelley

with irregular to lightly serrate edges, expanding again at 3/4 length
into broad, thickened irregularly convoluted projection on dorsal side,
distal extremity of stem thin and lamellate, apical margin gently concave,
lateral side bent strongly dorsad, lying transversely to axis, apically
acuminate, medial side not bent, apically rounded. Acropodite arising
on caudolateral side of prefemur, angling anteromediad over ventral
surface of prefemoral process, looping around medial surface of latter,
and curling onto shelf, obscured by shield, curving distad along dorsal
face of process to acuminate tip.

Female syntype — Body highly fragmented, length unmeasurable,
maximum width 4.2 mm, D/W ratio 75.6%. Agreeing essentially with
males in structural details except paranota more declined, giving appear-
ance of slightly more vaulted body. Valves (Fig. 8) relatively large,
moderately hirsute, with large medial lobes protruding through mid-
line of aperture, subtending deep central depression. Receptacle large,
lightly hirsute along ventral margin, cupped around valves, extending
slightly more on caudal side than on anterior.

Variation — There is considerable variation in color. The pattern
is generally bimaculate with colored paranota and uniformly black metaterga,
but two males that I collected at Elk Creek Falls Recreation Area,
Clearwater County, Idaho, displayed broad yellow bands, concolorous
with the paranotal markings, along the caudal margins of tergites 12-
17. The paranotal spots vary in color being white to cream-colored on
specimens that I encountered in Multnomah County, Oregon, reddish-
orange on ones from Stevens County, Washington, and bright lemon
yellow on ones from Latah and Clearwater counties, Idaho.

On the gonopods, the size and degree of irregularity of the distal
projection from the dorsal face of the prefemoral process varies consider-
ably. In some males, the structure is much larger and more flattened
than shown in figures 4-7, becoming a second, more distal, and unprotected
shelf; in others, it is more irregular and convoluted, and occasionally
the side margins curl upwards, forming lips and demarcating a narrow,
elongate trough or groove. In still others, the tip of the projection
bends or curves laterad. The apical gonopodal margin is as shown in
figures 4-7 in all specimens except those from Stevens County, Wash-
ington, in which the entire margin, not just the lateral side, is bent
dorsad. As there is only one sample with this configuration, I include
it under C. armata, but future workers with access to more material
may conclude that this condition is significant enough to warrant specific
recognition.

In females, the size of the medial projection of the valves varies,
some being very large and overshadowing the rest of the valves. The

The Chonaphini 135

receptacle in some females is larger than that of the syntype, extending
ventrad along the lateral surface of the valves and partly enclosing
the operculum as well as being cupped around the anterior and caudal
surfaces. These receptacles are saddle or bowl-shaped, with the valves
situated on top in a ventral depression.

Ecology^The specimens that I collected in Multnomah County,
Oregon, were wandering over the substrate in deciduous forests; those
from Stevens County, Washington, were under partly buried, "stuck,"
pine and fir logs and detached pieces of bark beside a dirt driveway
in a predominantly pine woods; and those from Latah and Clearwater
counties, Idaho, on and under logs, moss, and leaves. Specimens from
Chelan County, Washington, were also found "active on ground."
Labels with preserved samples indicate that C. armata has been en-
countered in willow litter, under rocks and fungi, under logs near
streams, under rotten logs, on moist hillsides, and under cow dung.

Distribution — Occurring in five segregated areas in the northwestern
United States: the lower Willamette Valley of Oregon, the eastern
slope of the Cascade Mountains in central Washington, the western
slope of the Blue Mountains, northern Idaho and western Montana,
and an isolated site in Stevens County, Washington (Figs. 68, 70).
Specimens were examined as follows:

OREGON: Benton Co., ca. 16 mi (25.6 km) N Corvallis, Finley
Natl. Wildlife Ref., 7M, F, 31 March 1972, L. Russell (VMNH) and
2M, F, 26-28 April 1978, D. R. Breakey (WU). Polk Co., Zena, 4-H
Camp, M, 23 April 1983, D. R. Breakey (WU). Marion Co., 2.5 mi
(4.0 km) W Mehama, 4M, 2F, 30 March 1969, R. L. Westcott (UCD).
Clackamas Co., West Linn, MM, FF, 6 May 1972, L. Russell (VMNH).
Multnomah Co., Portland, 2M, 19 June 1882, L. Henshaw (MCZ); South
Portland, M, May 1905, J. E. Benedict (NMNH); and Columbia River
Gorge, Oneonta Trail between Horsetail Falls and Oneonta Gorge, M,
2 June 1991, R. M. Shelley (NCSM) and John B. Yeon St. Pk., along
trail to Elowah Falls, 4M, 3 June 1991, R. M. Shelley (NCSM). Umatilla
Co., jet. N and S Forks, Umatilla R., M, 6 July 1967, F. A. Coyle
(WAS); 3.0 mi (4.8 km) NE Gibbon, along Umatilla R., M, 7 July
1967, F. A. Coyle (WAS); 10.0 mi (16.0 km) E Ukiah, Bear Wallow
Cpgd. along OR hwy. 244, M, 7 July 1967, F. A. Coyle (WAS). Grant
Co., vie. of Canyon City, M, 2F, October 1871, O. Harger, G. H.
Collier (PMNH) TYPE LOCALITY.

WASHINGTON: Stevens Co., along Springdale to Hunters Rd.,
13.5 mi (21.6 km) W jet. WA hwy 231, 6M, 4F, 1 June 1993, R. M.
Shelley (NCSM). Chelan Co., Soda Spring, ca. 7.9 mi (12.6 km) W

136 Rowland M. Shelley

Telma, 47.858°N, 120.968«W, M, 20-22 June 1974, R. Crawford (UWBM);
Wenatchee Ridge, ca 7.2 mi (11.5 km) W Telma, 47.860«N, 120.959°W,
M, F, 20 June 1974, R. Crawford (UWBM); along Roaring Creek, ca
6.5 mi (10.4 km) W Entiat, 47.687°N, 120.335°W, M, 15 June 1985,
R. Crawford (UWBM); along Entiat Summit Road, ca 7.9 mi (12.6
km) N Cashmere, 47.638°N, 120.440«W, 2M, 2F, 12 June 1982, R.
Crawford (UWBM); along US hwy. 2 ca 11.9 mi (19 km) N Leavenworth,
4M, 24 April 1988, C. S. Guppy (RBCM); and along US hwy. 2, ca
10 mi (16 km) N Leavenworth, Tumwater Cpgd., 6M, 4F, 18 August
1990 R. M. Shelley (NCSM). Kittitas Co., Cle Elum, 47.15°N-25°N,
120.90°W, M, 9 May 1953, J. J. Gevers (UWBM); Thorp, 47.0°N,
120. 6°W, M, 22 May 1954, Kilpatrick (UWBM); 5 mi (8 km) W
Ellensburg, F, 4 June 1940, E. F. Dailey (UWBM); beside Yakima R.
opposite Ellensburg, 3M, 8 July 1882, S. Henshaw (MCZ); and mouth
of Moonlight Cyn., ca. 7 mi (11.2 km) W, 3.4 mi (5.4 km) N Thorp,
47.119°N, 120.822°W, M, 25 May 1986, R. Crawford (UWBM). Yakima
Co., Taylors, Wenas Valley, M, 6 July 1882, S. Henshaw (MCZ); and
White Swan, M, 7 May 1938, W. W. Baker (NMNH).

IDAHO: Kootenai Co., locality unspecified, M, September 1890,
Leiberg (NMNH); 4th of July Cyn., M, 12 August 1929, R. V. Chamberlin
(NMNH); and Medimont, 3M, F, 23 June 1957, H. S. Dybas (FMNH).
Benewah Co., 4 mi (6.4 km) SE Emida, M, 3F, 16 April 1987, R. S.
Zack (WSU). Shoshone Co., mouth of Eighty Day Cr., 1.3 mi (2.1
km) up Steamboat Cr. from mouth at Coeur d'Alene R., M, 23 May
1975, F. W. Grimm (CMN); and 8 mi (12.8 km) ENE Clarkia, Hobo
Cedar Grove, 4M, 12 June 1985, W. Suter (UWY). Latah Co., Laird
Park, 8 mi (12.8 km) NE Harvard, Idaho Panhandle Nat. For., M, 9
June 1982, R. S. Zack (WSU) and 3M, F, 4 June 1993, R. M. Shelley
(NCSM); 4.5 mi (7.2 km) N, 8.5 mi (13.6 km) E Harvard, Cleveland
Gulch, M, F, 16 September 1978, A. K. Johnson (NCSM); 3 mi (4.8
km) N, 6.5 mi (10.4 km) E Harvard, along Blue Jacket Cr., M, 15
September 1978, A. K. Johnson (NCSM); nr. Harvard, Idaho Pan-
handle Nat. For., 2M, 22 July 1934, B. Malkin (CAS, WSU); E of
Bovill, along Bob's Cr., F, 30 May 1985, C. Rogers (WSU); and 6 mi
(9.6 km) E Bevrit, along Roger's Cr., 4M, F, 20 May 1986, along
Roger's Cr., 4M, F, 20 May 1986, R. S. Zack (WSU). Clearwater
Co., Elk Cr. Falls Rec. Area E of Bovill, 2F, 8 August 1991, and 4M,
2F, 4 June 1993, R. M. Shelley (NCSM); 3.5 mi (5.6 km) N, 6.4 km
(4.0 mi) E Headquarters, along Middleton Cr., 13M, 13 August 1978,
A. K. Johnson (NCSM); 3.5 mi (5.6 km) N, 7.0 mi (11.2 km) W
Pierce, 8M, 2F, 25 June 1978, A. K. Johnson (NCSM); and Greer, 2F,
30 August 1959, W. J. Gertsch, V. Roth (NMNH). Idaho Co., 6.0 mi

The Chonaphini 137

(9.6 km) E, 9.0 mi (14.4 km) S Pierce, 0.5 mi (0.8 km) N Austin Ridge
Lookout, 4M,2F, 30 June 1978, and 15M, 2F, 29 July 1978, A. K.
Johnson (NCSM); 10.5 mi (16.8 km) E, 6.0 mi (9.6 km) S Pierce,
Knoll Cr. Cyn., 2M, 3F, 25 August 1978, A. K. Johnson (NCSM);
13.0 mi (20.8 km) SSE Pierce, Eldorado Ridge, 2M, 18 July 1978, and
2M, 3F, 25 August 1978, A. K. Johnson (NCSM); 3.0 mi (4.8 km) W
Lowell, M, 18 July 1963, W. F. Barr (FSCA); and Lowell, M, F, 4 July
1949, C. O. Bowles (NMNH).

MONTANA: Sanders Co., 1.0 mi (1.6 km) W Noxon, 2M, F,
date unknown, R. Schmitt (NMNH, FSCA); Thompson Falls, 5M, F,
16 August 1967, J. R. Heifer (UCD); and 3.0 mi (4.8 km) SSE Thompson
Falls, Clark's Peak, Lolo Nat. For., 5M, 4 July 1950, B. Malkin
(CAS, NMNH, WSU). Missoula Co., Missoula, M, 7 July 1950, B.
Malkin (NMNH).

The following literature record is considered accurate and indicated
by the open symbol in figure 68.

WASHINGTON: Whitman Co., Madson (misspelled as Madison),
ca. 3 mi (4.8 km) W, 1 mi (1.6 km) S of Viola, Latah Co., ID
(Chamberlin 1911).

Remarks — Hoffman (1979) suggested that the then five nominal
species of Chonaphe — armata, cygneia, patriotica, remissa, and
serratus — might be subspecifically related, and I (Shelley 1990) suggested
that the genus might be monotypic with all these names synonyms of
armata. However, the unpublished record that I cited from King County,
Washington, is referrable to C. remissa, and the one from Stevens
County, Washington, is a misidentification of a new species, C.
schizoterminalis.

The newly cited records from Umatilla County, Oregon, are in
the Blue Mountains of eastern Oregon and corroborate my conclusion
(Shelley 1993^) that the type locality is in this range. In May 1993 I
spent two days in the vicinity of Canyon City, Grant County, the
presumptive type locality, attempting without success to confirm the
occurrence of C. armata in the southern Blue Mountains. This area
has been drastically altered by lumbering, and the only remaining
hardwoods are narrow willow and alder thickets along creeks. The
predominant vegetation is pine, and most of the southern Blue Mountains
are unsuitable for xystodesmids, which occur primarily in hardwood
associations. When Harger collected the types in October 1871, there
was surely an extensive hardwood community in the broad John Day
River Valley near the present towns of Mt. Vernon, John Day, and

138

Rowland M. Shelley

Fig. 9-14. Chonaphe remissa. 9, gonopods in situ, ventral view of
male from King County, Washington. 10, left gonopod of holotype,
medial viev/. 11, telopodite of the same dorsal view. 12, the same,
lateral view. 13, the same, ventral view. 14, left cyphopod of female
from Skamania County, Washington, caudal view. Abbreviations as in
Figs. 2-8. Scale line for Fig. 9 = 1.00 mm; line for other Figs. =
1.30 mm for 10-13, 1.00 mm for 14.

Prairie City, and also along Canyon Creek, a fast-running tributary
that flows through Canyon City and enters the main river at John
Day. Staying in Canyon City (Shelley 1993^), Harger probably did
not have to go far to find his new millipeds and centipede, but in
May, I drove all the major highways and many back roads in the
vicinities of these towns without finding any of the species or even
suitable habitat. Consequently, the record of C. armata from Grant
County in the southern Blue Mountains (Fig. 68) may reflect more of
the historical range than present occurrence.

The Chonaphini 139

Chonaphe remissa Chamberlin

Figs. 9-14

Chonaphe armata: Attems, 1931:65-67, figs. 100-101; 1938 (in part):156,

fig. 177. Verhoeff, 1941 :fig. 5.
Chonaphe remissa Chamberlin, 1949:127, figs. 4-5. Chamberlin and

Hoffman, 1958:27. Kevan, 1983:2968.

Type specimen — Male holotype (NMNH) collected by W. W.
Baker, 2 May 1933, at Puyallup, Pierce County, Washington.

Diagnosis — Dorsal surface of prefemoral process with broad,
thickened, irregular projection distal to shelf; apical margin slanting
laterad, lateral side with vertical lamina lying along axis, marginally
serrate; shield connecting with inturned lateral margin.

Color in life — Unknown, but evidence of light paranotal spots
on preserved specimens.

Holotype — Length 24.5 mm, maximum width 5.3 mm, W/L ratio
21.6%, D/W ratio 75.5%.

Somatic features agreeing with those of C. armata, with following
exceptions:

Epicranial suture distinct, terminating just above interantennal
region. Width across genal apices 3.2 mm, interantennal isthmus 1.1
mm. Antennae reaching back to midlength of 4th tergite, relative
lengths of antennomeres 2>3>4>5=6>1>7. Genae with distinct central
impressions. Facial setae as follows: epicranial 2-2, suprantennal 1-1,
interantennal 1-1, subantennal 1-1, frontal 1-1, genal 1-1, clypeal
about 11-11, labral about 14-14, merging with clypeal series and con-
tinuing for short distance along genal margins, about 3 setae per side.

Sides of metazonites with low, variable ridges above coxae on
pregonopodal segments. 4th sternum with moderately large, medially
coalesced projections between 3rd legs, length subequal to widths of
adjacent coxae; 5th sternum with two shorter, widely separated pro-
jections between 4th legs and slight depression between 5th legs.
Prefemora of legs on segments 15-18 with distinct distoventral spines,
longest on segments 16-17.

Gonopodal aperture extending caudad between 9th legs, 1.8-mm
wide and 0.9-mm long at midpoint, without indentations, sides elevating
strongly caudad and continuing onto caudal extension. Gonopods in
situ (Fig. 9, not this specimen) extending anteriad from aperture over
segment 6 and the caudal part of segment 5. Gonopod structure as
follows (Figs. 10-13): Coxae with linear fields of 5-8 macrosetae;
sterna with moderate-size lobes subtending coxae. Prefemoral process
arising from anteromedial surface, narrow basally, expanding greatly

140 Rowland M. Shelley

at 1/4 length into ledge or shelf on dorsal side, outer margin extending
beyond surface of ledge as translucent shield, tapering toward and
connecting with inturned lateral margin, stem of prefemoral process
continuing distad with lightly serrate margins, with broad, thickened,
irregularly convoluted projection from caudal surface, apical margin
slanting laterad, with perpendicular, marginally serrate lamina projecting
from dorsal surface lateral to midline. Acropodite arising on caudolateral
side of prefemur, looping around prefemoral process and curling onto
shelf, curling distad and terminating behind lateral extension of shield.

Female from Skamania County, Washington — Body fragmented,
length unmeasurable, maximum width 5.9 mm, D/W ratio 71.4%.
Agreeing closely with holotype in structural details, except paranota
more strongly declined, creating appearance of more vaulted body.
Valves (Fig. 14) moderately large, moderately hirsute, medial corners
strongly elevated, subtending deep central depression. Receptacle small,
situated directly beneath or dorsal to valves on lateral side, hirsute
along ventral surface, extending slightly around caudal side of latter
but not around anterior side.

Variation — As in C. armata, the size and degree of irregularity
of the distal projection of the prefemoral process varies, being larger
and more convoluted in some males. The sides of the projection occasion-
ally have variable teeth that project above the flattened distal surface,
which may be level or slant downward. The shield curves distad and
continues for varying distances beyond the shelf, connecting with the
inturned lateral margin. In some males, this continuation extends to
the distal extremity of the prefemoral process, where it is very narrow
and indistinct; in others, the continuation ends near the level of the
distal projection, and the inward curving lateral margin distal to this
point is separate.

Ecology — The specimen from Mason County was taken from
sabal/alder litter at the edge of a marsh; that from Grays Harbor
County was found on a canyon wall; and those from Ape Cave, Skamania
County, were recovered from a pitfall trap just inside the main entrance.

Distribution — Washington west of the crest of the Cascade Mountains.
Specimens were examined as follows:

WASHINGTON: King Co., Seattle, M, 9 April 1936, E. F. Dailey
(UWBM); and Snoqualmie Falls, M, 19 May 1933, M. H. Hatch (FSCA).
Pierce Co., Puyallup, M, 2 May 1933, W. W. Baker (NMNH) TYPE
LOCALITY. Mason Co., Dennie Ahl Seed Orchard, ca. 5 mi (8 km)
W Potlach, 47.379«N, 123.257-259°W, M, 6 May 1986, R. Crawford
(UWBM). Grays Harbor Co., along Canyon River, ca. 7.0 mi (11.2
km) WNW Matlock, nr. Mason Co. line, 47.262°N, 123.526°W, M, 29

The Chonaphini 141

August 1976, J. T. Thompson (UWBM). Skamania Co., Ape Cave, ca.
4.4 mi (7.0 km) E, 4.0 mi (6.4 km) N Cougar (in Cowlitz Co.),
46.109°N, 122.210°W, M, F, 13-30 May 1983, R. Crawford (UWBM).

The following literature record of C. armata is referrable to C.
remissa.

WASHINGTON: Thurston Co., farm near Olympia (Attems 1931);
Verhoeff's gonopod illustration (1941, fig. 5) is of this specimen.

Remarks — Most prior records of C. armata actually refer to C.
remissa, as no one realized that the Cascade Mountains in Washington
form a distributional boundary, with a different species on each side
of the crest. An addition to the above record is the general comment
by Loomis and Schmitt (1971) that C. armata occurs in the Puget
Sound area. However, south of the Columbia River in Oregon, C.
armata occurs in the Willamette Valley and eastern slope of the Coast
Range, and both the genus and tribe are absent from the Cascades.

The prefemoral processes of C. armata and remissa are very
similar and can be confused but the apical lamina is vertical or
coaxial with the stem in the latter and horizontal or transverse to it in
the former. Additionally, the thickened, convoluted, distal projection
is more proximal in C. remissa, being closer to the shelf/shield and
farther from the tip; in C. armata, it is situated closer to the tip,
occurring almost in the middle between the shelf and tip. The shield is
longer in C. remissa, extending well along the lateral side of the stem
of the prefemoral process; it terminates at the level of the shelf in C.
armata.

Chonaphe evexa, new species
Figs. 15-20

Type specimens — Male holotype and 1 male and 1 female paratypes
(UCD), and 1 male paratype (NCSM), collected by J. S. Buckett and
M. R. Gardner, 11 March 1968, 9 mi (14.4 km) north of Agness, Coos
County, Oregon.

Diagnosis — Dorsal surface of prefemoral process without projection
distal to shelf, with angular ridge; apical margin broadly rounded,
continuing downward to blunt termination on medial side, not divided
but with subterminal lateral margin inturned to narrowly rounded,
subacuminate tip; shield not connecting with lateral margin.

Color in life — Unknown, but evidence of paranoial spots in preserved
specimens.

Holotype — Length 21.5 mm, maximum width 3.4 mm, W/L ratio
15.8%, D/W ratio 82.4%.

142

Rowland M. Shelley

Fig. 15-20. Chonaphe evexa. 15, gonopods in situ, ventral view of
paratype. 16, left gonopod of holotype, medial view. 17, telopodite of
the same, dorsal view. 18, the same, lateral view. 19, the same, ven-
tral view. 20, left cyphopod of female paratype, caudal view. Abbre-
viations as in Fig. 2-8. Scale line for Fig. 15 = 1.00 mm; line for
other Fig. = 1.00 mm for 16-18, 1.20 mm for 19, 0.60 mm for 20.

The Chonaphini 143

Somatic features agreeing with those of C. armata, with following
exceptions:

Epicranial suture faint, becoming more distinct and deeply
impressed near termination above interantennal region. Width across
genal apices 1.9 mm, interantennal isthmus 0.8 mm. Antennae reaching
back to caudal margin of 3rd tergite, relative lengths of antennomeres
2>3>4>6>5>1>7. Genae with faint impressions. Facial setae as follows:
epicranial 2-2, interantennal 1-1, frontal 1-1, genal 3-3, clypeal about
7-7, labral about 11-11.

Collum moderately broad, ends terminating just above those of
succeeding tergite. Paranota moderately developed, broadest on segments
1-5, angling ventrolaterad and continuing slope of dorsum, anterior
corners blunt on segments 2-3, rounded on remaining tergites, caudo-
lateral corners with suggestions of teeth on 2-3, rounded on remaining
segments. Peritremata moderately distinct, moderately elevated above
paranotal surface; ozopores located near midlength, opening laterad.

Sides of metazonites smooth, without ridges or impressions. 4th
sternum with two minute, widely segregated knobs; 5th sternum with
two flattened ridges between 4th legs and moderate depression between
5th legs. Prefemor of legs on segments 14-17 with short, ventrodistal
spines.

Gonopodal aperture without caudal extension between 9th legs,
1.0-mm wide and 0.5-mm long at midpoint, without indentations, an-
terior margin flush with metazonal surface, sides elevating caudad
and continuing onto caudal margin. Gonopods in situ (Fig. 15, of
paratype) with telopodites extending anteriad from aperture in parallel
arrangement, overhanging 6th sternum. Gonopod structure as follows
(Figs. 16-19): Coxae relatively small, with two macrosetae, sternum
with a short lobe subtending each coxa. Prefemoral process arising
from anterior margin, narrow basally and curving slightly bisinuately,
expanding at 1/4 length into shelf, outer margin extending beyond
shelf as translucent shield, latter angling upward and merging imper-
ceptibly with lateral margin of stem, stem without distal projection,
with curvilinear ridge angling distolaterad from level of shelf to near
lateral margin, outer/lateral margin curving to broadly rounded apex,
subterminal edge slightly inturned to subacuminate tip, inner margin
broadly indented, apical margin curving downward to narrowly rounded
tip. Acropodite arising from caudal edge of prefemur, looping around
prefemoral process and curling above shelf, displaced distad, normally
curling onto shelf and terminating behind shield.

Male paratypes — The male paratypes agree with the holotype in
all particulars.

144 Rowland M. Shelley

possible. Agreeing closely with males in somatic features, except paranota
more strongly declined, creating appearance of more highly arched
body. Valves (Fig. 20) relatively small, corners extending slightly ventrad
for equal distances, subtending deep central depression, with dense
hair patches arising from medial and lateral projections. Receptacle
relatively small, flat, and inconspicuous, with long hairs arising from
ventral margin, extending slightly beyond anterior and caudal surfaces
of valves, but not cupped around latter.

Distribution — Known only from the type locality.

Remarks — This intriguing species exhibits several distinct somatic
features. As shown by the lower W/L ratio, C. evexa is longer and
narrower than either C. armata or remissa, a distinction that is made
even more dramatic in side by side comparisons because of its smaller,
more rounded paranota. The gonopodal aperture does not extend caudad
between the 9th legs, in marked contrast to the conditions in both C.
armata and remissa, and the gonopods are oriented slightly differently
in situ. In C. evexa, the acropodite arises almost centrally on the
caudal side, as opposed to a sublateral origin in the other species.
Consequently, the telopodite is rotated slightly counter-clockwise and
is situated differently on the coxa. Occurring in the Coast Range of
southwestern Oregon, C. evexa is well isolated from the main generic
distribution, as the most proximate locality for C. armata, in Benton
County, is some 144 mi (230 km) to the north-northeast.

Chonaphe schizoterminalis, new species
Figs. 21-26

Type specimens — Male holotype, one female paratype, and an
additional segment of a male with gonopods (MCZ) collected by R.
V. Chamberlin on an unknown date at Springdale, Stevens County,
Washington.

Diagnosis — Dorsal surface of prefemoral process without thickened
projection distal to shelf, with angular, laminate flap overlapping shield;
apical margin gently curved and continuing downward to acuminate
termination on medial side, strongly divided with lateral side extending
inward to subacuminate tip as dactyliform projection; shield connecting
with inturned lateral margin.

Color in life — Unknown, specimens completely blanched by
preservative.

Holotype — Length 16.1 mm, maximum width 2.9 mm, W/L ratio
18.0%, D/W ratio 82.8%.

Somatic features agree with those of C. armata, with the following

The Chonaphini

145

Fig. 21-26. Chonaphe schizoterminalis . 21, gonopods in situ, ventral
view of paratype. 22, left gonopod of holotype, medial view. 23, telopodite
of the same, dorsal view. 24, the same, lateral view. 25, the same,
ventral view. 26, left cyphopod of female paratype, caudal view. Ab-
breviations as in Figures 2-8. Scale line for Figure 21 = 1.00 mm;
line for other Figures = 1.25 mm for 22 and 24-25, 1.66 mm for 23,
1.00 mm for 26.

146 Rowland M. Shelley

Somatic features agree with those of C. armata, with the following
exceptions:

Epicranial suture thin but distinct, terminating just above
interantennal region. Width across genal apices 2.2 mm, interantennal
isthmus 0.5 mm. Antennae broken at articulations of 1st and 2nd
articles. Genae with shallow central impressions. Facial setae as follows:
epicranial 2-2, interantennal 1-1, subantennal 1-1, frontal 1-1, genal
3-3, dypeal about 12-12, labral about 16-16.

Collum broad, ends terminating above those of succeeding tergite.
Paranota moderately developed, broadest on segments 1-5, angling
ventrolaterad and continuing slope of dorsum, anterior corners blunt
on segments 2-3, rounded at best on remaining tergites, caudolateral
corners blunt on 2-3, rounded on remaining segments. Peritremata
distinct, strongly elevated above paranotal surface; ozopores located
near midlength, opening laterad.

Sides of metazonites with variable shallow grooves and impressions,
without ridges. 4th sternum with two paramedial projections, medially
coalesced, slightly longer than widths of adjacent coxae; 5th sternum
with two shorter, widely segregated projections subtending 4th coxae,
shorter than widths of latter, moderately depressed between 5th legs.
Prefemora of all legs without traces of spines.

Gonopodal aperture extending slightly caudad over sternum between
9th legs, about 1.3-mm wide and 0.6-mm long at midpoint, with slight
anteriolateral indentations, anterior margin flush with metazonal surface,
sides elevating strongly caudad and continuing onto caudal margin,
lowering slightly in midline. Gonopods in situ (Fig. 21) with telopodites
extending anteriad from aperture in parallel arrangement, overhang-
ing 6th sternum. Gonopod structure as follows (Figs. 22-25): Coxae
relatively small, with 2 macrosetae; sternum with short lobe subtending
each coxa. Prefemoral process arising from anterior margin, broad
basally, expanding at 1/4 length into shelf, outer margin extending
beyond latter as translucent shield, latter angling upward toward lateral
margin and fading out imperceptibly, distal part of stem with angular
laminate flap folding over shield to form enclosed tube, stem continu-
ing distad, apically divided and curving broadly apically, distomedial
margin strongly indented, distolateral margin expanding distad, leaning
over stem, and continuing as acuminate dactyliform projection, subequal
in length to that of medial margin. Acropodite arising from caudal
margin of prefemur, looping above medial surface of prefemoral process
and curling onto shelf, extending through enclosure formed by flap
and shield, emerging distad and terminating behind inturned lateral
margin.

The Chonaphini 147

Paratype gonopods — Agreeing with those of holotype in all details.

Female paratype — Length about 24.7 mm, maximum width 3.4
mm, W/L ratio 13.8%, D/W ratio 82.4%. Agreeing closely with male
in somatic features, except paranota shorter and more strongly declined,
creating appearance of more vaulted body. Valves (Fig. 26) moderate-
size, hirsute along ventral margins with moderate-size lobes at midlength
of ventral surfaces, without depressions. Receptacle small flat, and
inconspicuous, located directly beneath valves, not cupped around
sides of latter, with a few hairs from ventral margin.

Distribution — Known only from the type locality.

Remarks — Neither of the complete specimens is in good condition,
having softened after years in preservative. There is also an additional,
loose reproductive segment of a male with both gonopods intact. I
have labeled these as "male paratype gonopods."

The smallest species, C. schizoterminalis is somewhat broader
in proportion to its length than C. evexa. It is unique in the complete
absence of prefemoral spines on the walking legs, and like C. evexa,
the gonopodal telopodites are rotated counterclockwise on the coxa,
in contrast to the condition in C. armata and remissa. However, C.
schizoterminalis shares the caudal expansion on the aperture with the
last two species. Northeastern Washington has been very poorly sampled,
as most collectors have focused on the wet coastal forests west of the
crest of the Cascades. Additional unexpected millipeds may therefore
await discovery in the Okanogan, Colville, and Kaniksu National Forests
in Okanogan, Ferry, Stevens, and Pend Oreille counties, and C. schizoterminalis
could conceivably occur across the International Border near Rossland
and Trail, British Columbia, only 71.5 mi (113.8 km) to the north. I
visited the Springdale area in May 1993 to try to obtain more material,
but encountered C. armata, not C. schizoterminalis, approximately
13.5 mi (21.6 km) west of the village along the Springdale to Hunters
road.

Genus Semionellus Chamberlin
Semionellus Chamberlin, 1920:97. Attems, 1938:200. Chamberlin and

Hoffman, 1958:47. Jeekel, 1971:287. Hoffman, 1979:157. Kevan,

1983:2968.

Type species — Polydesmus (Leptodesmus) placidus Wood, 1864,
by original designation.

Diagnosis — Paranota present on all segments but becoming
progressively less distinct caudad, caudal part of body appearing sub-
cylindrical; epiproct without constriction; gonocoxae widely segregated
by narrow sternal band, latter with lateral lobes; telopodal elements

148 Rowland M. Shelley

subparallel for most of lengths, arising distad on prefemur; prefemoral
process long, expanded and laminate, elaborately ornamented, with strong
medial spine distal to midlength and numerous hairs on lateral surface
and inner distal and apical margins; acropodite acicular, passing through
essentially a single vertical plane, sublinear for most of length, lying
over medial face of prefemoral process; cyphopod valves flattened ventrad,
without lobes, projections, or cavities.

Description — A genus of moderate-size to large chonaphine
Xystodesminae with the following characteristics:

Body composed of head and 20 segments in both sexes. Head of
normal appearance, smooth. Epicranial suture sharp, distinct. Antennae
relatively short and broad, with 4 conical, terminal sensory cones; no
other sensory structures apparent. Facial setae with epicranial, sub-
antennal, frontal, clypeal, and labral series; interantennal and genal
absent.

Terga smooth, polished; strictures broad, distinct. Collum large
and broad, terminating at same level as succeeding tergite. Paranota
present on all tergites, becoming distinctly shorter around midbody
and progressively more so caudad, moderately declined, continuing
slope of dorsum. Peritremata distinct; ozopores opening sublaterad.

Caudal segments normal for family.

Sides of metazonites smooth, polished, with at most only shallow
impressions. Pregonopodal sterna of males glabrous; 4th with two
long, diverging projections, overhanging adjacent coxae; 5th with two
short lobes between 4th legs, flattened between 5th; 6th sternum strongly
depressed between both leg pairs. Postonopodal sterna glabrous, flat,
and unmodified, with only shallow transverse grooves originating be-
tween leg pairs. Coxae unmodified; prefemora with broadly rounded,
ventrodistal lobes, becoming progressively smaller caudad.

Gonopodal aperture large, subtrapezoidal, with broad caudal
extension between 9th legs. Gonopods in situ with telopodites in parallel
arrangement. Coxae with 2-5 macrosetae, connected to each other by
narrow sternal band, latter with lobes subtending coxae. Telopodal
elements subparallel for most of lengths, arising distad on prefemur;
latter relatively broad; prefemoral process large, expanded and laminate,
elaborately ornamented, with broad, cupulate, irregularly serrate basal
flange, bending abruptly dorsad and expanding into lateral and another
medial flanges, former marginally smooth, narrowing or expanding
distad, with or without thickened margin, latter cupulate basally, curling
around acropodite in some males, marginally irregular and with strong
distal spine, stem of process narrowing then expanding greatly distal
to flanges to subacuminate tip on inner, distal corner, inner margin
and lateral surface hirsute, becoming progressively denser distad, distal

The Chonaphini 149

margin indented. Acropodite long, slender, and aciciular, demarcated
from prefemur by strong constriction, bending anteriad basally, sublinear
for most of length, curving broadly distad, about 3/4 as long as prefemoral
process. Prostatic groove arising in pit in prefemur, running down
medial side of latter and curving onto lateral side of acropodite, curving
around latter and continuing to terminal opening.

Cyphopod aperture long and narrow, encircling 2nd legs, sides
and caudal margin elevated above metazonal surface, more so medial-
ly. Cyphopods in situ with valves oriented transversely in aperture,
common surface visible in opening. Valves moderate-size, moderately
hirsute, without marginal extensions and central depression. Receptacle
moderate-size, ventrally hirsute, located on dorsomedial side of valves,
not alate and cupped around latter. Operculum large, located on dorso-
lateral side of valves, with long apical hairs.

Distribution — Occurring in four segregated areas east of the Central
Plains including parts of Minnesota and Wisconsin; the lower peninsula
of Michigan, northwestern Ohio, and Indiana; one site in eastern Ohio;
and parts of Maryland, West Virginia, and Virginia.

Species — One.

Remarks — The only tribal component east of the Plains, Semionellus
is evidence of prior faunal linkage through the gap of some 1,053 mi
(1,684 km) in the northcentral United States. The Black Hills in western
South Dakota, an island of forested mountains in the midst of intervening
grasslands, is a plausible site for an undiscovered, relict, chonaphine
population that might bridge anatomical gaps between Semionellus
and Chonaphe. However, my field trip there in May 1986 produced
only parajulids and introduced species, and disclosed no xystodesmids,
possibly because of unseasonably cold weather (Shelley 1990). Hardwood-
fir environments in this area, particularly in Spearfish Canyon, the
north slope of Mount Harney, and the Iron Creek drainage above the
Needles Section of Custer State Park, should be reinvestigated in
warmer weather to ensure that the Black Hills do not harbor xystodesmids.
Inselberg ranges in Montana east of the Continental Divide like the
Big and Little Belt Mountains and the Absaroka and Bighorn Ranges
are other plausible areas for undiscovered chonaphines that would be
remnants of the ancient faunal connection.

Semionellus placidus (Wood)

Figs. 27-32

Polydesmus (Leptodesmus) placidus Wood, 1864:9; 1865:225, fig. 56.
Polydesmus (Leptodesmus) floridus, var.? Wood, 1864:9; 1865:226.

150

Rowland M. Shelley

Fig. 27-32. Semionellus placidus. 27, 3rd sternum of male from Mid-
land County, Michigan, caudal view. 28, gonopods in situ, ventral view
of male from Tucker County, West Virginia. 29, gonocoxae and ster-
num of male from Midland County, Michigan, caudal view. 30, left
gonopod of the same, medial view. 31, the same, lateral view. 32,
left cyphopod of female from Pepin County, Wisconsin, caudal view.
Abbreviations as in Fig. 2-8. Scale line for Fig. 28 = 1.00 mm; line
for other Fig. = 1.40 mm for 27, 3.50 mm for 29-31, 1.00 mm for
32.

The Chonaphini 151

Leptodesmus placidus: Bollman, 1888:406; 1893:122. Carl 1903:549-

551, Taf. 17, fig. 18. Williams and Hefner, 1928:110, fig. lib.
Leptodesmus borealis Bollman, 1893:183-184.

Trichomorpha placida: Attems, 1938:119-120, fig. 139. Causey, 1952:9.
Chonaphe michigana Chamberlin, 1946:31-32, figs. 1-2.
Semionellus placidus: Chamberlin, 1947:24, fig. 3. Johnson, 1954:248,

pi. Ill, fig. 19. Chamberlin and Hoffman, 1958:47-48. Kevan,

1983:2968.
Semionellus michiganus: Chamberlin, 1948:258.

Type specimen — Lost. The type locality is Michigan without further
specification (Wood 1864, 1865).

Diagnosis — With the characters of the genus.

Color in life — Paranota red, metaterga black with concolorous
red bands along the caudal margins.

Male from Midland County, Michigan (Holotype ofC. michigana) —
Length 28.3 mm, maximum width 4.8 mm, W/L ratio 16.7%, D/W
ratio 77.1%.

Head capsule smooth, polished; epicranial suture distinct, terminating
in interantennal region. Antennae appearing relatively short and broad,
reaching back to caudal margin of 3rd tergite; relative lengths of
antennomeres 2>3>6>4=5>1>7. Genae not margined laterally, with
distinct central impressions, ends broadly rounded and extending slightly
beyond adjacent cranial margins. Facial setae as follows: epicranial
2-2, interantennal, frontal, and genal not detected, clypeal about 10-
10, labral about 14-14.

Collum moderately broad, ends not extending below those of
succeeding tergite. Paranota well developed on anteriormost segments,
becoming noticeably shorter on segment 9 and progressively shorter
caudad, moderately declined, angling ventrolaterad and continuing
slope of dorsum, anterior corners rounded, caudolateral corners blunt
through segment 9, rounded thereafter. Peritremata broad, distinct,
strongly elevated above apranotal surface; ozopores located caudal to
midlength, opening sublaterad.

Sides of metazonites generally smooth, polished, with variably
curved, shallow impressions. Strictures distinct. Gonapophyses moderately
long and stout, extending moderately from 2nd coxa. Pregonopodal
sterna glabrous, variably modified; 4th sternum with two long, diverging
projections extending ventrolaterad and overlying 3rd coxa (Fig. 27);
5th sternum with two short, subconical, paramedian knobs between
4th legs, flat between 5th legs; 6th sternum strongly depressed to
accommodate telopodites. Postgonopodal sterna glabrous, with bicruciform
impressions, caudal margins gently curved, without modifications.

152 Rowland M. Shelley

Coxae without projections; prefemora with broadly rounded lobes on
outer surface, becoming smaller and less pronounced caudad; tarsal
claws sublinear. Hypoproct broadly rounded, paraprocts with margins
strongly thickened.

Gonopodal aperture very broad, subtrapezoidal, extending caudad
between 9th legs and overhanging caudal margin of segment, with
strong shelf in caudal extension, extending anteriad nearly to anterior
margin of prozonum, latter with only thin sclerotized strip along anterior
margin to maintain structural integrity, 2.4-mm wide and 2.1-mm
long at midpoint, without indentations, sides elevated slightly above
metazonal surface, inner margin continuing smoothly around bases of
coxae, outer margin continuing into strong elevations along caudolateral
and caudal margins, latter strongly thickened and slightly flared. Gono-
pods in situ (Fig. 28, not this specimen) with telopodites extending
anteriad from aperture in parallel arrangement, overhanging 6th ster-
num. Gonopod structure as follows (Figs. 29-31): Coxa relatively
large, with 2-5 macrosetae above cannula, connected to opposite member
by narrow sternal band, latter with two moderate lobes subtending
coxae. Prefemur relatively long and broad, expanding slightly distad,
with large, laminate prefemoral process arising distad from anterior
surface, expanded basally into broad, cupulate flange with strong spur
on medial margin, several minute teeth in indentation on distal margin,
and short, irregularly serrate projection laterally, stem of prefemoral
process curving slightly then bending abruptly dorsad and expanding
into second medial and a lateral flanges, medial flange widest basally
and curving dorsad with minute marginal teeth and strong distal spine,
lateral flange broadly rounded basally, extending distad into broad
ledge overhanging tip, ledge with numerous short hairs, overhanging
denser hairs on stem of prefemoral process, latter narrowing greatly
distal to bases of flanges then expanding greatly on inner surface,
inner margin hirsute and becoming progressively more so distad, expanding
apically into subacuminate tip at inner distal corner, directed dorsad,
apical margin broadly indented. Acropodite arising on caudomedial
side of premur, demarcated by strong constriction, bending abruptly
anteriad and continuing sublinearly parallel to stem of prefemoral
process, curving broadly dorsad distally and narrowing to acuminate
tip, overall length about 3/4 that of prefemoral process.

Female from Pepin County, Wisconsin — Length 27.4 mm,
maximum width 4.4 mm, W/L ratio 16.1%, D/W ratio 84.1%. Agreeing
closely with males in somatic details except paranota more strongly
declined, creating appearance of more highly arched body. Cyphopods
in situ with valves oriented transversely in aperture, common surface

The Chonaphini 153

visible in opening. Valves (Fig. 32) relatively large, subequal, ventral
surfaces flat, without depression, margins not extended, moderately
hirsute. Receptacle moderately large, located at dorsomedial corner
of valves, not alate, not cupped around latter, with long hairs arising
from ventral surface.

Variation — The only noteworthy gonopodal variation concerns
the degree of serration on the medial flanges, which can be more or
less than on the illustrated specimen and quite jagged on occasional
males, and the thickness of the distolateral ledge, whose margin is
thickened on some males and expanded slightly into a rim lying perpen-
dicular to the axis of the prefemoral process. On a few males the
acropodite lies closer to the stem of the prefemoral process, running
through the curvature of the second medial flange. Facial setae on a
male from Tucker Co., WV, are epicranial 2-2, interantennal absent,
subantennae 1-1, frontal 1-1, clypeal about 12-12, labral about 18-18,
merging with clypeal series and continuing for short distance along
genal border, about 4 setae per side.

Ecology — I have collected S. placidus in West Virginia in deciduous
forests and cove habitats under moist leaves near water sources.

Distribution — The only tribal representative occurring in the east,
S. placidus inhabits four segregated areas. The western population
traverses the Mississippi River in the Central Lowland Physiographic
Province from southeastern Minnesota to southeastern Wisconsin and
may not be contiguous, as linkage has not been demonstrated between
the samples taken along the Mississippi and those in southern and
eastern Wisconsin. The central population, also in the Central Lowlands,
extends from the central part of the lower peninsula of Michigan,
near the base of the "thumb," through western Ohio to western and
south central Indiana. The eastern population, located in the Appalachian
Plateau, Ridge and Valley, and Blue Ridge Physiographic Provinces,
ranges from western Maryland through eastern West Virginia, to western
Virginia, reaching its eastern periphery in Shenandoah National Park.
There is also a single record from southeastern Ohio along the Ohio
River. Approximate areas of the populations are 273 mi (437 km)
east/west and 45 mi (72 km) north/south for the western, 245 mi (392
km) east/west and 332 mi (531 km) north/south for the central, and
98 mi (157 km) east/west and 137 mi (219 km) north/south for the
eastern. The western and central populations are separated by around
156 mi (250 km), and the central and eastern populations are segregated
by about 234 mi (374 km); the southeastern Ohio record is intermediate
between the last two areas. No specimens have been encountered in

154 Rowland M. Shelley

about 117 mi (187 km) from the closest site in Ontario, and S. placidus
may eventually be discovered in Essex County, as was the rhysodesmine
species Pleuroloma flavipes Rafinesque (Shelley 1988). However, in
two days of searching in July 1986, including extensive investigations
in Point Pelee National Park, I did not encounter it. Specimens were
examined as follows:

MINNESOTA: Rice Co., Northfield, Carleton College, M, F,
Fall 1955, P. Jensen (VMNH). Wabasha Co., Goodhue, Lake Pepin,
2M, 2F, 25 March 1931, W. J. Gertsch (VMNH). Winona Co., 2 mi
(3.2 km) NE Elba, F, 14 October 1973, B. Cutler (UMN); Whitewater
St. Pk., M, 12 June 1961, H. W. Levi (MCZ); and Winona, juv., date
unknown, Holzinger (NMNH). Houston Co., Houston, M, 25 May
1940, C. E. Mickel (UMN).

WISCONSIN: Pepin Co., Lake Pepin, 2M, 2F, 25 July 1931, W.
J. Gertsch (NMNH). Dane Co., Madison, M, 22 February 1914, A. S.
Pearse (NMNH); and Bascom Woods nr. Madison, F, 30 September
1947, H. W. Levi (MCZ). Ozaukee Co., Mud Lake, 2M, F, 9 April
1979, Hildebrandt, Plonczynski (MPM). County Unknown, "Bob's Cabin,"
2M, 2F, 21 October 1973, P. Riemer (TMM).

MICHIGAN: Midland Co., Midland, 2M, 25 May 1942 and 7
May 1943, R. R. Dreisbach (NMNH). Ingham Co., East Lansing, Sanford
Woodlot, M, F, 3 October 1955, collector unknown (FSCA). Washtenaw
Co., 5 mi (8.0 km) W Ann Arbor, M, 6 July 1948, G. C. Wheeler
(FSCA). Monroe Co., Monroe, 6M, 4F, July 1965, collector unknown
(UCD).

INDIANA: Benton Co., Boswell, M, F, date unknown, Mattier
(NMNH). Greene Co., Richland Cr., exact location unknown, 2F, 25
March 1952, collector unknown (FSCA). Monroe Co., locality not
specified, M, 12 August 1953, B. G. Owen (FSCA); Green's Bluff,
exact location unknown, F, 1 November 1953, B. G. Owen (FSCA);
along Wylie L., M, 12 July 1953, B. G. Owen (FSCA); and Morgan-
Monroe St. For., 2F, 15 April 1952, collector unknown (FSCA). Union
Co., Whitewater St. Pk., 2F, 7 August 1953, B. G. Owen (FSCA).

OHIO: Monroe Co., 1.8 mi (2.9 km) N Rinard Mills, nr. Knowlton
Covered Bridge, F, 8 July 1986, D. R. Whitehead (NMNH).

MARYLAND: Garrett Co., exact location unknown, M, July
1907, W. Stone, T. D. Keim, H. W. Fowler (ANSP).

WEST VIRGINIA: Tucker Co., Lanesville, 5.5 mi (8.8 km) E
WV hwy. 32, Monongahela Nat. For., 3M, F, 23 August 1978, R. M.
Shelley, C. P. Withrow (NCSM); and Dolly Sods, IIM, IIF, 20 July
1986, D. R. Whitehead (NMNH). Randolph Co., 2.1 mi (3.3 km) W
Alpena, along US Hwy. 33, 14M, 5F, 23 August 1978, R. M. Shelley,

The Chonaphini 155

Alpena, along US Hwy. 33, 14M, 5F, 23 August 1978, R. M. Shelley,
C. P. Withrow (NCSM); and S of Harman, along Dry Fork Rd., 7 mi
(11.2 km) SW jet. Job Whitmer Rd., F, 23 June 1986, D. R. Whitehead
(NMNH). Pendleton Co., 4.5 mi (7.2 km) W Judy Gap, along Briery
Gap Rd., M, 2F, 21 August 1988, D. R. Whitehead, L. A. Pereira
(NMNH). Pocahontas Co., Monongahela Nat. For., exact location
unknown, M, 2F, 24 September 1972, W. A. Shear (WAS); and Hills
Cr. Falls Scenic Area, along WV hwy. 55 E Greenbrier Co. line, F,
19 June 1972, W. A. Shear (WAS).

VIRGINIA: Rockingham Co., 1 mi (11.2 km) NNW Rawley Springs,
Tomahawk Mtn., F, 17 June 1988, K. A. Buhlmann (VMNH). Warren
Co., N end of Skyline Dr., Shenandoah Nat. Pk., M, 24 September
1943, collector unknown (NMNH). Page Co., Luray, F, July 1966, C.
Ewing (NCSM); Skyland, Shenandoah Nat. Pk., M, date unknown, J.
P. E. Morrison (NMNH); and along Skyline Dr. at Stony Man Mtn.,
Shenandoah Nat. Pk., 7M, 3F, 29 May-11 June 1950, B. D. Burks
(NMNH). Page I Rappahannock Cos., along Skyline Dr., 1-2 mi (1.6-
3.2 km) S Panorama, M, F, 21 June 1938, H. F. & E. M. Loomis
(VMNH). Madison Co., along Skyline Dr. above Hemlock Spgs. Overlook,
Shenandoah Nat. Pk. 3M, 2F, 26 June 1953, L. Hubricht (VMNH);
along Skyline Dr. at milepost 39, Shenandoah Nat. Pk., F, 13 July
1957, Highton, Barry (VMNH); and "Limberlost," Shenandoah Nat.
Pk., 6M, 2F, 2 juvs., 27 May 1990, C. A. Pogue (VMNH). Greene
Co., along Skyline Dr. nr. milepost 71, Shenandoah Nat. Pk., F, 13
July 1957, Highton, Barry (VMNH). Augusta Co., W side of Humpback
Mtn., along Blue Ridge Pkwy. S of 1-64, M, F, 17 June 1947, (VMNH).
Botetourt Co., nr. Sugarland, Apple Orchard Mtn., 2M, F, 27 May
1962, R. L. Hoffman (VMNH) and M, 14 October 1962, collector
unknown (VMNH).

The following additional literature records are deemed valid and
are denoted by open symbols in figure 69.

WISCONSIN: Milwaukee Co., locality not specified (Causey 1952).

OHIO: Allen, Hardin, Seneca, and Wood cos., localities not specified
(Williams and Hefner 1928).

Deletions — The following literature records to either S. placidus
or a synonym refer to other polydesmoids and hence are deleted.

KANSAS: Jefferson Co., cited as Polydesmus floridus (Cragin
1885) and Leptodesmus placidus (Gunthorp 1913, 1921). No modern,
authentic records are available from anywhere near Kansas, as shown
in figures 1 and 69. These records of P. and L. floridus probably refer
to a eurymerodesmid and were cited as such by Shelley (1989).

156 Rowland M. Shelley

NEBRASKA: West Point, Roca, Rulo, and LaPlatte, cited as L.
floriaius (Kenyon 1893«) and L. floridus (Kenyon 18936). Shelley
(1989) synonymized this usage of L. floridus with E. mundus.

GEORGIA: Ft. Benning (Chamberlin 1951). As noted by Shelley
(1990), this record is a misidentification of Oxidus gracilis (Koch).

ILLINOIS: Union Co., Rich's Cave near Cobden (Causey 1952).
As this site is along the Mississippi River in southern Illinois, far
from the range of S. placidus, this record of a juvenile surely refers
to another xystodesmid.

Remarks — As noted by Shelley (1990), the northern- and western-
most record of S. placidus is that from Rice County, Minnesota, and
the southern- and eastern-most are in Botetourt and Madison counties,
Virginia, respectively. These records span a distance of some 858 mi
(1,381 km) in the east-west dimension and 429 mi (688 km) in the
north-south. The Monroe County, eastern Ohio record, shown by a
dot in figures 1 and 69, was obtained since my 1990 paper and lies
nearly midway between the central and eastern faunal areas. This
record may indicate the existence of a small additional population
along the northern/western side of the Ohio River.

In addition to the narrower body, females of S. placidus can be
distinguished from ones of other eastern taxa by the hirsute receptacle,
as this cyphopodal structure is glabrous in the other eastern tribes.

Genus Montaphe Chamberlin
Montaphe Chamberlin, 1949:127. Chamberlin and Hoffman, 1958:38.

Jeekel, 1971:273. Hoffman, 1979:157. Kevan, 1983:2968.

Type species — Leptodesmus (Chonaphe) elrodi Chamberlin, 1913,
by original designation. -

Diagnosis — Paranota present and distinct on all segments; epiproct
without constriction; gonocoxae narrowly segregated by narrow sternal
band, latter either with two paramedian lobes adjacent to coxae or
single, broad, central lobe; telopodal elements parallel or subparallel
for most of lengths, arising proximad on prefemur; prefemoral process
variable, either long, extending well beyond acropodal loop, and narrowly
blade-like, with strong distal barbules and a broad projection proximal
to midlength, or short, terminating inside acropodal loop, and apically
expanded and deeply divided; acropodite narrowly blade-like basally,
acicular to subacicular distad, typically looping over prefemoral process,
either without projections or expanding basally and with lateral spiniform
projection; cyphopods with medial valves expanding gently to strongly
ventrad, constituting at most only short lobes, subtending at most
only moderate central cavity.

The Chonaphini 157

Description — ^A genus of moderately large chonaphine Xystodesminae
with the following characteristics.

Body composed of head and 20 segments in both sexes, essentially
parallel sided, tapering at both ends.

Head of normal appearance, smooth, polished. Epicranial suture
faint or distinct, terminating above epicranial region. Antennae relatively
short and broad, becoming progressively more hirsute distad, with 4
conical sensory cones on ultimate anticle, no other sensory structures
apparent. Genae not margined laterally, with faint or distinct central
impressions, ends broadly rounded and extending slightly beyond adjacent
cranial margins. Facial setae with epicranial, genal, clypeal, and labral
series, with or without frontal and subantennal series, without inter-
antennal series.

Terga smooth, polished. Collum broad. Paranota well developed,
broadest on anteriormost segments, moderately declined, angling ventro-
laterad and continuing slope of dorsum, anterior corners rounded,
caudolateral corners variable, either blunt, squared, rounded, or angling
caudad. Peritremata distinct; ozopores located caudal to midlength.
Caudal segments normal for family.

Sides of metazonites smooth or granular, with or without variable
ridges above leg coxae. Strictures broad but distinct. Gonapophyses
moderately long and stout, extending moderately from 2nd coxae.
Pregonopodal sterna glabrous; 5th sternum with or without low elevations
between anterior (4th) legs, with moderate depression between 5th
legs; 6th sternum strongly depressed between both legs to accommodate
stems of telopodites. Postgonopodal sterna glabrous, unmodified. Coxa
and prefemora with or without short projections on certain legs; tarsal
claws gently curved. Hypoproct broadly rounded, paraprocts with mar-
gins strongly thickened.

Gonopodal aperture ovoid, extending caudad between 9th legs,
with or without slight anteriolateral indentations, anterior margin and
sides flush with metazonal surface, caudal margin elevated. Gonopods
in situ with telopodites extending generally forward in subparallel
arrangement, overhanging 6th and caudal half of 5th sterna. Coxae
moderate-size, with macrosetal tufts above and below cannula, narrow-
ly segregated by narrow sternal band, with either two lateral or one
central lobes. Telopodal elements parallel or subparallel for most of
lengths, arising basally from prefemur; latter relatively long and slender;
prefemoral process narrow and blade-like basally, curving anteriad at
midlength to 2/3 length, with or without broad, toothed projection
from anterior surface proximal to curve, either expanding distad into
deeply divided, bifurcate termination, or blade-like, extending sublinear-

158 Rowland M. Shelley

ly and curving apically to subacuminate tip, with or without variable
numbers of long, slender barbules scattered along stem distal to midlength
curve and clustered apically. Acropodite either acicular, looping over
prefemoral process, and without projections, or expanding near midlength
and with broad spiniform projection, stem looping thereafter and becoming
subacicular. Prostatic groove arising in pit in prefemur, running along
medial surface of prefemur onto dorsal or inner surface of acropodite,
continuing to terminal opening.

Cyphopodal aperture broad, encircling 2nd legs, sides and caudal
margin elevated above metazonal surface. Cyphopods in situ with
valves oriented transversely in aperture, common surface visible in
aperture. Valves variable in size, subequal, lightly to moderately hirsute,
medial corners gently to strongly extended, subtending slight to moderate
central depression. Receptacles moderate-size, alate, with long hairs
arising from ventral margin. Operculum moderate-size to large, located
lateral to valves.

Distribution — Occurring in the western periphery of the Columbia
Plateau Physiographic Province in central Washington and the northern
Rocky Mountains and environs from eastern Washington to western
Montana.

Species — Two.

Remarks — I (Shelley 1990) stated that in western Canada, the
Xystodesmidae did not occur east of the crest of the Cascade Mountains,
although there was an outside possibility that the family might be
found around Rossland and Trail where the forests seem more moist
and extensive. As Montaphe is now known less than 5 mi (8.0 km)
south of the International Border in Idaho, it seems certain that this
genus is represented in the adjoining part of the interior of British
Columbia.

As envisioned here, Montaphe is heterogeneous; the only features
shared by both component species is the rust color, the narrowly
segregated gonocoxae, and the general curvature pattern of the elements
of the telopodite. To some degree, M. paraphoena represents an abbreviated
or shortened version of M. elrodi, with the prefemoral process terminating
before the acropodal loop rather than after. The acropodite of M.
paraphoena is much broader and the only one in the tribe with a
secondary projection, but I think the general pattern of the gonopodal
elements of M. paraphoena is similar enough to that of M. elrodi to
be accommodated at least temporarily under the same genus, as opposed
to erecting another monotypic taxon. The two species are roughly
115.5 mi (184.8 km) apart, and the substantial anatomical differences
suggest that additional forms may await discovered in central Washington.

The Chonaphini

159

Fig. 33-39. Montaphe elrodi. 33, gonopods in situ, ventral view of
male from Spokane County, Washington. 34, gonocoxae and sternum,
caudal view of male syntype. 35, left gonopod of the same, medial
view. 36, telopodite of the same, lateral view. 37, the same, subventral
view. 38, distal extremity of telopodite of male from Pend Oreille
County, Washington, medial view. 39, right cyphopod of female syntype,
caudal view. Abbreviations as in Fig. 2-8. Scale line for Fig. 33 =
1.00 mm; line for other Fig. = 1.06 mm for 34 and 36, 1.00 mm for
35 and 37-39.

160 Rowland M. Shelley

Montaphe elrodi (Chamberlin)
Figs. 33-39
Leptodesmus (Chonaphe) elrodi Chamberlin, 1913:424-426, fig. 17.
Amphelictogon elrodi: Attems, 1938:159, fig. 179.
Montaphe elrodi: Chamberlin, 1949:127. Causey, 19546:82. Chamberlin

and Hoffman, 1958:38. Loomis and Schmitt, 1971:113-114. Kevan,

1983:2968.

Type specimens — Male lectotype (NMNH) and 9 male and 4
female paralectotypes (MCZ, NMNH) collected by C. C. Adams in
the summer of 1912 at an unknown site on Flathead Lake, Flathead/
Lake counties, Montana.

Diagnosis — Prefemoral process long and narrow, blade-like, bent
abruptly dorsad near midlength, with broad, irregular, and usually
subdivided projection proximal to bend and 5-13 strong, distal barbules;
acropodite narrowly blade-like basally, acicular distad, narrowing constant-
ly throughout length, without projections, usually looping over pre-
femoral process near bend of latter.

Color in life — Paranota rust-colored, metaterga black with broad,
rust-colored bands along caudal margins.

Lectotype — Length 27.5 mm, maximum width 4.1 mm, W/L ratio
14.9%, D/W ratio 70.1%. Body essentially parallel sided throughout
length, tapering at anterior and posterior ends.

Head capsule smooth, polished; epicranial suture shallow, indistinct.
Antennae relatively short and broad, reaching back to anterior part of
4th tergite; relative lengths of antennomeres 2>3=4=5>6>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial 1-1,
interantennal and frontal not detected and presumed absent, genal 1-
1, clypeal about 10-10, labral about 12-12.

Collum broad, ends extending slightly below those of adjacent
tergite. Paranota well developed throughout body, broadest on segments
1-6, caudolateral corners squared on segments 2-3, blunt on 4-5,
rounded on 6-16, angling caudad on 17-18. Peritremata broad, distinct,
strongly elevated above paranotal surface; ozopores opening dorsolaterad.

Sides of metazonites smooth, polished. 5th sternum moderately
depressed between caudal (5th) legs to accommodate apices of telopodites.
Postgonopodal sterna with broad, shallow central impressions and narrow
transverse grooves originating between leg pairs, caudal margins gently
curved. Coxae and prefemora without projections.

Gonopodal aperture ovoid, 1.7-mm wide and 0.6-mm long at
midpoint, without indentations, sides and anterior margin flush with
segmental surface, caudal margin strongly elevated into two broadly
rounded, caudolateral lobes, lower in midline. Gonopods in situ (Fig.

The Chonaphini 161

33, not this specimen) with telopodites extending anteriad from aperture
and overhanging anterior margin, acropodites curling mediad dorsal
to prefemoral processes and overlaying each other in midline, prefemoral
processes angling toward each other and extending forward over 6th
sternum, apices bent dorsad. Gonopod structure as follows (Figs. 34-
37): Coxa moderate size, with macrosetal tufts above and below cannula,
narrowly sepaiated from opposite member and connected by narrow
sternum, latter with small paramedian lobes subtending coxae. Prefemoral
process long and narrow, arising basally on medial side and extending
forward in sublinear fashion, with broad, cupulate medial projection
at 1/3 length, latter narrow basally then expanding broadly with 5
sharply acute teeth, 2 basally and 3 distad, and a second, more distal
projection, also with terminal teeth, arising from the first, stem of
prefemoral process bent strongly dorsad at 2/3 length, extending sub-
linearly and curving downward distad, apically subacuminate, with 8
slender barbules arising from inner margin distal to dorsal bend, spaced
more or less equidistantly, clustered at distal curve. Acropodite arising
lateral to prefemur, long, slender, and acicular, looping over prefemoral
process at level of caudal bend, curling over medial surface of latter
and overhanging medial projection, apically acuminate.

Male paralectotypes — The medial projection of the prefemoral
process varies with more or fewer teeth than in the lectotype. The
number of barbules varies from 5-13, which are generally clustered
distad near the distal curve and evenly spaced back to the bend at 2/3
length. One paralectotype has a barbule proximal to the bend.

Female paralectotype — Length 23.6 mm, maximum width 4.5
mm, W/L ratio 19.1%, D/W ratio 80.0%. Agreeing closely with lectotype
in structural details except paranota slightly shorter and more declined,
creating appearance of more vaulted body. Valves (Fig. 39) moderately
large and subequal, moderately hirsute, medial corners extending ventrad
well below level of lateral corners, subtending central cavity. Receptacle
relatively small, alate, cupped below medial corner of valves, extending
slightly up anterior and caudal surfaces of latter, with long hairs
arising from ventral margins. Operculum large, located laterad to valves.

Variation — As in the paralectotypes, variation among nontypical
males primarily involves the configuration of the medial projection
of the prefemoral process and the number and arrangements of barbules.
The medial projection is flattened and plate-like in some males, while
it is lobate in others; in the western part of the range its margins are
smoother and less serrate than in the types (Fig. 38). The barbules
vary as in the paralectotypes, clustering distad, being rather evenly
spaced proximad, and typically arising distal to the bend at 2/3 length.

162 Rowland M. Shelley

Additionally, the acropodite occasionally does not loop over the prefemoral
process, but lies roughly parallel to it (Fig. 38).

Ecology — According to labels with preserved samples, M. elrodi
has been encountered in a field under stones and logs, under rocks,
and in moss near a small woodland stream. The specimens that I
collected in Boundary and Bonner counties, Idaho, were found in
moist deciduous litter in hardwood patches in predominantly coniferous
forests; those from Idaho County were taken from litter in a ditch
beside a walkway in a large area of deciduous trees; and that from
Pend Oreille County, Washington, was encountered under a log and
rock on a talus slope in a cool, moist cove along a stream. Loomis
and Schmitt (1971) encountered the milliped between and under rocks,
moss, logs, cedar bark, in rotten wood, between rocks just above the
splash zone of cascading water in a stream, at the base of a talus
slide, and in a red cedar, Douglas-fir forest. Sites were typically in
creek bottoms with thick humus layers, and a number of specimens
were taken in cedar groves.

Distribution — Spokane and Stevens counties, Washington, to Lake
County, Montana, and from just south of the International Border in
Boundary County, Idaho, and Pend Oreille County, Washington, to
northern Idaho County, Idaho, north of the transverse stretch of the
Salmon River. The area is approximately 242 mi (387 km) in the
east-west dimension and 216 mi (346 km) in the north-south. The
Whitman County, Washington, record is from the Columbia Plateau,
and most of the other sites are in the Northern Rocky Mountains
Physiographic Province. Specimens were examined as follows:

WASHINGTON: Pend Oreille Co., ca. 4 mi (6.4 km) N Metaline
Falls, along feeder stream to Sullivan Cr., ca. 1.7 mi (2.7 km) E WA
hwy. 31, M, 31 May 1993, R. M. Shelley (NCSM); ca. 9 mi (14.4 km)
SSE Metaline Falls, Noisey Cr. Cpgd., Sullivan L., 2M, 2F, 10 July
1988, R. W. Baumann, Wells, Whiting (BYU); and Gypsy Meadow
NE Metaline Falls, 48.903°N, 117.080«W, F, 13 June 1986, R. Crawford
(UWBM). Spokane Co., Four Lakes, Granite L., 3M, F, 30 May 1947,
M. H. Hatch (FSCA); Mt. Spokane St. Pk., Deadman Cr., F, 10 July
1988, R. W. Baumann, Wells, Whiting (BYU); and Spokane, M, F, 22
July 1882, S. Henshaw (MCZ). Whitman Co., Ewan, 3M, F, 27 August
1932, M. H. Hatch (FSCA, UWBM).

IDAHO: Boundary Co., 4 mi (6.4 km) SW Porthill, along Canyon
Cr., 2M, 3F, 12 August 1991, R. M. Shelley (NCSM). Bonner Co.,
7.5 mi (12.0 km) N Priest River, along ID hwy. 57, M, F, 11 August
1991, R. M. Shelley (NCSM). Shoshone Co., N Kellogg, 1.3 mi (2.0

The Chonaphini 163

km) up Steamboat Cr. from Coeur d'Alene R., 8M, 3F, 15 May 1975,
F. W. Grimm (CMN). Clearwater Co., 5.2 mi (8.3 km) N, 0.5 mi (0.8
km) E Headquarters, 4M, 9 July 1978, A. K. Johnson (NCSM); and
8.7 mi. (13.9 km) E, 5.7 mi. (8.0 km) N Pierce, French Mtn. Rd.,
19M, 17F, 2 and 4 July 1978, A. K. Johnson (NCSM). Idaho Co., 13
mi (20.8 km) SSE Pierce, Eldorado Ridge, M, 12 July 1978, A. K.
Johnson (NCSM); 4 mi (6.4 km) SW Lolo Pass, US hwy. 12 at Russian
Cr., IIM, IIF, 3 September 1978, A. K. Johnson (NCSM); 11 mi
(17.6 km) SW Lolo Pass, along US hwy. 12, M, 25 June 1968, G. B.
Wiggins, Yamamoto, Smith (ROM); 24.3 mi (38.9 km) E Lowell,
along US hwy. 12, 3M, 3F, 10 June 1981, R. M. and S. B. Shelley, P.
D. Hardister (NCSM); Lowell, M, 2F, 4 July 1949, C. O. Bowles,
(NMNH); 1.1 mi (1.8 km) S Stites, Nez Perce Ind. Res., M, 15 May
1975, F. W. Grimm (CMNH); and 3 mi (4.8 km) E Syringa, Middle
Fork Clearwater R., 2M, F, 15 May 1975, F. W. Grimm (CMN).

MONTANA: Sanders Co., 1 mi (1.6 km) W Noxon, 3M, 4F, 2
May 1965, R. Schmitt (FSCA). Missoula Co., Clinton, Hell Gate Run,
3M, 3F, August 1930, collector unknown (NMNH). Flathead Co., Big
Fork, M, F, 24 August 1957, H. W. and L. L. Levi (MCZ). Flathead/
Lake cos., along Flathead L., 4M, F, 22 June 1909, collector unknown
(EIL); and Flathead L., lOM, 4F, summer 1912, C. C. Adams (MCZ,
NMNH) TYPE LOCALITY. Lake Co., 6 mi (9.6 km) E St. Ignatius,
Mission Cr. Canyon, 4M, 3F, 2 September 1978, A. K. Johnson (NCSM).

The following additional literature records are considered valid
and are denoted by open symbols in figure 68. Those from Idaho and
Montana are by Loomis and Schmitt (1971).

WASHINGTON: Stevens Co., Evans (Causey 19546).

IDAHO: Idaho Co., 3.1 mi (5.0 km) W Surveyor Cr.

MONTANA: Lake Co., along Big Fork River and North Crow,
Dog, Jocko, Schmidt, and Hell Roaring Creeks; canyon near Mission
Falls; St. Ignatius and 4.0 mi (6.4 km) E St. Ignatius; 8.5 mi (13.6
km) SE Swan Lake guard station; along Goat Cr., 10.0 mi (16.0 km)
S Swan Lake; along Big Knife Cr., 5.0 mi (8.0 km) from Arlee;
Methodist Camp, Rollins; McDonald Peak; Mission and McDonald
cirques; Lake Mary Ronan; and Yellow Bay. Missoula Co., along
Smith Cr. near Condon ranger station; 1.0 mi (1.6 km) below Smith
Cr.; along Crazy Horse Cr.; Pattee Cyn. Rec. Area; along Miller Cr.
and Mt. Sentinel, Missoula; and along Nine Mile Cr., Huson.

Remarks — The black base color of M. elrodi is restricted to the
prozonum and a narrow area on the anterior edge of the metaterga, so
the rust-colored bands cover most of the latter and dominate the milliped's
appearance. At first glance, it appears almost uniformly rust colored.

164

Rowland M. Shelley

Fig. 40-44, Montaphe paraphoena. 40, gonopods in situ, ventral view
of paratype. 41, gonocoxae and sternum, caudal view of paratype. 42,
left gonopod of holotype, medial view. 43, the same, lateral view. 44,
left cyphopod of female paratype, caudal view. Abbreviations as in
Figs. 2-8. Scale line for Fig. 40 = 1.00 mm; line for other Figs. =
3.20 mm for 41, 2.22 mm for 42-43, 1.00 mm for 44.

The Chonaphini 165

In August 1991 I visited Glacier National Park and searched for
M. elrodi at lower elevations on the west side of the Park, near Apgar,
McDonald Lake, and the cedar grove near Avalanche Campground.
As with Loomis and Schmitt (1971), I did not encounter M. elrodi in
these areas or in regions outside the Park near Whitefish, Columbia
Falls, and Hungry Horse Reservoir. The species therefore seems to be
absent from areas north/northeast of Flathead Lake.

Montaphe elrodi is the dominant species in the eastward xystodesmid
faunal extension into western Montana. Extremely abundant between
eastern Washington and Flathead Lake, M. elrodi is now known from
about 3 mi (4.8 km) south of the International Border near Porthill,
Boundary County, Idaho, and should therefore be expected near Creston,
British Columbia, only about 7 mi (11.2 km) north of Porthill. Records
from Metaline Falls, Pend Oreille County, and Evans, Stevens County,
Washington, are only about 10 and 20 mi (16 and 32 km) south of the
border and suggest the occurrence of M. elrodi near Walneta and
Nelway, British Columbia. It may also occur near Rossland and Trail,
although they are just west of the Columbia River, which may constitute
a distributional barrier. When collected, M. elrodi will represent a
new genus and species for Canada.

Montaphe paraphoena, new species
Figs. 40-44

Type specimens — Male holotype and 13 male, 6 female, and 1
juvenile paratypes (UWBM) collected by R. Crawford, K. Dorweiler,
and J. P. Pelham, 27 April 1991, in Tichenal Canyon, 7.5 mi (12.0
km) south, 1.5 mi (2.4 km) east of Waterville, (47.540°N, 120.035°W),
Douglas County, Washington. One male and one female paratypes
deposited in the NCSM.

Diagnosis — Prefemoral process short, expanding distad to irregular,
deeply divided, bi-lobed termination within curvature of acropodite;
latter expanding basically with thickened medial margin and spiniform
lateral projection, narrowing thereafter, becoming subacicular apically.

Color in life — Paranota rust-colored, metaterga black with broad,
rust-colored bands along caudal metatergal margins.

Holotype — Length 23.6 mm, maximum width 4.1 mm, W/L ratio
17.4%, D/W ratio 78.0%.

Somatic features agreeing with those of M. elrodi, with following
exceptions:

Epicranial suture distinct, accentuated by narrow black line. Width
across genal apices 2.6 mm, interantennal isthmus 0.9 mm. Relative
lengths of antennomeres 2>3>6>4>5>1>7. Genae with distinct central

166 Rowland M. Shelley

impressions. Facial setae as follows: epicranial 2-2, interantennal not
detected and presumed absent, subantennal 1-1, frontal 1-1, genal 4-
4, clypeal about 30-30, labral about 16-16.

Sides of metazonites granular, with ventral ridges just above leg
coxae on segments 1-14, higher and more distinctly elevated above
metazonal surface on segments 1-4, becoming progressively lower
caudad. 5th sternum with two low, widely segregated, paramedian
knobs between anterior (4th) legs, shorter than widths of adjacent
coxae, and moderate depression between 5th legs. Coxae with small
medial lobes on legs 3-5; prefemora with short, indistinct ventrodistal
spines on legs on segments 10-16.

Gonopodal aperture generally ovoid, extending slightly caudad
between 9th legs, 1.6-mm wide and 0.9-mm long at midpoint, indented
slightly anteriolaterad, anterior margin and sides flush with metazonal
surface, caudal margin becoming elevated at caudolateral corner, rising
slightly to midline. Gonopods in situ (Fig. 40, of paratype) with telopodites
extending generally anteriad from aperture and overhanging 6th sternum,
acropodites and prefemoral processes angling toward each other and
overlapping in midline. Gonopod structure as follows (Figs. 41-43).
Coxa moderately large, with 2 dorsal and 4 ventral macrosetae, narrowly
segregated from opposite member by narrow sternal band, latter with
medial lobe. Prefemoral process short and broad, arising from dorsal
surface, curving broadly at 2/3 length and expanding distad, deeply
divided apically and terminating inside loop of acropodite in two
acuminate determinations, medial one broader. Acropodite arising
ventrad on prefemur, curving over prefemoral process in subparallel
arrangement, expanding broadly with thickened medial margin and
with strong projection from lateral surface distal to midlength, latter
spiniform, extending nearly to level of terminus of prefemoral process,
stem of acropodite narrowing slightly distad and looping dorsad be-
yond terminus of prefemoral process, curling to acuminate tip.

Male paratypes — The male paratypes agree with the holotype in
all particulars.

Female paratype — Length 25.6 mm, maximum width 4.0 mm,
W/L ratio 15.6%, D/W ratio 87.5%. Agreeing essentially with the
males in somatic features except paranoia more strongly declined,
creating appearance of more vaulted body. Valves (Fig. 44) small,
subequal, lightly hirsute, becoming slightly higher mediad but without
distinct lobes or projections, slightly depressed centrally. Receptacle
small, alate, cupped around medial side of valves, with long hairs
arising from outer surface. Operculum relatively large, located laterad
to valves.

The Chonaphini 167

Ecology — The types were found under rocks on soil on a barren
slope in a heavily grazed, grassland area.

Distribution — Known only from the type locality, which is on
the eastern side of Badger Mountain and in the western periphery of
the Columbia Plateau Physiographic Province.

Genus Tubaphe Causey
Tubaphe Causey, 1954«:222. Chamberlin and Hoffman, 1958:52. Jeekel,

1971:291. Hoffman, 1979:159. Kevan, 1983:2968.

Type species — Tubaphe levii Causey, 1954, by original designation.

Diagnosis — Paranota present only on segments 1-4, remaining
segments appearing nearly juloid, with at most only slight ozopore
swellings; epiproct with distal constriction; gonocoxae narrowly segre-
gated, attached by membrane only, without trace of sclerotized band;
telopodal elements diverging, not parallel, prefemoral process arising
near midlength of prefemur, acropodite arising distad; prefemoral pro-
cess subacicular, gently curved, with minute distal barbules; acropodite
narrowly blade-like to subacicular, in form of broad, open loop curving
through a single vertical plane; cyphopod valves with medial corners
projecting distinctly ventrad, subtending central cavity.

Description — A genus of small, narrow, and subcylindrical
chonaphine Xystodesminae with the following characteristics.

Body composed of head and 20 segments in both sexes. Head of
normal appearance, smooth. Epicranial suture sharp, distinct. Antennae
moderately long, with 4 conical, terminal sensory cones, no other
sensory structures apparent. Facial setae with epicranial, subantennal,
and genal series present or absent, with clypeal and labral series.

Terga smooth, polished; strictures broad, distinct. Collum broad,
ends terminating above those of succeeding tergite. Paranota present
only on segments 1-4, strongly declined, succeeding segments with
only ozopore swellings. Peritremata indistinct on segments 2-4, absent
thereafter; ozopores opening sublaterad on swellings. Epiproct apically
truncate, distal portion demarcated by constriction.

Sides of metazonites smooth, polished, with slight lobes on segments
2-4. Pregonopodal sterna glabrous, without modifications, strongly
depressed on segment 6. Postgonopodal sterna glabrous, flat, and un-
modified, with only shallow transverse grooves originating between
leg pairs. Coxae with short tubercles on legs of segments 7-14; pre-
femora with ventrodistal spines on legs of segments 9-18.

Gonopodal aperture ovoid, without caudal extension. Gonopods
in situ with telopodites in parallel arrangement. Coxae with macrosetae
varying from 2-10 in two tufts, above and below cannula, attached by

168

Rowland M. Shelley

Figs. 45-49. Tubaphe levii. 45, profile of anterior segments of male
from Jefferson County, Washington, dorsal view. 46, gonopods in situ,
ventral view of male from Vancouver Island, British Columbia. 47,
left gonopod of male from Jefferson County, Washington, medial view.
48, telopodite of the same, lateral view. 49, left cyphopod of female
from Jefferson County, caudal view. Abbreviations as in Figs. 2-8.
Scale line for Fig. 46 = 1.00 mm; line for other Figs. = 0.5 mm for
45, 1.00 mm for 47-48, 1.30 mm for 49.

The Chonaphini 169

membrane only, without trace of sternal band. Telopodal elements
diverging, not parallel; prefemur long and narrow; prefemoral process
arising near midlength of prefemur, long, narrow, and subacicular,
gently curved, extending beyond level of distal extremity of acropodal
curvature, with 3-4 minute, distal barbules. Acropodite arising distad
on prefemur, long and narrow, blade-like to subacicular, demarcated
from prefemur by narrow constriction, configuration a broad, open
loop curving over prefemoral process to acuminate tip. Prostatic groove
arising in pit on prefemur, angling to lateral side of prefemur and
extending onto base of acropodite, continuing to terminal opening.

Cyphopod aperture relatively narrow, encircling 2nd legs, sides
and caudal margin elevated above metazonal surface. Cyphopods in
situ with valves oriented transversely, common surface visible in aper-
ture. Valves relatively large, subequal, and lightly hirsute, medial
corners extending strongly ventrad, subtending deep central cavity.
Receptable moderate-size, subtriangular, located below medial corners
of valves, not cupped around latter, with numerous long hairs. Operculum
large, located lateral to valves, with numerous long hairs.

Distribution — Along the Pacific Coast on the southwestern periphery
of Vancouver Island, British Columbia, and on the western and south-
western slopes of the Olympic Mountains, Washington.

Species — One.

Tubaphe levii Causey

Figs. 45-49

Tubaphe levii Causey, 1954^:223, figs. 2-4. Chamberlin and Hoffman,

1958:52. Kevan, 1983:2968.
Metaxycheir pacifica Shelley, 1990:2311-2313, figs. 1-5.

Type specimens — Male holotype (AMNH) and 2 female paratypes
(FSCA) collected by H. W. and L. L. Levi, 12 July 1951, at Graves
Creek Campground, Olympic National Park, Jefferson County, Wash-
ington. One gonopod of the holotype is lost and the other is broken.

Diagnosis — With the characters of the genus, as illustrated in
figures 45-49.

Description — Recently collected males from Jefferson County,
Washington, conform closely to the detailed anatomical account by
Shelley (1990) of the synonym, M. pacifica; the following supplemental
observations are the only significant additions.

Facial setae: epicranial generally not detected and presumed absent;
one male with one seta per side. Subantennal 1-1 and genal 2-2 on
most males.

170 Rowland M. Shelley

The gonocoxae are loosely joined by membrane with no trace of
a sternal remnant. The coxal macrosetae are in two clusters, one
above, and one below, the cannula, and vary in numbers from 2 to a
cluster of 8-10.

Ecology — In Washington, T. levii is restricted to the wet rain
forests on the western and southwestern slopes of the Olympic Moun-
tains. I did not encounter the milliped in the wettest areas during my
1990 field trip, for example around the Hoh Visitor Center, Olympic
National Park, because the vegetation is so dense that there are few
areas with exposed litter. My success came in slightly drier areas,
where T. levii was typically encountered in association with decaying
deciduous logs, usually under bark. This contrasts with the situation
on Vancouver Island, where I found the milliped in deciduous leaf
litter in August 1989 (Shelley 1990).

Distribution — The western periphery of Vancouver Island, Canada,
from the vicinities of Bamfield to China Beach Provincial Park up to
3-6 mi (5-10 km) inland, and western Clallam, and western and southern
Jefferson, counties, Washington, from Bogachiel State Park to Graves
Creek Campground, Olympic National Park. Canadian localities are
detailed in Shelley (1990); American localities are as follows:

WASHINGTON: Clallam Co., Bogachiel St. Pk., M, 6F, 23 August
1990, R. M. Shelley (NCSM). Jefferson Co., Hoh rain forest, Olympic
Nat. Pk., M, F, 5 May 1991, K. A. Buhlmann (VMNH); along Hoh
River Rd. just outside boundry of Olympic Nat. Pk., ca. 7.4 mi (11.8
km) E jet. US hwy. 101, 3M, 4F, 24 August 1990, R. M. Shelley
(NCSM); along Queets River Rd., Olympic Nat. Pk., 0.6 mi (0.9 km)
inside park boundary, M, 4F, 2 juvs., 24 August 1990, R. M. Shelley
(NCSM) and 4.8 mi (7.7 km) inside park boundary, M, F, 24 August
1990, R. M. Shelley (NCSM); and Graves Cr. Cpgd., Olympic Nat.
Pk., M, F, 12 July 1951, H. W. & L. L. Levi (AMNH, FSCA) TYPE
LOCALITY.

Remarks — Among American xystodesmids, the absence of paranota
and the cylindrical, nearly julidan body form caudal to segment 4 is
unique to T. levii. There are species in which the paranota are reduced
and thus appear somewhat cylindrical, for example S. placidus, but
no others in which these structures are entirely absent. Consequently,
T. levii probably occupies a distinct ecological niche apart from that
of the sympatric xystodesmid Harpaphe h. haydeniana (Wood). It is
noteworthy that there is much greater variation in body form among
the western, or western-related, xystodesmids than in the eastern tribes,
for in addition to the cylindrical and subcylindrical T. levii and S.
placidus, there are two highly convex species, Isaphe convexa Cook

The Chonaphini 171

and Thrinaphe hargeri Shelley (Shelley 1993c, d), and the extremely
flat species of Sigmocheir in the Sierra Nevada Mountains of California.
In contrast, although the eastern forms, representatives of the tribes
Apheloriini, Rhysodesmini, Nannariini, and Pachydesmini, differ in
the degree of convexity, they demonstrate similar overall body forms
without nearly the somatic differences of the western species. Adaptive
radiation of western xystodesmids has manifested itself in general
body form to a much greater degree than has that of the eastern
fauna.

Genus Metaxycheir Buckett and Gardner
Metaxycheir Buckett and Gardner, 1969:67. Hoffman, 1979:157. Kevan,
1983:2968.

Type species — Metaxycheir prolata Buckett and Gardner, 1969,
by original designation.

Diagnosis — Paranota present and distinct on all segments; epiproct
without constriction; gonocoxae narrowly segregated by narrow sternal
band, latter with central lobe; telopodal elements not parallel, prefemoral
process arising near midlength of prefemur, acropodite arising distad;
prefemoral process narrowly blade-like, slightly bisinuate, without
projections; acropodite narrowly blade-like, in form of narrow, open
loop, curving through more than one vertical plane; cyphopod structure
unknown.

Description — A genus of small to moderate-size chonaphine
xystodesminae with the following characteristics:

Body composed of head and 20 segments in both sexes. Head of
normal appearance, smooth. Epicranial suture sharp, distinct. Antennae
moderately long, with 4 conical, terminal, sensory cones and microsensilla
on penultimate antennomeres. Facial setae with epicranial, interantennal,
frontal, genal, clypeal, and labral series; genal setae arranged among
3 groups.

Terga smooth, polished; strictures broad, distinct. Collum large
and broad, ends terminating above those of succeeding tergite. Paranota
present on all tergites, broadest anteriorly, strongly declined, continuing
slope of dorsum and creating appearance of vaulted body. Peritremata
moderately distinct; ozopores opening sublaterad.

Caudal segments normal for family.

Sides of metazonites smooth, polished. Pregopnopodal sterna of
males with small lobes between anterior legs of 5th segment (4th
legs), moderate depression between 5th legs; 6th sternum strongly
depressed between both legs. Postgonopodal sterna flat, glabrous, and

172

Rowland M. Shelley

Figs. 50-53a. Metaxycheir prolata. 50, gonopods in situ, ventral view
of male from Latah County, Idaho. 51, gonocoxae and sternum of
male from Benewah County, Idaho, caudal view. 52, left gonopod of
the same, medial view. 53a, telopodite of the same, lateral view. Ab-
breviations as in Figs. 2-8. Scale line for Fig. 50 = 1.00 mm; line
for other Figs. = 1.20 mm for 51, 1.00 mm for 52-53.

The Chonaphini 173

unmodified, with only shallow transverse grooves originating between
leg pairs. Coxae of legs 3-7 moderately enlarged ventrad, swelling of
3rd coxae angular on anterior surface; prefemora without trace of
spines.

Gonpodal aperture generally ovoid, with caudal extension between
9th legs. Gonopods in situ with telopodites in parallel arrangement.
Coxae with macrosetae fields in two general tufts, attached to each
other by narrow sternal band, latter with central lobe. Telopodal elements
not parallel; prefemur long and narrow; prefemoral process arising
near midlength of prefemur, narrowly blade-like, apically acuminate,
bisinuately curved, without projections. Acropodite arising distad from
prefemur, long and blade-like, curving in form of narrow arc, acuminate.
Prostatic groove arising in pit in prefemur, running along medial face
of prefemur, angling onto lateral surface of acropodite and continuing
to terminal opening.

Females unknown.

Distribution — Whitman County, Washington, to Benewah and
Latah counties, Idaho.

Species — One.

Remarks — Among the three genera with long prefemoral processes,
Metaxycheir has the simplest gonopod, consisting of an unmodified
blade-like prefemoral process and a blade-like acropodite. I show
them as an unresolved trichotomy in figure 72, but Metaxycheir may
be the sister lineage to Tubaphe + Montaphe.

Metaxycheir prolata Buckett and Gardner

Figs. 50-53a

Metaxycheir prolata Buckett and Gardner, 1969:67-70, figs. 1-6. Kevan,
1983:2968.

Type specimen — Male holotype and 2 juvenile paratypes (UCD)
collected by R. L. Westcott, 16 May 1965, 7 mi (11.2 km) NE Moscow,
Latah County, Idaho. The vial label and citation in Buckett and Gardner
(1969) incorrectly state Nez Perce County, but this site is actually in
Latah County.

Diagnosis — With the characters of the genus.

Color in life — Unknown; the specimens that I collected in Whitman
County, Washington, were freshly molted and lacked pigmentation.

Male from Benewah County, Idaho — The following notes on somatic
features supplement the complete characterization of the holotype by
Buckett and Gardner (1969); for consistency in terminology with previous
accounts, gonopodal features are described in detail.

174 Rowland M. Shelley

Length 20.4 mm, maximum width 3.6 mm, W/L ratio 17.6%,
D/W ratio 75.0%.

Width across genal apices 2.1 mm, interantennal isthmus 0.7 mm.
6th antennomere with minute distal microsensilla. Facial setae as follows:
epicranial 2-2, interantennal 1-1, frontal 2-2; genal with four groups, a
central group of 4-4, one lateral to this of 1-1, one beneath antennae of
1-1, and one submarginal of 4-4; clypeal about 24-24; labral about IS-
IS.

Dorsum smooth, polished. Paranota well developed throughout
body, broadest on anterior segments, strongly declined, angling sharply
ventrad and creating appearance of strongly convex body, anterior
corners rounded on all segments, caudolateral corners blunt on segments
1-4, produced slightly caudad beginning on 7 and continuing thusly
to caudal end of body. Peritremata moderately distinct, moderately
elevated above paranotal surface; ozopores located caudal to midlength,
opening sublaterad.

Sides of metazonites smooth, polished. Gonapophyses short and
broad, only slightly extending from 2nd coxae. 5th sternum with 2
short projections subtending 4th coxae, moderately depressed between
5th legs; 6th sternum strongly depressed between both leg pairs to
accommodate stems of telopodites. Coxae of legs 3-7 swollen ventrad,
remaining coxae unmodified, prefemora without projections. " -

Gonopodal aperture ovoid, extending strongly caudad between
9th legs with shelf in extension, l.S-mm wide and O.S-mm long at
midpoint, without indentations, anterior margin flush with metazonal
surface, sides elevating strongly caudad to caudolateral corner, dropping
slightly on caudal extension but still well elevated above metazonal
surface. Gonopods in situ (Fig. 50) with telopodites extending anteriad
in parallel arrangement over 6th sternum, prefemoral processes curv-
ing toward each other and nearly meeting in midline. Gonopod struc-
ture as follows (Figs. 51-53a): Coxa moderate size, with sublinear
field of S-10 macrosetae, connected to opposite member by moderately
sclerotized sternum, latter with medial lobe. Prefemoral process long,
narrowly blade-like, arising anteriolaterad on prefemur, curving broadly
mediad basally then anteriad in generally bisinuate appearance, apically
acuminate. Acropodite blade-like, curving broadly anteriad then dorsad
distally, sides narrowing gradually to acuminate tip, forming narrow
loop.

Ecology — The Whitman County, Washington, specimens were
found under thin layers of moist leaves on relatively hard substrate in
a deciduous thicket beside the picnic area at Steptoe Butte. According

The Chonaphini 175

to the vial label, the Benewah County, Idaho, specimen was collected
from under cow dung.

Distribution — Same as that of the genus, a small area of about
25.1 mi (40.2 km) east/west and 16.9 mi (27.0 km) north/south. Specimens
were examined as follows:

WASHINGTON: Whitman Co., Steptoe Butte, M, 2F, juv., 3
June 1993, R. M. Shelley (NCSM).

IDAHO: Benewah Co., 4.0 mi (6.4 km) SE Emida, M, 16 April
1987, R. S. Zack (WSU). Latah Co., 7.0 mi (11.2 km) NE Moscow,
M, 2 juvs., 16 May 1965, R. L. Westcott (UCD) TYPE LOCALITY;
and 3.0 mi (4.8 km) SE Harvard, Laird Park, M, 1 May 1971, W. A.
Turner (WSU).

Remarks — No females have been collected of M. prolata, so the
cyphopod structure is unknown.

This species has a much broader, less acicular and more blade-
like, acropodite than do the other chonaphine species with long prefemoral
processes. However, the aperture and sternum conform to those of the
species of Chonaphe, Montaphe, and Semionellus.

Selenocheir, new genus

Type species — Selenocheir sinuata, new species.

Diagnosis — Paranota present and variably distinct on all segments;
epiproct without constriction; gonocoxae moderately separated, attached
by membrane only, without trace of sclerotized band; telopodal elements
not parallel, prefemoral process arising proximad on prefemur, acropodite
arising distad; prefemoral process short, less than half as long as
acropodite, generally sublinear, apically variable; acropodite variably
blade-like to subacicular, subtending broad, variable arc, curving through
single vertical plane; cyphopod valves without lobes, extensions, or
cavities; receptacle very large, partly enveloping valves.

Description — A genus of moderately large chonaphine
Xystodesminae with the following characteristics.

Body composed of head and 20 segments in both sexes, essentially
parallel sided, tapering at both ends.

Head of normal appearance, smooth, polished. Epicranial suture
distinct, terminating in interantennal region. Antennae relatively long
and broad, becoming progressively more hirsute distad, with 4 conical
sensory cones and microsensilla on outer distal margin of penultimate
antennomere. Genae not margined laterally, with shallow central impres-
sions, ends narrowly rounded and extending just beyond adjacent cranial
margins. Facial setae with epicranial, genal, clypeal, and labral series
present, with or without interantennal, subantennal, and frontal series.

176 Rowland M. Shelley

Terga smooth, polished. Collum relatively broad, ends terminating
below or at same level as those of following tergite. Paranota well
developed, broadest on anteriormost segments, moderately declined,
continuing slope of dorsum, anterior corners with variably small denticles
on segments 2-4, rounded thereafter, caudolateral corners variable,
either angular, blunt, or rounded. Peritremata broad, moderately distinct,
moderately elevated above paranotal surface; ozopores located caudal
to midlength, opening laterad to sublaterad. Caudal segments normal
for family.

Sides of metazonites with irregularly serrate to jagged ridges
subtending leg coxae in anteior half of body. Strictures distinct. Gona-
pophyses moderately long, extending moderately from 2nd coxae. Pre-
gonopodal sterna glabrous; 4th sternum unmodified; 5th sternum with
variable low knobs or elevated areas between 4th legs, flat, depressed,
or with low, elevated areas between 5th; 6th sternum either depressed
between both legs or with widely separated lobes between anterior
pair. Postgonopodal sterna glabrous, unmodified, with at most only
shallow transverse grooves, caudal margins smooth and gently curved.
Coxae without projections; prefemora with variably short, indistinct
ventrodistal spines in caudal half of body; tarsal claws gently curved.
Hypoproct broadly rounded, usually slightly extended in midline; para-
procts with margins strongly thickened.

Gonopodal aperture generally broadly ovoid, with at most only
very slight caudal extension of caudal margin, without anteriolateral
indentations, anterior margin flush with metazonal surface, sides ele-
vating caudad and continuing onto caudal margin, latter flared. Gonopods
in situ with telopodites directed either anteriad or anteromediad from
aperture, angling toward each other, lying parallel or crossing in midline,
extending forward over anterior margin of aperture and 6th sternum.
Coxae moderate-size, with 2 macrosetae above cannula and variably
dense cluster below, connected by membrane only, without trace of
sclerotized sternal band. Telopodal elements not parallel; prefemur
relatively short to moderately long; prefemoral process short, arising
proximad on anterior side, latter less than half as long as acropodite,
generally sublinear basally, with or without slight curve or bend at
midlength, apically broad or narrow, variably divided or simple. Acropo-
dite arising distad from prefemur, either smoothly or demarcated by
constriction, directed anteriad or anteromediad, relatively long, blade-
like basally, narrowing distad and in some cases becoming subacicular,
curving downward or dorsad distally and subtending variable arch,
with or without variably sinuate to uncinate apical bends or curves,
apically acuminate. Prostatic groove arising in pit in prefemur, running

The Chonaphini 177

along medial surface of latter and crossing to lateral side at base of
acropodite, crossing back to inner surface, or continuing along outer,
to terminal opening.

Cyphopodal aperture broad, encircling 2nd legs, sides and caudal
margin elevated above metazonal surface, latter rising to peak in
midline. Cyphopods in situ with valves oriented transversely in aperture,
common surface visible in opening. Valves variable in size, subequal
or anterior valve slightly larger, without distinct lobes or cavities,
situated nearly entirely on top of (ventral to) receptacle. Latter large,
alate, cupped around valves, more so on anterior side, with long hairs
arising from ventral margin. Operculum small to large, located laterad
to valves, with numerous long hairs.

Distribution — From Curry and Jackson counties, in southwestern
Oregon, to Mendocino and El Dorado counties, California, in the
northern Coast Range, the northern Sacramento Valley, and the Sierra
Nevada foothills, an area of about 176 mi (282 km) east/west and 273
mi (437 km) north/south.

Species — Three are currently recognized.

Remarks — The Chonaphini is the best placement for this genus,
as the only other option is a new tribe. It displays narrowly blade-like
to acicular acropodites that are similar to those of Tubaphe and Metaxycheir,
and shares the absence of a sternum with Tubaphe. The short prefemoral
process is plesiomorphic, and Selenocheir tends to bridge the anatomical
gap between the Chonaphini and Harpaphini in having a larger coxa
in relation to the overall bulk of the telopodite, and in the anteromedially
directed acropodites, particularly in S. directa, reminiscent of the
condition in Isaphe (Shelley 1993t/).

Key to Species of Selenocheir, based on adult males.

1. Acropodite subtending very broad, poorly defined arc, without

distinct distal curvature (Figs. 60-61); Tehama to Humboldt

and Mendocino counties, California arcuata, new species

Acropodite curving definitely downwards or dorsad distally, with
relatively distinct distal curve or bend 2

2. Distal part of acropodite, distal to distal curve/bend, variably

sinusoid (Figs. 54-57); Shasta to Mendocino and El Dorado

counties, California sinuata, new species

Distal part of acropodite without trace of sinusoid pattern (Figs.
65-66); Curry and Jackson counties, Oregon, to Del Norte
County, California directa, new species

178

Rowland M. Shelley

53b

Figs. 53b-58. Selenocheir sinuata. 53b, gonopods in situ, ventral view
of male from El Dorado County, California. 54, left gonopod of holo-
type, medial view. 55, telopodite of the same, lateral view. 56, distal
extremity of acropodite of left gonopod of male from Humboldt County,
California, medial view. 57, distal extremity of acropodite of left gonopod
of male from W of Burney, Shasta County, California, medial view.
58, left cyphopod of female paratype, caudal view. Abbreviations as in
Figures 2-8. Scale line for Figure 53b = 1.00 mm; line for other
Figures = 1.33 mm for 54-55, 1.14 mm for 56-57, 1.00 mm for 58.

The Chonaphini 179

Selenocheir sinuata, new species

Figs. 53b-58

Hybaphe tersa (nee Cook, 1904): Causey, 1954fl:222, fig. 1; 1955:91.
Chamberlin and Hoffman, 1958 (in part):36. Buckett, 1964:8-9.

Type specimens — Male holotype and 3 male, 7 female, and 3
juvenile paratypes (UCD) collected by J. S. Buckett and M. R. Gardner,
20 December 1966, 2 mi (3.2 km) southwest of Dales, Tehama County,
California. One male and one female paratypes deposited at NCSM.

Diagnosis — Acropodite directed generally anteriad from coxa,
distal curvature sharp, well defined, distal part variably sinusoid, blade-
like throughout length.

Color in Life — Paranota variably red to orange, metaterga black
with concolorous red to orange bands along caudal margins or middorsal
semilunar blotches.

Holotype — Length 27.6 mm, maximum width 9.5 mm, W/L ratio
34.4%, D/W ratio 33.7%.

Epicranial suture strong, distinct, terminating in interantennal
region. Width across genal apices 2.7 mm, interantennal isthmus 1.2
mm. Antennae relatively long and broad, reaching back to midlength
of 4th tergite; relative lengths of antennomeres 2>3>6>4>5>1>7; 6th
antennomere with microsensilla on outer distal margin. Genae with
shallow central impressions. Facial setae as follows: epicranial 2-2,
interantennal, subantennal, and frontal not detected and presumed absent,
genal 1-1, clypeal about 8-8, labral about 12-12.

Collum broad, ends terminating slightly below those of succeeding
tergite. Tergites smooth, polished. Paranota well developed throughout
body, broadest on anterior segments, becoming slightly narrower caudad,
moderately depressed, continuing slope of dorsum, anterior corners
with small but distinct denticle on segments 2-4, broadly rounded on
remaining segments, caudolateral corners angular on 2-3, blunt on 4-
5, rounded on remaining tergites, becoming progressively angled caudad
in caudal half of body. Peritremata moderately distinct, moderately
elevated above paranoial surfaces; ozopores located caudal to midlength,
opening sublaterad.

Sides of metazonites granular, with distinct, irregularly serrate
to jagged ridges subtending coxae through segment 10, becoming
progressively indistinct thereafter. Gonapophyses moderately long,
apically rounded. Pregonopodal sterna glabrous; 5th sternum with two
low, paramedian knobs between 4th legs, much shorter than widths of
adjacent coxae, and lower, flattened elevated areas between 5th legs;
6th sternum strongly depressed between both leg pairs to accommodate

180 Rowland M. Shelley

curvatures of telopodites. Postgonopodal sterna glabrous, flat, and
unmodified, with at most only shallow transverse grooves between
leg pairs, caudal margins gently curved. Coxae without projections;
prefemora of legs on segments 9-18 with ventrodistal spines, longer
and more distinct on segments 11-15; tarsal claws gently curved.
Hypoproct broadly rounded, slightly extended in midline; paraprocts
with margins strongly thicked.

Gonopodal aperture ovid, without indentations, 1.4-mm wide and
0.7-mm long at midpoint, anterior margin flush with metazonal surface,
sides elevating caudad and continuing onto caudal margin, latter flared
slightly but not extending caudad. Gonopods in situ (Fig. 53b) with
acropodites extending forward in parallel arrangement, overhanging
anterior margin of aperture, apices curling laterad then mediad. Gonopod
structure as follows (Figs. 54-55): Coxae moderate-size, connected
by membrane only, without trace of sclerotized sternal remnant; with
2 macrosetae above cannula and dense cluster of a dozen or so setae
on protuberance below latter. Prefemur moderately long and broad,
with short prefemoral process arising near midlength on anterior side,
latter extending to just beyond level of tip of acropodite, broadest
basally, sides narrowing and tapering to subacuminate tip on lateral
margin, directed toward midpoint of arc of acropodite. Latter arising
from prefemur at narrow constriction, relatively long, blade-like for
most of length, tapering smoothly and continuously, more so distad,
to subacuminate tip, in form of broad open arch, flattened at highest
point, curving broadly anteriad at 1/3 length and bent more strongly
dorsad at 2/3 length, distal part curving in markedly bisinuate fashion.
Prostatic groove arising in pit in prefemur, running along medial
surface of latter and crossing to lateral side at base of acropodite
returning to inner surface at peak of arch and continuing to terminal
opening.

Male Paratypes — In two individuals the acropodites cross in situ
at the midline instead of lying parallel, and in one male the prefemoral
process is shorter in relation to the acropodite.

Female paratype — Length 29.0 mm, maximum width 4.5 mm,
W/L ratio 15.5%, D/W ratio 82.2%. Agreeing essentially with males
excepting the presence of interantennal, subantennal, and frontal setae,
1-1 each, and paranota more strongly depressed, imparting more vaulted
appearance to body. Cyphopod aperture encircling 2nd legs, sides and
caudal margin flush with metazonal surface, elevated only midline.
Cyphopods in situ lying transversely in aperture, common surface
visible in opening. Valves (Fig. 58) relatively small, unequal, anterior
valve larger, situated nearly entirely on top of receptacle, slightly

The Chonaphini 181

distended mediad but not subtending distinct cavity, with numerous
long hairs. Receptacle large, alate, nearly completely enveloping valves,
extending halfway up anterior and caudal valvular surfaces, with long
hairs arising from ventral margin primarily from medial side. Operculum,
moderate-size, closely appressed to lateral side of valves.

Variation — The prefemoral process is noticeably shorter in the
male from El Dorado County and rises to a peak in the midline rather
than on the outer or lateral margin. The acropodite therefore terminates
well above the latter in these individuals, and the distal bend is more
gradual, thereby imparting a somewhat sigmoid pattern to the distal
curvature rather than sinuate. The sinuate condition is noticeably dimin-
ished in males from west of Burney, Shasta County, and Richardson
Grove, Humboldt County (Figs. 56-57).

Ecology — The specimens that I collected in El Dorado County
were encountered under wet hardwood litter beside a stream in a cove
along California highway 49. Those from Shasta County were found
under deciduous litter, mostly poison oak, on a bank above the Sacra-
mento River. The male from Butte County was found under a pine log
in a pine/fir association near a creek. According to Buckett and Gardner
(1968), S. sinuata, cited as Hybaphe sp., occurs in association with
luxuriant vegetation near the bottom of the south wall of the American
River Canyon, Placer County, and is thus ecologically segregated
from Wamokia discordis Buckett and Gardner, which occurs high on
the canyon wall, where the slope is steeper, the soil leached, and the
vegetation thin.

Distribution — The northern periphery of the Sacramento Valley
in Shasta County to the Pacific Ocean in Humboldt County and the
foothills of the Sierra Nevada Mountains in El Dorado County, a
distance of about 147 mi (236 km) north/south and 154 mi (246 km)
east/west. Specimens were examined as follows:

CALIFORNIA: Shasta Co., W of Burney, Moose Camp. Hatchet
Cr., M, 3F, 14 May 1966, A. A. Grigarick (UCD); along McCloud R.,
exact location unknown, 2M, F, 4 May 1929, E. C. Van Dyke (UCD);
Project City N of Redding, M, 10 December 1982, B. Miller (SDMNH);
Redding, along Sacramento R. trail, 2M, 3F, juv., 28 April 1991, R.
M. Shelley (NCSM); 21 mi (33.6 km) W Redding, F, 21 December
1966, J. S. Buckett, M. R. Gardner (UCD); and Inwood, 4 mi (6.4
km) NW Shingletown, F, 20 December 1966, J. S. Buckett, M. R.
Gardner (UCD). Humboldt Co., Richardson Grove St. Pk., M, 15 May
1966, H. Wilson (UCD). Tehama Co., 2 mi (3.2 km) S Dales, off CA
hwy. 36, 5M, 8F, 3 juvs., 20 December 1966, J. S. Buckett, M. R.

182

Rowland M. Shelley

Figs. 59-63. Selenocheir arcuata. 59, gonopods in situ, ventral view
of paratype. 60, left gonopod of holotype, medial view. 61, the same,
lateral view. 62, distal extremity of acropodite of male from 2 mi
(3.2 km) N Pashkenta, Tehama County, California, medial view. 63,
left cyphopod of female paratype, caudal view. Abbreviations as in
Figs. 2-8. Scale line for Fig. 59 = 1.00 mm; line for other Figs. =
1.60 mm for 60-61, 1.33 mm for 62, 1.00 mm for 63.

The Chonaphini 183

Gardner (UCD) TYPE LOCALITY; and 22 mi (35.2 km) W Red Bluff,
2M, 3F, 22 December 1966, J. S. Buckett, M. R. Gardner (UCD).
Butte Co., Forest Ranch, M, 27 April 1991, R. M. Shelley (NCSM);
and Chico, 4M, 3F, 7 May 1968, T. Komo, R. Wilkey, W. Wiard
(UCD). Placer Co., 1 mi (1.6 km) E Auburn, 4M, 28 January 1965,
M. R., R. C, J. L., B. W., and K. B. Gardner (UCD). El Dorado Co.,
cove along CA hwy. 49, 1 mi (1.6 km) S Placer Co. line, 4M, juv., 27
April 1991, R. M. Shelley (NCSM); and nr. confluence of North &
Middle Forks, American R., M, 26 March 1965, J. S. Buckett, M. R.
Gardner (UCD).

The following literature records of Hybaphe tersa from Shasta
County, in close proximity to each other, are believed to refer to S.
sinuata and are indicated by the open star in Figure 68. They are the
basis for Chamberlin and Hoffman's citation (1958) of Shasta County
for H. tersa and for Hoffman's citation (1979) of California for Hybaphe.
The Placer County record is of "an undescribed species of Hybaphe''
(Buckett and Gardner 1968).

CALIFORNIA: Shasta Co., Mt. Brock (Causey 1954fl); Low Pass
Cr. (Causey 1954^, 1955, Buckett 1964); and Madison Cr. (Causey
1955, Buckett 1964). Placer Co., American River Canyon near Auburn
(Buckett and Gardner 1968).

Remarks — Buckett and Gardner (1968) also recognized that this
species occurs in Shasta and Tehama counties.

Selenocheir arcuata, new species
Figs. 59-63

Type specimens — Male holotype and 9 male and 2 female paratypes
(UCD) collected by C. Smith, J. Clover, and F. Ennik, 15 May 1972,
at Black Rock Camp along Mill Creek, 18.8 mi (30.0 km) northeast
of Red Bluff, Tehama County, California. One male paratype deposited
in NCSM.

Diagnosis — Acropodite blade-like basally, becoming subacicular
apically, directed anteromediad, subtending broad, poorly defined arc,
distal curvature broad, indistinct, distal extremity either uncinate or
with short bisinuate section.

Color in life — Unknown.

Holotype — Length 30.8 mm, maximum width 4.8 mm, W/L ratio
15.6%, D/W ratio 68.8%.

Somatic features agreeing with those of S. sinuata, with following
exceptions:

Width across genal apices 3.2 mm; interantennal isthmus 0.9
mm. Relative lengths of antennomeres 2>3>6>4>5>1>7. Genae with

184 Rowland M. Shelley

distinct central impressions. Facial setae as follows: epicranial 2-2,
interantennal and subantennal not detected and presumed absent, frontal
1-1, genal 2-2, clypeal about 12-12, labral about 20-20.

Caudolateral corners of paranota angular on segments 2-3, blunt
on 4-8, rounded on remaining tergites, becoming progressively angled
caudad in caudal half of body.

Sides of metazonites with irregularly serrate ridges subtending
coxae through segment 7, becoming progressively indistinct thereafter.
5th sternum with very low, indistinct lobes between 4th legs, flat and
unmodified between 5th; 6th sternum with two distinct, widely separated
subtriangular lobes subtending anterior (6th) coxae, depressed centrally
to accommodate telopodites. Prefemoral spines present on legs on
segments 11-18, short and indistinct.

Gonopodal aperture broadly rounded, without indentations, 1.2-
mm wide and 0.9-mm long at midpoint, anterior margin flush with
metazonal surface, sides elevating strongly caudad and continuing
onto caudal margin, latter flared slightly and extending slightly caudad
in midline. Gonopods in situ (Fig. 59, of paratype) with acropodites
leaning laterad then curving broadly anteromediad, tips nearly overlapping,
extending well beyond anterior margin of aperture and over 6th sternum.
Gonopod structure as follows (Figs. 60-61): Coxae moderate-size, connected
by membrane only, with 2 macrosetae above cannula, dense cluster of
innumerable setae below cannula, and 4-5 setae above latter cluster.
Prefemur moderately long and broad, with short prefemoral process
arising anteriad, latter narrow basally, extending distad a short distance,
then expanding to shallowly divided tip, medial side longer and more
acute. Acropodite arising imperceptibly from prefemur, without con-
striction, long and narrowly blade-like for most of length, becoming
subacicular distad, in form of very broad, open arch, gently curved at
highest point, curving broadly anteriad basally and continuing at
essentially same curvature to just before tip, where it bends suddenly
in uncinate fashion, narrowing thereafter to subacuminate tip. Pro-
static groove arising in pit in prefemur, running along medial side of
latter and curving onto lateral side of acropodite, angling back onto
medial surface at level of base of uncinate curve and continuing to
terminal opening.

Male paratypes — The male paratypes agree closely with the
holotype except for the prefemoral process, which is slightly different
on nearly every individual. Its length and the apical division vary, as
the terminations are more unequal than in the holotype. In one paratype.

The Chonaphini 185

the lateral termination is absent, and the medial one arises from an
otherwise flat surface.

Female paratype — Length 35.1 mm, maximum width 5.2 mm,
W/L ratio 14.8%, D/W ratio 71.2%. Agreeing closely with males in
somatic features except paranota more strongly declined, creating appear-
ance of more highly arched body. Cyphopod aperture with sides and
caudal margin elevated above metazonal surface, former leaning inward
over opening, latter rising to peak in midline. Cyphopods in situ
lying transversely in aperture, common surface visible in opening.
Valves (Fig. 63) relatively small, subequal, situated nearly entirely
on top of receptacle, without lobes, moderately hirsute. Receptacle
very large, alate, situated to medial side but still nearly directly beneath
valves and nearly enveloping latter, extending well up both anterior
and caudal valvular surfaces, anterior "wing" larger than caudal, ex-
tending nearly completely up side of anterior valve, with numerous
long hairs arising from sides and ventral margins. Operculum relatively
small, closely appressed to lateral surface of valves.

Variation — The non-typical male from Tehama County and that
from Mendocino County have a slight bisinuate stretch instead of the
uncinate curve (Fig. 62), and the prefemoral process expands distad
and terminates in a central point. There is no suggestion of bifuration;
the distal expansion imparts a general clavate shape to the structure.
In the males from Humboldt County, the acropodite is directed strongly
submediad.

Ecology — Unknown; the vial labels do not provide habitat
information. The sample from Dead Mule Spring was at an altitude of
about 5,150 ft; those from Lake County were collected at 3,910 ft. by
ultraviolet light.

Distribution — The northern Sacramento Valley and Coast Range
of California, an area of about 84 mi (134 km) east/west and 119 mi
(190 km) north/south. Specimens were examined as follows:

CALIFORNIA: Tehama Co., Black Rock Camp along Mill Cr.,
18.8 mi (30 km) NE Red Bluff, lOM, 2F, 15 May 1972, C. Smith, J.
Clover, F. Ennik (UCD) TYPE LOCALITY; and Dead Mule Spring,
along unnamed rd., 2 mi (3.2 km) N Paskenta/Covelo rd., M, F, 29
August 1972, H. B. Leech (CAS). Mendocino Co., 6 mi (7.6 km) N
Potter Valley, M, 28 January 1967, J. S. Buckett, M. R. Gardner
(UCD). Humboldt Co., 5 mi (8.0 km) N Willow Creek (town), Tish
Tang Rec. area, 5M, 21 February 1976, and 2M, 3F, 20 December
1979, A. K. Johnson (NCSM). Lake Co., nr. clear Lake, N side Bartlett

186

Rowland M. Shelley

Figs. 64-67. Selenocheir directa. 64, gonopods in situ, ventral view
of paratype. 65, left gonopod of holotype, medial view. 66, telopodite
of the same, lateral view. 67, left cyphopod of female paratype, cau-
dal view. Abbreviations as in Figs. 2-8. Scale line for Fig. 64 = 1.00
mm; line for other Figs. = 1.00 mm for 65-66, 0.80 mm for 67.

Mtn. Summit, Mendocino Nat. For., M, F, 28 April 1969, F. Emmik,
M. Knudson (UCD).

Selenocheir directa, new species

Figs. 64-67 "' ■ •''
Type specimens — Male holotype and 20 male, 10 female, and 3
juvenile paratypes (NMNH) collected by A. K. Johnson, 22 December
1977, at Patrick Creek Campground, along US highway 199, 7 mi

The Chonaphini 187

(11.2 km) northeast of Gasquet, Del Norte County, California. Seven
male and 4 female paratypes deposited in NCSM.

Diagnosis — Acropodite directed strongly submediad, blade-like
throughout length, narrowing slightly distad, curving distinctly downward
or dorsad distally, without trace of sinusoid curvature.

Color in life — Unknown.

Holotype — Length 24.6 mm, maximum width 3.9 mm, W/L ratio
15.9%, D/W ratio 76.1%.

Somatic features agreeing with those of S. sinuata, with following
exceptions:

Width across genal apices 2.8 mm; interantennal isthmus 0.9 mm.
Relative lengths of antennomeres 2>3>6>4>5>1>7. Facial setae as follows:
epicranial 2-2, interantennal 1-1, subantennal 1-1, frontal 1-1, genal
2-2, clypeal about 13-13, labral about 18-18.

Collum terminating at same level as succeeding tergite. Caudolateral
corners of paranota angular on segments 2-3, blunt on 4-6, rounded
on remaining tergites, becoming progressively angled caudad in caudal
half of body.

Sides of metazonites with irregularly serrate ridges subtending
coxae through segment 9, becoming progressively indistinct thereafter.
5th sternum with low, rounded elevated areas between 4th legs, slightly
depressed between 5th; 6th sternum depressed between both legs to
accommodate telopodites. Prefemoral spines present on legs of segments
13-18, short and indistinct.

Gonopodal aperture broadly rounded, without indentations, 1.2-
mm wide and 0.6-mm long at midpoint, anterior margin flush with
metazonal surface, sides elevating slightly caudad and continuing onto
caudal margin, latter extended slightly caudad. Gonopods in situ (Fig.
64, of paratype) with acropodites curving broadly submediad, tips nearly
overlapping, extending well beyond anterior margin of aperture and
overhanging 6th sternum. Gonopod structure as follows (Figs. 65-66):
Coxae moderately-large, nearly equal to telopodite in overall bulk,
connected by membrane only, with 2 macrosetae above cannula and
moderate cluster of setae below. Prefemur relatively short and broad,
with moderately long prefemoral process arising anteriad, about half
as long as acropodite, broad basally, bent slightly dorsad and twisted
at 2/3 length, broad apically. Acropodite arising at slight constriction
of prefemur, moderately broad basally, directed submediad, con-
figuration a moderately broad arch, curving broadly at midlength, sides
narrowing smoothly and continuously to subacuminate tip. Prostatic
groove arising in pit in prefemur, running along medial side of latter

188 Rowland M. Shelley

and curving onto lateral margin of acropodite, continuing to terminal
opening.

Male paratypes — Except for minor variation in the length, degree
of bend, and broadness of the tip, the male paratypes agree with the
holotype.

Female paratype — Length, 28.1 mm, maximum width 5.1 mm,
W/L ratio 18.1%, D/W ratio 74.5%. Agreeing essentially with males
in somatic features except paranota more strongly declined creating
appearance of more highly arched body. Cyphopodal aperture very
broad, sides and caudal margin elevated above metazonal surface,
latter rising to peak in midline. Cyphopods in situ lying transversely
in aperture, common surface visible in opening. Valves (Fig. 67)
moderately large, subequal, without distinct lobes, moderately hirsute,
situated nearly entirely on receptacle, nearly completely enveloped
by latter. Receptacle large, alate, situated to medial side but nearly
directly beneath (dorsal to) valves, extending nearly completely up
anterior side of latter, caudal "wing" shorter, with numerous long
hairs arising from ventral margin. Operculum relatively large, closely
appressed to lateral sides of valves.

Variation — Aside from differences in the length and apical
configuration of the prefemoral process, the nontypical males agree
closely with the holotype.

Ecology — Unknown.

Distribution — The adjacent corners of southwestern Oregon and
northwestern California, an area of about 78 mi (125 km) east/west
and 59 mi (94 km) north/south. Specimens were examined as follows:

OREGON: Curry Co., 1 mi (1.6 km) N Gold Beach, on opposite
bank of Rogue R., 2M, 30 January 1967, A. Jung (UCD). Jackson
Co., Shady Cove, M, 28 March 1972, J. Schuh (FSCA); 10 mi (16
km) E, 6 mi (9.6 km) N Gold Hill, along OR Hwy. 234, M, 22
January 1972, E. M. Benedict (WAS); and 10 mi (16 km) NW Central
Point, along OR hwy. 234, M, 22 January 1972, E. M. Benedict (WAS).

CALIFORNIA: Del Norte Co., 1 mi (11.2 km) NE Gasquet,
Patrick Cr. Cpgd. along US hwy. 199, 28M, 14F, 3 juvs., 22 December
1977, and 3M, 3F, 21 December 1979, A. K. Johnson (NMNH, NCSM)
TYPE LOCALITY.

Remarks — Selenocheir directa bridges the anatomical gaps between
the Chonaphini and the Harpaphini, and future workers may conclude
that this necessitates synonymizing the tribes. The coxa in S. directa
is larger in comparison to the telopodal elements than in any of the
congeners or any other chonaphines, and the acropodite is directed

The Chonaphini

189

Fig. 68. Distributions of chonaphine genera and species in western
North America, dots, Chonaphe armata; triangles, C. remissa', horizon-
tal oval (southwestern Oregon), C evexa; vertical oval (northeastern
Washington), C. schizoterminalis; squares, Montaphe elrodi; asterisk (central
Washington), M. paraphoena; vertical rectangles, T. levii; horizontal
rectangles (Idaho), Metaxycheir prolata; stars, Selenocheir sinuata; vertical
half-shaded dots, S. arcuata; horizontal half-shaded dots, S. directa.
Open symbols denote literature records deemed reliable; the larger,
open star signifies three literature sites clustered in the same area.

190

Rowland M. Shelley

Fig. 69. Distributions of the Chonaphini, Semionellus, and S, placidus
in the eastern United States. Open symbols denote literature records
considered accurate.

strongly submediad on the coxa. These traits approximate those diagnostic
for the Harpaphini (Shelley 1993^).

DISTRIBUTION

Species and genera — As shown in Figure 70, the western chonaphine
species occupy mutually exclusive ranges, segregated from each other,
aside from the region in and around Idaho, where C. armata and
Montaphe elrodi overlap broadly; the former also overlaps Metaxycheir
prolata, and the latter completely envelops C. schizoterminalis. The
sympatry in California between S. sinuata and arcuata may be an
artifact because so few records are available from the Coast Range in
Lake, Mendocino, Trinity, and Humboldt counties. The northernmost,
Humboldt County, population of S. arcuata may be allopatric and
detached from the rest of the species, which occurs generally to the
south-southwest of S. sinuata. I have assumed that these populations
connect, which accounts for the range overlap with S. sinuata; if they
do not overlap, S. arcuata and sinuata are parapatric. Both S. sinuata
and Montaphe elrodi show southward projecting fingers from their

The Chonaphini

191

Fig. 70. Comparative distributions of chonaphine species in western
North America. 1, C. armata; 2, C. remissa; 3, C. evexa; 4 C.
schizoterminalis; 5, Montaphe elrodi; 6, M. paraphoena; 7, T. levii;
8, Metaxycheir prolata; 9, 5. sinuata; 10, S. arcuata; 11, S. directa.

192 Rowland M. Shelley

main ranges; the former is thus the southernmost chonaphine in the
West, and the latter, the northern- and eastern-most.

Chonaphe armata and S. placidus are the species with the most
interesting distributions; they occur in five and four areas, respectively
(Figs. 68, 69). In both cases, an ancestral range has fragmented, leaving
allopatric populations that have undergone little anatomical divergence
and are hence conspecific. Additionally, the distances between the
five populations of C. armata are much greater than those between it
and C. remissa, which is effectively parapatric, being segregated only
by the Columbia River and the spine of the Cascade Mountains. Chonaphe
remissa replaces C. armata west of the Cascades in Washington, as
the latter occurs only on their eastern slope, but in Oregon, C. remissa
is absent, and C. armata occurs only west of the Cascades, in the
lower Willamette Valley and eastern foothills of the Coast Range as
far south as Benton County. Chonaphe evexa is thus an allopatric,
southern species, detached from the main generic range by some 125
mi (200 km); C. schizoterminalis, in northeastern Washington, is essen-
tially parapatric with a population of C. armata.

The picture at the generic level (Fig. 71) is identical to that at
the specific, with the exception of Selenocheir in California and Chonaphe
in Washington. Selenocheir covers a broad area in southwestern Oregon
and northern California with a finger extending southward through
the Sierra Nevada foothills. In Chonaphe, the ranges of C. armata in
western Washington and Oregon join with that of C. remissa to form
a large area with a finger extending southward down the eastern slope
of the Coast Range and the western Willamette Valley. Ranges are
also mutually exclusive except in Idaho and environs, where they
overlap as in the species.

RELATIONSHIPS

Tribal — What seemed to be a straightforward study from such
specialized genera as Chonaphe and Semionellus, with their apomorphic
acicular acropodites and elaborate prefemoral processes, rapidly became
complex as I pondered forms like T. levii, Montaphe paraphoena, and
Metaxycheir prolata, and the meaning and significance of such inconsis-
tencies as the presence of lateral versus medial sternal lobes, and the
presence or absence of a sternum. Because the acropodites of its
species are as narrowly blade-like or acicular as those of T. levii and
M. prolata, the question arose as to whether Selenocheir is also a
chonaphine, but with a short, instead of a long, prefemoral process.
No other established tribe can accommodate Selenocheir, and a monobasic
category would be undefinable, so I place it in the Chonaphini. Selenocheir

The Chonaphini

193

Fig. 71. Comparative distributions of chonaphine genera in western
North America. 1, Chonaphe; 2, Montaphe; 3, Tubaphe; 4, Metaxycheir;
5, Selenocheir.

194 Rowland M. Shelley

Selenocheir Chonaphe Semionellus Metaxycheir Tubaphe Montaphc

Figure 72. Relationships in the Chonaphini.

sinuata was previously misidentified as Isaphe tersa (Cook), a member
of the Harpaphini (Causey 1955, Buckett 1964), because of key similarities
between Selenocheir, particularly S. directa, and Isaphe (= Hybaphe).
In /. tersa, as in all harpaphines, the coxa is larger than the telopodal
elements in overall bulk, and the latter is oriented transversely on the
prefemur so as to project directly mediad; the telopodal elements
therefore extend directly toward the viewer in medial aspect, so as to
provide a head-on perspective rather than a profile (Shelley 1993^).
In S. directa, the coxa is proportionately the largest in the genus,
being only slightly smaller than the telopodite, and the telopodal
elements are directed anteromediad, or about midway between a head-
on view, as in the Harpaphini, and a profile, as in chonaphine genera
like Tubaphe and Montaphe. Consequently, S. directa, and to a lesser
extent the entire genus Selenocheir, span the anatomical gaps between
the Chonaphini and Harpaphini, and if future workers conclude that
they should be merged, the former name holds priority. As the Asiatic
representatives of the Harpaphini have not been reviewed, and their
characteristics are poorly known in contrast to the American forms, I
believe a decision on this potential merger should be deferred until
the Oriental genera are better understood. Suffice it to say for now
that the Chonaphini is closely related to the Harpaphini, and that the
component taxa of both groups demonstrate a broad range of expressions
of a suite of anatomical features.

Generic and Specific — Two obvious chonaphine lineages are
apparent — one with expanded, laminate prefemoral processes and acicular
acropodites {Chonaphe + Semionellus), and one in which the prefemoral
process is narrow and blade-like, and the acropodite broader and less

The Chonaphini 195

acicular (Montaphe + Tubaphe + Metaxycheir). Selenocheir, with its
short, plesiomorphic prefemoral process, represents a separate, sister
line. There are no intermediate forms with partly expanded prefemoral
processes or ones of intermediate lengths, an anatomical gap that
suggests age and the extinction of intermediate forms. Age is also
indicated by the substantial geographical gaps — between intraspecific
populations of both S. placidus and C. armata, between species of
Chonaphe, between Metaxycheir and Tubaphe, and most especially
between the western faunal regions and Semionellus. Chonaphe evexa
and schizoterminalis share an angular elevation on the dorsal face of
the prefemoral process distal to the shield, this being a low ridge in
the former and a laminate flap in the latter, and appear to be peripheral
relicts of an early lineage that has been supplanted by the younger
branch leading to C. armata and remissa, which possesses the thicker,
distal projection at the location of the ridge. Although evolving more
recently, the C. armata + remissa lineage is old enough to have undergone
substantial fragmentation, with lacunae between the populations of C.
armata.

Regarding the forms with narrow, blade-like prefemoral processes,
the restricted distributions of Metaxycheir prolata and T. levii also
suggest age, and the former in particular seems to hold relict status
because it is known only from 4 samples and 6 adults despite inhabiting
the most heavily sampled area of Idaho. Montaphe elrodi, occupying
a broad, cohesive area in the western interior, evolved more recently,
and M. paraphoena, an enigmatic species, is a possible relict from an
intermediate line between the narrow, blade-like and expanded, laminate
forms. I am unable to resolve the relationships between these genera
and show an unresolved trichotomy in Figure 72.

My overall impression of the Chonaphini is thus one of age. The
geographical and anatomical lacunae contrast markedly with the situa-
tions in the Eastern and Meso-American tribes Apheloriini, Nanariini,
Pachydesmini, and Rhysodesmini, which lack such gaps and evolved
more recently (Shelley and Whitehead 1986). The Chonaphini appears
to be the second oldest xystodesmid assemblage next to the Orophini,
which has an even larger geographical hiatus between its only nearctic
representative, Orophe, in Idaho and Montana, and the only other
known genera Pamelaphe and Kiulinga, in China (Hoffman 1964,
Shelley 1993^). Perhaps the Xystodesmidae, or more properly the
subfamily Xystodesminae, experimented at an early age with long,
twisted gonopodal telopodites and ones with long, slender, comparatively
simple acropodites and complex prefemoral processes, before settling

196 Rowland M. Shelley

on the pattern of complex, elaborate acropodites and relatively short,
simple prefemoral processes. The descendants of such early experimenta-
tions survive today in discontinuous ranges as the tribes Orophini and
Chonaphini, respectively.

ACKNOWLEDGEMENTS— Field work for this study was
supported by grant number 4495-91 from the National Geographic
Society; anatomical examinations took place in January 1992, while I
was working at the NMNH on a Mid-Career Fellowship from the
National Science Foundation. Access to the types of C. cygneia,
michigana, patriotica, and serratus (holotype), Leptodesmus borealis
and L. (C.) elrodi, and nontypical material at the NMNH, was courtesy
of J. A. Coddington; the type of T. levii and nontypical material at
the AMNH were loaned by N. I. Platnick; the holotype of Metaxycheir
prolata (UCD) was donated by L. S. Kimsey; paratypes of C. serratus
and other samples at the FSCA were provided by G. B. Edwards; and
syntypes of Polydesmus armatus (PMNH) were loaned by C. L. Reming-
ton and R. J. Pupedis. Samples from the following collections were
provided by the indicated curators and collection managers: BYU, R.
W. Baumann; CAS, W. J. Pulawski; CMN, P. F. Frank and J. A.
Fournier; EIL, R. C. Funk; FMNH, D. A. Summers; MCZ, H. W.
Levi; MPM, J. P. Jass; RBCM, R. A. Cannings; ROM. D. C. Darling;
SDMNH, D. K. Faulkner; TMM, J. R. Reddell; UBC, S. G. Cannings;
UCD, the late R. O. Schuster; UMN, R. Hozenthal; UWBM, R. Crawford;
UWY, S. R. Shaw; VMNH, R. L. Hoffman; WSU, R. S. Zack; and
WU, D. R. Breakey. My colleague W. A. Shear provided valuable
samples from his personal collection, and R. Crawford (UWBM) suggested
the name, paraphoena, for the new species of Montaphe. Figures 2-
67 were prepared by R. G. Kuhler, NCSM scientific illustrator; Cathy
Wood graciously handled repeated word processing chores.

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poda: Diplopoda, Chilopoda). Brimleyana 18:1-13.

Shelley, R. M. 1993c. A new xystodesmid milliped genus and
species from Washington and Oregon (Polydesmida). Myria-
podologica 2:91-100.

Shelley, R. M. 1993^. The milliped genus Isaphe Cook (Polydesmida:
Xystodesmidae). Canadian Journal of Zoology 71:1161-1168.

Shelley, R. M. 1993e. The milliped genus Orophe Chamberlin
(Polydesmida: Xystodesmidae). Insecta Mundi 7:175-182.

Shelley, R. M., and D. R. Whitehead. 1986. A reconsideration of
the milliped genus Sigmoria, with a revision of Deltotaria and
an analysis of the genera in the tribe Apheloriini (Polydesmida:
Xystodesmidae). Memoirs of the American Entomological Soci-
ety Number 35.

Verhoeff, K. W. 1941. Uber Gruppen der Leptodesmiden and neues
System der Ordo Polydesmoidea. Archiv fur Naturgeschichte
10:399-415.

Williams, S. R., and R. A. Hefner. 1928. The millipedes and
centipedes of Ohio. Bulletin of the Ohio Biological Survey Num-
ber 18.

Wood, H. C. 1864. Descriptions of new species of North Ameri-
can Polydesmidae. Proceedings of the Academy of Natural Sci-
ences of Philadelphia 16:6-10.

Wood, H. C. 1865. The Myriapoda of North America. Proceed-
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200 Rowland M. Shelley

Wood, H. C. 1867. Notes on a collection of California Myriapoda,
with the descriptions of new eastern species. Proceedings of
the Academy of Natural Sciences of Philadelphia 19:127-130.

Received 19 May 1993
Accepted 17 February 1994

201

DATE OF MAILING

Brimleyana 19 was mailed on 25 May 1994.

ERRATA

In Brimleyana Number 19, page 51, the numbers 10 and 11 were
inadvertently reversed in the caption to figure 32. Number 10 actually
denotes Scytonotus virginicus intergrades, and number 11 is for S. v.
virginicus. Numbers are correct on the figure.

202

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BRIMLEYANA NO. 20, JUNE 1994
CONTENTS

Review of Biologically Significant Caves and their Faunas in Florida and South

Georgia. Richard Franz, Judy Bauer, and Tom Morris 1

The Chonaphini, a Biogeographically Significant Milliped Tribe in Eastern and
Western North America (Polydesmida: Xystodesmidae).
Rowland M. Shelley Ill

Miscellany 201