Freshwater Triclads (Turbellaria)

of North America,

IX: The Genus Sphalloplana

ROMAN KENK

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 246
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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 246

Freshwater Triclads (Turbellaria)

of North America,

IX: The Genus Sphalloplana

Roman Kenk

ISSUED

2 1977,

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRACT

Kenk, Roman. Freshwater Triclads (Turbellaria) of North America, IX: The

Genus Sphalloplana. Smithsonian Contributions to Zoology, number 246, 38

pages, 62 figures, 1 table.—A revision of the North American species of the

genus Sphalloplana is presented. Apart from the nine species already known

(S. percoeca, S. georgiana, S. kansensis, S. virginiana, S. weingartneri, S. buchanani, S. pricei, S. hubrichti, and S. mohri), seven new species are introduced and

described: 5. evaginata, S. californica, S. culveri, S. consimilis, S. subtilis, S.

holsingeri, and S. chandleri.

OFFICIAL PUUJCATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Kenk, Roman, 1898-

Freshwater triclads (Turbellaria) of North America, IX, the genus Sphalloplana.

(Smithsonian contributions to zoology ; no. 246)

Bibliography: p.

1. Sphalloplana. 2. Platyhelminthes—North America. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 246.

QL1.S54 no. 246 [QL391.P7] 591'.08s [595M23] 76-44832
Contents

Page

Introduction 1

Sphalloplana (Sphalloplana) percoeca (Packard, 1879) 3

Sphalloplana (Sphalloplana) georgiana Hyman, 1954 5

Sphalloplana (Sphalloplana) evaginata, new species 6

Sphalloplana (Sphalloplana) kansensis Hyman, 1945 8

Sphalloplana (Sphalloplana) californica, new species 8

Sphalloplana (Sphalloplana) culveri, new species 10

Sphalloplana (Sphalloplana) consimilis, new species 11

Sphalloplana (Sphalloplana) subtilis, new species 12

Sphalloplana (Speophila) virginiana Hyman, 1945 14

Sphalloplana (Speophila) holsingeri, new species 14

Sphalloplana (Speophila) weingartneri Kenk, 1970 17

Sphalloplana (Speophila) buchanani (Hyman, 1937) 17

Sphalloplana (Speophila) pricei (Hyman, 1937) 18

Sphalloplana (Speophila) hubrichti (Hyman, 1945) 20

Sphalloplana (Speophila) chandleri, new species 21

Sphalloplana (Speophila) mohri Hyman, 1938 23

Literature Cited 25

Figures 28

iii

Freshwater Triclads (Turbellaria)

of North America,

IX: The Genus Sphalloplana

Roman Kenk

Introduction

The genus Sphalloplana was established by

Beauchamp (1931:321) for a white, blind planarian

from Mammoth Cave in Kentucky, reported earlier

by Packard (1879:142) from the same locality under

the name of Dendrocoelum percoecum, new species.

Beauchamp stressed the fact that the species has

the external appearance of a dendrocoelid, particularly by the undulated body margins of the

preserved specimens. Anatomically, however, it

could not be placed in the family Dendrocoelidae,

as its internal pharyngeal musculature consisted of

two distinct layers, an inner circular and an outer

longitudinal one, while in the dendrocoelids it is

composed of one layer of intermingled circular and

longitudinal muscle fibers.

Additional species of Sphalloplana were described

by Hyman in several papers, all collected in caves

or springs and placed into a new family, Kenkiidae.

Hyman (1937:473) also introduced a related genus,

Speophila, which differed from Sphalloplana by

having a more highly developed adhesive organ. In

a later paper, however, she (1945:481) stated that

"it is probable that all gradations in the develop

ment of the adhesive organ exist among the Kenkiidae, and hence that eventually it will be impossible to draw any definite line between the two

genera. I therefore anticipate that when our

Roman Kenk, Department of Invertebrate Zoology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

cavernicolous planarians are sufficiently known,

Speophila will become a synonym of Sphalloplana."

This union of the two genera was finally proposed

by Mitchell (1968:614-615) and has been accepted

by Kawakatsu (1969:76), Carpenter (1970), and

myself (1972:13).

The genus Sphalloplana may be defined as Kenkiidae, in which the internal pharyngeal muscle

zone consists of two layers arranged as in the

Planariidae, an internal layer of circular fibers

surrounded by a layer of longitudinal muscles. In

the other two genera of the family, the outer zone

of the internal pharyngeal muscles has an intermingled network of longitudinal and circular fibers.

This is known for the genus Macrocotyla (Kenk,

1975:326). Kawakatsu, who had an opportunity of

examining Kenkia rhynchida Hyman from its typelocality, informed me (pers. comm.) that Kenkia

has the same muscular condition as Macrocotyla.

Sphalloplana may be divided into subgenera as

proposed by Carpenter (1971:1284): subgenus

Sphalloplana (type-species, Dendrocoelum percoecum Packard) with weak adhesive organ, and

subgenus Speophila (type-species, Speophila pricei

Hyman) with highly differentiated, deeply invaginated adhesive organ (with some gradations between

the two subgenera). The third subgenus proposed

by Carpenter, Polypharyngea (type-species, Sphalloplana mohri Hyman), based on the presence of

many pharynges, is of doubtful value, since polypharyngy occurs in other genera (Phagocata,

Crenobia) as a specific or even subspecific charac-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

teristic. Therefore I place S. mohri in the subgenus

Speophila.

Concerning the distinguishing characteristics of

the species of Sphalloplana, Carpenter (1970) assumes that they are subject to great variation.

Nevertheless, in worms preserved by a proper technique there is a great uniformity in the characters

of specimens from the same locality, with only little

variability. Kenkiid planarians are rather difficult

to preserve. The usual methods of relaxation (by

anesthetics) or extension (e.g., with diluted nitric

acid), which are appropriate for many planariids,

frequently result in great distortions of the internal

muscular organs (pharynx, penis, etc.). The same

is true for Carpenter's (1969) rapid freeze technique

that, however, preserves the external shape of the

animals beautifully (see also Kawakatsu and

Miyazaki, 1972:87-88, and the photographs in

Carpenter, 1970).

I have obtained the best and most uniform

results by instantaneous killing of the worms with a

hot fixative. A worm is placed in a small dish with

only a few drops of water. When it is well extended, a hot (almost boiling) fixative is rapidly

poured over it. I use as fixative a saturated (about

7%) solution of corrosive sublimate (HgCl2) in

water or physiological saline solution. After killing,

the fixative is acidulated with a few drops of

diluted acetic acid. After several hours, the specimen is washed in water for one hour, then transferred to increasing concentrations of ethyl alcohol.

The last traces of the sublimate are removed by

adding small quantities of tincture of iodine to the

alcohol.

The characters used for the differentiation of

the species of Sphalloplana are both external and

anatomical ones. Externally, the shape of the anterior end is very characteristic, some species

bearing anterolateral auricular projections, others

no auricles. The anterior border of the intestinal

area is either rounded or shows anterior extensions

along both sides, with a median recess, producing

the shape of the letter V, as is seen in some of the

species with well-developed adhesive organ. Anatomically, the location of the essentially prepharyngeal testes, whether dorsal, ventral, or both, is

important. In the copulatory apparatus, the shape

of the penis, the proportional size between the

penis bulb and penis papilla, the shape and histology of the penial lumen, and the location of the

mouth of the ejaculatory duct are to be noted. The

vasa deferentia in some species enter the penis bulb

anteroventrally or laterally, in others they curve

upward and enter the bulb posterodorsally. The

presence or absence of a distinct vagina on the

bursal duct and the location of the mouth of the

common oviduct are other distinguishing features.

Some species show a characteristic distribution of

infranucleate epithelia in the copulatory complex.

The present report deals chiefly with those

species of Sphalloplana that I have been able to

obtain alive. There are, in all, 19 species of the

genus reported in the literature, of which 15 occur

in the United States, two in Japan, and one each

in South Korea and Siberia. Five of the North

American species are considered to be identical and

one, S. hoffmasteri (Hyman), has been transferred

to the genus Macrocotyla (Kenk, 1975:333). The

Siberian species, S. ductosacculata Livanov and

Zabusova (1940), has the testicular zone extending

to the posterior end and may not belong to this

genus. One of the two undescribed Japanese species, S. sp. from the Yatsu-gadake Mountains

(Ichikawa and Kawakatsu, 1967:512), is a two-eyed,

darkly pigmented planarian and has not been

studied anatomically. In addition to the valid nine

species of Sphalloplana of the United States, seven

new species are described in the present paper.

A comparison of the American cavernicolous triclad fauna with that of Europe is very interesting

from the zoogeographical standpoint. The American Kenkiidae, split into many closely related

species, occupy the same ecological niches as do

the numerous subterranean species of Dendrocoelum and related genera in Europe (see particularly Beauchamp, 1932, and Gourbault, 1972).

The differentiation of both faunas apparently is

of relatively recent geological origin. There is no

kenkiid known to occur in Europe and no Dendrocoelum in the Americas.

ACKNOWLEDGMENTS.—My study of the genus

Sphalloplana would not have been possible without

the generous aid of many collaborators, particularly speleologists who kindly provided me with

materials from various caves. I wish to express to

them my deep appreciation of their valuable cooperation. It would be difficult to list all persons who

have sent me specimens. They are mentioned under

the collection data for the individual species. I

may name here only the most outstanding contrib-
NUMBER 246

utors: Dr. John R. Holsinger (Old Dominion

University), Mr. Jerry Lewis (Southern Illinois

University), Dr. Russell M. Norton (Yale University), Mr. Arnold Norden, and Mr. William W.

Torode. Dr. John C. Harshbarger (Smithsonian

Institution) kindly permitted me the use of his

laboratory facilities for the preparation of photographic illustrations. Thanks are due to the

American Museum of Natural History, which

kindly lent me some of L. H. Hyman's slides for

examination. I also wish to express my gratitude to

various publishers who permitted me to reproduce

some, in part copyrighted, figures from their publications: McGraw-Hill Book Company, American

Microscopical Society, Biological Society of Washington, and the publishers of American Midland

Naturalist.

The location of the type materials of the individual species is indicated by the following abbreviations: AMNH for American Museum of Natural

History, New York; and USNM (former United

States National Museum), for materials in National

Museum of Natural History, Smithsonian Institution, Washington, D.C.

Sphalloplana (Sphalloplana) percoeca

(Packard, 1879)

FIGURES 1, 11, 30, 45

Dendrocoelum percoecum Packard, 1879:141.

Fonticola percoecum.—Hyman, 1931:328.

Sphalloplana percaeca.—Beauchamp, 1931:321.

Dendrocoelum percaecum.—Giovannoli, 1933:622.

Sphalloplana percaeca.—Castle and Hyman, 1934:156.

Fonticola percaeca.—Hyman, 1935:345.

Sphalloplana percoeca.—Hyman, 1937:469.

Sphalloplana (Sphalloplana) percoeca.—Carpenter, 1971:1284.

Sphalloplana alabamensis Hyman, 1945:476.

TYPE MATERIAL.—Packard's original material

apparently is not preserved. Hyman's syntypes of

S. alabamensis, one whole mount and two slides

of sagittal sections (USNM 20639).

HISTORY.—The early history of the discovery and

description of the species has been reviewed by

Buchanan (1936:194). It was first depicted by

Packard (1879:141) in a small, rather primitive

illustration that was also reproduced in later publications of the same author (1880:141 and later

editions of his Zoology; 1888:28). Illustrations of

preserved specimens were furnished by Beauchamp

(1931:322) and drawings and photographs of living

animals by Buchanan (1936:196) and Carpenter

(1970, fig. 11). Descriptions of the species were

given by Packard (1888:28), Beauchamp (1931),

Buchanan (1936:195-196), Hyman (1937:470-471),

and Carpenter (1970:69-75). Hyman's description

of S. alabamensis is based on badly preserved materials and contains several inaccuracies. In the

following, only the essential characters of the

species will be discussed.

EXTERNAL FEATURES (Figures 1, 11).—Sphalloplana percoeca is a purely white species, without

any pigmentation. Mature animals generally are

about 10-12 mm long and about 2 mm wide, but

may reach a length of 16 mm. The anterior end

has a bulging frontal margin and conspicuous

rounded auricles projecting anterolaterally. The

situation of the adhesive organ is seen as an opaque

spot behind the center of the frontal margin. Behind the auricles, the lateral margins of the body

narrow to some extent, then widen again. Characteristic is the outline of the anterior border of the

intestinal area that is rounded and not forming the

two lateral extensions seen in some species with

more highly developed adhesive organs.

ANATOMY.—The adhesive organ (Figure 30) is a

shallow subterminal pit lined with an infranucleate

epithelium that is pierced by many eosinophilic

gland ducts and equipped with a system of muscle

fibers (see Beauchamp, 1931:323-325; Hyman,

1937:470).

In the reproductive system, examined by me

in four sets of sagittal serial sections, the ovaries

are situated at the level of the second or third

lateral branch of the anterior intestinal ramus. The

testes, in moderate number, are dorsal and prepharyngeal, occupying on either side a longitudinal

zone. The copulatory complex (Figure 45) has been

described and illustrated by Beauchamp (1931, fig.

8) and Hyman (1937, fig. 13), but needs further

analysis. The genital atrium is divided distinctly

into an anterior male atrium (am) surrounding

the penis papilla and a posterior chamber that

appears to be a combination of the common atrium

(ac) and the expanded terminal part of the bursal

duct or vagina (v). The penis has a rather small

bulb (bp) with weak musculature and a large, generally rounded, plug-shaped papilla (pp). This corresponds to the penis shape illustrated by

Beauchamp (1931, fig. 8) rather than to Hyman^s

(1937, fig. 13) diagram of the copulatory apparatus.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The papilla is covered with a cuboidal epithelium

with an underlying coat of muscle fibers, circular

ones adjoining the epithelium and longitudinal

ones below them. The bulb contains a small cavity,

the seminal vesicle (vs), of rounded shape in a

medial section, but extending laterally to both

sides. From this cavity originates a duct (de) passing

into the papilla, rather narrow in its anterior part

and of variable diameter posteriorly, sometimes

widening in a funnel-like fashion, and opening at

the tip of the papilla. This lumen apparently corresponds to an ejaculatory duct. It is surrounded

by a muscle layer consisting chiefly of longitudinal

fibers. The sperm ducts or vasa deferentia (vd)

approach the sides of the penis bulb, enter the

bulb laterally, and open into the lateral extensions

of the seminal vesicle. Eosinophilic gland ducts

enter the penis bulb from the surrounding mesenchyme, traverse the penis, and open on the outer

surface of the papilla. The two oviducts unite

above the male atrium and the common oviduct

(ode) thus formed opens into the dorsal roof of the

male atrium, far removed from the gonopore (gp).

The copulatory bursa (b) is a sac of variable shape

located between the posterior wall of the pharyngeal chamber and the penial bulb. Its outlet, the

bursal duct (bd), starts as a rather narrow, straight

canal running posteriorly above the penis and male

atrium to a level behind the gonopore. There it

turns ventrally and expands into a voluminous

cavity, the vagina (v), which is in wide connection

with the cavity of the common genital atrium (ac).

The vagina has a thick muscle coat of intermingled

circular and longitudinal fibers. It was not possible

to analyze the musculature of the anterior, thin

part of the bursal duct.

All epithelia of the copulatory apparatus are

nucleate except, perhaps, the lining of the male

atrium, which appears to be, at least in part, infranucleate in some of the specimens studied.

DISTRIBUTION AND ECOLOGY.—Sphalloplana percoeca is a troglobitic species and appears to be

widely distributed in the states of Kentucky, Tennessee, and Alabama, possibly extending also into

West Virginia and Georgia. Localities that need

verification are preceded by a question mark (?) in

the following list.

ALABAMA. JACKSON COUNTY: (I) Dinky Pit, Alabama Cave

Survey No. 756 (Carpenter, 1970:71). (2) Fern Cave, Survey

No. 597, located just above the town of Paint Rock; about 25

specimens collected by William W. Torode, many mature, 14

December 1975. (3) Graham Pit, Survey No. 943 (Carpenter,

1970:71). (4) "Old Saltee Cave" ( = Sauta Cave), Survey No.

50, near town of Lim Rock, type-locality of S. alabamensis

(Hyman, 1945:477; Carpenter, 1970:71; Holsinger, 1966:85).

(5) Williams Saltpeter Cave, Survey No. 590, located about 20

miles east of Huntsville; 30 specimens collected by W. W.

Torode, some mature, 27 October 1975.

KENTUCKY. CALDWELL COUNTY: (1) PLisanby Cave (Carpenter, 1970:69). CARTER COUNTY: (1) PCascade Cave (Dearolf,

1953:226). EDMONSON COUNTY: (1) PDiamond Cave (Packard,

1888:28). (2) PGanter Cave (Dearolf, 1953:226). (3) Great

Onyx Cave (Carpenter, 1970:71); specimens kindly sent to

me by J. H. Carpenter in February 1971. (4) Mammoth

Cave, type-locality of Dendrocoelum percoecum, in Shaler's

Brook, Gothic Avenue, Richardson's Spring, Annette's

Dome, Audubon Avenue, and Rafinesque Hall (Packard,

1888:28; Bolivar and Jeannel, 1931:308; Beauchamp, 1931:

317; Giovannoli, 1933:622; Buchanan, 1936:194; Barr, 1968:

155); one specimen collected by the writer in Annette's Dome,

August 1933. (5) PWhite Cave (Dearolf, 1953:226; Carpenter,

1970:71; Barr and Kuehne, 1971:71). ESTILL COUNTY: (1) Pearson Cave (Carpenter, 1970:71). HART COUNTY: (1) PMammoth

Onyx Cave (Dearolf, 1953:226). JACKSON COUNTY: (1) Clemons

Cave, (2) John Rogers Cave, (3) Morning Hole Cave, and (4)

Wind Cave (Carpenter, 1970:71). WAYNE COUNTY: (1) Jesse

Cave (Carpenter, 1970:71).

TENNESSEE (all records need verification). CAMPBELL

COUNTY: (1) Meredith Cave (Barr, 1961:31). DAVIDSON COUNTY:

(1) Mill Creek Cave (McRitchie, 1959:31). PUTNAM COUNTY:

(1) Harve Petty Cave (Barr, 1961:31). RUTHERFORD COUNTY: (1)

Herring Cave (McRitchie, 1959:31). VAN BUREN COUNTY: (1)

Creeping Cave (Barr, 1961:31). WHITE COUNTY: Indian Cave

(Carpenter, 1970:71).

WEST VIRGINIA. PENDELTON COUNTY: (1) PBlowhole Cave

(Carpenter, 1970:69).

Sphalloplana georgiana, a problematic species,

may also be identical with S. percoeca. The localities of that species are listed under S. georgiana.

Some ecological data for S. percoeca were presented by Buchanan (1936): The temperature of the

waters in Mammoth Cave was in the range of 12°C

to 14°C, the pH, 7.6. Holsinger (1966:85) observed

a large population of "S. alabamensis" in Sauta

Cave in a rimstone pool where large amounts of

decaying wood had accumulated; he speculated

that the flatworms may be able to feed on particulate organic matter or on smaller organisms that

inhabit such pools or on both types of food. Barr

(1968:155) and Barr and Duehne (1971:71) reported

that the species in White Cave occurred in temporary rimstone pools soon after the pools filled

with water dripping from stalactites or trickling

down and concluded that they had survived in the

silts of the dried pools by encysting in the layers

of the hygroscopic deposits at the bottom of the
NUMBER 246

pools. A similar survival of cave planarians had

been reported by Ginet and Puglisi (1964) for

Phagocata notadena (Beauchamp) and by Gourbault (1965:477) for Dendrocoelopsis chattoni

(Beauchamp) in two caves in France. The same

mode of survival during the desiccation of the

habitat is known also for some epigean planarians

(see, for example, Lascombe, 1971:31-32).

Carpenter (1970 and 1973) reported that cocoons

of S. percoeca were found in the caves in spring

and summer and hatched in the laboratory at 13°C

in about three months, each cocoon yielding 2-17

young.

PHYSIOLOGICAL STUDIES.—Buchanan (1935 and

1936) studied the behavior of the species in the

cave and the laboratory, its locomotion, reactions

to light, tolerance of acidity and alkalinity (pH

6.6 to 8.0) and of temperature (it may tolerate 24°C

for hours), and its limited regenerative ability.

Wells and Harris (1954) reported briefly on its sensitivity to ultraviolet irradiation.

TAXONOMIC POSITION.—The weakly developed

adhesive organ places S. percoeca in the subgenus

Sphalloplana of which it is the type-species. Other

specific characteristics are the presence of conspicuous auricles, the rounded anterior border of the

intestinal area, the dorsal position of the testes,

the shape of the penis (with small bulb and large,

plug-shaped papilla), the lateral entry of the vasa

deferentia, the opening of the penis lumen on the

tip of the papilla, the confluence of the common

genital atrium and the vagina of the bursal duct,

and the location of the mouth of the common oviduct, far removed from the genital aperture. From

the externally similar S. consimilis it differs by its

penial structure, from S. evaginata by the absence

of diverticula on the common genital atrium, and

from S. holsingeri by the dorsal position of the

testes and the anatomy of the copulatory complex.

Sphalloplana (Sphalloplana) georgiana

Hyman, 1954

FIGURE 46

Sphalloplana georgiana Hyman, 1954:566.

Sphalloplana (Sphalloplana) georgiana.—Carpenter, 1971:1284.

TYPE MATERIAL.—Holotype, one whole mount

(USNM 24614). Paratypes, one whole mount of

anterior part of the body (USNM 24614) and four

sets of sections on 16 slides (AMNH 705).

The species is known for certain only from the

type-locality and Hyman's (1954:566-570) description.

EXTERNAL CHARACTERS.—A blind, white species,

rather slender, with truncate anterior end bearing

a central adhesive organ. Size of preserved specimen, about 8 mm long.

ANATOMY.—Hyman states that the four specimens studied were in a very bad histological condition. The small adhesive organ is rather simple,

forming a cuplike depression with eosinophilic

glands and attached retractor muscles.

The testes are small, arranged on either side in a

rather short band anterior to the pharynx. Hyman

indicates that they are ventral; however, they seem

to me to be in a dorsal position if I interpret

Hyman's paratype slides correctly. In the copulatory organs (Figure 46), the penis has a rounded,

muscular bulb (bp) and a conical papilla covered

with a tall epithelium. The two sperm ducts (vd)

enter the bulb anterolaterally and open separately

into the sides of the rounded bulbar cavity (seminal

vesicle, vs), from which an ejaculatory duct (dc)

proceeds to open at the tip of the papilla. The

common oviduct (ode), surrounded by esosinophilic

shell glands, opens into the roof of the male atrium,

far removed from the gonopore (gp). The copulatory bursa was not observed. The bursal duct (bd)

runs as a slender canal dorsally to the male atrium

to behind the level of the gonopore, then curves

downward and anteroventrally and expands into a

large vagina (v) with thick walls, which is not

separated from the common genital atrium.

DISTRIBUTION AND ECOLOGY.—The type-locality

of S. georgiana is Howard's Waterfall Cave, Dade

County, Georgia, where it was collected by C. E.

Mohr on 8 December 1950. J. H. Carpenter (1970:

76) visited this locality again in 1969 and collected

two sickly, immature specimens that could not be

used for closer examination, but appeared externally identical with S. percoeca.

Dr. J. R. Holsinger transmitted to me planarians

from two caves located in Dade County near the

type-locality of S. georgiana: Byers Cave (1.5 miles

southwest of Rising Fawn) and Johnson Crook

Cave (near Rising Fawn), collected on 18 and 19

June 1967 by J. R. Holsinger, S. Peck, A. Fiske,

and R. Barody (see Holsinger and Peck, 1971:28).

Unfortunately, the specimens were not properly

preserved. Their recognizable anatomical charac-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ters (weak adhesive organ, predominantly dorsal

testes, far dorsal entry of the common oviduct,

general configuration of the penis and bursal duct)

are identical with the features observed in S.

percoeca.

TAXONOMIC POSITION.—Because of the bad condition of the type specimens, the exact position of

the species cannot be determined. All characters of

the copulatory apparatus are compatible with

S. percoeca (which occurs also in the neighboring

state of Alabama). The only essential difference

appears to be in the location of the testes, which

in S. percoeca are dorsal and in S. georgiana, according to Hyman's description, ventral. Hyman's

statement, however, is probably erroneous. The

identity of the two species has already been suspected by Carpenter (1970:77).

Sphalloplana (Sphalloplana) evaginata,

new species

FIGURES 2, 12, 20, 31, 47

TYPE MATERIAL.—Holotype, series of sagittal

sections of anterior region (6 slides) and of posterior region (12 slides), USNM 53427. Paratypes,

three sets of sagittal and transverse sections on 41

slides, USNM 53428-53430; one whole mount,

USNM 53431.

EXTERNAL FEATURES (Figures 2, 12).—Sphalloplana evaginata is a blind, rather large and plump

species, measuring up to 30 mm in length and 6 mm

in width when fully extended. The truncate anterior end has a convex frontal margin and bears

a pair of rather prominent rounded auricular

lobes protruding anterolaterally. Behind the head

is an insignificant narrowing or neck, then the

lateral body margins diverge, run parallel up to

the level of the mouth, then narrow again in the

postpharyngeal region. The tail end is bluntly

pointed. The root of the pharynx lies at about the

middle of the body and its length amounts to onesixth the body length. The copulatory complex is

situated in the anterior half of the postpharyngeal

region. The intestinal system ends anteriorly in

an arc (Figure 20) without forming the V-shaped

extension of the lateral branches which is characteristic of some species of the subgenus Speophila.

The color of the living specimens is a light pink,

due to a diffuse nongranular pigment permeating

the subepidermal tissues. This pigment is not

evenly distributed but is more concentrated in the

head, the pharynx, the copulatory organs, and in a

pair of longitudinal bands along both sides of the

midline in the posterior two-thirds of the prepharyngeal region (corresponding to the site of

the testes). A similar pinkish tint of the body is

seen in Macrocotyla glandulosa Hyman, which,

however, shows the denser organs (adhesive organ,

pharynx, and copulatory apparatus) in a lighter

hue than the general body coloration.

The movements of the worm are rather sluggish.

Upon mechanical stimulation, the adhesive organ

may be protruded temporarily as a pointed, conical

projection in the center of the frontal margin.

ANATOMY.—The marginal zone with tall epidermal cells and long rhabdites is developed typically. The adhesive organ (Figure 31) appears in

the slides as a subterminal shallow depression in

the middle of the frontal margin. The central part

of the organ may be invaginated, but not as deeply

as is seen in representatives of the subgenus Speophila. The epidermis lining the depression is

infranucleate, free of rhabdites, and pierced by

numerous gland ducts containing a granular eosinophilic secretion. The glands forming this secretion

are located in the mesenchyme some distance

behind the organ. A network of muscle fibers is

attached to the epithelium, functioning as retractors. The anatomy of the pharynx shows no

peculiarities.

The two ovaries are situated behind or below

the first to third lateral branches of the anterior

intestinal ramus. In fully mature animals the zone

of yolk glands or vitellaria begins a certain distance

anterior to the ovaries. The testes are numerous

and small, located dorsally, arranged in a pair of

broad longitudinal rows, beginning some distance

behind the ovaries and extending to the level of

the pharyngeal root or somewhat behind it. In the

zone of the testes, the thin sperm ducts or vasa

deferentia run on top of the ventral nerve cords,

generally medial to the oviducts or, in some places,

lateral to them. They connect with the testes by

long efferent ductules. At the level of the pharynx

and behind it, the vasa deferentia form the usual

expanded, tortuous spermiductal vesicles or false

seminal vesicles.

The copulatory complex (Figure 47) was studied

in series of sagittal sections of nine individuals.

The gonopore {gp) leads into a voluminous com-
NUMBER 246

mon atrium (ac), which connects anteriorly with

the male atrium (am) and posteriorly with the

widely expanded terminal part of the bursal duct

or vagina (v). The wall of the ventral section of

the common atrium forms 8-19 (average, 13) cylindrical diverticula or evaginations (da), which project into the surrounding mesenchyme in all directions, but predominantly laterally. The lining of

the genital atria is generally a cuboidal epithelium

underlaid with a thin layer of circular muscle fibers

followed by a layer of longitudinal muscles. Only

the dorsal part of the common atrium has a muscle

coat of interlaced circular and longitudinal fibers.

The penis has a rather small bulb (bp) and a

large, generally plug-shaped, papilla (pp) of variable outline, subject to contraction and distortion

when the animal is being preserved. The papilla

has a very feeble musculature, confined chiefly to

the rather thin layers of circular and longitudinal

muscles underlying its cuboidal epithelial cover,

and circular muscles surrounding the penial lumen,

while its general tissue is parenchymal, with considerable plasticity. The lumen of the penis (pi),

also variable in shape, is uniform both morphologically and histologically, not differentiated into a

seminal vesicle and an ejaculatory duct. It begins

in the ventral part of the penis bulb, proceeds

first dorsally, then arches posteriorly toward the

papilla. In some of the slides, this lumen appears

funnel-shaped, narrow anteriorly and enlarging

toward the papilla, with a wide opening into the

male atrium. In the specimen depicted in Figure

47, the distal part of the lumen was compressed

dorsoventrally, simulating a canal in the sagittal

section. The two vasa deferentia (vd), enlarged

as spermiductal vesicles, approach the base of the

penis bulb, enter it ventrolaterally, diminish in

diameter, and acquire a distinct layer of circular

muscles. They penetrate the bulb, unite in the

midline, and connect with the anterior end of the

penial lumen.

The two oviducts unite in the space between

the genital atrium and the bursal duct to form a

rather short common oviduct (ode) that opens into

the atrium from the dorsal side near the transition

between the male and common atria. The copulatory bursa (b), variable in size, is a rounded sac in

the usual position between the pharyngeal pouch

and the penial bulb. Its outlet, the bursal duct

(bd) runs first as a narrow straight canal in the

midline above the penis to a level behind the

gonopore. Then it curves sharply anteroventrally

and expands into a voluminous cavity, the vagina

(v), which fuses with the common atrium. The

anterior part of the duct has a rather thin muscular

layer, whereas the covering of the vagina consists of

a thicker, but rather loose, coat of intermingled

circular and longitudinal muscle fibers. The

epithelium of the duct is cuboidal and nucleate.

None of the epithelia of the entire copulatory

apparatus are infranucleate.

DISTRIBUTION AND ECOLOGY.—Sphalloplana evaginata is known to occur in four caves in Perry

County, Missouri:

Klump Cave (type-locality), situated east of Perryville; 3 February 1968, 2 specimens collected in

still pools in the stream bed in shallow water,

given to me by Dr. Joseph A. Beatty of Southern

Illinois University; 8 April and 30 September 1973,

17 specimens collected by Jerry Lewis and Brent

Opell, together with the isopod, Asellus antricolus

(Creaser), and the amphipod, Gammarus troglophilus Hubricht and Mackin.

Crevice Cave, near Perryville; 23 September 1961

and 9 June 1964, 8 specimens collected by Stewart

Peck and others, transmitted to me by Russell M.

Norton.

Berome Moore Cave, 4 October 1964, 3 specimens collected by R. A. Brandon and A. Altig, sent

to me by Russell M. Norton.

Garbage Hole Cave, part of the Rimstone Cave

System, southeast of Perryville; 3 February 1973, 2

specimens collected by Jerry Lewis, together with

amphipods and isopods, sent to me alive.

TAXONOMIC POSITION.—Sphalloplana evaginata is

placed in Carpenter's subgenus Sphalloplana on

account of its feebly developed adhesive organ,

although the occasional appearance of a short

invagination in the center of the adhesive pit

indicates a transition to the subgenus Speophila.

It shares other distinctive characters, such as the

dorsal location of the testes, the configuration of

the anterior border of the intestinal system, the

development of a common genital atrium, etc.,

with other species of both subgenera. The most

conspicuous feature of the species is the presence

of cylindrical coeca or evaginations on the wall of

the common genital atrium, which is expressed also

in the specific name evaginata. Similar differentiations have been reported only in 5. kansensis
8

Hyman (1945:479), where the evaginations are

described and depicted as "irregular epithelial

outgrowths." In other details (shape of the penis,

length of the common oviduct, etc.) S. kansensis

differs from S. evaginata.

Sphalloplana (Sphalloplana) kansensis

Hyman, 1945

FIGURE 48

Sphalloplana kansensis Hyman, 1945:478.

Sphalloplana (Sphalloplana) kansensis.—Carpenter, 1971:1284.

TYPE MATERIAL.—Holotype, set of serial sections

on 3 slides, AMNH 311-313.

The species was described by Hyman (1945:478-

479) from one preserved specimen. An examination

of her slides showed that the worm was badly contorted and not suitable for a detailed anatomical

analysis. Another specimen was sent to the writer

by Russell M. Norton, unfortunately also in a

badly preserved state, which made its exact study

impossible. The following description must, therefore, be based on the data presented by Hyman.

EXTERNAL FEATURES.—No illustration of the living animal is available. It is a rather large (20 mm

long), broad, and thin planarian, white and eyeless,

with a truncate anterior end.

ANATOMY.—The adhesive organ is a depression

equipped with eosinophilic glands and a moderate

musculature. It is deeper than is generally seen

in species of the subgenus Sphalloplana, but not as

highly differentiated as in representatives of the

subgenus Speophila, a feature which has been

noticed also by Carpenter (1970:81). Hyman states

that the rhabdites in the marginal zone are only

little larger than elsewhere.

The testes are dorsal. The penis has a slightly

developed bulb and a short, rounded papilla (Figure 48, pp). The vasa deferentia unite outside the

bulb to form the ejaculatory duct (de) that first

ascends dorsally, then bends posteriorly and opens

at the tip of the papilla. The male atrium (am) is

elongated and lined with a very tall epithelium,

the common atrium very small and its wall bears

irregular small blind epithelial outgrowths (da)

that are very characteristic of the species. The

bursal canal (fed) is very long and slender, the bursa

(b) being separated from the penis bulb by a considerable distance, as is the case also in my speciSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

men. The terminal portion of the bursal duct turns

ventrally and is considerably widened. The long

common oviduct (ode) enters the roof of the male

atrium.

DISTRIBUTION.—Sphalloplana kansensis is known

only from the type-locality, Purity Springs, a walled

spring located about 4 miles east of Augusta, Butler

County, Kansas. Hyman's specimen was collected

there on 18 March 1942 by Leslie Hubricht, another specimen by Russell M. Norton on 12 June

1964. I visited this locality on 22 May and 13 July

1967 and again on 10 June 1970 without finding

any planarians. Jerry Lewis was so kind as to

expose in the spring traps baited with fresh shrimp

in August 1973, also without results.

TAXONOMIC POSITION.—By its moderately developed adhesive organ S. kansensis is a species intermediate between the subgenera Sphalloplana and

Speophila, as Carpenter (1970:81) pointed out.

Many aspects of its morphology are unclear, and

Hyman's slides show a great deal of distortion of

the structures of the copulatory complex due to

muscular contractions and possibly some artifacts.

The epithelial outgrowths on the wall of the common atrium are reminiscent of the atrial diverticula

of 5. evaginata, but their shape appears to be quite

different. A clearer picture will ensue only upon

reexamination of properly preserved specimens.

Sphalloplana (Sphalloplana) calif arnica,

new species

FIGURES 4, 21, 32, 39, 49

TYPE MATERIAL.—Holotype, set of sagittal sections on 5 slides, USNM 53432. Paratypes, two sets

of sagittal and transverse sections on 8 slides,

USNM 53433-53434.

EXTERNAL FEATURES (Figures 4, 21).—The largest

specimen, when quietly gliding, measured 17 mm

in length and 2.5 mm in width. The anterior end

is bluntly truncate, with a conspicuous indentation

in the center of the frontal margin. The lateral

edges of the head are rounded, without auricular

projections. No distinct constriction or neck is seen

behind the head, at most a very slight transitory

narrowing during locomotion. Then the body

gradually widens and the lateral margins become

parallel up to the postpharyngeal region. The

posterior end may appear rounded or bluntly

pointed during gliding locomotion. The pharynx
NUMBER 246

is rather short, about one-eighth the body length,

and its root lies somewhat posterior to the middle

of the body. The motion of the animal is either a

smooth gliding or, upon mechanical stimulation,

"crawling." During gliding movement, the central

part of the frontal margin is rather sharply elevated

above the substrate. The worm is purely white,

rather transparent. The intestinal area ends anteriorly with a rounded outline, not showing any

V-shaped extensions of the gut branches.

ANATOMY.—The adhesive organ (Figure 32)

appears in the sections as a slightly depressed, subterminal field of infranudeate epithelium, pierced

by numerous eosinophilic gland ducts and provided

with a moderate number of muscle fibers (retractors). There is no invagination of the adhesive

epithelium into the mesenchyme, which places the

species in the subgenus Sphalloplana.

The testes (Figure 39) are numerous, essentially

ventral, some of them extending into the mesenchymal spaces between the intestinal branches or

bridging the entire dorsoventral diameter of the

body. They occupy, on either side, a longitudinal

zone beginning behind the third or fourth pair of

lateral intestinal branches and reaching to a level

some distance anterior to the insertion of the

pharynx, located mainly medially to the ventral

nerve cord. The testes are in wide communication

with the thin anterior vas deferens that runs along

the medial side of the nerve cord. The ovaries are

positioned below the second pair of intestinal

branches. In the testicular region, the vitellaria or

yolk glands lie mainly in the lateral regions of the

mesenchyme.

The copulatory apparatus (Figure 49) was studied

in two sets of serial sections, one cut longitudinally,

the other transversely. The gonopore (gp) leads

directly into the male atrium (am) and into the

expanded terminal portion of the outlet of the

copulatory bursa, the vagina (v). There is no common atrium developed. The lining of the atrium

is a flattened epithelium with the usual two muscle

layers, a circular and a longitudinal one.

The penis has a large, spherical, highly muscular

bulb (bp) and a rather long, conical papilla (pp).

The papilla is covered by a flattened epithelium

underlaid by a layer of fine circular fibers (/) followed by longitudinal muscles. The fibrous layer

is particularly thick at the base of the papilla,

where it forms a continuation of the circular

muscle layer of the atrial wall. The penis lumen

consists of a rounded cavity (pr) in the bulb, lined

with tall secretory cells filled with an eosinophilic

secretion. From this cavity a narrow canal, the

ejaculatory duct (de) runs posteriorly through the

penis papilla and opens at its tip. The vasa deferentia (vd) enter the penis bulb posteroventrally,

proceed within the bulb anterodorsally, and open,

on either side, into a rounded cavity (us) that

empties into the anterior part of the ejaculatory

duct. This pair of cavities apparently functions as

seminal vesicles, although morphologically it does

not correspond to the unpaired seminal vesicle of

other species of the genus. The latter is represented

in our species by the glandular lumen of the penis

bulb which may have the function of a prostate,

since no sperm passes through it but it apparently

adds some secretions to the sperm fluid.

The two oviducts unite in the space between the

male atrium and the bursal duct, the common

oviduct (ode) proceeding ventrally and opening

into the posterior part of the male atrium, very

close to the gonopore. The copulatory bursa (b)

is a rounded sac, somewhat compressed in the specimen studied. The bursal duct (bd) runs above the

penis and genital atrium as a narrow, straight

canal, then bends ventrally and expands considerably into a vagina (v) that histologically, however,

does not differ from the narrow part of the duct.

The rather thin muscle coat of the duct, not

clearly analyzable, appears to consist of an inner

circular and an outer longitudinal layer.

All epithelia of the copulatory apparatus are

nucleate.

DISTRIBUTION AND ECOLOGY.—The material of S.

californica was furnished to me by the courtesy of

a group of enthusiastic cave explorers, calling

themselves the "Bower Cave Diving Group." The

collections were made in the lake, which fills the

greater part of Bower Cave, by scuba diving, chiefly

by Messrs. Paul Hara, Bill Kruse, and Steven J.

Shimek.

Bower Cave, 8 miles east of Coulterville, Mariposa County, California (type-locality). 16 and 18

July 1971: 4 specimens collected and shipped to

me (only one, immature, arrived alive). 22 October

1971: water temperature 52°F (11°C), 3 immature

worms collected at a depth of about 70 feet (21 m).

2 April 1972: 1 sexually mature specimen from

about 70 feet (21 m) depth. 10 October 1973: 1
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

specimen collected at depth of 100 feet (30 m),

disintegrated in transit. 11 August 1974: 2 specimens, one of them semimature, collected at depth

of 30 feet (9 m).

The species lives in the cave together with

amphipods (Stygobromus wengerorum Holsinger),

which apparently are its natural food. Mr. Hara

informed me that in laboratory cultures the worm

does not accept liver as food, but ingests dead

amphipods and, very eagerly, crayfish meat. He

was able to maintain animals in a culture for over

one year and to raise one specimen to maturity.

TAXONOMIC POSITION.—Sphalloplana californica

is characterized by a very feebly developed adhesive

organ and a somewhat aberrant structure of its

copulatory apparatus, particularly the anatomy of

the penis. The paired seminal vesicles opening into

the ejaculatory duct and the development of a

prostatic cavity are unique within the genus as

far as we know today. Other characteristics, such as

the differentiation of a marginal zone with thickened epithelium and large rhabdites, and the prepharyngeal location of the testes are typical features

of the genus Sphalloplana.

Sphalloplana (Sphalloplana) culvert, new species

FIGURES 17, 33, 40, 50

TYPE MATERIAL.—Holotype, set of sagittal serial

sections on 5 slides, USNM 53435.

. EXTERNAL FEATURES.—I had only one sexually

mature specimen at my disposal, which measured,

in the preserved state, 5.5 mm in length and 1.9

mm in width. In life, the animal must have been

somewhat larger. Unfortunately, I had no opportunity of photographing or drawing the living

worm, only making a few notes on its external

appearance. The anterior end (Figure 17) was

truncate, with a slightly convex frontal margin,

showing occasionally a small median notch when

gliding. The lateral corners had small rounded

auricles projecting anterolaterally. Behind the

auricles was an insignificant narrowing of the body,

then the lateral margins diverged as is usually the

case in related species. The animal was unpigmented, white. The pharynx was inserted at about

the middle of the body and measured approximately one-sixth the body length.

ANATOMY.—The marginal epithelium with large

rhabdites is typically developed. The adhesive

organ (Figure 33) is a rather shallow subterminal

pit lined with an infranucleate epithelium, with

eosinophilic gland ducts opening through the bottom of the pit. The muscles attached to the organ

are chiefly longitudinal retractor muscles.

The ovaries are situated at the level of the

second or third lateral branch of the intestine. The

testes (Figure 40) are arranged in two longitudinal

rows, beginning a certain distance behind the

ovaries and ending at about the level of the

pharynx root. They are ventral, located above and

medially to the ventral nerve cords, opening widely

into the anterior vas deferens. The expanded and

tortuous spermiductal vesicles on the sides of the

pharynx are typically developed.

In the copulatory apparatus (Figure 50), the

gonopore (gp) leads into a small common genital

atrium (ac) that connects anteriorly with the male

atrium (am) and posteriorly with the vaginal end

portion (v) of the bursal duct. The penis consists

of a relatively small bulb (bp) with weak musculature and a large, plug-shaped papilla (pp) with

rounded tip. Near the tip is a rounded depression

that may or may not be an artifact caused by a

muscular contraction at the time of preservation.

The papilla has a coat of muscles at the periphery,

below the outer epithelium, a sheet of circular

fibers underlaid by a layer of longitudinal muscles.

Only a few longitudinal fibers run through the

center of the papilla from the penis bulb. The

epithelial cover of the papilla is infranucleate.

except for a ring of normal nucleate epithelium

at the base of the papilla. The bulbar lumen is a

seminal vesicle (vs), appearing rather small and

dorsoventrally compressed in the median section,

but transversely elongated and provided with a

muscular coat. It is lined with a cylindrical epithelium, apparently of secretory nature, and receives

the outlets of faintly eosinophilic glands. From its

median portion extends a narrow, nonglandular

duct, the ejaculatory duct (de), with cuboidal epithelium, which opens on the dorsal side of the

papilla close to its base. This duct also has a coat

of chiefly longitudinal muscle fibers. The two vasa

deferentia (vd) approach the penis bulb from its

sides, bend upward, enter the bulb laterally, diminishing in diameter, and open each into the lateral

extensions of the seminal vesicle.
NUMBER 246 11

The male atrium (am) duplicates the shape of

the penis papilla. It narrows posteriorly and connects with the common atrium (ac). The two oviducts unite in the space between the male atrium

and the bursal duct to form a common oviduct

(ode), which opens into the atrium from the dorsal

side close to the connection between the male and

common atria. The copulatory bursa (b) is of moderate size. Its outlet, the bursal stalk (bd), proceeds

posteriorly to a level behind the gonopore. It

widens gradually, arches anteroventrally, and expands to a voluminous cavity, the vagina (v),

which narrows again and opens into the common

atrium from the left side. The anterior part of the

bursal duct up to the level of the gonopore is lined

with a normal, nucleate epithelium, while the posterior portion, including the vagina, has an infranucleate lining. The muscular coat of the bursal

duct consists of intermingled circular and longitudinal fibers.

Remarkable is the great extent of infranucleate

epithelia in the copulatory complex. Such epithelia

are seen in the greater part of the cover of the penis

papilla, the terminal portion of the busal stalk, and

the lining of the oviduct. The male and common

genital atria and the penial lumen have normal

nucleate linings.

DISTRIBUTION AND ECOLOGY.—Sphalloplana cnlveri is known from only one locality, Harper Cave

in Tucker County, West Virginia, located 5 miles

southeast of Hendricks. One sexually mature specimen was collected by John R. Holsinger and David

C. Culver on 19 May 1973, together with two

Macrocotyla hoffmasteri (Hyman) and several specimens of a white Phagocata sp. Another specimen,

small and immature, was taken in the same cave 2

January 1974 by Arnold Norden and Beth Ball.

TAXONOMIC POSITION.—The principal characters

of S. culvert are the feeble development of the adhesive organ, presence of small auricles, ventral

position of the testes, dorsal mouth of the ejaculatory duct, and presence of a distinct vagina. It

shares some of these characters with S. holsingeri

from Virginia, which, however, differs by having a

more highly developed, deeply invaginated adhesive

organ with different histological structure. The

species is named in honor of one of its collectors,

Dr. David C. Culver of Northwestern University,

Evanston, Illinois.

Sphalloplana (Sphalloplana) consimilis, new species

FIGURES 5, 22, 34, 41. 51

TYPE MATERIALS.—Holotype, set of sagittal sections on 7 slides, USNM 53407. Paratypes, eight sets

of sagittal sections on 46 slides, USNM 53408-

53415.

EXTERNAL FEATURES (Figures 5, 22).—Fully grown

specimens are up to 14 mm long and 2 mm wide.

The head has a bulging frontal margin and rather

well-developed rounded auricles projecting anteriorly and laterally. In the center of the frontal

margin, a moderately developed adhesive organ is

discernible, which may be protruded as a conical,

pointed projection. Behind the auricles, the body

narrows somewhat, forming a necklike constriction,

then widens again. In the greater part of the body,

the lateral margins run parallel, tapering again in

the postpharyngeal region to meet at the more or

less pointed posterior end. The rather long pharynx, measuring about one-sixth of the body length,

is inserted behind the middle of the body. The copulatory complex occupies the anterior half of the

postpharyngeal region. The color of the worm is

purely white in most specimens. Animals from Buis

Saltpeter Cave, however, showed a light pinkish

hue in the neck region and in the area of the copulatory organs. The anterior margin of the intestinal

area does not form the V-shaped lateral extensions

seen in some species that have a more pronounced

adhesive organ.

ANATOMY.—The thickened marginal epithelium

is typically developed. The adhesive organ (Figure

34) appears in the slides as a shallow subterminal

pit, part of which is invaginated, but not as deeply

as in representatives of the subgenus Speophila.

Both the pit and the invagination receive numerous

eosinophilic gland ducts and are connected with

muscle fibers that presumably act as retractors.

In the reproductive system, the ovaries are

located behind or below the second to fourth lateral

branches of the anterior intestinal trunk. The

rather numerous testes (Figure 41), of moderate

size, are in a strictly dorsal position. They occupy,

on either side of the midline, a longitudinal zone

beginning at a considerable distance posterior to

the ovaries (behind the sixth to eighth lateral

intestinal branches) and extending to the level of

the pharynx root. The thin anterior vasa deferentia
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

run parallel to the ventral nerve cords, somewhat

removed from them, medially to the oviducts.

The copulatory apparatus (Figure 51) was analyzed in sagittal sections of 10 specimens. The genital atrium is divided into an anterior male atrium

(am) containing the penis and a posterior common

atrium that receives the outlet of the copulatory

bursa. Below the infranudeate atrial epithelium

are the usual two muscle layers, a circular one and

a longitudinal one. The penis has a weakly developed, not very muscular bulb and a short, rounded

or plug-shaped papilla, which is covered by a flattened to cuboidal, infranudeate epithelium with

an underlying very feeble muscle layer. Faintly

eosinophilic gland ducts (gl) enter the penis bulb

from the surrounding mesenchyme and penetrate

the penial tissues to open on the surface of the

penis papilla. The two vasa deferentia (vd), after

widening in the pharyngeal region to form the

usual spermiductal vesicles, approach the ventral

side of the penis bulb, proceed medially, and open

into the anterior end of the tubular penis lumen.

There is no seminal vesicle developed. The lumen

consists of a short canal (de), surrounded by a

muscle coat, which begins in the anteroventral

portion of the penis bulb, proceeds first posterodorsally, then arches posteroventrally and opens on

the ventral side of the penis papilla very near its

base. It apparently acts as an ejaculatory duct.

The common oviduct (ode), provided with

numerous eosinophilic shell glands, opens into the

atrium from the dorsal side near the transition

between the two atria, rather far removed from

the gonopore (gp). The copulatory bursa (b) is

round or ovoid. Its outlet, the bursal duct (bd), first

runs posteriorly, above the penis, as a rather narrow canal, then bends ventrally and sometimes

expands into a very wide compartment, the vagina

(v). The musculature surrounding the duct consists

of intermingled circular and longitudinal fibers.

Remarkable is the wide occurrence of infranudeate epithelia in the copulatory complex of the

species at full sexual maturity. The covering of

the penis papilla and the linings of the atria, the

bursal canal (including the vagina), and the common oviduct are all infranudeate.

DISTRIBUTION AND ECOLOGY.—Sphalloplana consimilis has been collected in several caves in western

Virginia and northeastern Tennessee.

TENNESSEE. CLAIBORNE COUNTY: (1) Buis Saltpeter Cave,

northeast of Tazewell, 3 specimens collected 19 August 1972

by J. R. Holsinger and D. C. Culver and sent to me alive.

VIRGINIA. LEE COUNTY: (1) Bowling Cave, east of Ben

Hur, 1 specimen collected 28 July 1967 by J. R. Holsinger.

(2) Cope Cave, south-southeast of Jonesville, 6 specimens

collected 23 August 1969 by J. R. Holsinger. (3) Gallohan

Cave No. 1, type-locality, 8 specimens collected 26 August

1971 by J. R. Holsinger, D. C. Culver, and J. M. Beck and

sent to me alive; they were associated with the isopods,

Asellus recurvatus and Lirceus usdagalun, the amphipod,

Crangonyx antennatus, and the snail, Fontigens sp. (see Holsinger and Bowman, 1973:267). (4) Gregory's Cave, east-southeast of Rose Hill, several specimens collected 27 October

1967 by J. R. Holsinger. (5) McClure's Cave, west of Jonesville, 9 August 1973, 5 specimens; 31 July 1974, about 50

specimens collected by J. R. Holsinger and D. C. Culver

from a huge population of perhaps 300 worms, sent to me

alive.

TAXONOMIC POSITION.—By the anatomy of the

moderately developed adhesive organ, S. consimilis

forms a transition between the subgenera Sphalloplana and Speophila. The most outstanding characteristic of the species is the configuration of the

penis, specifically the lack of a seminal vesicle and

the opening of the ejaculatory duct on the ventral

side near the base of the short, pluglike papilla. A

similar feature is seen in S. virginiana, which

species, however, has the testes positioned ventrally,

while in S. consimilis they are located on the dorsal

side. The species name, consimilis (Latin, similar),

refers to the great similarity of the copulatory

complex with that of S. virginiana.

Sphalloplana (Sphalloplana) subtilis, new species

FIGURES 6, 13, 27-29, 35, 42, 52

TYPE MATERIAL.—Holotype, set of sagittal sections on 5 slides, USNM 53444. Para types, two sets

of sagittal and transverse sections on 9 slides,

USNM 53445-53446.

EXTERNAL FEATURES (Figures 6, 13).—This is a

very slender, unpigmented species, up to 16 mm

long and 1 mm wide when extended. The head end

is truncated, with almost straight or slightly convex

frontal margin and rounded lateral edges, which

may extend very little anteriorly and laterally.

There is no conspicuous adhesive organ discernible

in life. In the quietly gliding animal, no necklike

construction is seen behind the head, at most a

very insignificant narrowing. The lateral margins

of the body run almost parallel for the greater part

of the body length, tapering behind the level of the
NUMBER 246 13

copulatory complex to meet at the posterior end.

The short pharynx, measuring about one-ninth of

the length of the body, is inserted far back at about

the beginning of the third fifth of the body length.

The copulatory apparatus is located in the anterior

half of the postpharyngeal region.

ANATOMY.—The marginal zone with thickened

epithelium and large rhabdites, characteristic of

the genus, is developed typically (Figure 29). The

adhesive organ (Figure 35) appears in the sections

as a small subterminal pit in the center of the

frontal margin, lined with an infranucleate epithelium pierced by numerous eosinophilic gland ducts.

It has a very feeble musculature, principally fibers

that may act as retractors. No protrusion of the

organ has been observed in living specimens upon

stimulation, such as is seen in many other representatives of the genus.

The anterior ramus of the intestine bears over

20 pairs of lateral branches, each posterior ramus

about 25 short branches. The anterior border of the

intestinal area is rounded. Behind the copulatory

apparatus, the two posterior rami unite to a single

trunk.

The two ovaries are located behind the third

to fifth lateral branches of the anterior intestinal

ramus. The testes, in moderate number, form a

pair of longitudinal rows extending through the

posterior three-fifths of the prepharyngeal region.

They are ventral, lying below the intestinal

branches (Figure 42), above and medial to the

ventral nerve cords. Many testicular follicles are

connected directly to the thin anterior vasa deferentia that run along the upper border of the nerve

cords, medial to the oviducts.

The copulatory apparatus (Figure 52) was analyzed in two series of sagittal sections. It occupies

the anterior half of the postpharyngeal region. The

genital aperture (gp) leads into two cavities, anteriorly into the male atrium (am) and posteriorly

into the expanded outlet of the copulatory bursa

or vagina (v). The penis consists of a rather small

bulb (bp) embedded in the mesenchyme and a

larger, plug-shaped papilla (pp). Its lumen (pi)

forms an arched canal, histologically rather uniform, without a differentiation into a seminal

vesicle and an ejaculatory duct. This canal, lined

with a cuboidal epithelium, opens into the atrium

on the ventral side of the penis papilla. The two

sperm ducts or vasa deferentia (vd), which at the

level of the pharynx expand to form the spermiductal vesicles, approach the penis bulb and enter

it anterolaterally, diminish in diameter, acquire

each a muscle coat, and open into the anterior part

of the penial lumen independently from the sides.

The musculature of the penis is very weakly

developed. That of the penis bulb consists of a

loose network of muscle fibers, while the papilla has

a thin muscle layer developed below the external

epithelium. The penial lumen is enclosed in a

coat of circular muscle fibers. Many faintly eosinophilic gland ducts enter the bulb from the surrounding mesenchyme and empty into the penis

lumen as well as into the atrium in a field on the

dorsal side of the penis papilla.

The two oviducts or ovovitelline ducts unite in

the space between the male atrium and the bursal

duct, forming a common oviduct (ode), which

runs ventrally and opens into the posterior part of

the male atrium. As is usual, the terminal parts of

the paired oviducts and the common oviduct receive the outlets of strongly eosinophilic so-called

shell glands. The copulatory bursa (b) is a voluminous sac lying anterior to the penis bulb. Its

outlet, the bursal duct (bd), begins as a rather

narrow, straight canal passing posteriorly above

the penis and male atrium to a level posterior to

the gonopore. There it bends abruptly anteroventrally and widens into a large expansion, the

vagina (v). Close to the gonopore, the vagina narrows again and joins the atrial outlet. The epithelial lining of the vaginal sac is infranucleate,

while the epithelia of the narrow proximal bursal

canal and the narrow terminal parts of the vagina

contain nuclei. The vagina has a coating of muscles

consisting of intermingled circular and longitudinal

fibers, the circular ones predominating. The musculature of the thin anterior part of the bursal

canal is very feeble and could not be analyzed in

my preparations.

DISTRIBUTION AND ECOLOGY.—Sphalloplana subtilis is known only from a spring on the property

of J. W. Biggers at 6278 East Edsall Road, Fairfax

County, Virginia, close to the Alexandria city line.

The spring (Figures 27, 28) is enclosed in a brick

structure with a removable concrete cover. The

water is 6 feet (1.8 m) deep, its level reaching to 2

feet (0.6 m) below the rim of the casing. The species

was obtained by J. R. Holsinger, W. Biggers, and

the writer by baiting with fresh marine shrimp on
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

several occasions between 18 March and 12 August

1973. The water temperature on 25 March was

8.6°C, on 12 August 17.7°C (while the temperature

of the surface layer was 22.1°C), the pH 5.4 on

both dates. A total of 10 specimens were collected,

three of them sexually mature. Besides S. subtilis,

the spring contained another species of Sphalloplana, S. holsingeri, which was much more abundant (over 80 specimens were collected).

TAXONOMIC POSITION.—The weakly developed

adhesive organ of S. subtilis places the species in

the subgenus Sphalloplana. The testes are ventral

as they are in some other species of the genus. The

most important specific characters are in the configuration of the copulatory organs. The short penis

with very weak bulb and plug-shaped papilla and

the ventral opening of the penial lumen show

some similarity with S. consimilis (which has dorsal

testes) and S. virginiana (which has a more highly

developed adhesive organ). Another outstanding

character, the development of a voluminous vagina

with infranucleate epithelium, is shared with S.

holsingeri, from which it differs by the anatomy of

the penis and of the adhesive organ. The specific

name, subtilis (Latin, slender) alludes to the elongated shape of the species.

Sphalloplana (Speophila) virginiana Hyman, 1945

FIGURE 53

Sphalloplana virginiana Hyman, 1945:477.

Sphalloplana (Sphalloplana) virginiana.—Carpenter, 1971:

1284.

TYPE MATERIAL.—Holotype, whole mount of one

specimen, AMNH 314. Paratypes, set of sagittal

sections on 5 slides, MNH 315-319, and whole

mount of three specimens on one slide, AMNH 706.

The following discussion is based on Hyman's

(1945:477-478) description and a reexamination of

Hyman's paratype sections.

EXTERNAL FEATURES.—A slender, white species,

preserved specimens measuring to 12 mm in length.

ANATOMY.—The adhesive organ, according to

Hyman, consists of an irregular depression lined

with eosinophilic gland cells, from which a small

band of retractor muscles proceed dor sally to join

the dorsal subepidermal muscle layers. The paratype slide, however, shows a distinct invagination

of the organ, as has already been noticed by

Carpenter (1970:79).

In the reproductive system, the numerous testes

are subventral, lying below the intestinal branches

or extending into the mesenchymal spaces between

the branches. Carpenter (1970:78) stated that the

testes appeared to him to be dorsal rather than

ventral on Hyman's slides; however, I see them

there clearly in a ventral position.

The copulatory complex (Figure 53) is located

at a considerable distance posterior to the pharyngeal pouch. The penis has a slightly developed

bulb and a papilla (pp) of moderate size, somewhat

more elongated than shown in Hyman's figure.

The penial lumen lacks a distinct seminal vesicle

and consists of a short, rather wide canal, the

ejaculatory duct (de), which begins in the penis

bulb, runs posteroventrally, and opens on the

ventral side of the papilla close to its base. The two

sperm ducts (vd) enter the penis bulb anteriorly

and connect with the anterior end of the ejaculatory duct. The mouth of the common oviduct (ode)

is on the posterodorsal roof of the male atrium

(am). The rather small copulatory bursa (b) is

situated above the penis bulb, with its outlet, the

bursal duct (bd), running posteriorly and curving

ventrally to connect with the common atrium (ac).

DISTRIBUTION AND ECOLOGY.—The type-locality of

S. virginiana is Showalter's Cave in Rockbridge

County, Virginia, where numerous specimens were

collected in a pool on 30 October 1943 (Hyman,

1945:478). Carpenter (1970:80) visited this locality

in 1969 and J. R. Holsinger at several occasions,

both without finding any planarians.

TAXONOMIC POSITION.—Sphalloplana virginiana

is here placed in the subgenus Speophila on account of its invaginated adhesive organ, as has

already been suggested by Carpenter (1970:79). Its

outstanding specific characteristics are the ventral

position of the testes, the lack of a seminal vesicle,

and, particularly, the ventral opening of the ejaculatory duct. The species appears to be rather close

to S. subtilis (which, however, has a very weak

adhesive organ) and to S. consimilis (which has the

testes positioned dorsally). A closer comparison will

be possible only after the reexamination of wellpreserved specimens from the type-locality.

Sphalloplana (Speophila) holsingeri, new species

FIGURES 7, 14, 18, 27, 28, 36, 43, 54

TYPE MATERIAL.—Holotype, set of sagittal sec-
NUMBER 246 15

tions on 5 slides, USNM 53436. Paratypes, eight

sets of sagittal and transverse sections on 35 slides,

USNM 53437-53443, 53478.

EXTERNAL FEATURES (Figures 7, 14).—Sphalloplana holsingeri is a blind, white species, measuring, when gliding quietly, up to 15 mm in

length and 1.5 mm in width. The anterior end is

truncate, with a bulging frontal margin flanked by

a pair of short, rounded auricles projecting anterolaterally. In the center of the frontal margin, the

location of the adhesive organ is visible as an

opaque spot. When protruded, the organ appears

as a short conical projection. Behind the auricles,

the body narrows slightly, then widens again to

attain its greatest width a short distance behind

the head. In the greater part of the body, the

lateral margins run parallel up to the region of

the copulatory complex, where they converge again

to meet at the pointed posterior end. An opaque

marginal rim, corresponding to the modified marginal epithelium, is conspicuous in the living

animal. The pharynx is inserted behind the middle

of the body and measures in length about oneseventh the length of the body. The copulatory

apparatus occupies the anterior half of the postpharyngeal region. The anterior end of the intestine forms a median projection behind the adhesive

organ (Figure 18); there is no V-shaped extension

of the lateral branches to the sides of the organ

such as is seen in some other species of the subgenus Speophila, e.g., S. pricei (Hyman) (Figure 19).

ANATOMY.—The epidermal epithelium of the

lateral margins is thickened and contains rhabdites

considerably larger than those of either the dorsal

or ventral epidermis, as is seen generally in the

genus Sphalloplana.

The adhesive organ (Figure 36) is well developed,

consisting of a rounded subterminal depression,

the central part of which forms a deep invagination when the organ is retracted. The muscular

differentiations of the organ correspond to those

described in other species of Sphalloplana with

conspicuous adhesive organs, such as S. pricei (see

Hyman, 1937:464-465), S. mohri Hyman (see

Mitchell, 1968:611-613), and S. weingartneri Kenk

(see Kenk, 1970:315). The glandular equipment of

the organ, however, differs from the conditions

seen in other species of the genus. Two kinds of

glands open through the epithelial lining: eosinophilic gland ducts passing through the infranucleate

epithelium of the invaginated portion and cyanophilic glands through the remaining field of the

depression surrounding the inverted canal. The

eosinophilic secretions are of a homogeneous or

coarsely granular nature, while the cyanophilic

secretions, staining deeply with hematoxylin,

appear as threadlike or rod-shaped filaments. The

gland cells producing these threads form voluminous masses occupying both the dorsal and ventral

portions of the mesenchyme and the spaces between

the intestinal branches in the medial part of the

anterior two-thirds of the prepharyngeal region.

In other species of Sphalloplana, only eosinophilic

secretions are known to occur. It is difficult to

interpret the functions of the two types of glands.

The eosinophilic secretions in the central, eversible

part of the organ presumably correspond to those

described in other triclads that are equipped with

adhesive organs, e. g., Procotyla finviatilis Leidy,

where the secretions dissolve into a sticky mucous

mass that is used in the capture of living prey. The

function of the cyanophilic secretions, so prominent

in S. holsingeri, is not clear. It is interesting to note

that a similar differentiation of the glands of the

adhesive organ has been observed in a European

planarian, Dendrocoelum album (Steinmann),

where erythrophilic glands open through the

epithelium of the adhesive pit and cyanophilic

gland ducts at the peripheral rim of the organ

(Steinmann, 1910:190).

The musculature of the pharynx follows the plan

of the genus Sphalloplana. The internal pharyngeal

epithelium is surrounded by a thick layer of circular muscle fibers, followed by a thinner layer of

longitudinal muscles.

In the reproductive system, the two ovaries are

situated on the medial sides of the ventral nerve

cords, behind the third or fourth lateral branch

of the anterior intestinal trunk. The numerous

testes (Figure 43) are arranged in two longitudinal

rows, one on either side of the midline, occupying

the posterior half of the prepharyngeal region.

They are located essentially ventral, medial to the

nerve cord. Only a few testes may be pushed toward

the dorsal side in the "septa" between the intestinal

branches, or may extend through the entire dorsoventral diameter of the body. Many follicles are

attached directly to the sperm ducts or vasa deferentia without the mediation of efferent ductiiles.

In the region of the testes, the vasa deferentia run
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

parallel to the ventral nerve cords, somewhat removed from the ventral integumental muscle layers.

At the level of the pharynx, they widen to form the

sinuous spermiductal vesicles (or false seminal

vesicles) that keep their expanded shape up to their

entrance into the penis bulb.

The copulatory apparatus (Figure 54) was analyzed in 12 series of sagittal sections. The penis

consists of a relatively small bulb (bp) with rather

feeble musculature, embedded in the mesenchyme,

and a larger, generally plug-shaped, papilla, sometimes constricted at its base, protruding into the

genital atrium. The shape of the papilla is rather

variable in the specimens studied, as it is easily

distorted when the animal is being killed. It lacks

any rigid supporting structures and its musculature

is confined to two rather thin layers of muscle

fibers, a circular layer underlying the external

epithelium and a weak longitudinal layer below

it. Only a few scattered longitudinal fibers run

through the parenchyma of the papilla. The bulb

contains an elongated cavity, the seminal vesicle

(vs), lined with an epithelium pierced by gland

ducts with a finely granular, faintly cyanophilic

secretion that enter the bulb from the surrounding

mesenchyme. Toward the penis papilla, the cavity

continues as a narrow, nonglandular canal, the

ejaculatory duct (de), which opens into the atrium

on the dorsal side of the papilla, about halfway

between its base and its tip.

The genital aperture or gonopore (gp) leads into

two cavities, anteriorly the male atrium (am) and

posteriorly the cavity of the large vagina (v). There

is no common atrium developed.

The two vasa deferentia (vd), which approach

the penis bulb as enlarged spermiductal vesicles,

enter the bulb ventrolaterally, diminishing in diameter, and open separately into short lateral extensions of the anterior part of the seminal vesicle.

The two oviducts, in the region of the copulatory

complex, ascend from their course above the ventral nerve cords, pass to the midline, and unite

above the male atrium to form the common oviduct (ode), which arches ventrally and opens into

the posterior part of the atrium. The copulatory

bursa (b) is an elongated sac lying close to the anterior face of the penis bulb. Its outlet is a rather

narrow, straight canal (bd) running in the midline

dorsally to the penis and atrium and extending

rather far posteriorly. Behind the level of the

gonopore, the canal curves ventrally and widens

into a voluminous cavity, the vagina (v). In the

preserved specimens, the outline of the vagina is

very variable, usually irregularly folded. In fully

mature worms, the lining of the vagina is infranucleate while that of the narrow part of the

bursal canal contains nuclei. In young specimens,

however, nuclei are present in the vaginal epithelium; it appears, therefore, that the nuclei pass

from the cellular lining only at the approach of

full sexual maturity. The vagina is surrounded by

a thick coat of interlaced circular and longitudinal

muscle fibers. The musculature of the anterior,

narrow part of the bursal canal is very feeble,

probably also made up of intermingled longitudinal and circular muscles.

DISTRIBUTION AND ECOLOGY.—Sphalloplana hoisingeri is known from only one locality, the spring

on the property of J. W. Biggers in Fairfax County,

Virginia, the same spring where S. subtilis occurs

(see above and Figures 27, 28). Two specimens,

both mature, were collected first by Dr. John R.

Holsinger on 23 May 1965 but were fixed in alcohol, which made an analysis of their anatomy

rather difficult. Additional (over 80) specimens

were obtained by Dr. Holsinger, Bill Biggers, and

the writer by baiting with shrimp meat on 18

March, 25 March, 2 April, and 12 August 1973,

among them many mature animals.

TAXONOMIC POSITION.—The presence of a deeply

invaginated, protrusible adhesive organ places the

species in the subgenus Speophila. Among the outstanding characteristics of the species is the differentiation of the glands of the adhesive organ,

eosinophilic glands in the inverted part, surrounded

by a field of cyanophilic gland openings. In the

reproductive system, it shares the ventral position

of the testes with several other species of the genus.

The most important specific characters are in the

anatomy of the copulatory complex: the opening

of the ejaculatory duct on the dorsal side of the

penis papilla and the development of a voluminous

vagina in the terminal part of the bursal duct. The

species is named in honor of my distinguished

colleague, Dr. John R. Holsinger, who collected the

first specimens and collaborated in the procurement of additional material of the species.

When the manuscript of this description was

completed, I received notice that the American

Museum of Natural History had, among the ma-
NUMBER 246 17

terials left by the late Libbie H. Hyman, a set of

three slides of sagittal sections marked "Sphalloplana holsingeri." The condition of the sections is

rather bad, so that no identification of the species

can be made. Apparently, Hyman's S. holsingeri

is an unpublished manuscript name and has no

nomenclatorial standing.

Sphalloplana (Speophila) weingartneri Kenk, 1970

FIGURES 23, 55

TYPE MATERIAL.—One set of 2 slides of sagittal

sections, USNM 41181.

A description of this species was given by Kenk

(1970), based on a single specimen. The description will not be repeated here, only the essential

characteristics may be listed.

EXTERNAL FEATURES (Figure 23).—This is a rather

small species, 6 mm long and 1.2 mm wide, white.

Anterior end truncate, without auricles. The anterior border of the intestinal zone is rounded,

without a V-shaped median recess. The pharynx is

inserted somewhat behind the middle of the body,

measuring about one-seventh the body length. The

region of the copulatory apparatus is visible in life

as a transparent field with an opaque spot in the

center.

ANATOMY.—The deeply invaginated adhesive

organ and the tall marginal epidermal cells with

long rhabdites are typically developed. The testes

are located both dorsally and ventrally. In the

copulatory apparatus (Figure 55), the penis consists

of a large, spherical, very muscular bulb (bp) and

a small, rather slender papilla (pp)- The vasa deferentia (vd) enter the bulb ventrolaterally. The penis

lumen is divided into an elongated seminal vesicle

(vs) with a secretory epithelial lining and a narrow

ejaculatory duct (de) that opens at the tip of the

penis papilla. The common oviduct (ode) empties

into the posterior part of the male atrium (am),

close to the gonopore (gp).

DISTRIBUTION AND ECOLOGY.—One specimen was

collected by Lawrence Weingartner in Bronsons

Cave in Spring Mill State Park, Lawrence County,

Indiana, on 15 August 1969. Dr. Jerry Carpenter

informed me that he obtained additional material

of the species from the type-locality at a later visit

(in 1970) and kindly sent me photographs of a

sexually mature specimen.

TAXONOMIC POSITION.—Sphalloplana weingartneri appears to be rather closely related to 5. pricei

of Pennsylvania. It resembles it by the shape of the

anterior end (absence of auricles), the location of

the testes (dorsal and ventral), and the ventrolateral

entry of the vasa deferentia into the penis bulb. It

differs from S. pricei by its smaller size, the configuration of the anterior intestinal border, and

particularly by the size relation between the bulb

and the papilla of the penis.

Sphalloplana (Speophila) buchanani

(Hyman, 1937)

FIGURES 15, 56

Sphalloplana percoeca.—Beauchamp, 1931:321 [in part].

Speophila buchanani [nomen nudum] Hyman, 1936:129.

Speophila buchanani Hyman, 1937:468.

Sphalloplana buchanani.—Mitchell, 1968:615.

Sphalloplana (Speophila) pricei.—Carpenter, 1971:1283 [in

part].

TYPE MATERIAL.—Syntypes, 6 specimens (2

whole mounts on one slide and 4 sets of serial

sections on 11 slides), AMNH 648.

I had no opportunity of examining living specimens of this species, only Hyman's type slides that

are histologically very poor, in part with badly

faded stain. Sphalloplana buchanani was described

originally by Hyman (1937:468-469) from preserved specimens obtained by Prof. J. W. Buchanan

in Mammoth Cave in Kentucky. Carpenter (1970,

1971) considers the species to be identical with

S. pricei from Pennsylvania. In the absence of an

exact analysis of its anatomy, however, we may

consider this identity to be still an open question.

The following account is based chiefly on the

literature data available.

EXTERNAL FEATURES (Figure 15).—Hyman (1937,

fig. 16) published an illustration of the aspect of

the species in life, prepared by Buchanan. It varies

in length from 7 to 15 mm. The anterior end is

shown to be truncate with rounded lateral edges,

lacking auricular projections. The protrusible adhesive organ appears as a retracted conical structure. The intestinal area terminates anteriorly with

a V-shaped outline. The pharynx, measuring about

one-eighth of the body length, is inserted somewhat

posterior to the middle of the body, and the copulatory complex occupies the anterior third of the

postpharyngeal region. All these features are compatible with the conditions obtaining in S. pricei.

ANATOMY.—The deeply invaginated adhesive
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

organ and the lateral marginal zone with large

rhabdites are developed typically. The testes, in

moderate number, are prepharyngeal and dorsal

(in S. pricei they are dorsal, ventral, and intermediate). Illustrations of the copulatory organs (Figure 56) have been given by Hyman (1937, fig. 19)

and probably by Beauchamp (1931, fig. 6), who

confused the species with the sympatric S. percoeca.

Both figures show the aspect of the organs in dorsal

view; however, Hyman's slide of horizontal sections,

from which her figure is drawn, shows the bursal

duct to be on the right side of the penis rather

than on the left side. The penis has a rather

muscular bulb (pb) and a cylindrical to conical

papilla (pp). The vasa deferentia enter the bulb

laterally, traverse its tissues, and unite to a rather

narrow canal, the ejaculatory duct (pi), which runs

through the center of the papilla and opens at its

tip. There is no seminal vesicle developed, but in

Hyman's figure a widening of the ejaculatory duct

is shown near the tip of the papilla. Carpenter

(1970:92) also stated that in his specimen of "S.

pricei" from Mammoth Cave no dilated seminal

vesicle was present. The common oviduct (ode)

opens into the male atrium close to the entrance

of the bursal stalk (bd), which is there enlarged to

form a vagina (v) with somewhat muscular walls.

DISTRIBUTION AND ECOLOGY.—The only reliably

documented habitat of S. buchanani is Mammoth

Cave, Edmonson County, Kentucky, where it was

collected first in 1928 (Bolivar and Jeannel, 1931:

38, in Audubon Avenue, together with S. percoeca),

by Buchanan and others in October and November

1935 (in the less accessible and deeper parts of the

cave, Hovey's Cathedral Domes, Fox's Alley, and

Becky's Alley, see Hyman, 1937:469), and again by

Carpenter (1970:92, 180).

Additional localities recorded in the literature

need further confirmation. McRitchie (1959:24)

reported "Speophila buchanani" from Mill Creek

Cave, Davidson County, Tennessee. Carpenter

(1970:92) found a Sphalloplana which he believed

to be identical with the Mammoth Cave form in

Cathedral Cave, Edmonson County, Kentucky, a

few miles from Mammoth Cave. His specimens,

however, had a well-developed seminal vesicle and

the vasa deferentia entering the penis bulb posterodorsally (see his fig. 51). Finally, Barr and Kuehne

(1971:70) observed S. buchanani in a stream in

Great Onyx Cave, Edmonson County, Kentucky, a

locality where Carpenter (1970:180) had collected

only S. percoeca.

A few physicochemical parameters of the habitat

of S. buchanani are given by Hyman (1937:469).

The temperature of the water in Mammoth cave

was 55°F [12.8°C], the pH, 7.6. Barr (1968:155)

observed the species in temporary drip pools and

assumes that it survives the periods of dryness by

encystment in the hygroscopic deposits at the bottom of the pools, as does S. percoeca.

TAXONOMIC POSITION.—Sphalloplana buchanani

appears to be very close to S. pricei, from which it

differs by the location of the testes and the anatomy

of the penis, particularly the absence of a seminal

vesicle and the morphology of the ejaculatory duct.

Sphalloplana (Speophila) pricei (Hyman, 1937)

FIGURES 8, 19, 24, 37, 44, 57

Speophila pricei Hyman, 1937:462.

Sphalloplana pricei.—Mitchell, 1968:615.

Sphalloplana (Speophila) pricei.—Carpenter, 1971:1284 [in

part].

TYPE MATERIAL.—Holotype, one set of 6 slides of

sagittal sections, USNM 20228. Paratypes, 5 specimens in alcohol, USNM 20236. Six specimens (one

whole mount and 5 sets of serial sections mounted

on 31 slides), designated as syntypes, AMNH 649

(see Feinberg, 1970:49).

EXTERNAL FEATURES (Figures 8, 24).—Mature

specimens are up to 28 mm long and 3.5 mm wide,

but maturity is observed also in smaller individuals.

The color in life is a pure white apart from the

intestinal contents, which may shine through the

body wall. The anterior end has a straight or very

slightly bulging frontal margin, with a median

notch frequently seen when the animal is gliding.

No auricular projections are developed, as the

rounded lateral edges of the head do not extend

anteriorly. Behind the head, the body first narrows,

forming a necklike constriction, then widens again.

In the greater part of the body, the lateral margins

run parallel, converging again behind the region

of the pharynx and meeting in the rather pointed

tail end.

The anterior intestinal border shows a V-shaped

indentation, the lateral branches extending somewhat anteriorly to both sides of the adhesive organ

(Figure 19). The pharynx is inserted at about the

middle of the body, its length amounting to
NUMBER 246 19

approximately one-seventh the body length. The

copulatory apparatus, visible in life as an elongated

transparent field with a central opaque spot, is

situated in the anterior half of the postpharyngeal

section.

ANATOMY.—The structure of the adhesive organ

(Figure 37), which in the retracted state forms a

deep, folded invagination, is described well by

Hyman (1937:464-465). No "snout-like" projection

was visible, however, in specimens fixed witH hot

mercuric chloride solution. The modificatioits of

the marginal epithelium with tall cells and large

rhabdites are also analyzed by Hyman.

I have studied the reproductive organs of the

species in eight sets of serial sections, including

some of the type specimens. The ovaries are situated behind the third or fourth pair of lateral

branches of the intestine. The numerous, rather

small testes are not confined to the dorsal region,

as Hyman indicates, but are located dorsally,

ventrally, and in intermediate positions (Figure

44). The thin anterior vas deferens on either side

runs along the ventral nerve cord, parallel to the

oviduct and on the medial side of it. It expands at

the pharyngeal level as the usual spermiductal

vesicle. The copulatory apparatus appears rather

much contracted in Hyman's figure 11. In specimens killed in the extended state by a hot fixative,

the apparatus is generally more elongated (Figure

57). The genital aperture (gp) leads into a common

atrium (ac), which communicates widely with the

anterior male atrium (am). The penis has a muscular bulb of moderate size and a large, very pliable

papilla (pp), which is very variable in appearance,

from conical to finger-shaped, often bent in various

directions. The constriction at the middle of the

papilla, mentioned by Hyman, is not a regular

occurrence. The papilla is provided with a very

feeble musculature, chiefly two thin layers of

muscle fibers, one circular, the other longitudinal,

below the external epithelium of the papilla. The

two vasa deferentia (vd) enter the penis bulb

ventrolaterally and proceed posterodorsally toward

the midline. They open into the penis lumen either

separately and close together or after first uniting

to a short common vas deferens. The penial lumen

is an elongated cavity, arching from the bulb

toward the papilla and opening at the tip of the

latter. The anterior part of the cavity has a glandular epithelial lining, the cells of which, staining

deeply with eosin, project in some of the specimens

into the lumen in a villus-like fashion. This anterior part is frequently widened and corresponds

to a seminal vesicle (vs). Posteriorly, the cavity continues as a canal with somewhat variable diameter,

the ejaculatory duct (de). Gland ducts with a homogeneous, faintly cyanophilic secretion enter the

penis bulb from the surrounding parenchyma and

open into the anterior portion of the ejaculatory

duct. The common oviduct (ode) opens from the

dorsal side into the atrium near the junction of the

male and common atria. The bursa copulatrix (b)

shows no peculiarities. Its outlet, the bursal duct

(bd), starts as a straight, narrow canal but widens

considerably in its posterior section, forming sinuous convolutions and acquiring a thicker muscle

coat. This widened terminal part may be termed

a vagina (v).

None of the epithelia of the copulatory complex

are infranucleate.

DISTRIBUTION AND ECOLOGY.—Sphalloplana pricei

inhabits several caves in Pennsylvania.

DAUPHIN COUNTY: Brownstone Cave: One specimen collected by John W. Price, 23 May 1936 (Hyman, 1937:466).

LANCASTER COUNTY: Refton Cave (type-locality): several specimens collected by John W. Price, 20 January and 17 April

1936 (Hyman, 1937:466); several specimens collected by Jerry

H. Carpenter, 20 May 1971; 10 specimens collected by Arnold

Norden and Beth Ball, 4 February 1973. MIFFLIN COUNTY:

Upper Johnson Cave: specimens collected by Charles E. Mohr

and Kenneth Dearolf, 23 January and 27 February 1937

(Hyman, 1937:471; Dearolf, 1941:170).

TAXONOMIC POSITION.—The distinguishing characters of S. pricei are the following: Adhesive organ

deeply invaginated, anterior intestinal border Vshaped, testes dorsal and ventral, penial papilla

large with ejaculatory duct opening at tip, and

entry of the vasa deferentia into the penis bulb

ventrolateral. Carpenter (1970:85) considers 5.

pricei to include three more of Hyman's species of

Speophila, S. buchanani, S. hubrichti, and 5. hoffmasteri, which he lists as synonyms of S. pricei. An

analysis of the morphological characters of these

forms, however, justifies their standing as good

separate species. Sphalloplana buchanani appears

to have only dorsal testes (Hyman, 1937:469), but

will need a more detailed reexamination; S.

hubrichti differs from our species by having dorsal

testes and not showing the V-shaped recess in the

anterior border of the digestive system; S. hoffmasteri, by the arrangement of the muscles in the
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

interior pharyngeal muscle zone, is to be removed

from the genus Sphalloplana and placed into

Macrocotyla (Kenk, 1975:333).

Sphalloplana (Speophila) hubrichti (Hyman, 1945)

FIGURES 9, 25, 58, 59

Speophila hubrichti Hyman, 1945:479.

Sphalloplana hubrichti.—Mitchell, 1968:615.

Sphalloplana (Speophila) pricei.—Carpenter, 1971:1284 [in

part].

TYPE MATERIAL.—Holotype, whole mount,

AMNH 320. Paratypes, whole mount and sets of

serial sections, AMNH 321-324 and 703 from

Kolms Cave; whole mount and set of serial sections,

AMNH 704, from Morrisons Cave.

EXTERNAL FEATURES (Figures 9, 25).—This is a

rather large species, measuring up to 20 mm in

length and 3 mm in width, in life of purely white

color. The head is truncate, with slightly bulging

frontal margin, frequently showing a median notch

at the site of the adhesive organ. The lateral edges

of the head are rounded, without auricular projections, but somewhat protruding laterally. The

adhesive organ is visible in the living specimen as

an opaque spot. The pharynx is situated behind

the middle of the body and its length is about oneeighth the body length. The copulatory organs

occupy the anterior half of the post pharyngeal

region. The testes are visible in life as transparent

spots, arranged in a pair of zones, several testes

wide, to the right and left of the midline, each zone

occupying the posterior half of the prepharyngeal

region, extending posteriorly to the level of the

anterior part of the pharynx. The intestinal zone

ends anteriorly in a transversal border, not showing

any V-shaped formation.

ANATOMY.—Apart from the materials of this species deposited in the American Museum of Natural

History and in the United States National Museum,

I had an opportunity of studying several live specimens from the type-locality in Missouri and from

localities in Illinois.

The adhesive organ is a deep pit with irregularly

folded walls and is provided with a complex musculature, as described by Hyman (1945:479).

The reproductive system was examined in the

para type slides and in 12 sets of serial sections preserved by a better method than that used by L.

Hubricht for his original collection. The testes are

of moderate size, dorsal or subdorsal. The ovaries

are located usually at the level of the third or

fourth lateral branch of the anterior intestinal

ramus. Hyman's description of the copulatory complex needs some emendations. The genital pore

(Figure 58, gp) leads into a rather small common

genital atrium that connects anteriorly with the

male atrium (am) and dorsally with the widened

outlet (v) of the copulatory bursa. The penis consists of a rounded, muscular bulb and a large, generally finger-shaped papilla. Each vas deferens (vd),

after expanding to form the convoluted spermiductal vesicle, enters the penial bulb ventrolaterally, proceeds toward the midline and opens into

a rather large seminal vesicle (vs). What Hyman

(1945:480) describes as "narrow ejaculatory duct" is

actually the anteroposteriorly compressed, rather

voluminous seminal vesicle that extends laterally

to both sides. The epithelial lining of this vesicle is

of a glandular nature, with club-shaped cells protruding into the lumen. In the median section, the

vesicle generally arches from the anteroventral part

of the bulb dorsally and posteriorly and, at the

transition between bulb and papilla, connects with

a duct that runs through the axis of the papilla to

open at its tip. This canal, the true ejaculatory

duct (de), is lined with a normal, nonglandular epithelium and varies considerably in its width and

shape. It appears to have no surrounding muscular

layer. The outer surface of the penis papilla is

covered by a cuboidal epithelium, below which

there is a well-developed layer of fine circular fibers,

followed by a layer of coarser longitudinal muscles.

In some specimens, the tip of the penis papilla was

invaginated into the ejaculatory duct (Figure 59), so

that the distal part of the duct showed the same

muscular layers as the outer covering of the papilla.

It is difficult to decide whether this invagination

corresponds to the shape of the resting male organ

that may extend at the moment of preservation by

a contraction of the musculature of the penis papilla.

The spaceous male atrium (am) duplicates the

shape of the penis. At its posterior end it narrows

and connects with the small common atrium. The

copulatory bursa (6) shows no peculiarities. Its

outlet, the bursal duct or stalk (bd), begins as a

narrow, straight canal running posteriorly above

the penis bulb, then gradually widens, becomes

more convoluted, and finally bends downward as an
NUMBER 246 21

expanded seetion (v) situated to the left of the midline.

The two oviducts or ovovitelline ducts, which

accompany the ventral nerve cord, ascend dorsally

and medially at the level of the copulatory complex, each giving off a short posterior vitelline duct

running toward the yolk glands of the tail region.

The oviducts unite in the space above the male

atrium and proceed posteroventrally as common

oviduct (ode), provided with the usual eosinophilic

shell glands, which opens into the common atrium

from the dorsal side.

None of the epithelia of the copulatory complex

are infranucleate.

DISTRIBUTION AND ECOLOGY.—Sphalloplana hubrichti inhabits a number of caves and springs in

Missouri and Illinois. The following localities may

be taken as habitats of the species.

MISSOURI. JEFFERSON COUNTY: Spring near Kimmswick

and spring near Selma, collected by Leslie Hubricht in June

1937 (Hyman, 1945:480). SAINTE GENEVIEVE COUNTY: Kolms

Cave (this is the correct spelling, rather than Kohn's Cave as

Hyman indicates), type-locality of the species; many specimens collected by Leslie Hubricht in June 1937 (Hyman,

1945:480); about 35 specimens collected by Leslie Hubricht

13 September 1941 (alcohol material, USNM 50907); 6 specimens collected by Jerry Lewis and Margaret Meister 21 July

1974 and 12 April 1975.

ILLINOIS. JACKSON COUNTY: A walled spring in Happy

Hollow, Fountain Bluff, south of Gorham, a total of 17

specimens collected by Jerry Lewis and Margaret Meister 10

February, 2 March, 14 April, 14 September, and 20 October

1974. MONROE COUNTY: Burksville Cave (also called Morrisons

Cave, Eckerts Cave, Illinois Caverns, or Mammoth Cave of

Illinois), collected by Leslie Hubricht, June 1937 (Hyman,

1945:480); about 25 specimens collected by Jerry Lewis 24

November 1974, occurring together with 2 species of Asellus

and 4 species of amphipods. UNION COUNTY: Richs Cave, east

of Cobden, 5 specimens, collected by Jerry Lewis, 2 February

1974, besides Phagocata gracilis (Haldeman).

Other distributional data for S. hubrichti given

in the literature either need reexamination or are

misidentifications caused by Hyman's defective description of the species. Beatty's (1966:10) "Cave

near Cobden" is probably Richs Cave in Union

County, Illinois, mentioned above. Beatty's Tom

Moore Cave, Perry County, Missouri has Macrocotyla lewisi rather than S. hubrichti (see Kenk,

1975:333). McRitchie's (1959:20, 21) localities in

Tennessee are probably all erroneous: the species

occurring in the spring at Stokes Lane in Nashville

is 5. chandleri, described in this paper (p. 21).

Specimens from the spring "at junction of U.S.

routes 70N and 70S, Cheatham County," and from

Herring Cave, near Lascassas, Rutherford County,

are not further described by McRitchie; S. chandleri from the Stokes Lane spring in Nashville is

also mentioned by Darlington and Chandler (1972:

160) and by Chandler (1972:62) as "Speophila hubrichti."

TAXONOMIC POSITION.—By the development of a

conspicuous adhesive organ, 5. hubrichti belongs to

the subgenus Speophila. Carpenter (1970:85) placed

the species in synonymy with S. pricei, from which

it may be distinguished in life by a slightly different

shape of the head and by the transverse or rounded

anterior border of the intestinal zone (which in S.

pricei shows a V-shaped formation) and in the

anatomy by the location of the testes (dorsal in S.

hubrichti, dorsal and ventral in 5. pricei). From the

very similar S. chandleri it differs by the shape of

the anterior intestinal border and by the entry of

the vasa deferentia into the penis bulb (ventrolateral in S. hubrichti, dorsal after recurving in S.

chandleri). The remaining American species of the

subgenus differ from S. hubrichti chiefly in the location of the testes and the configuration of the

copulatory complex.

Sphalloplana (Speophila) chandleri, new species

FICURES 10, 26, 38, 60, 61

Speophila hubrichti.—McRitchie, 1959:20 [misidentification].

Sphalloplana (Speophila) pricei.—Carpenter, 1970:96 [in part].

TYPE MATERIALS.—Holotype, set of sagittal sections on 7 slides, USNM 53416. Paratypes, 10 sets

of sagittal and transverse sections on 62 slides,

USNM 53417-53426.

EXTERNAL FEATURES (Figures 10, 26).—A purely

white species of moderate size, up to 18 mm long

and 3 mm wide. Smaller individuals (12 mm long)

may also be sexually mature. The anterior end is

truncated, with a somewhat convex frontal margin

and rounded lateral corners lacking prominent

auricular projections. The protrusible adhesive organ is seen as a slightly opaque area and its outlet

may be noticeable as a small notch in the midline

of the frontal margin when the animal is gliding

quietly. There is no definite constriction or neck behind the anterior end, only a gentle narrowing of

the body margins when the animal is in locomotion. Then the body widens gradually, to narrow
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

again in the postpharyngeal region to a rounded or

bluntly pointed posterior end. The anterior portion

of the intestinal area forms a V-shaped border, the

lateral branches of the intestine reaching farther

forward to both sides of the adhesive organ than

the median intestinal trunk. The root of the

pharynx is located at about the middle of the body

length or slightly behind the middle. Its length is

about one-eighth the length of the body.

ANATOMY.—The adhesive organ (Figure 38), in

its retracted state, is a deep invagination with

folded walls, as is typical for the subgenus Speophila. The zone of marginal epithelium is also typically developed.

The two ovaries are situated below or behind

the first to third pair of lateral intestinal branches.

The numerous, rather small, rounded testes are

strictly dorsal, occupying, on either side of the midline, a longitudinal zone beginning some distance

behind the level of the ovaries and ending near the

pharyngeal root. The copulatory complex (Figure

60) was studied in 14 sets of serial sections. The

genital aperture or gonopore (gp) leads into the

common genital atrium (ac) that is not very clearly

marked off from the terminal part of the outlet (bd)

of the bursa copulatrix (b). Anteriorly, the common

atrium connects with the male atrium (am) surrounding the penis papilla. The penis appears in

the slides in various stages of muscular contraction

indicating the great plasticity of this organ. The

rounded, muscular penis bulb is of moderate size.

The penis papilla, when extended (Figure 60), is

long, finger-shaped, and larger than the bulb. In

several of the specimens, however, the posterior part

of the papilla is inverted into the penis lumen to a

varying degree or even telescoped (Figure 61). The

musculature of the penis papilla is restricted to a

two-layered coat underlying the external epithelium, while the penis lumen seems to be devoid of a

musculature of its own. The penial lumen consists

of a rather large cavity, lined with a tall, secretory

epithelium, the seminal vesicle (vs), located in the

penis bulb and extending into the basal part of the

penis papilla, and a canal (de) of variable diameter

running through the rest of the papilla and opening at its tip. This canal, corresponding to an

ejaculatory duct, has a nonglandular, cuboidal or

flattened epithelial lining. The two vasa deferentia

(vd) approach the copulatory apparatus as expanded, twisted spermiductal vesicles filled with

masses of sperm, run posteriorly to the level of the

male atrium, curve dorsally, enter the penis bulb

posterodorsally, and open into the seminal vesicle.

This recurving of the vasa deferentia is very characteristic and was seen in all specimens examined,

regardless of the state of contraction of the muscular organs. The common oviduct (ode) opens into

the posterior part of the male atrium from the

dorsal side, generally at a considerable distance

from the gonopore (in the specimens from Indiana

and Virginia more closely to the gonopore). The

copulatory bursa (b) is a rounded sac of variable

size. Its outlet, the bursa stalk (bd), widens gradually as it proceeds posteriorly, then becomes contorted and thrown into loops, and bends ventrally

to unite with the common atrium. The terminal

part of the bursal stalk does not change its histological characteristics. The muscle layer surrounding the stalk consists of intermingled circular and

longitudinal fibers.

None of the epithelia of the copulatory apparatus are infranucleate.

DISTRIBUTION AND ECOLOGY.—Sphalloplana chandleri was collected by Dr. Clay Chandler under

rocks in a stream issuing from a cave near Stokes

Lane in Nashville, Davidson County, Tennessee, on

22 August 1973 (water temperature, 16°C) and in

May 1975. A total of about 35 specimens were sent

to me alive. Some were placed in cultures maintained at about 14°C. Two cocoons were laid in the

cultures soon after the collection dates. The cocoons

are spherical, unstalked, W/2 mm in diameter. The

locality has been known to turbellarian workers for

some time. The presence of "Speophila hubrichti"

in the Stokes Lane spring had been reported by

McRitchie (1959:20) and by Darlington and Chandler (1972:60). McRitchie states that he collected a

very large specimen, 3 cm in length, on 5 May 1959.

Carpenter (1970:96), who considered S. hubrichti to

to be a synonym of S. pricei, listed S. pricei for the

same locality, collected in 1969. Other possible

habitats of "S. hubrichti" in Tennessee, mentioned

by McRitchie (1959:21) (spring at junction of U.S.

routes 70N and 70S, Cheatham County, and Herring Cave, near Lascassas, Rutherford County),

which also may refer to 5. chandleri, need further

examination.

INDIANA. Small spring next to the road on Edwardsville

Hill, west of New Albany, Floyd County. One specimen col-
NUMBER 246 23

lected by Jerry Lewis on 22 December 1974 (besides Phagocata gracilis).

VIRGINIA. Fallen Rock Cave, Tazewell County, located

5 miles south-southeast of Pounding Mill. A total of 7 specimens collected by John R. Holsinger and others on 13

October 1973, 27 July and 27 November 1974, and on 24

July 1976.

One cocoon, laid in a culture (kept at 14°C) in

June, hatched after about 12 weeks and nine young

emerged.

McRitchie (1959:20, 36, 37) comments on the gliding and leechlike locomotion and on the sensitivity

to light of the form from the Stokes Lane spring.

TAXONOMIC POSITION.—Sphalloplana chandleri,

apart from its well-developed adhesive organ that

places it in the subgenus Speophila, is characterized

by the lack of auricles, the V-shaped anterior intestinal border, dorsal position of the testes, recurving of the sperm ducts before dorsal entry into

the penial bulb, large penis papilla, nonmuscular

penial lumen, and lack of a distinctive vagina. In

life, it is very similar to S. pricei (lack of auricles,

V-shaped intestinal border), but differs from it by

the dorsal location of the testes and the recurving

of the vasa deferentia. Sphalloplana hubrichli,

which also has a similar outline of the anterior end,

has a different intestinal border and configuration

of the sperm ducts. It is interesting to note that

Carpenter (1970:88, 96) was aware of some of these

differences but did not consider them to be significant enough to be the basis of species separation.

The species is named in honor of the collector,

Dr. Clay M. Chandler of the Middle Tennessee

State University.

Sphalloplana (Speophila) mohri Hyman, 1938

FIGURES 16, 62

Sphalloplana mohri Hyman, 1938:137.

Sphalloplana kutscheri Mitchell, 1968:598.

Sphalloplana sloani Mitchell, 1968:600.

Sphalloplana zeschi Mitchell, 1968:604.

Sphalloplana reddelli Mitchell, 1968:607.

Sphalloplana (Polypharyngea) mohri.—Carpenter, 1971:1284.

TYPE MATERIAL.—Sphalloplana mohri, holotype,

whole mount, AMNH 650; paratypes, two specimens (one whole mount and one set of serial

sections on 24 slides), AMNH 651. S. kutscheri,

holotype, set of serial sections on 35 slides, USNM

38982; two paratypes. S. sloani, holotype, set of

serial sections on 36 slides, USNM 38980; three

paratypes. S. zeschi, holotype, set of serial sections

on 10 slides, USNM 38981; eight paratypes. S. reddelli, holotype, set of serial sections on 7 slides,

USNM 38979; four paratypes. The paratypes of

Mitchell's species are in the collection of Dr. R. W.

Mitchell.

Sphalloplana mohri was first briefly mentioned

by Hyman (1938:137) and later (1939:276-280)

more fully described. Mitchell (1968) established

four additional species on the basis of minor differences in the structure of the reproductive system,

which in my opinion do not justify their specific

differentiation from S. mohri. The published descriptions offer a sufficiently clear account of the

external aspects and the anatomy of the species.

The essential characters of the species are the following.

EXTERNAL FEATURES (Figure 16).—Sphalloplana

mohri is the largest species of the genus, mature

animals attaining a length of 20-30 mm and even

35 mm. Photographs of the species have been published by Mitchell (1974:413, a well-extended specimen, and pp. 415, 420, and 422, all "S. zeschi") and

earlier by Mohr (1948:17, "planarian"), and outline

drawings by Mitchell (1968:608). The anterior end

is truncate, with a straight or slightly bulging

frontal margin, the lateral edges bearing short

auricular projections extending laterally and anteriorly. The adhesive organ in the center of the

frontal margin may appear retracted or protruded

as a pointed conical structure. Behind the auricles,

the head narrows to some extent, then widens again

gradually to reach its maximum width. The anterior border of the intestinal area appears to have

a V-shaped outline as seen in Carpenter's (1970)

figure 13. An important character of the species is

its polypharyngy. Hyman (1939:276) gives the number of pharynges as being about 50, Mitchell (1968)

as 40 for all his species. The elongated pharyngeal

pouch, measuring about one-fourth to one-third the

body length, begins at a level anterior to the middle

of the body.

ANATOMY.—The adhesive organ, when retracted,

forms a rather deep invagination with irregular,

folded walls, as indicated by Mitchell (1968:612);

when protruded, it appears as a broad, pointed,

conical projection. Hyman (1939:276) states that

the epidermis along the lateral margins of the body

does not show the large rhabdites usually seen in
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the species of Sphalloplana. The marginal epithelium is thicker than elsewhere, however, and tapers

gradually toward the dorsal and ventral surfaces.

The testes, in moderate number, are arranged in

a pair of longitudinal rows beginning some distance

behind the head and ending at the level of the

more anterior pharynges. In Mitchell's type slides

they are seen in a predominantly dorsal position.

The copulatory apparatus (Figure 62) in the various

specimens studied by Mitchell appears to be subject to small variations, probably due to various

states of muscular contraction. The penis has a

small, not very muscular bulb and a larger, cylindrical or conical papilla with rounded or more or

less pointed tip. The vasa deferentia (vd) enter the

anterior side of the bulb, traverse it, and unite

within the papilla to a straight canal, the ejaculatory duct (de), which opens at the tip of the papilla.

There is no distinct seminal vesicle developed, although the ejaculatory duct may show some local

widening such as that seen in Hyman's figure 4,

which she interprets as a seminal vesicle. The common oviduct (ode) opens into the posterior part of

the male atrium from the dorsal side. The bursal

duct (bd) proceeds from the copulatory bursa (b)

posteriorly above the penis as a straight, narrow

canal, then bends ventrally at the level of the gonopore (gp), widens somewhat, and acquires a thick

muscular coat. This terminal section of the duct

(v) may be considered to be a vagina.

DISTRIBUTION AND ECOLOGY.—Sphalloplana mohri

has been reported from several caves in Texas.

Ezell's Cave (type-locality) in Hays County, collected by Charles E. Mohr and Kenneth Dearolf, 19

June 1938 (Hyman, 1939:280), also recorded by

Mitchell (1968:609). Mitchell's species are from four

other caves of the Edwards Plateau of Texas

(Mitchell, 1968: S. kutscheri, Spanish Wells Cave,

Travis County, collected by Mitchell, 9 June 1967;

S. sloani, Harrell's Cave, San Saba County, collected

by Mitchell, 14 October 1967; S. zeschi, Zesch

Ranch Cave, Mason County, collected by Mitchell,

10 June 1968 and other dates; and S. reddelli, Cascade Caverns, Kendall County, collected by James

Reddell, 8 April 1966). Eigenmann (1900:229 and

1902:84-85) reported the finding of an unidentified

flatworm in the outflow of an artesian well in San

Marcos, Hays County. Dr. Glenn Longley brought

me a preserved, slightly damaged specimen of a

polypharyngeal Sphalloplana from this well, collected 28 October 1974 by Joe Kolb. The specimen

was not fully mature sexually, but agreed in all

recognizable characters with S. mohri.

A few data on the habitat of S. mohri have been

presented by Hyman (1939:280), whose material

was from a shallow pool in Ezell's Cave with a

water temperature of 71°F-72°F [about 22°C].

Mitchell (1974) gives more data on the ecology of

his S. zeschi in Zesch Ranch Cave. The worms occur

there in an intermittent pool with a temperature

of 21°C-21.5°C, together with the amphipod, Stygonectes russelli Holsinger, and an ostracod, Candona sp. Their natural food appears to consist of

injured or moribund amphipods and of any other

arthropods (cave crickets, flies, etc.) that may have

fallen into the water.

LABORATORY STUDIES.—Mitchell (1974) performed

some very interesting experiments on S. zeschi. He

analyzed the types and speed of movement, the

righting time from an inverted position, attraction

to food and mode of feeding, negative rheotaxis,

responses to light, and temperature tolerance and

preference.

TAXONOMIC POSITION.—The well-developed adhesive organ places S. mohri in the subgenus Speophila. It shows, however, certain characters that

differ from the general features of this subgenus.

The absence of distinct zone of tall epithelial cells

on the lateral margins of the body appears to be

unique within the genus Sphalloplana. The polypharyngy, also a unique character, should not be

given great systematic value, as it occurs in other

genera (Phagocata, Crenobia) as a specific or even

subspecific feature. It is, however, a good distinguishing character of the species. The small differences in the anatomy of the forms described by

Mitchell (1968) are, in the writer's judgment, well

within the framework of the expected variations

caused by contractions and distortions of muscular

organs in the process of fixation, particularly when

different methods of killing (nitric acid, formalin)

are employed.
Literature Cited

Ban, T. C.

1961. Caves of Tennessee. Tennessee Division of Geology

Bulletin, 64: 569 pages, 1 folding map.

1968. Ecological Studies in the Mammoth Cave System of

Kentucky, I: The Biota. International Journal of

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Barr, T. C, and R. A. Kuehne

1971. Ecological Studies in the Mammoth Cave System of

Kentucky, II: The Ecosystem. Annales de Spileologie,

26:47-96.

Beatty, J. A.

1966. Key to the Land and Fresh-Water Triclad Turbellaria of North America (for Living Specimens Only).

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Beauchamp, P. de

1931. Biospeologica, LVI: Campagne speologique de C.

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113-380, plates 6-8.

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1931. Biospeologica, LVI: Campagne speologique dans

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Buchanan, J. W.

1935. The Eyeless Cave Flatworm, Sphalloplana percaeca.

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Carpenter, Jerry H.

1969. A New Killing and Fixing Technique for Small Animals. Transactions of the American Microscopical

Society, 88:450-451.

1970. Systematics and Ecology of Cave Planarians of the

United States. 213 pages. Ph.D. dissertation, University of Kentucky [unpublished, reproduced from

typewritten copy].

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United States. Dissertation Abstracts International,

(B)32:1283-1284. [This is the published abstract of

the above paper, Carpenter, 1970.]

1973. Systematics and Biogeography of Cave Flatworms.

Bulletin of the National Speleological Society, 35:15.

Castle, W. A., and L. H. Hyman

1934. Observations on Fonticola velata (Stringer), including a Description of the Anatomy of the Reproductive System. Transactions of the American Microscopical Society, 53:154-171.

Chandler, C. M.

1972. Planarians of Tennessee. Journal of the Tennessee

Academy of Science, 47:62.

Darlington, J. T., and C. M. Chandler

1972. A Survey of the Epigean Triclad Turbellarians of

Tennessee. American Midland Naturalist, 88:158-

166.

Dearolf, K.

1941. The Invertebrates of 35 Pennsylvania Caves. Proceedings of the Pennsylvania Academy of Science,

15:170-180.

1953. The Invertebrates of 75 Caves in the United States.

Proceedings of the Pennsylvania Academy of Science,

27:225-241.

Eigenmann, C. H.

1900. A Contribution to the Fauna of the Caves of Texas.

Proceedings of the American Association for the

Advancement of Science, 49:228-230.

1902. In C. J. Ulrich, A Contribution to the Subterranean

Fauna of Texas. Transactions of the American

Microscopical Society, 23:84-85.

Feinberg, Harold S.

1970. A Catalogue of Type Specimens in the Department

of Living Invertebrates, The American Museum of

Natural History, New York, New York, U.SA. The

Phyla Platyhelminthes and Gnathostomulida. 66

pages. New York: American Museum of Natural History [reproduced from typewritten copy].

Ginet, R., and R. Puglisi

1964. £cologie de Fonticola notadena de Beauchamp (Turbellaric, Triclade) dans la grotte de La Balme (Isere,

France); survie en periode de se'cheresse. International Journal of Speleology, 1:203-216.

Giovannoli, L.

1933. Invertebrate Life of Mammoth and Other Neighboring Caves. In V. Bailey, Cave Life of Kentucky,

Mainly in the Mammoth Cave Region. American

Midland Naturalist, 14:600-623.

Gourbault, N.

1965. Amyadenium chattoni, Triclade obscuricole: Donnees

preliminaires sur la biologie et la duree de la regeneration. Annales de Speh'ologie, 20:477-481.

1972. Recherches sur les Triclades paludicoles hypoges.

Memoires du Museum National d'Histoire Naturelle

(Paris), nouvelle se>ie, serie A (Zoologie), 73:1-249, 1

folding map, plates 1-3.

Holsinger, J. R.

1966. A Preliminary Study on the Effects of Organic Pollution of Banners Corner Cave, Virginia. International

Journal of Speleology, 2:75-89.

Holsinger, J. R., and T. E. Bowman

1973. A New Troglobitic Isopod of the Genus Lirceus

(Asellidae) from Southwestern Virginia, with Notes

25
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

on Its Ecology and Additional Cave Records for the

Genus in the Appalachians. International Journal of

Speleology, 5:261-271.

Holsinger, J. R., and S. B. Beck

1971. The Invertebrate Cave Fauna of Georgia. National

Speleological Society Bulletin, 33:23-44.

Hyman, L. H.

1931. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, IV:

Recent European Revisions of the Triclads, and

Their Application to the American Forms, with a

Key to the Latter and New Notes on Distribution.

Transactions of the American Microscopical Society,

50:316-335.

1935. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, VI:

A New Dendrocoelid from Montana, Dendrocoelopsis vaginatus n. sp. Transactions of the American

Microscopical Society, 54:338-345.

1936. Some Cave Planarians of the United States. Anatomical Record, 67, supplement 1:129.

1937. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, VIII:

Some Cave Planarians of the United States. Transactions of the American Microscopical Society, 56:

457-477.

1938. Additional North American Cave Planarians. Anatomical Record, 72(4), supplement, page 137.

1939. North American Triclad Turbellaria, X: Additional

Species of Cave Planarians. Transactions of the

American Microscopical Society, 58:276-284.

1945. North American Triclad Turbellaria, XI: New,

Chiefly Cavernicolous, Planarians. American Midland Naturalist, 34:475-484.

1954. North American Triclad Turbellaria, XIII: Three

New Cave Planarians. Proceedings of the United

States National Museum, 103(3333):563-573.

Ichikawa, A., and M. Kawakatsu

1967. Records of Two Planarian Species of the Family

Kenkiidae from Japanese Subterranean Waters.

Archiv fur Hydrobiologie, 63:512-519.

Kawakatsu, M.

1969. An Illustrated List of Japanese Freshwater Planarians in Color. Bulletin of Fuji Women's College

(Sapporo), series 2. 7:45-91.

Kawakatsu, M., and T. Miyazaki

1972. Effect of Different Fixatives on a Common Japanese

Freshwater Planarian, Dugesia japonica Ichikawa et

Kawakatsu. Bulletin of Fuji Women's College (Sapporo), series 2, 10:81-117.

Kenk, R.

1970. Freshwater Triclads (Turbellaria) of North America,

III: Sphalloplana weingartneri, New Species, from a

Cave in Indiana. Proceedings of the Biological Society of Washington, 83:313-320.

1972. Freshwater Planarians (Turbellaria) of North America. Biota of Freshwater Ecosystems, Identification

Manual. Number 1, ix + 81 pages. Washington:

Environmental Protection Agency.

1975. Freshwater Triclads (Turbellaria) of North America,

VII: The Genus Macrocotyla. Transactions of the

American Microscopical Society, 94:324-339.

Lascombe, Claude

1971. Recherches ecologiques et biogeographiques sur deux

especes jumelles de Planaires d'eau douce dans la

rigion lyonnaise. [vii] + 116 pages, 20 plates. Thesis,

Universite Claude Bernard (Lyon) [unpublished, reproduced from typewritten copy.]

Livanov, N. A., and Z. I. Zabusova

1940. Planarii basseina Teletskogo ozera i novye dannye o

nekotorykh drugikh sibirskikh vidakh (Paludicoles

du Lac Teletzkoe et nouvelles donnees sur quelques

formes sibcricnnes). Trudy Obshchestva Estestvoispytatelei pri Kazanskom Gosudarstvennom Universitete, 56(3-4):8S-159.

McRitchie, Robert Gerald

1959. The Kenkiidae of the Nashville Area, v + 43 pages,

10 plates. M.A. thesis, Vanderbilt University [unpublished, reproduced from typewritten copy].

Mitchell, R. W.

1968. New Species of Sphalloplana (Turbellaria: Paludicola) from the Caves of Texas and a Reexamination

of the Genus Speophila and the Family Kenkiidae.

Annales de Spiliologie, 23:597-620.

1974. The Cave-Adapted Flatworms of Texas: Systematics,

Natural History, and Responses to Light and Temperature. Pages 402-430 in Riser and Morse, editors,

Biology of the Turbellaria. New York, etc.: McGrawHill Book Company.

Mohr, C. E.

1948. Unique Animals Inhabit Subterranean Texas. National Speleological Society Bulletin, 10:15-21, 88.

Packard, A. S.

1879. Zoology for Students and General Readers, viii +

719 pages. New York: Henry Holt and Company.

1880. Zoology for High Schools and Colleges. Second edition, viii + 719 pages. New York: Henry Holt and

Company. [Identical with the preceding publication,

with changed title page.]

1888. The Cave Fauna of North America, with Remarks

on the Anatomy of the Brain and Origin of the

Blind Species. Memoirs of the National Academy of

Sciences (Washington), 4(1):3-159, plates 1-27,1 map.

Steinmann, P.

1910. Eine neue Gattung der paludicolen Tricladen aus

der Umgebung von Basel (Polycladodes alba n. g.

n. sp.). Verhandlungen der Naturforschenden Gesellschaft in Basel, 21:186-196.

Wells, P. H., and C. O. Harris

1954. Ultraviolet Sensitivities of Pigmented and Colorless

Flatworm Species. Anatomical Record, 120:793.
NUMBER 246 27

TABLE 1.—Comparison of the North American species of Sphalloplana

Sphalloplana

pereoeea

georgiana

evaginata

kansensis

californica

culvert

consimilis

subtilis

virginiana

holsingeri

weingartneri

buchanani

pricei

hubrichti

chandler!

mohri

bution

KY, TN

AL

GA

MO

KS

CA

WV

VA, TN

VA

VA

VA

IN

KY

PA

MO.IL

TN, IN

VA

TX

prominent

t

prominent

?

absent

small

prominent

very

small

?

small

absent

absent

absent

absent

absent

small

organ

weak

weak

weak

moderate

weak

weak

moderate

weak

moderate

strong

strong

strong

strong

strong

strong

strong

border

rounded

?

rounded

?

rounded

rounded?

rounded

rounded

?

rounded

rounded

V-shaped

V-shaped

square

V-shaped

?

Testes

dorsal

dorsal?

dorsal

dorsal

predom.

ventral

ventral

dorsal

ventral

ventral

ventral

dorsal &

ventral

dorsal

dorsal,

ventr.,4

interned.

predom.

dorsal

dorsal

predom.

dorsal

Penis

papilla

plugshaped

conical

large,

plugshaped

small

long,

conical

large,

plugshaped

plugshaped

plugshaped

plugshaped

plugshaped

small,

fingershaped

conical

or cylin.

long,

fingershaped

long,

fingershaped

long,

fingershaped

conical

or cylin.

Seminal

vesicle

small

small

absent

absent

paired

small

absent

absent

narrow

present

absent

narrow

present

present

absent

EJacul.

duct

opens

at tip

opens

at tip

opens

at tip

opens

at tip

opens

at tip

opening

dorsal

opening

ventral

opening

ventral

ventral

opening

dorsal

opens

at tip

opens

at tip

opens

at tip

opens

at tip

opens

at tip

opens

at tip

Oviduct

opening

far

dorsal

far

dorsal

far

dorsal

common

ov.long

close to

gonopore

intermediate

intermediate

intermediate

dorsal

close to

gonopore

close to

gonopore

?

intermediate

close to

gonopore

intermediate

intermediate

Vagina

wide

wide

widening

widening

widening

large

widening

.prominent

narrow

large

narrow

widening

convoluted

convoluted

convoluted

narrow

Remarks

pink color, cylindrical diverticula

on common atrium

irregular diverticula on common atrium

prostate in penis bulb,

double seminal vesicle

wide-spread infranucleate epithelia

very slender shape

no common atrium

large penis bulb

no common atrium

vasa defer, recurve,

enter bulb posterodorsally

large, polypharyngeal

a

ac

am

b

bd

bp

da

de

f

gl

gP

i

inv

m

ABBREVIATIONS USED IN

genital atrium

common genital atrium

male atrium

copulatory bursa

bursal duct

penis bulb

atrial diverticulum

ejaculatory duct

fibrous layer

gland duct

gonopore

intestine

invaginated part of

penis papilla

mouth

ILLUSTRATIONS

me marginal epithelium

od oviduct

ode common oviduct

pb penis bulb

ph pharynx or pharyngeal pouch

pi penial lumen

pp penis papilla

pr prostate

t testis

v vagina

vd vas deferens or sperm duct

ve efferent ductule

vi vitellarium or yolk gland

vs seminal vesicle
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

n

i

n <n

FIGURES 1-10.—Sphalloplana, photographs of living specimens: 1, S. percoeca from Williams

Saltpeter Cave, Alabama, X 4.4; 2, S. evaginata from Garbage Hole Cave, Missouri, X 3.7; 3,

S. californica, immature, from Bower Cave, California, X 4; 4, S. californica, mature, from

Bower Cave, California, X 4.9; 5, S. consimilis from Gallohan Cave No. 1, Virginia, X 4.8; 6,

S. subtilis from Biggers Spring, Virginia, X 4.4; 7, S. holsingeri from Biggers Spring, Virginia,

X 4.9; 8, S. pricei from Refton Cave, Pennsylvania, X 3.5; 9, S. hubrichti from Kolms Cave,

Missouri, X 3.6; 10, S. chandleri from Nashville, Tennessee, X 4.7.
NUMBER 246 29

13

18 19 20

lmm

FIGURES 11-20.—Sphalloplana, outline drawings of living specimens, indications of pharynx

and anterior intestinal border: 11, S. percoeca, redrawn from Buchanan (1936:196), X

6; 12, S. evaginata, X 3.6; 13, S. subtilis, X 6; 14, S. hohingeri, X 6; 15, S. buchanani, redrawn

from Hyman (1937, fig. 16), X 7.3; 16, S. mohri, redrawn from Mitchell (1974:413, "5. zeschi"),

used by permission of McGraw-Hill Book Company, X 3. 17, S. culvert, anterior end; 18,

5. hohingeri, anterior end, showing intestinal area; 19, S. pricei, anterior end, showing intestinal

area; 20, 5. evaginata, anterior end, drawn from whole mount.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 21-26.—Sphalloplana, outline drawings of living specimens, with indications of pharynx

and anterior intestinal border: 21, 5. californica, X 6; 22, 5. consimilis, X 6.6; 23, S. weingartneri (redrawn from Carpenter's photograph), X 11.7; 24, S. pricei, X 4.3; 25, 5. hubrichti,

X 4.3; 26, S. chandleri, X6.
NUMBER 246 91

f<-V,?\ • '

me

31 32

FIGURES 27-32.—Biggers Spring, Fairfax County, Virginia, type-locality: 27, 28, Sphalloplana subtilts and S. holsingeri. Transverse section of prepharyngeal region, showing differentiation of

the marginal epithelium, X 186: 29, Sphalloplana subtilis. Adhesive organs in sagittal section:

SO, S. percoeca, X 153; 31, S. evaginata, X 55; 32, S. califomica, X 175.
32

a.UTHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 33-38.—Adhesive organs in sagittal section: 33, 5. culveri, X 139; 34, 5. consimilis, X

109; 35, S. subtilis, X 180; 36, S. hohingeri, not fully retracted, X 56; 37, S. pricei, X 62; 38,

S. chandleri, X 64.
NUMBER 246

sirtoj^

l ^

•

43 vd

FIGURES 39-44.—Sphalloplana, parasagittal section

of prepharyngeal region, showing location of testes

39, S. californica, X 61; 40, S. culvert, X 174; 41, S

consimilis, X 61; 42, 5. subtHis, X 146; 43, S. hoi

singeri, X 73; 44, S. pricei, X 77. 44
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0.1mm

45 b

vd bp

ode ac

FIGURES 45-48.—Semidiagrammatic views of copulatory apparatus in sagittal section: 45, S.

percoeca from Great Onyx Cave, Kentucky; 46, 5. georgiana, after Hyman (1954:569), modified;

47, 5. evaginata from Klump Cave, Missouri; 48, S. kansensis, after Hyman (1945:483), modified.
NUMBER 246 35

0.5mm

gP

0.5mm ode

50

51

0.5 mm bd ode

vd gl de PP gP

FIGURES 49-51.—Semidiagrammatic views of copulatory apparatus in sagittal section: 49, S.

californica from Bower Cave, California; 50, S. culvert from Harper Cave, West Virginia; 51,

S. consimilis from Gallohan Cave No. 1, Virginia.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0.5 mm am ode bd

ode

54

FIGURES 52-55.—Semidiagrammatic views of copulatory apparatus in sagittal section: 52, S. subtilis from Biggers Spring, Virginia; 53, S. virginiana, after Hyman (1945:483), modified; 54, S.

holsingeri from Biggers Spring, Virginia; 55, S. weingartneri, after Kenk (1970:316).
NUMBER 246 37

b pb vd bd pi od ode pp v vd vs bp de pp inv

0.5 mm

bd de ode

am PP ac gP

0.5 mm

58

FIGURES 56-59.—Semidiagrammatic views of copulatory apparatus: 56, S. buchanani, after Hyman (1937:467), dorsal view; 57, S. pricei from Refton Cave, Pennsylvania, sagittal section; 58,

S. hubrichti from Richs Cave, Illinois, sagittal section; 59, S. hubrichti from Kolms Cave, Missouri, penis, sagittal.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0.5 mm

gp ode bd

0.5 mm

m gp ode bd

62 ode gp

FIGURES 60-62.—Semidiagrammatic views of copulatory apparatus in sagittal section: 60, S.

chandleri from Nashville, Tennessee; 61, S. chandleri from Nashville, Tennessee; 62, S. mohri,

after Mitchell (1968:606, S. reddelli), modified.


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