Biota of Freshwater Ecosystems
Identification Manual No. 1
FRESHWATER PLANARIANS (TURBELLARIA) OF NORTH AMERICA
by
Roman Kenk
Research Associate
Department of Invertebrate Zoology
Smithsonian Institution
Washington, D. C. 20560
for the
ENVIRONMENTAL PROTECTION AGENCY
Project # 18050 ELD
Contract # 14-12-894
February 1972
For sale by the Superintendent of Documents, VS. Government Printing Office
Washington, D.C. 20402 - Price $2.50
Stock Number 5501-0365
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EPA Review Notice
This report has been reviewed by the Environmental
Protection Agency and approved for publication.
Approval does not signify that the contents
necessarily reflect the views and policies of
the EPA, nor does mention of trade names or
commercial products constitute endorsement
or recommendation for use.
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Reports should be directed to the Chief, Publications Branch
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Protection Agency, Washington, DC 20460.
ENVIRONM21TTA1 PROTECTION AGENCY
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FOREWORD
"Freshwater Planarians (Turbellaria) of North America" is the
first of a series of identification manuals for selected taxa
of invertebrates occurring in freswater systems. These docu-
ments prepared by the Oceanography and Limnology Program,
Smithsonian Institution, for the Environmental Protection
Agency will contribute toward improving the quality of the
data upon which environmental decisions are based.
Additional manuals will include, but not necessarily be limit-
ed to, freshwater representatives of the following groups:
amphipod crustaceans (Gammaridae), branchiuran crustaceans
(Argulus'), isopod crustaceans (Asellidae), decapod crayfish
crustaceans (Astacidae), leeches (Hirudinea), polychaete worms
(Polychaeta), aquatic dryopoid beetles (Dryopoidea), and
freshwater clams (Sphaeriacea).
111
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ABSTRACT
A key is presented for the identification of the species of North
American freshwater triclads or planarians known at present.
Introductory chapters deal with the collecting, culturing, pres-
ervation, study, and general organization and life cycle of
planarians. The key is followed by a listing of the species and
subspecies, giving their distinguishing characteristics, ecological
requirements, and geographic ranges. Illustrations depict the
external appearance and diagrams of the reproductive organs of the
individual taxa. The principal literature for each species is
indicated and listed in the appended bibliography of 65 items. An
index of the generic and specific names and synonyms concludes the
report. One new subspecies, Polyoelis ooronata brewipenis , is es-
tablished for L. H. Hyman's Polyoel-is
This report was submitted in fulfillment of Project #18050ELD,
Contract #14-12-894 under the sponsorship of the Environmental
Protection Agency,
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CONTENTS
Section Page
I Introduction 1
Collecting 2
Transporting and Culturing 3
Preservation 4
Preparation for Anatomical Study 4
General Features 5
Reproduction 9
Characters Used in Identification 11
II Key to the Genera 13
III Key to the Species 15
IV Species Characteristics and Ranges 19
Not Recognizable Species 66
V Abbreviations Used on Figures 69
VI Acknowledgments 71
VII References 73
VIII Index of Scientific Names 79
VII
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FIGURES
Page
1 Baiting of planarians in lakes 2
2 Digestive system of a planarian 5
3 Reproductive system 6
4 Egg capsules of planarians 9
5 Muscle layers of pharynx in cross section 13
6,7 Cura foremanii 19
8-10 Dugesia tigrina 20
11-12 Dugesia dorotoeephala 22
13,14 Dugesia antillana 23
15,16 Dugesia polyehora 24
17,18 Polyoelis coronata ooronata 25
19 Polyoelis oovonata brevipenis new subspecies 26
20 Poly delis oovonata bovealis 27
21,22 Planaria daotyligera daotyligera 28
23 Planaria daotyligera rrusoulosa 29
24,25 Planaria ooeulta 30
26,27 Eymanella retenuova 31
28,29 Phagoaata oregonensis 32
30 Phagoeata monophavyngea 33
31,32 Phagoeata vevnalis 34
33,34 Fhagocata velata 35
35,36 Phagooata bulbosa 36
37,38 Phagoeata bursaperforata 37
39,40 Phagocata tahoena 38
41,42 Phagoeata erenophila 39
43,44 Phagooata morgani morgani 40
45 Phagooata morgani polyoelis 41
46,47 Phagooata nivea 42
48,49 Phagoeata graoilis 43
50,51 Phagooata woodaorthi 44
52,53 Sphalloplana perooeoa 45
54,55 Sphalloplana aldbamensis 46
56,57 Sphalloplana geovgiana 47
58,59 Sphalloplana virginiana 48
60 Sphalloplana kansensis 49
61,62 Sphalloplana pricei 50
63,64 Sphalloplana buehanani 51
65,66 Sphalloplana hubriohti 52
67,68 Sphalloplana hoffmasteri 53
69,70 Sphalloplana weingavtneri 54
71,72 Sphalloplana mohri 55
73-75 Kerikia rhynohida 56
76,77 Macroeotyla glandulosa 57
viii
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78,79 Dendroooelopsis vaginata 58
80,81 Dendrocoelopsis piviformis 59
82,83 Dendroeoelopsis ataskensis 60
84 Dendroaoelopeis hymanae 61
85-87 Dendrocoelopsis amevioana 62
88,89 Proeotyla fluviatilis 63
90,91 Proeotyla typhlops 64
92,93 Reetoeephala exotica 65
94 Planaria simplex 66
95 Planaria unionicola 66
96 Hydpolimaac bruneus 67
IX
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SECTION I
INTRODUCTION
The present report is a review of the species and subspecies of the
North American freshwater triclads or planarians known at the present
time. It should not be considered to give a definitive picture of
the planarian fauna of the geographic area covered, north of Mexico,
as many parts of the continent are still little explored with regard
to lower aquatic invertebrates. This is particularly true for large
sections of the West and South of the United States and the greater
part of Canada.
It was considered appropriate not to limit the review to a simple key.
When attempting to identify a given species through a key, one ends up
with a specific name which one must accept on good faith. Should one
happen to have an undescribed species at hand, it would, in most cases,
key out to a known species. In order to avoid this pitfall, every
species is treated separately, its distinguishing external appearance
and anatomical features are described, and the principal literature
sources which may lead to a more detailed description of its morphology
and ecology are listed. In this way it should be possible to avoid the
uncertainties inherent in the use of simple dichotomic keys and to have
a means of either confirming or questioning the correctness of the
identification.
In addition to the valid names of the individual species, all synonyms
which have been used for the American planarians are listed in the
species descriptions, giving the name of the author and the date of
the first publication of each synonym.
Planarians have played a certain role in the biological assessment of
water quality in Europe. Some species have been found to be very sen-
sitive to organic and inorganic pollution of their habitats while others
tolerate mild degrees of pollution. In general it may be said that
planarians are intolerant to the presence of heavy metal salts in the
water. Little has been done in the study of American turbellarians with
regard to their behavior toward pollutants. The American fauna of
planarians differs from that of Europe, and no native species is common
to both continents. One American species was introduced in Europe at the
beginning of this century, and one imported European species was dis-
covered recently in North America. It is, therefore, not possible to
apply the results obtained by European workers to the American planarian
fauna without detailed field observations.
It is hoped that the present report will be useful in any future system-
atic investigation using the presence or absence of planarians in the
evaluation of water quality in North America.
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COLLECTING
Planarians are, in general, negatively phototactic animals, hiding in
daytime in dark places. They may be collected by examining the under-
sides of flat stones and other objects (fallen leaves, pieces of wood
etc.) or the stems of submerged or partially submerged water plants.
The planarians are then removed from the substrate with a soft paint-
brush or the moistened ball of a finger (carefully, without exerting
any pressure), placing them immediately in a jar containing water taken
from the same habitat. Where there is dense vegetation or an accumu-
lation of leaf litter or other debris, samples of these may be taken in
a glass jar filled with water; after keeping the jar in a cool place,
usually overnight, planarians, if present, will tend to accumulate in
the upper layer of the water and may be collected with a wide-mouthed
syringe fitted with a rubber bulb. Many planarian species can be
attracted by bait (a piece of liver, meat, a dead frog, fish, crushed
snails or earthworms) placed under a flat stone and examined after
several hours. A very effective method is that of placing the bait in
a glass or plastic jar with a lid bearing many small round perforations
(of 3-5 mm diameter). The jar is then submerged in a shaded location in
a stream, pond, or lake and left for some time. Upon retrieval, usually
after 24 hours, planarians will be found inside the jar while larger scav-
engers, such as crayfish, have been kept out. This method is applicable
also in lakes at greater depths if the bait jar is submerged, together
Fig. 1. Baiting of planarians in
lakes, ba, canvas bag with stones;
fl, float; j, jar with perforated
lid containing bait.
with some anchoring ballast, and attached by a line to a float (Fig. 1).
Not all planarian species will be attracted to dead bait. Some species,
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e.g. Procotyla fluviatilis and some subterranean planarians, apparently
do not react to chemical food stimuli but rather to vibrations produced
by their living prey, such as small crustaceans and insect larvae. Other
methods of securing deep-water planarians are by means of a bottom grab
sampler or by a trawling dredge equipped with a canvas bag which skims
the upper layer of mud or silt; the planarians may be separated by using
sieves of proper mesh size.
TRANSPORTING AND CULTURING
Some species of planarians can be indentified by an experienced collector
with reasonable certainty in the field. In many cases, however, it is
necessary to transport them to a laboratory or even to send them,
preferably alive, to a specialist. As many planarians, particularly
those living in cold springs, deep lakes, or in subterranean habitats
(caves, groundwater), are very sensitive to temperature fluctuations
(stenothermic), they should be transported in well-insulated containers.
Vacuum-insulated (thermos) bottles are quite appropriate and may be used
also for the shipping by air mail (special delivery) across the continent,
provided they are taken care of immediately at the point of destination
(they should not arrive at a laboratory on a weekend).
Some planarians cannot be identified by their external features alone
but require the study of anatomical characters, specifically an analysis
of their reproductive organs. Unfortunately, many of the specimens col-
lected in the field are sexually immature. It is important, therefore,
to select the largest individuals from a given population for further
study and identification. Should there be no mature animals available,
it is often possible to raise young animals to maturity in laboratory
cultures.
Culturing is carried out in shallow glass aquaria (finger bowls) or in
enameled pans kept in the dark or in dim light at proper temperatures,
according to the tolerance limits of the species. The water should be
changed every two or three days when kept at room temperature and about
once a week in refrigerated cultures. Spring water or filtered pond
water may be used, but chlorinated tap water should be avoided unless
the chlorine has been removed by appropriate chemicals (e.g., sodium
thiosulfate, the photographic "hypo"), by letting it stand for 24 hours
in an open container, or by bubbling air through it. The planarians may
be fed once or twice a week. Most species will accept small pieces of
liver, meat, clotted blood, cut-up earthworms, or living food such as
the oligochaete Tubifex (obtainable in pet stores as fish food). A few
species refuse dead food but attack living small crustaceans (Gammarua^
Asellus, Dophnia, etc.). The food is left in the aquarium for several
hours, then removed, the water changed, and, if necessary, the aquarium
cleaned. Planarians are capable of surviving extended periods of
starvation, several months, particularly when kept at low temperature.
When starving, they gradually grow smaller, their anatomical makeup is
simplified, their entire reproductive system is reduced, and they become
indistinguishable from young animals.
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PRESERVATION
The soft, very delicate and flexible body of planarians can easily be
injured and distorted. It is very difficult to preserve its natural
shape, particularly the shape of the anterior end or head, which is of
some taxonomic importance. Animals killed with alcohol or formalin are
often badly contracted and twisted and are difficult to study. The
fixing method should preserve the animal well extended. The method
recommended by Hyman (1953a:128) is to place the planarians in a very
small amount of water, kill them stretched out with 2% nitric acid
followed by 70% alcohol (or any other fixative). De Beauchamp
(1932:126) uses a mixture of ethyl alcohol (70%, 7 parts), formalin
(2 parts) and glacial acetic acid (1 part) into which the animals are
dropped individually; after several hours the fixative is replaced with
70% alcohol. Recently Carpenter (1969b) has developed a method of rapid
freezing of the extended animals. My favored method is to kill the ex-
tended planarians, while they are in gliding motion: in very little water,
by pouring over them a hot, almost boiling, fixative, preferably a
saturated solution of corrosive sublimate (mercuric oxide, HgCl2) in
water or saline, with a subsequent addition of a few drops of diluted
acetic acid; after fixing for 4-24 hours the animals are first washed in
water, then transferred to increasing strengths of alcohol until a concen-
tration of 70% to 80% is reached in which they may remain until further
treatment; the last traces of sublimate are removed by adding small
amounts of tincture of iodine to the alcohol until the color remains
stable.
None of these methods gives ideal results since there will always be some
contraction or distortion of the body or of internal organs. For ana-
tomical study, simple shrinkage is better than twisting.
PREPARATION FOR ANATOMICAL STUDY
Whole mounts of planarians, stained or unstained, may be prepared by the
usual techniques but are of limited value in the analysis of anatomical
structures. They will show the number and arrangement of the eyes, the
configuration of the digestive system, and possibly some parts of the
reproductive system (testes, ovaries, copulatory organs). Details of
the anatomy must be studied in microtome sections.
The specimens are embedded in paraffin or a combination of celloidin and
paraffin and serial sections of 5-10 microns thickness are prepared.
For the purpose of species identification, sagittal sections are most
suitable, transversal and horizontal sections may be useful if sufficient
material is available. The choice of histological stains is optional
according to the preference of the investigator. A good staining method
for the general study and the analysis of muscles and glands is a combi-
nation of hematoxylin and eosin.
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GENERAL FEATURES
Planarians or freshwater triclads are elongated, flattened Turbellaria,
the native American species measuring in length from about 2 mm (freshly
hatched) to perhaps 40 mm. In the extended, quietly gliding animal one
distinguishes an anterior portion or head, often characteristically
shaped, bearing important sense organs. The lateral margins of the head
may protrude as auricles (which have chemical and mechanical sensory
receptors). There are usually two dark eye spots on the head, each
located near the medial margin of a white area, often accompanied by
smaller accessory or supernumerary eyes. Some species have numerous
eyes and subterranean planarians are often eyeless or blind. A median
adhesive organ may be differentiated on the frontal margin of the head.
The body may or may not be pigmented, often exhibiting various patterns
of pigment arrangement. The mouth (m), which serves also as anus, is
situated in the midline on the ventral surface, far removed from the
head; the gonopore or genital aperture is found in the space between the
mouth and the posterior end.
The body is covered with a ciliated epidermis containing rhabdites, small
rod-like structures which, when discharged, produce abundant mucus. The
-PI
Fig. 2. Digestive system of a
planarian. at, anterior intes-
tinal ramus; au, auricle; e, eye;
tn, mouth; ph., pharynx; pi, poste-
rior intestinal ramus.
digestive system (Fig. 2) is a gastrovascular cavity with three ma.?
divisions (one anterior and two posterior intestinal rami, ai and
from which many lateral branches originate. In the prolongation
anterior ramus is a muscular cylindrical pharynx (ph) lying in
arate chamber, the pharyngeal pouch. When feeding, the pharyp
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traded through the mouth opening. A few American species have, besides
the principal pharynx, multiple pharynges attached to the anterior por-
tions of the posterior intestinal rami, which project into the common
pharyngeal chamber (polypharyngeal species).
The space between the epidermis with its underlying basement membrane
and the gastrodermis or intestinal epithelium is filled with a loose
parenchymatous tissue or mesenchyme in which the remaining organs are
embedded. The central nervous system if formed by a pair of cerebral
ganglia or brain in the head, from which a pair of ventral nerve cords
extend to almost the posterior end, connected by numerous transverse
commissures. From these parts originate many nerves running to various
organs and to a nerve plexus located below the epidermis.
pp od am sph ac gp
Fig, 3. Reproductive system. A, dorsal view (female gonads
indicated on left, male gonads on right side); B, generalized
diagram of the copulatory apparatus in sagittal section.
ac, common genital atrium; ad, adenodactyl; am, male atrium;
b, copulatory bursa; bd, bursal duct; br, brain or cerebral
ganglion; gp, gonopore; m, mouth; od, oviduct; ode, common
oviduct; ov, ovary; p, penis; pa, parenchyma; pb, penial
bulb; ph, pharynx; pp, penis papilla; sph, sphincter; sv,
spermiductal vesicle; t, testis; v, vagina; vd, vas deferens;
vs, seminal vesicle.
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GENERAL FEATURES
Planarians or freshwater triclads are elongated, flattened Turbellaria^
the native American species measuring in length from about 2 mm (freshly
hatched) to perhaps 40 mm. In the extended, quietly gliding animal one
distinguishes an anterior portion or head, often characteristically
shaped, bearing important sense organs. The lateral margins of the head
may protrude as auricles (which have chemical and mechanical sensory
receptors). There are usually two dark eye spots on the head, each
located near the medial margin of a white area, often accompanied by
smaller accessory or supernumerary eyes. Some species have numerous
eyes and subterranean planarians are often eyeless or blind. A median
adhesive organ may be differentiated on the frontal margin of the head.
The body may or may not be pigmented, often exhibiting various patterns
of pigment arrangement. The mouth (m), which serves also as anus, is
situated in the midline on the ventral surface, far removed from the
head; the gonopore or genital aperture is found in the space between the
mouth and the posterior end.
The body is covered with a ciliated epidermis containing rhabdites, small
rod-like structures which, when discharged, produce abundant mucus. The
---PI
Fig. 2. Digestive system of a
planarian. ai, anterior intes-
tinal ramus; au, auricle; e, eye;
m, mouth; ph, pharynx; pi, poste-
rior intestinal ramus.
digestive system (Fig. 2) is a gastrovascular cavity with three main
divisions (one anterior and two posterior intestinal rami, ai and pi)
from which many lateral branches originate. In the prolongation of the
anterior ramus is a muscular cylindrical pharynx (ph) lying in a sep-
arate chamber, the pharyngeal pouch. When feeding, the pharynx is pro-
-------
truded through the mouth opening. A few American species have, besides
the principal pharynx, multiple pharynges attached to the anterior por-
tions of the posterior intestinal rami, which project into the common
pharyngeal chamber (polypharyngeal species).
The space between the epidermis with its underlying basement membrane
and the gastrodermis or intestinal epithelium is filled with a loose
parenchymatous tissue or mesenchyme in which the remaining organs are
embedded. The central nervous system if formed by a pair of cerebral
ganglia or brain in the head, from which a pair of ventral nerve cords
extend to almost the posterior end, connected by numerous transverse
commissures. From these parts originate many nerves running to various
organs and to a nerve plexus located below the epidermis.
b-l--
od--V--,
pa
pp od am sph ac gp
Fig. 3. Reproductive system. A, dorsal view (female gonads
indicated on left, male gonads on right side); B, generalized
diagram of the copulatory apparatus in sagittal section.
ac, common genital atrium; ad, adenodactyl; am, male atrium;
b, copulatory bursa; bd, bursal duct; br, brain or cerebral
ganglion; gp, gonopore; m, mouth; od, oviduct; ode, common
oviduct; ov, ovary; p, penis; pa, parenchyma; pb, penial
bulb; ph, pharynx; pp, penis papilla; sph, sphincter; sv,
spermiductal vesicle; t, testis; v, vagina; vd, vas deferens;
vs, seminal vesicle.
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The protonephridial excretory system consists of a network of fine tubes
with branches terminating in flame cells and with many pores on the
surface of the body.
Muscular fibers are found in several layers underlying the epidermis
(integumental muscles), individual fibers traversing the parenchyma in
various directions, and variously arranged fibers in the pharynx, the
adhesive organ when present, and in parts of the reproductive system.
No separate circulatory system is developed, its functions being taken
over by the gastrovascular cavity and the system of interstitial spaces
of the parenchyma.
The parenchyma contains the cell bodies of many unicellular glands, the
ducts of which open through the epidermis to the outside, through the
surface of the adhesive organ, the tip of the pharynx, and various parts
of the reproductive organs.
The taxonomically most important structures are those of the reproductive
system. All freshwater planarians are hermaphroditic, both male and
female genital organs developing in the same individual (Eig. 3). The
paired ovaries or germaries (ov) lie in the anterior portion of the
body adjacent to the ventral nerve cord. From each ovary originates a
canal, the oviduct or ovovitelline duct (od) which proceeds posteriorly
along the dorsal side of the nerve cord. At the boundary between the
ovary and the oviduct is a small enlargement of the canal, the seminal
receptacle. Many vitellaria or yolk glands (vi) branch out from the
oviduct, consisting of large eosinophilic cells with yolk inclusions.
Cell masses, representing a transition between oogonia and yolk cells,
may be attached to the ovaries, forming the so-called parovaria.
The male gonads are numerous testicular follicles (t) located on either
side in a zone extending from behind the ovaries to the level of the
pharynx, the mouth, or almost to the posterior end (exceptionally, the
testes may fuse into one elongated organ on either side). Each testis
is connected to the sperm duct or vas deferens (vd) by a thin canal, the
vas efferens. The sperm ducts run posteriorly parallel to the nerve
cords. In the region of the pharynx they usually widen, forming twisted
enlargements filled with sperm, the false seminal vesicles or spermi-
ductal vesicles (sv).
Behind the pharynx is the copulatory apparatus or complex consisting of
the terminal portions of the gonoducts and various accessory structures.
The genital aperture (gp) leads into a cavity, the genital atrium or
antrum, which may be subdivided into several parts separated by stric-
tures: a male atrium (am) enclosing the penial papilla, a female atrium
receiving the openings of parts of the female complex, and a common
atrium (ac) which connects with the gonopore. The nomenclature of the
atria is frequently inconsistent, as the mouth of the oviduct may be in
the posterior part of the "male" atrium.
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The male copulatory organ or penis is a muscular organ consisting of two
parts, the anterior penis bulb (pb) embedded in the parenchyma and the
penis papilla (pp) protruding into the male atrium. Generally the two
sperm ducts (vd) enter the penis bulb, separately or combined as a common
vas deferens, and open into a cavity, the seminal vesicle (vs) or bulbar
cavity. From this cavity a narrower duct proceeds into the penis papilla,
the ejaculatory duct (de). There are, however, several modifications of
this plan found in the different species.
The oviducts or ovovitelline ducts usually approach the midline above the
genital atrium and unite to form a common oviduct (ode) which generally
opens into the atrium from the dorsal side. A usually sac-shaped
accessory organ, the copulatory bursa (b), lies in the space between the
pharyngeal pouch and the penis bulb and is connected to the atrium by a
muscular tube, the bursal duct or canal (bd), running dorsal to the
atrium either in the midline or to one side of it. In a few species the
bursa is absent but the bursal canal is developed, either ending blindly
or connecting with a branch of the intestine. In the genera Cura and
Dugesia, the paired or united oviducts open into the distal part of the
bursal canal instead of the atrium.
Species of the genus Planaria have a hollow very muscular organ attached
to the posterior portion of the copulatory complex, the adenodactyl (ad)
or musculo-glandular organ.
Gland ducts originating in the surrounding parenchyma open into various
parts of the copulatory complex: the penial cavity, the terminal parts
of the oviducts ("shell glands", sg), the common atrium (cement glands),
the adenodactyl, etc.
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REPRODUCTION
Reproduction of planarians is either sexual or asexual. Sexual repro-
duction usually involves copulation of two individuals during which the
sperm of one partner are deposited in the copulatory bursa of the other
(autofecundation has been observed only in Cura foremani-i among the
American planarians). From the bursa the sperm are expelled through the
bursal duct, enter the oviducts, and are finally stored in the seminal
receptacles. The ova are fertilized as they pass through the receptacles
on their way to the atrium where they accumulate. They are deposited in
a cocoon, a rather large (diameter 0.5 to over 3 mm) spherical or
ellipsoidal capsule with a shell of horny consistency, attached to the
substrate by a gelatinous secretion or, more rarely, by a thin stalk
(Fig. 4).
A B
Fig. 4. Egg capsules of planarians. A, unstalked cocoon;
B, stalked cocoon.
Each capsule contains several (2-20, sometimes more) zygotes and
thousands of yolk cells which serve as food for the developing embryos.
When the young hatch from the cocoon one to several weeks after its
deposition, they have more or less the shape of the adults, without
passing through a larval stage and metamorphosis. They lack, however,
any trace of the reproductive system which will develop only when they
are almost fully grown.
Asexual reproduction is by fission or by fragmentation. Fission occurs
usually at a postpharyngeal level and consists of a tearing apart of the
posterior portion of the body. Both parts then regenerate the missing
organs, the anterior part a new tail and the posterior part a new head,
pharynx, etc. In fragmentation the body breaks up into several smaller
portions each of which then encysts, i.e., acquires a rounded shape and
encloses itself in a layer of transparent slime which usually adheres to
the substrate. After a certain rest period small planarians hatch from
the cysts, resembling the young hatched from an egg capsule. Asexual
reproduction is very common in some species, and certain populations have
been cultured through several generations without developing sexual
structures.
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CHARACTERS USED IN THE IDENTIFICATION OF PLANARIANS
It is not always possible to identify individual species and even genera
by their external characteristics, and the final decision must often be
made by examining anatomical features. In a well-explored geographic
region one may be reasonably sure to recognize certain species which bear
distinctive characters and are known to occur locally, but it is always
advisable to have the identification confirmed by anatomical study.
Of the external features, the shape of the anterior end of the quietly
gliding animal is often characteristic. It may be pointed (triangular),
truncate, or rounded. The lateral edges may protrude as auricular ap-
pendages. There may or may not be a constriction or neck behind the head.
An adhesive organ may be developed on the frontal margin. The number and
arrangement of the eyes sometimes furnish useful specific characters. It
is best to prepare an outline sketch of the anterior end of the specimens
in gliding motion.
The pigmentation of the body, including both shade and pattern on the
dorsal and ventral surfaces, should be noted. The pigments are either
granular, enclosed in subepidermal pigment cells, or nongranular and
diffused in various tissues. Granular pigments can usually be recognized
in microscopical sections, while diffused pigments easily bleach in the
reagents used for fixing and storing. Thus, some pigmented planarians
have been described from preserved specimens as "white" (e.g., the dark
Dencbcoooelopsis vaginata and the pink Maovoeotyla glandulosd). In life
unpigmented planarians may appear colored from the contents of the
intestine visible through the body wall. In this case, however, the
areas not penetrated by the intestine and its branches are white: head,
body margins, and spots above the pharynx and copulatory complex.
Some anatomical features may be examined on living specimens in squash
preparations or on preserved specimens after clearing the tissues or in
whole mounts. Detailed studies of the anatomy, however, demand the
preparation of serial sections. The characters most frequently used are
the structure of the adhesive organ; the configuration of the lateral
edge of the body (development of a distinct marginal zone with special
differentiations of glands and rhabdites); and an analysis of the repro-
ductive system. The posterior limits of the bands of testes should be
determined as well as their location close to the dorsal or to the ven-
tral side. In the structure of the copulatory organs, the size and shape
of the penis and the course of the sperm ducts are important. The ovi-
ducts may open into the atrium or into the bursal duct. Special sphinc-
ters may be developed on the terminal part of the bursal canal or in
other places. Further details concerning the copulatory complex are
discussed in the descriptions of the individual species.
11
-------
SECTION II
KEY TO THE GENERA OF NORTH AMERICAN FRESHWATER TRICLADS
epe
~~
~ me '"
epi
Fig. 5. Muscle layers of pharynx in cross section. A, Planariidae;
B, Dendrocoelidae. epe, external epithelium; epi, internal epithe-
lium; me, external muscle layer; mi, internal muscle layer.
1 Internal muscle zone of pharynx consists of two distinct
layers, a circular and a longitudinal one (Fig. 5A):
Family Planariidae 2
Internal muscle zone of pharynx consists of one layer of
intermingled circular and longitudinal fibers (Fig. 5B):
Family Dendrocoelidae 9
2 (1) Oviducts, separate or united, open into end part of bursa
stalk 3
Oviducts unite, the common oviduct opening into the genital
atrium 4
3 (2) Zone of testes extends to level of pharynx: Cura
Zone of testes extends to posterior end: Dugesia
4 (2) Eyes numerous, forming a band around anterior end: .. Polyoetis
Eyes two or wanting, if numerous not on head margin 5
5 (4) Adenodactyl present Planaria
No adenodactyl 6
6 (5) Anterior end with adhesive organ: 7
Anterior end without adhesive organ: 8
7 (6) Body elongated, flat, with well-developed postpharyngeal
region: Sphalloplana
Body turtle-shaped, with reduced postpharyngeal region: Kenkia
8 (6) Penis normal, with bulb and papilla: Phagooata
Penis without bulb, papilla rudimentary: Hymanel'la
9 (1) Anterior end with deeply invaginated adhesive organ:Macvocoty1a
Anterior end with adhesive disc or without adhesive organ .. 10
13
-------
10 (9) Penis with bulb and papilla 11
Penis with large bulb, no papilla; .., Rectooephala
11 (10) Penis bulb rounded, containing seminal vesicle:
Dendpocoe lops-Is
Penis bulb elongated, containing a prostatic vesicle (Fig.89):
Pvoootyla
14
-------
SECTION III
KEY TO THE SPECIES OF NORTH AMERICAN FRESHWATER TRICLADS
1 Anterior end triangular 2
Anterior end truncate 7
Anterior end rounded 35
2 (1) Head rather acutely pointed, with prominent auricles, pharynx
pigmented 3
Head bluntly pointed, auricles absent or rounded, pharynx
unpigmented 5
3 (2) Auricles long and pointed, ventral side pigmented 4
Auricles short, ventral side usually unpigmented:
Dugesia tigrina (p. 20)
4 (3) No diverticula on ejaculatory duct; Dugesia dorotooep'hala (p. 22)
Ejaculatory duct with a pair of diverticula:
Dugesia antillana (p. 23)
5 (2) Auricles rounded, penis with rounded bulb and a single seminal
Vesicle, bursal sac absent Cuva foremanii (p. 19)
Auricles absent, bursal sac developed 6
6 (5) Penis with divided bulb and two seminal vesicles:
Dugesia polyahroa (p. 24)
Penis without bulb, with rudimentary papilla:
Hymanella vetenuova (p. 31)
7 (1) Frontal margin without adhesive organ 8
Frontal margin with adhesive organ 28
8 (7) Body pigmented 9
Body without pigment, white 21
9 (8) Eyes normally two (with occasional accessory eye spots),
close together 10
Eyes numerous, on margin of head (genus Polyaelis') 19
10 (9) Pharynx single (monopharyngeal) 11
Pharynges multiple (polypharyngeal) 18
11 (10) Adenodactyl present 12
No adenodactyl present 13
12 (11) External muscle layer of penis papilla thickened, adenodactyl
opens into a separate chamber of the genital atrium:
Planaria dactyligera museulosa (p. 29)
External muscle layer of penis papilla not thickened, adeno-
dactyl opens into atrium near gonopore:
Planopia daetyligera daetyligera (p. 28)
15
-------
13 (11) Penis papilla with terminal lobe, ejaculatory duct opening
on ventral side of papilla .... Phagooata orenophila (p. 39)
Penis papilla without terminal lobe, ejaculatory duct
opening at tip of papilla 14
14 (13) Penis rudimentary, without bulb and with small papilla:
Hymanella retenuova(p. 31)
Penis well-developed, with bulb and papilla 15
15 (14) Ejaculatory duct with blind diverticulum:
Phagooata velata (p. 35)
Ejaculatory duct without diverticulum 16
16 (15) Copulatory bursa rounded, sac-shaped 17
Copulatory bursa U-shaped, testes in part longitudinally
fused Phagooata veimalis (p. 34)
17 (16) Penis with round bulb and narrow ejaculatory duct:
Phagooata bulbosa (p. 36)
Penis with small bulb and wide undivided penial lumen:
Phagooata monopharyngea (p. 33)
18 (10) Penis papilla long, conical, with fibrous layer below outer
epithelium, penis lumen with two seminal vesicles:
Phagooata gpaoilis (p. 43)
Penis papilla short, truncate, with intermingled circular
and longitudinal muscle fibers below outer epithelium,
penis lumen not subdivided .... Phagooata woodworthi. (p. 44)
19 (9) Bursa duct with anterior glandular and posterior non-glan-
dular sections 20
Bursa duct non-glandular throughout:
Polyoelis ooronata bovealis (p. 27)
20 (19) Penis papilla as large as, or larger than, penis bulb:
Polyoel-Ls ooronata covonata (p. 25)
Penis papilla smaller than penis bulb:
Polyoelis ooTonata brevipenis (p. 26)
21 (8) Head with slender, pointed auricles:
Phagooata bursaperforata (p. 37)
Head without pointed auricles 22
22 (21) Polypharyngeal Phagooata graoilis (p. 43)
Monopharyngeal 23
23 (22) Eyes normally two (smaller accessory eye spots may occur) .. 24
Eyes wanting Pvoeotyla typhlops (p. 64)
Eyes many, in two longitudinal rows:
Phagooata morgani polyoelis (p. 41)
16
-------
24 (23) Eyes separated by about 1/3 the width of the head, about
equidistant from frontal and lateral margins:
Dendrocoelopsis alaskensis (p. 60)
Eyes close together, removed farther from frontal than from
lateral margins 25
25 (24) Anterior intestinal ramus extending to a level in front of the
eyes Planaria oeeulta (p. 30)
Anterior intestinal ramus starts behind level of eyes 26
26 (25) Opening of ejaculatory duct at tip of penis papilla:
Phagooata ovegonensis (p. 32)
Opening of ejaculatory duct on ventral side of penis
papilla 27
27 (26) Tip of penis papilla with a muscular, wart-like structure,
testes extend to level of mouth:
Phagoaata movgani movgani (p. 40)
Tip of penis papilla without muscular wart, testes extend to
posterior end Phagooata nivea (p. 42)
28 (7) Inner pharyngeal muscle zone with separate layers of cir-
cular and longitudinal fibers 29
Inner pharyngeal muscle zone with one layer of intermingled
circular and longitudinal fibers 30
29 (28) Body turtle-shaped, with cylindrical snout, postpharyngeal
region reduced Kenkia rhynehida (p. 56)
Body elongated, flat, in life without snoutlike extension,
postpharyngeal region well-developed: genus Sphalloplana
(The systematics of this genus is undergoing great changes
as more material is made available to specialists by the
efforts of speleological investigators. For in part un-
reliable data on individual species see the descriptive
part of the report, pages 45-55.)
30 (28) Adhesive organ forms a deep subterminal invagination on the
head, copulatory complex excessively glandular:
Maeroaotyla glandulosa (p. 57)
Adhesive organ in form of a disc or shallow depression 31
31 (30) Body pigmented 32
Body unpigmented, white 34
32 (31) Penis with voluminous bulb, no papilla:
Reotooephdla exotica (p. 65)
Penis with normal bulb and papilla 33
33 (32) Testes ventral, to level of gonopore:
Dendrocoelopsis vaginata (p. 58)
Testes dorsal, to posterior end:
Dendroeoelopsis piriformis (p. 59)
17
-------
34 (31) Eyes wanting Dendroeoelopsis hymanae (p. 61)
Eyes two, or two groups of scattered eyes:
Proootyla fluwLatilis (p- 63)
Eyes multiple, arranged in two longitudinal rows:
Dendroooelopsis amerioana (p. 62)
35 (1) Penis reduced, without bulb and with rudimentary papilla:
Eymanella vetenuova (p. 31)
Penis normal, with bulb and papilla 36
36 (35) Ventral surface unpigmented Phagooata tahoena (p. 38)
Ventral surface pigmented Dugesia polydhroa (p. 24)
18
-------
SECTION IV
SPECIES CHARACTERISTICS AND RANGES
Cura foremanii (Girard, 1852)
Synonyms: Dugesia foremanii Girard, 1852; Planar-La foremanii: Stimpson, 1857;
P. simplissima Curtis, 1900; P. simplioissima: Morgan, 1904; P. lugubris:
Morgan, 1901 (not 0. Schmidt, 1861); Curtisia simplioissima: Graff, 1916; C.
foremani: Kenk, 1930; IPlanaria gonooephala: Pearl, 1903 (not Duges, 1830);
tDugesia modesta Girard, 1893.
sg ode
vs vd
Fig. 7 after Kenk (1935).
gl
gp
Length 7-15 mm, generally rather broad and thick. Head bluntly tri-
angular, with rounded, only slightly protruding auricles. Color uni-
formly gray or brown to almost black, ventrally lighter. Besides the
(normally two) white eye fields there is a light oblique dash on the
dorsal side of each auricle and the mouth and gonopore appear as white
spots. Pharynx unpigmented (white), which distinguishes the species
from the common American species of Dugesia (this may be checked on the
freshly extirpated pharynx). Testes very few, dorsal, between ovaries
and level of mouth. Penis relatively small, with moderately developed
bulb containing the rounded seminal vesicle and a finger-shaped papilla
traversed by a straight ejaculatory duct opening at its tip. No bursal
sac is developed, the bursal canal attaches to a branch of the intestine.
The two oviducts unite behind the copulatory apparatus and open into the
bursal duct which also receives the shell glands. Cocoon spherical,
attached to substrate by a thin, flexible stalk. Inhabitant of cool
streams. Reproduces only sexually. Eastern half of North America, from
New Brunswick to Louisiana and westward to Minnesota and Arkansas.
Principal literature: Curtis (1900), Stevens (1904), Kenk (1935 and
1944).
19
-------
Dugesia tigrina (Girard, 1850)
Synonyms: Planaria maoulata Leidy, 1847; P. tigrina Girard, 1850; P. lata
Sivickis, 1923; Dugesia maoulata: Girard, 1851; D. lata: Hyman, 1951;
Euplanaria maoulata: Kenk, 1930; E. lata: Kenk, 1930; E. tigrina: Kenk, 1935;
E. novangliae Hyman, 1931; IE. microbursalis Hyman, 1931; tDugesia
miovobursal-is: Hyman, 1939; ?P. gonoaephaloides Girard, 1850; "?Planaria
gonocephaloides: Stimpson, 1857; ?P. gonoaephala: Woodworth, 1897 (not Duges,
1830).
8
Fig. 9 after Kenk (1935).
10
Fig. 10. Pigment patterns (after Hyman, 1939b).
20
-------
A very common species, widely distributed and polytypic. Mature animals
6-18 mm long and 1-3 mm wide. Head triangular, with bluntly pointed or
somewhat rounded anterior tip. Auricles short and broad. Eyes normally
two, situated close together, anterior to the level of the auricles.
Coloration very variable, sometimes appearing almost uniformly brown to
the naked eye or composed of spots of various shades of brown with ir-
regularly dispersed white splotches (spotted type); or showing a pair
of dark longitudinal stripes separated by a light middorsal streak
(striped type), both types sometimes occurring in the same population.
Ventral surface usually unpigmented. Pharynx pigmented, with white
tip. Anatomically very similar to D. dorotoaephala. Testes ventral,
reaching close to posterior end. Penis with round bulb and short con-
ical papilla directed posteroventrally. The vasa deferentia ascend to
the dorsal side before entering the bulb, then curve posteroventrally,
each widening into an elongated seminal vesicle. They unite at the base
of the papilla and proceed to its tip as a narrower ejaculatory duct.
Occasionally the lumen of the penis is expanded as a single seminal
vesicle, apparently a transitory condition. Bursal duct forms a rather
sharp angle at the entrance of the separate or united oviducts.
Shell glands open into the bursal duct below the mouths of the oviducts.
The small common genital atrium frequently shows a posterior divertic-
ulum. Reproduction sexual and asexual. Cocoon round, stalked. Widely
distributed, generally in warm ponds, lakes, and rivers in the United
States and southern Canada, from the Atlantic to the Pacific coasts.
Principal literature: Curtis (1902), Sivickis (1923), Hyman (1939b),
Kenk (1935, 1944).
21
-------
Dugesia dorotoaephala (Woodworth, 1897)
Synonyms: Planapia dovotocephdla Woodworth, 1897; P. agiHs Stringer, 1909;
Euplanari-a dorotocephala: Kenk, 1930; E. agili-s: Kenk, 1930; E. philadelphiea
Hyman, 1931; Dugesia. agilis: Hyman, 1939; D. diabolis Hyman, 1956.
11
12
O.lmi
gp
A polytypic species, in nature up to 30 nun long and about 3.5 mm wide
(may grow larger in laboratory cultures). Head triangular, with slightly
convex sides and rather pointed tip. A pair of elongated, sharply
pointed auricles extends laterally, during gliding usually held elevated.
Eyes normally two, close together, just anterior to the base of the
auricles. Color brown to almost black dorsally, somewhat lighter ven-
trally, occasionally with a darker middorsal streak behind the pharynx.
Under magnification small white and dark spots are seen on the general
background color. Pharynx pigmented light gray, with a short white tip.
Testes numerous, ventral, extending to posterior end. Penis with spher-
ical bulb and short conical or rounded papilla pointing posteroventrally.
Vasa deferentia enter bulb dorsolaterally, bend down toward the axis of
the papilla, and increase in diameter, each duct forming a seminal vesi-
cle. Distally they unite to a short ejaculatory duct opening at the tip
of the papilla. Under certain conditions the two seminal vesicles may
appear fused, forming a larger, round vesicle. Bursal duct running pos-
teriorly above the penis, then curving ventrally toward the small common
atrium. At the turning point it receives the two oviducts from the
posterolateral sides. Below these openings (which are close together)
is a short zone where numerous outlets of shell glands open into the
bursal canal (none into the oviducts). Sexual or asexual reproduction
may predominate in different populations. Cocoon spherical, stalked.
Distributed in running and standing waters across the continent in the
United States and southern Canada. Principal literature: Woodworth
(1897), Hyman (1925, 1929, 1931b), Kenk (1944).
22
-------
Dugesia antillana Kenk, 1941
13
14
Fig. 14 after Kenk (1941).
Very slender, up to 19 mm long and 0.7 mm wide. Head triangular, rather
broad, with long, pointed auricles. Dorsal side grayish-brown, sometimes
with an indistinct darker streak in the prepharyngeal midline, ventrally
somewhat lighter. Eyes usually two, lying close together. The two eye
fields, the auricles and a spot at the site of the gonopore lack pigment.
Pharynx pigmented. Testes numerous, predominantly dorsal, starting
behind the ovaries and extending to near the posterior end. At full ma-
turity, some testes may occupy the entire dorsoventral diameter. Vasa
deferentia expand at the level of the mouth into spermiductal vesicles,
then narrow again as they approach the penis. The gonopore leads into
a common atrium which dorsally receives the bursal duct and anteriorly
connects with the male atrium. Penis with rather small bulb and short,
conical papilla. The vasa deferentia enter the bulb dorsolaterally and
widen within the bulb into a pair of spindle-shaped cavities which unite
in the papilla to a short canal, the ejaculatory duct, which opens at
the tip of the papilla. Close to the union of the two sperm ducts, a
pair of tubular diverticula extend toward the penial bulb (very charac-
teristic of the species). Copulatory bursa voluminous, bursal stalk
running dorsal to the penis and bending ventrally behind it to approach
the common atrium. Oviducts open separately into the bursa stalk. Below
their opening, the stalk receives numerous eosinophilic shell glands.
Inhabits mountain streams. Only sexual reproduction has been observed.
Known only from Puerto Rico. Literature: Kenk (1941).
23
-------
Dugesia polydhvoa (0. Schmidt, 1861)
Synonyms: Planaria polychroa 0 Schmidt, 1861; P. lugubris: Bohmig, 1909-
Dugesia lugubris: Ball, 1969 (in part).
15
m
Fig. 16 after Bohmig (1909).
Mature specimens up to 20 mm long and about 3 mm wide. Head more or less
rounded, occasionally showing a small median projection. A gentle nar-
rowing or neck behind the lateral edges. Eyes usually two, close to the
frontal end. Color variable, usually a shade of brown, either uniformly
distributed or somewhat spotty. A pair of lighter elongated auricular
sense organs is seen near the lateral margin of the head some distance
behind the eye level. Pharynx unpigmented. Testes numerous, dorsal,
extending to posterior end. The penis consists of a very muscular bulb
and a generally conical papilla. The bulb is divided into two sections,
an anterodorsal part which contains the rounded anterior seminal vesicle
and a posterior part enclosing the elongated posterior seminal vesicle.
The two vesicles are joined by a narrower canal. From the posterior ves-
icle the ejaculatory duct proceeds through the papilla to its tip. The
vasa deferentia open separately into the anterior seminal vesicle, the
two oviducts into the posterior part of the bursa stalk. Reproduction
only sexual. Cocoon spherical, stalked.' This species, widely distrib-
uted in Europe, has been introduced to the St. Lawrence River system in
eastern Canada (Ontario) and New York State. Principal literature:
Bohmig (1909), Komarek (1925), Ball (1969), Reynoldson and Bellamy
(1970).
24
-------
PolyoeUs eoronata oovonata (Girard, 1891)
Synonym: Phagoeata coronata Girard,
17 T8& 18 «••«•«
1891.
bg
ode bd
ph m b
bp Ip
9P
Mature animals up to 19 mm long and 2.4 mm wide. Head truncate, with
convex frontal margin and a pair of triangular auricles with rounded
tips extending anterolaterally. Eyes multiple, irregularly scattered on
the dorsal side of the head in a band which parallels the margin of the
head, narrows posteriorly, and extends over the anterior quarter of the
prepharyngeal region. Dorsal side brown to almost black, ventral side
lighter. The individual eyes have no pigmentless fields. Testes ventral,
prepharyngeal. Penis with very muscular bulb and more or less conical
papilla, the size relations of the two parts being subject to great vari-
ation according to the state of contraction of the organ (generally the
papilla is as long as the bulb). The male atrium duplicates in shape the
penis papilla and receives in its posterior part the -mouth of the common
oviduct. Bursa usually large, with a wide bursal duct proceeding back-
ward to one side (usually left) of the midline, lined with the same
glandular epithelium as the bursal sac and covered by a very feeble mus-
cle layer. Only in its posterior fourth the histological aspect of the
epithelial lining changes suddenly, the canal narrows, and its muscular
coat thickens. Reproduction sexual and asexual. Inhabits springs and
cold creeks in southern Wyoming, Utah, Montana, New Mexico. Principal
literature: Girard (1893), Braithwaite (1962).
25
-------
Polyoelis oovonata brevipenis new subspecies
Synonym: Polyaelis eoronata: Hyman, 1931,
19
i I :
am ac bd
Fig. 19 after Kenk (1952).
Externally indistinguishable from the subspecies eovonatat differing
from it chiefly in the anatomy of the copulatory apparatus. 13-20 mm
long when mature. The penis bulb is very large and elongated, the penis
papilla short, plug-like. The penial cavity or seminal vesicle is volum-
inous, stretched out in the longitudinal axis, and receives the two sperm
ducts laterally, often asymmetrically. The ejaculatory duct is rather
wide and short. The bursal canal, which lies on the left side of the
penis, has a wide anterior portion which is lined with a glandular epi-
thelium similar to that lining the bursal sac. Where the duct turns
downward at the level of the male atrium, its histological nature changes
abruptly. The epithelium becomes nonglandular and the duct acquires a
strong muscle coat (this division of the bursal duct into an anterior
glandular and a posterior muscular part is seen also in the subspecies
ooronata but not in borealis). Reproduction sexual and by fission. Re-
ported from South Dakota and Colorado, but is probably more widely
distributed. Literature: Hyman (1931a), Kenk (1952).
26
-------
Polyaelis ooTonata borealis Kenk, 1953
Synonym: Polyoelis borealis Kenk, 1953.
20
m b vd vs
Fig. 20 after Kenk (1953).
Not distinguishable from the nominate subspecies by external features.
Up to 20 mm long and 2.5 mm wide. Differs from the subspecies aoronata
and brevipenis by the anatomy of the bursal duct which is evenly muscular
throughout its length and is not divided into glandular and muscular
sections. The penis has a well-developed, conical papilla (in brevipenis
the papilla is reduced in size, blunt, containing a short ejaculatory
duct). Reproduction sexual and asexual. In springs, cold streams, also
in mountain lakes, Alaska, northern Wyoming, Idaho. Principal literature:
Kenk (1953).
27
-------
Planaria dactyHgera dactyligera Kenk, 1935
Synonym: Planapia daotyl-igera Kenk, 1935.
21 Hi 22
/ i i i i i i ; i i i
vd gl bd vs od am de ode gp ad gl
Fig. 22 after Kenk (1935).
Length of mature animals up to 13 mm, width 1.75 mm. Head truncate, with
almost straight frontal margin and rounded lateral edges. Behind the
head is an insignificant narrowing or neck. Eyes usually two, their
distance from each other about 1/3 the width of the head and their dis-
tance from the frontal margin much greater than from the lateral margins.
Dorsal side darkly pigmented, gray, brown, or black; ventral surface
lighter. Externally indistinguishable from Eymanella and some species
of Phagoaata. Testes predominantly ventral, their zone on either side
reaching to the level of the mouth. The gonopore leads into the genital
atrium which has only one compartment, the male atrium, as the openings
of the common oviduct, the bursal canal, and the adenodactyl are in the
immediate vicinity of the gonopore. The sperm ducts enter the penis bulb
anterolaterally and open into the seminal vesicle with a common opening.
The ejaculatory duct emerging from the vesicle tapers posteriorly and
open at the tip of the penis papilla. The thickness of the muscle coat
underlying the outer epithelium of the papilla is about equal to that
adjoining the epithelium of the male atrium. Behind the gonopore is the
hollow adenodactyl. Cocoon ellipsoidal or spherical, unstalked. Only
sexual reproduction has been observed. Inhabitant of springs and creeks
in Virginia. Literature: Kenk (1935).
28
-------
Planar-la daoty'li,geva musculosa Kenk, 1969
23
Fig. 23 after Kenk (1969).
Differs from the subspecies daatyligera mainly in the anatomy of the penis
and of the genital atrium. The outer muscle zone of the penis papilla
(mp) consists of a very thick layer of circular fibers followed by a
normal layer of longitudinal muscles. The adenodactyl, instead of having
its opening at the gonopore, connects with a separate compartment (aa) of
the atrium lying posterior to the genital aperture. In streams and springs,
North Carolina, southern Virginia, and Louisiana. Literature: Kenk (1969).
29
-------
Planaria oaculta Kenk, 1969
24
25
0.5mm
m vd vs
am gp
Fig. 25 after Kenk (1969).
A small, white planarian up to 9 mm long and 1.5 mm wide. Head truncated,
with slightly convex frontal margin (which, in the gliding animal, may
present an indication of a bulge in the center). Eyes two, their distance
from each other amounting to 1/4 the width of the head, each eye farther
removed from the frontal than from the lateral margin. Externally very
similar to the eastern Phagocata movgani and the western P. oregonensis.
However, in P. oaoulta the tip of the anterior intestinal ramus extends
between the two eyes whereas in the adult P. morgani and P. oregonensis
its anterior tip is behind the eye level. Testes predominantly ventral,
stopping posteriorly at the level of the mouth. Penis with medium large
bulb and elongated conical papilla. Vasa deferentia opening independ-
ently into a glandular seminal vesicle. The highly convoluted ejaculatory
duct ends at the tip of the penial papilla and appears to be capable of
eversion. The common oviduct and, behind it, the stalk of the copulatory
bursa connect with the genital atrium close to the genital aperture.
Posterior to the aperture, the atrium extends into a small compartment
which receives the opening of the adenodactyl. This is a hollow organ
with a thick and dense muscle shell. The most outstanding characteristics
of the species are the structure of the penis and the presence of the
adenodactyl. So far found only in a well in the western part of Virginia.
Literature: Kenk (1969).
30
-------
26
fi
, u
Hymanella retenuova Castle, 1941
27
Fig. 27 after Hyman (1955).
Mature animals 7-14 ram long and 1.3-2.5 mm wide. Head of gliding animal
truncate with bulging frontal margin, at times appearing low triangular.
Lateral corners of head rounded, no distinct necklike constriction behind
them. Eyes two, farther removed from front end than from lateral margins.
Pigmentation grayish or brown. In life not distinguishable from pigmented
species of Phagoaata or Planaria. Testes not well analyzed, possibly
dorsolateral and prepharyngeal (or ventral and longitudinally fused?).
Vasa deferentia behind the spermiductal vesicles ascend to the penis and
unite. Penis very small, without bulb, consisting of a small papilla
traversed by the ejaculatory duct formed by the union of the sperm ducts.
Genital atrium very large, represented only by the male atrium which at
its posterior end receives the opening of the common oviduct from the
dorsal side. Copulatory bursa of varying size, its duct rather narrow,
running above the atrium and posteriorly arching down to the gonopore.
No separate vagina is differentiated. Cocoon ellipsoidal, unstalked, is
carried in the atrium up to four weeks before being deposited. In vernal
pools and seepage springs, apparently widely distributed in the east of
North America, from Massachusetts to North Carolina and Louisiana and
west to Ontario. Principal literature: Castle (1941), Hyman (1955).
31
-------
Phagooata oregonensis Hyman, 1963
Synonym: Fontioola oregonensis: Ball, 1969
28 m 29
0.5mm
ph m b
pi vd am ode gp
Fig. 29 after Kenk (1970a).
A small, white species up to 8 mm long and 1.5 mm wide, Head truncate,
frontal margin with slightly protruding central and lateral portions.
Lateral edges rounded, extending laterally, causing a constriction to
appear behind the head. Eyes usually two, but smaller supernumerary eye
spots are not rare. The distance between the two principal eyes is 1/5
to 1/4 the width of the head at eye level, their distance from the frontal
margin greater than from the lateral margins. The tip of the anterior
intestinal ramus is behind the level of the oyes. Externally similar to
the Alaskan P. nivea and the eastern P. morgani. Testes ventral, extend-
ing posteriorly behind the copulatory complex (in P. movgani only to the
level of the mouth). Penis consists of a medium size bulb and a short,
conical, pointed papilla. The sperm ducts, after expanding at the sides
of the pharynx as spermiductal vesicles, continue posteriorly to the level
of the male atrium, then turn forward again to enter the penis bulb later-
ally. They open into the penial lumen independently, without first unit-
ing. The lumen is an elongated cavity, not differentiated into seminal
vesicle and ejaculatory duct, and opens at the tip of the penis papilla.
The openings of the common oviduct and of the canal of the copulatory
bursa are close to the gonopore. No common genital atrium is developed.
Known only from a spring in Portland, Oregon. Literature: Hyman (1963),
Kenk (1970a).
32
-------
Phagooata monopharyngea Hyman, 1945
Synonyms: Phagooata gracilis monopharyngea Hyman, 1945; Fontioola graoilis
monopharyngea: Ball, 1969.
vd bp
pl PP
Fig. 30 after Hyman (1945).
A problematic species. Preserved specimens up to 15 mm long, rather
broad. Head truncate, with slightly indicated auricles, without adhesive
organ. Pigmentation dark gray. Eyes normally two, frequently with small
accessory eyes. No testes seen. Penis with small bulb and large, con-
ical papilla. Vasa deferentia enter and traverse the bulb and open sep-
arately into the wide elongated penial lumen which is situated in the
papilla and extends to its tip. Common oviduct opens into posterodorsal
wall of atrium. Copulatory bursa sac-shaped, bursal duct narrow in its
anterior part, widens gradually as it proceeds posteriorly to join the
common atrium. Known from the outlet of a tile drain, Iowa. Literature:
Hyman (1945).
33
-------
Phagoeatajoernalis Kenk, 1944
Synonym: Fpntiaola vernalis: Ball, 1969
32
SH,
Fig. 32 after Kenk (1944).
Length up to 12 mm, width 1.5 mm. Head truncate, somewhat variable in
shape during locomotion. Lateral edges of frontal margin rounded, behind
them a very slight narrowing of the lateral margins (neck). Dorsal side
gray, often with brownish hue, sometimes with darker longitudinal streaks;
ventral surface lighter. Pigment is lacking in the ocular spaces and at
the oral and genital apertures. Pigmentation may be almost absent in very
young specimens and in animals ready for asexual reproduction. Eyes two,
farther removed from the frontal than from the lateral margins. Cannot
be distinguished from several other species by external characters. Rare-
ly found sexually mature. Testes few, ventral, prepharyngeal, in part
longitudinally fused. Penis with rather small bulb and elongated, conical
papilla. Vasa deferentia enter bulb and unite to form a rather narrow
longitudinal canal, the ejaculatory duct, opening at the tip of the
papilla. No seminal vesicle is developed. Copulatory bursa U-shaped,
narrow in the midline, the two horns extending anteroventrally and ending
below the intestine. Oviducts unite in the space between the male atrium
and the bursal duct and open into the atrium anterior to the mouth of the
bursal duct. Reproduction is principally asexual, by fragmentation fol-
lowed by encystment. Found usually in temporary ponds in winter and early
spring. Distributed chiefly in the Mid-West (Michigan, Ontario, probably
Illinois and Indiana) and Tennessee. Principal literature: Kenk (1944).
34
-------
Phagooata velata (Stringer, 1909)
Synonyms: Planaria velata Stringer, 1909; Fontioola velata: Kenk, 1930,
33 iPi 34
I!
U
vd gl am
Fig. 34 after Kenk (1944).
Length up to 20 mm, width 2 mm. Head truncated, with slightly convex
frontal margin and rounded lateral edges. Behind the head a faint
constriction (neck) may occur in gliding locomotion. Dorsal surface
usually dark gray to almost black, occasionally with a darker streak
along the postpharyngeal midline. Ventral side lighter. Young speci-
mens and individuals before fragmentation may be very lightly pigmented.
Eyes usually two, farther distant from the frontal than from the lateral
margins. Externally not distinguishable from several other American
planariids. Testes numerous, predominantly dorsal, distributed almost
to the posterior end. Genital atrium divided into common and male
atria. Penis with small bulb and conical papilla. Vasa deferentia enter
bulb ventrolaterally and open separately into a bulbar cavity, the sem-
inal vesicle, from which the ejaculatory duct proceeds to the tip of the
papilla. Close to its opening, a ventral diverticulum branches off,
extending toward the base of the penial papilla. Copulatory bursa sac-
shaped, its canal opening into the common atrium. Reproduction sexual
or asexual, the latter by fragmentation and encystment. In springs,
streams, and spring-fed ponds, apparently all across the continent.
Principal literature: Stringer (1909), Castle (1928), Castle and Hyroan
(1934), Kenk (1944).
35
-------
Phagocata bulbosa Kenk, 1970a
35
36
0.5mm
vd
vs
am de
ode gp
Fig. 36 after Kenk (1970a).
Mature animals up to 12 nun long and 1.7 mm wide. Head truncate, with
straight or centrally convex frontal margin and rounded lateral edges.
Behind the head is a very slight narrowing of the body. Eyes normally
two, rather close together and removed from the anterior end. Dorsal
surface uniformly gray or somewhat mottled, ventral side lighter.
Cannot be separated by its external features from some other pigmented
species of the same genus or from Planaria daotyligera and Hymanella
Yetenuava, Testes numerous), predominantly ventral, situated on either
side in a broad zone extending from behind the ovary to the level of
the mouth. There is no common genital atrium developed. Penis with a
spherical, muscular bulb and a conical, rather stiff, pointed papilla.
Vasa deferentia enter bulb anteroventrally retaining their expanded
shape as spermiductal vesicles, form a few convolutions within the bulb,
and open separately into the seminal vesicle. This cavity tapers pos-
teriorly to a narrow straight canal, the ejaculatory duct, which opens
at the tip of the penial papilla. The openings of the common oviduct and
of the bursa stalk are close to the gonopore. Cocoon ellipsoidal,
unstalked. Only sexual reproduction observed. In seepage springs, North
Carolina. Literature: Kenk (1970a).
36
-------
phagoaata bursapepforata Darlington, 1959
Sfynonym: Fontiaola bursaperforata: Ball, 1969.
37 YT 38 b bp de bd
ode
sv
I I / I
od am af gp sph
Fig. 37 after Carpenter (1970).
Fig. 38 after Darlington (1959).
A slender species, sexually mature specimens measuring 10-14 mm in length
and about 1 mm in width. Head truncate, with slightly convex frontal
margin and a pair of rather long, thin, pointed auricles extending
anterolaterally, with a slight constriction behind them. Eyeless and
unpigmented, white. Pharynx situated rather far back. Testes in moder-
ate number, prepharyngeal, may bridge the entire dorsoventral diameter
of the body. Penis consists of a relatively small bulb and a cylin-
drical papilla tapering toward the tip. The vasa deferentia narrow
behind the spermiductal vesicles, enter the penis bulb ventrolaterally,
and unite within the bulb to a straight canal, the ejaculatory duct,
without passing through an enlarged cavity or seminal vesicle. The duct
widens only moderately just before its opening at the tip of the penis
papilla. Genital atrium divided into male and female atria. The outlet
of the common oviduct is on the dorsal wall of the male atrium, rather
far removed from the female atrium. The copulatory bursa appears sac-
shaped in sagittal section but extends laterally toward the right poste-
rior intestinal ramus and communicates with it. Bursal canal narrow in
its anterior part, widening as it descends behind the male atrium, and
equipped with a terminal muscular sphincter. In springs, rivulets, and
a cave (J. H. Carpenter, personal communication), Georgia and Alabama.
Literature: Darlington (1959).
37
-------
Phagocata tahoena Kawakatsu, 1968
Synonyms: Phagooata nivea tahoena Kawalcatsu, 1968; Fontiaola nivea tahoena'
Ball, 1969.
39 ^ 40
bd ode v i
sv
vd pi am pp gl
Fig. 40 after Kawakatsu (1968),
A rather plump and thick planarian reaching over 12 mm in length and about
2 mm in width (but may be sexually mature at half that size). Anterior
end rounded, frontal margin with a small median projection when the animal
is gliding. No neck constriction. Eyes two, at a distance from each
other of about 1/3 the width of the head or less. Dorsal side with a
brown pigment which fades out toward the lateral margins and leaves a
crescent-shaped area along the frontal margin white. Ventral surface
unpigmented, white. Testes predominantly ventral, extending from the
level of the ovaries to the posterior end. Penis with well-developed
bulb and conical or finger-shaped papilla. Penial lumen wide, traversing
the penis from the bulb to the tip of the papilla, not clearly differen-
tiated into seminal vesicle and ejaculatory duct. Sperm ducts open
separately into the anterior part of the penis lumen. Oviducts uniting
above the atrium and opening into its posterior section. Bursa and bursa
stalk lined with a very thick epithelium, the stalk running on the left
side of the penis and lacking a differentiated vagina. Known only from
Lake Tahoe, California and Nevada, at depths of 15 to 1632 feet (4.6 to
500 m). Literature: Kawakatsu (1968), Kenk (1970a).
38
-------
41
Phagooata crenophila Carpenter, 1969a
b 0.5mm
de am ode pp gp
bd
Fig. 42 after Kenk (1970a).
Up to 22 mm long and 2 mm wide. Head truncated, with straight, convex,
or slightly wavy frontal margin. Lateral edges rounded, protruding
laterally to some extent so that there is a narrowing or neck behind
them, rather conspicuous in some populations. Eyes normally two, their
distance from each other being about 1/3 the width of the head, the
distance of each eye from the lateral margin smaller than from the fron-
tal margin. Dorsal side gray to black, ventral side lighter. Dis-
tinguishable from other similar species only by anatomical characters.
Testes predominantly ventral, extending posteriorly to almost the tail
end of the body. Penis consists of a weakly muscular bulb and a short,
rather plump papilla the tip of which is drawn out into a flattened lobe.
Sperm ducts enter the bulb anteroventrally and unite within the bulb to
a short common vas deferens. This opens into a small cavity, the seminal
vesicle, from which a narrow ejaculatory duct proceeds in a postero-
ventral direction to open on the ventral side of the papilla. The common
oviduct opens into the posterodorsal part of the male atrium, the duct of
the copulatory bursa ends very close to the gonopore. The most outstand-
ing specific characteristics are the lobular tip of the penis papilla, the
configuration of the penial lumen, and the postpharyngeal extension of the
testicular zone. In cold mountain springs and streams in the Rocky
Mountains and the Sierra Nevada. Literature: Carpenter (1969a), Kenk
(1970a).
39
-------
Phagoaata mopgani morgani (Stevens § Boring, 1906)
Synonyms: Planaria trunoata Leidy, 1851; P. movgani Stevens § Boring, 1906;
P. albissima: Kepner $ Rich, 1918 (not Vejdovsky, 1883); Dendroeoelum
truneatim: Girard, 1893; Fonticola truncata: Hyman, 1931; F. morgani: Castle
§• Hyman, 1934; F. morgani movgani: Ball, 1969; Phagoeata morgani: Hyman,
1937; IDendroeoelwn superbum Girard, 1850; IGaleooephala supevba: Stimpson,
1857; IPhagooata aavernioola Hyman, 1954; IFontiaola oavem-Lcola: Ball, 1969.
43
44
Fig. 44 after Kenk (1935).
Average length of sexually mature specimens 14 mm, width 2 mm. Anterior
end truncate, frontal margin straight, convex, or concave when the animal
is in motion. No adhesive organ. Lateral edges of head rounded, with a
very slight narrowing behind them. Eyes normally two, close together and
far removed from the frontal margin. Unpigmented, white. Anterior ramus
of intestine ends behind the eye level in adult specimens. Externally
similar to P. nivea ( for differences see that species) and Planavia
oooulta (which has the intestine ending in front of the eyes). Penis with
rather small bulb and conical or rounded papilla. Characteristic for the
species is a muscular, wart-like structure with thin, flattened epithelium
at the tip of the penis papilla. The sperm ducts enter the penial bulb
laterally and unite to a generally tubular ejaculatory duct which opens
into the atrium on the ventral side of the penis papilla. There is no
distinct seminal vesicle developed. Bursa sac-shaped, its duct runs
posteriorly, usually to one side (generally left) of the midline, widens
gradually without forming a pronounced vagina, and ends close to the
gonopore. The common oviduct enters the posterior part of the male atrium
near the outlet of the bursal duct. Reproduction sexual and by fission.
Cocoon ellipsoidal or spherical, unstalked. Inhabitant of springs and
cold creeks of eastern North America from New Brunswick to North Carolina
and west to Wisconsin and Kentucky, also found in Taylor Creek, El Dorado
County, California (here apparently introduced). Principal literature:
Stevens $ Boring (1906), Kenk (1935, 1944).
40
-------
Phagoeata morgani polyoelis (Kenk, 1935)
Synonym: Font-ioola movgani var. polyoelis Kenk, 1935.
45
Fig._45 after Kenk (1935)
Differs from the subspecies morgani chiefly in the number and arrangement
of the eyes. The eye spots are packed closely together, arranged in a
pair of almost parallel longitudinal rows in front of the area covered
by the intestine. Their number increases during the growth of the animal
from a minimum of 3 on one side in freshly hatched specimens to a maximum
of about 40 on the adults. There is no difference between the two sub-
species in the anatomy of the reproductive system. Recorded from Virginia
and Ontario. Principal literature: Kenk (1935).
41
-------
Phagocata nivea Kenk, 1953
Synonyms-.Phagoaata nivea nivea Kawakatsu, 1968; Fontioola nivea nivea: Ball,
1969.
46
47
Fig. 47 after Kenk (1953).
Up to 8 nun long and 1.5 nun wide. Head truncated, with slightly bulging
frontal margin and rounded lateral corners. No distinct neck. Body
unpigmented, white. Eyes normally two, close together, rather far
removed from the frontal margin. The species resembles P. movgani ex-
ternally and can be separated from it only by anatomical characters.
Testes numerous, predominantly ventral although a few testes may move
toward the dorsal side between the branches of the intestine. They are
arranged on either side in a zone extending to almost the posterior end
(in P. movgani this zone ends at the level of the mouth). The penis
consists of a spherical bulb and a bluntly conical papilla. The two
sperm ducts open separately into the elongated penial cavity which is
not clearly divided into a seminal vesicle and an ejaculatory duct,
though it gradually narrows posteriorly. It opens into the atrium on
the ventral side of the papilla. There is no special muscular differ-
entiation or wart at the tip of the papilla (such as is seen in P. mov-
gani~). The atrium may appear divided into two chambers, the male and
common atria, but this division is not always evident. The two oviducts
unite in the space above the male atrium, the common oviduct opening
into the atrium posterodorsally. Copulatory bursa large, bursal duct
displaced to the left of the midline and surrounded by a coat of inter-
mingled circular and longitudinal muscle fibers (in P. morgani it has
two separate layers, a circular one adjoining the epithelium and a lon-
gitudinal layer). Cold-stenothermic, occurring in mountain streams in
Alaska. Literature: Kenk (1953).
42
-------
Phagooata graeilis (Haldeman, 1840)
Synonyms: Planavia gvaailis Haldeman, 1840; Euptanaxn-a gpaeilis: Kenk,
1930; Fontioola gvaoilis: Castle § Hyman, 1934; Phagoaata grao-Llis
graeil-is Hyman, 1945; Fontteola gvaoilis graoilis: Ball, 1969; Phagooata
subterTanea Hyman, 1937; Fontieola subtevTanea\ Ball, 1969.
48
49
0.5 mm
ode
Fig. 49 after Kenk (1970c).
A polytypic species, Mature specimens 8^30 mm long and 1.5-6 mm wide.
Head truncated with straight or slightly bulging frontal margin.
Generally there is a pair of rounded auricles protruding laterally, but
in some populations (in Virginia) these may be subdued. Color generally
a shade of gray or brown, darker dorsally than ventrally. In subter-
ranean populations the pigmentation may be almost entirely lacking.
Eyes normally two, supernumerary eyes occasionally seen, also eyeless
specimens in some subterranean habitats. Pharynges multiple. Testes
numerous, predominantly ventral, extending to posterior end. Vasa
defe entia with a characteristic backward loop before entering the penis
bulb. Penis with highly muscular bulb and elongated pointed papilla.
Below the outer epithelium of the papilla is a layer of fine fibers,
apparently of an elastic nature, to which longitudinal muscles coming
from the bulb attach. Penis lumen divided into two cavities, an anteri-
or nonglandular one and a larger glandular vesicle. The sperm ducts
open into the anterior vesicle separately or united. The ejaculatory
duct, of variable diameter, proceeds from the posterior vesicle to the
tip of the papilla. The muscle coat of the bursal stalk consists of
two layers, circular and longitudinal, throughout its length. For
differences from P. woodttorthi- see that species. Cocoon spherical or
ellipsoidal, unstalked. Reproduction only sexual. Eastern United
States south of the Delaware River and west to Missouri. Principal
literature: Peaslee (1910), Kenk (1935, 1970c), Hyman (1937a).
43
-------
Phagocata woodworthi Hyman, 1957a
Synonyms: Planaria gpaeilis: Girard, 1850 (not Haldeman, 1840); Phago-
cata gracilis: Girard, 1850 (and other authors before 1937); Euplanaria
gracilis: Hyman, 1931 (in part); Phagocata gracilis woodworthi Hyman,
1951; Fontioola gracilis woodworthi: Ball, 1969.
50
51
0.5mm
bd ode sph
I
gp
b vd mp pi od
Fig. 51 after Kenk (1970c).
Adult specimens 15-30 mm long and 2^4.5 mm wide. Head truncate, frontal
margin assuming a straight, convex, or concave shape during locomotion.
Lateral edges of the head rounded, very little protruding laterally.
Dorsal side dark gray, brown, or almost black, ventral side lighter.
Eyes normally two, situated at a distance of about 1/3 the head width.
Pharynges multiple. Cannot be distinguished externally from
P. gvao-tlis. Testes numerous, mainly ventral (but also dorsal) to the
intestine. Vasa deferentia behind the spermiductal vesicles approach
the penis bulb directly without forming a backward loop. Penis with
feebly muscular bulb and a stout, short, usually truncate papilla. The
outer musculature of the papilla consists of a thick layer of inter-
mingled circular and longitudinal fibers very characteristic for the
species. Sperm ducts enter bulb anterodorsally or anterolaterally,
unite in the bulb and open into the wide penial lumen which is not
differentiated into seminal vesicle and ejaculatory duct. The very
wide bursal canal is displaced to the right of the midline. It is
coated, in the anterior part, by the usual two layers of muscle fibers,
circular and longitudinal. As it approaches the atrium near the gono-
pore, its muscular envelope becomes very thick and consists of inter-
mingled longitudinal and circular fibers. Reproduction probably only
sexual. Northeastern parts of the U. S, north of the Delaware River
and eastern Canada as far west as Ontario. Principal literature:
Woodworth (1891, misidentified as P. gvao^Hs), Hyman (1937a), Kenk
(1970c).
44
-------
Sphalloplana perooeoa (Packard, 1879)
Synonyms: DendTOQoelwn pereoeeum Packard, 1879; Fontioola percoeaum: Hyman,
1931.
co bp bd ode
JO , i I
I I I
52
1
[
V
1
1
I
d
i
i
i
PP
gp
Fig. 52 after Carpenter (1970), modified.
Fig. 53 after Hyman (1937b), modified.
Length, up to 16 mm, width about 3 mm. Head truncate, with convex frontal
margin and a pair of prominent, rounded auricles projecting antero-
laterally. A weak adhesive organ is present in the center of the anterior
margin, consisting of a subterminal depression covered with an eosino-
philic glandular epithelium to which retractor muscle fibers attach.
Eyeless and without pigment, white. Lateral margins of the body with a
zone of tall epithelial cells containing long rhabdites. Testes dorsal,
prepharyngeal. The penis has a rounded bulb and a conical papilla. The
vasa deferentia enter the bulb anterolaterally and open separately into
the elongated penial lumen which is not distinctly divided into a seminal
vesicle and an ejaculatory duct and opens at the tip of the papilla. The
common oviduct enters the genital atrium from the dorsal side at the
transition between the male atrium and the small common atrium. The
copulatory bursa is sac-shaped, its stalk rather narrow in the anterior
portion which passes above the penis and widens only near its opening
into the common atrium behind the mouth of the common oviduct. Caves in
Kentucky and probably in neighboring states. Principal literature:
De Beauchamp (1931), Buchanan (1936), Hyman (1937b).
45
-------
Sphalloplana aldbamensis Hyman, 1945
54
ph bp vd
\i
\ ll i I i
de pp am bd ode
ac
i
KP
Fig. 54 and 55 after Hyman (1945),
modified.
Preserved specimens 5-6 mm long. Anterior end truncate, without auric-
ular extensions. Eyeless, in life presumably white. Body margins with
large rhabdites. Adhesive organ at center of frontal margin, consisting
of a depression of the epidermis through which eosinophilic glands open,
surrounded by an arrangement of mostly radial muscle fibers. Testes
rather few, in a pair of longitudinal rows, prepharyngeal. Copulatory
complex excessively glandular. Vasa deferentia open into a small chamber
(seminal vesicle?) apparently before entering the penis bulb. From this
chamber the narrow ejaculatory duct traverses the bulb and the short,
rounded penis papilla, widening in the papilla. Epithelium of the
papilla tall and glandular. Male atrium separated from the common atrium
by a constriction. The common oviduct opens into the posterior part of
the male atrium. Copulatory bursa rather small, its duct with irregular
enlargements, glandular in its distal portion. Cave in Alabama.
Literature: Hyman
46
-------
Sphalloplana geovg-iana Hyroan, 1954
56
vd bp
Fig. 56 and 57 after Hyman (1954),
modified.
Described from two defective specimens. Preserved animals about 8 mm
long. Head truncate, with a central adhesive organ. Blind and un-
pigmented. Adhesive organ is a subterminal cuplike depression with
outlets of eosinophilic glands and longitudinal retractor muscles.
Testes small, ventral, forming on either side a band occupying the
posterior half of the prepharyngeal region. Penis with muscular bulb
and conical papilla. The vasa deferentia enter the bulb ventrolaterally
and open separately into a rounded seminal vesicle. This continues
posteriorly as a straight ejaculatory duct to the tip of the papilla.
The common oviduct opens into the roof of the male atrium. Copulatory
bursa not studied. Bursal duct narrow in its anterior portion, pro-
ceeding posteriorly beyond the gonopore, then turning abruptly antero-
ventrally and widening considerably. This widened portion, interpreted
best as a vagina, is surrounded by a thick layer of chiefly circular
muscle fibers. Known from a cave in Georgia. Literature: Hyman (1954).
47
-------
58
Sphalloplana virginiana Hyman, 1945
59
am pp
Fig. 58 and 59 after Hyman (1945),
modified.
Preserved specimens up to 12 mm long, with bluntly rounded head provided
with a central weak adhesive organ. Eyeless and unpigmented. The ad-
hesive organ forms an irregular depression with glandular openings and a
band of muscle fibers serving as retractors. Body margins with a rim
containing very large rhabdites. Testes ventral and prepharyngeal.
Penis with feebly developed bulb and medium-sized, rounded papilla. Vasa
deferentia enter separately into the penis lumen which is not subdivided
and opens on the ventral side of the papilla. Common ovovitelline duct
opens into posterior part of male atrium. Copulatory bursa rather small,
bursal duct initially narrow, widening slightly as it approaches the
atrium. Known only from a cave in Virginia. Literature: Hyman (1945).
48
-------
Sphalloplana kanaensis Hyman, 1945
60
\
pp am da gp ode
Fig. 60 after Hyman (1945), modified.
Preserved specimens 20 ram long, eyeless, white. Anterior end truncate,
with conspicuous adhesive organ. Marginal rhabdites only slightly
larger than those of the general epidermis. Adhesive organ consists of
a depression with openings of eosinophilic glands and muscular differ-
entiations. Testes dorsal, prepharyngeal. Penis bulb slightly devel-
oped, penis papilla rounded, with weak musculature. Sperm ducts unite
outside the penis bulb to form a canal, the ejaculatory duct, which
after a short sinuous course opens at the center of the penis papilla.
Male atrium with very tall epithelium, receiving at its posterior border
the long common oviduct. The small common atrium, which connects with
the bursal canal, has irregular blind diverticula (important specific
characteristic). Bursal canal long, narrow in its anterior section but
widening as it approaches the atrium. Known only from a spring in
Kansas. Literature: Hyman (1945).
49
-------
Sphalloplana prioei (Hyman, 1937b)
Synonym: Speophila pricei Hyman, 1937.
61
bd ode
1
SV V
, «, C _ , ' r * — ^7-^, ~^', ~~~
\
j
1
s pp
- ~~ ' ~4
1
1
1
am
*«*
\
\
\
m
Fig. 62 after Hyman (1937b).
Mature animals up to 28 mm long and 3.5 mm wide. Head truncate, with
gently bulging frontal margin and rounded lateral edges (auricles) which
in quiet gliding protrude somewhat anteriorly and laterally. Behind the
auricles is a constriction or neck. A well-developed adhesive organ
forming a deep protrusible invagination with glandular and muscular
differentiations is located in the center of the frontal margin. Pigment-
less (white) and blind. Lateral margins with a zone of modified epithe-
lium containing very large rhabdites. Testes dorsal, prepharyngeal.
Penis with rounded bulb and short, generally conical papilla. Vasa de-
ferentia form spermiductal vesicles at the level of the pharynx, then
narrow again as coiled canals and enter the penis bulb separately. In
the bulb they unite and soon widen to an elongated seminal vesicle from
which an ejaculatory duct proceeds to the tip of the penis papilla. The
common oviduct opens into the posterior part of the male atrium. Copu-
latory bursa of moderate size, bursa duct initially narrow but gradually
widens in the posterior portion which opens into the atrium close to the
gonopore. Caves in Pennsylvania. Literature: Hyman (1937b).
50
-------
Sphal'lop'Lana buchanani (Hyman, 1937b)
Synonym: Speophila buehanani Hyman, 1937.
63 r^ 64
pi od ode pp v
Fig. 63 after Carpenter (1970), modified.
Fig. 64. View from dorsal side (after Hyman, 1937b).
Up to 15 mm long. Anterior end truncate, with a well-developed, protru-
sible adhesive organ in the middle of the frontal margin, and rounded
lateral edges. No distinct narrowing or neck behind the head. Pig-
mentless and blind. Lateral margins with large rhabdites. Testes
dorsal, prepharyngeal. Penis with muscular bulb and well-developed,
cylindrical or conical papilla. The sperm ducts enter the bulb and
unite to a common canal representing the ejaculatory duct. There is no
histologically distinct seminal vesicle, but the duct may show a small
widening in its course. Common ovovitelline duct opening into the pos-
terior part of the male atrium. Bursal duct displaced to one side of
the midline, seems to be widened in its terminal section. Caves in
Kentucky. Principal literature: Hyman (1937b).
51
-------
Sphalloplana hubriehti (Hyman, 1945)
Synonym: Speophila hubriahti Hyman, 1945.
65 ^TX 66
vd
PP Pi
bd
m ode
Fig. 65 and 66 after Hyman (1945), modified.
Preserved specimens up to 17 mm long, with broadly rounded anterior end,
white, eyeless. Adhesive organ conspicuous, consisting of a deep pit
surrounded by many eosinophilic glands and provided with a complex system
of muscle fibers. Marginal rhabdites very large. Testes dorsal,
prepharyngeal. Penis with muscular bulb and long, finger-shaped papilla.
Vasa deferentia unite at the anteroventral border of the penis bulb and
traverse the bulb as a narrow canal, the common vas deferens. This opens
into a widened, elongated cavity, the penial lumen, which proceeds to the
tip of the papilla. Copulatory bursa large. Its outlet runs first above
the penis as a narrow canal, then widens considerably while being dis-
placed to the right side of the midline. Caves and springs in Illinois
and Missouri. Literature: Hyman (1945).
52
-------
Sphalloplana hofftnasteri (Hyman, 1954)
Synonym: Speophila hofpnasteri Hyman, 1954.
67 XDv 68
Fig. 67 and 68 after Hyman (1954), modified.
Preserved specimens about 11 mm long, plump. Anterior end rounded, with
adhesive organ. Eyeless and white. Lateral margins of the body with a
zone of large rhabdites. Adhesive pit is a deep epithelial invagination,
with glandular outlets and retractor and protractor muscles. Testes
prepharyngeal, rather large, situated on either side of the midline in a
band in the middle regions of the sections. Penis with large bulb and
elongated, conical papilla. The vasa deferentia enter the bulb and pro-
ceed to the base of the papilla where they unite to a short common vas
deferens. This empties into the elongated penial lumen which first widens,
then narrows again, and opens at the tip of the papilla. The shape of
the copulatory bursa is not definitely known. The bursal duct is rather
narrow in its anterior course while its distal part, behind the atrium,
is differentiated as a vagina with tall epithelial cells and what seem
to be muscular structures. Caves in West Virginia. Literature: Hyman
(1954).
53
-------
Sphalloplana weingartnevi Kenk, 1970b
69
70
ph
vd vs bp am pp
gp ode
Fig. 70 after Kenk (1970b).
Mature specimens 6-9 mm long and 1.2-1.8 ram wide. Anterior end truncate,
with slightly bulging frontal margin and rounded lateral edges, without
a neck constriction. Eyeless and unpigmented, white. Body relatively
transparent, showing in life the position of the anterior adhesive
organ, the pharynx, and in the postpharyngeal part the copulatory complex
as a round or elliptical transparent area with an opaque spot in the
center. The adhesive organ consists of a deep invagination of the
epithelium with irregular outline, which receives the outlets of many
eosinophilic glands and to which a system of retractor muscles is
attached. Testes numerous, both dorsal and ventral, prepharyngeal.
Penis has a large, spherical bulb and a short, finger-shaped papilla.
Vasa deferentia enter bulb ventrolaterally and empty separately, but
close together, into the elongated, glandular seminal vesicle. From this
vesicle a narrower, somewhat curved (contraction?), nonglandular ejacu-
latory duct proceeds to the tip of the penis papilla. Genital atrium
rather small, receiving the common oviduct in its posterior part. Copu-
latory bursa a rounded sac, bursal duct narrow in its anterior portion,
gradually widening as it curves down toward the gonopore but showing no
histologically distinct vagina nor a terminal sphincter. Found in a cave
in Indiana. Literature: Kenk (1970b).
54
-------
Sphalloplana mohri- Hyman, 1939c
Synonyms: Sphalloplana kutscheri Mitchell, 1968; S. sloani Mitchell, 1968;
S. zesahi Mitchell, 1968; S. reddelli Mitchell, 1968.
72
ode gp
Fig. 71 after Carpenter (1970), modified.
Fig. 72 after Mitchell (1968), modified.
Mature animals 20-35 mm long and 3-5 mm wide. Anterior end truncate,
with laterally protruding auricles. The glandular and muscular adhesive
organ, deeply invaginated when at rest, may be extended as a conical
projection through its opening at the center of the frontal margin.
Behind the auricles the head narrows to a neck, then the body widens
again. Eyeless and without pigment, white. Polypharyngeal, with up to
about 50 pharynges. Testes relatively few, situated between the in-
testinal branches, extending back to the level of the anterior pharynges.
Penis consists of a bulb of moderate size and a larger cylindrical or
conical papilla, both with feeble musculature and easily distorted in
preservation. The vasa deferentia enter the bulb anterolaterally and
unite in the penis papilla. The common vas deferens may show a widened
section, the equivalent of a seminal vesicle, and continue as ejaculatory
duct to the tip of the papilla. The common ovovitelline ducts open into
the posterior part of the male atrium (there is hardly any common atrial
cavity developed). Bursal duct with a narrower anterior section and a
wider, thick-walled, glandular and muscular terminal portion (vagina).
Caves in Texas. Principal literature: Hyman (1939c), Mitchell (1968).
55
-------
Kenkia rhynohida Hyman, 1937b
73
ode v
Fig. 73 and 74. Ventral and lateral views
of preserved specimen (after Hyman, 1937b),
Fig. 75 after Hyman (1937b).
Has not been studied in detail in life. Living specimens about 1/4 inch
(6 nun) long, of oval-domed shaped, white, with, a creamy center. Pre-
served animals 2-4 mm long, of oval outline, with a convex dorsal and
a concave ventral surface, described as being of the shape of a minia-
ture turtle. At the anterior end is a snout-like projection, cylindrical
in cross section, containing a deep epithelial invagination with glandular
and muscular differentiations, apparently serving as an adhesive organ.
The postpharyngeal region is much reduced in size. Eyes are absent. The
lateral margins of the body have a thick band of very large rhabdites.
Testes few, large, forming on either side a group located dorsolaterally
in the middle of the prepharyngeal region. Copulatory apparatus close to
posterior end. Penis with spherical bulb and elongated papilla which is
conical at its base and cylindrical in the distal part. The vasa defer-
entia enter the penis bulb, unite, and continue as a narrow canal, the
ejaculatory duct, to the tip of the penis papilla without forming an
enlargement or seminal vesicle. Copulatory bursa small, with narrow
stalk enlarged only in the distal portion close to the gonopore. Common
oviduct opens into posterior part of male atrium. Known only from
Malheur Cave, Harney County, Oregon. Literature: Hyman (1937b).
56
-------
76
Maorocotyla glandulosa Hyman, 1956
77
bd
ode gp
(Hyman's description emended by study of additional material). A large
species, up to 30 mm long and 6 mm wide. Anterior end truncate, with
straight or slightly waved frontal margin and rounded lateral edges
which protrude only little to the sides, causing a faint incurving to
appear behind them. A protrusible adhesive organ may be projected from
the center of the frontal margin during searching movements or upon
stimulation. Eyes lacking. Color in life a very light reddish-brown or
orange. The pigment is not granular but is diffused in the subepidermal
tissues, including the head and the body margins, absent only in the
pharynx, the copulatory complex, and the denser parts of the adhesive or-
gan. Lateral margins with large rhabdites. Testes ventral, prepharyngeal.
Penis with large bulb and conical papilla, flanked by voluminous masses
of eosinophilic glandular tissue which empty their secretions into the
penis lumen. The penis bulb contains a large cavity with irregular folded
outline, the seminal vesicle. From this cavity a rather wide canal, the
ejaculatory duct, proceeds to the tip of the papilla. The sperm ducts
open into the seminal vesicle independently, without uniting. Common
oviduct connects to the posterior part of the male atrium. Copulatory
bursa relatively large, with wide outlet opening near the gonopore. Re-
production sexual, with spherical egg capsules of over 3 mm diameter,
unstalked. The species resembles a Sphalloplana in all essential char-
acteristics except the arrangement of the pharyngeal muscle fibers,
which is of the dendrocoelid type (see p. 13). Caves (J. H. Carpenter,
personal communication) and spring in Missouri and Iowa. Literature:
Hyman (1956).
57
-------
DendroGoelopsis vaginata Hyman, 1935
78
%'!'•&
IP
de f
ac gp ode
Mature specimens 14-22 mm long and about 3 mm wide. Head truncate, with
gently bulging frontal margin, a weak subterminal adhesive organ, rounded
corners, and a slight narrowing (neck) behind them. Eyes usually two,
their distance from each other about 1/3 the width of the neck, the dis-
tance of each from the frontal margin a little larger than from the later-
al margin. Dorsal side gray or brown, ventral surface somewhat lighter
(the pigment bleaches easily after preservation). Pigment absent from
the two eye fields, a point near the mouth, and a circular area around the
gonopore. A pair of somewhat lighter oblique streaks on the head laterally
to the eyes. Similar to D. piriformis but with different body outline
(largest width is reached in the prepharyngeal region). Testes numerous,
ventral, reaching to level of gonopore (in D. piviformis dorsal, to pos-
terior end). Penis with large spherical bulb and finger-shaped papilla.
The bulb contains a large, round, highly glandular seminal vesicle and
receives the two sperm ducts ventrolaterally. From the vesicle a narrower
ejaculatory duct proceeds to the tip of the papilla. Below the flattened
outer epithelium of the penis papilla is a dense fibrous layer followed
by a layer of longitudinal muscles. The outlet of the copulatory bursa
widens as it proceeds posteriorly and gradually acquires a folded outline.
It opens into the atrium close to the gonopore. Egg capsule spherical,
about 1.6 mm in diameter. Springs, streams, and lakes in Montana, Oregon,
and Washington. Literature: Hyman (1935).
58
-------
80
Dendpoooelopsis piriformis Kenk, 1953
81
m
Fig. 81 after Kenk (1953).
A rather broad and plump species, up to 15 mm long and 3 mm wide. Head
truncated, with a bulging central portion of the frontal margin indi-
cating the site of the adhesive organ. In quiet gliding the greatest
width of the body is reached at the beginning of the last third. A pair
of rounded auricles protrude only little laterally, with a shallow con-
striction behind them. When resting, the body appears pear-shaped, the
lateral margins often showing a ruffled outline. Eyes usually two,
situated at a distance of about 1/3 the width, of the head. Pigmentation
dorsally brown or gray, with a dark field between the eyes and extending
to the frontal margin on both sides of the adhesive organ which itself
is unpigmented. There may be one median and often a pair of additional
lateral longitudinal dark stripes along the body. Ventral side lighter
gray. Pharynx short, of the dendrocoelid type. Testes numerous, dorsal,
arranged in a pair of wide bands reaching close to the posterior end (in
the similar D. vaginata the testes are ventral). Penis with large,
spherical bulb and long, finger-shaped, pointed papilla. Sperm ducts
enter the bulb anterolaterally and open into the seminal vesicle inde-
pendently, each on a conical projection. Seminal vesicle large, with
glandular wall of irregularly lobed outline. Ejaculatory duct narrow and
long, opening at tip of penis papilla. Common oviduct opens into atrium
close to gonopore. Bursa and bursal canal without peculiarities. In-
habitant of lakes and streams in Alaska and northwestern Canada. Principal
literature: Kenk (1953), Holmquist (1967).
59
-------
82
DpndroGoelopsis alaskensis Kenk, 1953
83
vd vs
i i
a ode gp bd
Fig. 83 after Kenk (1953),
A large, white species, not yet studied at full maturity. Largest speci-
men examined 20 mm long and 4 mm wide. Anterior end truncated, with
slightly convex central part of the frontal margin flanked by rounded
auricles protruding anteriorly and laterally. No distinct adhesive or-
gan. Eyes normally two, separated by about 1/3 the width of the head and
nearly equidistant from the frontal and lateral margins. There sometimes
are smaller accessory eyes developed in front of or behind the principal
eyes. The description of the reproductive system is based on the exami-
nation of one specimen which was not fully mature. Primordia of the
testes predominantly ventral, traceable posteriorly to the level of the
mouth (they possibly may extend farther back in fully mature animals).
Penis with spherical bulb and short papilla. Vasa deferentia open sepa-
rately into the penial lumen. Penis cavity in wide communication with
the undivided genital atrium. Common oviduct opens into atrium anterior
to the opening of the bursal duct. In springs and cold creeks in Alaska.
Literature: Kenk (1953).
60
-------
Dendroooelops-is hymanae Kawakatsu, 1968
bp vs bd ode gl
de pp
Fig. 84 after Kawakatsu (1968).
Has not been studied in life. Description was prepared from one mature
and one immature specimen in defective condition. Unpigmented (in life
"translucent pink"), eyeless, up to 14 mm long and 2 mm wide. Anterior
end with subterminal adhesive organ forming a depression with glandular
and muscular differentiations. Testes numerous, of moderate size,
ventral, extending posteriorly to the base of the pharynx. Penis with
highly muscular spherical bulb and conical pointed papilla. Seminal
vesicle divided into a pair of elongated cavities, each receiving one of
the sperm ducts anterolaterally. The two cavities unite at the base of
the papilla from where a rather narrow ejaculatory duct proceeds to the
tip of the papilla. The common oviduct opens into the posterior part of
the male atrium from the dorsal side. There is no common atrium devel-
oped. The outstanding specific characteristics are the lack of eyes
and the division of the seminal vesicle. Known only from Lake Tahoe,
California and Nevada, from a depth of about 1600 feet. Literature:
Kawakatsu (1968).
61
-------
Dendroooelopsis americana (Hyman, 1939a)
Synonym: Soroaelis ameriaana Hyman, 1939.
85 I.,"/ 86
ode
gp bd
Fig. 87. Spiny wart of the penis sur-
face, enlarged (after Hyman, 1939c).
Up to 18 nun long and 2 mm wide. Anterior end of quietly gliding animal
truncate, with convex frontal margin, laterally and anteriorly protruding
auricular appendages, and a distinct narrowing or neck behind the auri-
cles. Adhesive organ subterminal, weakly developed. Eyes numerous,
arranged in two almost parallel longitudinal rows placed at a distance
from each other of about 1/3 the width of the neck region. Number of
eyes increases during growth. Body unpigroented, white. Testes not
numerous, predominantly dorsal, extending posteriorly to behind the cop-
ulatory complex. Sperm ducts unite at a variable distance anterior to
penis. Penis with muscular rounded bulb and conical papilla. The common
vas deferens enters the bulb anteriorly, traverses the bulb as a straight
narrow canal and opens into a highly glandular cavity with irregular out-
line, the seminal vesicle. From there the narrow ejaculatory duct runs
to the tip of the penis papilla. Cells of the outer epithelium of the
papilla appear as wartlike eminences, each studded with colorless,
pointed spines (this can be seen on the freshly extirpated penis). Com-
mon oviduct opens into the posterior end of the male atrium. Copulatory
bursa absent, bursal canal blindly closed and covered by a thick layer of
muscles. Cocoon almost spherical, unstalked. Inhabits caves and springs
in Oklahoma and Arkansas. Literature: Hyman (1939a, 1939c).
62
-------
Procotyla fiuv-iatilis Leidy, 1857
Synonyms: Vendroooelim superbum: Leidy, 1851 (not Girard, 1850); D.
laeteum: Woodworth,1896 (not 0. F. Miiller, 1774); D. graffi Wilhelmi,
1909; W. pulchervimum Girard, 1850; lOligooelis puldherrima:
Stimpson, 1857.
88
89
0.5 mm
bd de ode
I I
...
Mature animals 12-20 igm long and 2-5 mm wide. Head truncated, rather
variable in shape during locomotion, Subterminal adhesive organ
clearly visible in living animal, bulging out slightly during gliding,
flanked by two rounded auricular projections with a constriction or
neck behind them. Eyes of variable number, in the northern distribu-
tional area 1-8 on either side, in the sourthern states usually only 1.
The groups of eyes are rather far removed from each other, by about 1/2
the width of the head. Testes numerous, dorsal, extending posteriorly
to level of copulatory complex. Vasa deferentia unite at the level of
the penis and enter the penis bulb anteroventrally as common vas
deferens. The penis consists of an elongated muscular bulb with a large
cavity and a short conical papilla. The bulb is composed of several
layers surrounding the cavity: an outer, very thick layer of longitudi-
nal muscles, a fibrous layer, an inner layer of chiefly longitidinal
muscle fibers (in P. typhlops these are circular), and the glandular
epithelial lining of the cavity or prostate. The common vas deferens
runs along the ventral wall of the penial cavity and connects with it
at the base of the penis papilla. Beyond this junction the lumen con-
tinues into the penis papilla as ejaculatory duct and opens at its tip.
The common oviduct and the outlet of the sac-shaped copulatory bursa
open close to the gonopore. Cocoon ellipsoidal, unstalked. The species
refuses liver or Tubifex as food but takes living amphipods, isopods,
and aquatic insect larvae. Inhabits ponds, lakes, streams and springs
in the eastern part of North America, from Maine to Louisiana and west
to Ontario, Wisconsin, and Illinois. Principal literature: Woodworth
(1897), Hyman (1928), Kenk (1944).
63
-------
Proootyla typhlops Kenk, 1935
90
vc
c v
1
me
d
|
mi
vc
t i
1 i
epr !
c pr
de ' od
am
gp I bd
gl
Fig. 91 after Kenk (1935).
A small, slender species, up to 12 mm long and 1.3 mm wide. Head truncate,
with straight or slightly convex frontal margin, rounded lateral edges,
and an insignificant narrowing behind them. No distinct adhesive organ.
Eyeless and white. Testes numerous, predominantly dorsal, extending
posteriorly to the region of the pharynx. Vasa deferentia, after passing
through the spermiductal vesicles, unite outside the penis bulb to a com-
mon sperm duct which is also sinuous and expanded. The penis consists of
a large, elongated, muscular bulb and a short, conical papilla. The com-
mon vas deferens enters the anterior part of the bulb and continues the
entire length of the bulb ventrally to a large, glandular cavity with
which it communicates at the base of the papilla. The role of this cavity
is probably that of a prostatic gland adding its secretions to the sperm
passing through the sperm duct. The wall of the penis bulb has three
layers: an outer, thick layer of approximately longitudinal muscle fibers,
a thin fibrous layer, and an inner layer of circular muscles. The lumen
of the penis papilla, representing the ejaculatory duct, tapers toward the
tip of the papilla where it opens into the atrium. The copulatory bursa
is typical, sac-shaped, its duct at first rather narrow but gradually
widening posteriorly without a histologically differentiated vagina. Ap-
parently a subterranean species found in springs and groundwater pools in
Virginia (a two-eyed form alleged to be this species has been reported
from Florida, but this record needs confirmation). In the laboratory the
species accepts liver and Tubifex as food. Literature: Kenk (1935).
64
-------
Eeatocephala exotica Hyman, 1953b
93
vdc pb am gp
Fig. 92 and 93 after Hyman (1953 b).
Preserved specimen 14 nun long. Anterior end appears co be truncate, with
a central adhesive organ. Eyes two, Pigmentation uniformly black. Ad-
hesive organ forms an irregular invagination of glandular epithelium,
equipped with a complex system of muscle fibers. Ovaries rather far back
from the head, testes not seen. Copulatory complex consists chiefly of
a large cavity lined with a tall glandular epithelium which forms villus-
like projections. This cavity is interpreted by Hyman as an enlarged
bulbar lumen. The sperm duct (called ejaculatory duct by Hyman), appar-
ently originating from the union of the paired ducts, enters the anterior
wall of the cavity and projects into its lumen. The common oviduct opens
into the posterior sector of the male cavity which is lined by a normal,
nonglandular epithelium (interpreted as the remnant of a male atrium).
The copulatory bursa, situated behind the pharyngeal pouch, has an exceed-
ingly long, narrow canal running backward in the dorsal midline, widening
only in its posterior portion, then narrowing again and opening into the
atrium near the gonopore. Only one specimen has been studied, collected
in the Shaw Lily Ponds (now Kenilworth Aquatic Gardens), Washington,
District of Columbia, apparently introduced with exotic water plants.
Literature: Hyman (1953b).
65
-------
NOT RECOGNIZABLE SPECIES
Planaria simplex Woodworth, 1896a . Described from one immature specimen
of 4 mm length and 1.8 mm width, of ovate
94 shape. "Anterior end rounded, set off from the
rest of the body by slight lateral indentations
at the level of the eyes, i. e. at about 1/10
total length from the anterior end. No evi-
dence of cephalic appendages. ... Pigment
located in spots of nearly uniform size, dis-
tributed uniformly over all parts of the body;
no clear areas surrounding eyes or at sides of
head. Color of alcoholic specimen ochre-yellow."
Dredged off New York Point, Lake Michigan (on
Grand Traverse Bay, Michigan). Literature:
Woodworth (1896a, 1896b).
After Woodworth (1896b).
Planaria fuKginosa Leidy. 1851 . Synonyms: Planc&ia (Typhlolepta?)
fuliginosus Leidy, 1851; AnoaeUd fuHginosa: Stimpson, 1857. "Body
oval, dilated; inferiorly flat; superiorly moderately convex,
fuliginous. Eyes none: in their ordinary position a slightly greater
accumulation of black pigment upon the upper surface. Mouth inferior,
a little posterior to the centre; oesophagus simple, cylindrical, white,
1 line long by 1/2 line broad." Length 8 mm, breadth 6 mm (extended?).
Rancocas Creek near Pemberton, Burlington County, New Jersey. Litera-
ture: Leidy (1851), Girard (1893).
Planaria unioniaola Woodworth> 1897 . Described from one specimen which
apparently had been sketched in life by the
collector and had been brownish red, mottled
95 with purplish dots (color of intestinal con-
tents?). According to the sketch the head is
truncate, with a sinuous frontal margin. The
two eyes show large circular periocular fields.
The posterior end is blunt, suggesting either
an injury or recent fission. After preser-
vation the specimen was much contracted and
shriveled, 2.8 mm long and 1.8 mm broad. Col-
lected on the mantle of Vnio in the Illinois
River near Havana, Illinois. Literature:
Woodworth (1897).
\J
After Woodworth (1897).
66
-------
Hydrolimax bruneus Girard, 1891 . Length 19 ram, width 3 mm. Head ob-
o, tusely truncate, with a neck constriction
'O behind it. Eyes two, Body dark brown, head
somewhat lighter. Intestinal coecuro reaches
anteriorly between the eyes. Genital open-
ing anterior to the middle of the body, with
a thickening at its level caused by the pres-
ence of eggs or cocoons. Found in a small
stream in Fairmount Park, Philadelphia, Penn-
sylvania. This may be one of the brown spe-
cies of Phagocata., or Hymanella3 or may not
be a triclad. Literature: Girard (1891, 1893)
After Girard (1893).
67
-------
SECTION V
ABBREVIATIONS USED ON FIGURES
The figures of the copulatory apparatus are diagrams reconstructed
from adjoining sagittal sections. Their orientation is with the an-
terior parts to the left. In the outline drawings, pigmented species
are indicated by shading, unpigmented species are white.
Abbreviations used are as follows:
a genital atrium or antrum i
aa atrium of adenodactyl m
ac common atrium or antrum me
ad adenodactyl mi
af female atrium or antrum mp
ai anterior intestinal ramus n
am male atrium or antrum od
ao adhesive organ ode
au auricle
avs anterior seminal vesicle ov
b copulatory bursa or bursa pa
copulatrix pb
bd bursal duct, canal, or stalk ph
bg glandular portion of bursal pi
canal pi
bp penial bulb pp
br brain or cerebral ganglion pr
eg cyanophilic glands pvs
co copulatory apparatus or r
complex sg
de ejaculatory duct sn
dp diverticulum of penial sph
lumen sv
eg eosinophilic glands
epe external epithelium t
epi internal epithelium v
epr epithelium of prostate vd
f fibrous layer vdc
gl glands vi
gp gonopore or genital aperture vs
intestine or gut
mouth or oral aperture
external muscle layer
internal muscle layer
muscles of penis papilla
ventral nerve cord
oviduct or ovovitelline duct
common oviduct or common
ovovitelline duct
ovary
parenchyma
penis bulb
pharynx
posterior intestinal ramus
penial lumen
penis papilla
prostate
posterior seminal vesicle
margin with large rhabdites
shell glands
snout
sphincter
spermiductal vesicle or
false seminal vesicle
testis
vagina
vas deferens or sperm duct
common vas deferens
vitellaria or yolk glands
seminal vesocle
69
-------
SECTION VI
ACKNOWLEDGMENTS
The cooperation of various publishers who kindly permitted the repro-
duction of figures from their in part copyrighted publication is
gratefully acknowledged: American Microscopical Society; the pub-
lishers of American Midland Naturalist; American Museum of Natural
History; University of Michigan; and Washington Academy of Sciences.
Mr Jack R. Schroeder and Mrs Carolyn B. Gast, Scientific Illustrators,
and Mr Victor E. Krantz of the Smithsonian photographic laboratories
provided valuable assistance in the preparation and copying of the
illustrations.
71
-------
SECTION VII
REFERENCES
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(1932). Biospeologica. LVI /should be LVIIIJ. Turbellaries,
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(1852). Descriptions of Two New Genera and Two New Species of
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Girard, C. (1891). Deux Especes Nouvelles de Planaires Americaines.
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(1928). Studies on the Morphology, Taxonomy, and Distribution of
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Commonly but Erroneously Known as Dendroaoelum lacteum. Trans-
actions of the American Microscopical Society, 47:222-255.
(1929). Studies on the Morphology, Taxonomy, and Distribution of
North American Triclad Turbellaria. II. On the Distinctions be-
tween Planaria agilis an Planaria dorotocephala, with Notes on the
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of the American Microscopical Society, 48:406-415.
(1931a). Studies on the Morphology, Taxonomy, and Distribution
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(Girard). Transactions of the American Microscopical Society,
50:124-135.
(1931b). Studies on the Morphology, Taxonomy, and Distribution
of North American Triclad Turbellaria. IV. Recent European Re-
visions of the Triclads, and Their Application to the American
Forms, with a Key to the Latter and New Notes on Distribution.
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(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.
i (1937a). Studies on the Morphology, Taxonomy, and Distribution of
North American Triclad Turbellaria. VII. The Two Species Confused
under the Name Phagocata gracilis, the Validity of the Generic
Name Phagocata Leidy 1847, and Its Priority over Fonticola Komarek
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56:298-310.
(1937b). 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
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Hyman, L. H. (1939b). North American Triclad Turbellaria. IX. The
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(1939c). North American Triclad Turbellaria. X. Additional
Species of Cave Planarians. Transactions of the American
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(1953a). Turbellaria (Flatworms). In: R. W. Pennak, Freshwater
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(1953b) • North American Triclad Turbellaria, XIV. A New,
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(1954). North American Triclad Turbellaria. XIII. Three New Cave
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(1963). North American Triclad Turbellaria. XVI. Fresh-Water
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Kawakatsu, M. (1968). North American Triclad Turbellaria. XVII. Fresh-
water Planarians from Lake Tahoe. Proceedings of the United States
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Mitchell Scientific Society, 51:79-125, 8 plates.
(1941). A Fresh-Water Triclad from Puerto Rico, Dugesia antillana,
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Kenk, R. (1970b). Freshwater Triclads (Turbellaria) of North America.
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31:1-16, 1 plate.
77
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SECTION VIII
INDEX OF SCIENTIFIC NAMES
agilis (Dugesia)3 22
- (Euplanaria) 3 22
- (Planaria) 3 22
aldbamensis (Sphalloplana) 3 46
alaskensis (Dendroeoelopsis) 3 60
albissima (Planaria) 3 40
ameriaana (Dendroooelopsis ) 3 62
- (Sovoaelis)3 62
Anooelis fuliginosa3 66
antillana [Dugesia) 3 23
borealis (Poly celts )3 27
- (Polyaelis aoronata) 3 27
brevipenis (Polyeelis ooronata)3 26
bruneus (Hydrolimax) 3 67
buahanani (Speophila), 51
- (.Sphalloplana) 3 51
bulbosa (.Phagoaata) , 36
bursaperforata (Fontieola) 3 37
- (Phagoeata) 3 37
oavernicola (Fontieola) f 40
- (Phagoeata), 40
coronata (Phagoeata) , 25
- (Poly cells )3 26
- borealis (Polyaelis) 3 27
- brevipenis (Polyoelis)3 26
- ooronata (PolyoeHs)3 25
orenophila (Phagooata) 3 39
Cura, 13
19
Curtisia fovemani, 19
- simplioi.ssima3 19
daotyligeva (Planaria) 3 28
- cfactz/ Hgera ' (Planaria) 3 28
- musoulosa (Planaria) 3 29
Dendvoaoelopsis3 14
- alaskensis3 60
- americana3 62
- hymanae3 61
- piriformis3 58,59
- vaginata3 12, 58, 59
Dendvoooelwn graffi3 63
- Iaoteum3 63
- peTaoeeum, 45
- pulaherrimm3 63
- superbim3 40, 63
- truneatum, 40
diabolis (Dugesia)3 22
dorotooephala (Dugesia)3 22
(Euplanaria)3 22
(Planaria), 22
Dugesia3 13
agilis3 22
antillana3 23
diabolis3 22
dorotocephala, 22
foremaniif 19
gonocephaloides, 20
Zata, 20
lugubris, 24
maeulata3 20
miavobur'salts3 20
modes"ta, 19
polyahroa3 24
tigrina, 20
Euplanaria agilis3 22
cZorotocepha^a, 22
graeilis3 43, 44
— Zata, 20
maoulata3 20
miorobursalis3 20
novangliaet 20
philadelphiaa3 22
tigrina3 20
exotica (Reotocephala), 65
fluviatilis (Proeotyla); 63
Fontieola bursaperforata3 37
cayerntcjo^aj 40
graeilis3 43
gracilis gvaoilis, 43
graoilis monopharyngea} 33
graailis woodwovthi3 44
movgani, 40
morgani movgani, 40
morgani polyoelis, 41
nivea3 42
n£uea nivea3 42
nivea tahoena3 38
ovegonensis3 32
percoeoum3 45
subterranea3 43
40
35
vermalis3 34
79
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foremanii (Cura), 19
(Curtisia), 19
(Dugesia), 19
(Planaria) , 19
fuliginosa (Anoaelis), 66
(Planaria)3 66
fuliginosus (Planaria), 66
(Typhlolepta), 66
Galeoaephala superba, 40
georgiana (Sphalloplana)3 47
glandulosa (Maoroaotyla), 12, 57
gonoeephala (Planaria)3 19, 20
gonooephaloides (Dugesia), 20
(Planaria)3 20
graailis (Euplanaria), 43, 44
(Fontiaola)3 43
(Phagooata)3 43, 44
(Planaria), 43, 44
graoilis (Fontioola), 43
graailis (Phagooata), 43
monopharyngea (Fontiaola)3 33
monopharyngea (Phagoaata)3 33
woodworthi (Fontioola)3 44
woodworthi (Phagooata)3 44
graffi (Dendrocoelum)3 63
hoffmasteri (Speophila), 53
(Sphalloplana), 53
hubriehti (Speophila)3 52
(Sphalloplana)3 52
Hydrolimax bruneus3 67
hymanae (Dendroooelopsis)3 61
Hymanella3 13
retenuova, 31
kansensis (Sphalloplana)3 49
Kerikia3 13
vhynohida3 56
kutsoheri (Sphalloplana)3 55
laateim (DendvoGoelum)3 63
Zata (Dugesia)3 20
(Euplanaria), 20
(Planaria)3 20
lugubri-s (Dugesia)3 24
(Planaria)3 19, 24
MaoToootyla, 13
glandulosa., 12, 57
maoulata (Dugesia), 20
(Euplanaria)3 20
(Planaria)3 20
miarobursalis (Dugesia)3 20
(Euplanaria)3 20
modesta (Dugesia)3 19
mohri (Sphalloplana)3 55
monopharyngea (Phagooata), 33
(Phagooata graeilis)3 33
morgani (Fontioola)3 40
(Phagooata)3 40
(Planaria)3 40
morgani (Fontioola)3 40
movgani (Phagoaata)3 40
polyoelis (Fontioola)3 41
polyoelis (Phagooata)3 41
musculosa (Planaria daotyligera)3 29
nivea (Fontioola)3 42
(Phagooata), 42
nivea (Fontioola), 42
nivea (Phagooata), 42
tahoena (Fontioola), 38
tahoena (Phagooata), 38
novangliae (Euplanaria), 20
oooulta (Planavia), 30
Oligooelis puloherrima, 63
oregonensis (Fontioola), 32
(Phagooata), 32
percoeca (Sphalloplana), 45
pevcoeoim (Dendroooelum), 45
(Fontiaola), 45
Phagooata, 13
bulbosa, 36
bursapevforata, 37
cavernioola, 40
coronataj 25
avenophila, 39
graoilis, 43, 44
gvacilis gvaailis, 43
gvaoilis monopharyngea, 33
gfaoilis woodworthi, 44
monophavyngea, 33
morgani, 40
morgani morgani,40
morgani polyoelis, 41
nivea, 42
nivea nivea, 42
nivea tahoena, 38
oregonensis, 32
su&terranea, 43
tahoena, 38
velata, 35
vemalis, 34
woodworthi, 44
philadelphioa (Euplanaria), 22
piriformis (Dendroooelopsis), 59
Planaria, 13
80
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Planaria agilis, 22
albissima, 40
daotyligera328
daotyligera daotyligera, 28
daotyligera musoulosa, 29
dovotoeeghalaj 22
foremanii, 19
fuliginosa, 66
fuliginosus, 66
gonooephala, 19, 20
gonooephaloides, 20
graoilis, 43, 44
lota, 20
lugubris, 19, 24
maaulata, 20
morgani, 40
oooulta> 30
polyohroa, 24
simplex, 66
simplioissima, 19
simplissima, 19
tigrina, 20
trunoata, 40
unionioola, 66
velata, 35
polyoelis (Fontioola morgani), '41
(Phagooata morgani), 41
Polyaelis, 13
borealis, 27
corona ta., 26
ooronata borealis, 27
eoronata brevipenis, 26
coronata eoTonata, 25
polyahcoa (Dugesia)3 24
(Planaria)3 24
priaei (Speophila)3 50
(Sphalloplana), 50
Proaotyla3 14
fluviatilis3 2, 63
typhlops3 63, 64
pulaherrima (Oligooelis)3 63
pulcherrimum (Dendrocoelwn)3 63
Reotooephala.3 13
exotica, 65
veddelli (Sphalloplana)3 55
retenuova (Hymanella)3 31
rhynahida (Kenkia)3 56
simplex (Planavia), 66
simplioissima (Curtisia)3 19
(Planaria)3 19
simplissima (Planaria)3 19
sloani (Sphalloplana) 3 55
Sovooelis amevioana3 62
Speophila buchanani, 51
- hoffmasteri3 53
- hvl>viohti3 52
iaei3 50
Sphalloplana3 13
- alabamensis 3 46
- buohananij 51
Sphalloplana geofgiana3 47
- hoffmasteri, 53
- hubriohti, 52
- kansensisj 49
- kutschevi, 55
j 55
j 45
priee3 50
reddelH3 55
sloani, 55
vivginiana, 48
weingavtnevi , 54
zesehit 55
subterranea (Fontieola), 43
- (Phagooata), 43
supevba (Galeooephala) 3 40
superbum (Dendroaoelwi)3 40j 63
tahoena (Phagocata)3 38
- (Phagoaata nivea), 38
tigrina (Dugesia), 20
- (Euplanaria) 3 20
- (Planaria), 20
trunaata (Fontioola), 40
- (Planaria), 40
truncation (Dendroeoelum), 40
Typhlolepta fuliginosus 3 66
typhlops (Procotyla), 63, 64
unioniaola (Planaria) , 66
vaginata (Dendrocoelopsis) , 58, 59
velata (Fontioola), 35
- (Phagooata), 35
- (Planar ia ) 3 35
vernalis (Fontioola) 3 34
- (Phagooata), 34
virginiana (Sphalloplana), 48
weingartneri (Sphalloplana), 54
woodworthi (Fontioola graoilis), 44
- (Phagooata) 3 44
- (Phagooata graoilis )3 44
zesohi (Sphalloplana), 55
81
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4. Title Biota of Freshwater Ecosystems Identification
Manual No. 1 FRESHWATER PLANARIANS (TURBELLARIANS) OF NORTH
AMERICA,
SELECTED WATER
RESOURCES ABSTRACTS
INPUT TRANSACTION FORM
1. Report No.
3. Accession No.
w
7. Author(s)
Kenk, R
9. Organization
Smithsonian Institution
Washington, D. C.
5. Report Date
8. Per for wing Organization
Report No.
10. Project No.
18050 ELD
//. Contract I Grant No.
14-12-894
}£. Type of Report and
Period Covered
12. Sponsoring Organization
15. Supplementary Notes
IS. Abstract
A key is presented for the identification of the species of North American
freshwater triclads or planarians known at present. Introductory chapters
deal with the collecting, culturing, preservation, study, and general organisation
and life cycle of planarians. The key is followed by a listing of the species
and subspecies, giving their distinguishing characteristics, ecological requirements,
and geographical ranges. Illustrations depict the external appearance and diagrams
of the reproductive organs of the individual taxa. The principal literature for
each species is indicated and listed in the appended bibliography of 65 items. An
index of the generic and specific names and synonyms concludes the report. One
new subspecies, Polycelis ooronota brevipenis is established for L. H. Hyman's
Polyeelis coronata.
I7a.Descriptors *Aquatic fauna, Life cycles, Preservation, Distribution.
I7b. identifiers *i(jentification manual, *Illustrated key, *Freshwater planarians,
*Triclads, *Turbellaria, *North America, Species list, Collection, Culture,
Anatomy, Distinguishing characteristics, Ecological requirements.
17c. COWRR Field & Group
10A
IS. Availability
19. Security Class.
(Report)
20. Security Class.
(Page)
21. »o.«f
Pages
22. Price
Send To:
WATER RESOURCES SCIENTIFIC INFORMATION CENTER
US DEPARTMENT OF THE INTERIOR
WASHINGTON. D C 20240
Abstractor Roman Kenk
institution Smithsonian Institution
WRSIC 102 (REV. JUNE 1971)
ft U S. GOVERNMENT PRINTING OFFICE : 1972 O - 467-101
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