Mmrm voiixrnon cowraoi luiyKiirJCH sEs^iEy • isoso EID02/72 'Bl(ym. OF FRESHWATER MciixnaJ. FRESHWATER nHlililK TlllBELLARIA) («i V ^»»,r: ,■ iiM»u..ui. uw^ IB, mmmM^m^m. mottcno)^ agemcy 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 Stocli Number 5501-0365 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. WATER POLLUTION CONTROL RESEARCH SERIES ITie Water Pollution Control Research Series describes the results and progress in the control and abatement of pollution in our Nation's waters. They provide a central source of information on the research, development, and demonstration activities in the water research program of the Environmental Protection Agency, through inhouse research and grants and contracts with Federal, State, and local agencies, research institutions, and industrial organizations. Inquiries pertaining to Water Pollution Control Research Reports should be directed to the Chief, Publications Branch (Water), Research Information Division, R&M, Environmental Protection Agency, Washington, DC 20460. 11 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 (ArguZus) , isopod crustaceans (Asellidae), decapod crayfish crustaceans (Astacidae) , leeches (Hirudinea) , polychaete worms (Polychaeta) , aquatic dryopoid beetles (Dryopoidea) , and freshwater clams (Sphaeriacea) . Ill 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, Polyaetis aovonata brevipen-is , is es- tablished for L. H. Hyman's Polyaetis ooronata. This report was submitted in fulfillment of Project #18050ELD, Contract #14-12-894 under the sponsorship of the Environmental Protection Agency, 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 Vll 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 Cvra fovemanii 19 8-10 Dugesia tigrina 20 11-12 Dugesia dorotoaephala 22 13,14 Dugesia antiltana 23 15,16 Dugesia polychora 24 17,18 Polyaelis oovonata aoronata 25 19 Polyaelis aoronata brevipenis new subspecies 26 20 Polyoetis aoronata borealis 27 21,22 Planaria daotyligeva daotytigera 28 23 Planaria daatytigera musaulosa 29 24,25 Planaria occulta 30 26,27 Eymanella retenuova 31 28,29 Phagooata oregonensis 32 30 Phagoaata monopharyngea 33 31,32 Phagooata vemalis 34 33,34 Phagooata velata 35 35,36 Phagooata bulbosa 36 37,38 Phagooata bursaperforata 37 39,40 Phagooata tahoena 38 41,42 Phagooata orenophila 39 43,44 Phagoaata morgani morgani 40 45 Phagoaata morgani polycelis 41 46,47 Phagooata nivea 42 48,49 Phagooata gracilis 43 50,51 Phagooata woodworthi 44 52,53 Sphalloplana percoeoa 45 54,55 Sphalloplana alabamensis 46 56,57 Sphalloplana georgiana 47 58,59 Sphalloplana virginiana 48 60 Sphalloplana kansensis 49 61,62 Sphalloplana priaei 50 63,64 Sphalloplana buohanani 51 65,66 Sphalloplana hubriahti 52 67,68 Sphalloplana hoffmasteri 53 69,70 Sphalloplana weingartneri 54 71,72 Sphalloplana mohri 55 73-75 Kenkia rhynahida 56 76,77 Macroootyla glandulosa 57 viii 78,79 Dendrocoelopsis vaginata 58 80,81 Dendrocoelopsis piriformis 59 82,83 Dendrocoelopsis alaskensis 60 84 Dendrocoelopsis hymanae 61 85-87 Dendrocoelopsis americana 62 88,89 Procotyla fluviatilis 63 90,91 Procotyla typhlops 64 92,93 Rectocephala exotica 65 94 Planaria simplex 66 95 Planaria unionicola 66 96 Hydrolimax bruneus 67 IX 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 conmion 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. 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. e.g. Proootyta fZuviatitis 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 {GammcccuSt Asellus^ Daphnia, 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. 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. GENERAL FEATURES Planarians or freshwater triclads are elongated, flattened Turbellariaj 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 Fig. 2. Digestive system of a planarlan. 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. 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. 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 (Fig. 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. 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 Cuva 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. 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 foremanii 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 . 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. Theie 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 Dendroaoelopsis vaginata and the pink Maoroaotyla 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""" 11- nii -|; -— ^^ epi A "-^ ^^ B 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: .. Polyoelis 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: Phagoaata Penis without bulb, papilla rudimentary: Hymanella 9 (1) Anterior end with deeply invaginated adhesive organ -.Maoroaoty la 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; Reotooephala 11 (10) Penis bulb rounded, containing seminal vesicle: Dencboooelopsis Penis bulb elongated, containing a prostatic vesicle (Fig. 89); ProGotyla 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 Axxzt: Dugesia dorotooephala (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 Cura 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 retenuova (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 PoZyaelis) 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 musaulosa (p. 29) External muscle layer of penis papilla not thickened, adeno- dactyl opens into atrium near gonopore: Planaria dactyligera dactyligera (p. 28) 15 13 (11) Penis papilla with terminal lobe, ejaculatory duct opening on ventral side of papilla .... Phagoaata crenophila (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: Hymanelta retenuovaip. 31) Penis well-developed, with bulb and papilla 15 15 (14) Ejaculatory duct with blind diverticulum: Phagoaata 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 Phagoaata vemalis (p . 34) 17 (16) Penis with round bulb and narrow ejaculatory duct: Phagoaata bulbosa (p. 36) Penis with small bulb and wide undivided penial lumen: Phagoaata monopharyngea (p. 33) 18 (10) Penis papilla long, conical, with fibrous layer below outer epithelium, penis lumen with two seminal vesicles: Phagoaata graailis (p. 43) Penis papilla short, truncate, with intermingled circular and longitudinal muscle fibers below outer epithelium, penis lumen not subdivided .... Phagoaata woodworthi (p. 44) 19 (9) Bursa duct with anterior glandular and posterior non-glan- dular sections 20 Bursa duct non-glandular throughout: Potyoelis aoronata boreatis (p. 27) 20 (19) Penis papilla as large as, or larger than, penis bulb: Polyaelis aoronata aoronata (p. 25) Penis papilla smaller than penis bulb: Polyaelis aoronata brevipenis (p. 26) 21 (8) Head with slender, pointed auricles: Phagoaata bur sap erf or at a (p. 37) Head without pointed auricles 22 22 (21) Polypharyngeal Phagoaata graailis (p. 43) Monopharyngeal 23 23 (22) Eyes normally two (smaller accessory eye spots may occur) . . 24 Eyes wanting Proaotyla typhlops (p. 64) Eyes many, in two longitudinal rows: Phagoaata morgani polyaelis (p. 41) 16 24 (23) Eyes separated by about 1/3 the width of the head, about equidistant from frontal and lateral margins: Dendroooelopsis alaskensis Cp- 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 oaaulta (p . 30) Anterior intestinal ramus starts behind level of eyes 26 26 (25) Opening of ejaculatory duct at tip of penis papilla: Fhagooata oregonensis (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 morgani (p. 40) Tip of penis papilla without muscular wart, testes extend to posterior end Phagoaata 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 rhynohida (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: Maovocotyla 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: Reatoaephala exotica (p. 65) Penis with normal bulb and papilla 33 33 (32) Testes ventral, to level of gonopore: Dendroooelopsis vaginata (p. 58) Testes dorsal, to posterior end: Dendroooelopsis piriformis (p. 59) 17 34 (31) Eyes wanting Dendroooelopsis hymanae (p. 61) Eyes two, or two groups of scattered eyes: Proaotyla fluviatilis (p- 63) Eyes multiple, arranged in two longitudinal rows: Dendroooelopsis cmerioana (p. 62) 35 (1) Penis reduced, without bulb and with rudimentary papilla: Eymanelta vetenuova (p. 31) Penis normal, with bulb and papilla 36 36 (35) Ventral surface unpigmented Phagooata tahoena (p. 38) Ventral surface pigmented Dugesia polyohroa (p. 24) / 18 SECTION IV SPECIES CHARACTERISTICS AND RANGES •t^ura foremanii (Girard, 1852) Synonyms: Dugesia foremanii Girard, 1852; Planaria foremanii: Stimpson, 1857; P. simplissima Curtis, 1900; P. simpliaissima: Morgan, 1904; P, lugubris: Morgan, 1901 (not 0. Schmidt, 1861); Curtisia simpliaissima: Graff, 1916; C. foremani: Kenk, 1930; IPlanaria gonoaephala: Pearl, 1903 (not Duges, 1830); IDugesia modesta Girard, 1893, sg ode vs vd Fig. 7 after Kenk (1935). 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 maculata Leidy, 1847; P. tigvina Girard, 1850; P. lata Sivickis, 1923; Dugesia maculata'. Girard, 1851; D. lata: Hyman, 1951; Euplanaria maculata: Kenk, 1930; E. lata: Kenk, 1930; E. tigrina: Kenk, 1935; E. novangliae Hyman, 1931; ?E , microbursalis Hyman, 1931; IDugesia microbupsalis : Hyman, 1939; ?D. gonooephaloides Girard, 1850; ?Planaria gonooephaloides : Stimpson, 1857; ?P. gonocephala: Woodworth, 1897 (not Duges , 1830). 8 vd de am ac gp Fig. 9 after Kenk (1935). 10 Fig. 10. Pigment patterns (after Hyman, 1939b). 20 A very conunon 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. dorotooephala. 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 dorotoaephata (Woodworth, 1897) Synonyms: Planaria dovotoaephala Woodworth, 1897; P. agilis Stringer, 1909; Euplanaria dorotooephala: Kenk, 1930; E. agilis: Kenk, 1930; E. philadelphioa Hyman, 1931; Dugesia agilis: Hyman, 1939; B. diabolis Hyman, 1956. 11 12 0.1 mm A polytypic species, in nature up to 30 mm 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 dorsolateral ly, bend down toward the axis of the papilla, and increase in diameter, each duct forming a seminal vesi- cle. Distal ly 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 ventral ly 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 -K 14 III 'I III .' I pb pp am ac ^p 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 dorsal ly 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 polyahroa (0. Schmidt, 1861) Synonyms: Planaria polyahroa 0 Schmidt, 1861; P. lugubris: Bbhmig, 1909; Dugesia lugubris: Ball, 1969 (in part). 15 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 Polyoelis aoronata aoronata (Girard, 1891) Phagocata aoronata Girard, 1891. 18 0.5 mm ode bd ph m b 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 aoronata brevipenis new subspecies Synonym: Polycelis ooronata: Hyman, 1931. 19 Fig. 19 after Kenk (1952). Externally indistinguishable from the subspecies aoronata, 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 Potyoelis ooronata boreatis Kenk, 1953 Synonym: Polyoelis horealis Kenk, 1953. 20 bd ode 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 ooronata 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 daatyligera daatyligera Kenk, 1935 / Synonym: Planaria daatyligera Kenk , 1935 21 m 22 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 Rymanella and some species of Phagooata. 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 anterolateral ly 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 Planaria daotyligera musoulosa Kenk, 1969 23 . Fig. 23 after Kenk (1969). Differs from the subspecies daotyligera 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 'Plccnaria oooulta Kenk, 1969 24 25 0.5 mm 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 Phagoaata morgani and the western P. oregonensis. However, in P. oaautta 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 Hymanella retenuova Castle, 1941 26 27 Fig. 27 after Hyman (1955), Mature animals 7-14 mm 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 Phagoaata oregonensis Hyman, 1963 Synonym: Fontioola oregonensis: Ball, 1969 28 rri 29 . O.Smm 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 -syes. Externally similar to the Alaskan P. nivea and the eastern P. morgani. Testes ventral, extend- ing posteriorly behind the copulatory complex (in P. morgani 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 Phagoaata monoTpharyngea Hyroan , 1945 Synonyms: Phagocata graoilis monopharyngea Hyman, 1945; Fontioola graoilis monopharyngea: Ball, 1969. vd bp 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 Phagoaata ■Qemalis Kenk, 1944 Synonym: Fpntiaola vermalis: Ball, 1969. 31 32 ij 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 CStringer, 1909) Synonyms: Planaria velata Stringer, 1909; Fontioola velata: Kenk, 1930, r\ r\ ^ ^ O i4 bp vs de od S9 bd vd gl I I / I am dp ac gp 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 mjirgins . 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 Hyman (1934), Kenk (1944). 35 Phagoaata bulbosa Kenk, 1970a 35 36 Fig. 36 after Kenk (1970a). Mature animals up to 12 mm long and 1 . 7 ram 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 daatyligera and Eymanella vetenuova. 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 Synonym: 37 phagooata bursaperforata Darlington, 1959 Fontioola bursaperforata: Ball, 1969, de bd sv I 1 / I , I I / I I od am at 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 Phagoaata tahoena Kawakatsu, 1968 Synonyms: Phagoaata nivea tahoena Kawalcatsu, 1968; Fontioola nivea tahoena- Ball, 1969. 39 bd ode V i sv vd pi am pp gl Fig. 40 after Kawakatsu (1968) A rather plump and thick planarian reaching over 12 ram 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 500m). Literature: Kawakatsu (1968), Kenk (1970a). 38 41 Phagoaata arenophila Carpenter, 1969a b 0.5 mm 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 C1969a) , Kenk (1970a). 39 Phagoaata morgani morgani (Stevens % Boring, 1906) Synonyms: Planaria trunaata Leidy, 1851; P. morgani Stevens § Boring, 1906; P. alhissima: Kepner 5 Rich, 1918 (not Vejdovsky, 1883); Dendroaoelvm trunaatum: Girard, 1893; Fontioola trnmaata: Hyman, 1931; F. morgani: Castle § Hyman, 1934; F. morgani morgani: Ball, 1969; Phagoaata morgani: Hyman, 1937; "^.Dendroooelum superbum Girard, 1850; ?Galeoaephala superba: Stimpson, 1857; ?Phagoaata aavemiaola Hyman, 1954; IFontioola oavemioola: Ball, 1969, 43 44 pp g de od am mp ode bd gp Fig. 44 after Kenk (1935). Average length o£ 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 Planaria oaoulta (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 Phagoaata morgani polyoelis (Kenk, 1955) Synonym: Fontioola morgani var. potycelis Kenk, 1935. 45 -ph 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 Phagooata nivea Kenk, 1953 Synonyms '.Phagooata nivea nivea Kawakatsu, 1968; Fontiaola nivea nivea: Ball, 1969. 46 47 ode bd vd pb Fig. 47 after Kenk (1953). gp Up to 8 mm long and 1.5 mm 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. mor- 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 Phagoaata graoilis (Haldeman, 1840) Synonyms: Plancwia gracilis Haldeman, 1840; Euplanaria graoilis: Kenk, 1930; Fontioola gracilis: Castle § Hyman, 1934; Phagocata gracilis graoilis Hyman, 1945; Fontioola gracilis gracilis: Ball, 1969; Phagooata subterranea Hyman, 1937; Fontioola siibterranea: Ball, 1969. 48 49 0.5 mm 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. woodworthi 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 Phagooata woodworthi Hyman, 1957a Synonyms: Planaria gracilis: Girard, 1850 (not Haldeman, 1840); Phago- oata gracilis: Girard, 1850 (and other authors before 1937); Euplanaria gracilis: Hyman, 1931 (in part); Phagooata gracilis woodworthi Hyman, 1951; Fontioola gracilis woodworthi: Ball, 1969. 50 51 0.5 mm bd ode sph b vd mp pi od gp Fig. 51 after Kenk (1970c). Adult specimens 15-30 mm long and 2t^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. gracilis. 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 anterolateral ly, 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. gracilis), Hyman (1937a), Kenk (1970c). 44 Sphalloplana peraoeaa (Packard, 1879) Synonyms: Dendroaoelum percoeaion Packard, 1879; Fontioola peraoeoum: Hyman, 1931. en (^ , t;*^ bp bd ode 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 atriiom. 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 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 atriiun 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 (1945). 46 Sphaltoptana georgiana Hyman, 1954 56 \i6 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 Sphalloplana virginiana Hyman, 1945 58 59 am pp ode 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 kansensis Hyman , 1945 b bd de pp am da gp ode Fig. 60 after Hyman (1945), modified. Preserved specimens 20 mm 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 SphaZloplana priaei (Hyman, 1937b) Synonym: Speophita priaei Hyman, 1937, 61 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 Sphalloplana buahanani (Hyman, 1937b) Synonym: Speophila buahanani Hyman, 1937. 63 r^ 64 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 hubviahti (Hyman, 1945) Synonym: Speophila hubviahti Hyman, 1945 65 rr-K 66 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 Sphalloptana hoffmasteri (Hyman, 1954) Synonym: Speophita hoffmasteri Hyman, 1954. 67 /dK, 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 Sphaltoptana weingartneri Kenk, 1970b 69 70 vd vs bp am pp gP ode Fig. 70 after Kenk (1970b), Mature specimens 6-9 ram long and 1.2-1.8 mm 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 kutsaheri Mitchell, 1968; S. sloani Mitchell, 1968; S. zesohi 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 anterolateral ly 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 rhynahida Hyman, 1937b 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 mm) 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 Maaroootyla glandulosa Hyman , 1956 77 pi 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 Dendroaoelopsis vaginata Hyman, 1935 m b vd vs bd de f am 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. piriformis 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 Dendpoaoelopsis piriformis Kenk, 1953 80 r^ 81 :^ bd ode gp 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). N 59 Df-ndroooelopsis alaskensis Kenk , 1955 82 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). y 60 DendroGoelopsis hymanae Kawakatsu, 1968 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 Dendroaoelopsis ameriaana (Hyman, 1939a) Synonym: Sorooelis amerioccna Hyman, 1939 85 ) ( 86 gp bd Fig. 87. Spiny wart of the penis sur- face, enlarged (after Hyman, 1939c). Up to 18 mm 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 unpigmented, 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 fluviatilis Leidy, 1857 Synonyms: Vendroooelwn superbum: Leidy, 1851 (not Girard, 1850); D. laateum: Woodworth, 1896 (not 0. F. Miiller, 1774); D. graffi Wilhelmi, 1909; ?D. pulaherrimum Girard, 1850; lOligooelis pulahervima: Stimpson, 1857. 88 89 0-5 mm vdc pp gp Mature animals 12-20 mm long and 2-5 ma. 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 sourthem 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 typhtops Kenk , 1935 90 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 Reotooe-phdla exotica Hyman, 1953b 93 ode bd vdc pb am gp Fig. 92 and 93 after Hyman (1953b). 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 Planccria 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 C1896a, 1896b). After Woodworth (1896b). Planaria fuliginosa Leidy, 1851 . Synonyms: Planccria (Typhlolepta? ) fuliginosus Leidy, 1851; Anoaelis fuliginosa: 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 unionioola Woodworth. 1897 . Described from one specimen which ~ ~~~~ ' apparently had been sketched in life by the collector and had been brownish red, mottled 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 shriveled, 2.8 mm long and 1 lected on the mantle of Unio River near Havana, Illinois. Woodworth (1897). contracted and 8 mm broad. Col- in the Illinois Literature: After Woodworth (1897). 66 Hydrolimax hruneus Girard, 1891 . Length 19 mm, width 3 mm. Head ob- Q, tusely truncate, with a neck constriction ^O behind it. Eyes two. Body dark brown, head somewhat lighter. Intestinal coecura 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 Phagoaata, or Hymanella^ 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 aa atrium of adenodactyl ac common atrium or antrum ad adenodactyl af female atrium or antrum ai anterior intestinal ramus am male atrium or antrum ao adhesive organ au auricle avs anterior seminal vesicle b copulatory bursa or bursa copulatrix bd bursal duct, canal, or stalk bg glandular portion of bursal canal bp penial bulb br brain or cerebral ganglion eg cyanophilic glands CO copulatory apparatus or complex de ejaculatory duct dp diverticulum of penial lumen eg eosinophilic glands epe external epithelium epi internal epithelium epr epithelium of prostate f fibrous layer gl glands gp gonopore or genital aperture i intestine or gut m mouth or oral aperture me external muscle layer mi internal muscle layer mp muscles of penis papilla n ventral nerve cord od oviduct or ovovitelline duct ode common oviduct or common ovovitelline duct ov ovary pa parenchyma pb penis bulb ph pharynx pi posterior intestinal ramus Pl penial lumen PP penis papilla pr prostate pvs posterior seminal vesicle r margin with large rhabdites sg shell glands sn snout sph sphincter sv spermiductal vesicle or false seminal vesicle t testis V vagina vd vas deferens or sperm duct vdc common vas deferens vi vitellaria or yolk glands vs 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 Carol>'n B, Cast, 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 Ball, I. R. (1969). Dugesia luguhris (Tricladida: Paludicola) , a Euro- pean Immigrant into North American Fresh Waters. Journal of the Fisheries Research Board of Canada, 26:221-228. Beauchamp, P. de (1931). Biospeologica. LVI. Campagne speologique de C. Bolivar et R. Jeannel dans I'Araerique du Nord (1928). 2. Tur- bellaries Triclades. Archives de Zoologie Experimentale et Generate, 71:317-331. (1932). Biospeologica. LVI ^should be LVIIIJ. Turbellaries, Hirudinees, Branchiobdellides , deuxieme serie. Archives de Zoo- logie Experimentale et Generate, 73:113-380, plates 6-8. Bohmig, L. (1909). Tricladida. In: Die Susswasser fauna Deutschtands, Heft 19:143-176. Braithwaite, L. F. (1962). 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(1923). Studies on the Physiology of Reconstitution in Planaria lata, with a Description of the Species. Biological Bulletin, 44:113-152. Stevens, N. M. (1904). On the Germ Cells and the Embryology of Planaria simplissima. Proceedings of the Academy of Natural Sciences of Philadelphia, 56:208-220, plates 13-16. Stevens, N. M. , and Boring, A. M. (1906). Planaria morgani n, sp. Proceedings of the Academy of Natural Sciences of Philadelphia, 58:7-9, plate 1. Stringer, C. E. (1909). Notes on Nebraska Turbellaria with Descriptions of Two New Species. Zoologisoher Anzeiger, 34:257-262. Woodworth, W. M. (1891). Contributions to the Morphology of the Tur- bellaria. I. On the Structure of Phagocata gracilis, Leidy. Bulletin of the Museum of Comparative Zoology at Harvard College, 21:1-44, plates 1-4. (1896a). Preliminary Report on Collections of Turbellaria from Lake St. Clair and Charlevoix, Michigan. Bulletin of the Michigan Fish Commission, 6:94-95. (1896b). 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Bulletin of the Museum of Comparative Zoology at Harvard College, 31:1-16, 1 plate. 77 SECTION VIII INDEX OF SCIENTIFIC NAMES agilis (Dugesia) , 22 (Euplanaria) J 22 (Plcmaria)j 22 atabamensis (Sphalloplana) ^ 46 alaskensis (Dendrocoelopsis) , 60 albissima (Planaria) ^ 40 ameriaana (DendroGoetopsis) ^ 62 (Sovoaetis) , 62 Anooetis fuliginosa, 66 antillana (Dugesia) , 23 boTealis (Polyaelis) ^ 27 (Polyaelis ooronata) , 27 bvevipenis (Polyaelis ooronata) , 26 bruneus ( Hydro Umax )j 67 buohanani (Speophila) , 51 (Sphalloplana) 3 51 bulbosa (Phagoaata) y 36 bursaperforata (Fontiaola) , 37 (Phagoaata) J 37 oavermiaola (Fontioola) , 40 (Phagoaata) , 40 aoronata (Phagoaata), 25 (Polyaelis ) J 26 borealis (Polyaelis) ^ 27 brevipenis (Polyaelis) ^ 26 aoronata (Polyaelis), 25 avenophila (Phagoaata) , 39 Citr-aj 13 foremaniiy 19 Curtisia foremani, 19 simplioissima, 19 daotyligeva (Planaria) , 28 dactz/ ligera ' (Planaria) , 28 rmsaulosa (Planaria) , 29 Dendroaoelopsisj 14 alaskensis , 60 ameriaana, 62 hymanae, 61 piriformis, 58,59 vaginata, 12, 58, 59 Dendroaoelum graffi, 63 laoteum, 63 peraoeaum, 45 pulaherrimum, 63 superbum, 40, 63 trunaatum, 40 diabolis (Dugesia) , 22 dorotoaephala (Dugesia), 22 (Euplanaria), 22 (Planaria), 22 Dugesia, 13 agilis, 22 antillana, 23 diabolis, 22 dorotoaephala, 22 foremanii, 19 gonoaephaloides , 20 lai:;a, 20 lugubris, 24 maaulata, 20 miarobursalis , 20 modes ta, 19 polyahroa, 24 tigvina, 20 Euplanaria agilis, 22 dorotoaephala, 22 graailis, 43, 44 Zataj 20 maaulata, 20 miarobursalis , 20 novangliae , 20 philadelphiaa, 22 tigrina, 20 exotiaa (Reatoaephala) , 65 fluviatilis (Pvoaotyla) , 63 Fontiaola bursaperforata, 37 aavemiaola, 40 graailis, 43 graailis graailis, 43 graailis monopharyngea, 33 graailis woodjorthi, 44 morgani, 40 morgani morgani, 40 morgani polyaelis, 41 nivea, 42 nivea nivea, 42 nivea tahoena, 38 ore^onensisj 32 peraoeaum, 45 subterranea, 43 trunaata, 40 velata, 35 vemalis, 34 79 foremanii (Cura), 19 (Curtisia) , 19 (Dugesia) 3 19 (Planaria) J 19 fuliginosa (AnoQelis), 66 (Planaria) , 66 fuliginosus (Planaria) , 66 (Typhlolepta) , 66 Galeooephala supevha^ 40 geovgiana (Sphalloplana) ^ 47 glandulosa (Maoroootyla) , 12, 57 gonooephala (Planaria) , 19, 20 gonoaephatoides (Dugesia)j 20 (Ptanaria) 3 20 gracilis ( Eupt anuria ) ^ 43, 44 (Fontiaola) , 43 (Phagooata) , 43, 44 (Planavia), 43, 44 graoilis (Fontiaola) , 43 gracilis (Phagocata) , 43 monopharyngea (Fonticola) ^ 33 monopharyngea (Phagooata) , 33 woodworthi (Fontioola) ^ 44 woodworthi (Phagooata) , 44 graffi (Dendroooelum) , 63 hoffmasteri (Speophila) , 53 (Sphalloplana) , 53 hubriohti (Speophila) , 52 (Sphalloplana) , 52 Hydrolimax bruneus, 67 hymanae (Dendroaoelopsis) , 61 Eymanella, 13 retenwoya, 31 kansensis (Sphalloplana) , 49 Kenkia^ 13 rhynohida, 56 kutscheri (Sphalloplana) ^ 55 lacteim (Dendtrocoelum) , 63 Zata (Dugesia) 3 20 (Euplanaria) 3 20 (Planaria) 3 20 lugubris (Dugesia), 24 (Planaria) , 19, 24 Macrocotyla, 13 glandulosa, 12, 57 maoulata (Dugesia) , 20 (Euplanaria), 20 (Planaria) , 20 miorobursalis (Dugesia), 20 (Euplanaria) i 20 modesta (Dugesia), 19 mohri (Sphalloplana) , 55 monopharyngea (Phagocata), 33 (Phagooata gracilis), 33 morgani (Fonticola) , 40 (Phagocata) , 40 (Planaria), 40 morgani (Fonticola) , 40 morgani (Phagocata) , 40 polyoelis (Fonticola) , 41 polycelis (Phagocata) , 41 musGulosa (Planaria dactyligera) , 29 nivea (Fonticola) , 42 (Phagocata) , 42 nivea (Fonticola) , 42 nivea (Phagocata) , 42 tahoena (Fonticola) , 38 tahoena (Phagocata) , 38 novangliae (Euplanaria) , 20 occulta (Planaria), 30 Oligooelis pulcherrima, 63 oregonensis (Fonticola) , 32 (Phagocata) , 32 percoeca (Sphalloplana) , 45 percoecum (Dendrocoelum) , 45 (Fonticola) , 45 Phagocata, 13 bulbosa, 36 bursaperforata, 37 oavemicola, 40 ooronata, 25 arenophila, 39 gracilis, 43, 44 gracilis gracilis, 43 gracilis monopharyngea, 33 gracilis woodoorthi, 44 monopharyngea, 33 morgani, 40 morgani morgani, ^0 morgani polycelis, 41 nivea, 42 nivea nivea, 42 nivea tahoena, 38 oregonensis, 32 subterranea, 43 tahoena, 38 velata, 35 vemalis, 34 woodworthi, 44 philadelphica (Euplanaria), 22 piriformis (Dendrocoelopsis), 59 Planaria, 13 80 Plccnaria agitis, 22 alhissima, 40 daotyligera, 28 daatyligera daatyligera^ 28 daatyligera musoulosa, 29 dorotoaephalat 22 foremanii, 19 fuliginosa, 66 fuliginosusy 66 gonoaephala, 19, 20 gonocephaloides, 20 graoilis, 43, 44 lataj 20 lugubrisj 19, 24 maculata, 20 morgani, 40 oaaultay 30 polyahroa^ 24 - simplex^ 66 simpliaissima^ 19 simplissima, 19 tigrina, 20 trunoata, 40 union-iooZa, 66 velata, 35 polyaelis (Fontioola morgani), "41 (Phagoaata movgani) , 41 Polyaelis J 13 borealis , 27 aovonata, 26 aoronata borealis , 27 aoronata brevipenis , 26 aoronata aoronata^ 25 polyahroa (Dugesia), 24 (Planaria)j 24 prioei (Speophila) , 50 (Sphalloplana) , 50 Proaotyla, 14 fluviatilis, 2, 63 typhlops, 63, 64 pulaherrima (Oligoaelis) , 63 puloherrimwn (Dendroaoelum) , 63 Reatoaephala, 13 exotiaa, 65 reddelli (Sphalloplana) ^ 55 retenuova (Hymanella), 31 rhynahida (Kenkia) , 56 simplex (Planaria) , 66 simpliaissima (Curtisia) , 19 (Planaria) J 19 simplissima (Planaria)j 19 sloani (Sphalloplana) , 55 Soroaelis ameriaana, 62 Speophila buahanani, 51 hoffmasteri, 53 hiibriahti, 52 prioei, 50 Sphalloplana, 13 alabamensis , 46 buahanani, 51 Sphalloplana georgiana, 47 hoffmasteri, 53 hubriahti, 52 kansensis , 49 kutsoheri, 55 mohriy 55 percoecaj 45 priaei, 50 reddelli, 55 sloani, 55 virginiana , 48 weingartneri, 54 zesahi, 55 subterranea (Fontioola) , 43 (Phagoaata) , 43 superba (Galeoaephdla) , 40 superbum (Dendroaoelum) , 40j 63 tahoena (Phagoaata) , 38 (Phagoaata nivea), 38 tigrina (Dugesia) , 20 (Euplanaria) , 20 (Planaria) , 20 trunaata (Fontioola) , 40 (Planaria) , 40 trunaatum (Dendroaoelim) , 40 Typhlolepta fuliginosus, 66 typhlops (Proaotyla), 63, 64 unioniaola (Planaria) , 66 vaginata (Dendroaoelopsis), 58, 59 velata (Fontioola) , 35 (Phagoaata) , 35 (Planaria) , 35 vemalis (Fontioola) , 34 (Phagoaata) , 34 virginiana (Sphalloplana) , 48 weingartneri (Sphalloplana) , 54 woodworthi (Fontioola graailis) , 44 (Phagoaata) , 44 (Phagoaata graailis), AA zesahi (Sphalloplana), 55 81 4. 7";f/egj^g^^ q£ Freshwater Ecosystems Identification Manual No. 1 FRESHWATER PLANARIANS (TURBELLARIANS) OF NORTH AMERICA^ SELECTED WATER RESOURCES ABSTRACTS INPUT TRANSACTION FORM ;. Report No. J. Accession No. w 7. Author(s) Kenk, R 9. Organization Smithsonian Institution Washington, D. C. 5. Report Date 6. 8. Performing Organization Report No. 10. Project No. 18050 ELD II. Contract/Grant No. 14-12-894 13. Type of Report and Period Covered 12. Sponsoring Organization IS. Supplementary Notes 16. 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, Polyaelis coronata brevipenis is established for L. H. Hyman's Polyaelis coronata. na. Descriptors *k(\uatic fauna. Life cycles, Preservation, Distribution. nb. Identifiers *itientif ication manual, *Illustrated key, *Freshwater planarians, *Triclads, *Turbellaria, *North America, Species list. Collection, Culture, Anatomy, Distinguishing characteristics. Ecological requirements. I7C.COWRR Field & Croup lOA IS. Availability 19. Security Class. (Report) 20. Security Class. (Page) 21. No. of Pages 22. Price Send To : WATER RESOURCES SCfENTIFIC INFORMATION CENTER US DEPARTMENT OF THE INTERIOR WASHINGTON. D C 20240 Abstractor Roman Kenk ^Institution Smithsonian Institution WRSIC102{REV JUNE 1971) o U. S. GOVERNMENT PRINTING OFFICE : 1972 O - 467-101