Bijdragen tot de Dierkunde, 49 (1): 153-161 1979

New freshwater triclads from Tasmania

(Platyhelminthes, Turbellaria)

by

Ian R. Ball & Tran Thi Vinh-Hao

Department of Biology, University of Waterloo, Ontario, Canada

and

Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands

Abstract characters that usually require the preparation of serial

in often entire sections are given parentheses; however, Three new freshwater triclads are described from Tasmanian cleared specimens will suffice for the of lakes. Two of first records of recognition diag- these represent the the genus nostic characters. Spathula in Tasmania and the third belongs to the endemic

genus Romankenkius. The taxonomic affinities of the species

are discussed and keys to all the Tasmanian genera and SYSTEMATIC DESCRIPTIONS

These be species are provided. keys are to regarded as Family DUGESIIDAE Ball, 1974a provisional since it is likely that there remain many planarian

species to be discovered and described from Tasmania. to the Tasmanian of freshwater Key genera plan-

arians INTRODUCTION

1. Anterior margin with two auricular streaks and four pairs three Only species of freshwater planarians, or of sensory fossae; (testes reduced to one or two pairs;

triclads have hitherto been described from Tas- male atrium with radial muscle plates; penial papilla

and Cura see 1974b) long slender) . . (1 species, Ball, mania (Ball, 1974b). Two of these belong to Anterior with two of margin one or pairs sensory fossae, the endemic Romankenkius the genus and third, and two ciliated pits (testes numerous and close-packed;

Cura is in stout 2 pinguis, a species widespread Australia, male atrium normal; penis a cone or cylinder) .

2. Anterior ramus of intestine without diverticula New Zealand, and New Caledonia. The absence preocellar caudal and shell (oviducts, with at most a tiny branch, of of the any representatives genus Spathula, the glands enter a posterior diverticulum of the bursal diverse most Notogaean genus, has been held to canal) Romankenkius of be Anterior ramus intestine with one or two pairs of a zoogeographical anomaly (Ball, 1977a) but preocellar diverticula (oviducts, with a long caudal it is here resolved by of two new descriptions branch, and shell glands enter the bursal canal direct- A species from the island. third new species of the ly) Spathula

endemic Romankenkius is also described. genus Genus Romankenkius Ball, 1974b to the Tasmanian and Keys genera species are

and facilitate further to the known of Romankenkius provided so as to encourage Key Species

work the fauna of this on interesting region. 1. Colour fawn with two dark longitudinal strips (testes

extend to the R. bilineatus But caution must be exercised in attempting to copulatory apparatus) . sp. nov. Uniformly and darkly pigmented (testes prepharyngeal) fit discovered newly specimens into these pro- 2 visional For old the keys. a group as as planarians 2. (Testes dorsal; penis with prostatic vesicle) kenki Ball, a total of six Tasmanian is R. 1974b species a meagre one (Testes ventral; penis without prostatic vesicle) . . . . and in all there probability are many more species R. pedderensis Ball, 1974b to be discovered.

Romankenkius bilineatus sp. nov. (figs. 1, 4, 5)

MATERIALS AND METHODS Type material. — Tasmanian Museum and Art Gallery, For the available Hobart. present study only preserved material was Holotype: sagittal sections on one slide (K847). and in all essentials the methods employed were those Paratypes: sagittal sections on three slides (K848), sagittal described in studies of Australian freshwater sections slide transverse sections previous plana- on one (K849), of an

rians refer to (Ball, 1977a). Locality references the Tas- immature specimen on one slide (K850), frontal sections of

manian Lands Series. In the immature Department Tasmap keys an specimen on one slide (K865). 154 I. R. BALL & V. H. T. TRAN - TRICLADS FROM TASMANIA

Figs. 1-3. External features of preserved specimens of three Tasmanian planarians: 1, Romankenkius bilineatus sp. nov.;

dittae Scale lines 2, Spathula sp. nov.; 3, Spathula ochyra sp. nov. 1 mm.

— Preserved The is The Description. specimens up to 5.5 X copulatory apparatus variable. penis

2.2 mm. In life the animal is fawn or light brown is always large, though only weakly muscular, and

is rather shallow. The with two longitudinal dark brown stripes that pass the bulb usually papilla

each side of the mid-line and unite be on posteriorly takes the form of a wide cylinder and may and anteriorly. In preserved specimens the ground suspended vertically in the atrium (K847, K849,

is white. be colour The head is pointed, without fig. 4) or more normally disposed (K848,

and and In the is tentacles, the body broad leaf-like with a fig. 5). former case there little space

rounded end. There is of the and the and posterior one pair eyes between penis pharyngeal pocket

close and the the set together distant from the anterior so the bursa lies displaced to right side of

each and margin, and one pair of ciliated pits copulatory apparatus. The vasa deferentia enter the

fossae. The anterior of the intestine bulb from to sensory ramus penis separately above, enlarging the vesicles the extends between eyes without forming pre- form false seminal at point of entry.

ocellar diverticula. The pharynx is large, its root There is no true seminal vesicle, or expanded

of the its bulbar and wide duct being at about 55% body length and tip cavity, the ejaculatory opens at 70-75%. centrally on the tip of the papilla. In two of the - 1979 155 BIJDRAGEN TOT DE DIERKUNDE, 49 (1)

there dilations DP also specimens are glandular of the grid réf. 951564). Fulton (in litt.) re-

proximal part of the ejaculatory duct (K848, ports this species from Great Lake and an out-

K849). All the epithelia of the male apparatus wardly similar species has been recorded by

wall atrium Leonard Timms from Lake and are nucleate. The posterior of the male & (1974) Sorrell

be this is furnished with especially tall epithetial cells, Lake Crescent to the south-east. It may that

is in Tasmania because have or cushion cells, as is the facing epithelium of the species widespread we

The received from C. Patterson and H. B. N. penis papilla. testes are very numerous, being reports

situated both dorsally and ventrally to the intestine Hynes of the occurrence of a distinctly striped

and extending from behind the ovaries to the level triclad in other Tasmanian waters. According to

of the copulatory apparatus, often in a rather Fulton this species is found at depths of 7-15

lateral show substrates position. They a tendency to fusion. meters on ranging from sand to clay

The bursa copulatrix lies to the right of the and silt. it lie between copulatory apparatus (fig. 4) or may

the penis and the pharynx (fig. 5) albeit still to

the right of the mid-line. The bursal canal is an

wide duct that into the side unusually opens right

the atrium Its of near to the gonopore. nucleate

epithelium is underlain by a thin layer of longi-

tudinal muscle fibres and a thin layer of circular

the has muscle fibres; ectal region an outer re-

inforcing layer of thin longitudinal fibres. Near

to its junction with the atrium the bursal canal

throws small off a diverticulum, either posteriorly

or obliquely to the left. This diverticulum, which

receives the separate openings of the unbranched

tall oviducts, has a nucleate epithelium, but the

surrounding musculature is much reduced. The

into and shell glands open this diverticulum also

into the neighbouring region of the bursal canal

proper.

The situated paired ovaries are ventrally just

the brain. there be behind Adpressed to these may

a small clump of darkly staining vitelline cells on

the antero-dorsal face of each. Otherwise the vitel-

laria are sparsely distributed throughout the body

length, principally in the ventral half of the body.

— distin- Diagnosis. The new species is easily

guished from the two other species of the genus

its by distinctive striped colour pattern and by the

extension of the testes to the level of the bursa.

— 4-5. Romankenkius bilineatus sections Distribution. Romankenkius bilineatus has been Figs. sp. now, sagittal of the copulatory viewed from the left side. 4, which is apparatus confirmed only from the type locality bc = bursa Holotype (K847); 5, Paratype (K848). copu- Arthurs Lake in Tasmania. The were specimens latrix; bd = bursal canal; di = diverticulum of bursal canal;

collected ed = = glands; = by W. Fulton, 4 April 1977 (East Lake, ejaculatory duct; eg eosinophil go gono-

= male atrium; od = left oviduct; = pore; ma pe penis; South Arthurs Lake, grid ref. DP 964511) and = = shell vd = deferens. 4 ph pharynx; sg glands; vas Fig. 22 1977 Lake, North Arthurs Lake, May (East has been reversed from the original slide. 156 I. R. BALL & V. H. T. TRAN - TRICLADS FROM TASMANIA

refers the be fixation artifacts. Our own Etymology. — The specific epithet to explained as

is definitive distinctive striped colour pattern of this species. material insufficient, however, for a solution to this puzzle and further analysis must be DISCUSSION OF THE GENUS left to Australian zoologists with access to live

ROMANKENKIUS material.

for two The genus Romankenkius was erected

Tasmanian R. kenki and R. dugesiids, pedderen- Genus Spathula Nurse, 1950 sis, characterized by the opening of the oviducts Key to the Tasmanian species of Spathula and the shell glands into a posterior diverticulum Anterior with of inter- 1. margin one pair sensory fossae; eyes of the bursal canal. This arrangement was dittae symmetrical; (ejaculatory duct normal) . S. sp. now manifestation of as one an evolutionary Anterior with of preted margin two pairs sensory fossae; eyes trend in the of the aquatic planarians to separation wide apart; (ejaculatory duct wide, irregular, and ever-

sible) S. ochyra sp. nov. copulatory and glandular functions of the female

genital canal, or vagina (Ball, 1974b, 1977c). dittae Spathula sp. nov. (figs. 2, 6) Romankenkius bilineatus seems to be an inter-

material. — of and Inverte- mediate form because the separation is not entirely Type Department Entomology brate Zoology, Royal Ontario Museum, Toronto, Canada. complete in that whereas most of the shell glands Holotype: sagittal sections on one slide (ROM Cl39a).

into the diverticulum some into the empty empty Paratype: sagittal sections on one slide (ROM Cl39b).

bursal canal itself. Tasmanian Museum and Art Gallery, Hobart. Paratypes:

three wholemounts on one slide (K845); sagittal sections on The new species differs from the other two one slide (K846). in species the genus not only in the lesser develop-

of ment the bursal canal diverticulum with its Description. — Maximum size 3.25 X 1.0 mm. reduced musculature, but also in the extension of Pigmented, dorsal surface a uniform light brown,

the testes the the ventral is beyond pharyngeal root to copu- surface somewhat paler. The head

and in of ductus latory apparatus the absence a rounded, the body broad with parallel margins, the

genito-intestinalis. The anatomy and histology of tail broad and blunt. There are two relatively large

the has been considered in detail the genus previously eyes set at a distance equal to that between

and in most other in- and the frontal and lateral There (Ball, 1974b) particulars, eyes margins.

the of the is ciliated and cluding general asymmetry copulatory one pair of pits one pair of sensory

the is in organs, new species agreement with that fossae. The anterior ramus of the intestine extends

One is between the and forms of analysis. exception particularly noteworthy eyes two pairs pre-

however. The of small of vitel- is presence a clump ocellar diverticula. The pharynx relatively large,

line cells the is in its the and its adpressed to ovary paralleled a root at 40% of body length tip at

primitive Victorian species, Eviella hynesae Ball, 75%; thus the tail of the worm is relatively short. 1977b. Other similarities between these two species The copulatory complex is distinctive. The penis

the broad leaf-like and are appearance the fusion, is large and consists of a well-developed, elongate partial in R. bilineatus and complete in E. hynesae, and muscular bulb from which the broad plug-like

the of testicular follicles. papilla extends into the genital atrium. The vasa

The variation in the copulatory apparatus of deferentia, showing little or no recurvature, enter

R. bilineatus is remarkable and bulb from the sides into (figs. 4, 5) quite the separately and open an it be that there may are in fact two closely related, expanded seminal vesicle of elongate shape that is

It is also with The yet sympatric, species. plausible to argue lined a tall nucleate epithelium. wide that the specimens with the vertically disposed ejaculatory duct travels straight through the penis

and bursa The penis laterally displaced (K847, K849) papilla and opens slightly subterminally. are more contracted than the specimen illustrated distal half of the penial papilla has reduced epi-

But in in 5. even the latter the bursa thelial musculature and in the it is fig. specimen holotype sepa-

and its canal lie the of the mid-line and rated to right from the proximal half by a lip or fold. The

are not convinced that all the differences muscles of we can mesenchymal the papilla are well BIJDRAGEN TOT DE DIERKUNDE, 49 (1) - 1979 157

dittae the of viewed left side, Fig. 6. Spathula sp. nov. Sagittal section through copulatory apparatus the holotype from the

bc = bursa bd = bursal ed = = = male od = left copulatrix; canal; ejaculatory duct; go gonopore; ma atrium; oviduct;

= = = = deferens. pe penis; sg shell glands; sv seminal vesicle; vd vas

and developed. The testes are very numerous, being tudinal muscles a thicker layer of prominent

packed closely together in the ventral regions of circular fibres. The musculature is most strongly

the body and distributed from just behind the developed in the medial regions of the canal.

ovaries to the level of the bursa. The paired ovaries are ventrally situated imme-

The bursa copulatrix is a small sac lying diately behind the brain. The oviducts arise from

the On the anterior face of each and then between the penis and pharyngeal pouch. ovary curve

both the right and the left side it is in close contact laterally around them before travelling ventrally

the tail animal. with diverticula of the intestine but a true genito- to of the However, as they pass

intestinal connectionwas not discerned. The bursal the bursal canal each throws off a short branch

canal arises rather the anterior face that into the canal before it into ventrally on opens just opens of the bursa, twists and ascends vertically, and the atrium. Because of the relatively posterior

of then curves over the copulatory apparatus as an position the copulatory apparatus the caudal

duct oviducts ever-widening that bends forward into the branches of the are relatively short. Shell

wall of the atrium above the into the bursal canal between the posterior just gono- glands empty

The of the and the atrium. pore. nucleate cuboidal epithelium oviductal openings The vitellaria

is bursal canal underlain by a thin layer of longi- lie dorsally to the intestine, are most extensive in 158 I. R. BALL & V. H. T. TRAN - TRICLADS FROM TASMANIA

of the and extends between the and forms of the middle regions body, are sparse eyes two pairs

and There few diverticula. The is about one anteriorly posteriorly. are a pre- preocellar pharynx

ovarian follicles. fifth of the body length, its root being a little less

than halfway down the body.

Diagnosis. — In general habitus, in its possession The copulatory complex is unusual in the of ventral testes distributed to the copulatory structure of the male apparatus. The penis is large and in the subterminal of the apparatus, opening and and consists of an elongate and muscular bulb dittae is most similar to ejaculatory duct, Spathula blunt that into the male a broad, papilla projects S. from Victoria 1977a: It foeni (Ball, 22). may which in the atrium. The vasa deferentia, pharyn- be from this most distinguished species easily by and with geal region are greatly enlarged packed the musculature of the bursal canal and strong the sperm, enter penis bulb separately as narrow by the The ventral of the plug-like penis. opening tubes with little or no recurvature. Within the bulb

bursal canal into the bursa is unusual as are the they unite to form a broad seminal vesicle. The diverticula anterior the preocellar of the ramus of duct from this vesicle to at ejaculatory runs open intestine, this latter feature being one shared only the tip of the penis papilla. It is the structure and with the next species. arrangement of this duct that make this species

unusual for it seems that the central core of the Distribution. — Known only from the type penis is eversible, although it should be borne in locality which is Lake St. Clair in Tasmania (grid mind that the following interpretation is based ref. DP 345370). The specimens were collected three mature March Dr. Prince and upon only fully specimens. on 18 1974 by G. forwarded

In the the central core of the Dr. D. A. of La holotype (fig. 7) to us by Hay Trobe University. penis is fully everted so as to form a flared exten-

— The is for out- sion of the The of the Etymology. specific epithet an copulatory organ. flaring

standing friend, Dr. Dittie Rebel of Amsterdam. extension is unequal so that there is an accentuated

lip dorsally and to the right, immediately beneath

which the duct This Spathula ochyra sp. Nov. (figs. 3, 7, 8, 9, 10) ejaculatory opens. terminal

of the duct is surrounded Type material. —Tasmanian Museum and Art Gallery, Ho- part by eosinophil gland bart. sections slides Holotype: sagittal on two (K866). cells. The ejaculatory duct and the large seminal

Paratypes: sagittal sections on three slides (K867), sagittal vesicle are lined with a tall nucleate epithelium sections of slide a specimen not fully mature on one (K868), underlain a of ar- frontal sections on one slide (K869), frontal sections of a by prominent layer irregularly specimen not fully mature on one slide (K870). In the last muscle fibres. In both ranged fully mature para- series the ribbons are arranged vertically, rather than hori- types the eversion is only partial and involves only zontally as is our usual custom with frontal sections. the dorsolateral part of the eversible structure.

— Preserved to 7 2 In these the seminal vesicle Description. specimens up X specimens appears as a mm. Dorsal surface dark-brown to black, pigment large cavity lined with a tall, irregular, and

This often blotchy, ventral surface paler. The head is nucleate epithelium. narrows to a short slim

There duct which almost rounded, the body broad. are two eyes set immediately widens once more wide the distance between them to form the duct The apart, exceeding ejaculatory proper. ejacula- the distance between each and its duct is wide but it is eye respective tory now very partially body margin. On the body margin just anterior occluded by a large flap from its dorsal (fig. 8)

lateral wall. The to the eyes on each side there is a ciliated pit as or right (fig. 9) principal passage found in all of the In of the duct is beneath and species genus. the present ejaculatory to the left

this is of this occlusion and before species, however, an unusually deep sensory entally, narrowing

and in one it forms tube to the seminal it receives organ specimen a long prior entering vesicle, running ventro-caudad almost to the plane of the the secretions of numerous eosinophil glands. It is

itself. Between the ciliated and the anterior that this is with the eye pits postulated region homologous of the there 'terminal' of the tip margin are sensory fossae, two on glandular part ejaculatory duct

it each side. The anterior ramus of the intestine as appears when fully everted (fig. 7). BIJDRAGEN TOT DE DIERKUNDE, 49 (l) - 1979 159

of viewed from Figs. 7-9. Spathula ochyra sp. nov. 7, sagittal section through the copulatory apparatus the holotype (K866)

the right side; 8, sagittal section through the penis of a paratype (K867) viewed from the right side; 9, horizontal section

from bc = bursa bd = bursal through the penis of a paratype (K869) viewed the dorsal aspect, copulatrix; canal;

= = = = = eversible of male od ed ejaculatory duct; eg eosinophil glands; ev part ejaculatory duct; ma atrium; oviduct;

= = = of = = shell = seminal pb penis bulb; pe penis; ph pore pharyngeal pocket; pp penis papilla; sg glands; sv vesicle;

vd = vas deferens.

follicles situated and the somewhat the left of the The testes are very numerous pharynx, to mid-

the line. The is duct that in the ventral region of body and distributed bursal canal a wide curves

the level the and the from just behind ovaries to the of over the copulatory apparatus opens into bursa side of above the copulatrix. right the atrium gonopore.

the atrium The bursa is a large sac lying between the penis Immediately prior to entering the 160 1. R. BALL & V. H. T. TRAN - TRICLADS FROM TASMANIA

be bursal canal receives the oviducts, and beneath Diagnosis. — Spat hula ochyra may readily

from all other of the these the openings of the shell glands. The canal distinguished species genus is lined with tall that is its structure. the a nucleate epithelium only by unique penial Externally new conspicuously ciliated entally. The epithelium is species resembles S. fontinalis from New Zealand,

and the other with of overlain by layers of longitudinal then strong only species two pairs sensory fossae. circular muscle fibres which may be reinforced The latter species, however, has a strong

muscle the canal that is ectally by a few longitudinal fibres. The muscula- sphincter on bursal lacking

is ventral side the ture thickest on the of the canal. in new species, and the position and distribu-

The bursa is itself largely overlain by a thin layer tion of the testes are quite different. The preocellar

of of encircling fibres extending the circular layer of diverticula the intestine are peculiar to the two the canal. One noteworthy feature of the holotype Tasmanian Spathula species described in this is the of in the bursal canal agglomeration sperm paper.

its with the The at junction bursa copulatrix. sperm mass is attached to the epithelium by the heads Distribution. — Known only from the type of which is Arthurs Lake in Tasmania the sperm (fig. 10). locality (grid

The imme- DP paired ovaries are ventrally situated ref. 922567). The specimens were collected

oviducts March and diately behind the brain and the follow in Cowpaddock Bay on 29 1977 22 the usual course to the copulatory apparatus. Here May 1977 by W. Fulton. The species was found each branches sending one branch caudad to the at shallow sites of 3-5 metres on a substrate of

and black posterior vitellaria, a dorsal branch curving coarse mud, with large amounts of Elodea

to into the bursal canal above the and other In this upwards open canadensis, some plants, present. zone of the shell glands. The vitellaria extend lake Spathula ochyra is less abundant than is Ro- throughout the body, with a few pre-ovarian folli- mankenkius bilineatus (Fulton, in litt.).

extensive in cles, and they are especially the two

Etymology. — The is from the fully mature paratypes. specific epithet

Greek, ochyros, meaning firm, stout, strong, and

allusion the of is an to both general appearance

and the the animal appearance of the male copu-

latory organ.

DISCUSSION OF THE GENUS SPATHULA

The erected genus Spathula originally was by

Nurse (1950) for two species from New Zealand

but it was not accepted by later specialists. Re-

cently, however, Ball (1977a) resurrected, revali-

and redefined the add six dated, genus so as to

and new species from New Zealand southeastern

Australia. The geographical distribution of the

held be it genus was to anomalous because was

in unrepresented Tasmania even though well

represented in the mountains of Victoria (Ball,

1977a: 33). With the discovery of the new species

from Lake St Clair and Arthurs Lake this bio-

geographical anomaly is removed.

Both of the new species are unusual for their

10. Spat hula ochyra nov. section the Fig. sp. Sagittal through possession of preocellar diverticula to the anterior

canal of the = circular bursa/bursal holotype. cm muscle; ramus of the intestine. This is a primitive feature ep = epithelium; lm = longitudinal muscle; lu = lumen of

within the as = a whole bursa; sm sperm mass. Haploneura (Meixner, BIJDRAGEN tot de dierkunde, 49 (l) - 1979 161

in marine between these will have be 1928), and whereas it is widespread the relationships genera to

species it is rare in the freshwater planarians. repeatedly re-assessed as new species are discovered A comparable situation within the Australian and described.

fauna is to be found only in Eviella hynesae, a ACKNOWLEDGEMENTS primitive retrobursal planarian from Victoria Dr. D. A. Hay (La Trobe University, Melbourne) and Mr. This is blind but the (Ball, 1977b). species W. Fulton (Inland Fisheries Commission, Hobart, Tasmania)

anterior extends far from the the based. ramus forwards provided specimens upon which this paper is

of Some the necessary histological preparation was carried out brain and ends in a series of small diverticula. in the Royal Ontario Museum, Toronto, Canada but most was The testes are also of comment. In both worthy performed by Greet Vooren in Amsterdam. To all the above

the Tasmanian the ventral testes form we our thanks. The species a express study was partly supported by Operating Grant A0016 of the National Research Council continuous zone from just behind the ovaries to of Canada. shared with the copulatory bursa, a feature S. foeni

from This of Victoria (Ball, 1977a: 29-30). type REFERENCES

distribution, found also in Romankenkius bili- BALL, I. R., 1974a. A contribution to the phylogeny and biogeography of the freshwater triclads (Platyhelmin- neatus, is also characteristic of some species of thes, Turbellaria). In: N. W. RISER & M. P. MORSE

a with in South Neppia, genus representatives eds., Biology of the Turbellaria: 339-401 (McGraw-Hill, America, South Africa, and New Zealand (Ball, New York).

1974b. A new of freshwater triclad from Tas- , genus 1974b). It is of interest, therefore, that our first with mania, reviews of the related genera Cura and tentative identification of S was that pathula ochyra Neppia (Turbellaria, Tricladida). Life Sei. Contr., R.

it to this austral The Ont. Mus., 99: 1-48. belonged genus. general 1977a. A monograph of the Spathula , (Platy- the genus appearance of copulatory complex conforms helminthes, Turbellaria, Tricladida). Aust. J. Zool., with that of some of in species Neppia especially Suppl. Ser., 47: 1-43.

the with a wide 1977b. A new and retrobursal from large penis, complex ejaculatory , primitive planarian Australian fresh (Platyhelminthes, Turbellaria, duct with waters and the wide bursal canal strong mus- Tricladida). Bijdr. Dierk., 47: 149-155. culature that diminishes However the entally. , 1977c. On the phylogenetic classification of aquatic

planarians. In: T. G. KARLING & presence of sensory fossae and of branched ovi- M. MEINANDER eds., The Alex Luther Centennial Turbellaria. Symposium on ducts preclude the specimens from belonging to Acta zool. Fenn., 154: 21-35.

our criteria and deter- Neppia by (Ball, 1974b) LEONARD, B. V. & B. V. TIMMS, 1974. The littoral rock mine their of the S fauna of lakes membership genus pathula (Ball, three highland in Tasmania. Papers

Procs. Roy. Soc. Tasmania, 108: 151-156. 1977a). The phylogenetic and biogeographical MEIXNER, J., 1928. Der Genitalapparat der Tricladen und relationships of austral planarians have been dis- seine Beziehungen zu ihrer allgemeinen Morphologie, cussed in at length a number of recent papers Phylogenie, Ökologie und Verbreitung. Z. Morphol.

Oekol. Tiere, 11: 570-612. (Ball, 1974a, 1974b, 1977a) and need not be NURSE, F. R., 1950. Freshwater triclads new to the fauna of repeated here. Nonetheless it is clear that the New Zealand. Trans. Roy. Soc. N.Z., 78: 410-417.

Received: 8 March 1979