Bijdragen tot de Dierkunde, 54 (1): 101-126 — 1984
On the species of the Duges ia gonocephalagroup
(Platyhelminthes, Turbellaria, Tricladida) from Greece
by
Elizabeth J. de Vries
Institute of Taxonomie Zoology, University of Amsterdam,
P. 0. Box 20125, 1000 HC Amsterdam, The Netherlands
Summary The of groups or superspecies. status Dugesia
of and is the the genus, Dugesia gonocephala a species group, comprising gonocephala, type-species
which differ from each the best has numerous closely related species, supposedly one known, previously
other in and In this re-assessed Vries morphological karyological aspects. been (De & Ball, 1980). known and described ofthe paper presently species, group The family Dugesiidae, of which Dugesia is from the eastern Mediterranean region, are reviewed. the type-genus, was erected by Ball (1974a) who Hitherto the triclads ofthis region have been poorly known also made the first strict in with Yet there is phylogenetic analysis comparison their western counterparts. of the Within the it a wealth of species in the area. Six well-delimited new group. genus Dugesia as was
of the described and species Dugesia gonocephala group are then known, Ball recognized a number of
one further species is rescued from The wealth synonymy. units which he phylogenetic gave subgeneric of material available has also made possible an assessment rank. One of the largest of these was the of the taxonomic of characters con- validity many usually
sidered be of within this difficult nominate to importance group. subgenus, Dugesia, exemplified by Dugesia gonocephala and its allies.
Résumé In his phylogenetic revision, Ball (1974a)
that each of the de suggested subgenera recognized Dugesia gonocephala est un groupe d’espèces, contenant
distin- him was of rank but that nombreuses espèces étroitement apparentées, se by worthy generic
de l’autre au de vue et elevation guant l’une point morphologique should await a thorough analysis of
Dans le article les karyologique. présent on passe en revue each This he has for the group. done genera du décrites espèces groupe connues ou comme nouvelles, Neppia (Ball, 1974b) and Spathula (Ball, 1977). de la région est-méditerranéenne. Jusqu’à présent les Now research is being carried out in Amster- Triclades de cette région restaient mal connus en com- dam in the of the avec néanmoins hope paraison ceux occidentaux; cette région se elucidating phylogenetic
abondance On décrit six and caractérise par son d’espèces. status of, phylogenetic relationships within, espèces nouvelles bien délimitées du groupe Dugesia the subgenus Dugesia, especially the D.
gonocephala, et une autre espèce, considérée auparavant s.l. is gonocephala group. Dugesia gonocephala a ici validée. L’abondance du comme synonyme, est a number of matériel aussi créé la de vérifier la superspecies, comprising closely disponible a possibilité related which resemble each other in validité taxonomique de plusieurs caractères qui sont species ex-
dans ternal but differ from in généralement regardés comme importants ce groupe appearance, each other
difficile. internal and of their anatomy aspects karyology
within (Benazzi, 1955). The taxonomy this
group has proved to be complicated. This is INTRODUCTION because here with of we are dealing a group
This is of series towards related paper one a leading a phylogenetically closely species, of
revision of the sensu which some have evolved than phylogenetic genus Dugesia species more
stricto, the of the others. There are several that type-genus family Dugesiidae. species possess
Distributed throughout the Old World this markedly different features in their copulatory
has been divided into number of so that their identification genus a species apparatus, poses no
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whereas other difficulties, species are very welcomed of studying an extensive triclad col-
similar to each other, differing only in details, lection from the fresh waters of the Mediterra-
for the making it hard nonspecialist to nean region. The specimens were collected by
distinguish them. Dr. H. Malicky of the Biological Station of
Species of the gonocephala group occur Lunz, Austria, as a part of an ecological survey
the Old World. Several deal of the Mediterranean from throughout papers region. Apart a
with members of the from number of the group specific species already known, samples
contained several of the D. geographical areas. The Asiatic species were new species
discussed Ball and Kawakatsu from Greece the Greek by (1970) by et gonocephala group and
al. has In materialfrom several (1976). Lepori (1951) given an account islands. addition, new
of the Mediterranean known Greek which in western species at localities, was already present
that time, and Dahm (1971) has dealt with the the Zoölogisch Museum, Amsterdam, has been
African Vries Ball studied. The representatives. De & (1980), new species are here described,
their taxonomie assessed and and more recently De Vries (in prep.), have re- position the
D. from used assessed the status of gonocephala s. str. validity of some commonly taxonomie
European locations. characters discussed.
In order obtain the into to necessary insight Moreover, it is here felt that workers in this
the of variation within the will benefit from range morphological geographic area a complete
number of from different overview of the of the group, a large species species gonocephala group
studied. As result in this Therefore the geographical regions were a occurring area. descrip-
it became that the triclad of this study apparent tions of D. cretica and D. gonocephala s. str. (sensu
of of the Mediterranean Vries also included. In fauna a large area was De & Ball, 1980) are this
still unknown. It is that the Italian the of the entire true penin- way paper serves as a synopsis
in sula and the Italian islands are relatively well subgenus Greece.
analysed, due to the work of Benazzi and his
school (Benazzi, 1961), but the eastern
ABBREVIATIONS IN THE FIGURES Mediterranean, and in particular Greece and USED
the Greek "terra in- islands, are effectively ad adenodactyl mf muscular fibres cognita" as far as their triclad fauna is concern- af atrial fold mr muscular ridge ed. at atrium mz muscular zone
There are only two species of the subgenus be bursa copulatrix nz nuclear zone known from Dugesia Greece, viz. Dugesia sagitta bs bursal canal ov oviduct
atrium (Schmidt, 1861), from Corfu and Cephalonia, ca common pb penis bulb dp diaphragm pf penial fold and Dugesia cretica (Meixner, 1928) from Crete. duct ed ejaculatory pg penial glands Schmidt did not a sufficient Unfortunately, give fd fold pp penis papilla of D. and morphological description shell sagitta, go gonopore sg glands
this with seminal vesicle subsequently species was synonymized lu lumen sv
male atrium vd deferens D. gonocephala by Komarek (1925). There have ma vas also been reports on the karyology of two lucida sketches. Drawings were made from camera On from Dugesia species Corfu one of (Ball, 1979), each the bar 200 figure scale represents /tm. the species belonging to the subgenus Schmidtea, and the other to the subgenus Dugesia. Further-
more, there are some karyological data known MATERIALS AND METHODS
from D. cretica (see Dahm, 1958; Benazzi &
For sectioned Benazzi-Lentati, 1976). Apart from these data, morphological study specimens were serially at 8 intervals, stained by Mallory-Heidenhain, from the Greek /im the Dugesia species region are Phosphotungstic acid haematoxylin, Masson's trichome or unknown. and mounted DePeX. each first Azan, in On slide the sec- For all these the reasons was tion is the when the label is the opportunity on upper left corner slide to
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and sections the ribbons right. For sagittal transverse were inner in the extra longitudinal layer outer mus- arranged vertically, and for frontal sections the ribbons cle zone. The bulk of the material has were arranged horizontally. The paired ovaries are ventrally located just been deposited in the Zoölogisch Museum of the Universi-
Additional material loaned behind the brain. the oviducts arise ty of Amsterdam (ZMA). was Typically
by the Museum für Naturkunde der Humboldt- from the dorsal side of the ovaries from where
Berlin Universitàt, (ZMB). caudad the of the they run ventrally to level
dorsad copulatory apparatus. Here they turn
and into the of the open separately vaginal area SYSTEMATIC SECTION bursal canal just above the shell glands. The
abundant vitellaria are distributed both dorsally All the species described in this belong to paper and ventrally throughout the body length, the D. and the gonocephala thus are closely group first follicles occurring just in front of the related. This that have several implies they ovaries.
features in These morphological common. and dor- The follicular testes are numerous
features best reference to are expressed by sal, extending throughout the body length from itself. In this Dugesia gonocephala way un- the a position just beside ovaries. avoided the necessary repetition is in species The bursa copulatrix is lined with a high descriptions, and in the latter the individual glandular epithelium. Stalked spermatophores
variations on the morphological "groundplan" often are to be seen in the bursa. The
will be given. To facilitate comparison a musculature of the bursal canal is of the re- character table is table I. provided as versed type, with inner, subepithelial,
longitudinal muscles overlain with circular
fibres. The epithelium of the bursal canal Dugesia gonocephala (Dugès, 1830) may 1) be either infranucleate or nucleate. Typically, (Fig. there of but not always, is an extra outer layer Planaria 1830: 83. gonocephala Dugès, fibres in the longitudinal vaginal area forming Dugesia gonocephala; Girard, 1850: 265. an ectal reinforcement of the bursal canal. This Euplanaria gonocephala; Hesse, 1897: 549. feature occurs in all the species to be described Dugesia (Dugesia) gonocephala; Ball, 1974a: 376. below.
— animals 18 The bulb embed- Description. Living up to mm penis comprises a muscular
long and 3 mm broad. Head of low triangular ded in the atrial wall and a free intromittent
with each of Characteristic is the of form two eyes consisting a papilla. diaphragm,
multicellular varied size and which the in- pigment cup containing many shape, separates
retinal cells. Supernumerary eyes, one or two trabulbar seminal vesicle from the ejaculatory
The colour of be divided on either side, sometimes occur. duct. The genital atrium may or un-
the dorsal surface is shade of divided. usually some grey
the in or brown, pigment varying intensity;
the — sometimes the animals appear black to na- Remarks. Traditionally any freshwater
is somewhat the above has ked eye. The ventral surface paler planarian matching description
than the dorsal surface. The been D. De marginally placed assigned to gonocephala sensu lato.
auricular streaks, just behind the eyes, are free Vries & Ball (1980) and De Vries (in prep.),
of pigment. from a detailed review of the old literature and
the material The inner musculature of pharynx com- from studies of existing and new from
inner circular and the demonstrated prises separate longitudinal original localities, have that
muscle the earlier of layers. Typically outer pharyngeal concepts D. gonocephala s. str. are
musculature also consists of two muscle layers, misleading, and thus its geographical distribu-
an inner circular and outer tion is difficult to These authors have an longitudinal one, assess.
but in members of the there is an a new characterization of the some group given Dugesia
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reconstruction ofthe Fig. 1. Dugesia gonocephalas. str. (Dugès, 1830), diagrammatic sagittal copulatory apparatus (ZMA
V.Pl.617.1).
of in its restricted but this character is found to within gonocephala Dugès (1830) zone, vary
sense. populations (De Vries & Ball, 1980).
In Dugesia gonocephala s. str. (Dugès, 1830)
De Vries 1980 sensu & Ball, (fig. 1) the penis Dugesia sagitta (Schmidt, 1861)
bulb is muscular. The is of an 2 & very penis papilla (Figs. 9A)
elongate, conical shape, often protruding Planaria sagitta Schmidt, 1861: 13 the through gonopore. A large, funnel-shaped Planaria gonocephala; Komarek, 1925: 323 seminal vesicle diaphragm the from 1979: separates Dugesiagonocephala; Ball, 187; De Vries & Ball, 1980 the ejaculatory duct and this diaphragm often 349.
extends far into the penis papilla. At the base of Type-material. — Messonghi River, just west of the penis papilla there is a penial fold. The Messonghi, Corfu, Greece. Coll. I. R. Ball, 12 May 1978.
distinctness of this character on the depends Neotypes: four sets of sagittal sections (ZMA
of contraction relaxation of the frontal state or penis V.PI.624.1-4), one set of sections (ZMA
of sections papilla. However, the longitudinal muscle V.PI.624.5), one set transverse (ZMA V.PI.624.7), alcohol specimens (ZMA V.PI.624). fibres separating the penis papilla and the
fold penial are always distinguishable. Additional material. — Spring at Marbella Beach,
The atrium is divided into a male and a com- Corfu, Greece. Coll. I. R. Ball, 13 May 1978. Two atrium annular mon by a well-developed specimens in alcohol (ZMA V.P1.625). One set of sagittal muscular ridge in the atrial wall. This muscular sections (ZMA V.P1.625.1).
Stream is near Mesaria, Corfu, Greece, 39°44'N ridge always present and is an important 19°44'E. Coll. H. Malicky, 11 June 1977 and 1 May 1979 character for the identification of this species. (Korfu 1 + 2). Three sets of sagittal sections (ZMA The often pharynx possesses an extra V.PI.626.1 + 4; V.P1.627.1). One set of frontal sections muscle in the muscle of longitudinal layer outer (ZMA V.PI.626.2) and one set transverse sections
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V. Pl. in alcohol (ZMA 626.3), specimens (ZMA The the left of bursal canal runs slightly to V.PI.626). the copulatory complex before opening into the Corfu, Greece. Coll. J. Wilhelmi, June 1906. Nine atrium. The lumen of the bursal canal is lined specimens in alcohol. One set of sagittal sections in the with infranucleate The ectal Museum für Naturkunde, Berlin (ZMB Vermes 5385). an epithelium.
vaginal reinforcement extends about halfway to
The original type-material from Corfu could the bursa copulatrix.
not located. In this I be re-establishing species The penis bulb is weakly muscular, and
have selected the best materialfor the The preserved encloses a slightly lobed seminal vesicle.
and thus the is here vesicle lined with neotype type-locality lumenof the seminal is a glan-
restricted to the dular in Messonghi River, Corfu, epithelium; some specimens these
Greece. This is and a large, fast-flowing perma- granular glands have discharged their contents
nent river, where the planarians were ex- into the lumen of the seminal vesicle. The
when ceedingly abundant originally collected. rounded conical penis papilla protrudes into the
undivided atrium and is only weakly muscular.
The base of the penis papilla is surrounded by
— which Description. Living specimens up to an asymmetric penial fold, is always
10 and wide and olive- than mm long 3 mm grey- to more developed dorsally ventrally, or even brown in colour. totally absent from the ventral side. The
The ovaries are located ventrally in the area ejaculatory duct follows a central course of the seventh intestinal branch. its Many large through the penis papilla and opens at tip.
and meiotic The seminal vesicle is from the oocytes are present, phases are to separated be in number of duct valve-like seen a specimens. ejaculatory by a well-developed
The testes begin just in front of the ovaries. diaphragm.
The vasa deferentia form small extra-bulbar or A characteristic feature of the male
vesicles is its false seminal before entering the penis copulatory apparatus extremely glandular
The surround of the bulb to open into the seminal vesicle very close nature. penial glands part
to the diaphragm. seminal vesicle inside the penis bulb, in places
Fig. 2. Dugesia sagitta (Schmidt, 1861), diagrammatic sagittal reconstruction of the copulatory apparatus (ZMA V.Pl.624.1).
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penetrating into the surrounding tissue. From Dugesia cretica (Meixner, 1928)
the bulb the extend into the F penis glands penis ( ig. 3)
papilla where they narrow down to form a glan- Euplanaria cretica Meixner, 1928: 585. dular the duct. zone surrounding ejaculatory Euplanaria (Euplanaria) cretica; Kenk, 1930b: 292.
The whole zone is in me- glandular V-shaped Dugesia (Dugesia) cretica; Ball, 1974a: 376. dian sections. The glands along the ejaculatory
duct those in the — are more granular than Material. In stream near Kakopetros, Crete, Greece, diaphragm, often staining differently. 35°24'N 23°45'E. Coll. H. Malicky, 13 May 1979 sections slides (K.611). Sagittal on four (ZMA V.PI.648.1), specimens in alcohol (ZMA V.PI.648).
— the Karyology. Material from neotype- Brackish-water spring, Georgiupolis, Crete, Greece, has been Ball All locality analysed by (1979). 35°22'N 24°15'E. Coll. H. Malicky, 28 September 1972 the specimens were sexual diploids, (K. 237). One set of sagittal sections (ZMA V.PI.651.1)
and of 2n=2x=16, with only metacentric chromo- two sets frontal sections (ZMA V.PI.651.2-3), specimens in alcohol (ZMA V.PI.651). somes.
Stream near Topolia, Crete, Greece, 35°25'N
23°41'E. Coll. H. Malicky, 3 October 1972 (K. 258). One
Remarks. — Komarek declared the (1925) set of sagittal sections (ZMA V.PI.653.1), alcohol Planaria of Schmidt be with sagitta to conspecifïc specimens (ZMA V.PI.653). Streamlet 37°34'N D. gonocephala (Dugès) and this view has been near Karia, Tinos, Greece, 25°10'E. Coll. H. Malicky, 8 June 1979 (Tinos 4). Two accepted without exception (Kenk, 1974). D. sets of sagittal sections (ZMA V.PI.652.1-2), one set of differs from D. s. str. in its sagitta gonocephala frontal sections (ZMA V.PI.652.3), one set of transverse weak bulb, its valve-like extremely penial sections (ZMA V.PI.652.4), specimens in alcohol (ZMA and diaphragm, the cone of prominent glands V.PI.652). within the penis. Diagnostic features of D.
The type-material was collected by H.J. Wichmann in such the gonocephala s. str., as funnel-shaped May 1927 in a stream near Meskla, Theristo, on Crete, and annular diaphragm, elongate penis papilla, and is in the Greece, now collections of the U.S. National of ridge muscles in the atrial wall, are absent Museum, Washington D.C. from the Corfu material. Therefore, I consider
Schmidt's species to be valid.
The visibility of the characteristic penial Description. — The preserved animals reach glands depends on the staining, which in its a length of 8 mm and a width of 2 mm, a
the is found in animals turn is greatly influenced by type of fixative copulatory apparatus even used. all the which small Nevertheless, recently preserved are relatively (5 mm).
fixed in different ovaries normal in size and and material, fixatives, clearly The are shape
the which shows glandular areas are important positioned in the area of the fifth intestinal characteristics of the species. Even the material branch. The oviducts arise from the dorsal wall collected in 1906 by Wilhelmi, which proved to of the ovaries. hold the stain with difficulty, showed the The testes begin at the level of the ovaries
albeit less and The deferen- specific glandular areas, conspicuous- are normally developed. vasa ly. In this specimen from Wilhelmi's collection, tia form small extra-bulbar seminal vesicles the is because of before the bulb to into the penis very extended, probably entering penis open the fixative used, so that the penial fold is not seminal vesicle.
The bursal canal the left of the very distinct. runs to penis
Schmidt recorded D. also from and from into the (1861) sagitta opens a posterior position
but the the atrium in male- Cephalonia, present study strongly sug- atrium, thereby dividing a
that this record different and atrium. The lumen of the bursal gests concerns a a common
with nucleate species (D.( aenigma n. sp.). The name D. sagitta canal is lined a epithelium. In the is here reserved for the from the first of species vaginal area a well-developed layer mentioned locality, Corfu. longitudinal muscles forming an ectal reinforce-
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of the Fig. 3. Dugesia cretica (Meixner, 1928), diagrammatic sagittal reconstruction copulatory apparatus (ZMA
V.Pl.648.1).
be about double of viz. distinct of ment can seen, extending halfway to a layer muscles, a layer
the bursa copulatrix. inner longitudinal muscles, overlain by a thick
of circular muscles. A lumen is The penis consists of a muscular penis bulb outer layer pres-
small in the of the in enclosing a and lobed seminal vesicle, and ent posterior part adenodactyl;
this lumen be filled with a muscular conical penis papilla. The some specimens may
ejaculatory duct runs centrally through the secretion.
and funnel- penis papilla opens at its tip. A
the — There shaped diaphragm separates ejaculatory Karyology. are two different
duct from the seminal vesicle. Some penial chromosome numbers known of D. cretica:
glands are present. A large finger-like muscular Dahm (1958) has reported a chromosome
arises from the number of individual glandular organ or adenodactyl, 2n= 2x= 16 in one of D.
base of whereas Benazzi of the penis bulb ventrally the penis cretica, & Benazzi-Lentati
have recorded of papilla and protrudes into the atrium alongside (1976) a karyotype consisting
the all penis papilla. In the specimens studied, 16 bivalents and some univalents in different
the of the which attributed adenodactyl was positioned ventrally specimens they to D. cretica (cf.
In of the the of remarks D. penis. most specimens papilla on minotauros).
the adenodactyl extended somewhat to the left
of the whereas in it — The penis papilla, one specimen Remarks. first detailed morphological
found into atrium Kenk was to protrude the slightly to description was given by (1930a). The
the right of the penis papilla, so that the papilla material studied here conforms with Kenk's of the be of in but dif- adenodactyl seems to a structure description most aspects, is found to
limited The is it in that the atrium is flexibility. adenodactyl as large fer from here described as
and sometimes than the Kenk describes it as even larger penis being divided, whereas as un-
papilla, its tip often entering the vaginal area of divided. However, as will be pointed out below,
the bursal canal. The consists of the division of adenodactyl the atrium is a difficult diagnostic
glandular parenchymatic tissue surrounded by character, and has to be dealt with cautiously.
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records the of the The which Furthermore,'Kenk papilla ovaries, are positioned between
adenodactyl ventrally to the right of the penis the fourth and fifth intestinal branch, are large
the material normal papilla, whereas in the majority of though not hyperplastic; many oocytes
of observed. studied here, it is situated ventrally to the left were
the It that the of the The contain but penis papilla. seems papilla testes many spermatogonia
adenodactyl protrudes into the atrium in a also ripe spermatozoa. The vasa deferentia
form small false then slightly flexible way, although it invariably seminal vesicles and bend
the the before downwards from arises from penis bulb ventrally of penis up recurving to open a
papilla. dorsal aspect into the seminal vesicle.
D. cretica is known with The canal the left of certainty to occur on bursal runs slightly to
the the the Greek islands Crete and Tinos (De Vries, in copulatory apparatus and opens into
of is divided into male and prep.). There are records this species from atrium, which a a com-
Rumania atrium. The bursal canal is lined with (Natasescu, 1972), Turkey (Kosswig mon a
Greece and Persia overlain distinct & Battalgil, 1942; 1943), (De nucleate epithelium, by a layer
Beauchamp, 1953) and Israel (Bromley, 1974), of longitudinal muscles surrounded by circular
muscles. In the but from the given descriptions it is doubtful vaginal area the bursal canal
whether these records refer D. cretica widens and becomes the to or to a very spacious, sur-
muscles closely related species. rounding circular are more developed
De Beauchamp (1936) recognized two Per- and some outer longitudinal fibres can be seen,
he third sian forms which thought attributable to D. extending to about one of the distance to
cretica , viz. B and C, A being the original species the bursa. Surrounding the musculature of the
from Crete. Livanov bursal canal there is distinct of nuclei. In (1951) however, has a zone demonstrated is in that form C, which does not one specimen a spermatophore present
is the bursa. possess an adenodactyl, a distinct species: D.
iranica which The consists of Livanov, 1951. Form B, possesses penis a muscular, rigid-
of variable size and without which an adenodactyl a looking, conical papilla and a penis bulb,
is different It is is formed thin of muscles surround- lumen, most likely a species. by a layer clear that these forms described from outside ing the seminal vesicle. The ejaculatory duct
Crete and Tinos are in need of taxonomie runs slightly ventrally through the penis
but its A valve- clarification. papilla, opens at tip. glandular
like diaphragm separates the ejaculatory duct
from the seminal vesicle. Dorsally and laterally
Dugesia malickyi sp. Nov. the left dorsal extension to is a of the penis
(Figs. 4 & 9B) papilla, which protrudes slightly into the
atrium. The tissue inside this structure or fold, Type-material. — Spring, Polydroson, Greece, is little that in the 38°36'N 22°34'E. Coll. H. Malicky, 30-31 May 1976 a more parenchymatic than
sections on five slides and (M. 23). Holotype: sagittal (ZMA penis papilla at the atrial side it is sur- V.P1.635.1). Paratypes: One set ofsagittal sections (ZMA rounded by a thin layer of muscular fibres, of V. PI.635.2), one set of frontal sections (ZMA V. PI.635.3), nuclei be few which the can clearly seen. A one set of transverse sections (ZMA V.P1.635.4), alcohol muscle fibres be from can seen running specimens (ZMA V.PI.635). sparse the bulb the penis to fold, no glands are present.
Etymology. — This species is named after The dorsal side of the penis papilla is more
muscular and than the ventral side. Dr. Hans Malicky, who collected most of the glandular material described in this paper.
Remarks. — Characteristic of this species is
— The Description. preserved specimens are the strong muscularity of the bursal canal,
a of to 13 in the area. In contrast the very large, reaching length up mm especially vaginal
width of to 3 and a up mm. male copulatory organ is but weakly muscular.
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4. reconstruction of the Fig. Dugesia malickyi sp. nov., diagrammatic sagittal copulatory apparatus (ZMA V.Pl.635.1).
5. damoae reconstruction of the Fig. Dugesia sp. nov., diagrammatic sagittal copulatory apparatus (ZMA V.Pl.630.1)
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The ventral ejaculatory duct is also unusual
within the group.
Dugesia damoae sp. nov.
(Figs. 5, 6 & 9C-D)
— Stream Type-material. near Mandates, Samos,
Greece, 37°47'N 26°49'E. Coll. H. Malicky, 27-29 May
1979 (Samos 8). Holotype: sagittal sections on five slides
(ZMA V.PI.630.1). Paratypes: one whole mount (ZMA
V.PI.630.2), two sets of sagittal sections (ZMA
V.PI.630.6-7), two sets of frontal sections (ZMA
of sections V.PI.630.4 & 630.8), one set transverse (ZMA
V.PI.630.5), specimens in alcohol (ZMA V.PI.630).
Etymology. — The species is named after the philosopher Damo from Samos, daughter of
Pythagoras and Theano.
D — The escription. animals are rather large, their length in preserved condition measuring
10 and their width 3 up to mm up to mm.
The ovaries normal in size and and are shape
between the third they are positioned ventrally and fifth branch of the intestines.
The testes begin in front of the ovaries. The vasa deferentia form large false seminal vesicles, before entering the penis bulb where
into the seminal vesicle. they open 6. damoae frontal Fig. Dugesia sp. nov., diagrammatic The bursa is thin copulatrix enveloped by a reconstruction of the copulatory apparatus (ZMA but distinct of muscles. In some layer oblique V.Pl.630.3). Bursal canal drawn slightly to the left for specimens a spermatophore is present in the reasons of clarity.
The canal the left bursa. bursal runs slightly to of the and rather copulatory apparatus opens
into the the dorsal atrial posteriorly atrium, thereby dividing is suspended on the wall, extending atrium into a male and a common atrium. The laterally to the left of the papilla and consisting lumen of the bursal canal is lined with an in- of tissue with parenchymatic some sparse franucleate in epithelium; not fully mature glands, surrounded by a sheet of longitudinal specimens however, the epithelium is nucleate. and circular muscles. The musculature of the
The musculature of the is well vaginal area penis bulb reaches to the dorsal side of the
and ectal reinforcement is developed an present adenodactyl. The whole aspect of the and extends about the is of halfway to bursa. copulatory apparatus one great musculari-
bulb The penis is formed by a distinct but ty- thin muscle layer enveloping the seminal vesi- cle. The conical is and ariadnae penis papilla large Dugesia sp. nov
The duct 8 muscular. ejaculatory runs centrally (Figs. 7, & 9E-F)
the its and is in through papilla, opens at tip Type-material. — Stream northeast of Koronis, places surrounded by glands. A valve-like, Naxos, Greece, 37°09'N 25°33'E. Coll. H. Malicky, 21
diaphragm separates the ejaculatory 1976 glandular May (Naxos 5). Holotype: sagittal sections on three from the seminal vesicle. duct An adenodactyl slides (ZMA V.PI.628.1). Paratypes: one whole mount
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7. ariadnae reconstruction of the Fig. Dugesia sp. nov., diagrammatic sagittal copulatory apparatus (ZMA V.PI.628.1).
V.Pl. two sets of sections (ZMA 628.4), sagittal (ZMA The bursal canal runs from the bursa to the
V.PI.628.5-6), one set of frontal sections (ZMA atrium, being situated slightly to the left of the V.PI.628.2), one set of transverse sections (ZMA penis, and the lumen is lined with an in- V.PI.628.3). franucleate epithelium in mature animals; in
Other material. — Stream above Koronis, Naxos, not fully mature animals it is nucleate. The in-
Greece, 37°08'N 25°32'E. Coll. H. 26-28 Oc- Malicky, ner longitudinal muscle layer of the bursal tober 1980 (Naxos 26). One set of sagittal sections (ZMA canal is very distinct. An ectal reinforcement is V. PI.629.1), oneset offrontal sections (ZMA V.PI.629.2), which about the present, extends halfway to alcohol specimens (ZMA V.PI.629). bursa.
The bulb consists of thin but distinct Etymology. — Named after the mythological penis a of muscles the and figure of Ariadna, daughter of king Minos from layer enveloping spacious
who behind Theseus folded seminal vesicle. The seminal vesicle is Crete, was left by on the isle of Naxos after having helped him kill the separated from the ejaculatory duct in the penis
Minotaur. papilla by a distinct valve-like diaphragm. The
duct the ejaculatory runs centrally through
Description. — The preserved specimens rounded conical penis papilla and it is in places reach a length of 8 mm and a width of 2 mm. very dilated, especially directly behind the
normal in size and and in The ovaries are shape diaphragm. There are some penial glands the situated ventrally between the fourth and fifth diaphragm and surrounding the ejaculatory intestinal branch. duct. The penis papilla protrudes somewhat
The testes begin at the level of the ovaries. obliquely into the atrium. Two adenodactyls,
The deferentia form each side of the vasa large extra-bulbar one at penis, are suspended seminal vesicles before entering the penis bulb from the dorsal atrial wall close to the base of where they open into the seminal vesicle. the penis papilla. Both adenodactyls consist of
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Dugesia aenigma sp. nov
(Figs. 10 & 13A-B)
Planaria 1861: sagitta Schmidt, 14 (in part; Cephalonia material).
Type-material. — Stream north of Charaktion,
Cephalonia, Greece, 38°10'N 20°41'E. Coll. H.
Malicky, 1 June 1977 (Kef. 2). Holotype: sagittal sections
on 3 slides (ZMA V.PI.642.1). Paratypes: one whole
mount (ZMA V.PI.624.7), four sets of sagittal sections
(ZMA V.PI.642.2-5), one set of horizontal sections on 3
slides (ZMA V.PI.624.6), specimens in alcohol (ZMA
V.PI.624).
Other material. — Kulurata, Cephalonia, Greece,
38°12'N 20°40'E. Coll. H. Malicky, 31 May 1977 and
27-28 September 1980 (Kef. 1). Five sets of sagittal sec-
tions (ZMA V.PI.643.1-5), specimens in alcohol (ZMA
V.PI.643).
Pastra, Cephalonia, Greece, 35°05'N 20°45'E. Coll.
H. Malicky, 27 September 1980 (Kef. 25). Two sets of
sagittal sections of not fully mature specimens (ZMA
V.PI.644.1-2), specimens in alcohol (ZMA V.PI.644).
Agrostoli, Cephalonia, Greece. Coll. J. Wilhelmi, June
1908. One set of sagittal sections (ZMB 5384), specimens
in alcohol (ZMB 5384).
— Dedicated friend Etymology. to a pictured
within.
8. ariadnae frontal Fig. Dugesia sp. nov., diagrammatic
— Preserved reconstruction of the Description. specimens reach a copulatory apparatus (ZMA V.Pl.628.2). Bursal canal drawn slightly to the left for length of 8 mm and a width of 2 mm. reasons of clarity. ovaries The are large but normal and posi-
tioned between the fifth and sixth intestinal
branch. parenchymatic tissue with some scattered The testes begin in front of the ovaries. The
surrounded form glands, by a muscular sheet. vasa deferentia small extra-bulbar seminal
The parenchymatic tissue of the adenodactyls vesicles before entering the penis bulb where
into the tissue. The into the seminal vesicle close the may penetrate surrounding they open to atrium is rather wide so that it can contain all diaphragm. these structures. The atrial wall is folded and The bursa copulatrix is large and often con-
Some of these atrial folds divide the tains the bursal canal muscular. a spermatophore, runs
into male and atrium. almost the atrium a a common medially over copulatory apparatus.
Bundles of muscle fibres in the In the of the are present majority specimens the bursal tissue surrounding the atrium. canal is lined with an infranucleate epithelium,
although in some specimens this epithelium is
— of In ectal Remarks. The morphology the nucleate. the vaginal area some rein-
is is forcement be in the copulatory apparatus complicated, which can seen vaginal area. The in caused the of adenodac- atrium is divided into male and part by presence two a a common tyls, but also by the asymmetrical setting of the atrium.
in the The of thin penis papilla atrium, all of which make it penis bulb consists a layer of difficult to interpret the serial sections. muscles surrounding the spacious and lobed
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ofsections the of: Fig. 9. Microphotographs through copulatory apparatus A, Dugesia sagitta (Schmidt, 1861), sagittal sec-
tion viewed from the left side the left side (ZMA V.Pl.624.1). B, Dugesia malickyi sp. nov., sagittal section viewed from
damoae viewed from the left side (ZMA V.Pl.635.1). C, Dugesia sp. nov., sagittal section (ZMA V.Pl.630.1). D, Dugesia
damoae section viewed from anterior E, ariadnae sp. nov., transverse to posterior (ZMA V.Pl.630.5). Dugesia sp. nov.,
section viewed from the left side ariadnae section viewed from sagittal (ZMA V.Pl.628.1). F, Dugesia sp. nov., transverse anterior All scale bars indicate 200 posterior to (ZMA V.Pl.628.3). μm.
seminal vesicle. The penis papilla hangs down The diaphragm is valve-like, there are excessive
in the atrium and is short, blunt and plug- penial glands in the diaphragm and the papilla,
and in the of the but do form distinct shaped, invaginations tip penis they not such a glandular
papilla are often to be seen. The ejaculatory zone as in D. sagitta. The dorsal side of the
which in is duct is places dilated, runs centrally papilla very glandular.
the and has terminal The of the through papilla a opening. morphology copulatory apparatus
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10. reconstruction Fig. Dugesia aenigma sp. nov., diagrammatic sagittal (ZMA V.Pl.642.1).
is complex because of the structure of the atrial Remarks. — Schmidt (1861) recorded D. wall. Close to the base of the papilla and begin- sagitta from Corfu and Cephalonia, but his
the lateral median is identificationof the from ning at level, there a specimens Cephalonia
The muscular and glandular ridge-like fold in the was only based on external appearance.
Schmidt atrialwall which protrudes into the atrium. The present survey suggests that was prob-
with The appearance of this fold varies considerably, ably dealing two separate species.
between the D. for the from even specimens belonging to same name sagitta is reserved species
In the fold the first mentioned viz. population. most specimens runs locality by Schmidt,
the left atrial wall the and the from has along dorsally over penis Corfu, species Cephalonia papilla to the right atrial wall, but it does not been given a new species name.
the ventral of extend over atrial wall. Dorsally
minotauros median of it is Nov. the axis the penis papilla not so Dugesia sp. well that it is obvious 11 & developed, so not on me- (Figs. 13C-D) dian sections. In some specimens the fold is on- Type-material. — Stream about 8 km south of on one side of the whereas in ly present penis, Rethimnon, Crete, Greece. Coll. J. P. Duffels, 25 May
other individual the muscular and one glan- 1982. Holotype: one set of sagittal sections (ZMA
of sections dular fold has invaginated into the atrial wall V.PI.636.1). Paratypes: one set sagittal (ZMA
V. one set of frontal sections (ZMA V.PL636.3), in PI.636.2), (fig. 13B). Nevertheless, a muscular area the of sections one set transverse (ZMA V.PI.636.4), dorsal side of the atrial wall close to the papilla specimens in alcohol (ZMA V.PI.636). is always visible, even in not fully mature
material. — specimens. Other Stream near Kaminaki, Crete,
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Greece, 35°09'N 25°27'E. Coll. H. Malicky, 12 May
1971 (K. 71), two sets of sagittal sections (ZMA
V.PI.637.1-2).
Stream near Agios Nicolaos, Crete, Greece, 35°12'N
24°28'E. Coll. H. Malicky, October 1971 (K. 282), one set of sagittal sections (ZMA V.PI.638.1), specimens in alcohol (ZMA V.P1.638).
Stream east of Agios Ioannis, Crete, Greece, 35°03'N
25°53'E. Coll. H. Malicky, 8 May 1979 (K. 603), one set
of sagittal sections (ZMA V.PI.639.1), specimens in
alcohol (ZMA V.PI.639).
Stream near Kalamafka, Crete, Greece, 35°04'N
25°40'E. Coll. H. Malicky, 23 September 1972 (K. 217),
of sections of immature one set sagittal specimen (ZMA
V.PI.640.1), specimens in alcohol (ZMA V.PI.640).
Stream near Arkadi, Crete, Greece, 35°18'N 24°37'E.
Coll. H. Malicky, 4 October 1972 (K. 268), one set of
sagittal sections (ZMA V.PI.641.1), specimens in alcohol
(ZMA V.PI.641).
Etymology. — Named after the minotaur
which from Crete, a mythological figure was
half bull, half human.
— in Description. The animals are large,
state lengths of about 12 preserved reaching 11. minotauros frontal Fig. Dugesia sp. nov., diagrammatic
their width to 3 mm, reaching up mm. reconstruction of the copulatory apparatus (ZMA
The ventral ovaries the Bursal canal drawn to the left for are very large, ma- V.Pl.636.3). slightly
of reasons clarity. jority of the animals studied possessed
hyperplastic ovaries with clusters of small
the frontal of the and the oocytes scattered through part diaphragm between the seminal vesicle
animal. and funnel- Normally developed oocytes were only ejaculatory duct is well developed
The rarely found. shaped. penis papilla is a muscular and
the The testes are normally developed with many elongate cone; ejaculatory duct is surround-
The deferentia form ed and ripe spermatozoa. vasa by many penis glands runs centrally
small extra-bulbar seminal vesicles before the to through papilla open terminally.
dorsal entering the penis bulb to open from a On the left of the base of the penis papilla,
into the seminal vesicle. and level the position at a slightly higher than seminal
In several which specimens a spermatophore was vesicle, there is a fold-like protuberance,
found in the bursa copulatrix. The bursal canal is both muscular and glandular. The
to musculature the atrial wall close runs to the left of the copulatory apparatus lining to the
into the undivided atrium. In "fold" is more than in other open most developed parts of
specimens the lumen of the bursal canal is lined the atrial wall. The centre of the "fold" consists with of tissue with muscular fibres a nucleate epithelium, but some specimens parenchymatic
infranucleate The ectal it and in the possess an epithelium. running through places paren-
reinforcement is almost completely confined to chymatic tissue from the "fold" penetrates the the vaginal area. surrounding tissue.
of muscular The penis consists a penis bulb,
seminal and Remarks. — The ovaries in this enclosing a spacious vesicle, a hyperplastic
it known muscular penis papilla which protrudes species are noteworthy since is that in somewhat obliquely into the atrium. The individuals with hyperplastic ovaries,
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reconstruction of the Fig. 12. Dugesia elegans sp. nov., diagrammatic sagittal copulatory apparatus (ZMA V.Pl.645.1).
polyploidy and aberrant forms of sexual May 1975 (R. 5). One set of sagittal sections (ZMA V.PI.646.1). reproduction may occur (Benazzi & Grasso,
well 1977; Bromley, 1977). Polyploidy as as
— The is derived recorded for D. cretica section Etymology. specific epithet diploidy are (cf.
from the Latin neat or of D. and there is distinct elegans meaning elegant on karyology cretica) a and refers to the slender and elegant external possibility that the polyploid individuals in- of the appearance species. vestigated (Benazzi & Benazzi-Lentati, 1976)
have here may belonged to D. minotauros, newly
— animals described. Description. The are sexually
mature at very small sizes: animals of 5 mm
length and 1.5 mm width were com- Dugesia elegans sp. nov. already Several animals (Figs. 12 & 13E-F) pletely sexually developed.
in the and were cultured laboratory they reach-
Type-material. — Stream in the Butterfly near Valley ed of 10 their width remain- a length up to mm, Petaludes, Rhodes, Greece. Coll. E. J. de Vries, 25 May ing 1.5 mm, giving them a very slender and 1982. Holotype: one set of sagittal sections (ZMA
appearance. V.PI.645.1). Paratypes: one whole mount (ZMA elegant and V.PI.645.10), four sets of sagittal sections (ZMA The ovaries are normal well developed
one set of horizontal sections V.PI.645.2-645.5), (ZMA and situated in the area of the fifth intestinal
V.PI.645.9) and two sets of transverse sections (ZMA meiotic be in ramus; phases are to seen many V.PI.645.7 & 8). specimens. The oviducts leave the ovaries from
of from — the lateral side instead the dor- Other material. 2 km southwest of Laerma, posterior
Rhodes, Greece, 36°08'N 27°55'E. Coll. H. Malicky, 6 sal wall, as in the other species.
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The testes begin behind the level of the penis is only weakly muscular and has its
the ovaries. The vasa deferentia form large extra- ejaculatory duct ventrally. Furthermore,
bulbar seminal before sizes of the differ vesicles, bending up to body two species significantly,
enter the where into the D. much smaller and thinner than penis bulb, they open elegans being
seminal vesicle. D. malickyi.
The bursa is and often contains compact a
spermatophore. The bursal canal runs to the DISCUSSION the left of copulatory apparatus and opens
somewhat laterally into the undivided atrium. The wealth of morphological data obtained here The lumen of the bursal canal is in most from the Greek species makes possible some animals lined with infranucleate an epithelium, the of general statements on taxonomie validity in although other specimens a nucleate several morphological characteristics from the In epithelium is observed too. the vaginal area D. gonocephala group as a whole and from the the bursal musculature is well very developed Mediterraneanrepresentatives in particular. As and some fibres of ectal reinforcement are pres- of of a consequence our increasing knowledge which confined the ent are mostly to vaginal D. gonocephala s.l. it is clear that the validity of area. several features, traditionally thought to be of The penis bulb is formed by a well-developed taxonomie importance, is questionable. muscle layer surrounding the spacious seminal
vesicle. The penis papilla is blunt and muscular Pharyngeal musculature and protrudes obliquely into the atrium, so that
the whole length of the ejaculatory duct is not A morphological feature which was commonly
in sections. The believed be of taxonomie value seen medial sagittal ejaculatory to is the extra
duct is central with terminal The muscle in the a opening. longitudinal layer outer
is Dorsal the muscle Kenk for diaphragm funnel-shaped. to penis pharyngeal zone. (1930a), ex-
it and somewhat to the left of is a blunt, ample, laid stress on this feature for
muscular projection into the atrium. The bulk distinguishing immature specimens of D. cretica
of this projection is formed by muscles and from D. gonocephala and he thought that it might
some glands, but parenchymatic tissue can be prove a good defining character for the latter
in into the Other authors seen too, which places penetrates species. frequently have referred
surrounding tissue. The muscles of the dorsal to the discriminatory value of this character
"fold" extend into the dorsal side of the papilla, even, as in some cases, when the collections
here the dorsal of the under giving part papilla a more study were entirely asexual (e.g. Marcus,
muscular than the ventral There aspect part. 1953, 1955; Dahm, 1967; Ball, 1970; Young &
are penial glands surrounding the diaphragm Young, 1974). On the other hand, De and the ejaculatory duct. Beauchamp (1936) has always stressed the
variability, and hence unreliability, of this
Karyology. — Material of D. elegans from the character, even within the single "species" D.
type-locality has been analysed karyologically gonocephala . and all the studied be specimens proved to sex- The evidence now favours De Beauchamp's
ual diploids, 2n=2x=16, all of the opinion. In their characterization of D.
chromosomes being metacentric. gonocephala s. str., De Vries & Ball (1980) noted
whereas that, this third muscle layer was usual-
Remarks. — characters in Although some are ly present the species, it could not always be
shared between D. and D. the demonstrated in individual of malickyi elegans, every a popula-
be the in it two species can distinguished by tion; some individuals, moreover, was so
and the of the diffuse be muscularity shape penis papilla as to demonstrable in only some of
in the latter species, whereas in D. malickyi the the sections. The statement by these authors
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of sections of: section Fig. 13. Microphotographs through the copulatory apparatus A, Dugesia aenigma sp. nov., sagittal viewed from the left side section viewed from the left side (ZMA V.Pl.642.1). B, Dugesia aenigma sp. nov., sagittal (ZMA
minotauros frontal sections viewed from dorsal ventral D, V.Pl.642.4). C, Dugesia sp. nov., to (ZMA V.Pl.636.3). Dugesia minotauros section viewed from the left side E, sec- sp. nov., sagittal (ZMA V.Pl.636.1). Dugesia elegans sp. nov., sagittal
viewed from the left side section viewed from tion (ZMA V.Pl.645.1). F, Dugesia elegans sp. nov., transverse posterior to indicate anterior (ZMA V.Pl.645.10). All scale bars 200 μm.
is absent been recorded from of D. that this layer in the species as known some populations
from Corfu be since these Kawakatsu A distinct may now ignored japonica (see et al., 1976).
referred D. has also found in specimens are herein to sagitta extra longitudinal layer been
(Schmidt). Several members of the D. two populations of D. iberica Gourbault &
from from Mallorca H. gonocephala group Africa are now known to Benazzi, 1979, (coll.
this character and it has it not recorded in the possess (Dahm, 1971) Malicky) although was
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TABLE I
within the Greek of the Distribution of character states species Dugesia gonocephala group.
stricto gonocephala sagitta cretica malickyi damoae ariadnae aenigma minotauros elegans
D. sensu D. D. D. D. D. D. D. D.
Character
Oviducts arise from ovaries:
— dorsally + + + + + + + + —
— — laterally — — — — — — — +
Bursal canal epithelium:
— nucleate + — + + + + + + +
— — infranucleate + + — + + + + +
Atrium :
— — — divided + — + + + + +
— — — — — — + + — undivided +
Penis bulb:
— — — — — — — — — muscular very +
— muscular — — + — + + — + +
— — — — weakly developed — + — + — +
Shape of penis papilla:
— — conical — — + + + — — —
— — — — — — — — rounded conical + +
— elongate conical + — — — — — — + —
— — — — — plug shaped — — — — +
— — — — — -— — — + — blunt
Shape of penis diaphragm:
— — — — — — funnel + + + +
— valve — + — + + + + — —
Adenodactyl: — — + — + + — — —
Fold:
— atrial — — — ? — — + ? ?
— — — — — ? penial + + ? ?
original description of other populations from Epithelium of the bursal canal
Mallorca (Gourbault & Benazzi, 1979).
Evidently the classificatory significance of The taxonomie importance of the nucleation of
this character needs It the the bursal canal has also now re-assessment. may epithelium lining
be stated in far of much discussion. however, that so as the Mediter- been the subject Marcus
ranean species are concerned the extra layer of (1953) suspected that this character might be
which is confirmed longitudinal muscles in the outer muscle zone of variable, by the findings of
the is absent with the of De Vries Ball in of pharynx exception & (1980) populations D.
some populations of D. iberica from Mallorca gonocephala s. str. from western Europe. The and possibly some individuals of D. ariadnae data derived here from the Greek species show
in the latter scattered the of the (from Naxos); case some unequivocally that nucleation
fibres in were to be seen only one individual. epithelium depends, as would be expected, on
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of of the the state development copulatory ap- The nature and origin of these "folds" is
D. and D. cretica often paratus. malickyi consistently unclear, especially if they are situated
nucleate in D. close possessed a epithelium, whilst to the base of the penis papilla. In princi-
infranucleate could be sagitta only an epithelium was ple such questions resolved by study-
but other the of present, in all the species type ing the morphological development of the
of maturi- epithelium depended upon the state copulatory apparatus during maturation. If the
while in the "fold" ty: fully mature specimens is present before the penis papilla has
of the bursal canal in- it is epithelium was developed, most likely to be a direct
in less it derivative of the atrial wall franucleate, mature specimens was rather than a penial
situations nucleate. Intermediate were observed fold (Ball, 1970). Unfortunately, however,
in of the some cases wherein the epithelium enough developmental stages are not always
vaginal region was infranucleate whereas closer available in existing collections, thereby
the bursa itself it nucleate. studies. In to was precluding developmental such cases
It follows from these observations that the the morphological and histological structure of
state of nucleation of the epithelium of the bur- the "fold" must be compared with that of the
sal canal is of taxonomie when and significance only atrial wall penis papilla, but even so in
assessed mature individuals. reliable conclusion be on fully many cases a cannot
drawn.
In two of the species described here, D. ariad- Adenodactyls and similar structures and D. nae damoae, the structures are highly dif-
the most difficult Probably morphological ferentiated so that it seems justified to classify features the which to interpret are structures are them as adenodactyls. D. ariadnae is easily iden-
in the present atrium together with the penis tified by its two adenodactyls. D. damoae
Often it is uncertain whether the ac- like papilla. possesses one large adenodactyl, D. etrusca
is companying structure an adenodactyl, a monoadenodactyla Lepori, 1947, but this species
fold atrial fold. penial or an differs from D. damoae in that the adenodactyl
differentiated arises from the ventral side of the Adenodactyls are highly penial base,
instead of from the dorsal atrial in of uncertain wall, as D. musculoglandular organs function,
it is assumed that have damoae. though generally they a stimulatory function during copulation. A In D. sagitta, the histology and position of the classic discussion of such has fold it be fold. organs been given clearly proves to a penial In D.
Kenk He three distinct atrial it by (1930b). recognized aenigma, the fold is obviously an fold as types, the hollow Planaria torva type, the solid is situated quite independently of the base of the
Polycelis tenuis type, and a complex hollow Den- penis. In D. minotauros, D. malickyi and D.
lacteum The drocoelum type. details of these need elegans however, "folds" are present that cannot
because for classified. muscular not concern us here the most part unambiguously be The
found in of the left of the "adenodactyls" some species protrusion laterally to the penis the D. do conform with in D. be direct gonocephala group not his papilla minotauros could equally a classification. Adenodactyls with a lumen, such derivative of the atrial wall or of the penis
found in D. since is rather as are bactriana De Beauchamp, papilla, the whole area muscular.
from and cretica The of the "fold" resembles the 1959, Afghanistan, D. (cf. structure one
Kenk, 1930a) are exceptional within the D. described for D. ilvana Lepori, 1948, from Elba,
For the the that in latter gonocephala group. most part except the species the two
of "adenodactyls" the Mediterranean species "adenodactyls" are found laterally on both
be folds or of varied sides of the whereas appear to protuberances penis papilla, in D. size, shape, position and structure, and the ter- minotauros the single "fold" is confined to the
of these has become left side of the In D. minology structures con- penis papilla. malickyi and fused. D. is fold dorsal In elegans there a to the penis.
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D. the tissue of the "fold" is like After studied of malickyi more having an extensive number
that of the penis papilla than that of the atrium. specimens of the Mediterranean D. gonocephala
In D. the dorsal "fold" consists of of the elegans a s.l., it can be concluded that the shape
muscular from which the muscles area, extend atrium is greatly influenced by the state of
into the dorsal side of the that relaxation of the penis papilla, so or contraction penis papilla, by
it is connected and almost the of fixed animals closely with, a part curvature the and even by
the of, latter. Developmental studies are the fixative used. Consequently, it is obvious
for final elucidation of the of that necessary a nature the division of the atrium into a male and a
the or in these three "folds", adenodactyls, common atrium is arbitrary in many species.
In the the criteria species. present paper following
from the mentioned the atrial Apart species already concerning morphology are used: if
there are two other Mediterranean the bursal canal in the roof species opens high of the
described D. that the atrium is united as having "adenodactyls" : etrusca atrium, so one cavity,
Benazzi, 1944, from northern has two via the in D. Italy, opening directly gonopore, as sagit-
dorsal D. minotauros and D. it is called large parenchymatic adenodactyls, one ta, elegans, un-
and the and one ventral, to penis papilla, D. divided. If the bursal canal runs to a position
benazzii from the of the Lepori, 1951, Tyrrhenian very posterior atrium, opening more or
has muscular Islands, a adenodactyl, the posi- less independently of the atrium into the
tion of which varies in (De Beauchamp, 1953). gonopore, as D. cretica, D. damoae, D. malickyi
In numbers and D. it some classificatory schemes the aenigma, is classified as divided.
and positions of the "adenodactyls" have In D. ariadnae the morphology of the
role played an important (Lepori, 1951; Dahm, copulatory apparatus is more complex. In this
1958). But as has been demonstrated above, the species the atrial wall forms muscular folds
of these is which in interpretation structures hazardous one place form a division of the
without studies. atrium. The ofthe division developmental appearance depends
on the contraction of the muscles and can
therefore between vary specimens. Morphology of the atrium the During present survey, comparative
of material of other Mediterranean has The anatomy the genital atrium, insofar as it species
be considered undivided divided been studied, others several may or into a among specimens
male and has also assumed of D. etrusca the a common part, monoadenodactyla. According to
taxonomie in studies of criteria the atrium of this is some importance past given here, species
the D. to be gonocephala group (Lepori, 1951; Dahm, regarded as divided, although Lepori
it 1958). However, in dealing with this (1947) describes as being undivided. This
feature needs characteristic within this has take re-assessment the en- group, one to throughout
into that tire D. and its account the division of the atrium is gonocephala group validity as a
taxonomie not as distinct as it is found to be in other character needs further discussion.
genera, such as Polycelis (cf. Ball & Reynoldson,
1981: figs. 19, 20, 21) and Bdellocephala (Von The penis Graff, 1912-1917).
It is that there In true are species of the contrast to the morphological features of
in which divided discussed gonocephala group a atrium is doubtful diagnostic value, as above,
in the unambiguously present, as D. gonocephala s. anatomy of the penis provides several
where annular muscular in the characters which do not much str., an ridge vary within the atrial wall divides the atrium (De Vries & Ball, species and are therefore of taxonomie validity.
1980). Yet the criteria by which this feature Even though the shape of the penis is inevitably
be ascertained have well de- its may never been influenced by state of contraction and relaxa-
fined for the members of the it is form differences group. tion, possible to recognize
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between the species. Thus, the penis papillae of Kawakatsu, 1972a, 1972b; Kawakatsu & Basil,
the Greek be classified into five Kawakatsu et The species may 1975; al., 1980). glandular
in I. the between the categories as shown table nature of diaphragm varies
The position of the ejaculatory duct is equally species: in D. sagitta the diaphragm is extremely
character since it does whereas in D. str. the a valid not seem to vary glandular, gonocephala s.
In D. in On the within the species. malickyi the ejaculatory glands the diaphragm are sparse.
duct somewhat the basis of these data it is that the runs ventrally through most likely
will have for papilla, whereas in all other species it follows a diaphragm important implications
central The duct in all of within course. opens terminally an analysis phylogenetic relationships
species. the gonocephala group.
The musculature of the penis bulb is here
found to be a consistent feature within the Oviducts
between it wide species, though species shows a In all but one of the species here described the of conditions within the D. range gonocephala oviducts arise from the dorsal ovarial wall. D. group. This range extends from the well- elegans is the exception and in this species they developed, muscular penis bulb as found in D. arise from the lateral posterior wall. The tax- gonocephala s. str. and D. benazzii (Lepori, 1951), onomie value of this finding is not known to the extremely weakly developed penis bulbs because usually it is not mentioned in species of D. sagitta and D. malickyi. In between there descriptions. From the comparative anatomical of of the are different degrees development undertaken this similar study during survey a musculature, ranging from a thin but compact condition, of oviducts arising from the dorsal muscle the seminal vesicle layer surrounding as ovarial wall, is known to occur in D. benazzii,D. in D. damoae, to the more scattered muscles of iberica and D. gonocephala s. str. Yet again we the penis bulb of D. ariadnae and D. aenigma. Of have a feature which needs re-assessment the Greek species described here the penis bulb the entire throughout group. of D. elegans is the most strongly developed.
According to Ball (1974a) the diaphragm
separating the seminal vesicle from the THE DUGESIA GONOCEPHALA GROUP IN ejaculatory duct is an evolutionarily derived, or GREECE apomorph, character shared by all the species of
the which he with the In the gonocephala group, equates eastern Mediterranean, specifically
subgenus Dugesia. A diaphragm is unrecorded Greece and the Greek islands, the Dugesia
in two oriental assumed to is only species belong gonocephala group now represented by eight
to the subgenus, viz. D. izuensis Kato, 1943 (see species, but undoubtedly more species remain
Kawakatsu, 1983) and D. annandalei Kaburaki, to be discovered. The type-species of the
of uncertain D. defined 1918, both species being taxonomie subgenus, gonocephala s. str., as by
The and size of the De status. shape diaphragm Vries & Ball (1980) has its major area of
between it be well distribution in but it has been may vary species; can western Europe,
and in D. recorded from hence it be developed, large funnel-shaped, as Bulgaria, may ex-
be discovered in Greece if gonocephala s. str. and D. benazzii, occasionally pected to carefully
extending far into the penis papilla, whereas in looked for. The geographic distribution of the
most of the Mediterranean species it is less Greek D. gonocephala species is presented in fig.
and like valve for in- 14. developed shaped a as,
stance, in D. sagitta. In the Asiatic and in the Table I gives the distribution of several
African species, the diaphragm is funnel- character states, thought to be of taxonomie im-
much smaller than in D. in the of D. shaped, though portance, Greek species the
and much gonocephala s. str., more posteriorly gonocephala group.
situated in the papilla (Ball, 1970; Dahm, 1971; The geographic distribution, as it is known at
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14. distribution of the Greek of the Fig. Geographic species Dugesia gonocephala group.
present, and the diagnostic characters of the D. sagitta (Schmidt, 1861)
summarized species, are below. Known only from Corfu. This species can be
distinguished from the other species of the
D. str. its characteristic in the gonocephala s. (Dugès, 1830) group by glandular zone
Mainly distributed throughout western Europe, penis, and by the asymmetric fold at the base of but also recorded from Bulgaria (De Vries, in the penis. It differs especially from D.
As De Vries Ball also in its lack of prep.). characterized by & gonocephala s. str. a true funnel-
this be its and (1980), species may recognized by shaped diaphragm, elongate papilla, an- well-developed and elongate conical penis nular muscle ridge of the atrial wall.
with papilla a large funnel-shaped diaphragm separating the seminal vesicle from the D. cretica (Meixner, 1928)
the muscular This ejaculatory duct, penis bulb, and species, as redescribed by Kenk (1930), is by the annular muscular ridge of the atrial wall known with certainty only from Crete, and also
atrium. from Tinos in D. giving a clearly divided (De Vries, prep.). cretica is
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easily identified by its large, muscular muscular fold laterally to the left of the penis
the adenodactyl situated ventrally from penis papilla at a level slightly higher than the
papilla. Typically, this adenodactyl possesses a seminal vesicle.
lumen in the posterior part.
D. elegans sp. nov.
D. Recorded from Rhodes. Characterized its malickyi sp. nov. by
Known only from Polydroson, Greece. This muscular penis bulb, the blunt penis papilla
has conical in which and the muscular dorsal fold. species a large penis papilla penial
the duct follows ventral This shares similarities with D. ejaculatory a slightly species many
differences in course, a dorsal extension of the penis papilla, malickyi. The principal reside
valve-like The bulb size is much and a diaphragm. penis and body (D. elegans smaller and slim-
and the of penis papilla are extremely weakly developed, mer), strong muscularity the penis
the which is bulb and dorsal fold of this in contrast to vaginal area, very penial species. (See
muscular. also D. malickyi.)
shares characteristics with D. malickyi many
D. elegans, but differs from it principally in penis ACKNOWLEDGEMENTS
of the shape and muscularity copulatory ap- Dr. Hans Malicky (Biological Station of Lunz, Austria), Furthermore, D. reaches paratus. malickyi Prof. Dr. Ian R. Ball (Institute of Taxonomie Zoology, sizes and has significantly larger a much University ofAmsterdam) and Dr. J. P. Duffels (ITZ) are
thanked for the material which this is plumper body shape than D. elegans. providing on study
based. The author is grateful to Dr. G. Hartwich of the
Museum für Naturkunde in Berlin for his cooperation in D. damoae nov. sp. of much Berlin and for the loan material in his care. I am
Recorded from Samos. This can be only species indebted to Prof. Ball for his critical comments and helpful from the other members of the distinguished discussions and to Dr. P. J. H. van Bree (ITZ) for his col-
legiality and advice. The photographs were made by Mr. group by the presence of one dorsal adenodac- L. der Laan the dorsal atrial wall. van (ITZ). tyl, suspended on
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Received: 13 April 1984
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