Content uploaded by Eduardo Mateos
Author content
All content in this area was uploaded by Eduardo Mateos
Content may be subject to copyright.
Accepted by W. Sterrer: 31 Jan. 2008; published: 2 Apr. 2008 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1739: 1–20 (2008)
www.mapress.com/zootaxa/
Terrestrial planarians (Platyhelminthes, Tricladida, Terricola) from the Iberian
Peninsula: new records and description of three new species
MIQUEL VILA-FARRÉ1,2, EDUARDO MATEOS3, RONALD SLUYS4 & RAFAEL ROMERO1,5
1Departament de Genètica, Facultat de Biologia, Universitat de Barcelona
2E-mail: mvilafarre@gmail.com
3Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona; E-mail: emateos@ub.edu
4Institute for Biodiversity and Ecosystem Dynamics & Zoological Museum, University of Amsterdam, P. O. Box 94766, 1090 GT
Amsterdam, The Netherlands. E-mail: sluys@science.uva.nl
5E-mail: rromero@ub.edu
Abstract
Little is known about the taxonomy and distribution of terrestrial planarians on the Iberian Peninsula. Few studies have
tried to investigate the local diversity of these animals, due to both their lack of economic interest and their low abun-
dance. In this study we have made extensive searches and collections of terrestrial planarians from the Iberian Peninsula,
thus gathering new information on their taxonomy and biogeography. The study includes the description of three new
species of the genus Microplana, viz. Microplana aixandrei sp. nov., Microplana grazalemica sp. nov., and Microplana
gadesensis sp. nov. We present distribution maps summarizing published and new records of land planarians. The present
work substantially increases our knowledge on this group of animals in Spain and Portugal and at the same time also evi-
dences the scarcity of data and studies on the biology of these organisms.
Key words: Platyhelminthes, Tricladida, Rhynchodemidae, Microplana, Rhynchodemus, Iberian Peninsula, Andalusia,
Grazalema, biogeography, taxonomy, spermatophore
Introduction
Although a considerable amount of taxonomic and ecological knowledge on several European faunal groups
is currently available, there are still some autochthonous elements of Europe that remain virtually unknown.
Such is the case for terrestrial planarians (Platyhelminthes, Tricladida, Terricola). Research efforts on some
relatively recently introduced terrestrial flatworm species has been considerable, due to their potential eco-
nomic impact (Boag & Yeates, 2001; Murchie et al., 2003), but resources for the study of autochthonous ter-
restrial planarians are virtually nonexistent. This apparent lack of interest of the scientific community results
from different factors. First, autochthonous terrestrial flatworms have no economic impact. Second, these spe-
cies are usually rare and hard to find, as is reflected in their taxonomic literature: 7 of the 19 known autochtho-
nous European species are described on the basis of a single specimen. In addition, some of this material was
lost, destroyed or is poorly preserved (Minelli, 1977). Finally, specimen identification requires an elaborate
process that involves the production of histological sections, which have to be examined under a compound
microscope. As a result, the general knowledge on this group of invertebrates in Europe is rather poor.
The Iberian Peninsula presents a low number of records for terrestrial planarians, as compared to other
groups of animals in this area or to terrestrial planarians in some other European regions (Sluys, 1999). Fur-
thermore, some of these planarian records correspond to unidentified species or to specimens with doubtful
identity (H. Jones, personal communication).
VILA-FARRÉ ET AL.
2 · Zootaxa 1739 © 2008
Magnolia Press
Simroth (1881) provided the two first records for Iberian terrestrial planarians, concerning two species
collected in the area of Oporto, Portugal. Von Graff (1899) considered one of these two species to be M.
pyrenaica (von Graff, 1899) (Jones, 1998). It was only after a long period of time that a third record was
reported for the Iberian Peninsula when Filella (1983) collected Bipalium kewense Moseley, 1878, an intro-
duced species, from the area of Barcelona. This worker failed in finding new animals in later visits (E. Filella,
personal communication). Mateos et al. (1998), described the new species M. nana Mateos et al., 1998 from
Les Alberes (Eastern Pyrenees) and reported the presence of another species, M. terrestris (Müller, 1774),
from the area of Barcelona. More recently, Boix and Sala (2001), collected Rhynchodemus sylvaticus Leidy,
1851 from the Estany d’Espolla (NE Spain) and Mateos et al. (2004) described the presence of two unidenti-
fied terricolans from three localities in the Montnegre-Corredor (NE Spain).
FIGURE 1. Distributional records of autochthonous terrestrial planarians in the Iberian Peninsula. A. Iberian Peninsula
excluding Catalonia. B. Catalonia. Numbers indicate the total number of localities in each area. Arrow points to the type
locality of the three new species.
Zootaxa 1739 © 2008 Magnolia Press · 3
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
In the present paper we describe three new species of the genus Microplana from Andalusia, in the most
southern part of Spain. Furthermore, we provide distribution maps and a table specifying 55 new localities
where terrestrial planarians were found. These data substantially increase our knowledge on the diversity of
the Iberian land planarians but at the same time reveal the scarcity of studies on this group of animals.
Material and methods
Between 1998 and 2006 different areas from the Iberian Peninsula were sampled for terrestrial planarians
(Fig. 1). Forested and potentially humid areas, usually close to rivers and streams, were selected in order to
maximize detection probability. Oak, beech, and gallery forests were the main prospected areas. Planarians
were collected with a fine paint brush from underneath stones and logs. Specimens were individually trans-
ported to the laboratory in little plastic containers with humid soil. Live specimens were photographed. For
histological studies specimens were killed (under field or laboratory conditions) using Steinmann’s fluid
(Sluys, 1989a) and were subsequently fixed in 70% ethanol. Fixed specimens were cleared in clove oil and
then embedded in paraffin, cut at intervals of 5–7 µm, and mounted on gelatine-coated slides. The sections
were stained in Mallory-Heidenhain (Sluys, 1989a) and mounted in DePeX. In addition to this routine over-
sight stain, several series of sections were stained with the alcian blue-periodic acid-Schiff (AB-PAS) tech-
nique to detect polysaccharides (Romeis, 1989). The identification of R. sylvaticus has not been based on
histological sections but on its special external morphology, typically showing two dark longitudinal stripes, a
medial black circular spot, and two large eyes situated a little distant from the anterior tip (Ball & Reynoldson,
1981). The material examined is deposited in the Centre de Recursos de Biodiversitat Animal, Facultat de
Biologia, Universitat de Barcelona.
Abbreviations used in figures
bl, bulbar lumen; br, brain; dep, dorsal epidermis; e, eye; ed, ejaculatory duct; fgd, female genital duct;
gid, genito-intestinal duct; gp, gonopore; h, head; i, intestine; cm, circular muscles; lm, longitudinal muscles;
mf, muscular fibres; od, oviduct; ov, ovary; pb, penis bulb; ph, pharynx; phc, pharyngeal cavity; pp, penis
papilla; sg, shell glands; sp, sperm; sph, sphincter; spt, spermatophore; t, tail; te, testis; va, vacuolated paren-
chymal area; vap, vacuolated epidermis; vd, vas deferens; vep, ventral epidermis; vnc, ventral nerve cord.
Systematic account
Order Tricladida Lang, 1884
Suborder Terricola Hallez, 1892
Family Rhynchodemidae von Graff, 1896
Subfamily Microplaninae Pantin, 1953
Genus Microplana Vedjovsky, 1890
Microplana aixandrei sp. nov.
Material. Holotype, CRBA435, Llano del Berral (lat. 36.75428, long. -5.45399; alt. approx. 657 m) in the
central sector of the Sierra de Grazalema, Cádiz (Spain), 5 December 2004, sagittal sections on 1 slide.
Paratypes: CRBA436, ibid., sagittal section on 1 slide; CRBA437, ibid., horizontal sections on 1 slide.
VILA-FARRÉ ET AL.
4 · Zootaxa 1739 © 2008
Magnolia Press
Diagnosis. Microplana aixandrei sp. nov. can be distinguished from its congeners by its small size (up to
10 mm long), cylindrical body tapering anteriorly to a blunt point, bluntly pointed tail, and hyaline body sur-
face. Regarding anatomical features, the species differs from its congeners in the following features: presence
of two ventral testes on either side of the body; spherical penis bulb provided with a strong musculature and a
distinct bulbar lumen; short and vertically oriented penis papilla; atrium divided in a cup-shaped cavity and a
tubular distal cavity; wide and obliquely orientated bursal canal with a sphincter at its proximal end; a copula-
tory bursa without genito-intestinal connections; use of spermatophore in the transfer of sperm.
Ecology and distribution. The species is known only from the type locality. In contrast to other Iberian
terrestrial planarians, M. aixandrei can be considered a relatively common species of the soil fauna at the type
locality. During mating the sperm is transferred to the copulatory bursa of the partner aggregated in a sper-
matophore.
Etymology. The specific epithet is based on the nickname of Vila’s grandfather, Miquel Farré Servent,
who lived in a house known as “casa l’Aixandre” in his hometown Salàs del Pallars.
Description. The living, sexually mature specimen measured 10 mm in length and about 0.5 mm in
width, in elongated state (Fig. 2A). The cylindrical body tapers anteriorly to a blunt point; tail also bluntly
pointed. The body surface is hyaline and therefore the species appears white in colour due to the content of the
intestine. The anterior and posterior ends, where the intestine is absent, are transparent. The ventral surface is
hyaline. The hyaline colouration and the tiny size of the preserved specimens prevented us to adequately
observe the creeping sole.
The two small eyes (eye cup diameter 11 µm in sections) are located at a short distance in front of the
brain (Fig. 2B). In living and preserved specimens the eyes are only clearly visible under observation through
a dissecting microscope.
The anterior body region is filled with vacuolated parenchymal tissue (Fig. 2B, C) that is reduced between
the eyes. This region with vacuolated tissue extends backwards to the level of the testes. The adjacent ventral
epidermis is also vacuolated and thick (about 25–30 µm in longitudinal sections while approximately 18 µm
in the adjacent non-vacuolated ventral and dorsal epidermis); this vacuolated part of the epidermis (Fig. 2C)
extends to almost the level of the ovaries.
The subepidermal longitudinal fibres of the body musculature are weak. In the ventral body region numer-
ous longitudinal fibres are distributed in two parenchymal bands, which are especially strong over and under
the ventral nerve cords. The scarce dorsal longitudinal parenchymal fibres are very weak and apparently dis-
continuous.
The cylindrical pharynx is about one-eighth of the body-length (0.3 mm) and is situated in the posterior
third of the animal, in an almost horizontal position. The outer epithelium, which is ciliated only at the poste-
rior part of the pharynx, is underlain by a layer of longitudinal muscles, followed by a layer of circular mus-
cles (Fig. 3A), intermingled with some additional longitudinal fibres (not represented in Fig. 3A). Close to the
dorsal insertion of the pharynx, the inner epithelium is underlain by a thin layer of circular muscles that
becomes thicker at the posterior end of the pharynx. At the tip of the pharynx this circular layer narrows again
before reaching the lumen. This circular layer is followed by a thick layer of longitudinal muscles. The mouth
is situated at the posterior portion of the pharyngeal pocket, close to the hind wall of the pharyngeal pouch. In
specimen CRBA435 the mouth is situated at 1.63 mm from the tip of the body and 0.15 mm from the gonop-
ore.
There are two pairs of ventrally located, oblong testes follicles, occupying about one-fourth of the dorso-
ventral diameter in the prepharyngeal part of the body. The testes are located in the posterior third of the ante-
rior body region. In specimen CRBA435 there is an additional immature third testis situated on one side of the
animal.
Zootaxa 1739 © 2008 Magnolia Press · 5
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
FIGURE 2. Microplana aixandrei. Holotype. CRBA435. A. Living animal. B. Sagittal section through the head. C. Sag-
ittal section through front end. D–E. Sagittal sections of the copulatory apparatus; anterior to the left. F. Sagittal recon-
struction of the copulatory apparatus; anterior to the left.
VILA-FARRÉ ET AL.
6 · Zootaxa 1739 © 2008
Magnolia Press
The strongly muscular, spherical penis bulb is covered with intermingled longitudinal and circular muscle
fibres and is provided with a well-developed bulbar lumen (Fig. 2D). This bulbar lumen tapers gradually to
form an ejaculatory duct that opens at the tip of the penis papilla. The bulbar lumen and ejaculatory duct are
lined with a nucleated epithelium that is underlain with a layer of circular muscle fibres. At the level of the
posterior section of the pharyngeal pocket the thin vasa deferentia (diameter about 6
μ
m at the mid-level of the
pharyngeal pocket) enlarge to form spermiducal vesicles, which narrow before entering the penis bulb. After
having penetrated the penis bulb, the vasa deferentia increase in diameter, subsequently opening separately
into the dorsal part of the seminal vesicle.
The vertically oriented penis papilla is short and conical. The papilla is covered with a thin, nucleated epi-
thelium that is underlain with a thick, subepithelial layer of circular muscle bound by a layer of longitudinal
fibres.
The atrium consists of a dorsal cup-shaped cavity and a distal tubular part. The lining epithelium of the
atrium is underlain with a subepithelial circular muscle layer, thickened at the posterior wall of the tubular
part, followed by a thin layer of longitudinal muscles.
The ovaries are situated immediately above the ventral nerve cords. They lie at about one-third of the dis-
tance between the anterior end of the body and the root of the pharynx, occupying about one–fifth of the
dorso-ventral diameter. The oviducts arise from the ventral side of the ovaries. In running backwards, the
ducts follow the course of the ventral nerve cords. Behind the gonopore the ducts turn dorsally and open sepa-
rately into the bursal canal.
The copulatory bursa is an irregular sac-shaped structure, lined with tall vacuolated cells (Fig. 2E). Sev-
eral muscle fibres traverse in dorso-ventral direction the parenchyma between the posterior wall of the copula-
tory bursa and the adjacent intestinal branch (Fig. 2F). The copulatory bursa is not connected with the gut.
All three of the animals examined had in their copulatory bursa remnants of an irregular structure (37x21
µm in specimen CRBA435) formed by a blue homogeneous substance partially enveloped by a thin, brown
layer, most likely of sclerotic nature (Fig. 2E). We have not found the origin of this substance in penial or
atrial glands, but the colour and texture resemble the wall of a cocoon capsule. The location and nature of this
structure suggest that it forms part of a spermatophore.
The wide bursal canal, which receives the opening of the shell glands at the same level as it receives the
oviducts, is lined with nucleated cells. The distal section of the oviducts, just before communicating with the
bursal canal, also receives the secretion of the shell glands. The bursal canal is surrounded by a subepithelial
layer of circular muscles and some scattered longitudinal muscles fibres. A sphincter, consisting of circular
muscle fibres, surrounds the proximal section of the bursal canal (Fig. 2F).
Discussion. Among the approximately 19 species of native land planarians known from Europe, three of
which are newly reported in the present paper, M. aixandrei stands apart from the other species by a combina-
tion of external features and the anatomy of its genital apparatus.
A hyaline body colouration is also found in M humicola Vedjovsky, 1890. However, this species shows a
greenish anterior end, in contrast to M. aixandrei. Regarding anatomical features, M. humicola has dorsal tes-
tes and a genito-intestinal duct, while a copulatory bursa is absent (Schneider, 1935). In contrast, M. aixandrei
presents ventral testes and a copulatory bursa, but lacks a genito-intestinal duct.
A copulatory bursa that is devoid of any connection with the intestine occurs also in M. mahnerti Minelli,
1977, M. styriaca (Freisling, 1935), and M. grazalemica sp. nov. described below.
Microplana mahnerti shows about twenty testes on each side of the body and an elongated penis papilla.
In contrast, M. aixandrei shows two testes on each side of the body and a short and conical penis papilla. In M.
mahnerti the bursal canal runs from the wall of the atrium parallel to the body surface and the oviducts open at
its distal section. A sphincter is absent (Minelli, 1977). In contrast, in M. aixandrei the bursal canal is a
obliquely running structure that receives the oviducts at its central region, while a sphincter is present in the
proximal section of the canal. With respect to external features, M. mahnerti shows a grey colouration and big
Zootaxa 1739 © 2008 Magnolia Press · 7
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
size (13 mm in preserved specimens), contrasting with the hyaline colouration and reduced size of M. aixan-
drei (about 4 mm in preserved specimens).
In M. styriaca (Freisling, 1935) an expanded bulbar lumen is absent, while the penis bulb and papilla are
elongated. In contrast, in M. aixandrei a distinct bulbar lumen is present, while the penis bulb is rounded and
the penis papilla is short and conical. In M. styriaca the oviducts form a short common oviduct before they
enter the bursal canal (Freisling, 1935), contrasting with the oviducts in M. aixandrei that open separately into
the bursal canal. Furthermore, M styriaca is dark-brown, whereas M. aixandrei is hyaline.
The new species M. grazalemica sp. nov. is not hyaline, contrasting with the hyaline colouration of M.
aixandrei. With respect to anatomical features, M. grazalemica possesses about 15 testes on each side of the
body, in contrast to the two pairs present in M. aixandrei. The presence of a spermatophore in the copulatory
bursa of the three specimens of M. aixandrei suggests that the animals are adult worms and not juveniles of M.
grazalemica sp. nov. The use of spermatophores represents a unique feature of this species, not previously
recorded in any species of the Microplaninae. A vacuolated parenchymal tissue in the anterior part of the body
is absent in M. grazalemica but present in M. aixandrei.
There are various explanations for the vacuolation of the epidermis and parenchyma observed in M. aix-
andrei: vacuolation as an artefact of tissue fixation, processing and subsequent histological preparative treat-
ment, apparent vacuolation due to the chromophobicity of the contents of the vacuoles, normally vacuolated
planarian tissues, or vacuolation due to a disease process.
A heavy infection of gregarine parasites can result in pathological peri-intestinal histolytic vacuolation of
the mesenchyme. In this condition the epidermis is not involved; rather gregarine gamonts and zygocysts are
present in the parenchyma surrounding the gut trunk, branches and diverticula, and gamonts are generally
present in the gut lumen and mesenchyme (L. Winsor, personal communication). No gregarines were present
in the specimens of M. aixandrei examined here, and both epidermis and parenchyma exhibit vacuolation.
Normally vacuolated and vesicular tissues in planarians are associated with the reproductive organs, such
as the phagocytic cells of sperm resorptive tissues in various bursae, vitelline follicles, ovarian tubae, and
parovarian tissues (Cernosvitov, 1931; Sluys, 1989b; Winsor, 2006). The vaculoate epidermis and paren-
chyma in M. aixandrei are located at the anterior tip and are not associated with reproductive structures. Nor
do they exhibit the fine cytological characteristics of resorptive tissues.
In addition to the routine oversight stain, sections were stained with the alcian blue-periodic acid-Schiff
(AB-PAS) technique to detect hexose-containing and sialic acid-containing mucosubstances, and alciano-
philic carbohydrates with carboxylated and sulphate ester groups, which in the adhesive musculo-glandular
organ in the terricolan Pimea can be weakly basiphil to chromophobic (Winsor, 1991). The AB-PAS gave
negative results. It is, therefore, concluded that nothing is present in the vacuoles. The most likely explanation
is that something has been lost during fixation or processing. Strongly acidic fixatives such as Heidenhain’s
SUSA and Bouin’s dissolve or fail to stabilize certain secretory elements such as types of acidophil or
“zymogen” granules (Drury & Wallington, 1980; Leal-Zanchet & Hauser, 1999). Nitric acid is a protein-coag-
ulant fixative, a suitable concentration in a fixative for which is 0.5M (Baker, 1958). However the concentra-
tion of nitric acid in Steinmann’s fixative is approximately 5M, making the reagent very strongly acidic. When
additional specimens of M. aixandrei are available they will be fixed in non-acid and formaldehyde-based fix-
atives for further histochemical studies.
Microplana grazalemica sp. nov.
Material. Holotype, CRBA438, CRBA439, Llano del Berral (lat. 36.75428, long. -5.45399; alt. approx. 657
m) in the central part of the Sierra de Grazalema, Cádiz (Spain), 5 December 2004, sagittal sections on 2
slides.
VILA-FARRÉ ET AL.
8 · Zootaxa 1739 © 2008
Magnolia Press
Other material examined: CRBA440, CRBA441, CRBA442, Río Majaceite (lat. 36.77368, long. -
5.486587; alt. approx. 278 m) in the western part of the Sierra de Grazalema, Cádiz, 6 December 2004, sagit-
tal sections on 3 slides.
Diagnosis. With respect to external features, M. grazalemica sp. nov. can be distinguished from its conge-
ners by its size (12 mm), cylindrical body tapering anteriorly to a blunt point, bluntly pointed tail, brown dor-
sal surface with dark spots, and anterior end without conspicuous eyes. Anatomically the species is
characterized by the presence of a large and rounded penis bulb, obliquely oriented conical penis papilla, a
copulatory bursa not connected to the intestine and communicating with the atrium through a slightly
obliquely orientated bursal canal.
Ecology and distribution. The species is known only from two localities in the Sierra de Grazalema.
Etymology. The specific epithet is based on the name of the mountain system from where the specimens
were collected, i.e. the Sierra de Grazalema in Southern Spain.
Description. In elongated state the living, sexually mature specimens measured 10–12 mm in length, with
a width of about 1 mm (Fig. 3B). The preserved holotype specimen measured 3.45 x 0.68 mm. The cylindrical
body tapers anteriorly and posteriorly to form blunt points. The dorsal surface is brown with darker spots all
over the surface; the anterior end is dark brown. The anterior end of the body is slightly invaginated. The
creeping sole is less than one-third of the body width.
The two small eyes (eye cup diameter 13–16 μm in sections) are located at a short distance anterior to the
brain and are only clearly visible under observation through a dissecting microscope. In CRBA440-442 a third
reduced eye is present close to the anterior body margin.
The subepidermical longitudinal fibres of the body musculature are weak. In the ventral body region
numerous longitudinal fibres are especially present over and under the ventral nerve cords. The scarce dorsal
longitudinal parenchymal fibres are very weak and apparently discontinuous.
In specimen CRBA438-439, the cylindrical pharynx is about one-eight of the total body length (0.3 mm)
and placed in a posterior horizontal position, while in specimen CRBA440-442 the pharynx represents about
one-twelfth of the body length (0.4 mm) and is situated slightly posterior to the central region. The outer epi-
thelium is ciliated and underlain by a layer of longitudinal muscles followed by a layer of circular muscles.
Underneath the inner pharynx epithelium lies an outer layer of circular muscles, followed by an inner layer of
longitudinal muscles fibres. The mouth is located in the middle of the pharyngeal pouch in specimen
CRBA438-439 at 2.2 cm from the tip of the body, but in specimen CRBA440-442 it is situated in the posterior
portion of the pharyngeal pocket, somewhat anterior to the hind wall of the pharyngeal pouch at 2.7 cm from
the tip of the body. In specimens CRBA438-439 and CRBA440-442 the gonopore is situated at 0.23 and 0.6
mm from the mouth, respectively.
There are 13 to 15 testes situated on either side of the body (CRBA438-439 and CRBA440-442, respec-
tively). The rounded or oval-shaped, irregularly sized follicles occupy approximately one-eight of the dorso-
ventral diameter and are arranged in ventral longitudinal rows, extending anteriorly from about the root of the
pharynx up to two-fifth of the distance between the root of the pharynx and the ovaries.
The vasa deferentia open separately into the bulbar lumen, which communicates with an ejaculatory duct
that opens at the tip of the penis papilla (Fig. 4A). The bulbar lumen and the ejaculatory duct are lined with a
nucleated epithelium that is underlain with a layer of circular muscle fibres.
The conical penis papilla has a relatively broad base. It has an oblique ventro-caudal orientation and in
both specimens the tip projects into the bursal canal. The outer wall of the penis papilla is covered with a thin
epithelium, which is underlain with a layer of circular muscles at its distal section; on the middle and more
proximal parts of the papilla the circular muscle layer is considerably thicker and bounded by a layer of longi-
tudinal muscles.
The genital atrium is lined with a nucleated epithelium that is underlain with a subepithelial circular mus-
cle layer, followed by a longitudinal one.
Zootaxa 1739 © 2008 Magnolia Press · 9
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
The small ovaries are situated close to the ventral nerve cord, occupying about one-fifth of the dorso-ven-
tral diameter. The ovaries are located at approximately one-third of the distance between the first testis and the
brain. Somewhat posterior to the gonopore, the oviducts turn towards the middle of the body and open sepa-
rately into the bursal canal.
The copulatory bursa is rounded and slightly flattened; it is lined with tall, vacuolated cells, with the
nuclei mainly in peripheral position (Fig. 4B). The bursa is connected with the atrium by means of a some-
what obliquely running bursal canal (Fig. 5A, B). The lining epithelium of the canal bears long cilia and con-
sists of nucleated cells surrounded by a subepithelial layer of circular muscle fibres. Shell glands could not be
discerned.
FIGURE 3. A. Diagrammatic transverse section through the pharynx of Microplana aixandrei showing the arrangement
of the rows of longitudinal and circular muscles. B. Microplana grazalemica. Living animal. Scale bar 1mm.
VILA-FARRÉ ET AL.
10 · Zootaxa 1739 © 2008
Magnolia Press
Discussion. This species can be distinguished from the other European native land planarians by a combi-
nation of external features and anatomical characteristics of the genital apparatus.
The dorsal colouration pattern of M. grazalemica resembles that of M. nana and M. gadesensis sp. nov,
the latter described below. However, in both M. nana and M. gadesensis a copulatory bursa is absent.
There are three European species that are provided with a copulatory bursa and lack a genito-intestinal
duct, viz. M. mahnerti, M. styriaca, M. aixandrei. In M. mahnerti the bursal canal runs parallel to the body
surface and the oviducts open at its most distal end. In contrast, in M. grazalemica the bursal canal is an
obliquely running structure with the opening of the oviducts at its central region. (Minelli, 1977). In M. styri-
aca the penis bulb is very large and elongated, while an expanded bulbar lumen is absent, in contrast to the
rounded penis bulb from M. grazalemica that is provided with a distinct bulbar lumen. In M. styriaca the ovi-
ducts form a short common oviduct that enters the bursal canal (Freisling, 1935), contrasting with the oviducts
of M. grazalemica, which open separately into the bursal canal.
Differences between M. aixandrei and M. grazalemica were detailed above.
FIGURE 4. Microplana grazalemica. CRBA438-A439.A–B. Sagittal sections of the copulatory apparatus; anterior to
the left.
Zootaxa 1739 © 2008 Magnolia Press · 11
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
FIGURE 5. Microplana grazalemica. CRBA438-A439. A. Sagittal reconstruction of the copulatory apparatus; anterior
to the left. CRBA440-442. B. Sagittal reconstruction of the copulatory apparatus; anterior to the left.
Microplana gadesensis sp. nov.
Material: Holotype, CRBA443, CRBA444, Llano del Berral (lat. 36.75428, long. -5.45399; alt. aprox. 657
m) in the central part of the Sierra de Grazalema, Cádiz (Spain), 5 December 2004, sagittal sections on 2
slides. This specimen was accidentally broken in two pieces while it was being embedded in paraffin.
Paratype: CRBA445, ibid., sagittal sections on 1 slide.
Diagnosis. Microplana gadesensis sp. nov. can be distinguished anatomically from its congeners by a
muscular penis bulb provided with a very thick layer of circular muscle fibres, rounded bulbar lumen, large
penis papilla provided with a wide ejaculatory duct, genito-intestinal duct that links the female genital duct to
the gut, and absence of a copulatory bursa.
VILA-FARRÉ ET AL.
12 · Zootaxa 1739 © 2008
Magnolia Press
Ecology and distribution. The species is known only from the type locality.
Etymology. The specific epithet is based on the name of the region where the specimens were collected,
Cádiz, named Gades in Latin, in southern Spain.
Description. When the animals were fixed in the field and later prepared for sectioning, they were consid-
ered to be morphologically similar to M. grazalemica, and therefore no particular attention was paid to their
external features. It may be assumed that their external appearance did not differ much from that of M. graza-
lemica. From the sections of the holotype it could be determined that the animal was about 10 mm long.
The two small eyes (eye cup diameter 16 µm in sections) are located at a short distance in front of the
brain (Fig. 6) and are only clearly visible under observation through a dissecting microscope.
The subepidermical longitudinal fibres of the body musculature weak. In the ventral body region numer-
ous strong parenchymal longitudinal fibres are present especially dorsally and ventrally of the ventral nerve
cords. This ventral musculature becomes especially strong at the cephalic region, where it attaches at the epi-
thelium of an invaginated area (Fig. 6). This longitudinal musculature partially encloses the eyes.
The short pharynx is about one-twelfth of the total body length (0.3 mm), its root being located slightly
posterior to the middle of the body. The outer epithelium is ciliated from the root to the tip of the pharynx. It is
underlain by a thin layer of longitudinal muscles followed by a thin layer of circular muscles. The inner epi-
thelium of the pharynx is underlain with a very thick outer layer of circular muscles, followed by a longitudi-
nal muscles layer, the latter in some areas intermingled with the circular layer. In specimen CRBA445 the
mouth is situated close to the posterior wall of the pharyngeal pouch at 2.5 cm from the tip of the body and 0.9
mm from the gonopore.
The number of elongated testes varies from four (CRBA445) to six (CRBA443-444) on either side of the
body. The follicles occupy approximately one-fourth of the dorso-ventral diameter and are arranged in longi-
tudinal rows, extending from some distance behind the ovaries to the root of the pharynx.
The penis bulb is provided with a rounded bulbar lumen, leading to a very wide ejaculatory duct that runs
centrally through the penis papilla and gradually narrows before opening at the tip of the papilla (Fig. 7A).
The bulbar lumen and the ejaculatory duct are lined with a nucleated epithelium. The bulbar lumen is sur-
rounded by a thick layer of circular muscle fibres that extends on the proximal part of the penis papilla; this
coat of circular muscle is highly developed on the ventral side of the seminal vesicle.
The large and elongated penis papilla is highly muscular; it is oriented parallel to the body surface. At the
level of the penis bulb, the two vasa deferentia turn dorsally and open separately into the anterior portion of
the bulbar lumen.
The penis papilla projects into the female genital atrium. The gonopore opens into the mid-ventral section
of the atrium.
The small ovaries occupy one-fifth of the dorso-ventral diameter, lying on top of the ventral nerve cords.
The ovaries are located at approximately one-sixth of the distance between the brain and the root of the phar-
ynx. The oviducts arise from the ventral side of the ovaries. The rather narrow female genital duct is provided
with distinct cilia and receives the openings of the oviducts at its posterior end. The duct is lined with nucle-
ated epithelium that is underlain with a thick, subepithelial layer of circular muscles followed by a layer of
longitudinal muscle fibres. The female genital duct of specimen CRBA445 receives the openings of the shell
glands anteriorly to the openings of the oviducts. Secretion granules are present in the same area in the holo-
type, but not the glands itself. A long genito-intestinal duct arises from the distal end of the female genital
duct (Fig. 7B) and immediately recurves to run approximately parallel to the dorsal wall of the atrium, subse-
quently communicating with the gut. The genito-intestinal duct is lined with a ciliated epithelium and is sur-
rounded by a layer of circular muscles (Figs. 8A, B).
The wall of the elongated atrium is lined with a thin epithelium, provided with a thin circular muscular
layer, surrounded by a thin longitudinal layer of muscles.
Zootaxa 1739 © 2008 Magnolia Press · 13
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
FIGURE 6. Microplana gadesensis. CRBA443-444. Sagittal section through the head.
Discussion. Microplana gadesensis can be distinguished from the other known native European land pla-
narians species by the anatomy of its copulatory apparatus.
Regarding the external features, similar body colouration is found in M. nana and M. grazalemica. How-
ever, M. nana and M. grazalemica lack a genito-intestinal duct and also the thick layer of circular muscle
fibres around the bulbar lumen.
Among the known European species of Microplana a genito-intestinal duct has been reported also for M.
humicola, M. pyrenaica, M. howesi (Scharff, 1900), M. giustii Minelli, 1976, M. henrici (Bendl, 1908), M.
attemsi (Bendl, 1909), M. peneckei (Meixner, 1921), M. scharffi (von Graff, 1899), M. monacensis (Heinzel,
1929), and M. terrestris. However, M. humicola presents a hyaline colouration (Schneider, 1935), instead of
the pigmented dorsal colouration pattern of M. gadesensis. With respect to anatomical features, M. humicola
has two testes on either side of the body, while its penis papilla is vertically oriented, in contrast to the 4-6 tes-
tes and horizontally oriented penis papilla of M. gadesensis. M. pyrenaica and M. howesi are bigger species
(up to 5 cm long) with a poorly developed penis papilla (Minelli, 1977), in contrast to the large and well
defined papilla of M. gadesensis. In M. giustii, the testes are present in a postpharyngeal position (Minelli,
1976), contrasting with the prepharyngeal testes of M. gadesensis. M. henrici, M. attemsi, and M. peneckei
show atrial folds (Minelli, 1977), whereas atrial folds are absent in M. gadesensis. In M. scharffi the vasa def-
erentia fuse to a common vas deferens at the base of the penis papilla (Ball & Reynoldson, 1981). In M. gade-
sensis the vasa deferentia open into the bulbar lumen, far anterior to the root of the penis papilla. With respect
to external features, M. scharffi is a very long (maximum 90 mm) animal with a pale colouration, whereas M.
gadesensis is shorter and with darker colouration. In M. monacensis (Heinzel, 1929) and M. terrestris
(Minelli, 1977) a bursa is present, whereas such a structure is absent in M. gadesensis.
VILA-FARRÉ ET AL.
14 · Zootaxa 1739 © 2008
Magnolia Press
FIGURE 7. Microplana gadesensis. CRBA443-444. A–B. Sagittal sections of the copulatory apparatus; anterior to the
left.
Biogeography and biodiversity
In the present study, we present 54 new locality records for 202 individuals (Fig. 1, Table 1) from the Iberian
Peninsula, with an undetermined number of species per record. The intrinsic complexity of the identification
of planarian specimens (see Introduction) presently prevents us to provide an extensive list of species col-
lected at each locality. Since this is a work in process, further results will be presented in future publications.
However, the external features of the specimens collected suggest that usually more than one species is
present at a single locality.
The southern sector of the Iberian Peninsula has been sampled in three distinct areas, yielding 7 new
localities and 43 specimens. We have focused our analysis on the region of the Sierra de Grazalema, which
has the maximum average annual rainfall (more than 2000 mm) of the entire Iberian Peninsula, thus poten-
tially forming a suitable area for terrestrial planarians. The material that has been analysed so far yielded three
Zootaxa 1739 © 2008 Magnolia Press · 15
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
TABLE 1. List of localities with new records of autochthonous terrestrial planarians in the Iberian Peninsula. The first
locality of the unidentified species belongs to Portugal while the rest belongs to Spain. Latitude and longitude are
expressed in decimal degrees (Datum European 1979). *: locality with populations of the three new species described in
the present paper (M. aixandrei, M. grazalemica, M. gadesensis).
Locality Province Latitude Longitude Species
Eo river Asturias 43.42404956 -7.13722351 Rynchodemus sylvaticus
Luarca Asturias 43.53122231 -6.54066729 Rynchodemus sylvaticus
Serra del Corredor, riera Can Rimbles Barcelona 41.594097 2.46444 Rynchodemus sylvaticus
Serra del Corredor, riera de Canyamars Barcelona 41.598317 2.44302 Rynchodemus sylvaticus
Montjuïc mountain, Vivers dels Tres
Pins Barcelona 41.36903802 2.16342328 Rynchodemus sylvaticus
Collsacabra, Puig de Rajols Girona 41.996119 2.488137 Rynchodemus sylvaticus
Sierra de Grazalema, Río Majaceite Cádiz 36.773681 -5.486587 Microplana grazalemica
sp.nov
Sierra de Grazalema, Llano del Berral* Cádiz 36.75428 -5.45399 three newly described
Microplana
Mata da Margaraça, Serra do Açor Coimbra 40.21681 -7.90961 unidentified
Sant Llorenç del Munt, Sot Teixoneres Barcelona 41.66168 1.99706 unidentified
Collserola, La Rierada Barcelona 41.437904 2.056765 unidentified
Collserola, Can Catà Barcelona 41.474183 2.147384 unidentified
Serra del Corredor, torrent de
Canyamars Barcelona 41.605197 2.45639 unidentified
Serra de l'Obac Barcelona 41.649761 1.97323 unidentified
Montnegre, Sot la Massaneda Barcelona 41.668975 2.602379 unidentified
Montnegre, Pantà la Brinxa Barcelona 41.678624 2.587049 unidentified
Montnegre, àrea Ramió, Sot de Can
Masó Barcelona 41.708653 2.616559 unidentified
Montseny mountains, Tordera river
shore Barcelona 41.728985 2.419703 unidentified
Montseny mountains, Tordera river
shore Barcelona 41.733335 2.415153 unidentified
Montseny town Barcelona 41.758725 2.399094 unidentified
Montseny mountains, el Molar Barcelona 41.763575 2.365855 unidentified
Montseny mountains Barcelona 41.768404 2.467023 unidentified
Montseny mountains, Sot les Mines Barcelona 41.777115 2.353205 unidentified
Font del Vidre Barcelona 42.151601 1.93114 unidentified
Los Alcornocales Cádiz 36.091333 -5.6222 unidentified
Sierra de Grazalema, Llanos del
Republicano Cádiz 36.696744 -5.376621 unidentified
Sierra de Grazalema, salto del cabrero
path Cádiz 36.706868 -5.421502 unidentified
Sierra de Grazalema, Las Covezuelas Cádiz 36.711273 -5.3684 unidentified
La Miel Cádiz 36.11933041 -5.47959327 unidentified
Fragas do Eume A Coruña 43.417185 -8.063563 unidentified
Joanetes Girona 42.116900 2.400990 unidentified
Puigpardines Girona 42.142877 2.42516 unidentified
VILA-FARRÉ ET AL.
16 · Zootaxa 1739 © 2008
Magnolia Press
new species, suggesting that upon further research the species richness of this area might turn out to be higher
than in other sections of the Iberian Peninsula.
The presence of R. sylvaticus is reported for new areas of the northern sector of the Iberian Peninsula, viz.
the Cantabric and the Mediterranean regions (Fig. 1), where other elements of the Centro-European fauna are
also usually present. It should be noted that one of the records corresponds to a plant nursery (Viver dels Tres
Pins, Barcelona, NE Spain). However, the others correspond to natural areas.
General discussion
For practical purposes we have adopted in this paper the higher classification of the land planarians that is cur-
rently available (Kawakatsu et al., 2003 and references therein). However, we are well aware of the fact that
the higher classification of triclads in general and that of land planarians in particular, is in need of revision.
Since a revised classifcation of the triclad flatworms is in preparation and will be published elsewhere and the
primary purpose of the present paper is the description of three new species, we have adopted the currently
classical higher taxonomy of the terrestrial planarians.
There are three genera of autochthonous European land planarians, viz. Rhynchodemus (subfamily Rhyn-
codeminae), Geobenazzia and Microplana (subfamily Microplaninae). We have assigned the three new spe-
cies to the genus Microplana on the basis of the taxonomic definition of this genus presented in Kawakatsu et
al. 2003: “Microplaninae with an elongate, rounded body (preserved worms may have rounded anterior with
greatest diameter anterior to mouth); with two small eyes; male copulatory organ consists of well-developed
Fageda d'en Jordà Girona 42.149366 2.519729 unidentified
Ansó valley Huesca 42.886305 -0.805996 unidentified
Serra del Tallat, Montblanquet Lleida 41.481613 1.124143 unidentified
Montsec, Sant del Bosc Lleida 42.066583 0.901788 unidentified
Talarn Lleida 42.18945694 0.89633052 unidentified
Capdella, camí Filià Lleida 42.47181881 0.98914475 unidentified
Capdella Lleida 42.47638271 0.99366981 unidentified
Sierra Cebollera, Puente Ra Logroño 42.04626 -2.68545 unidentified
Monasterio de Valvanera Logroño 42.23156 -2.85017 unidentified
Isaba, Belabarte creek shore Navarra 42.877835 -0.862635 unidentified
Roncal valley, Belagua river shore Navarra 42.890385 -0.892624 unidentified
Anduña river shore Navarra 42.921576 -1.011301 unidentified
Irati forest Navarra 42.988866 -1.098518 unidentified
Irati forest Navarra 43.007606 -1.201696 unidentified
Road NA174, Artesiaga river shore Navarra 43.113056 -1.538889 unidentified
Tximista river shore Navarra 43.233825 -1.659663 unidentified
Etxalar, water channel margin Navarra 43.240646 -1.662193 unidentified
Road NA 4400 from Etxalar to Sare Navarra 43.242546 -1.62907 unidentified
Road NA 4000 from Lesaka to
Endara's marsh Navarra 43.269574 -1.749217 unidentified
Tail of Endara's marsh Navarra 43.273254 -1.775105 unidentified
Els Ports de Beceït Tarragona 40.707546 0.236715 unidentified
Ergoyen Vizcaya 43.32776899 2.81787795 unidentified
Zootaxa 1739 © 2008 Magnolia Press · 17
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
penis with muscular bulbus, bulbar cavity and elongate penis papilla projecting into short antrum; female
organ has one simple canal (vagina) entering common genital antrum; typically a genito- intestinal canal is
present arising from vagina or behind the common ovovitelline duct; seminal bursa can be present with a short
connection (bursasteles, bursal canal, or proximal duct) to female tract, with or without connections to intes-
tine, or bursa is blind lacking connections to intestine; when bursa is absent a genito-intestinal connection is
often present; without adenodactyls, tentacles or suckers.”
FIGURE 8. Microplana gadesensis. CRBA443-444. A. Sagittal reconstruction of the copulatory apparatus; anterior to
the left. CRBA445. B. Sagittal reconstruction of the copulatory apparatus; anterior to the left.
A genito-intestinal duct is reported here only for M. gadesensis. The taxonomic characters of this species
are in complete agreement with those mentioned in the definition of the genus Microplana (Kawakatsu et al.,
2003). Microplana aixandrei and M. grazalemica possess a copulatory bursa (or seminal bursa) that is con-
nected to the female tract but not to the intestine. Kawakatsu et al. (2003) consider M. nana and M. perereca
VILA-FARRÉ ET AL.
18 · Zootaxa 1739 © 2008
Magnolia Press
Marcus & Marcus, 1959 to be good species of the genus Microplana, albeit that these species lack a genito-
intestinal duct. Therefore we assign M. aixandrei, M. grazalemica, and M. gadesensis to the genus
Microplana and not to Rhynchodemus. In Rhynchodemus a genito-intestinal duct and a well developed penis
papilla are absent and the subepithelial longitudinal muscle fibres of the strong cortical musculature are
grouped into large, definite bundles, in contrast to species of Microplana, including the three described as new
in the present paper.
In the present work we have not considered the synonymic treatment proposed in Minelli (1977) and
Kawakatsu et al. (2003). In this paper we follow the advice of H. Jones (personal communication), who has
examined part of the original slides corresponding to the synonymized species and considers that some of
them must maintain their original generic assignments.
Presence of a spermatophore is extremely rare in terrestrial flatworms. It has been described only for the
geoplanid Pimea monticola Winsor, 1991 and the Australian rhynchodemid Platydemus victoriae (Dendy,
1898), a member of the subfamily Rhynchodeminae (Winsor, 1998). The spermatophore in P. monticola dif-
fers in size (500 µm) and composition (multiple layers that completely envelope sperm) from the structure
observed in M. aixandrei. In P. victoriae the spermatophore presents a thin red single layer enveloping the
sperm similar to the single layered structure observed in M. aixandrei, but probably of different nature.
Although we did not observe sperm in the putative spermatophore of M. aixandrei the existence of a sper-
matophore of sclerotic structure is reported also for the freshwater planarians (Sluys, 1989a). Similary, Sluys
and Ball (1989) suggested the existence of a spermatophore in Vatapa tumidosa (Sluys & Ball 1989), a marine
planarian, on the basis of similar reasons (localization in the copulatory bursa and sclerotic nature), although
sperm was not observed. The recurrent observation of this structure only in the copulatory bursa of M. aixan-
drei reinforces our idea that it represents a spermatophore. This spermatophore in M. aixandrei represents the
first report of such a structure in a member of the subfamily Microplaninae.
The amphiatlantic species R. sylvaticus is widely distributed in continental Europe and the British Isles,
including an isolated record in northern Spain (Minelli, 1977; Boix & Sala, 2001). On the basis of our new
findings, R. sylvaticus is probably widely distributed in the northern sector of the Iberian Peninsula. Further-
more, its special strategy of locomotion in which the animals leave a dotted slime-trail and thus use less
mucus (mainly composed of water) allows the species to live in relative dry areas (Pantin, 1950). Such areas
are present in southern Spain and Portugal (with Mediterranean climatic characteristics), where the species
has not yet been collected.
Sluys (1999) published several maps of the world summarizing the biogeographic data on species richness
and endemism of land planarians. Only few data have been added to the European fauna since then, thus this
work still holds at the present time.
In his map of species richness, Sluys (1999, Fig. 1), the two grid cells corresponding to the northern sector
of the Iberian Peninsula and the Balearic Islands have two and three records, corresponding with at least four
different species in total, while no record is provided for the southern sector. The new data presented in this
paper suggest that this low specific richness is due to a collector’s artefact.
In contrast to the few records available in the literature, our intensive sampling efforts show that land pla-
narians are widely distributed across the Iberian Peninsula. The ecological conditions required for their pres-
ence (Kawaguti, 1932; Ball & Reynoldson, 1981), especially high humidity, are present on the Iberian
Peninsula in forested areas and near shores of rivers. The extremely low density of the worms, their cryptic
habitus, and their specific habitat requirements ask for a thorough sampling methodology (detailed searches of
covered surfaces of stones and adjacent ground) and an intensive sampling effort for successful fieldwork.
The results presented in this study show that future studies on terrestrial planarians most likely will yield other
new and interesting information on this group of animals on the Iberian Peninsula.
Zootaxa 1739 © 2008 Magnolia Press · 19
THREE NEW TERRICOLA FROM THE IBERIAN PENINSULA
Acknowledgements
We are grateful to Edgar Vila Farré, Sergi Rodríguez, Oscar Gordo, Mar Sancho Prat, Salvador Carranza and
Cristina Cabrera for their help in the field and to Eduard Filella for his information on the current status of
Bipalium kewense on the Iberian Peninsula. We are indebted to Hugh Jones and Manuel Irimia for their valu-
able suggestions during the preparation of this paper and to Kay Eckelt and Michael Lang for their help in the
translation of the German texts. We are grateful to Dr. L. Winsor and Prof. Dr. M. Kawakatsu for reviewing
and commenting on the penultimate version of the manuscript. Dr. L. Winsor also kindly assisted with the his-
tological interpretation of the vacuolated tissues in one of the species. MVF acknowledges financial support
from SYNTHESYS, a program of the European Commission under the Sixth Research and Technological
Development Framework Programme “Structuring the European Research Area”, which enabled MVF to
work at the Zoological Museum of the University of Amsterdam during March 2005 (grant number NL-TAF-
179). This work is also supported by grant BFU2004-05015/BFI (to R.R.) from the “Ministerio de Uducación
y Ciencia” of Spain. Prof. Dr. M. Kawakatsu and Dr. T. Kifune are thanked for nomenclatural advice.
References
Baker, J.R. (1958) Principles of biological microtechnique. Methuen, London, 357 pp.
Ball, I.R. & Reynoldson, T.B. (1981) British Planarians. Cambridge University Press, Cambridge, VII+142 pp.
Boag, G. & Yeates, W. (2001) The potential impact of the New Zealand Flatworm, a predator of earthworms, in Western
Europe. Ecological Applications, 11, 1276–1286.
Boix, D. & Sala, J. (2001) Presència del gènere Rhynchodemus (Platyhelminthes, Tricladida, Terricola) a la Península
Ibèrica. Scientia gerundenesis, 25, 31–32.
Cernosvitov, L. (1931) Studien über die Spermaresorption. III Die Samenresorption bei den Tricladen. Zoologisches
Jahrbuch, 54, 295–332.
Drury, R. A. & Wallington, E. A. (1980) Carleton's Histological Techniques (5th ed.). Oxford University Press, Oxford,
520 pp.
Filella, E. (1983) Nota sobre la presència de la planària terrestre Bipalium kewense Moseley, 1878 a Catalunya. Butlletí
de la Institució Catalana d’Història Natural, 49, 151.
Freisling, J. (1935) Über Rhynchodemiden (Turbellaria, Tricladida Terricola) der Gattung Rhynchodemus Leidy und
Artiocotylus Graff und ihre weiblichen Geschlechtsgänge. Mitteilungen des Naturwissenschaftlichen Vereins für
Steiermark, 75, 5–22.
Heinzel, L. (1929) Zur Kenntnis der Rhynchodemiden. Zoologische Jahrbücher, Abteilung für Systematik, 56, 425–462.
Jones, H.D. (1998) The African and European land planarian fauna, with an identification guide for field workers in
Europe. Pedobiologia, 42, 477–489.
Kawaguti, S. (1932) On the physiology of land planarians. Memoirs of the Faculty of Science and Agriculture, Taihoku
Imperial University, 7, 15–55.
Kawakatsu, M., Froehlich, E.M., Jones, H.D., Ogren, R.E. & Sasaki, G. (2003) Additions and corrections of the previous
land planarian indices of the world (Turbellaria, Seriata, Tricladida, Terricola). Additions and corrections of the pre-
vious land planarian indices of the world–11. The Bulletin of Fuji Women's University, 41, 89–114.
Leal-Zanchet & A.M., Hauser, J. (1999) Penis glands of the dugesiid planarian Girardia schubarti (Platyhelminthes, Tri-
cladida, Paludicola). Invertebrate Biology, 118, 35–41.
Mateos, E., Giribet, G. & Carranza, S. (1998) Terrestrial planarians (Platyhelminthes, Terricola) from the Iberian Penin-
sula: first records of the family Rhynchodemidae, with the description of a new Microplana species. Contributions
to Zoology, 67 267–276.
Mateos, E., Vila, M. & Romero, R. (2004) Presència de planàries terrestres del gènere Microplana (Platyhelminthes, Tri-
cladida, Terricola) al Parc del Montnegre i el Corredor. In: Barcelona, D.d. (Ed.), IV Trobada d’Estudiosos del Mont-
negre i el Corredor, 101–104.
Minelli, A. (1976) Planarie terrestri dell'Arcipelago Toscano (Studi sulla Riserva Naturale dell'Isola di Montecristo. III).
Lavori della Società Italiana di Biogeografia, 5, 93–98.
Minelli, A. (1977) A taxonomic review of the terrestrial planarians of Europe. Bollettino di Zoologia, 44, 399–419.
Murchie, K.M., Moore, J.P., Walters, K. & Blacksaw, R.P. (2003) Invasion of agricultural land by the earthworm preda-
tor, Arthurdendyus triangulatus (Dendy). Pedobiologia, 47, 920–923.
Pantin, C.F.A. (1950) Locomotion in British terrestrial nemertines and planarians: with a discussion on the identity of
VILA-FARRÉ ET AL.
20 · Zootaxa 1739 © 2008
Magnolia Press
Rhynchodemus bilineatus (Mecznikow) in Britain, and on the name Fasciola terrestris, O. F. Müller. Proceedings of
the Linnean Society of London, 162, 23–37.
Romeis, B. (1989) Mikroskopische Technik. Ed. P. Böck, 17th edition, Urban und Schwarzenberg, München, Wien, Balti-
more, 697 pp.
Schneider, F.W. (1935) Zur Anatomie und systematischen Stellung von "Microplana" humicola (VEJD.). Zoologische
Jahrbücher, Abteilung für Systematik, 62, 179–194.
Sluys, R. (1989a) A monograph of the marine triclads. A. A. Balkema, Rotterdam & Brookfield, Rotterdam, XII + 463
pp.
Sluys, R. (1989b) Sperm resorption in triclads (Platyhelminthes, Tricladida). Invertebrate Reproduction and Develop-
ment, 15, 89–95.
Sluys, R. (1999) Global diversity of land planarians (Platyhelminthes, Tricladida, Terricola): a new indicator-taxon in
biodiversity and conservation studies. Biodiversity and Conservation, 8, 1663–1681.
Sluys, R. & Ball, I.R. (1989) A synopsis of the marine triclads of Australia and New Zeland (Plathyelminthes, Tricladida,
Maricola). Invertebrate Taxonomy, 2, 915–959.
Winsor, L. (1991) A new genus and species of terrestrial flatworm from the central highlands of New Caledonia (Tricla-
dida: Terricola). Mémoirs du Muséum nationale D’Histoire Naturelle. Series A, 149, 19–30.
Winsor, L. (1998) Aspects on taxonomy and functional histology in terrestrial flatworms (Tricladida: Terricola). Pedobi-
ologia, 42, 412–432.
Winsor, L. (2006) New and revised terrestrial flatworm taxa (Platyhelminthes: Tricladida: Terricola) of Australia and the
Subantarctic Islands of New Zealand. Tuhinga, 17, 81–104.
