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Journal of Natural History
Vol. 45, Nos. 15–16, April 2011, 869–891
Land planarians (Platyhelminthes: Tricladida: Geoplanidae)
from the Iberian Peninsula: new records and description of two new
species, with a discussion on ecology
Miquel Vila-Farré
a,b,c∗
, Ronald Sluys
d
, Eduardo Mateos
b
, Hugh D. Jones
e
and
Rafael Romero
a
a
Departament de Genètica, Facultat de Biologia, Universitat de Barcelona, Barcelona, Spain;
b
Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Barcelona,
Spain;
c
Departamento de Biología Evolutiva y Biodiversidad, Museo Nacional de Ciencias
Naturales-CSIC, Madrid, Spain;
d
Institute for Biodiversity and Ecosystem Dynamics &
Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The
Netherlands;
e
Department of Zoology, The Natural History Museum, London, Cromwell Road,
London, SW7 5B, UK
(Received 22 July 2010; final version received 27 October 2010; printed 31 March 2011)
Two new species of the genus Microplana are described from the Iberian Peninsula.
The new taxa are compared with congeneric species. Distributional records for two
other European species, Microplana monacensis (Heinzel, 1929) and Microplana
groga Jones et al. 2008, are presented and the presence of Microplana terrestris
(Müller, 1774) is confirmed on the Iberian Peninsula. A partial re-description of
Microplana nana Mateos, Giribet and Carranza, 1998 is provided. The finding
of a new and probably introduced, but unidentified, species of land planarian is
reported. Land planarians were generally found in the vicinity of deciduous trees
and rivers.
Keywords: Platyhelminthes; Tricladida; Microplana; Rhynchodemus;Iberian
Peninsula; taxonomy; ecology
Introduction
Detailed systematic studies on the terrestrial planarians of the Iberian Peninsula only
started as recently as the early 1980s (Filella-Subirà 1983). Earlier workers speculated
on the identity of observed specimens that were known only by their external appear-
ance (cf. Von Graff 1899), and the relevant literature is still limited to a few papers (cf.
Vila-Farré et al. 2008 and references therein; Mateos et al. 2009). Recently, Jones et al.
(2008) clarified the taxonomic status of some European species. Following this new
taxonomic information and the analysis of the Iberian terrestrial planarians initiated
by Vila-Farré et al. (2008), we describe two new species from Spain and provide the
second distributional records since their first descriptions for two European species,
Microplana monacensis and Microplana groga. Furthermore, we confirm the pres-
ence of populations of Microplana terrestris (Müller, 1774) on the Iberian Peninsula
and provide an account of the morphological differences between the various pop-
ulations. During our re-examination of the type material of the Spanish species
*Corresponding author. Email: mvilafarre@gmail.com
ISSN 0022-2933 print/ISSN 1464-5262 online
© 2011 Taylor & Francis
DOI: 10.1080/00222933.2010.536267
http://www.informaworld.com
870 M. Vila-Farré et al.
Microplana nana we noted some discrepancies between the anatomy of the specimens
and the account provided by Mateos et al. (1998). Therefore, we present a partial re-
description of this species. In addition, the presence of a new and probably introduced
species of land planarian is reported for the area.
Materials and methods
Between 1998 and 2006 different areas of the Iberian Peninsula were sampled for
terrestrial planarians. A distribution map of this area, the species discovered, and
an account of the methodological approach, is provided by Vila-Farré et al. (2008).
From 2007 to 2009 many of these areas have been revisited and, in addition, a
new locality in southern Spain has been sampled, namely Benamargosa, Málaga
(Table 1). Specimens were individually transported in small plastic containers (with
humid soil from the area where they were collected) to the laboratory, where they
were photographed using a digital camera (Nikon CoolPix 995). For anatomical
and histological studies, animals killed (under field or laboratory conditions) with
Steinmann’s fluid (cf. Sluys 1989a) were subsequently fixed in 70% ethanol. Fixed
specimens were cleared in clove oil, dehydrated in an ascending series of alcohol
concentrations, and then embedded in paraffin wax, cut at intervals of 5–7 µm, and
mounted on gelatine-coated slides. The sections were stained in Mallory–Heidenhain
(cf. Sluys 1989a) or Harris haematoxylin (cf. Humason 1967) and mounted in DePeX.
Reconstructions of the copulatory apparatus were obtained using a camera lucida
attached to a compound microscope. The identification of Rhynchodemus sylvati-
cus (Leidy, 1851) was not based on histological sections but on its peculiar external
morphology. The material examined is deposited in the Centre de Recursos de
Biodiversitat Animal, Facultat de Biologia, Universitat de Barcelona, Spain (CRBA),
the Zoological Museum Amsterdam, the Netherlands (ZMA), the Natural History
Museum, London, UK (NHML), and the Natural History Museum, Vienna, Austria
(NHMW).
Table 1. List of localities with new records of terrestrial planarians in the Iberian Peninsula.
Locality Province Longitude Latitude Species
Fragas do Eume A Coruña −8.0636 43.4172 Microplana robusta,
Microplana terrestris
Montseny,
el Molar
Barcelona 2.3658 41.7636 Microplana groga
Capdella Lleida 0.9937 42.4764 Microplana terrestris
Sant del Bosc,
Cellers
Lleida 0.9410 42.0535 Microplana hyalina
Benamargosa Málaga −4.1809 36.8248 Rhynchodemus sylvaticus;
introduced species
Etxalar Navarra −1.6622 43.2406 Microplana monacensis
Irurita Navarra −1.5389 43.1130 Microplana monacensis
Note: Latitude and longitude are expressed in decimal degrees (Datum European 1979).
Journal of Natural History 871
Abbreviations used in the figures: ae, anterior end; bc, bursal canal; bca, bul-
bar cavity; cb, copulatory bursa; cm, circular muscles; dep, dorsal epidermis; ed,
ejaculatory duct; fgd, female genital duct; gid, genito-intestinal duct; gp, gonopore;
i, intestine; is, intestinal structure; lm, longitudinal muscles; od, oviduct; pb, penis
bulb; ph, pharynx; phc, pharyngeal cavity; pp, penis papilla; scb, secondary copula-
tory bursa; sg, shell glands; sm, spongy mesenchyma; vd, vas deferens; vep, ventral
epidermis; vnc, ventral nerve cord.
Systematic account
Order TRICLADIDA Lang, 1884
Suborder CONTINENTICOLA Carranza et al. 1998
Family GEOPLANIDAE Stimpson, 1857
Genus Microplana Vejdovsky, 1890
Microplana terrestris (Müller, 1774)
(Figures 1, 2, Table 1)
Material examined
ZMA V.Pl. 6855.1, Parque Natural Fragas do Eume, province of A Coruña (north-
western Spain), 20 November 2004, sagittal sections on 12 slides; V.Pl. 6855.2, ibid.,
sagittal sections on 11 slides; V.Pl. 6855.3, ibid., sagittal sections on 14 slides. CRBA-
3775a-e, Parque Natural Fragas do Eume, province of A Coruña (northwestern
Spain), 20 November 2004, sagittal sections on five slides. CRBA-3774a-d, Capdella,
north of the province of Lleida (Spain), 23 March 2004, sagittal sections on four slides;
CRBA-3779a-l, ibid., sagittal sections on 12 slides. CRBA-3780a-l, Capdella, north
of the province of Lleida (northeastern Spain), unknown date, sagittal sections on
12 slides.
Comparative discussion
Among European land planarians M. terrestris has the largest distributional range; it
has been reported from the USA (cf. Ogren and Kawakatsu 1998). The first apparent
record for the Iberian Peninsula (Mateos et al. 1998) has been recently shown to be
a misidentification (Jones et al. 2008). In a recent study Mateos et al. (2009) reported
the species from the northern part of the Iberian Peninsula, this time identifying the
species only on the basis of molecular techniques. During our samplings many locali-
ties have yielded specimens with external traits corresponding to those of M. terrestris:
large size; grey or black colour; broad, white creeping sole. Two of these populations,
distributed in the north of the Iberian Peninsula (Figure 1), have been analysed. Earlier
papers on the species (Von Ke nnel 1882; Bendl 1908; Jones et al. 2008) have adequately
detailed the morphology of the copulatory apparatus and other anatomical structures.
We will here only mention those traits that were characteristic of each population.
First, animals from Parque Natural Fragas do Eume (Figure 2A, B) have two small
structures under the intestine (Figure 2B) and above the penis papilla, lined with an
epithelium that is histologically very similar to that of the copulatory bursa and the
genito-intestinal duct. These structures were found in all three specimens examined.
Although it is known that more than one connection may exist between the copulatory
872 M. Vila-Farré et al.
Figure 1. Distributional records of terrestrial planarians from the Iberian Peninsula used for
this study. Filled circles: localities with previously known species. Stars: localities with new
species.
bursa and the intestine (cf. Von Graff 1899; Bendl 1908) this structure in our specimens
is clearly independent, but could well be a remnant of a second genito-intestinal duct.
Second, specimens from Capdella (Figures 2C, D) have a vertically oriented penis
papilla (Figure 2C). The new records considerably extend the known distribution
range of the species.
Microplana groga Jones, Webster, Littlewood and Mc Donald, 2008
(Figures 1, 2, Table 1)
Material examined
The slides used for the identification of these animals from the area of el Montseny,
el Molar (Catalunya, Northeastern Spain, one specimen, 1 February 2004), unfortu-
nately have been lost.
Comparative discussion
Jones et al. (2008) described this species on the basis of five individuals previously
identified as M. terrestris by Mateos et al. (1998). The individual presented here is
attributed to M. groga and belongs to a population approximately 50 km from the
type locality. It presents characteristics similar to those of the animals described by
Jones et al. (2008) but it is externally different (Figure 3A). Instead of being “red-
dish to black” it is cream with small darker spots covering the entire dorsal surface.
However, the copulatory apparatus (Figure 3B) is similar to that of M. groga, albeit
that in our specimen the circular muscle layer surrounding the ejaculatory duct is
stronger. Although we have observed differences between the external morphology
of this specimen and M. groga we believe that both populations belong to the same
species because of the similarities in the structure of the copulatory apparatus. Only
after collecting new material will we be able to discern whether these are separate
species rather than one, variable species.
Journal of Natural History 873
Figure 2. Microplana terrestris. (A,B) Specimen from Fragas do Eume: (A) living animal;
(B) ZMA V.Pl. 6855.1, sagittal reconstruction of the copulatory apparatus; anterior to the left;
epithelia of bulbar cavity and ejaculatory duct indicated by dotted line because limits are poorly
defined in histological sections. (C,D) Specimen from Capdella; CRBA-3779a-l: (C) sagittal
reconstruction of the copulatory apparatus; anterior to the left; (D) sagittal sections of the
copulatory apparatus; anterior to the left.
Microplana monacensis (Heinzel, 1929)
(Figures 1, 4, Table 1)
Material examined
Holotype. NHMW 2849, transverse sections of the anterior part on two slides, trans-
verse sections of the middle part of the body on two slides, sagittal sections of the
posterior region on three slides.
Other material. ZMA V.Pl 6856.1, Etxalar, close to an irrigation channel, Navarra
(Northern Spain), 12 May 2006, sagittal sections on six slides. CRBA-3777a-b,
874 M. Vila-Farré et al.
Figure 3. Microplana groga. Specimen from El Molar, Montseny: (A) living animal, (B) sagittal
reconstruction of the copulatory apparatus; anterior to the left.
Etxalar, close to an irrigation channel, Navarra (Northern Spain), 12 May 2006, sagit-
tal sections on two slides. CRBA-3778, Irurita, in the shore of the river Artesiaga,
Navarra (Northern Spain), 14 May 2006, sagittal sections on one slide.
Comparative discussion
This species was described from Monaco on the basis of only one specimen
(Heinzel 1929). Minelli (1977a) considered M. monacensis to be a junior synonym
of M. scharffi. However, Jones et al. (2008), after the re-analysis of the type specimen,
pointed out the validity of the species. The species, whose external appearance has not
previously been described, is characterized by a well-developed penis bulb into which
the vasa deferentia enter separately and, subsequently, unite to form the ejaculatory
duct, which is surrounded by a very thick layer of circular muscle fibres. The narrow
bursal canal is lined with a ciliated, nucleated epithelium. After it has received the
separate openings of the oviducts, the canal curves dorsally and communicates with
the copulatory bursa, which communicates through a short genito-intestinal duct with
the gut.
We have collected specimens of M. monacensis from two localities in Navarra,
Etxalar and Irurita (Table 1, Figure 1). In elongated state the living, sexually mature
specimens measured 20 mm in length, with a width of about 2 mm (Figure 4A).
The dorsal surface is orange and the anterior end is dark brown. The very wide
creeping sole occupies about two-thirds of the body width. The internal anatomy
of these specimens has been compared with that of the type material housed in the
Naturhistorisches Museum Wien in Vienna.
We consider these specimens to belong to M. monacensis, albeit that they exhibit
a few differences when compared with the type material from Monaco. First, in the
anteroventral part of the body of the Iberian specimens (from about half way along
the pharynx to the anterior tip) there is a thick and conspicuous zone of tissue absent
in the holotype. It consists of mesenchyme interspersed with many cavities. The zone is
located in the lateral regions of the body and lies between the ventral epidermis and the
mesenchymal layer of longitudinal muscles (Figure 4B). We believe that this zone of
tissue corresponds to what has been described in the older literature for other species
Journal of Natural History 875
Figure 4. Microplana monacensis. (A–D) Specimens from Navarra: (A) living animal.
(B,C) Specimen ZMA V.Pl. 6856.1: (B) zone with spongy mesenchyma in the anteroventral
part of the body; (C) sagittal reconstruction of the copulatory apparatus; anterior to the left.
(D) ZMA V.Pl. 6856.1, extension of copulatory bursa approaching the ventral epidermis.
876 M. Vila-Farré et al.
as the spongy structure of parts of the ventral mesenchyma (“Bindegewebslücken”; cf.
Von Graff 1912–17: 2762). Second, the vasa deferentia open separately into a small
bulbar cavity (Figure 4C) surrounded by two distinct layers of circular muscle fibres
of different orientation. This cavity is not depicted in the reconstruction provided by
Heinzel (1929). Third, the gonopore lies under the middle of the ventral wall of the
male atrium, whereas it is placed in the common atrium in the animal from Monaco.
Fourth, the oviducts open close to the anterior section of the bursal canal. In the
type specimen they open in the central section of the bursal canal. Fifth, in specimens
ZMA V.Pl. 68.56.1 and CRBA-3777a-b the copulatory bursa extends laterally (as an
elongated tube) and eventually seems to open to the exterior, but the poor state of
the tissue prevents a proper observation of this feature. This presumed “opening”,
not observed in the type specimen, occurs on the dorsolateral side in one animal but
ventrally in the other (Figure 4D).
We consider that our specimens belong to M. monacensis despite the observed
anatomical differences. However, because of the lack of material from the type locality
we cannot rule out the possibility that our specimens belong to a different species. We
will be able to clarify this point only if new material from the type locality becomes
available.
The new records considerably extend the distribution range of the species, as it was
previously only known from Monaco.
Microplana robusta Vila-Farré and Sluys, sp. nov.
(Figures 1, 5, Table 1)
Material examined
Holotype. ZMA V.Pl. 6857.1, Parque Natural Fragas do Eume, province of A Coruña
(North We stern Spain), 3 July 2003, sagittal sections on 13 slides.
Other material. CRBA-3776a-j, Parque Natural Fragas do Eume, province of
A Coruña (North Western Spain), 12 April 2006, sagittal sections on 10 slides.
Etymology
The specific epithet is derived from the Latin adjective robustus, meaning strong, firm,
and alludes to the relatively large body size.
Diagnosis
Microplana robusta sp. nov. can be distinguished anatomically from its congeners by
a muscular penis bulb provided with intermingled longitudinal and circular muscle
fibres, short and cylindrical penis papilla, very short genito-intestinal duct t hat links
the long and narrow female genital duct to the gut, and absence of a copulatory bursa.
Description
In elongated state, living sexually mature specimens up to 50 mm long, 2–3 mm
wide (Figure 5A). Preserved holotype specimen 12.9 mm long, as determined from
Journal of Natural History 877
Figure 5. Microplana robusta. (A–D) Specimens from Fragas do Eume: (A), extended liv-
ing animal during movement; (B) living animal, resting; (C) diagrammatic transverse section
through the pharynx, showing the arrangement of the rows of longitudinal and circular mus-
cles; (D) holotype ZMA V.Pl. 6857.1, sagittal section of the copulatory apparatus; anterior to
the left. (E) Holotype ZMA V.Pl. 6857.1, sagittal reconstruction of the copulatory apparatus;
anterior to the left.
histological sections. Cylindrical body (flattened when resting, Figure 5B) tapers ante-
riorly and posteriorly to form blunt points. Dorsal surface orange with a lighter
anterior end and a darker posterior end in ZMA V.Pl. 6857.1. Creeping sole reaches
anterior end. Two black eyes (eye cup diameter 59 µm in sections) located laterally at
a short distance anterior to the brain. Numerous longitudinal muscle fibres present in
ventral body region.
Short cylindrical pharynx about one-tenth of total body length (0.9 mm in
sections) in holotype and placed in a central, almost horizontal position. Outer epithe-
lium of pharynx ciliated and underlain by layer of longitudinal muscles, followed by
layer of circular muscles. Thin layer of longitudinal muscle fibres under inner pharynx
epithelium, followed by intermingled layers of circular and longitudinal muscle fibres
and a layer of longitudinal fibres (Figure 5C). Mouth located approximately in middle
of pharyngeal pouch, 5 mm from tip of body in holotype. Spherical structure, origin
unknown, bounded by circular muscle fibres, in intestinal lumen of holotype.
Very thick subintestinal mesenchymal transverse muscle layer in holotype, dorsal
to brain and ventral nerve cords, ventral to intestine and continuing into posterior
end, where it becomes weaker.
More than eight testes situated on either side of body in specimen CRBA-3776a-
j. Small rounded, or oval-shaped, irregularly-sized follicles occupy approximately
one-fifth of the dorsoventral diameter of body, arranged in ventral longitudinal rows,
extending anteriorly from about root of pharynx to an unknown distance (because
frontal region of CRBA-3776a-j removed for use in DNA analysis). Sperm present in
878 M. Vila-Farré et al.
vasa deferentia; testes not observed in holotype. Ovaries not discovered in specimens
examined.
Strongly muscular, spherical penis bulb (Figure 5D) with intermingled longitu-
dinal and circular muscle fibres, especially well-developed in specimen CRBA-3776.
Narrow vasa deferentia run directly dorsally to the ventral nerve cords and open
separately into ejaculatory duct, which opens at tip of penis papilla (Figure 5E).
Ejaculatory duct lined with nucleated epithelium and surrounded by thick layer of
circular muscle fibres stronger at base of duct. Short and conical penis papilla of holo-
type has relatively broad base and an oblique, dorsoventral orientation. Conical penis
papilla larger in CRBA-3776a-j. Outer wall of penis papilla covered with thin, nucle-
ated epithelium, underlain with thin layer of circular muscles, bounded by layer of
longitudinal muscles. Genital atrium lined with nucleated epithelium underlain with
subepithelial circular muscle layer, followed by layer of longitudinal muscle. Gonopore
1.7 mm from mouth.
Rather narrow female genital duct with distinct cilia, openings of oviducts at
posterior end. Anterior end of female genital duct projects slightly into atrium in holo-
type. Duct lined with nucleated epithelium underlain with thick, subepithelial layer of
circular muscles, followed by layer of longitudinal muscle fibres. Shell glands open
into f emale genital duct at same level as oviducts. Very short genito-intestinal duct
arises from distal end of female genital duct (Figures 5D,E) and immediately com-
municates with gut. Genito-intestinal duct lined with ciliated epithelium by layer of
circular muscle.
Discussion
The absence of ovaries in the sectioned specimens suggests that they were not fully
mature. Despite this we suspect that the gross morphology of the copulatory appa-
ratus will not change substantially after maturity because the copulatory apparatus
of the type specimen has all the typical elements of the copulatory apparatus of a
member of the genus Microplana, including the genito-intestinal duct. This species
can be distinguished from other native European land planarians by a combination
of external features and anatomical characteristics of the genital apparatus. The dor-
sal colouration pattern of M. robusta resembles that of M. pyrenaica (Von Graff,
1893), M. scharffi and M. monacensis. Some individuals of M. groga can probably
also exhibit a similar colouration pattern. However, M. pyrenaica is provided with an
enormous penis bulb with a large intra-bulbar cavity and an extremely reduced penis
papilla. In contrast, the penis bulb of M. robusta is much smaller than in M. pyrenaica
and an intra-bulbar cavity is absent. Furthermore, its penis papilla is well developed.
In addition, the female genital duct of M. pyrenaica is wider and almost horizontal
and its genito-intestinal duct arises from its central section, whereas in M. robusta
this duct is very narrow and has an oblique ventro dorsal disposition. In M. robusta
the genito-intestinal duct communicates with the posterior end of the female genital
duct, whereas in M. pyrenaica the middle section of the female duct gives rise to the
genito-intestinal duct (cf. Heinzel 1929).
Microplana scharffi is yellow or cream when starved (McDonald and Jones 2007).
However, it is an extremely long and thin planarian (up to 9 cm), in contrast to
M. robusta, which is shorter and thicker. With respect to anatomical features, the type
specimen of M. scharffi has a very short genito-intestinal duct and possesses a copula-
tory bursa. Although a very short genito-intestinal duct is also present in M. robusta,
Journal of Natural History 879
the latter does not have a copulatory bursa. Further more, in M. scharffi the vasa defer-
entia fuse half-way along the elongated penis papilla, whereas in M. robusta the ducts
fuse within the penis bulb and the penis papilla is conical.
In M. monacensis a copulatory bursa is present (Heinzel 1929), whereas such a
structure is absent in M. robusta. In addition, the penis papilla of M. monacensis is
elongated, but it is conical in M. robusta.
Although M. groga was described originally as being “reddish to brown” (Jones
et al. 2008) we have also collected cream-coloured specimens similar to M. robusta.
However, M. groga is more slender than M. robusta. Regarding anatomical features,
in M. groga the vasa deferentia open separately at the base of the broad ejaculatory
duct, close to the base of the penis bulb. In contrast, in M. robusta the ducts open
separately at the base of a narrow ejaculatory duct, close to the base of the short
and conical penis papilla. In addition, in M. groga there is a short, horizontal female
genital duct (= vagina) of the same length as the genito-intestinal canal, whereas in
M. robusta the very long female genital duct is oblique and the genito-intestinal canal
is extremely short.
Among the known European species of Microplana a genito-intestinal duct
has also been reported for the following species: M. humicola Vejdovsky, 1889;
M. pyrenaica; M. giustii Minelli, 1976; M. henrici (Bendl, 1908); M. attemsi (Bendl,
1909); M. peneckei (Meixner, 1921); M. scharffi; M. monacensis; M. terrestris; M. nana;
M. kwiskea Jones et al., 2008; M. groga; M. gadesensis Vila-Farré et al. 2008.
Comparisons with the species M. pyrenaica, M. scharffi, M. monacensis and M. groga
have been presented above. Microplana humicola is a small species (4 mm) that presents
a hyaline colouration (cf. Schneider 1935), instead of the pigmented dorsal coloura-
tion pattern of the much larger M. robusta. Microplana humicola has two testes on
either side of the body and its penis papilla is elongated, in contrast to the abundant
testes and short conical penis papilla of M. robusta.InM. robusta the genito-intestinal
canal is also shorter. Microplana giustii is a black species with post-pharyngeal testes
(cf. Minelli 1976), contrasting with the orange-cream colouration of
M. robusta, with
its prepharyngeal testes. Microplana henrici, M. attemsi and M. peneckei present atrial
folds (cf. Minelli 1977a), whereas these are absent in M. robusta.InM. terrestris a
bursa is present, whereas such a structure is absent in M. robusta.InM. kwiskea (cf.
Jones et al. 2008) the very elongated penis papilla houses a wide ejaculatory duct; the
genito-intestinal canal is long. In M. robusta, in contrast, the penis papilla is short
and conical and the genito-intestinal canal is very short. Microplana gadesensis has
an elongated penis papilla and a very long genito-intestinal duct (cf. Vila-Farré et al.
2008), contrasting with the short and conical penis papilla and the very short genito-
intestinal duct of M. robusta. While in M. nana the penis papilla is elongated and
the penis bulb is narrow, in M. robusta the penis papilla is short and conical and the
spherical penis bulb is wide. The genito-intestinal canal is longer in M. nana than in
M. robusta.
A female copulatory apparatus provided with a genito-intestinal canal is found
only in two continental African species of Microplana, M. harea (Du Bois-Reymond
Marcus, 1953) and M. termitophaga Jones et al., 1990. In M. harea (1) the testes extend
to the level of the copulatory apparatus, (2) the almost spherical penis bulb is pro-
vided with a distinctive seminal vesicle and a reduced penis papilla, (3) the common
oviduct meets the female genital duct, with approximately the same length as the
genito-intestinal duct, at its distal part; all of these features are different in M. robusta.
With respect to external features, M. harea is dorsoventrally flattened and “the back
880 M. Vila-Farré et al.
is dark-gray with a fine, partially faded black middle stripe ...” and a “light area lies
between the small eyes ...” (Du Bois-Reymond Marcus 1953), contrasting with the
cylindrical body and the uniform orange-cream dorsal surface that is devoid of stripes
in M. robusta.
Microplana termitophaga is very similar to M. harea with only one important
exception that is relevant for the present discussion: the genito-intestinal duct of
M. termitophaga is extremely short. However, the differences between M. harea and
M. robusta pointed out above are also valid for M. termitophaga.
Among the known species of Microplana from Madagascar and neighbour-
ing islands a genito-intestinal duct has been reported for M. mediostriata (Geba,
1909), M. trifuscolineata (Kaburaki, 1920) (Mauritius) and M. tristriata (Geba, 1909)
(Comores). Unfortunately, the copulatory apparatus of M. gebavoeltzkowi Ogren and
Kawakatsu, 1988 (Comores) has never been described. Although M. mediostriata,
M. striata and M. trifuscolineata have a genito-intestinal duct the gross morphology
of their copulatory apparatus is completely different from that of M. robusta, notably
the presence of a penis sheath in the three first-mentioned species. Furthermore,
in M. tristriata the meeting point of the oviducts is far removed from the genito-
intestinal canal, whereas in M. robusta the point of communication of the oviducts
with the female genital duct coincides with the origin of the genito-intestinal canal.
Microplana gebavoeltzkowi is dirty yellow with a broad grey-brown median band,
whereas M. robusta is orange and without a median band.
Among the known Asian species of Microplana a genito-intestinal duct has only
been reported for M. unilineata (Frieb, 1923) and M. uniductus (De Beauchamp, 1930).
Microplana unilineata presents a mid-dorsal stripe, which is absent in M. robusta.
Regarding anatomical features, the penis bulb of M. unilineata houses a bulbar cav-
ity and the penis papilla is relatively long; the genito-intestinal duct is also relatively
long. In contrast, in M. robusta, there is no seminal vesicle and the penis papilla and
genito-intestinal duct are short. In addition, M. uniductus presents an elongated penis
bulb and a common oviduct, whereas in M. robusta the penis bulb is rounded and a
common oviduct is absent.
Microplana teres (Von Graff, 1899), also from Asia, is described on the basis
of an immature specimen, showing a conical, slender penis papilla. In contrast, in
M. robusta the penis papilla is not slender. The external colouration pattern is also
different in both species.
The Asian Microplana ruteocephala Kaburaki, 1922 is “uniformly light black with
a slight touch of reddish tint, in front grading over into a red or yellow tone at the
tip of the body. Extending over the surface is a fine black median stripe, which loses
itself in front” (Kaburaki 1922). In contrast, M. robusta
is uniformly orange cream. As
the description of M. ruteocephala is only based on external morphology we cannot
compare its anatomy with that of M. robusta.
Four species of the genus Microplana have been recorded from North America,
two of which (M. scharffi and M. terrestris) have been discussed above. The third
species, M. rufocephala (Hyman, 1954) is a long, black species provided with a long
genito-intestinal duct, contrasting with the features of M. robusta. Microplana atro-
cyanea (Walton, 1912), the fourth North American member of the genus, is a black
planarian with an elongated penis papilla and a long genito-intestinal duct (cf. Hyman
1954) and therefore also differs from M. robusta.
Journal of Natural History 881
Among the known Microplana species of South and Central America a genito-
intestinal duct has been reported for the following species: M. yaravi (Du Bois-
Reymond Marcus, 1957), M. montoyai (Fuhrmann, 1914) and M. cockerelli (Von
Graff, 1989). However, M. yaravi is up to 20 mm long, provided with two broad
longitudinal stripes and a copulatory bursa (“uterus” in Du Bois-Reymond Marcus
1957).
Mature specimens of M. montoyai are up to 40 mm long, with black dorsal
colouration, a white mid-dorsal stripe, a big bulbar cavity, and a short genito-intestinal
duct that arises from t he middle of the roof of the female genital duct (“vagina” in
Fuhrmann 1914). In contrast, M. robusta has orange dorsal surface, while a mid-
dorsal stripe is absent. Regarding anatomical features, a bulbar cavity is absent in
M. robusta and its genito-intestinal duct arises from the distal end of the genital duct.
Microplana cockerelli (Von Graff, 1899) has a bluish-black colouration with both
body ends sharply marked off from the rest of the body by a different, lighter coloura-
tion; it also has a pale mid-dorsal longitudinal stripe, two seminal vesicles, one of them
a presumed “prostatic organ” (Prudhoe 1949), and a relatively long genito-intestinal
canal, all being features that are different from those in M. robusta.
Specimens from las Fragas do Eume, with the external morphology of M. robusta,
have been studied f rom a molecular point of view by Mateos et al. (2009). One of the
animals analysed in the present paper, CRBA-3776a-j, was part of that study and its
sequence is available from GenBank (accession number, COI: FJ969962, 18S Tipus II:
FJ969990). In the study of Mateos et al. (2009) these specimens were grouped together
as morphotype M3, which formed a molecular clade consisting solely of specimens
from this locality. Although Mateos et al. (2009) included land planarians from several
areas of the Iberian Peninsula, this species did not occur at other localities and, conse-
quently, it is probably restricted to a few areas with special and appropriate ecological
conditions.
Microplana hyalina Vila-Farré and Sluys, sp. nov.
(Figures 1, 6, Table 1)
Material examined
Holotype. ZMA V.Pl. 6858.1, Montsec, camí del Sant del Bosc, province of Lleida,
Spain, 9 January 2002, sagittal sections on three slides.
Other material. ZMA V.Pl. 6859.1, Montsec, camí del Sant del Bosc, province of
Lleida, Spain, 11 May 2008, sagittal sections on one slide.
Etymology
The specific epithet is derived from the Greek adjective hyalinos, like glass, and alludes
to the hyaline colouration of the body.
Diagnosis
A small, hyaline Microplana species up to 5 mm long, with spherical penis bulb pro-
vided with a strong musculature and a distinct bulbar lumen; short and vertically
oriented penis papilla; atrium divided into a cup-shaped cavity and a tubular distal
882 M. Vila-Farré et al.
cavity; bursal canal with a distinctive, initially posteroventrally oriented loop-like
curvature communicating with the posterolateral section of the copulatory bursa.
Description
Living, sexually mature specimen ZMA V.Pl. 6858.1 about 5 mm long and about
0.5 mm wide, in elongated state (Figure 6A). Cylindrical body tapers anteriorly to
bluntly rounded point; posterior end also bluntly pointed. Dorsal and ventral body
surfaces hyaline. Colouration and tiny size of preserved specimens prevented adequate
observation of creeping sole. Two black eyes (eye cup diameter 19 µm in sections) a
short distance anterior to brain.
Figure 6. Microplana hyalina. (A–D) Specimens from Sant del Bosc: (A) living animal. (B–D)
holotype, ZMA V.Pl. 6858.1; (B) sagittal reconstruction of the copulatory apparatus; anterior
to the right. (C, D) sagittal sections of the copulatory apparatus; anterior to the right.
Journal of Natural History 883
Subepidermal longitudinal fibres of body musculature weak. In ventral body
region numerous longitudinal fibres distributed in two mesenchymal bands, especially
strong over and under ventral nerve cords. Dorsal longitudinal mesenchymal fibres
run over intestine.
Short, cylindrical pharynx about one-eighth of body length, situated in posterior
half of animal, in almost horizontal position. Outer epithelium of the pharynx ciliated,
underlain by thin layer of longitudinal muscles, followed by thicker layer of circular
muscles. Very thick outer layer of circular muscles underneath inner pharynx epithe-
lium, intermingled with inner layer of longitudinal muscles fibres. In specimen ZMA
V.Pl. 6859.1 posterior gut trunks meet at posterior end of body.
Mouth opening located at about one-third of distance between posterior wall of
pharyngeal pouch and root of pharynx. In holotype mouth situated 2.1 mm from
anterior end of body and 0.46 mm from gonopore.
About 12 ellipsoidal testes on each side of body. Follicles situated ventrally, occu-
pying approximately one-quarter of dorsoventral diameter of body. Testes arranged
in longitudinal rows, extending from some distance behind ovaries almost to root of
pharynx.
Vasa deferentia narrow at level of penis before entering penis bulb, opening sepa-
rately into lateral part of bulbar cavity (Figure 6B). Strongly muscular, spherical penis
bulb consists of intermingled longitudinal and circular muscle fibres. Bulb houses
spherical and very spacious bulbar lumen (Figure 6C) that narrows considerably
before communicating with ejaculatory duct. Bulbar lumen and ejaculatory duct lined
with nucleated epithelium underlain with layer of circular muscle fibres.
Vertically oriented penis papilla short and conical. Papilla covered with a thin,
nucleated epithelium underlain with very thick, subepithelial layer of circular muscles,
bounded by thick layer of longitudinal fibres. Male atrium consists of a dorsal cup-
shaped cavity and a lateral tubular part with diagonal orientation. Lining epithelium
of atrium underlain with subepithelial circular muscle layer, thickened at tubular part,
followed by thin layer of longitudinal muscles.
Ovaries situated above ventral nerve cords, occupying about one-quarter of
dorsoventral diameter of the body, positioned between one-quarter and one-sixth
of distance between brain and root of pharynx. Oviducts arise from ventral side of
ovaries, where spermatozoa are observed. Oviducts run backwards laterally to ventral
nerve cords; ducts turn dorsally and open separately into bursal canal.
Copulatory bursa spherical, lined with layer of tall vacuolated cells; not connected
with gut. One specimen, ZMA V.Pl. 6859.1, had in its copulatory bursa remnants of an
irregular structure (diameter 26 µm), presumably of a sclerotic nature. Origin of this
structure not found in penis or atrial glands, but colour and texture suggest a sclerotic
substance, suggesting that it may be part of a spermatophore. Sperm also present in
this bursa. Secondary bursa-like structure not connected to the copulatory bursa nor
to copulatory apparatus (Figure 6B) present in holotype. Shell glands open into bur-
sal canal mostly ventrally to openings of oviducts; bursal canal lined with nucleated
and ciliated cells and surrounded by subepithelial layer of circular muscles and some
scattered longitudinal muscle fibres. Bursal canal runs obliquely, i.e. posterodorsally,
towards copulatory bursa from point of communication with atrium. After open-
ings of oviducts, canal runs lateral to bursa with distinctive, initially posteroventrally
oriented, loop-like curvature (Figure 6B,D) before communicating with posterolateral
section of copulatory bursa.
884 M. Vila-Farré et al.
Discussion
Among the approximately 23 species of native land planarians known from Europe,
two of which are described as new in the present paper, M. hyalina stands apart from
the other species by the unique anatomy of its bursal canal, as well as by a unique
combination of its internal and external characters. A hyaline body colouration is
also found in M. humicola and M. aixandrei Vila-Farré et al., 2008. Microplanana
humicola, however, has a greenish anterior end, in contrast to M. hyalina. Regarding
anatomical features, M. humicola has two pairs of dorsal testes and a genito-intestinal
duct but a copulatory bursa is absent (cf. Schneider 1935). In contrast, M. hyalina
presents 12 pairs of ventral testes and a copulatory bursa, but lacks a genito-intestinal
duct.
Microplana aixandrei, a species endemic to the far southern part of the Iberian
Peninsula but probably present also in other areas, is similar in size and colouration to
M. hyalina and also produces spermatophores. The former has two testes on each side
of the body, in contrast to M. hyalina, which has 12 testes on each side. In M. aixandrei
the bursal canal is an obliquely running structure that communicates with the anterior
section of the irregularly sac-shaped copulatory bursa; a sphincter is present in the
proximal section of the canal. In contrast, in M. hyalina the canal runs laterally to
the bursa and also shows a distinctive, initially posteroventrally oriented, loop-like
curvature before communicating with the posterolateral section of the oval-shaped
copulatory bursa; a sphincter is absent. The outer epithelium of the pharynx is ciliated
only at the posterior part of the pharynx in M. aixandrei but completely ciliated in
M. hyalina.
In addition to M. aixandrei, a copulatory bursa devoid of any connection with
the intestine occurs also in M. howesi (Scharff, 1900), M. mahnerti Minelli, 1977,
M. styriaca (Freisling, 1935) and M. grazalemica Vila-Farré et al. 2008. However, in
these species the bursal canal communicates with the anterior section of the copulatory
bursa, whereas in M. hyalina the bursal canal communicates with the copulatory bursa
through its posterolateral section. The gross morphology of the copulatory apparatus
and the external appearance in these species are also very different from M. hyalina.
A dorsal connection of the bursal canal with the copulatory bursa is also present
in M. monacensis (Heinzel, 1928). However, in M. monacensis a genito-intestinal
duct is present, whereas it is absent in M. hyalina. In addition, the horizontal
penis papilla of M. monacensis is elongated but is conical and vertically inserted in
M. hyalina.
Among the nine native species of Microplana from Africa, Madagascar and neigh-
bouring islands, none has the external colouration and anatomy of the copulatory
bursa and bursal canal of M. hyalina.
Unfortunately, M. gebavoeltzkowi from Madagascar is known only from its exter-
nal appearance, namely dirty yellow with a broad grey-brown median band, which
contrasts with the body colouration of M. hyalina.
Only one Asian species, Microplana indica (Chaurasia, 1985), possesses a copula-
tory bursa without intestinal connections (cf. Chaurasia 1985). The creamy dorsal
surface of this species is marked with black spots making up a dorsal band. In
contrast, in M. hyalina
the dorsal surface is uniformly hyaline. Regarding anatomical
features, M. indica has post-pharyngeal testes and a cylindrical penis papilla, whereas
M. hyalina has pre-pharyngeal testes and a conical penis papilla.
Journal of Natural History 885
Microplana teres, also from Asia, is described on the basis of an immature spec-
imen with a slender penis papilla, contrasting with the short and blunt papilla of
M. hyalina. Furthermore, the external colouration pattern is also different in both
species.
The internal anatomy of the Asian M. ruteocephala is unknown but its external
appearance differs much from M. hyalina (see above).
Only one American species has a copulatory bursa that is not linked to the intes-
tine, M. costaricensis (De Beauchamp, 1913). This species has a very long cylindrical
penis and a bursal canal that communicates with the ventro-anterior section of the
bursa. In contrast, in M. hyalina the penis is conical and the bursal canal opens at the
posterolateral section of the copulatory bursa.
Various kinds of secretions may be associated with the sperm (cf. Souza and Leal-
Zanchet 2004) but only very rarely in land planarians is the ejaculate enveloped by
a true spermatophore of a clear sclerotic nature (Sluys 1989b and references therein;
Winsor 1998). Microplana hyalina is only the third species of land planarian for which
such a spermatophore is described (Winsor 1998). It has been suggested that in other
species of land planarian other types of secretion, which do not completely surround
the spermatozoa, may still function as spermatophore, facilitating the transfer of
spermatozoa (Souza and Leal-Zanchet 2004).
Microplana nana Mateos, Giribet and Carranza, 1998
(Figures 1, 7, Table 1)
Material examined
Holotype. NHML 1998.2.9.1, sagittal sections of one specimen on three slides
(labelled LA 2.1, 2.2, 2.3).
Paratype. NHML 1998.2.9.2, sagittal sections of one specimen on two slides (labelled
LA1.1, 1.2). For further details see Mateos et al. (1998).
Description
For details of external features, epidermis, musculature, nervous system and alimen-
tary system, see Mateos et al. (1998).
Testes as described by Mateos et al. (1998), discharge into sperm ducts (vasa
deferentia), which run posteriorly to copulatory apparatus. Sperm ducts not readily
discernable in sections but must be very narrow. (Duct labelled “D” on fig. 5 of Mateos
et al. 1998, is ovovitelline duct, not sperm duct.) Anterior to copulatory apparatus,
each sperm duct expands to about 40 µm in diameter and forms a sperm storage struc-
ture (spermiducal vesicle) on either side. They then narrow (duct is occluded but visible
over most of its length, but appears to be about 2.5 µm in diameter) and separately
enter penis bulb. Penis with basal diameter about 170–200 µm and total (retracted)
length about 450 µm. In both specimens, the distal end, about 150 µm, is reflected
with diameter about 40 µm. Sperm ducts discharge separately but close together into
anterior end of ejaculatory duct. Ejaculatory duct eccentric, in ventral half of penis,
maximum diameter about 50 µm, lined with tall columnar cells, about 5 µm diameter
in distal, reflected, portion.
886 M. Vila-Farré et al.
Figure 7. Microplana nana. (A) NHML 1998.2.9.1, sagittal section of the copulatory apparatus;
anterior to the right. (B) NHML 1998.2.9.2, sagittal section of the copulatory apparatus;
anterior to the right. (C) NHML 1998.2.9.1, sagittal reconstruction of the copulatory apparatus;
anterior to the right. (D) NHML 1998.2.9.2, sagittal reconstruction of the copulatory
apparatus; anterior to the right. Scale bars represent 200 µm in all figures.
Ovaries, oviducts and vitellaria as described by M ateos et al. (1998). Oviducts
extend posterior to penis and converge to separately join posterior extremity of female
genital duct. Duct short, about 50 µm long and Y-shaped. Base of Y opens into pos-
terior of atrium, each upper arm of the Y receiving one ovovitelline duct. Narrow
genito-intestinal duct (Figure 7) runs dorsally from female genital duct to branch of
intestine. Many eosinophilic glands, presumably “shell gland” secreting the cocoon
material, surround region around female genital duct.
Discussion
The type material has been re-examined because details of the copulatory appara-
tus differ f rom the description and diagram given by Mateos et al. (1998: fig. 2) and
accordingly the species is partially re-described above, as far as is necessary to correct
the details.
The differences between the description of Mateos et al. (1998) and the present
description are as follows. First, the sperm ducts enter the penis separately and remain
separate in the base of the penis until they discharge separately into the ejaculatory
duct. They are shown in fig. 2 of Mateos et al. (1998) fusing before entering the penis
bulb and discharging into the ejaculatory duct as a single duct. Second, the female
Journal of Natural History 887
genital duct i s about 50 µm long, whereas it is shown as about 300 µm long in fig. 2
of Mateos et al. (1998). Third, there is a genito-intestinal duct running dorsally from
the female genital duct. This was stated to be absent by Mateos et al. (1998).
Genus Rhynchodemus Leidy, 1851
Rhynchodemus sylvaticus (Leidy, 1851)
(Figure 1, Table 1)
Discussion
As was predicted in a previous paper (Vila-Farré et al. 2008) new sampling has yielded
one new locality in southern Spain, la Axarquía (Table 1, Figure 1), where this species
has been found under a flowerpot in a mango plantation. The origin of this popula-
tion, native or introduced, is uncertain because of the anthropogenic character of the
habitat.
Exotic species
Material examined
CRBA-3781a-b, Benamargosa, la Axarquía. province of Málaga (southern Spain),
25 December 2007, sagittal sections on two slides. CRBA-3782a-b, ibid., sagittal
sections on two slides.
Discussion
In the same mango plantation where some individuals of R. sylvaticus were found,
under the next flowerpot, we collected a number of asexual land planarians. They were
characterized by an external morphology very similar to that of the former species
but with only one dorsal line instead of the two of R. sylvaticus. Molecular analy-
sis performed by Marta Álvarez (personal communication) placed these individuals
within the genus Platydemus. Because of the anthropogenic origin of the habitat and
as Platydemus is mainly found in the Indo-Pacific region, with only one African species
(Jones 1998), we tentatively consider these animals to be an exotic component of the
Iberian fauna.
Ecology of Iberian terrestrial planarians
Land planarians are more sensitive than most other organisms to the humidity
of their environment because they lack any special mechanism of water conserva-
tion (Kawaguti 1932). The animals are not good burrowers because of the absence
of any fluid-filled body cavities so they have limited access to humid environ-
ments in deeper layers of the soil. However, the worms may use cracks and voids
made by other organisms to retreat to a certain depth in the soil and so avoid
desiccation.
The Iberian Peninsula is an area with an Atlantic climate in its northern sector
and a Mediterranean one in the other parts, with populations of land planarians
being widely distributed throughout the Peninsula (cf. Vila-Farré et al. 2008). Of
the 58 records of land planarians properly georeferenced, at least 59% are located
888 M. Vila-Farré et al.
in the proximity of a river or stream. The constant humidity provided by the water-
courses probably enables the establishment of populations in areas that are regularly
affected by long dry periods, i.e. the Mediterranean zone of the Iberian Peninsula. In
this region the animals usually occur in very low numbers (one to four animals col-
lected per survey journey, with two collectors searching the under surface of stones for
6 hours), and the specimens are also rather small and frequently white. Generally, big-
ger specimens have been collected from the Atlantic region of the Iberian Peninsula,
e.g. large specimens of M. terrestris and M. robusta from Galicia and unidentified
specimens up to 8 cm long from Navarra.
We have only observed relatively large concentrations of animals (10 or more
animals collected per survey journey) in the Sierra de Grazalema (southern Spain;
M. aixandrei) and in Sant del Bosc (northeastern Spain; M. hyalina). Both species
are very small, which is probably an advantage in Mediterranean areas that are sub-
jected to periodic drought. In areas where watercourses are absent (for example in
the Sant del Bosc, a slope in a holm oak forest with abundant vegetation and high
humidity; type locality of M. hyalina) and during long dry periods (from 2006 to April
2008) planarians can suddenly reduce their population size to such an extent that it
is impossible to find any specimens, although this area was visited each year three or
more times in spring, summer and autumn. Seasonal rarity may be the result of retreat
into less accessible crevices rather than of population shrinkage. After a rainy period
(May 2008), however, planarians were found again in large numbers (more than 10
individuals per survey journey in 2009, with two collectors searching the under sur-
face of stones for 3 hours), suggesting that the populations of M. hyalina are well
adapted to a Mediterranean climate. It is also possible that because of their small size
the animals could be hidden during dry periods and that they re-appear rapidly when
rain comes.
We have collected land planarians mostly in the vicinity of deciduous trees (oak,
holm oak, beech and gallery forests) under stones and, occasionally, also under tree
trunks, where they are more difficult to detect. We have never observed cocoons.
Although some European species have been collected in coniferous or mixed forests
(e.g. M. manherti Minelli, 1977, collected in the Jura Mountains in a mixed wood-
land with Carpinus betulus L. and Picea glauca (Moench) (cf. Minelli 1977b), in Spain
they have never been found in coniferous forests. We have only one record of a small
white land planarian in a wheat field with some Pinus trees around. As the wheat field
environment is extremely dry it may be that the animal was transported by recent
rains from adjacent areas or even may have actively moved to the area under humid
conditions.
Froehlich (1955) pointed out that land planarians cannot endure long periods of
submersion in water. Nevertheless, we have often collected specimens from the banks
of rivers in areas that are frequently flooded. One of the species, R. sylvaticus,on
one occasion was even collected from a temporary pool (Clot d’Espolla, Girona,
Northeastern Spain; Boix and Sala 2001). This species, although rare, is present in
a wide variety of ecosystems in Spain: river banks (two records); dry areas in villages
(Talarn, northeastern Spain; but only in very rainy periods and close to a very narrow
temporary water canal); seasonal streams (two records); plant nurseries (one record);
and mango plantations (one record).
Journal of Natural History 889
General discussion
In this paper we have followed the classification for the triclad flatworms published by
Sluys et al. (2009). In the comparative analyses of the new species presented above we
have included all the known taxa of the genus Microplana. We considered it particu-
larly important to analyse African species because the fauna of the Iberian Peninsula
has a strong African component (Ribera 2000; Pleguezuelos et al. 2008). In addition,
the accidental (Jones 1998) or voluntary (Kawakatsu et al. 1992) introduction of land
planarians is a well-known factor of dispersal in this group of animals. Therefore, an
alien origin of the new Iberian species has been discounted after comparison with all
currently known members of the genus.
A general overview of the distribution of Iberian land planarians has been
provided by Vila-Farré et al. (2008) and Mateos et al. (2009). The present paper
complements these analyses by reinforcing some of their conclusions and predictions.
First, Iberian populations composed of individuals of similar external morphology
and located in geographically distant areas, belong in fact to different species (e.g.
M. aixandrei in the southern part and M. hyalina in the northeastern part of the
Peninsula). Second, similar to the situation found in other animal groups, the Iberian
fauna of terrestrial planarians exhibits a high degree of endemism. Furthermore,
European species such as M. terrestris and M. monacensis are present in at least
the northern sector of the Peninsula. We would also expect to find North African
species during future samplings in the southern sectors of Spain and Portugal. Third,
the study of the land planarians of the Iberian Peninsula is already yielding new and
interesting ecological (innovation in reproductive strategies like the use of a true sper-
matophore) and biogeographic (high degree of endemism) information. In addition,
populations of well known European species, such as M. terrestris, display on the
Iberian Peninsula particular morphological characteristics suggesting the presence
of interpopulation variability and/or cryptic species, a situation that may be anal-
ysed better with the help of molecular techniques. Fourth, although the presence of
introduced and widely distributed species, such as Arthurdendyus triangulatus (Dendy,
1895) (cf. Boag and Yeates 2001), is still not established for the Iberian Peninsula,
we have here recorded introduced land planarians in areas where continued human
activity allows for their introduction.
Acknowledgements
We are grateful to Sisco Monjo, Edgar Vila, David Fuses, Blanca Olivet, Sergi Rodríguez, Mar
Sancho Prat, Juan Pascual Fontiveros, María del Carmen Anaya Ruiz, Juan Pascual Anaya,
Jaime Pascual Anaya, Susana Lozano and Mónica Lima for their help in the field and to
Miguel Lozano Calderón, Juana Jiménez Fortes and Francisco Lozano Calderón for permitting
sampling activity on their properties (Valle-Iglesias, El Soto in La Axarquía, Spain). M.V.F. is
indebted to Kay Eckelt and Michael Lang for their help in the translation of German texts.
M.V.F. acknowledges financial support from SYNTHESYS, a programme of the European
Commission under the Sixth Research and Technological Development Framework Programme
“Structuring the European Research Area”, which enabled M.V.F. to work at the Zoological
Museum of the University of Amsterdam during May 2009 (grant number NL-TAF 5333).
This work is also supported by grant BFU2004-05015/BFI (to R.R.) from the “Ministerio de
Educación y Ciencia” of Spain. We are grateful to Dr H. Sattmann ( Naturhistorisches Museum
Wien) for providing a loan of the type material of Microplana monacensis.DrM.Riutort
890 M. Vila-Farré et al.
(University of Barcelona) and Marta Álvarez are thanked for information on the molecular
analysis of the introduced presumed Platydemus specimen. Completion of the manuscript was
made possible by a grant from the Netherlands Centre for Biodiversity Naturalis to R. Sluys.
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