taxonID	type	description	language	source
03DB7724E2048361FC8DFA98FB12FCEB.taxon	materials_examined	Material examined: Holotype: ZMA V. Pl. 7114.1, river just before opening into the sea, Nagos, Chios, Greece, 38 ° 33 ′ 32.31 ″ N, 26 ° 4 ′ 59.42 ″ E, 30 April 2010, coll. M. Vila-Farré, sagittal sections on seven slides. Paratypes: ZMA V. Pl. 7114.2, ibid., sagittal sections on six slides; V. Pl. 7114.3 (RS 221 - 3), ibid., horizontal sections on three slides. Other material: ZMA V. Pl. 7115.1, river, Nagos, Chios, Greece, 38 ° 33 ′ 27.57 ″ N, 26 ° 4 ′ 51.61 ″ E, 30 April 2010, coll. M. Vila-Farré, sagittal sections on five slides; V. Pl. 7115.2, ibid., sagittal sections on five slides; V. Pl. 7115.3, ibid., horizontal sections on three slides. Etymology: The specific epithet is derived from the Latin adjective effusus, generous, abundant, and alludes to the highly glandularized penis papilla. seminal vesicle; highly glandularized penis papilla; a bursal canal that widens considerably at its communication with the atrium; ectal reinforcement of the bursal canal confined to the vaginal region. ph 1 mm Diagnosis: Dugesia effusa is characterized by the combination of the following features: presence of a small, dorsal penial fold; central ejaculatory duct; short, valve-like diaphragm; large, intrabulbar Ecology and distribution: The species is known only from two sites in the same river, i. e. the type locality close to the opening into the sea and another site further upstream. Description: Preserved specimens up to 9 × 2.25 mm, with low-triangular head with rounded auricles; tail obtusely pointed (Fig. 3). Dorsal surface pale brown; ventral surface pale. Two eyes, situated in pigmentfree patches. Pharynx situated in the mid-region of the body, measuring between one-quarter and one-sixth of the body length. Mouth opening located at the posterior end of the pharyngeal pocket. The testes are located dorsally and extend from the level of the ovaries into the posterior end of the body. The vasa deferentia penetrate the ventro-lateral wall of the penis bulb and open into the seminal vesicle at a point very close to the diaphragm. The ovoid or bc vd 100 µm pear-shaped seminal vesicle fills the major part of the penis bulb and is lined with a columnar, nucleated epithelium. Through a very narrow diaphragm this seminal vesicle opens into the funnel-shaped, proximal section of the ejaculatory duct (Fig. 4). The short, stubby lips of the valve-like diaphragm, as well as the funnel-shaped section of the ejaculatory duct, receive the finely granular and dark red staining secretion of erythrophil penis glands. The broad ejaculatory duct follows a slightly ventrally displaced course through the penis papilla and opens at the blunt tip of the penis papilla, the actual opening being rather narrow. Along the major part of its length the lining epithelium of the ejaculatory duct is pierced by the numerous openings of abundant penis glands that produce an orange – brown secretion. The plug-shaped penis papilla is lined with a nucleated epithelium and is provided with a subepithelial layer of circular muscles, followed by a layer of longitudinal muscles. A penial fold is located symmetrically at the dorsal base of the penis papilla; the fold is traversed by some longitudinal muscle fibres. The ovaries are situated directly medially to the ventral nerve cords and are located at one-third to one-quarter of the distance between the brain and the root of the pharynx. The oviducts are lined with an infranucleated epithelium and are surrounded by a well-developed coat of circular muscles. The oviducts open separately into the ventral-most, widened section of the bursal canal, close to the point where the canal communicates with the atrium. Shell glands discharge their secretion into the bursal canal ventrally to the oviducal openings. The bursal canal is lined with a nucleated, cuboidal-columnar epithelium. The diameter of the bursal canal increases considerably near its point of communication with the atrium. Notably the most ventral section of the canal, at the level of the oviducal openings, shows a widening into posterior direction (Fig. 5). The bursal canal is overlain with a thin layer of circular muscles, the latter being particularly developed in the vaginal region. Ectal reinforcement in the form of outer longitudinal muscle fibres is present in the vaginal area and extends towards the point where the bursal canal bends forwards. The copulatory bursa is a voluminous sac-shaped structure that fills the entire dorsoventral space of the body. In several specimens remnants of a spermatophore are present in the bursa. Discussion A dorsal penial fold of similar size and location as in this species D. effusa is present also in D. sagitta (some specimens have only one, dorsal fold), D. malickyi, D. benazzii Lepori, 1951, D. elegans and D. leporii Pala, Stocchino, Corso & Casu, 2000. In D. elegans the openings of the vasa deferentia into the seminal vesicle are far removed from the diaphragm, contrasting with the location of the openings immediately anterior to the diaphragm in all other species mentioned. In addition, the penial fold of D. elegans is more developed and more strongly muscular than in D. effusa. (cf. De Vries, 1984).	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E2048361FC8DFA98FB12FCEB.taxon	description	The GMYC analysis supports D. effusa as a different species (Fig. 2, Table 1), clearly delimitating the specimens from Chios as entity 20. Furthermore, D. effusa is not close to D. sagitta in the phylogenetic tree of Solà et al. (2013). Nevertheless, D. effusa shares with D. sagitta the ‘ V-shaped’ glandular zone that surrounds the ejaculatory duct (cf. De Vries, 1984: 106). In D. sagitta there are usually two penial folds, the ventral one being smaller than the dorsal one; the ventral fold may also be completely absent. However, in relation to the size of the penis papilla, the penial fold of D. sagitta is considerably bigger than that in D. effusa. Furthermore, the dorsal penial fold of D. sagitta is traversed by a cyanophilic secretion, which is discharged through its lining epithelium; such is not the case in D. effusa.	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E20B8364FCBCFC5EFB38FD73.taxon	description	(FIGS 6 – 9) Material examined: Holotype: ZMA V. Pl. 7116.1, Melanes, Naxos, Greece, 37 ° 5 ′ 3.38 ″ N, 25 ° 26 ′ 59.40 ″ E, alt. 199 m, 9 April 2009, coll. Eduardo Mateos & Eduard Solà, sagittal sections on nine slides. Paratypes: ZMA V. Pl. 7116.2, ibid., sagittal sections on ten slides; V. Pl. 7116.3, ibid., horizontal sections on four slides; V. Pl. 7116.4, ibid., sagittal sections on eight slides. 200 µm pg ed od vd Etymology: The specific epithet is derived from the Latin adjective improvisus, unexpected, and alludes to our surprise in finding a second and new species of Dugesia on such a small island as Naxos. Diagnosis: Dugesia improvisa is characterized by: an acentral, ventrally displaced ejaculatory duct, opening at the tip of the penis papilla; a short diaphragm; ectal reinforcement being confined to the posterior wall of the ascending portion of the bursal canal; vasa deferentia separately opening into the anterior section of the seminal vesicle, at a point close to the diaphragm; broad zone of abundant penis glands traversing the penial papilla and opening into the ejaculatory duct. Ecology and distribution: Specimens were collected from under stones in a small, shallow pool, receiving the outflow of water from a concrete pipe. The species is known only from this type locality. Description: Preserved specimens up to about 12.5 × 3 mm. Triangular head with distinct, blunt auricles. Posterior end obtusely pointed. Dorsal surface pale brown, with the pigment arranged in a finely reticulated pattern and with a concentration of pigment following the outline of the pharyngeal pocket (Fig. 6). Dorsal body margin and ventral surface pale. The two eyes are situated in conspicuous pigment-free patches. The pharynx is located in the posterior half of the body and measures about 1 / 8 th of the body length in preserved specimens. The mouth opening is located at the posterior end of the pharyngeal pocket. The testes are located dorsally and extend from the level of the ovaries to the posterior end of the body. The vasa deferentia penetrate the antero-lateral wall of the intrabulbar seminal vesicle; the ducts open separately into the vesicle at a position very close to the diaphragm (Fig. 7). The intrabulbar seminal vesicle is lined with an epithelium, consisting of columnar cells, that is pierced by the numerous openings of penis glands, the latter producing a granular, erythrophil secretion. At the free end of the lining epithelium of the seminal vesicle this secretion projects into the lumen as relatively large, pear-shaped, granular drops. Through a short, stubby diaphragm the seminal vesicle opens into the proximal, funnelshaped section of the ejaculatory duct. The diaphragm is short. The proximal funnelshaped section of the ejaculatory duct, immediately adjacent to the diaphragm, houses a sickle-shaped flap of tissue or secretion (Figs 7 – 9). This flap seems to be attached to the rest of the diaphragm by only a minute piece of tissue. The lining epithelium of the flap is pierced by the openings of the erythrophil penis glands that open into the seminal vesicle and also penetrate the epithelium of the rest of the diaphragm. The flap was observed in all four specimens examined and its histology suggested true mesenchyme, surrounded by an epithelium. The ejaculatory duct runs slightly acentrally, i. e. ventrally displaced, through the penis papilla, opening at its tip. The major portion of the ejaculatory duct receives the conspicuous, abundant and granular secretion of erythrophil penis glands, which are located outside of the penis. The penis papilla is a broad, pointed or blunt cone. The papilla is covered with a nucleated epithelium and is underlain with a subepithelial layer of circular muscles, followed by a layer of longitudinal muscles. The penis bulb is well developed and muscular. The small, paired ovaries are situated at about 1 / 3 rd of the distance between the brain and the root of the pharynx and are positioned directly medially to the ventral nerve cords. The oviducts arise from the dorsal surface of the ovaries and run backwards immediately dorsally to the ventral nerve cords. At the level of the copulatory apparatus the oviducts curve dorsomedially to open separately into the most proximal, posterior, section of the bursal canal, i. e. close to the point where the duct communicates with the atrium. Erythrophil shell glands open into the bursal canal immediately ventrally to the openings of the oviducts. The bursal canal is lined with a cuboidal, nucleated epithelium and is surrounded by a reversed musculature: a thin subepithelial layer of longitudinal muscle, followed by a thicker layer of circular muscle. Around the proximal, posterior, section of the bursal canal this circular muscle layer is rather thick, but it becomes gradually thinner towards the copulatory bursa. Ectal reinforcement of the bursal canal musculature is only present along the proximal section of the canal, i. e. from its opening into the atrium to about the point where the duct curves anteriad. However, this ectal reinforcement is only present as a single layer of longitudinal muscle along the posterior wall of the ascending portion of the bursal canal; it was not observed along the anterior wall of this part of the canal. The bursal canal communicates with a large, sac-shaped copulatory bursa, which occupies most of the dorso-ventral space of the body. In two specimens the bursa contained remnants of a sclerotic spermatophore. Discussion The presence of a peculiar flap of tissue on the diaphragm sets D. improvisa immediately apart from any of the known species of Dugesia. However, in specimens of other species of Dugesia a more or less crescent-shaped stretch of secretion may be present in precisely the same position, albeit less clearly attached to the epithelium, while in these specimens its staining properties clearly suggest a glandular origin. In these animals, and also in D. improvisa, this flap or stretch of secretion may be related to the formation of the spermatophore (which is formed in the ejaculatory duct) or to the transfer of sperm into the latter. However, in D. improvisa the flap did not resemble a spermatophore in statu nascendi but suggested true mesenchyme surrounded by an epithelium. We are hesitant to consider this feature as a diagnostic character of D. improvisa, but would first prefer to check the presence of this flap in another series of individuals of D. improvisa. Unfortunately, additional material is not presently available. However, D. improvisa also presents a combination of other characters that makes it different from its congeners. In the fact that the vasa deferentia open into the seminal vesicle at a point close to the diaphragm, D. improvisa resembles a good number of other species of Dugesia (cf. Sluys et al., 1998, table II). However, in other features these species differ much from D. improvisa, for example in the presence of penial or atrial folds, except Dugesia subtentaculata (Draparnaud, 1801) and D. burmaensis (Kaburaki, 1918). However, the atrium of D. subtentaculata shows a distinct musculo-glandular area (cf. De Vries, 1986), which is absent in D. improvisa. Furthermore, D. subtentaculata also possesses a ring of spongiose mesenchymatic tissue in the penis papilla that is absent in D. improvisa. In addition, in D. subtentaculata the ectal reinforcement along the bursal canal is much more developed and extends much farther anteriad. The gross morphology of the copulatory apparatus of D. burmaensis is very similar to that of D. improvisa. However, for D. burmaensis it has been reported that the oviducts arise from the anterolateral wall of the ovaries, contrasting with their dorsal origin in D. improvisa. Dugesia burmaensis resembles D. improvisa in the presence of highly developed penis glands, discharging their abundant secretion into the ejaculatory duct. Such a broad zone with abundant secretion traversing the penis papilla is also characteristic of D. sagitta from Corfu. However, there are a number of clear differences between D. sagitta and D. improvisa. In D. sagitta the penis papilla is blunt and provided with distinct, asymmetric penial folds at both the dorsal and the ventral side of its base (cf. De Vries, 1984), which are absent in D. improvisa. Furthermore, in D. sagitta the ejaculatory duct follows a central course through the penis papilla, whereas it has a ventrally displaced trajectory in D. improvisa. In addition, the ectal reinforcement of the bursal canal extends much farther anterior in D. sagitta. In all molecular analyses D. improvisa is the sister species of D. ariadnae (Fig. 2; Solà et al., 2013), the latter also restricted in its distribution to the island of Naxos. However, the two species are clearly delimited in the GMYC analysis, while morphologically D. ariadnae is very different from D. improvisa. In particular, D. ariadnae is characterized by two welldeveloped adenodactyls that are suspended from the dorsal atrial wall, one on either side of the base of the penis. On the basis of our comparative and integrative analysis, as presented above, we conclude that D. improvisa concerns a new species.	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E20E8367FC4AFD02FD33FC0C.taxon	description	(FIGS 10 – 12) Material examined: Holotype: ZMA V. Pl. 7117.1, 650 m before Kipouries (coming from Fita), Chios, Greece, 38 ° 30 ′ 43.31 ″ N, 25 ° 59 ′ 55.06 ″ E, 30 April 2010, coll. M. Vila-Farré, sagittal sections on 12 slides. Paratype: ZMA V. Pl. 7117.2, ibid., sagittal sections on nine slides. Etymology: The specific epithet is derived from the Latin naias, water nymph, and alludes to the small freshwater stream from which the specimens were collected. ed cs pp 200 µm sv pg ed cs cg sg Diagnosis: Dugesia naiadis is characterized by: vasa deferentia that open into the proximal, anterior section of the seminal vesicle; a short diaphragm; an acentral, ventrally displaced ejaculatory duct, opening terminally at the tip of a blunt penis papilla; a broad zone of cyanophilic secretion in the dorsal section of the penis papilla; oviducts that open symmetrically into the most proximal section of the bursal canal; a bursal canal provided with many irregular pleats and folds, surrounded by a welldeveloped coat of circular muscle and a zone of mesenchymatic, erythrophil gland cells; hyperplasic ovaries; lack of testes. Ecology and distribution: Specimens were collected from a small creek; the species is known only from the type locality. Description: Preserved specimens with low triangular head and rounded auricles (Fig. 10), measuring up to 11 mm in length and 2.5 mm in width. Dorsal body surface pale brown; ventral surface pale. A pair of eyes is present and somewhat smaller additional eyes are present also in the sectioned specimens. Pharynx located in the middle of the body, measuring about 1 / 6 th of the body length. The mouth opening is located at the posterior end of the pharyngeal pocket. Testes are completely absent. The ovaries are hyperplasic: ovarian tissue fills the entire dorsoventral space over a distance of about 750 μm. The midpoint of the hyperplasic ovaries is located at about 1 / 4 th the distance between the brain and the root of the pharynx. The oviducts open separately and symmetrically into the most proximal section of the bursal canal, i. e. close to the point where the canal communicates with the atrium (Fig. 11). Erythrophil shell glands discharge their secretion into the bursal canal, immediately ventrally to the oviducal openings. The bursal canal is lined with a nucleated epithelium; it follows a somewhat undulating course towards the copulatory bursa, while giving rise to a number of irregular pleats or folds that project into the surrounding mesenchyme (Fig. 12). The canal is surrounded by a very thin, subepithelial layer of longitudinal muscle, followed by a thick layer of circular muscle. Ectally to its surrounding coat of muscles the bursal canal is surrounded by a zone of mesenchymatic, erythrophil gland cells, which discharge their secretion into the lining epithelium of the canal. Only in specimen ZMA V. Pl. 7117.1 (Fig. 12) could ectal reinforcement by some longitudinal muscles be detected on the posterior wall of the bursal canal, in the region of the oviducal openings. In specimen ZMA V. Pl. 7117.1 (Fig. 12) the copulatory bursa is a large sac-shaped structure that fills the entire dorso-ventral space, but in ZMA V. Pl. 7117.2 (Fig. 11) the bursa is much smaller and also lined with cells with a more densely stained content. Although the oviducts run from the level of the copulatory apparatus to the ovaries, vasa deferentia could be traced only in the vicinity of the penis bulb. After having penetrated the ventro-lateral wall of the penis bulb, the vasa deferentia open separately into the proximal, anterior section of the seminal vesicle. The latter gradually narrows towards a small diaphragm, through which it communicates with the ejaculatory duct. Seminal vesicle and ejaculatory duct are positioned in the ventral region of the penis papilla, which therefore is asymmetrical: its dorsal section is much larger that the ventral section. The ejaculatory duct receives the secretion of numerous erythrophil penis glands and opens terminally at the blunt tip of the penis papilla. The latter is a plugshaped structure that fills most of the male atrium. The penis papilla is covered with a nucleated epithelium that is underlain by a thin layer of circular muscle, followed by an equally thin layer of longitudinal muscle. The dorsal section of the penis papilla is traversed by a broad zone of strands of cyanophilic secretion that does not seem to open into the ejaculatory duct or through the covering epithelium of the papilla. The spaces present in the penial mesenchyme, near the tip of the papilla, seem to result from clefts in torn tissue. Discussion Presence of hyperplasic ovaries and complete absence of testes are signs that these animals probably concern sexualized specimens from an otherwise asexually reproducing population. Such sexualization may be induced either spontaneously (as was the case with these animals from Chios) or experimentally and has been reported for 11 species of Dugesia (cf. Charni et al., 2004 and references therein; Stocchino, Sluys & Manconi, 2012; Harrath et al., 2013). Furthermore, hyperplasic ovaries and poorly developed testes have been found also in ex-fissiparous specimens of Phagocata morgani (Stevens & Boring, 1906; Benazzi & Ball, 1972). The fortunate circumstance that animals of an otherwise asexually reproducing population sometimes develop reproductive organs enables taxonomic identification of such specimens. In that context, the animals from Chios should be compared with other species for which a ventrally displaced ejaculatory duct has been reported, forming a presumably monophyletic subset within the genus Dugesia (Sluys et al., 1998). This comparison should be restricted to those species in which the ventrally displaced ejaculatory duct opens terminally at the tip of the penis papilla, thus excluding species with a subterminal opening. This immediately excludes D. sicula, D. aethiopica Stocchino et al., 2013 and Dugesia arabica Harrath & Sluys, 2013 as candidate species because these have a subterminal opening of the ejaculatory duct. However, both D. aethiopica and D. arabica resemble the Chios specimens in the presence of a bursal canal with many elaborate folds, a feature that has been reported also for D. biblica (cf. Benazzi & Banchetti, 1972), albeit that in the latter it is much less developed in comparison with D. aethiopica, D. arabica and the Chios specimens of D. naiadis. For D. biblica Benazzi & Banchetti (1972) describe the bursal canal as having ‘ ... un diametro alquante irregolare ... ’ [a considerably irregular diameter], which agrees with our observations on specimens from Israel (ZMA V. Pl. 698.1, V. Pl. 699.1). Another difference between the Chios animals and D. aethiopica and D. sicula concerns the openings of the oviducts into the bursal canal. In both D. sicula and D. arabica the oviducal openings are highly asymmetrical, in contrast to the symmetrical openings in D. naiadis (cf. Sluys, 2007; Harrath et al., 2013). In the specimens of D. aethiopica from Ethiopia the situation is different in that the oviducts open symmetrically into the ventral part of the horizontally running section of the bursal canal. In these type specimens the proximal section of the bursal canal approaches the atrium by running more or less parallel to the body surface, thus contrasting with the course of the canal in D. naiadis. In the presence of mesenchymal glands around the bursal canal and the patch of cyanophilic secretion in the penis papilla D. naiadis resembles D. sicula, D. biblica and the presumed biblica specimens from Bucak, Turkey (ZMA V. Pl. 813). However, in other features D. naiadis differs from these taxa. The phylogenetic analysis (Fig. 1) shows that D. naiadis belongs to the sicula – aethiopica clade (as defined in Lázaro et al., 2009) with maximum support (100 / 1), being the sister group of D. aethiopica and D. sicula. The fact that the GMYC method (Fig. 2, Table 1, entity 33) delimits the four specimens of D. naiadis as a differentiated species supports the description of this new species. Interestingly, D. naiadis does not present the duplication in the nuclear ribosomal internal transcribed spacer- 1 (ITS- 1) molecule that D. aethiopica and D. sicula share (data not shown; cf. Baguñà et al., 1999; Lázaro et al., 2009). On the basis of their gene identity we have been able to assign several asexual Dugesia populations from Chios to either D. naiadis or D. sicula (Table S 5).	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E20D8378FF18FBFEFF1DF8BB.taxon	description	(FIG. 13) Material examined: Holotype: ZMA V. Pl. 7118.1, Ermones, Corfu, Greece, 39 ° 36 ′ 37.98 ″ N, 19 ° 46 ′ 41.64 ″ E, somewhat higher upstream than ZMA V. Pl. 7119, 20 April 2009, coll. R. Sluys, sagittal sections on 13 slides. Paratypes: ZMA V. Pl. 7118.2, ibid., horizontal sections on eight slides; V. Pl. 7118.3, ibid., sagittal sections on six slides. Other material examined: ZMA V. Pl. 7119.1, Ermones, Corfu, Greece, 39 ° 36 ′ 41.93 ″ N, 19 ° 47 ′ 1.40 ″ E, outflow of river into the sea, 20 April 2009, coll. R. Sluys, sagittal sections on five slides; V. Pl. 7119.3, ibid., horizontal sections on six slides; V. Pl. 7119.4, ibid., sagittal sections on 18 slides, V. Pl. 7119.5, ibid., sagittal sections on 14 slides; V. Pl. 7119.6, ibid., sagittal sections on 17 slides. Etymology: The specific epithet is based on the prefix para (somewhat resembling, related to) and the specific epithet of the species D. sagitta. Diagnosis: The species differs morphologically from its closest relative, D. sagitta, in the presence of a very large dorsal penial fold, very small ventral fold and a ventrally displaced ejaculatory duct. Ecology and distribution: The species is known only from two sites in the same river. One site is close to the opening of this river into the sea, while the type locality is located slightly farther upstream. Comparative discussion: The taxonomic status of D. sagitta (Schmidt, 1861) from Corfu as a valid and separate species was clarified by De Vries (1984). Prior to her study, the Dugesia populations from Corfu were usually considered to be conspecific with D. gonocephala, following a conclusion reached by Komárek (1925). To avoid future taxonomic confusion, De Vries (1984) fortunately designated a series of neotypes for D. sagitta. Although the International Code of Zoological Nomenclature (ITZN, 1985; ICZN, 1999) restricts designation of a neotype to only one specimen that forms the new name-bearing type of a nominal species and thus does not allow it to be a series of animals, the neotype specimens specified by De Vries (1984: 104) represent a morphologically homogeneous set of animals. As neotype locality was chosen Messonghi River, just west of Messonghi. The Ermones population was first mentioned by Ball (1979), who attributed it to D. gonocephala. In the same paper the karyotype of presumed D. gonocephala from Corfu was analysed but it is not clear which population was studied, either the one from Ermones or the animals from Messonghi River. However, De Vries (1984) writes that animals from the neotype locality of D. sagitta, i. e. Messonghi River, were analysed. Our integrative analysis of the populations that we sampled from Corfu revealed an unexpected and interesting situation. Molecular analysis of both COI and ITS- 1 grouped the various populations sampled into three clades (cf. Solà et al., 2013). These three clades are also identified as separate entities in the GMYC analysis (Fig. 2, entities 1, 2 and 3). One clade was formed by populations 27, 28, 29 and 30 (i. e. north of the San Salvador mountain range). The second clade consisted of populations 33 and 34. The third clade consisted of two samples from basically the same locality, namely Ermones (localities 31 and 32) (Fig. S 2). On the basis of morphological analysis of the populations from Corfu we were able to differentiate between only two types. The majority of the populations sampled conformed to the classical diagnosis of D. sagitta, notably in the presence of a well-developed dorsal fold and a very small or absent ventral fold, and with a central ejaculatory duct. This also holds true for populations that we have not re-collected, but of which material is present in the collections of the NBC: Messonghi River, Marbella beach (now called Par. Ag. Ioannis Peristeron) and Mesaria. However, the population from Ermones (ZMA V. Pl. 7118 + V. Pl. 7119) is characterized by a very large dorsal penial fold, very small ventral fold and a ventrally displaced ejaculatory duct (Fig. 13). Thus, coincidence of molecular and morphological results suggests that at least the population from Ermones is well differentiated from other populations on Corfu. Therefore, we do here describe this population as the new species D. parasagitta. It remains remarkable that the populations that are geographically closest to the D. sagitta type locality, namely ZMA V. Pl. 7120 from near Vouniatades (locality 33) and ZMA V. Pl. 7121 from near Benitses (34) (entity 2, Fig. 2), differ molecularly so much from the populations in the northern part of the island (entity 1, Fig. 2), whereas morphologically they cannot be distinguished from each other, nor from the neotype population. After the separation and description of D. parasagitta, the nominal species D. sagitta actually forms a paraphyletic taxon, according to all molecular analyses done so far (cf. Solà et al., 2013; COI gene tree, Fig. 2). Furthermore, the geographical distribution of the various populations (Fig. S 2) suggests that these two units form two independent lineages. In view of the definition of a species as an independently evolving lineage, this suggests that these lineages are actually two different species. We do take a conservative approach to taxonomy and do not assign formal species status to these taxa, pending the availability of further data. However, we do suggest that entity 2 (from localities 33 and 34, i. e. in the proximity of the neotype locality of D. sagitta) is assigned to the nominal species D. sagitta, and that entity 1 (from localities 27 and 29) represents a UCS (Fig. S 2, Table 1).	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E2128378FC57FF18FC18FC51.taxon	diagnosis	Diagnosis: Dugesiidae with very slender body and rotund head. Asymmetrical penis papilla with oblique or almost vertical orientation, when non-extended. Ejaculatory duct with a distinctly subterminal opening at the anterior or antero-ventral side of the penis papilla and surrounded by a well-developed coat of circular muscle. Testes dorsal, distributed throughout the body length. Intrabulbar seminal vesicle surrounded by well-developed coat of interwoven muscle. Common oviduct, opening onto ventral, horizontal and broadened section of the bursal canal, which receives the openings of shell glands anteriorly to the oviducal opening. Bursal canal covered with a coat of circular muscle. Type species: Recurva postrema Sluys & Solà sp. nov. Etymology: The generic name is derived from the Latin adjective recurvus, bent backwards, and alludes to the situation that the ejaculatory duct curves backwards to such an extent that its opening is located at the antero-ventral side of the penis papilla. Gender: female.	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E212837AFC88FC1AFBC7F8BB.taxon	description	(FIGS 14 – 16) Material examined: Holotype: ZMA V. Pl. 7122.1, NE Laerma, Rhodes, Greece, 36 ° 10 ′ 6.76 ″ N, 27 ° 57 ′ 34.55 ″ E, alt. 135 m, 5 April 2009, coll. Eduardo Mateos and Eduard Solà, sagittal sections on six slides. Paratypes: ZMA V. Pl. 7122.2, ibid., sagittal sections on four slides (not fully mature specimen); V. Pl. 7122.3, ibid., sagittal sections on six slides; V. Pl. 7122.4, ibid., sagittal sections on four slides; V. Pl. 7122.5, ibid., sagittal sections on seven slides; V. Pl. 7122.6, ibid., horizontal sections on four slides; V. Pl. 7122.7, ibid., sagittal sections on six slides; V. Pl. 7122.8, ibid., sagittal sections on eight slides; V. Pl. 7122.9, ibid., sagittal sections on six slides. Etymology: The specific epithet is derived from the Latin adjective postremus, located posteriorly, and alludes to the far posteriorly located position of the copulatory apparatus. Diagnosis: Animals slender, with rotund head. Pharynx and copulatory apparatus situated in the far posterior end of the body. Dorsal testes, distributed throughout the body length but anteriormost testes located at a considerable distance behind the brain. Vasa deferentia open asymmetrically into intrabulbar seminal vesicle. Penis papilla asymmetrical, with more or less vertical orientation in the male atrium. Ejaculatory duct opening at the anterior or ventroanterior side of the penis papilla. Ventral or ventroanterior, muscular penial fold present at the point of insertion of the penis papilla. Ovaries located at about 1 / 4 th the distance between the brain and the root of the pharynx. Distal, posterior parts of the oviducts increase in diameter before communicating with an equally wide common oviduct. Bursal canal is surrounded by a well-developed coat of circular muscle. Ecology and distribution: The species is known only from the type locality, where it was collected from stagnant water in a rather dry creek. Specimens were found in high numbers, gliding on the substrate, together with other small, white flatworms of an unknown species. Description: Preserved specimens measure up to 9.5 mm in length and 2.25 mm in width. Notably live specimens are very slender (Fig. 14), with a rotund head that is provided with a pair of close-set eyes, situated in pigment-free patches. Each eye cup houses numerous retinal cells. Behind the eyes, along the lateral margins of the body, there is an auricular streak on either side, at the level of which the head narrows so that there is a more slender neck region. The dorsal surface is finely pigmented pale brown, with notable accumulations of pigment around the pharyngeal pocket. Ventral surface pale. The pharynx measures between 1 / 6 th and 1 / 8 th of the body length and is positioned far into the posterior part of the body. The musculature of the pharynx conforms to the planariid type. This highly posterior location of the pharynx means that the copulatory apparatus is pushed far into the tail end of the animal. The mouth opening is located at the posterior end of the pharyngeal pocket. The testes are located dorsally, extending from directly behind the ovaries to almost the posterior margin of the body. After having penetrated the penis bulb, the vasa deferentia open separately into the intrabulbar seminal vesicle. The openings of the seminal ducts are asymmetrical in that one vas deferens opens into the ventral section and the other in a more dorsal section of the seminal vesicle (Figs 15, 16). The latter, lined with a nucleated epithelium and surrounded by a coat of intermingled muscle, communicates with the ejaculatory duct, which in most of the specimens examined exhibits an S-shaped loop before curving downwards to follow its central course through the penis papilla. The papilla is more or less cylindrical in shape and has a more or less vertical orientation in the male atrium. The penis papilla is highly asymmetrical in the sense that in its distal, ventral section the ejaculatory duct shows a sharp, anteriorly directed, knee-shaped bend, after which it opens at the anterior or ventroanterior side of the penis papilla (Figs 15, 16). This course of the ejaculatory duct results in the situation that the anterior portion or lip of the penis papilla is shorter and smaller, in some specimens much shorter and smaller, than the posterior section. At the base of this anterior or ventro-anterior lip of the penis papilla, at its point of insertion, a penial fold is present. This fold is characterized by a more or less developed outbulging and is provided with its own musculature. It is a penial fold, in contrast to an atrial fold, because it is located entally to the point of attachment of the musculature of the penis bulb. The penis papilla is covered with a thin, nucleated epithelium. The ovaries are located at about 1 / 4 th the distance between the brain and the root of the pharynx. This implies that also the row of testes starts at a considerable distance posterior to the brain, as may be observed even in living specimens (Fig. 14). Directly posterior to the gonopore the oviducts turn dorso-medially, while their diameter increases sv cb considerably. Subsequently, the oviducts fuse to form a common oviduct, with an equally wide diameter, that opens into the ventral section of the bursal canal. The latter starts at the copulatory bursa as a rather narrow duct that gradually widens and posterior to the gonopore makes a sharp anteriorly directed bend before opening into the rather dorsal section of the atrium. The more or less horizontally running and widened part of the bursal canal receives the openings of the shell glands anteriorly to the opening of the common oviduct. The nucleated bursal canal is surrounded by a well-developed coat of circular muscle. The copulatory bursa sits immediately anterior to the penis bulb.	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
03DB7724E211837CFF20FF18FE18FA35.taxon	description	(FIG. 17) Material examined: Holotype: ZMA V. Pl. 7123.1, near Agios Georgios, Kefalonia, Greece, 38 ° 6 ′ 0.72 ″ N, 20 ° 44 ′ 55.50 ″ E, 26 April 2009, coll. R. Sluys, sagittal sections of the anterior, prepharyngeal end of the animal on six slides; V. Pl. 7123.1, ibid., sagittal sections of the posterior end (including the pharynx) of the same animal on six slides. Etymology: The specific epithet is derived from the Latin adjective coniunctu s, connected, and alludes to the genito-intestinal connection present in this species. Diagnosis: Animals slender, with rotund head. Dorsal testes, distributed throughout the body length. Vasa deferentia narrow when penetrating the ventro-lateral side of the penis bulb, subsequently expanding again and opening into the mid-lateral section of the intrabulbar seminal vesicle. Asymmetrical penis papilla, with an oblique, ventro-posterior orientation. Ejaculatory duct opening at the anteroventral side of the penis papilla. Common oviduct surrounded by a coat of circular muscle. Copulatory bursa communicating with a branch of the intestine. Bursal canal surrounded by a layer of circular muscle. Ecology and distribution: The species is known only from its type locality, where it was found under stones in an almost dry, muddy stream flowing beneath a concrete bridge. Description: In the field the two specimens collected (one immature) were identified as Schmidtea - like animals, i. e. with a rounded head. The animals were very slender, the holotype specimen measuring up to 2 cm in length when fully stretched and moving. Dorsal surface pigmented, ventral surface pale (as deduced from examination of the sections). Each eye cup houses numerous retinal cells. The pharynx measures about 1 / 9 th of the body length, its root being situated about half-way along the body length. The mouth opening is located at the posterior end of the pharyngeal cavity. The testes are situated dorsally, extending from directly behind the brain into the posterior end of the body. The vasa deferentia, which are expanded to spermiducal vesicles, narrow considerably when they penetrate the ventro-lateral side of the penis bulb. Once within the bulb, the ducts expand again in diameter and, subsequently, open into the mid-lateral section of the intrabulbar seminal vesicle. The latter is lined with a nucleated epithelium and surrounded by a rather thick coat of interwoven muscles. The ejaculatory duct arises from the dorsal section of the seminal vesicle and immediately thereafter pp cod sharply curves postero-ventrally to continue its more or less central course through the penis papilla. However, at some point the ejaculatory duct makes another sharp, hooked bend towards the anteroventral surface of the body (Fig. 17). This results in the situation that the duct opens at the anteroventral side of the penis papilla. The papilla has an oblique, ventro-posterior orientation and is covered with a nucleated epithelium. Because of the peculiar course of the ejaculatory duct, the distal section of the penis papilla is highly asymmetrical, with a short ventral lip and a bulky dorsal lip. In fact, the tip of the papilla is to some extent also curved towards the lateral side of the male atrium. Therefore, the opening of the ejaculatory duct is not only displaced towards the antero-ventral side of the penis papilla but also to a more lateral position. This lateral twist of the tip of the penis papilla may be due to a preservation artefact. The major portion of the ejaculatory duct is surrounded by a relatively thick layer of mostly circular muscle fibres. The paired ovaries are situated directly behind the brain. Immediately posterior to the gonopore the oviducts turn medially and fuse to form a common oviduct, which opens at the postero-ventral section of the bursal canal. The common oviduct is surrounded by a coat of circular muscle. The bursal canal arises as a broad duct from the mid-posterior wall of the atrium. This first, broad section of the canal runs more or less horizontally and receives the openings of the abundant shell glands, which open anteriorly to the opening of the common oviduct. This broad part of the bursal canal narrows considerably and, subsequently, curves forwards to continue its course immediately dorsally to the male atrium and the penis bulb. Half-way along its course the canal becomes even narrower before communicating with the copulatory bursa. The entire bursal canal is lined with a nucleated epithelium and is surrounded by a layer of circular muscle. The copulatory bursa lies immediately anterior to the penis bulb, while its ventral part is connected with a branch of the intestine.	en	Sluys, Ronald, Solà, Eduard, Gritzalis, Konstantinos, Vila-Farré, Miquel, Mateos, Eduardo, Riutort, Marta (2013): Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 169 (3): 523-547, DOI: 10.1111/zoj.12077, URL: http://dx.doi.org/10.1111/zoj.12077
