identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
206687B6FFD0FFBDFC78A512FE76071D.text	206687B6FFD0FFBDFC78A512FE76071D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dugesia milloti de Beauchamp 1952	<div><p>Dugesia milloti de Beauchamp, 1952</p><p>Material examined</p><p>ZMA V.Pl. 787.1, stream of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.15&amp;materialsCitation.latitude=-12.616667" title="Search Plazi for locations around (long 49.15/lat -12.616667)">Roussette</a>, Montagne d’Ambre, Madagascar (12°37 ʹ 0.00 ʹʹ S 49°8 ʹ 60.00 ʹʹ E), 17 September 1957, sagittal sections on nine slides .</p><p>Mf N F 217-c, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.916668&amp;materialsCitation.latitude=-19.033333" title="Search Plazi for locations around (long 47.916668/lat -19.033333)">Andrangolaoka</a>, Madagascar (19°1 ʹ 60.00 ʹʹ S 47°55 ʹ 0.00 ʹʹ E), coll. F. Sikora (specimens registered at the Berlin Museum on 25 March 1892), sagittal sections on nine slides; Mf N F 217-a, ibid., sagittal sections on eight slides; Mf N F 217-b, ibid., sagittal sections on five slides .</p><p>Geographical distribution</p><p>Known from the following localities: (i) the type locality Morafenobé, forêt de Mahajeby, stream of Roussette, northwest Madagascar; (ii) Parc National de La Montagne d’Ambre, Diana Region, northern Madagascar; (iii) Andrangolaoka, Central High Plateau, east to Mantasoa Lake; (iv) Cascade de Mutsamudu, Anjouan, Comoro Islands (Fig. 1).</p><p>Comparative discussion</p><p>As de Vries (1988) based her redescription of D. milloti primarily on specimens from Montagne d’Ambre (in our material represented by specimen ZMA V.Pl. 787.1) her account still is basically correct, although she did not provide specific information on the testes and the position of the mouth opening. Examination of our material learned that the small testes are situated dorsally and that the follicles extend from a short distance behind the ovaries to the posterior end of the body. The ovaries are located at a short distance (~400 μm) behind the brain, i.e. at about one-ninth of the distance between the brain and the root of the pharynx. The mouth opening is located at about one-sixth of the distance between the hind wall of the pharyngeal pocket and the root of the pharynx.</p><p>Characteristic for D. milloti are the two annular atrial folds or valves, the posterior valve receiving a bundle of longitudinal muscles that extends posteriorly from the penis bulb (cf. de Vries 1988: figs 9, 10; Fig. 3). These annular atrial folds are much less clear in the old specimens from Andrangolaoka, which suffer from preservation and contraction artefacts (Figs 4, 5). However, in specimen MfN F217-a, folds in the atrium are still visible, presumably representing the most posterior valve—also receiving the longitudinal muscles from the penis bulb—as the anterior fold sits at the root of the penis papilla (Fig. 6).</p><p>As compared with the other long-eared species of Dugesia (see below), the copulatory apparatus of D. milloti is characterized by a slim, conical penis papilla and a large, valve-like diaphragm, the latter receiving the erythrophil secretion of penial glands that contributes to the formation of the spermatophore (de Vries 1988: figs 9, 10; Figs 5, 6).</p></div>	https://treatment.plazi.org/id/206687B6FFD0FFBDFC78A512FE76071D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Stocchino, Giacinta Angela;Sluys, Ronald;Solà, Eduard;Riutort, Marta;Manconi, Renata	Stocchino, Giacinta Angela, Sluys, Ronald, Solà, Eduard, Riutort, Marta, Manconi, Renata (2024): The long-eared freshwater planarians of Madagascar form a separate phylogenetic clade within the genus Dugesia (Platyhelminthes: Tricladida), with the description of two new species. Zoological Journal of the Linnean Society (zlae 143) 202 (4): 1-29, DOI: 10.1093/zoolinnean/zlae143, URL: https://doi.org/10.1093/zoolinnean/zlae143
206687B6FFDDFFBAFF02A77CFA4E0723.text	206687B6FFDDFFBAFF02A77CFA4E0723.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dugesia crassimentula Sluys & Stocchino 2024	<div><p>Dugesia crassimentula Sluys &amp; Stocchino, sp. nov.</p><p>urn:lsid:zoobank.org:act: 544B4EB4-732B-415E-9753- 98D78F1C31D0</p><p>Material examined</p><p>Holotype: ZMA V.Pl. 786.1, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=46.885864&amp;materialsCitation.latitude=-22.48105" title="Search Plazi for locations around (long 46.885864/lat -22.48105)">Route Nationale d’Ivohibe</a> (700), south-east Madagascar (22°28 ʹ 51.78 ʹʹ S 46°53 ʹ 9.10 ʹʹ E), 1950, coll. J. Millot, sagittal sections on four slides.</p><p>Etymology</p><p>The specific epithet is based on the Latin adjective crassus, thick, stout, and the noun mentula, penis; it alludes to the plump penis papilla of the species.</p><p>Diagnosis</p><p>A Dugesia species characterized by the following features: high triangular head with a distinctly pointed auricle on either side; mouth opening at the hind end of the pharyngeal pocket; distinct layer of transverse muscles directly dorsally to the ventral nerve cords; plump, barrel-shaped penis papilla, provided with a large dorsal penial fold and a small ventral fold; bundles of longitudinal muscles traversing the penis papilla and attaching to the penial folds; large, conical diaphragm; distinct seminal vesicle absent; ectal reinforcement on bursal canal absent; highly asymmetrical openings of oviducts into bursal canal.</p><p>Geographical distribution</p><p>Known only from the type locality.</p><p>Description</p><p>As this specimen was originally assigned to the species Dugesia milloti (cf. de Vries 1988: 363), it is assumed that its external appearance is similar to that of the latter. Length of preserved specimens ~ 9 mm (as measured on histological sections).</p><p>Apart from the usual subepidermal body musculature there is a distinct layer of transverse muscle located directly dorsally to the ventral nerve cords and extending throughout the body length (Fig. 7).</p><p>Pharynx located in the middle of the body, measuring about a quarter of the body length (as measured on histological sections). No extra layer of longitudinal muscle in the outer pharynx musculature. Mouth opening at the hind end of the pharyngeal pouch.</p><p>Many small, dorsal testes are distributed throughout the body length, extending from directly behind the ovaries far into the posterior end of the body.</p><p>At about the level of the root of the pharynx, the vasa deferentia expand to form spermiducal vesicles, which are filled with sperm. Posteriorly to the pharyngeal pocket the sperm ducts turn dorso-medially, after which they first decrease in diameter and then separately penetrate the antero-lateral wall of the penis bulb. Hereafter, the sperm ducts traverse the penis bulb to separately open into the funnel-shaped, proximal, posterior portion of the lumen of the diaphragm (Fig. 8A). This lumen is lined with a cuboidal, nucleated epithelium that is penetrated by a fine-grained, cyanophil secretion.</p><p>The large, conical diaphragm is covered by a cuboidal, nucleated epithelium, which continues on the proximal, anterior half of the ejaculatory duct and is penetrated by the orange secretion of penis glands. The distal part of the wide ejaculatory duct shows a few folds before it opens at the blunt tip of the penis papilla. The latter is a plump, barrel-shaped structure that is provided with penial annexes (cf. Stocchino et al. 2017) in the form of two penial folds. One, small fold is located at the ventral surface of the papilla, rather close to its root, while the other fold occurs dorsally and is much larger (Figs 7 and 8A). In addition to these penial folds, the ventral lip of the penis papilla exhibits at its tip an invagination that may represent a non-permanent structure, presumably being based on a contraction artefact.</p><p>The musculature of the penis bulb extends posteriorly beyond the root of the penis papilla and attaches to two small, dorsally and ventrally located atrial folds, situated near the root of the papilla (Figs 7 and 8A). Bundles of well-developed longitudinal muscle fibres run from the penis bulb through the papilla and attach to the penial folds. The epithelium of the penis papilla is underlain by a thin layer of circular muscle.</p><p>The ovaries are situated at some distance (~500 μm) behind the brain, i.e. at about a quarter of the distance between the brain and the root of the pharynx. The infranucleated oviducts run posteriorly to about the level of the gonopore, from where they turn medio-dorsad. The left oviduct runs far towards the dorsal body surface and then bends antero-ventrally to open through the dorsal wall of the bursal canal. The right oviduct opens into the broad, basal portion of the bursal canal, not far from where it communicates with the atrium. Shell glands discharge an orange secretion into the bursal canal immediately ventrally to the oviducal openings.</p><p>The bursal canal arises from the postero-lateral wall of the atrium and at first has an oblique orientation into postero-dorsal direction before it curves sharply anteriad.The obliquely oriented portion of the bursal canal is already rather wide and after the bend in anterior direction it gradually enlarges even more until it is as wide as the copulatory bursa, thus passing smoothly into the latter (Fig. 8B). The bursal canal is lined with an infranucleated epithelium that is underlain by a single, subepithelial layer of longitudinal muscle, followed by an only slightly thicker layer of circular muscle; ectal reinforcement is absent. Bursal canal and copulatory bursa contain a large spermatophore of which the sac-shaped capsule is packed with sperm.</p><p>Comparative discussion</p><p>This specimen initially was identified by de Vries (1988) as D. milloti, as she was of the opinion that contraction of the longitudinal muscles running from the penis bulb and attaching to the posterior atrial fold would considerably shorten the atrium, so that the anterior fold would sit at the base of the penis papilla. Further, she argued that contraction of the penis bulb musculature would narrow the seminal vesicle and alter the shape of the diaphragm. However, we believe that the morphology of this specimen cannot be explained as merely due to contraction artefacts but that it exhibits a suite of essentially different character states concerning the following structures: penis papilla, penial and atrial folds, diaphragm, seminal vesicle, ectal reinforcement, oviducts, mouth opening, and transverse musculature. These differences between D. milloti and D. crassimentula will be discussed below.</p><p>An immediately obvious difference between D. milloti and D. crassimentula concerns the size and shape of the penis papilla, which in the former is always a slim cone (cf. de Vries 1988: figs 9, 10; Fig. 3A), whereas in the latter it is a plump, barrel-shaped structure that completely fills the male atrium (cf. de Vries 1988: fig. 14C; Fig. 7). Further, in D. crassimentula the penis papilla is provided with two folds, a large dorsal one and a small ventral fold, which are absent in D. milloti .</p><p>The atrium of D. milloti is provided with two annular folds or valves, with the posteriorly extending musculature of the penis bulb attaching to the posterior valve (cf. de Vries 1988: figs 9, 10; Fig. 8A, B). In contrast, D. crassimentula only has two small atrial folds, not valves, one dorsal and one ventral, receiving the longitudinal muscles coming from the penis bulb.</p><p>The diaphragm of D. milloti is of the stubby, valve-shaped type (cf. de Vries 1988: figs 9, 10; Figs 3A, B, 5), whereas that of D. crassimentula is of the pointed, conical type (Fig. 8A). Another aspect of the penis that differs between D. milloti and D. crassimentula is the presence of a well-developed seminal vesicle in the former (cf. de Vries 1988: figs 9, 10; Fig. 3A, B) and the virtual absence of such a vesicle in the latter (Fig. 8A).</p><p>In D. milloti the connection between copulatory bursa and bursal canal is established in the usual manner, while the diameter of the latter also shows the usual dimension (cf. de Vries 1988: figs 9, 10; Fig. 5). In contrast, in D. crassimentula the diameter of the bursal canal expands until it has reached the same size as the copulatory bursa, after which it almost imperceptibly grades into the bursa (Fig. 8B). With respect to the female copulatory apparatus, another difference between D. milloti and D. crassimentula resides in the development of the ectal reinforcement on the bursal canal. In D. milloti this extra layer of longitudinal muscles extends from the vaginal area to the copulatory bursa or at least to about halfway along the bursal canal, whereas in D. crassimentula ectal reinforcement is completely absent. Another aspect of the female reproductive system concerns the oviduct, which consists of nucleated cells in D. milloti and infranucleated cells in D. crassimentula, while in the former the ducts open symmetrically into the bursal canal, whereas in D. crassimentula the oviducal openings are highly asymmetrical.</p><p>In D. crassimentula the mouth opening is located at the posterior end of the pharyngeal pouch, whereas in D. milloti it is shifted anteriad, being located at about one-sixth of the distance between the hind wall of the pharyngeal pouch and the root of the pharynx (see D. milloti description).</p><p>De Vries (1988) already noted that the development of the transverse musculature, located directly dorsally to the ventral nerve cords, varies between species of Dugesia . In D. milloti it is poorly developed, whereas in D. crassimentula the transverse muscles are quite distinct (Fig. 7).</p><p>Thus, there are ample anatomical differences between D. milloti and D. crassimentula, supporting the separate specific status of the latter, as it neither conforms to any other known species of Dugesia .</p></div>	https://treatment.plazi.org/id/206687B6FFDDFFBAFF02A77CFA4E0723	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Stocchino, Giacinta Angela;Sluys, Ronald;Solà, Eduard;Riutort, Marta;Manconi, Renata	Stocchino, Giacinta Angela, Sluys, Ronald, Solà, Eduard, Riutort, Marta, Manconi, Renata (2024): The long-eared freshwater planarians of Madagascar form a separate phylogenetic clade within the genus Dugesia (Platyhelminthes: Tricladida), with the description of two new species. Zoological Journal of the Linnean Society (zlae 143) 202 (4): 1-29, DOI: 10.1093/zoolinnean/zlae143, URL: https://doi.org/10.1093/zoolinnean/zlae143
206687B6FFDAFFA3FC5BA78FFB4C0613.text	206687B6FFDAFFA3FC5BA78FFB4C0613.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dugesia insolita Stocchino & Sluys 2024	<div><p>Dugesia insolita Stocchino &amp; Sluys, sp. nov.</p><p>urn:lsid:zoobank.org:act: 9199C00D-BA4A-4DD4-BED2- 16A3A1884BBE</p><p>Material examined</p><p>Holotype: RMNH.VER. 21536.1 (field code: GS32-Dmad5-6) one set of sagittal sections on 18 slides, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.216667&amp;materialsCitation.latitude=-21.133333" title="Search Plazi for locations around (long 47.216667/lat -21.133333)">Central High Plateau</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.216667&amp;materialsCitation.latitude=-21.133333" title="Search Plazi for locations around (long 47.216667/lat -21.133333)">Western Slope</a>, small unnamed water course, tributary of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.216667&amp;materialsCitation.latitude=-21.133333" title="Search Plazi for locations around (long 47.216667/lat -21.133333)">Matsiatra River</a>, Upper Mangoky basin, 4 km upstream of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.216667&amp;materialsCitation.latitude=-21.133333" title="Search Plazi for locations around (long 47.216667/lat -21.133333)">Ambohimahasoa</a> (21°08 ʹ S 47°13 ʹ E), between Route National 7 to Fianarantsoa and Route national 25 to <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.216667&amp;materialsCitation.latitude=-21.133333" title="Search Plazi for locations around (long 47.216667/lat -21.133333)">Ranomafana National Park</a>, Madagascar, September 2011, coll. R. Manconi.</p><p>Other specimens: Nine individuals were preserved in absolute ethanol, two of which (Dins1 and Dins2) were used for DNA extraction and sequencing in the study of Solà et al. (2022). GenBank accession numbers of the obtained sequences are given in Table 2.</p><p>Etymology</p><p>The specific epithet, derived from the Latin adjective insolitus, unusual, uncommon, alludes to the remarkable situation that in this species the diaphragm has taken over the function of the penis papilla.</p><p>Diagnosis</p><p>A Dugesi a species characterized by the following features: high triangular head with a distinctly pointed auricle on either side; mouth opening at the posterior end of the pharyngeal pocket; vasa deferentia separately opening into anterior portion of the bulbar lumen; functional penis papilla a stubby, cone-shaped structure exhibiting an outbulging that provides it a shape in the form of a head of a spear; erythrophil penis glands discharging through the lining epithelium of the functional penis papilla; male atrium with three annular folds, the posterior-most one receiving longitudinal muscles that extend from the penis bulb; hyperplasic ovaries extending posterior to the level of the root of the pharynx; oviducts separately opening into the most proximal section of the bursal canal; poorly developed ectal reinforcement layer extending from about the knee-shaped bend in the bursal canal to about halfway between the bend and the copulatory bursa.</p><p>Geographical distribution</p><p>Known only from the type locality.</p><p>Habitat</p><p>Planarians were found under pebbles and boulders in shallow water of a rivulet in the relict Rainforests of the Atsinanana.</p><p>Reproduction and behaviour</p><p>At collection, all 20 specimens of D. insolita were asexual. After four months of rearing under standard laboratory conditions, a single asexual specimen underwent a sexualization process, thus producing an ex-fissiparous individual characterized by hyperplasic ovaries and presence of a copulatory apparatus. Asexual individuals exhibited post-pharyngeal fissioning.</p><p>Just after collection and during rearing, both in the field and in the laboratory, a particular kind of movement was observed. Besides the usual gliding type of locomotion, these planarians showed a swimming-like movement with up-and-down undulations of the body when they were not adhering to the substrate and moved, for example, from the water surface to the bottom of the container. Moreover, during their usual gliding the body margins showed an undulating movement.</p><p>Furthermore, on some occasions we observed a peculiar autotomy-like process when the worms were picked up softly with a paintbrush. Immediately after the planarians were touched, they quickly detached their tail, thus dividing at a post-pharyngeal level. The wound margins of the two resulting fragments were characterized by very irregular outlines, resembling those resulting from a forceful laceration. On one occasion, it happened that after this first division, touching of the anterior portion triggered another quick autotomy-like process, thus resulting in the detachment of another post-pharyngeal fragment. After these autotomy-like processes, each of the resulting fragments regenerated the missing body parts.</p><p>Description</p><p>Live animals with a high triangular head, provided with distinctly pointed auricles and two prominent eyes set in pigment-free patches (Fig. 9). Body size of live, fissiparous specimens ranged from 8 to 11 mm in length and 1.5–2 mm in width, while the single sexualized specimen measured 17 × 5 mm. Dorsal surface with numerous dark-brown specks and a dark mid-dorsal stripe that runs from behind the pharynx to the tip of the tail (Fig. 9). In some specimens this dorsal stripe runs from the anterior tip of the head to the very tip of the tail. Ventral surface pale and provided with dark brown specks, less numerous than on the dorsal surface.</p><p>Pharynx located at about halfway the length of the body, measuring about one-sixth of the body length. No extra layer of longitudinal muscles in the outer pharynx musculature. The mouth opening is located at the posterior end of the pharyngeal pocket.</p><p>The outer pharynx epithelium is underlain by a thin, subepidermal layer of circular muscle, followed by an equally thin layer of longitudinal muscle; this outer pharyngeal musculature is devoid of an extra, inner layer of longitudinal muscles. The lining epithelium of the pharyngeal lumen is underlain by a thick, subepithelial layer of circular muscles (~50 μm thick), followed by a much thinner layer of longitudinal muscles (Fig. 10A).</p><p>The well-developed testes are located dorsally and extend from about the anterior level of the hyperplasic ovaries far into the tail region of the body.</p><p>The vasa deferentia are expanded to form spermiducal vesicles—filled with sperm—which, in addition, may show considerable outbulgings. Upon penetrating the muscular penis bulb, the vasa deferentia gradually decrease in diameter and after having become very narrow, the ducts separately open into the anterior portion of the bulbar lumen or seminal vesicle. The wall of the latter is highly pleated, with the folds projecting into its lumen. One of the larger folds forms an elongated papilla that projects posteriad into the lumen of a structure that functions as penis papilla (see below); through this fold runs a minute duct by means of which the seminal vesicle communicates with the lumen of the functional penis papilla. The latter is a stubby, cone-shaped structure that has a more or less horizontal orientation and projects into the atrial lumen (Fig. 11). At about one-third of the distance between its root and the tip, the papilla exhibits a fold or outbulging, thus providing the papilla—in sagittal view—with a hastate shape or the form of a head of a spear.</p><p>Erythrophil penis glands discharge through the lining epithelium of the functional penis papilla, which is surrounded by the funnel-shaped portion of a sclerotic spermatophore (Figs 11, 12). It is here surmised that the secretion of these penial glands contributed to the formation of the spermatophore, in the same way that normally in Dugesia species the spermatophore emanates from the penial diaphragm (see below). Therefore, it is here also surmised that the presumed penis papilla of D. insolita is actually homologous to the diaphragm (see below). Therefore, in the above and below we refer to the papilla as the functional penis papilla instead of the true penis papilla.</p><p>The large penis bulb is strongly muscular and consists of intermingled circular and longitudinal muscle fibres. From the outer surface of the penis bulb longitudinal fibres run posteriad and attach to the posterior atrial fold (Fig. 11).</p><p>The major part of the male atrium is lined with a tall epithelium, which, however, becomes very thin on its most anterior section near the base of the functional penis papilla. The atrium is surrounded by a subepithelial layer of circular muscles that is well-developed on its anterior section but rather thin on more posterior portions. The male atrium is provided with three annular folds or constrictions. One of the folds envelopes the root of the functional penis papilla, a second fold is located just anterior to the tip of the papilla, while the third, posterior fold is positioned at the posterior end of the male atrium. This third fold may indeed be considered as the separation between the male and the female atrium, since the openings of shell glands penetrate the dorsal epithelium of that part of the atrium that lies immediately posterior to this third fold. Basically, the openings of erythrophilic shell glands surround the opening of the bursal canal through the roof of the female atrium (Fig. 11).</p><p>The highly hyperplasic ovaries extend from ~800 μm behind the eyes and the brain to a little posterior (~700 μm) to the level of the root of the pharynx, occupying almost the entire dorsoventral space of the body (Fig. 10B). The oocytes clearly exhibit a degenerative condition, as evidenced by vacuolization of their nuclei and cytoplasm.</p><p>The infranucleated oviducts run immediately dorsal to the ventral nerve cords; at the level of the gonopore the ducts turn medially to open—separately—into the most proximal section of the bursal canal, close to the opening of the latter through the roof of the female atrium. From the point of this opening, the bursal canal at first runs vertically towards the dorsal body surface and then exhibits a knee-shaped, anteriorly-directed bend, after which it follows its course more or less parallel to the dorsal and ventral body surfaces. The bursal canal communicates with a more or less ellipsoidal copulatory bursa that is located at a short distance behind the pharyngeal pocket and the mouth opening.</p><p>The bursal canal is lined with a tall, infranucleated epithelium. The long horizontal portion of the bursal canal is surrounded by a single, subepithelial layer of longitudinal muscles, followed by a slightly thicker layer of circular muscles. The short, vertical portion of the bursal canal is surrounded by a thin, subepithelial layer of circular muscles, followed by a layer of obliquely oriented fibres. An extra outer layer of longitudinal muscles forms a poorly developed ectal reinforcement layer that runs from about the knee-shaped bend in the bursal canal to about halfway between the bend and the copulatory bursa.</p><p>Very abundant and conspicuous cement glands discharge their orange-staining secretion into the short gonoduct.</p><p>Karyology</p><p>Metaphasic plates revealed that specimens constantly showed a set of 18 chromosomes. Chromosomes from six metaphasic plates could be arranged, according to their length, into nine groups of two chromosomes with a diploid chromosome set of 2n = 18; n = 9. The analysis within each pair of chromosomes revealed uniformity in both length and centromeric position, except for chromosome 1, in which the first element is always longer than its homologue. The first two chromosomes were clearly identifiable. Instead, from the third to the ninth pair, the differences in length were so small that contiguous chromosomes could be overlapping (such as chromosomes 3 and 5, 6 and 7, 8 and 9). Chromosomal length decreased gradually, with low standard deviation values. In contrast, the centromeric indices showed great variation, resulting in high standard deviations, as, for example, for chromosome numbers 4 and 6 (Fig. 13; Table 3).</p><p>The karyometric data indicate that the diploid chromosome complement is characterized by metacentric heterobrachial chromosomes, with the exceptions of chromosome 3, which is submetacentric, and chromosomes 1 and 9 that are metacentric, bordering on metacentric isobrachial (Table 3).</p><p>Comparative discussion</p><p>Dugesia insolita is included in a phylogenetic clade encompassing all long-eared Dugesia from Madagascar with full support (Fig. 2). However, none of the phylogenetic approaches lent good support for the relationship of D. insolita to the rest of the long-eared populations in this lineage. Thus, its precise position within this group remains unresolved.</p><p>The two specimens from the same population of D. insolita are clustered together with full support in both ML and BI phylogenetic trees, indicating that they most probably also belong to this species.</p><p>Sperm transfer by means of a sclerotic spermatophore is a common phenomenon among freshwater planarians (Sluys 1989, Sluys and Riutort 2018). Such spermatophores may be simply sac-shaped or may possess a long, narrow, and hollow stalk, the latter situation being the case in species of Dugesia, including D. insolita .</p><p>Spermatophores are formed in the ejaculatory duct running through the penis papilla, the secretions of various penial glands contributing to the formation of the spermatophore. In species of Dugesia, the diaphragm in the proximal section of the ejaculatory duct makes a different contribution to the formation of the spermatophore as compared with the middle and distal parts of the duct. The secretions discharged through the tip of the lining epithelium of the conus of the diaphragm form an antiagglutination layer at the inner surface of the stalk of the spermatophore, while the actual wall of the spermatophore is formed by secretions discharged through a more basal, anterior portion of the diaphragm, as well as the most proximal part of the ejaculatory duct (Vreys et al. 1999).</p><p>This construction method results in the situation that during the formation process the funnel-shaped basal portion of the stalk of the spermatophore envelopes the diaphragm (cf. Vreys et al. 1997: fig. 2; 1999: fig. 4), a situation that, over the years, we have regularly observed in histological preparations of species of the genus Dugesia . A similar situation is present in D. insolita (Figs 11, 12) in which the funnel-shaped base of the spermatophore covers a cone-shaped structure that resembles a short penis papilla, the covering epithelium of which is penetrated by the openings of erythrophil penial glands that contribute to the formation of the stalk of the spermatophore. We have never encountered the situation that in species of Dugesia the penis papilla contributed in this way to the formation of the spermatophore, as the latter is always formed within the ejaculatory duct. Therefore, it is here surmised (see above) that this cone-shaped, penis-like structure is actually homologous to the diaphragm in other species of Dugesia —as it is without doubt that D. insolita belongs to this genus (see molecular results)—albeit it may function as a penis papilla. It should be noted that the conspicuous fold projecting into the lumen of the functional penis papilla and through which runs a minute duct, cannot be seen as phylogenetically homologous to the diaphragm, nor as being functionally equivalent to it, since it does not receive the secretion of penial glands and does not have any connection with the spermatophore.</p><p>In this context it is noteworthy that the epithelium of the first annular fold in the atrium, located at the base of the functional penis papilla, is much lower than that of the rest of the epithelium. Perhaps this lower epithelium signals that this proximal, basal fold actually is the remnant of what once was a normal penis papilla?</p><p>Other notable features of D. insolita are (i) shell glands opening through the roof of the female atrium, (ii) the knee-shaped bend in the bursal canal, (iii) muscle fibres extending from the penis bulb to the posterior wall of the male atrium, (iv) the presence of hyperplasic ovaries, and (v) a highly triangular head with long and pointed auricles.</p><p>In species of Dugesia, shell glands usually open into the bursal canal, just ventral to the oviducal openings. However, frequently the openings of the shell glands are located very close to the point where the bursal canal communicates with the atrium, while occasionally part of the shell glands actually discharges into the female atrium, e.g. in D. hepta Pala et al., 1981, D. superioris Stocchino &amp; Sluys, 2013, D. effusa Sluys, 2013, and D. naiadis (see: Stocchino et al. 2005, 2013, Sluys et al. 2013). Dugesia insolita represents the most extreme condition, in which almost none of the shell glands open into the bursal canal and the majority of the glands open through the roof of the female atrium.</p><p>From its opening into the atrium, the bursal canal in species of Dugesia generally curves smoothly towards the anterior end of the body to open into the copulatory bursa. However, in some species the bursal canal bends much more abruptly in the anterior direction, as is the case in D. capensis Sluys, 2007, D. bifida Stocchino &amp; Sluys, 2014 (see: Sluys 2007, Stocchino et al. 2014), and also in D. insolita .</p><p>The situation in which longitudinal muscle fibres run posteriorly from the penis bulb and eventually attach to the wall of the atrium is an uncommon condition among species of Dugesia . This feature is more or less present in D. debeauchampi de Vries, 1988 and D. liguriensis de Vries, 1988 but a situation very similar to that in D. insolita is expressed in D. milloti de Beauchamp, 1952 (see: de Vries 1988).</p><p>Presence of hyperplasic ovaries is characteristic of normally asexually reproducing specimens that have sexualized, i.e. have developed reproductive organs, either in the field or under laboratory conditions. In a similar way, after 4 months of rearing, a single specimen in our laboratory culture of animals of the population from the small tributary of the Matsiatra River developed reproductive organs (specimen RMNH.VER.21536.1, described above), including aberrant hyperplasic ovaries. In our experience, hyperplasic ovaries and, sometimes, poorly developed testes are the only abnormalities in such sexualized individuals, so that their copulatory apparatus can be interpreted as representing the normal condition of the species.</p><p>A high triangular head with long, pointed auricles has been described as a uniquely characteristic feature of D. milloti (de Vries 1988), but now it is known also from D. insolita .</p><p>Despite the fact that D. insolita shares with D. milloti its characteristic head shape, the atrial folds, and the layer of longitudinal muscles that runs between the wall of the atrium and the penis bulb, the two species are distinctly different. Dugesia insolita is the only species of Dugesia in which the diaphragm has evolved into a functional penis papilla. Nevertheless, this diaphragm has retained its primary function, formation of the spermatophore (see above). In D. milloti, the ancestral condition is still present in which the spermatophore is formed within the ejaculatory duct, with the funnel-shaped base of the spermatophore enveloping the diaphragm (see: de Vries 1988: fig. 14A).</p><p>Karyology</p><p>As for karyology, D. insolita shows an eudiploid chromosome complement of 18 chromosomes with basic number n = 9. Among Dugesia species, this haploid number is shared by only six other species: D. sicula and D. maghrebiana Stocchino et al., 2009, from the Mediterranean Region; D. arabica from Yemen; D. aethiopica, D. afromontana Stocchino &amp; Sluys, 2012, and D. bifida Stocchino &amp; Sluys, 2014 from the Afrotropical Region (cf. Stocchino et al. 2002, 2004, 2009, 2012, 2014, Harrath et al. 2013) (Table 4).</p><p>Dugesia insolita shares with D. bifida the condition in which fissiparous individuals exhibit a diploid chromosome complement, while in all of the other above-mentioned five species the fissiparous populations are always triploid and/or mixoploid, while species with sexual populations are diploid (viz., D. sicula and D. arabica). However, D. insolita differs from D. bifida in that its diploid karyotype is characterized by metacentric chromosomes, excepting chromosome 3, which is submetacentric, while all chromosomes of D. bifida are metacentric. Dugesia insolita and D. bifida represent the easternmost records of species with a basic chromosomal number of n = 9.</p></div>	https://treatment.plazi.org/id/206687B6FFDAFFA3FC5BA78FFB4C0613	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Stocchino, Giacinta Angela;Sluys, Ronald;Solà, Eduard;Riutort, Marta;Manconi, Renata	Stocchino, Giacinta Angela, Sluys, Ronald, Solà, Eduard, Riutort, Marta, Manconi, Renata (2024): The long-eared freshwater planarians of Madagascar form a separate phylogenetic clade within the genus Dugesia (Platyhelminthes: Tricladida), with the description of two new species. Zoological Journal of the Linnean Society (zlae 143) 202 (4): 1-29, DOI: 10.1093/zoolinnean/zlae143, URL: https://doi.org/10.1093/zoolinnean/zlae143
