taxonID	type	description	language	source
03BFF20FD3644963FC81FC509F77FB8D.taxon	materials_examined	Material examined Neotype MZUSP PL 2074 (field number F 6387): Parque Estadual da Pedra Branca, Rio de Janeiro / RJ, Brazil (22 ° 56 ′ 13.5 ″ S, 043 ° 27 ′ 39.5 ″ W). F. Carbayo et al., collected (coll.), 13 December 2014. Cephalic region: horizontal sections on two slides; ovarian region: horizontal sections on eight slides; piece behind ovarian region: horizontal sections on ten slides; prepharyngeal region: transverse sections on ten slides; pharynx: sagittal sections on five slides; copulatory apparatus: sagittal sections on ten slides. SMF N 574: Barreira, Teresópolis, Rio de Janeiro / RJ, Brazil. E. Bresslau coll., 1914. Cephalic region preserved in balsam on one slide; pre-pharyngeal region: transverse sections on one slide; pharynx: sagittal sections on two slides; copulatory apparatus: sagittal sections on three slides; tail preserved in balsam on one slide. SMF N 628: Barreira, Teresópolis, Rio de Janeiro / RJ, Brazil. E. Bresslau, Coll. 1914. Cephalic region and tail preserved in balsam on one slide; pre-pharyngeal region: transverse sections on one slide; pharynx and copulatory apparatus: sagittal sections on three slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	discussion	Note We could not confirm conspecificity of some specimens studied by Riester (1938) and specimens examined by Froehlich (1955 a, 1956 a, 1958) (see Supporting Information, Table S 1).	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	distribution	Distribution Areas covered with Atlantic forest in the municipalities of Rio de Janeiro and Teresópolis, state of Rio de Janeiro.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median black band, bounded on either side by a fine white-yellow stripe, externally to which is a very thin black line, and this is externally bordered by a marginal reddish-iron band. Inner pharynx musculature with two layers. Anteroventral region of penis bulb more developed. Penis papilla with dorsal insertion strongly displaced anteriorly to level the ventral insertion.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	materials_examined	Type locality Rio de Janeiro, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	description	External aspect Live animals ~ 70 mm long and 4 mm wide. Elongated body, with margins approximately parallel; anterior end rounded, posterior pointed. Dorsum convex, ventral side slightly convex. Dorsal colour constituted by a median black band (one-third of body width), bounded on either side by a fine white-yellow stripe (twoninths), externally to which is a very thin black line (one-ninth), and this is bordered externally by a marginal reddish-iron band (one-third) (Fig. 4 A). Ventral body surface grey with dark margins at cephalic end, and saffron yellow on the remaining surface. After fixation, body colour faded slightly. Two types of eyes: (1) a conical type (zuckerhutförmig, sugarloaf shaped), 90 µm high and 40 µm in width; and (2) a cup-shaped type, 40 µm in diameter, with the former contouring the anterior end. Cup-shaped-type eyes spread as a band, one-third of body width on either side. In the posterior half of the body, the eyes even reach to the median black band. Sensory pits simple invaginations, 20 μm deep, located ventromarginally in a single row from the very anterior body tip up to at least the first one-quarter of body length. Relative position mouthto-body length, 48 %. Relative position gonoporeto-body length, 61 %. Internal morphology Creeping sole 97 % of body width. Very abundant rhabditogen cells piercing dorsal and marginal epidermis. Glandular margin absent. The cutaneous musculature comprises three layers, namely a subepithelial circular layer, followed by a double diagonal layer with decussate fibres, and then a well-developed longitudinal layer. Muscle fibres of the longitudinal, innermost layer (30 µm thick) arranged into bundles with 15 – 35 fibres each. CMI, 8 %. Three parenchymal muscle layers present, all weakly developed: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres (Fig. 4 B). Ventral nerve plate present. Mouth situated at a distance from the root of the pharynx equivalent to 67 % of pharyngeal pocket length. Pharynx cylindrical (Fig. 4 C). Oesophagus-topharynx ratio, 12 %. Outer pharyngeal musculature consisting of a subepithelial layer (6 µm) of longitudinal muscle, followed by a layer (30 µm) of circular fibres. Inner pharynx musculature consisting of a subepithelial layer of circular fibres (40 µm), followed by a layer of of longitudinal fibres (10 µm). Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance equivalent to 17 % of body length; posterior follicles near to the pharynx root. Penis bulb oval, relatively strongly developed, more developed ventro-anteriorly; it extends from 0.5 mm anterior to penis papilla to being level with the gonopore region (specimen MZUSP PL 2074). Penis bulb consists of tightly packed muscle fibres variously orientated anteriorly, and longitudinally and obliquely orientated posteriorly; in this posterior region, fibres on the right side run obliquely downwards to embrace the ventro-anterior portion of the female atrium and, subsequently, presumably anchor on ventral epidermis. Sperm ducts are dorso-internal to the ovovitelline ducts. Lateral to penis insertion, sperm ducts bend medially and then ventrolaterally to penetrate the bulb. The ducts subsequently run anteriad and join to continue posteriorly and sinuously as the ejaculatory duct (Fig. 4 D). Ejaculatory duct traverses central region of penis papilla to open at its tip. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 5 - µm-thick layer of circular muscles. Wall of ejaculatory duct stained reddish, consisting of a cuboidal, ciliated epithelium, surrounded by circular muscles, which constitute a muscular cylinder with a diameter equivalent to three times the diameter of the duct (Fig. 5 A). Penis papilla very long, 5 mm, with dorsal and ventral insertions at the same horizontal plane (Fig. 4 D). This papilla is cylindrical along most of its length; total length equal to 17 times its diameter, entirely occupying both male and female atria; dorsal insertion strongly displaced anteriorly and very ventrally, so that it projects from the anteroventral portion of the male atrium. Subepithelial musculature consisting of a dense, 42 - µm-thick layer of circular muscle, tending to decussate distally, followed by a single layer of longitudinal muscle. Epithelium of papilla pierced by reddish secretions. Poor condition of sections does not display further histological details. Male atrium long and ample, not folded. Ovaries oval shaped, 250 µm in maximal length, and situated at a distance from anterior end equivalent to one-fifth of body length. Ovovitelline ducts emerge from an unknown region of ovaries, lateral to female atrium. Then, they curve medially and join to form the common glandular ovovitelline duct, dorsally to the posterior third of this atrium (Fig. 4 D). Common glandular ovovitelline duct as long as one-third of female atrium length and communicates with the female genital duct; this duct is a projection of the posterior region of the female atrium and is dorso-anteriorly orientated. The female genital duct is lined with an epithelium provided with intraepithelial gaps (Fig. 5 B). Poor condition of sections does not display further histological details. Female atrium without folds, two to three times as long as male atrium, lined with a tall epithelium, which posteriorly presents intraepithelial gaps. Atrial epithelium surrounded by a thin layer of longitudinal muscle fibres, followed by a layer of muscles with crisscross arrangement; thickness of both layers equal to 25 µm, distally doubling in thickness.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3644963FC81FC509F77FB8D.taxon	discussion	Remarks The original description of the dorsal chromatic pattern of this species may be ambiguous (“ Sides and foot coloured dirty ‘ orpiment orange’; above with two stripes on each side of pale ‘ primrose-yellow’, edged externally with black; on centre of the back a stripe of glossy black ”; Darwin, 1844, p. 244). From Darwin’s description, Marcus deduced that ‘ an additional black lateral stripe should exist on each side, albeit only the external stripe was mentioned’ (Marcus, 1952 [original in Portuguese]). No specimen matching Marcus’ description was ever found, but this ambiguity is removed in Darwin’s field notes. In an entry on 17 June, 1832, he states: “ Colours: back with glossy black stripe; on each side of this a primrose white one edged externally with black; these stripes reach to extremities, & become uniformly narrower. sides & foot dirty ‘ orpiment orange’ ” (in Keynes, 2000, p. 47). We found this precise colour pattern displayed by the specimen MZUSP PL 2074 from the type locality. Hence, it is most probable that this specimen and that found by Darwin are conspecific. A conclusive comparison is not possible because there is no record of Darwin’s specimen in the Natural History Museum, London, where the material collected by Darwin is deposited (H. Jones, personal communication).	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36B4966FCB7FB7798D5F9F0.taxon	materials_examined	Material examined Holotype MZUSP PL 2072 (field number F 5236): Nova Iguaçú / RJ, Brazil. Cristiano Sampaio et al., coll., 4 – 6 April 2012. Cephalic region: horizontal sections on eight slides; ovarian region: sagittal sections on five slides; pre-pharyngeal region: transverse sections on ten slides; pharynx: sagittal sections on six slides; copulatory apparatus: sagittal sections on 11 slides. Paratype MZUSP PL 2090 (field number F 7001): Teresópolis, Rio de Janeiro / RJ, Brazil. E. M. Froehlich coll., 20 July 1952. Copulatory apparatus: sagittal sections on 18 slides. Paratype SMF N 460: Barreira, Teresópolis, Rio de Janeiro / RJ, Brazil. Ernest Bresslau coll., 25 February 1914. Cephalic region and tail preserved in balsam on one slide; pre-pharyngeal region: transverse sections on one slide; pharynx and copulatory apparatus: sagittal sections on two slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36B4966FCB7FB7798D5F9F0.taxon	distribution	Distribution Areas covered with Atlantic forest in the municipalities of Nova Iguaçú and Teresópolis, State of Rio de Janeiro, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36B4966FCB7FB7798D5F9F0.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median luminous orange band, bounded on either side by a black stripe, externally to which is a greenish-yellow stripe, and this again is bordered externally by a marginal black band. Inner pharynx musculature with two layers. Anterodorsal region of penis bulb more developed. Penis papilla with dorsal insertion strongly displaced anteriorly and very ventrally, as long as 18 times its diameter.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36B4966FCB7FB7798D5F9F0.taxon	etymology	Etymology The specific epithet is a Tupi (indigenous Brazilian tribe) word that means mound (Tibiriçá, 1984), referring to the hilly terrain where the species was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36B4966FCB7FB7798D5F9F0.taxon	description	External aspect Live animals 45 mm long when creeping. Body margins parallel; anterior end rounded, posterior end pointed. In preserved specimens, dorsum convex, ventral side slightly convex. Dorsal colour pattern of the preserved specimen F 5236 constituted by a median luminous orange band (one-third of body width), bounded on either side by a black stripe (1 / 12 th), externally to which is a greenish-yellow stripe (1 / 12 th), and this again is bordered externally by a marginal black band (one-sixth) (Fig. 6 A). Ventral body surface pale orange. In specimen F 7001 (preserved for> 60 years), black became brownish, remaining colours became cream. Two types of eyes: a conical type, 80 µm high and 30 µm wide; and a cup-shaped type, 50 µm in diameter. The former type of eye contours the anterior portion of the body. Four millimetres behind anterior end, cup-shaped-type eyes spread dorsally onto the yellowish stripes until rear end of body, where the eyes are sparsely distributed. Sensory pits simple invaginations, 25 μm deep, located ventromarginally in a single row and extending from the very anterior end up to at least the first one-fifth of body length. Relative position mouth-to-body length, 55 %. Relative position gonopore-to-body length, 67 %. Internal morphology Creeping sole 77 % of body width. Glandular margin absent. Cutaneous musculature same as in the type species of the genus in terms of organization. CMI, 7 %. Muscle fibres of the longitudinal layer (15 – 20 µm thick) arranged into bundles with ten to 30 fibres each. Parenchymal musculature same as in the type species of the genus in terms of organization and relative development (Fig. 6 B). Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 54 % of pharyngeal pocket length. Pharynx cylindrical (Fig. 6 E). Oesophagus-to-pharynx ratio, 18 %. Pharyngeal musculature (Fig. 6 D) same as in the type species of the genus in terms of organization and development, with the exception of the inner longitudinal layer, which reaches 35 µm in thickness. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 13 % of body length; posteriormost near the pharynx root. Penis bulb oval shaped, relatively strongly developed, consisting of tightly packed muscle fibres variously orientated. The bulb is more developed dorso-anteriorly, and extends from 0.5 mm anterior to penis papilla up to the gonopore region (specimen MZUSP PL 2072). Diagonal fibres surrounding distal section of penis bulb run obliquely downwards on the right side of the body to embrace ventrally the anterior portion of the female atrium. Sperm ducts run laterally to the ovovitelline ducts. Laterally to ventral insertion of penis papilla, sperm ducts bend anteriorly and medially, then penetrate penis bulb ventrolaterally, and maintain this course until they join each other. From this point onwards, the ejaculatory duct runs posteriorly, centrally traversing the penis papilla to open at its tip. Sperm present in extrabulbar portion of sperm ducts; in one specimen, sperm was also present at the anterior section of the intrabulbar lumen. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 5 - to 8 - µm-thick layer of circular muscles. Proximal half of ejaculatory duct lined with cuboidal, ciliated epithelium and pierced by gland cells producing weakly erythrophil, fine granules; distal half lined with squamous, non-ciliated epithelium pierced by gland cells producing erythrophil, coarse granules. Anterior half of this duct surrounded by circular muscles, which constitutes a muscular cylinder in the distal portion with a diameter equivalent to five times the diameter of this duct. Penis papilla thin, somewhat conical, because it narrows very gradually, and very long; total length equal to 18 times its diameter, occupying almost entirely both male and female atria; dorsal insertion strongly displaced anteriorly and very ventrally, so that it projects from anteroventral portion of male atrium. Squamous lining epithelium of the penis papilla pierced by numerous gland cells producing xanthophyl and erythrophil fine granules, and underlain by a dense 85 - µm-thick layer of circular muscle, distally tending to decussate, followed by a single layer of longitudinal muscle (Fig. 6 F). Male atrium not folded (Fig. 7 A, B), lined with a low, non-ciliated epithelium, and underlain by a 5 - µm-thick layer of circular muscles, followed by an equally thin layer of longitudinal fibres. A set of muscle fibres is also present as longitudinal fibres extending around the gonopore canal laterally to the male atrium, and being intermingled with those of the penis bulb. Ovaries oval shaped, 300 µm in maximal diameter, and situated at a distance from anterior end equivalent to 22.5 % of body length. Ovovitelline ducts emerge from the dorsolateral aspect of ovaries. The oviducts run laterally to the female atrium, curve medially and then join to form the common glandular ovovitelline duct, dorsally to this atrium. Common glandular ovovitelline duct as long as 40 % of female atrium length, communicating with female genital duct (Fig. 7 B); this duct is a projection of the posterior region of the female atrium, bent dorsally and lined with a squamous, non-ciliated epithelium. A cyanophil fibrillar mass, 5 – 10 µm thick, is immediately present underneath this epithelium of the female atrium, with very thin circular filaments (<1 µm) resembling muscle fibres in stain affinity and orientation, followed by a layer of longitudinal muscle fibres. Female atrium without folds, as long as three times that of male atrium, and lined with a cuboidal-tocolumnar epithelium, which is pierced by glands producing xanthophil granules and is underlain by a thin layer of muscle fibres in a criss-crossed arrangement, followed by a loose layer (25 µm) of longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	description	FIGS 8 AND 9	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	materials_examined	Material examined Holotype MZUSP PL 2089 (field number F 7000): Parque do Estado, São Paulo / SP, Brazil. O. Françoso and E. M. Froehlich, coll., 29 May 1985. Cephalic region: sagittal sections on ten slides; ovarian region: horizontal sections on 12 slides; pre-pharyngeal region: transverse sections on ten slides; pharynx: sagittal sections on 20 slides; copulatory apparatus: sagittal sections on 35 slides. Paratype Marcus’ specimen (1951) (from EMF collection): Eldorado neighborhood, West zone of the city of São Paulo / SP, Brazil. E. Marcus and E. M. Froehlich, coll., 1 November 1947. Copulatory apparatus: sagittal sections on five slides. Paratype MZUSP PL 2061 (field number F 1022): Mogi das Cruzes / SP, Brazil (23 ° 27 ′ 0 5.5 ″ S, 046 ° 13 ′ 40.8 ″ W). F. Carbayo, coll., 27 May 2006. Cephalic region: transverse sections on 13 slides; ovarian region: horizontal sections on seven slides; pre-pharyngeal region: transverse sections on five slides; pharynx: sagittal sections on eight slides; copulatory apparatus: sagittal sections on five slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	distribution	Distribution Areas covered with secondary Atlantic forest in the municipalities of São Paulo and Mogi das Cruzes, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the municipality where the species was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median orangeochre band, bounded on either side by a thin whitish stripe, and in turn this is bordered externally by a thin black line, externally to which is a grey band with dense white mottling, externally to which is a thin marginal black line. Longitudinal parenchymal muscle fibres disposed around intestine as a loose muscular tube. Diameter of muscular cylinder surrounding ejaculatory duct equivalent to 1.3 times the diameter of this duct.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36E4965FF5DF9869B69FB4D.taxon	description	External aspect Preserved animals 50 mm long and 4 mm wide. Body margins parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side flat. On histological slides, body 1.25 mm in height. In live animals, dorsal colour pattern constituted by a median orange-ochre band (one-third of body width) that passes into the black of the cephalic region (onefifth of body width), bounded on either side by a thin whitish stripe (one-ninth), and in turn this is bordered externally by a thin black line, externally to which is a grey band (the two latter totalling two-ninths) with dense white mottling, externally to which is a thin marginal black line (Fig. 8 A). This marginal band was discovered in a detailed examination of the preserved specimen; this band remained hidden in dorsal view owing to the strong convexity of the dorsum. Ventral body surface whitish, with blackish margins. In preserved specimen F 1022, whitish, blackish and greyish colours lost brightness; orange-ochre faded. In specimen F 7000, median band was ochre-brown bordered on either side by cream lines, in turn this was bordered by a brown green line, externally to which was a beige grey band. Two types of eyes: a conical type, 90 µm high and 40 µm in width; and a cup-shaped type, 40 µm in diameter. The former type contours the anterior tip of the body. Behind the first millimetre, cup-type eyes spread progressively onto the dorsum, reaching the whole dorsum from the pharynx region until posterior end. Sensory pits simple invaginations, 55 μm deep, located ventromarginally in a single row, at least along first third of body from very anterior end. Relative position mouth-tobody length, 55 %. Relative position gonopore-to-body length, 71 %. Internal morphology Creeping sole 78 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 9.5 % of body height. Muscle fibres of the longitudinal, innermost layer (40 µm thick) arranged into bundles with 18 – 20 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse fibres, and subintestinal layer with transverse fibres. Longitudinal fibres around intestine present, constituting a loose tube with fibres mixing with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 44 – 46 % of pharyngeal pocket length. Pharynx cylindrical, with dorsal insertion slightly posterior to ventral one (Fig. 8 B). Oesophagus-topharynx ratio, 15 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (70 µm) of circular fibres, the innermost of which are intermingled with longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (70 µm) of circular fibres and an innermost layer (30 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 22 % of length of body; posteriormost follicles near pharynx root. Penis bulb elongated, extending posteriorly from 0.5 mm (specimen MZUSP PL 2061) anterior to penis papilla to the level of the gonopore. The bulb consists of tightly packed muscle fibres, variously orientated. Diagonal fibres on the right side of the bulb run obliquely downwards to embrace ventrally the anterior portion of female atrium. These fibres apparently cross the nerve plate to anchor on the ventral epidermis. Sperm ducts run dorsally to the ovovitelline ducts. Laterally to penis insertion, sperm ducts bend medially and then penetrate the penis bulb ventrolaterally. Subsequently, the ducts recurve and separately open into the sinuous ejaculatory duct (Figs 8 C, 9 A). The latter traverses the central region of the penis papilla to open at its tip. Sperm present in sperm ducts, also along its course within penis bulb. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 20 - µm-thick layer of circular muscles. Ejaculatory duct lined with columnar, ciliated epithelium; epithelium of extrapenial portion pierced by glands producing weakly erythrophil, fine granules, and being surrounded by 60 - µm-thick layer of circular muscle fibres. Epithelium of intrapenial portion of ejaculatory duct richly pierced by glands producing erythrophil granules and surrounded by a 20 - µm-thick layer of circular muscles, which constitutes a muscular cylinder possessing a diameter equivalent to 1.3 times the diameter of this duct (Fig. 9 B). Penis papilla long and cylindrical along most of its length, with dorsal insertion slightly anterior to the ventral. Total length of papilla equal to ten times its greatest diameter, and occupying male atrium and almost entire female atrium; lined with a cuboidal, non-ciliated epithelium, pierced by two types of gland cells, producing coarse erythrophil granules and fine cyanophil granules, respectively. Erythrophil glands pierce anterior and distal third of penis; necks of cyanophil glands pierce dorsal epithelium of second third, and sometimes also the ventral one, and pack space immediately beneath epithelium so that subepithelial muscle fibres are apart from it (Fig. 9 C). Subepithelial musculature of penis papilla consisting of dense 80 - to 90 - µm-thick layer of circular muscle, the fibres of which are decussate distally, followed by a 10 - to 20 - µm thick layer of longitudinal muscle. Male atrium not folded, lined with a low, non-ciliated epithelium, pierced by some glands secreting amorphous substance, and underlain by 7 - µm-thick layer of circular muscles, followed by a nearly inconspicuous layer of longitudinal fibres. Ovaries elongated, 1.1 mm in maximal anteroposterior diameter, 0.2 mm in dorsoventral width, and situated at a distance from anterior end equivalent to 22 % of body length. Ovovitelline ducts emerge from dorsolateral aspect of ovaries laterally to female atrium, curve medially and join common glandular ovovitelline duct dorsally to this atrium. Common glandular ovovitelline duct as long as a third of female atrium length; it communicates with female genital duct, which is a dorso-anteriorly orientated projection of the posterior region of the female atrium (Fig. 9 A) and is lined with a 25 - µm-tall cuboidal, ciliated epithelium. Female atrium without folds, posterior portion funnel shaped; as long as 1.5 times that of male atrium; anteriorly lined with a cuboidal epithelium; posteriorly lined with a tall epithelium with stratified appearance and intraepithelial holes. Epithelium pierced by two types of glands, producing erythrophil granules and, less abundantly, xanthophil. Epithelium underlain by 5 - to 45 - µm-thick layer of circular-to-decussate muscle fibres, followed by some longitudinal fibres, and then a layer (50 – 150 µm thick) of thicker longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36D4958FF5EFB39982FFDD7.taxon	materials_examined	Material examined Holotype MZUSP PL 2071 (field number F 5134): Parque Jardim Previdência, São Paulo / SP, Brazil (23 ° 34 ′ 48.7 ″ S, 046 ° 43 ′ 39.3 ″ W). F. Carbayo et al., coll., 5 February 2006. Cephalic region: horizontal sections on four slides; ovarian region: horizontal sections on two slides; region behind ovaries: horizontal sections on ten slides; pharynx: sagittal sections on six slides; copulatory apparatus: sagittal sections on eight slides. Paratype MZUSP PL 2060 (field number F 0509): Parque Ecológico do Tietê, São Paulo / SP, Brazil (23 ° 29 ′ 27.1 ″ S, 046 ° 31 ′ 0 1.6 ″ W). F. Carbayo, coll., 27 May 2005. Pharynx and post-pharynx region: sagittal sections on six slides (incompletely mature). Paratype MZUSP PL 666 (field number F 1163): Parque Jardim Previdência, São Paulo / SP, Brazil (23 ° 34 ′ 46.8 ″ S, 046 ° 43 ′ 39.0 ″ W). O. M. Junior, coll., 17 June 2007. Pharynx: sagittal sections on 13 slides; copulatory apparatus: sagittal sections on seven slides (incompletely mature).	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36D4958FF5EFB39982FFDD7.taxon	distribution	Distribution Areas covered with secondary Atlantic forest and garden in the municipalities of São Paulo and Mongaguá, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36D4958FF5EFB39982FFDD7.taxon	etymology	Etymology The specific epithet is a Tupi (indigenous Brazilian tribe) word formerly referring to the city of São Paulo, where the species was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36D4958FF5EFB39982FFDD7.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median traffic red band, bounded on either side by a black stripe, externally to which is a white stripe, and in turn this is externally bordered by a black band, externally to which is a thin marginal white line. Dorsal and ventral insertions of penis papilla at the same transverse plane. Wall of female atrium not folded. Femaleto-male atrial length ratio, 2.0.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD36D4958FF5EFB39982FFDD7.taxon	description	External aspect Live animals 25 mm long and 4 mm wide. Body relatively short, with margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side slightly convex. Dorsal body surface constituted by a median traffic red band (1 / 3.5 th of body width), passing into black part of cephalic region (one-sixth of body length) and of posterior tip, bounded on either side by a black stripe, externally to which is a white stripe (one-seventh), and in turn this is bordered externally by a black band (one-seventh), externally to which is a thin marginal white line (Fig. 10 A). Ventral side whitish. After fixation, dorsal black colour faded, red passed into yelloworange and white into cream. Two types of eyes: a conical type, contouring only anterior end; and a cup-shaped type, spreading dorsally onto the black bands. Sensory pits simple invaginations, 25 μm deep, located ventromarginally in a single row from the very anterior end up to at least first one-fifth of body length. Relative position mouth-to-body length, 42.5 %. Relative position gonoporeto-body length, 64 %. Internal morphology Creeping sole ~ 85 % of body width, as measured on horizontal or sagittal sections. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 5 % of body height. Muscle fibres of the longitudinal, innermost layer (25 µm thick) arranged into bundles. Three parenchymal muscle layers present, all weakly developed: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse fibres. Additionally, longitudinal fibres surround intestine as a loose tube, the ventral fibres of which are mixed up with those of the subintestinal layer. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 50 – 65 % of pharyngeal pocket length. Pharynx cylindrical, with dorsal insertion slightly posterior to ventral one (Fig. 10 B). Oesophagus-to-pharynx ratio, 12 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (45 µm) of circular fibres, the innermost of which are intermingled with longitudinal ones. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (40 µm) of circular fibres and an innermost layer (10 µm) of longitudinal fibres (Fig. 10 C). Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 25.5 % of body length; posteriormost follicles a little anterior to pharynx root. Penis bulb elongated, somewhat oval, extending from 0.3 mm anterior to penis papilla; from thereon the bulb extends ventrally, whereas dorsally it extends posteriorly for only short distance. The bulb consists of variously orientated packed muscle fibres. Diagonal fibres on the right side of the bulb run obliquely downwards to embrace ventrally the anterior portion of the female atrium. These fibres apparently mix with the subintestinal parenchymal musculature, but their anchor point could not be discerned. Sperm ducts run dorsally to the ovovitelline ducts. Sperm ducts bend medially behind insertions of penis papilla and penetrate penis bulb ventrolaterally. The ducts subsequently recurve to join and thus form the ejaculatory duct, which sinuously posteriad, then traverses the penis papilla to open at its tip. Distal portion of sperm ducts containing sperm, along their course both outside and inside penis bulb. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 6 - µm-thick layer of circular muscles. Extrapapillar portion and anterior portion of intrapapillar section of ejaculatory duct lined with columnar ciliated epithelium, pierced by glands producing erythrophil fine granules. Remaining section of the ejaculatory duct also columnar and ciliated, pierced by numerous gland cells producing erythrophil granules. Entire ejaculatory duct surrounded by a layer of circular muscle, 35 µm thick around anterior portion, otherwise 10 µm thick (Fig. 10 D). Diameter of muscular cylinder surrounding ejaculatory duct is equivalent of 3.5 times the diameter of this duct. Penis papilla is long, horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 11 A, B). Total length of papilla equal to five times its greatest diameter, and completely filling both male and female atrium; papilla lined with a cuboidal epithelium, pierced by gland cells producing coarse erythrophil granules; glands producing cyanophil granules packed in space immediately beneath epithelium of the dorsal second third of penis papilla, so that subepithelial muscle fibres are separated from it (Fig. 10 D). Penial subepithelial musculature consisting of dense 90 - µm-thick layer of circular muscle, followed by a layer (25 µm thick) of longitudinal muscle. Male atrium ample, not folded, lined with a low, non-ciliated epithelium, pierced by glands secreting erythrophil amorphous substance and underlain by a 15 - µm-thick layer of circular muscles, followed by an inconspicuous layer of longitudinal fibres. Ovaries incompletely developed, 500 µm in length along anteroposterior body axis, located above ventral nerve plate and situated at a distance from anterior end equivalent to ~ 20.5 % of body length. Laterally to the female atrium, the oviducts curve medially and join to form a common glandular ovovitelline duct. This duct is as long as one-third of the female atrium length and communicates with the female genital duct, the latter being a funnel-shaped, anterodorsally orientated projection of the posterior region of the female atrium. Female atrium funnel shaped and without folds, as long as two times the male atrium length (Fig. 11 B); in its anterior portion it is lined with cuboidal epithelium, posteriorly with a 40 - µm-high epithelium with stratified aspect; pierced by two types of glands producing cyanophil and xanthophil granules, respectively. Epithelium underlain by a 5 - to 40 - µm-thick layer of circular-to-decussate muscle fibres, followed by a 25 - µm-thick layer of longitudinal fibres. Additionally, 4 - µm-thick longitudinal fibres around these muscles constitute a kind of muscle coat.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD350495FFF49FDAA98B3FEAF.taxon	materials_examined	Material examined Holotype MZUSP PL 2079 (field number F 6660): Cidade Turística de Paranapiacaba, Santo André / SP, Brazil (23 ° 46 ′ 38.9 ″ S, 046 ° 18 ′ 0 5.2 ″ W). F. Carbayo et al., coll., 5 June 2015. Cephalic region: horizontal sections on four slides; ovarian region: horizontal sections on nine slides; region behind ovaries: horizontal sections on 14 slides; pre-pharyngeal region: transverse sections on 16 slides; pharynx: sagittal sections on 17 slides; copulatory apparatus: sagittal sections on 20 slides. Paratype MZUSP PL 2087 (field number F 6998): Cidade Turística de Paranapiacaba, Santo André / SP, Brazil. E. M. Froehlich et al., coll., 13 May 1990. Copulatory apparatus: sagittal sections on 39 slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD350495FFF49FDAA98B3FEAF.taxon	distribution	Distribution Only known from the type locality.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD350495FFF49FDAA98B3FEAF.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the district where the type material was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD350495FFF49FDAA98B3FEAF.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median stripe of orange-ochre, bounded on either side by a black line, externally to which is a broader white line, and in turn this is externally bordered by a thin black line, externally to which is a grey band with dense white mottling, bordered by a thin marginal black line. Diameter of muscular cylinder round ejaculatory duct equivalent to 0.5 times the diameter of this duct. Female-tomale atrial length ratio, 3.5 – 4.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD350495FFF49FDAA98B3FEAF.taxon	description	External aspect Preserved animals up to 50 mm long, and 4 mm wide. Body margins nearly parallel; anterior end rounded, posterior end pointed. Dorsum strongly convex, ventral side slightly convex. Dorsal body surface constituted by a median stripe of orange-ochre (onequarter of body width) that passes into the black of cephalic region (one-fifth of body length), bounded on either side by a black line (1 / 16 th) posteriorly merging with each other, externally to which is a broader white line (one-eighth), and this is externally bordered by a black stripe (1 / 16 th), externally to which is a grey stripe (one-eighth) with dense white mottling, then bordered by a marginal thin black line (Fig. 12 A). Ventral body surface is whitish with blackish margins. In preserved specimen F 6660, dorsal colour slightly faded. In preserved specimen F 6998, stripes were varied from faded black grey to pearl orange. Two types of eyes: a conical type contouring only anterior end; and a cup-shaped type, spreading dorsally onto the black stripes. Sensory pits simple invaginations, 40 μm deep, located ventromarginally in a single row from the very anterior tip to at least the end of the anterior fifth of the body. Relative position mouth-to-body length, 61 %. Relative position gonopore-to-body length, 74 %. Internal morphology Creeping sole 78 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 8 % of body height at pre-pharyngeal region. Longitudinal muscle fibres of the innermost layer (35 – 50 µm thick) arranged into bundles with 17 – 30 fibres each. Three parenchymal muscle layers present: a dorsal layer of decussate diagonal fibres, a loose supraintestinal layer of transverse and longitudinal fibres, and a loose subintestinal layer with traverse and longitudinal fibres (Fig. 12 B, C). Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 66 % of pharyngeal pocket length. Pharynx cylindrical (Fig. 12 D). Oesophagus-topharynx ratio, 26 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (70 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (90 µm) of circular fibres and an innermost layer (15 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes (at a distance from anterior end equal to 22 % of body length) behind ovarian region, posteriormost near root of pharynx. Penis bulb elongated, strongly developed, consisting of tightly packed muscle fibres variously orientated in its anterior section, while posteriorly the fibres are orientated diagonally, extending from 1 mm (holotype) anterior to penis papilla to the level of the gonopore, even more posteriorly in the dorsal body region. Diagonal fibres on the right side of the bulb run obliquely downwards to embrace ventrally the anterior portion of the female atrium; further course of these fibres could not be determined owing to tissue damage. Sperm ducts run dorsolaterally to the ovovitelline ducts. Laterally to penis bulb, sperm ducts bend anteriorly and medially, and then ventrolaterally penetrate the bulb. The ducts subsequently recurve and join to form an unpaired, sinuous sperm duct directed posteriorly (Fig. 13 A, B). The unpaired portion continues as an ejaculatory duct inside the penis papilla, which traverses centrally to open at its tip. Sperm present in sperm duct, both along its course outside and in part of anteriorly directed portion inside penis bulb. Epithelium of sperm ducts and unpaired sperm duct pierced by glands producing minute weakly cyanophil granules; surrounded by a 7 - µm-thick layer of circular muscles. Ejaculatory duct lined by 15 µm, ciliated epithelium pierced by abundant glands producing erythrophil granules; surrounded by a layer of circular muscle fibres constituting a muscular cylinder with a diameter equivalent to 0.5 times the diameter of this duct. Penis papilla long, with dorsal and ventral insertions at the same transverse plane; it is horizontal and cylindrical, with a length (4 mm) equal to ten times its diameter, even occupying more than half of the female atrium. Papilla covered with a cuboidal, non-ciliated epithelium, pierced by gland cells especially distally, producing erythrophil granules (Figs 12 E, 13 D). Subepithelial penis musculature consisting of a dense 25 - to 105 - µm-thick layer of circular muscle, followed by a thin layer (12 µm) of longitudinal muscle. Male atrium not folded (Fig. 13 A, B), lined with a low, non-ciliated epithelium, pierced by glands producing an amorphous erythrophil substance and underlain by a 15 - µm-thick layer of circular muscles, followed by a thin layer of longitudinal fibres. Ovaries oval shaped, 550 µm in maximal anteroposterior diameter and 250 µm in dorsoventral diameter, and situated at a distance from anterior end equivalent to 16 % of body length. Ovovitelline ducts emerge from dorsolateral aspect of ovaries. The oviducts run laterally to the female atrium, then curve medially and join to form a common glandular ovovitelline duct, dorsally to this atrium. Common glandular ovovitelline duct as long as half of the female atrium length, communicates with female genital duct; the latter being a 40 - µm-long projection of the dorsoposterior region of the female atrium (Fig. 13 A, B, D). Female atrium long, without folds, as long as 3.5 – 4 times the male atrium length (Fig. 13 A, B, D) and lined with a tall epithelium (70 µm). Posterior section of female atrium lined with an epithelium with stratified aspect and some internal gaps; surrounded by a thin layer of longitudinal muscle fibres, followed by 70 - µm-thick layer of circular fibres, and a third layer (115 µm) of fibres in criss-cross arrangement. No evident distinction between atrial fibres and those of common muscle coat.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3574952FF5FFE599B45FEC2.taxon	materials_examined	Material examined All specimens were collected in Parque Estadual Serra do Mar, Caraguatatuba / SP, Brazil. Holotype MZUSP PL 2091 (field number F 7004; 23 ° 35 ′ 38.0 ″ S, 045 ° 25 ′ 47.1 ″ W): F. Carbayo et al., coll., 14 May 14 2016. Ovarian region: horizontal sections on eight slides; pre-pharyngeal region: transverse sections on seven slides; pharynx and copulatory apparatus: sagittal sections on 34 slides. Paratype MZUSP PL 2080 (field number F 6727; 23 ° 35 ′ 38.4 ″ S, 045 ° 25 ′ 46.8 ″ W): F. Carbayo et al., coll., 2 July 2015. Before sectioning, copulatory apparatus studied on images of orthogonal virtual sections obtained by µCT. Ovarian region: horizontal sections on eight slides; pre-pharyngeal region: transverse sections on nine slides; pharynx and copulatory apparatus: sagittal sections on eight slides. Paraype MZUSP PL 2126 (field number F 6976; 23 ° 35 ′ 38.0 ″ S, 045 ° 25 ′ 47.1 ″ W): F. Carbayo et al., coll., 1 January 2016. In absolute ethanol.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3574952FF5FFE599B45FEC2.taxon	distribution	Distribution Only known from borders of Atlantic forest in the municipality of Caraguatatuba, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3574952FF5FFE599B45FEC2.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the municipality where the type material was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3574952FF5FFE599B45FEC2.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median orangeochre band, bounded on either side by a black stripe, externally to which is a white stripe, and this in turn is externally bordered by a black stripe that merges into a grey band with white mottling, which fades gradually towards its outer margin, while this margin is bordered by a black stripe. Penis bulb extending anteriorly 0.8 mm from penis papilla. Wall of female atrium not folded.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3574952FF5FFE599B45FEC2.taxon	description	External aspect Live animals 60 mm long and 4 mm wide when extended. Body margins nearly parallel; anterior end rounded, posterior pointed. Dorsum convex, ventral side slightly convex. Dorsal body surface constituted by a median orange-ochre band (one-quarter of body width) that grades into the black of the cephalic region (one-ninth of body length) and of posterior end, bounded on either side by a black line (one-tenth) that join each other posteriorly, external to which is a white stripe (one-ninth), which join each other at the posterior end, and in turn this is externally bordered by a black stripe (one-ninth) that merges into a grey band (one-ninth) with white mottling and which gradually fades towards its outer margin. This margin is bordered by a black line (Fig. 14 A). Ventral body surface whitish with black margins. In preserved specimens, dorsal black colour became dark greyish, and grey became cream; whitish remained unchanged. Two types of eyes: a conical type contouring only the anterior end; and a cup-shaped type, spreading dorsally onto the surface external to the white stripes. Sensory pits simple invaginations, 50 μm deep, located ventromarginally in a single row from the very anterior end up to the end of first one-third of body. Relative position mouth-to-body length, 55 %. Relative position gonopore-to-body length, 69 %. Internal morphology Creeping sole 80 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 6 % of body height. Muscle fibres of the longitudinal innermost layer (15 µm thick) arranged into bundles with 20 – 30 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres rarely lateral to intestine. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 34 – 60 % of pharyngeal pocket length. Pharynx cylindrical, with dorsal insertion posteriorly placed (Fig. 14 B). Oesophagus-to-pharynx ratio, 14 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (55 µm) of circular fibres, intermingled with longitudinal ones at its innermost portion. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (35 µm) of circular fibres and an innermost layer (10 µm) of longitudinal fibres (Fig. 14 C). Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes (at a distance from anterior end equal to 25 % of body length) behind ovarian region, posteriormost near root of pharynx. Penis bulb elongated, extending from 0.8 mm anterior to penis papilla to the level of the gonopore region; strongly developed, consisting of tightly packed muscle fibres variously orientated, somewhat more developed dorsally than ventrally. Bulb fibres on the right side run obliquely downwards to embrace ventrally the anterior portion of the female atrium, subsequently anchoring presumably on ventral epidermis. Sperm ducts run dorsolaterally or dorsally to the ovovitelline ducts. Laterally to penis bulb, sperm ducts bend anteriorly and medially, and then ventrolaterally penetrate the bulb. The ducts subsequently join to form the ejaculatory duct, with initial section outside penis papilla, then crossing it centrally to open at its tip (Fig. 15 A). Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 5 - to 25 - µm-thick layer of circular muscles. Sperm contained in distal portion of sperm ducts, both along their course outside and inside penis bulb. Ejaculatory duct lined with tall, ciliated epithelium; pierced by numerous gland cells producing erythrophil granules, except in its first millimetre. Ejaculatory duct surrounded by a 25 - µm-thick layer of circular muscle fibres constituting a muscular cylinder with a diameter equivalent to 1.7 times the diameter of this duct. Penis papilla 5 mm long, horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 15 A – C). Total length of papilla equal to seven times its diameter. Papilla covered with a cuboidal epithelium pierced by gland cells producing erythrophil granules; dorsal epithelium of second third of papilla pierced by glands producing cyanophil granules; space immediately beneath this epithelium packed by necks of these cells (Fig. 15 D, E). Subepithelial musculature consisting of dense layer of circular muscle, 130 µm thick dorsally, 60 µm thick ventrally, followed by a 20 - µm-thick layer of longitudinal muscle. Male atrium ample, not folded, lined with a low, nonciliated epithelium and underlain by an 8 - µm-thick layer of circular muscles, followed by an 8 - µm-thick layer of longitudinal fibres. Additional longitudinal muscle fibres around atrium dorsal to gonopore canal. Ovaries are oval shaped, 450 µm in length in longitudinal axis, and situated at a distance from anterior end equivalent to one-quarter of body length. Ovovitelline ducts emerge from dorsolateral aspect of ovaries. Ovaries run laterally to posterior portion of female atrium, then curve medially to communicate with common glandular ovovitelline duct. The latter duct is as long as one-seventh of female atrium length and opens into the female genital duct, a projection of the dorsoposterior region of the female atrium. Female atrium without folds, cylindrical along most of its course, funnel shaped posteriorly; as long as 1.5 times that of male atrium (Fig. 15 A); funnelshaped portion lined with tall (20 – 25 µm) epithelium with stratified aspect, being pierced by two types of glands, producing erythrophil and xanthophil amorphous substance, respectively; cylindrical portion lined with cuboidal epithelium. Female atrium surrounded by 10 - to 35 - µm-thick layer of circular-to-decussate muscle fibres, followed by a 10 - to 65 - µm-thick layer of longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	description	FIG. 16	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	materials_examined	Material examined Holotype MZUSP PL 2066 (field number F 3562): Parque Estadual do Jurupará, Ibiúna / SP, Brazil. F. Carbayo, coll., February 2009. Cephalic region: horizontal sections on 14 slides; ovarian region: horizontal sections on 12 slides; pre-pharyngeal region: transverse sections on ten slides; pharynx and copulatory apparatus: sagittal sections on 33 slides. Paratype MZUSP PL 2081 (field number F 6907): Santana de Parnaíba / SP, Brazil (23 ° 26 ′ 57.2 ″ S, 047 ° 00 ′ 43.8 ″ W). F. Carbayo coll., 15 December 2015. Cephalic region: horizontal sections on 12 slides; ovarian region: horizontal sections on six slides; region behind ovaries: horizontal sections on six slides; prepharyngeal region: transverse sections on 19 slides; pharynx and copulatory apparatus: sagittal sections on 21 slides. Paratype MZUSP PL 2082 (field number F 6908): Santana de Parnaíba / SP, Brazil (23 ° 26 ′ 57.2 ″ S, 047 ° 00 ′ 43.8 ″ W). F. Carbayo coll., 12 December 2015. Pharynx and copulatory apparatus: sagittal sections on 11 slides. Paratype MZUSP PL 2083 (field number F 6909): Santana de Parnaíba / SP, Brazil (23 ° 26 ′ 57.2 ″ S, 047 ° 00 ′ 43.8 ″ W). F. Carbayo coll., 12 December 2015. Pharynx and copulatory apparatus (underdeveloped): sagittal sections on 17 slides. Paratype MZUSP PL 2088 (field number F 6999): Pico do Jaraguá, São Paulo / SP, Brazil (23 ° 27 ′ 28.7 ″ S, 046 ° 45 ′ 49.9 ″ W). E. M. Froehlich coll. 15 August 1951. Ovarian region: horizontal sections on 18 slides; prepharyngeal region: transverse sections on 12 slides; copulatory apparatus: sagittal sections on 43 slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	distribution	Distribution Secondary Atlantic forest in the municipalities of Ibiúna and Santana de Parnaíba, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of one of the municipalities where the type material was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median orangeochre stripe, bounded on either side by a black stripe, externally to which is a white stripe, and this in turn is externally bordered by a black band, externally to which is a thin marginal white line. Dorsal insertion of penis papilla strongly displaced anteriorly and very ventrally. Posterior wall of female atrium richly folded. Female-to-male atrial length ratio, 1.0.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35A4950FF48FEB69ED8FCA6.taxon	description	External aspect Live animals 40 – 50 mm long and 3 mm wide. Body margins nearly parallel; anterior end rounded, posterior pointed. Dorsum convex, ventral side slightly convex. Dorsal body surface constituted by a median orange-ochre band, varying from one-quarter to oneseventh of body width, that grades into the black of the cephalic region (one-ninth of body length) and posterior end, bounded on either side by a black stripe (onefifth), which merge posteriorly, externally to which is a white stripe (one-quarter), and this is bordered externally by a black band (one-third), externally to which is a thin marginal white line (Fig. 16 A). Ventral surface of the body whitish. In preserved specimens, the dorsal colour remained unchanged. Two types of eyes: a conical type, 100 µm high and 35 µm in width; and a cup-shaped type, 40 µm in diameter. The former contour the anterior end of the body. Cup-shaped-type eyes spread onto the dorsum, reaching whitish stripes all along the body. Sensory pits located ventromarginally in a single row, from the very anterior end up to the end of first quarter of body. Relative position mouth-to-body length, 50 – 58 %. Relative position gonopore-to-body length, 64 – 71 %. Internal morphology Creeping sole 82 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 6 – 8 % of body height. Muscle fibres of the longitudinal, innermost layer (30 µm thick) arranged into bundles with 15 – 25 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres around intestine constituting a loose tube whose fibres mix up with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 42 – 59 % of pharyngeal pocket length. Pharynx cylindrical (Fig. 16 B). Oesophagusto-pharynx ratio, 17 %. Outer pharyngeal musculature consisting of a subepithelial layer (8 µm) of longitudinal muscle, followed by a layer (85 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin (5 µm) subepithelial layer of longitudinal muscle, followed by a layer (75 µm) of circular fibres and an innermost layer (12 µm) of longitudinal fibres (Fig. 16 C). Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes behind ovaries, at a distance roughly equivalent to 10 % of body length; posteriormost follicles slightly anterior to root of pharynx. Penis bulb elongated, extending anteriad ~ 0.3 mm anterior to penis papilla; in the dorsal portion of the body, the penis bulb extends posteriorly to the level of the gonopore; in the ventral body region, the penis bulb extends posteriorly only to about halfway between the root of the penis papilla and the gonopore. The bulb is strongly developed, consisting of tightly packed muscle fibres variously orientated. Penis bulb muscle more densely packed dorsally than ventrally (Fig. 16 D, E). Bulb fibres on the right side run obliquely downwards to embrace ventrally the anterior portion of the female atrium, subsequently anchoring presumably on ventral epidermis. Sperm ducts run dorsolaterally to the ovovitelline ducts. Posteriorly to level of insertion of penis papilla, sperm ducts bend anteriorly and medially and then penetrate penis bulb ventrolaterally. The ducts subsequently recurve and join to form the ejaculatory duct. Extrapapillar portion of this duct considerably sinuous; traversing penis papilla centrally to open at its tip. Sperm present in distal portion of sperm duct, along its course both outside and inside penis bulb. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 20 - µm-thick layer of circular muscles. Anterior portion of ejaculatory duct lined with squamous-to-cuboidal, ciliated epithelium pierced by numerous gland cells producing erythrophil granules and surrounded by a coat of circular muscle fibres constituting a muscular cylinder with a diameter equivalent to three times the diameter of this duct. Penis papilla long, horizontal and progressively thinner towards its tip; this tip very thin. Dorsal insertion strongly displaced anteriorly and very ventrally, so that it projects from anteroventral portion of male atrium. Penis papilla completely fills male atrium and a variable part of the female atrium, probably owing to fixation artefacts. Total length of papilla equal to six to seven times its greatest diameter. Penis papilla covered with a cuboidal epithelium, pierced by gland cells producing fine erythrophil granules. Necks of these glands fill a moderate space immediately beneath epithelium of second third of penis papilla. Subepithelial musculature consisting of dense 45 - µm-thick layer of circular muscle, followed by a 30 - µm-thick layer of longitudinal muscle. Male atrium ample, not folded (Fig. 16 D, E), lined with a low epithelium and underlain by a 5 - µm-thick layer of decussate muscles. Circular muscle fibres surround both innermost portion of gonopore canal and nearby atrium. Ovaries oval shaped, 400 µm in length in longitudinal body axis and situated at a distance from anterior end equivalent to one-fifth of body length. Ovovitelline ducts emerge from dorsolateral aspect of ovaries. The oviducts run laterally to posterior portion of female atrium, then curve medially and join to form common glandular ovovitelline duct. This duct is as long as one-sixth of female atrium length and communicates with female genital duct; the latter is a very narrow projection of the posterior region of female atrium, lined with a 6 - µm-tall ciliated epithelium and surrounded by a small but distinct sphincter made up of circular muscle fibres. Female atrium narrow and long, somewhat funnel shaped; as long as male atrium (Fig. 16 D, E). Atrial wall very richly folded, either along anterior half or along almost all posterior half, thus narrowing its lumen. Lining epithelium mostly cuboidal, 12 µm high, posteriormost portion tall, 40 µm and with stratified aspect. Two types of gland cells pierce epithelium, producing cyanophil granules and erythrophil substance, respectively. Female atrium surrounded by a 5 - µm-thick layer of longitudinal muscle fibres, which do not occur on the posterior portion, followed by a 40 - µm-thick layer of decussate fibres. A loose muscle coat of thick (10 µm) longitudinal fibres envelopes this atrium.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	description	FIG. 17	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	materials_examined	Material examined All specimens were collected in Parque Estadual da Ilha do Cardoso, Cananeia / SP, Brazil (25 ° 05 ′ 28.4 ″ S, 047 ° 55 ′ 44.2 ″ W). J. Pedroni, coll., 18 October 2009. Holotype MZUSP PL 2067 (field number F 4096): Cephalic region: horizontal sections on five slides; ovarian region: horizontal sections on 12 slides; prepharyngeal region: transverse section on 12 slides; pharynx: sagittal sections on 17 slides; copulatory apparatus: sagittal sections on 45 slides. Paratype MZUSP PL 2068 (field number F 4097): pharynx: transverse sections on six slides; undeveloped copulatory apparatus: sagittal sections on three slides. Paratype MZUSP PL 2069 (field number F 4098): pharynx: sagittal sections on eight slides; copulatory apparatus: sagittal sections on 13 slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	distribution	Distribution Area covered with Atlantic forest in Cardoso Island, municipality of Cananeia, state of São Paulo.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the municipality where the species was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median yellow orange band, bounded on either side by a black stripe, externally to which is a white stripe, and this in turn is externally bordered by a black stripe, which merges into a grey band with white mottling, which gradually fades towards its outer margin, while this margin is bordered by a thin black stripe. Penis bulb extending anteriorly 1 mm from penis papilla. Stroma of the penis papilla with necks of glands gathered in bundles resembling an orange in cross-section. Wall of posterior half of female atrium richly folded.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3584957FCC6FC2C9923FEC0.taxon	description	External aspect Adults up to 66 mm long and 5 mm wide. Body margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side flat. Dorsal body surface constituted by a median yellow-orange band (approximately one-quarter of body width) that grades into the black of cephalic region (1 / 13 th of body length), bounded on either side by a black stripe (one-sixth) which join posteriorly, externally to wh ich is a white stripe (one-seventh), and this in turn is bordered externally by a black stripe, which merges into a grey band with white mottling (one-sixth), which gradually fades towards its outer margin, while this margin is bordered by a thin black stripe (Fig. 17 A). Ventral surface is whitish with black margins. The dorsal colours of preserved specimens remained unchanged. Two types of eyes: a conical type, in a single row around anterior tip; and a cup-shaped type, spreading onto dorsum, reaching whitish stripes except at the anterior extremity of the body. Sensory pits simple invaginations, 20 μm deep, located ventromarginally in a single row, from the very anterior end up to at least first one-sixth of body length. Relative position mouth-to-body length, 55 %. Relative position gonopore-to-body length, 70 %. Internal morphology Creeping sole 84 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 5 – 6 % of body height. Muscle fibres of the longitudinal, innermost layer (25 µm thick) arranged into bundles with 25 – 33 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres around intestine are present, constituting a loose tube whose fibres mix with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 29 % of pharyngeal pocket length. Pharynx cylindrical, with dorsal insertion slightly posterior to ventral one (Fig. 17 B). Oesophagus- to-pharynx ratio, 12 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (50 µm) of circular muscle fibres with longitudinal ones in its innermost region. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (50 µm) of circular fibres and an innermost layer (15 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes slightly behind ovarian region (equivalent to 20 % of body length), posteriormost follicles slightly anterior to the root of the pharynx. Penis bulb elongated, extending from 1 mm anterior to penis papilla to the level of the gonopore region; well developed, consisting of tightly packed muscle fibres variously orientated. Diagonal fibres on the right side of the bulb run obliquely downwards to embrace ventrally the anterior portion of the female atrium. These fibres apparently mix with the subintestinal parenchymal musculature. Sperm ducts run dorsolaterally to the ovovitelline ducts. Laterally to ventral insertion of penis papilla, sperm ducts bend anteriorly and medially and then ventrolaterally to penetrate the bulb. The ducts subsequently recurve, then join to form the ejaculatory duct. This wide canal (Fig. 17 C) initially curves dorsoposteriorly, then enters penis papilla and traverses it sinuously to open at its distal tip. Sperm present in distal portion of sperm ducts, along their course both outside and inside penis bulb. Ejaculatory duct lined with columnar, ciliated epithelium, 25 – 40 µm high in anterior portion, 40 – 50 µm high inside the papilla, and surrounded by a 40 - µm-thick layer of circular muscle fibres, which constitutes a muscular cylinder with a diamter equivalent to 1.5 times the diameter of this duct. Extrapapillar and anterior portion of ejaculatory duct pierced by numerous gland cells producing fine erythrophil granules; from that portion until nearly the tip of papilla the duct is pierced by cells producing coarse erythrophil granules. Penis papilla long, horizontal and cylindrical, with dorsal insertion slightly anterior to the ventral part; tip blunt. The papilla occupies the male atrium and anterior two-fifths of the female atrium. Total length of papilla equal to six times its greatest diameter. It is covered with a squamous epithelium, which is pierced by two types of very abundant gland cells, producing fine erythrophil and cyanophil granules, respectively, with the former being more abundant. Necks of these two types of glands progressively gather in larger and less numerous bundles; in the distal portion of the papilla, erythrophil necks are gathered in nine to 14 bundles, which are separated from each other by cyanophil glands, thus giving rise to a picture resembling an orange in cross-section (Fig. 17 D, E). Subepithelial musculature consisting of dense 50 - µm-thick layer of circular muscle, slightly decussate, followed by a 20 - µm-thick layer of longitudinal muscle. Male atrium ample, not folded (Fig. 17 E), lined with a squamous epithelium pierced by glands secreting an amorphous erythrophil substance, and underlain by a 5 - µm-thick layer of circular muscles, followed by a thin layer of longitudinal fibres. Circular muscle fibres surround both innermost portions of gonopore canal and nearby atrium. Ovaries oval shaped, 500 µm in length in the longitudinal axis, and situated at a distance from anterior end equivalent to one-fifth of body length. Ovovitelline ducts emerge from dorsolateral wall of ovaries. Laterally to hind region of female atrium, the ducts curve medially and join to form common glandular ovovitelline duct. This duct is as long as one-sixth of female atrium length; it first bends posteroventrally, then curves anteriorly to open into female genital duct; the latter is a projection of the posterior region of the female atrium and lined with a 12 - µm-high columnar, ciliated epithelium. Female atrium long, twice as long as male atrium (Fig. 17 E); anterior half spacious and lined with cuboidal epithelium; hind half has richly folded wall projected into its own lumen consisting of 25 - µm-high, stratified-like epithelium. Two types of gland pierce female epithelium, producing erythrophil and cyanophil granules, respectively, the second being more abundant in posterior half. Atrium surrounded by 25 - to 40 - µm-thick subepithelial layer of circular muscle fibres, followed by a layer of longitudinal fibres, partly interwoven with the circular ones.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35F494AFF44FEA898B5FEC2.taxon	materials_examined	Material examined All specimens were collected in Parque Nacional de Saint-Hilaire / Lange, Matinhos / PR, Brazil. Holotype MZUSP PL 2062 (field number F 1611): F. Carbayo et al., coll., 10 January 2008. Cephalic region: transverse sections on 21 slides; ovarian region: horizontal sections on 13 slides; pharynx: transverse sections on 18 slides; copulatory apparatus: sagittal sections on 15 slides. Paratype MZUSP PL 2063 (field number F 1635) (immature): F. Carbayo et al., coll., 11 January 2008. Cephalic region: horizontal sections on nine slides; pharynx: sagittal sections on five slides; portion behind pharynx: sagittal sections on four slides. Paratype MZUSP PL 1009 (field number F 1614): F. Carbayo et al., coll., 1 January 2008 (25 ° 45 ′ 51.7 ″ S, 048 ° 37 ′ 21.6 ″ W). Preserved in absolute ethanol.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35F494AFF44FEA898B5FEC2.taxon	distribution	Distribution Areas covered with Atlantic forest in Parque Nacional de Saint-Hilaire / Lange, Matinhos, state of Paraná, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35F494AFF44FEA898B5FEC2.taxon	etymology	Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the river flowing nearby the sampling site.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35F494AFF44FEA898B5FEC2.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median orange stripe, bounded on either side by a thin black stripe, externally to which is a broader white stripe, and this in turn is externally bordered by a black line, externally to which is a grey band with dense white mottling, externally to which is a marginal black line. Longitudinal parenchymal muscle fibres disposed around intestine as a loose muscular tube. Diameter of muscular cylinder surrounding ejaculatory duct equivalent to 12 times the diameter of this duct.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD35F494AFF44FEA898B5FEC2.taxon	description	External aspect Live animals 43 mm long and 3.5 mm wide. Body elongated, margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side flat. Dorsal colour constituted by a median orange band, bounded on either side by a thin black stripe (one-third of body width) that grades into the black of the cephalic region (1 / 12 th of body length), externally to which is a broader white stripe (one-sixth), and this in turn is externally bordered by a black line, externally to which is an area (one-sixth) coloured with a grey band with dense white mottling, externally to which is a marginal black line (Fig. 18 A). Ventral body surface cream, with black margins. The dorsal colours of preserved specimens remained unchanged. Two types of eyes: a conical type, around anterior tip; and a cup-shaped type, which are located from a short distance behind anterior tip to posterior end, extending dorsally onto whitish stripes. Sensory pits simple invaginations, located ventromarginally in a single row from the very anterior end up to at least a distance equivalent to 13 % of body length. Relative position mouth-to-body length, 50 %. Relative position gonopore-to-body length, 64 %. Internal morphology Creeping sole 80 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 6 % of body height. Muscle fibres of the longitudinal, innermost layer (6 – 12 µm thick) arranged into bundles with seven to 20 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres around intestine present, constituting a loose tube of fibres mixed up with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 47 % of pharyngeal pocket length. Pharynx cylindrical (Fig. 18 B). Oesophagus-topharynx ratio, 17 %. Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (65 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature (Fig. 18 C) consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (45 µm) of circular fibres and an innermost layer (10 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes slightly anterior to ovarian region, posteriormost follicles slightly anterior to root of pharynx. Penis bulb elongated, extending from 1.2 mm anterior to penis papilla to behind level of gonopore; very strongly developed dorsally to male atrium. It consists of tightly packed muscle fibres variously orientated anteriorly. Diagonal fibres on the right side of the bulb run obliquely downwards to embrace ventrally the anterior portion of the female atrium. Those fibres seem to cross the nerve plate and to anchor on ventral epidermis. Sperm ducts run dorsolaterally to the ovovitelline ducts. Laterally to ventral insertion of penis papilla, sperm ducts bend anteriorly and medially and then ventrolaterally to penetrate the bulb. The ducts subsequently join near root of penis papilla to form the ejaculatory duct. This duct traverses the central region of penis papilla to open at its tip (Fig. 18 D, E). Sperm present in distal portion of sperm ducts, along their course both outside and inside penis bulb. Sperm ducts lined with a cuboidal, ciliated epithelium, pierced by glands producing fine weakly erythrophil granules; sperm ducts and anterior portion of ejaculatory duct surrounded by a circular muscle. Distal portion of ejaculatory duct surrounded by a layer of circular-to-decussate muscles. Thickness of muscles surrounding the ejaculatory duct increases from 50 µm proximally to a maximum of 150 µm in mid-papilla, and constituting a muscular cylinder with a diameter equivalent to 12 times the diameter of this duct. Penis papilla long, horizontal and cylindrical, with dorsal and ventral insertions at the same transverse plane; tip pointed (Fig. 18 D, E). It occupies the entire male atrium and most of the female atrium. Total length of papilla equal to seven times its greatest diameter. Papilla covered with a cuboidal epithelium, pierced by two types of gland cells, producing erythrophil and cyanophil granules, respectively. Space immediately beneath mid-dorsal epithelium of papilla packed by necks of latter type of gland cells (Fig. 19 A, B). Subepithelial musculature consisting of dense layer of circular muscle, tending to decussate distally, 185 µm thick dorsally, 145 µm ventrally, followed by a 15 - µm-thick layer of longitudinal muscle. Distally, abundant decussate fibres in stroma. Male atrium ample, not folded, lined with a squamous epithelium, and underlain by a 5 - µm-thick layer of circular muscles, followed by an inconspicuous layer of longitudinal fibres. Ovaries oval shaped, 425 µm in diameter in longitudinal axis, and situated at a distance from anterior end equivalent to 17 % of body length. Ovovitelline ducts emerge from dorsolateral wall of ovaries. Laterally to female atrium, the oviducts curve medially and join to form common glandular ovovitelline duct. Common glandular ovovitelline duct as long as a quarter of female atrium length, after a knee-shaped bend communicating with a 30 - µm-long female genital duct lined with cuboidal, ciliated epithelium, the latter opening into the female atrium. Female atrium long, not folded (Figs 18 D, 19 C, D) and as long as two times the male atrium. Posterior, curved section lined with 20 - µm-tall stratified-like, non-ciliated epithelium and surrounded by a 30 - µmthick mass of very thin, seemingly circular muscle fibres, mixed with apparently minute erythrophil granules (Fig. 19 D). Anterior section of female atrium lined with cuboidal epithelium and surrounded by a 10 - to 40 - µm-thick layer of circular and longitudinal muscle fibres. A 10 - to 110 - µm-thick layer of longitudinal muscles embraces entire female atrium.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	materials_examined	Material examined All specimens were collected in Parque Estadual de Intervales, Ribeirão Grande / SP, Brazil (24 ° 16 ′ 0 9.8 ″ S, 0 4 8 ° 2 4 ′ 1 9.7 ″ W). F. Carbayo et al., coll., 1 3 December 2008. Holotype MZUSP PL 2065 (field number F 3166): Ovarian region: sagittal sections on 17 slides; prepharyngeal region: transverse sections on seven slides; pharynx: sagittal sections on 11 slides; male genital atrium: sagittal sections on 21 slides; female genital atrium: sagittal sections on nine slides. Paratype MZUSP PL 2064 (field number F 3132) (immature): Cephalic region: horizontal sections on three slides; pharynx and copulatory apparatus: sagittal sections on 12 slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	distribution	Distribution Areas covered with Atlantic forest in Parque Estadual de Intervales, Ribeirão Grande, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	etymology	Etymology The specific epithet is the name of another locality within the conservation area from where the species was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median melon yellow band, bounded on either side by a black band, externally to which is a white band divided longitudinally in half by a black line. Penis bulb extending anteriorly 2 mm from penis papilla. Sperm ducts join inside penis papilla. Diameter of muscular cylinder around ejaculatory duct equivalent to ten times the diameter of this duct.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	description	External aspect Live animals ~ 50 mm long and 4 mm wide. Body margins nearly parallel; anterior end rounded, posterior pointed. Dorsum and ventral side slightly convex. Dorsal body surface constituted by a median melon yellow band (one-fifth of body width) that grades into black of cephalic region (one-tenth of body length), bounded on either side by a black band (one-quarter), externally to which is a white band (one-fifth) that is longitudinally divided into half by a black line fading towards rear end (Fig. 20 A). Ventral surface white, with blackish margins. The dorsal colours of preserved specimens became slightly paler. Two types of eyes: a conical type, in a single row around anterior tip; and a cup-shaped type, spreading onto dorsum and reaching the black bands from immediately behind anterior end to the rear end. Sensory pits simple invaginations, 45 μm deep, located ventromarginally in a single row, from the very anterior end up to at least first one-quarter of body length. Relative position mouth-to-body length, 52 %. Relative position gonopore-to-body length, 67 %. Internal morphology Creeping sole 81 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 5 % of body height. Muscle fibres of the longitudinal, innermost layer (25 – 30 µm thick) arranged into bundles with 20 – 40 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres around intestine constituting a loose tube whose fibres mix with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 37 – 39 % of pharyngeal pocket length. Pharynx cylindrical, dorsal insertion slightly posterior. Oesophagus-to-pharynx ratio, 12 % (Fig. 20 B). Outer pharyngeal musculature consisting of a subepithelial layer (7 µm) of longitudinal muscle, followed by a layer (55 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (110 µm) of circular fibres and an innermost layer (40 µm) of longitudinal fibres. Testes located dorsally between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 22 % of body length; posteriormost follicles slightly anterior to root of pharynx. Penis bulb elongated, extending from 2 mm anterior to penis papilla to the level of the gonopore region; very strongly developed, mainly dorsally to male atrium. Penis bulb consists of tightly packed muscle fibres; fibres variously orientated proximally, and diagonally orientated distally. Distal fibres on the right side run obliquely downwards to embrace ventrally the anterior portion of the female atrium; these fibres seem to join fibres of the subintestinal parenchymal muscle layer, thus forming bundles; their anchor points could not be discerned. Sperm ducts run dorsally to the ovovitelline ducts. Shortly before attaining the level of penial insertion, sperm ducts bend medially and ventrolaterally penetrate the bulb. The ducts subsequently run posteriorly, penetrate penis papilla and join to form the ejaculatory duct; this traverses the central region of penis papilla to open, presumably, at its tip, but this tip is missing. Sperm present in distal portion of sperm ducts, along their course both outside and inside penis bulb and penis papilla. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 10 - µm-thick layer of circular muscles. Ejaculatory duct lined with cuboidal, ciliated epithelium containing fine cyanophil granules, and surrounded by layer of circular muscles, tending to decussate towards tip of penis; layer with a maximal thickness of 140 µm on mid papilla, and constituting a muscular cylinder with a diameter equivalent to ten times the diameter of this duct (Fig. 20 C). Penis papilla horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 21 A, B). Papilla occupies entire male atrium and most of female atrium. Total length of papilla equal to six times its greatest diameter. Penis papilla covered with a low epithelium pierced by two types of gland cells, producing erythrophil and cyanophil granules, respectively. A moderate space, 10 µm thick, immediately beneath mid-dorsal epithelium of papilla packed by necks of latter gland type (Fig. 20 C). Subepithelial musculature consisting of dense layer of circular muscle, tending to decussate distally, 200 µm thick dorsally, 100 µm thick ventrally, followed by a 10 - µm-thick layer of longitudinal muscle. Stroma rich in diagonal fibres, especially distally. Male atrium ample, not folded, lined with a cuboidal epithelium, underlain by a 12 - µm-thick layer of circular muscles, followed by a thin layer of longitudinal fibres. Ovaries oval shaped, ~ 200 µm in anteroposterior diameter, situated at a distance from anterior end equivalent to one-quarter of body length. Laterally to female atrium, the oviducts curve medially and join to form the common glandular ovovitelline duct. Common glandular ovovitelline duct as long as one-third of female atrium length, and communicating with posterior portion of female atrium, the latter being 200 µm long and curved towards dorsal body surface. Female atrium without folds, as long as three times that of male atrium (Fig. 21 B). The very short posterior portion (200 µm in length) of the female atrium is lined with a 25 - µm-high stratified-like, non-ciliated epithelium, which is surrounded by a 40 - µm-thick layer of very thin muscle fibres. Otherwise, the female atrium is lined with cuboidal epithelium and pierced by gland cells producing cyanophil granules. Epithelium of female atrium and that of female genital duct are surrounded by 5 - to 20 - µm-thick layer of circular-todecussate muscle fibres, followed by a 5 - to 55 - µmthick layer of longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	description	FIGS 22 AND 23	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	materials_examined	Material examined MZUSP PL 1068 (field number F 3249): Parque Estadual da Serra do Tabuleiro, Paulo Lopes / SC, Brazil (27 ° 58 ′ 53.0 ″ S, 048 ° 44 ′ 53.0 ″ W). F. Carbayo, et al., coll., 14 January 2009. Cephalic region: transverse sections on 11 slides; ovarian region: horizontal sections on six slides; region behind ovaries: sagittal section on nine slides; pre-pharyngeal region: sagittal sections on ten slides; pharynx: sagittal sections on 11 slides; copulatory apparatus: sagittal sections on 11 slides. MZUSP PL 2075 (field number F 6528): RPPN Vale das Pedras, Alfredo Wagner / SC, Brazil (27 ° 37 ′ 0 1.0 ″ S, 049 ° 20 ′ 52.0 ″ W). A. L. Almeida et al., coll., 16 January 2015. Cephalic region: horizontal sections on four slides; ovarian region: horizontal sections on five slides; pre-pharyngeal region: transverse sections on four slides; pharynx: sagittal sections on four slides; copulatory apparatus: sagittal sections on nine slides. MZUSP PL 2076 (field number F 6544): RPPN Vale das Pedras, Alfredo Wagner / SC, Brazil (27 ° 37 ′ 0 1.1 ″ S, 049 ° 20 ′ 53.3 ″ W). A. L. Almeida et al., coll., 18 January 2015. Cephalic region: horizontal sections on five slides; region behind ovaries: horizontal sections on four slides; pre-pharyngeal region: transverse sections on seven slides; pharynx: sagittal sections on five slides; copulatory apparatus: sagittal sections on seven slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	discussion	Note We could not confirm identification of a specimen studied by Froehlich (1955 b); see Supporting Information (Table S 1).	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	distribution	Distribution Areas covered with Atlantic forest in municipalities of Blumenau, Paulo Lopes, Alfredo Wagner, Eastern part of Santa Catarina state, South Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	diagnosis	Diagnosis Anterior third of dorsal body pure orange; remaining dorsal side constituted by a median light grey band, bordered on either side by a black band; entire dorsum sprinkled with conspicuous whitish specks. Penis bulb extending anteriorly 1 mm from penis papilla. Papilla as long as seven to nine times its diameter and with a very thin tip. A conspicuous mass of very thin circular muscle fibres around posterior section of female atrium.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD342494FFF7FFEAD9836FB4A.taxon	description	External aspect Live animal 30 mm long and 3 mm wide when extended. Body elongated, margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side slightly convex. Anterior third of dorsal body surface pure orange; remaining two-thirds with a median light grey wide band (one-third), bordered on either side by a black band (one-third), with the exception of the very anterior end; entire dorsum sprinkled with conspicuous whitish specks (Fig. 22 A). Anterior third of ventral body surface traffic white; rest of ventral surface squirrel grey. The dorsal colours of preserved specimens became slightly paler. Two types of eyes: a conical type, 80 µm high and 35 µm wide; and a cup-shaped type, 35 µm in diameter. The former contour the anterior third of the body. Cup-shaped type eyes in lateral bands, approximately one-third of body width. Sensory pits simple invaginations, 30 μm deep, located ventromarginally in a single row, from the very anterior end up to the pre-pharyngeal region. Relative position mouth-to-body length, 53 %. Relative position gonopore-to-body length, 67 %. Internal morphology Creeping sole 85 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 5 % of body height. Muscle fibres of the longitudinal, innermost layer (20 µm thick) arranged into bundles with 17 – 21 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and a subintestinal layer with transverse and longitudinal fibres. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 51 % of pharyngeal pocket length. Pharynx cylindrical. Oesophagus-to-pharynx ratio, 22 % (Fig. 22 B). Outer pharyngeal musculature consisting of a subepithelial layer (5 µm) of longitudinal muscle, followed by a layer (50 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (65 µm) of circular fibres and an innermost layer (20 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 27 % of body length; posteriormost follicles a little anterior to the root of the pharynx. Penis bulb elongated, extending from 1 mm anterior to penis papilla to level of gonopore region (Fig. 22 C, D); strongly developed, consisting of tightly packed muscle fibres variously orientated proximally, whereas diagonally orientated distally. Penis bulb more developed dorsally to male atrium than ventrally. Oblique muscular ring embraces dorsodistal portion of male atrium and anteroventral portion of female atrium; some fibres detach from the ring and continue downwards to ventral epidermis behind gonopore region (Fig. 23 A). Sperm ducts run dorsolaterally to the ovovitelline ducts. Laterally to penis bulb, sperm ducts bend anteriorly and medially and then ventrolaterally penetrate the bulb. The ducts subsequently join to form an unpaired duct that continues as the ejaculatory duct near the root of the penis papilla, and traverses its central region to open at its tip. Sperm present in distal portion of sperm ducts, along their course both outside and inside penis bulb. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 7 - to 10 - µm-thick layer of circular muscles. Ejaculatory duct lined with tall, ciliated epithelium, pierced by conspicuous and numerous gland cells producing erythrophil granules. It is surrounded by a layer of circular muscles, tending to decussate towards penis tip; the layer has maximal thickness of 75 µm in the posterior half of the papilla, constituting a muscular cylinder with a diameter equivalent to four times the diameter of this duct. Penis papilla long, horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane; the very tip of this papilla is thin and long. This papilla occupies the entire male atrium and most of the female atrium. Total length of papilla equal to seven to nine times its greatest diameter. The papilla is covered with a cuboidal, non-ciliated epithelium, pierced by gland cells producing erythrophil granules; glands progressively more densely distributed towards the distal portion of the penis papilla. Subepithelial musculature consisting of dense 90 - µmthick layer of circular muscle, tending to decussate distally, followed by a single layer of longitudinal muscle. Stroma of papilla traversed by numerous diagonal muscle fibres, especially distally. Male atrium ample, not folded; lined with a low, non-ciliated epithelium, and underlain by a 5 - µm-thick layer of circular muscles, followed by an inconspicuous layer of longitudinal fibres. Ovaries oval shaped, 260 µm in diameter in anterior – posterior axis, and situated at a distance from anterior end equivalent to one-quarter of body length. Ovovitelline ducts emerge from dorsolateral wall of ovaries. Laterally to female atrium, the oviducts curve medially and join to form common glandular ovovitelline duct. Common glandular ovovitelline duct as long as half of female atrium length, and communicating with female genital duct; this duct is a projection of the dorsoposterior region of thefemale atrium and is lined with a 25 - µm-tall stratified-like, ciliated epithelium, with an erythrophil-free surface. A cyanophil fibrillar mass 10 – 20 µm thick is present immediately underneath this epithelium, with very thin circular filaments (<1 µm); it might be of glandular nature, albeit their circular arrangement resembles that of muscle fibres (Fig. 23 B). Female atrium without folds (Fig. 22 C, D), twice as long as the male atrium, and lined with a cuboidal epithelium. This epithelium and that of female genital duct surrounded by 5 - to 20 - µm-thick layer of circularto-decussate muscle fibres, followed by a 5 - to 55 - µmthick layer of longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	description	FIG. 24	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	materials_examined	Material examined Original type material (from EMF collection): MZUSP PL 2092 (field number Specimen A): Curitiba / PR, Brazil. Froehlich, 21 November 1952. Copulatory apparatus: sagittal sections on two slides. MZUSP PL 2093 (field number Specimen B): Curitiba / PR, Brazil. E. M. Froehlich coll., 21 November 1952. Copulatory apparatus: sagittal sections on four slides. Specimen A or B: pre-pharyngeal region: transverse sections on two slides; pharynx: sagittal sections on one slide. Other specimens: MZUSP PL 2084 (field number F 6993): Morretes / PR, Brazil. E. M. Froehlich et al., coll., 17 June 1953. Pharynx and copulatory apparatus: sagittal sections on nine slides. Additional material: MZUSP PL 2086 (field number F 6995): Morretes / PR, Brazil. Magno C. Segalla, coll., 1 November 1987. Pharynx and copulatory apparatus: sagittal sections on 37 slides. MZUSP PL 715 (field number F 1710) (imature): Morretes / PR, Brazil (25 ° 28 ′ 10.6 ″ S, 048 ° 48 ′ 49.5 ″ W). F. Carbayo et al., coll., 13 January 2008. Pre-pharyngeal region: transverse sections on three slides; pharynx: sagittal sections on five slides; region behind pharynx: sagittal sections on four slides. MZUSP PL 716 (field number F 1730) (incompletely mature): Morretes / PR, Brazil (2 5 ° 2 8 ′ 1 0. 6 ″ S, 048 ° 48 ′ 49.5 ″ W). F. Carbayo et al., coll., 13 January 2008. Cephalic region: horizontal sections on four slides; pharynx and copulatory apparatus: sagittal sections on six slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	discussion	Note An unlabelled specimen from the original type material is not conspecific (see Discussion below and Supporting information, Table S 1).	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	distribution	Distribution Areas covered with Atlantic forest in the municipalities of Curitiba and Morretes, State of Paraná, South Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median mustard-coloured band, bounded on either side by a black band; entire dorsum sprinkled with conspicuous whitish specks. Penis bulb extending anteriorly 1.5 mm from penis papilla. Diameter of muscle muscular cylinder around ejaculatory duct equivalent to ten times the diameter of this duct. Female-to-male atrial length ratio, 2.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD347494DFEE1FB3999F3FB36.taxon	description	External aspect Preserved adults 58 mm long and 4 mm wide. Body margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side flat. Dorsal body surface of live specimens constituted by a median mustard-coloured band (one-third of body width) that grades into the black of cephalic region (one-sixth of body length), bounded on either side by a black band (one-third); entire dorsum sprinkled with conspicuous whitish specks (Fig. 24 A, B). Ventral surface whitish with blackish margins. The dorsal colours of preserved specimens collected in 2008 became slightly paler; the dorsal side of one specimen collected in 1952 consisted of a beige red median band bordered by greyish lateral bands. Two types of eyes: a conical type, in a single row around anterior tip; and a cupshaped type, spreading onto dorsum along lateral bands being one-third of body width. Relative position mouth-to-body length, 65 %. Relative position gonopore-to-body length, 76 %. Internal morphology Creeping sole 86 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, being as thick as 6 % of body height. Muscle fibres of the longitudinal, innermost layer (10 – 20 µm thick) arranged into bundles with seven to 30 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse fibres, and subintestinal layer with transverse fibres. Longitudinal fibres around intestine present, constituting a loose tube whose fibres mix with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 30 – 45 % of pharyngeal pocket length. Pharynx cylindrical; dorsal insertion posteriorly placed. Oesophagus-to-pharynx ratio, 14 % (Fig. 24 C). Outer pharyngeal musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (35 µm) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (38 µm) of circular fibres and an innermost layer (10 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes very slightly anterior to ovarian region, posteriormost testes near root of pharynx. Penis bulb elongated, extending from 1.5 mm anterior to penis papilla to level of gonopore region; very strongly developed, consisting of tightly packed muscle fibres variously orientated proximally, whereas diagonally orientated distally. Distal fibres on the right side run obliquely downwards to embrace ventrally the anterior portion of the female atrium and subsequently anchor on ventral epidermis. Penis bulb more developed dorsally to male atrium than ventrally. Sperm ducts run dorsally, or slightly lateral to the ovovitelline ducts. Sperm ducts lined with a cuboidal, ciliated epithelium and surrounded by a 10 - to 15 - µm-thick layer of circular muscles. Ejaculatory duct lined with cuboidal, ciliated epithelium, pierced by gland cells apparently producing erythrophil granules; surrounded by a layer of circular muscles with maximal thickness of 140 µm on the posterior half, and constituting a muscular cylin- der with a diameter equivalent to ten times the diameter of this duct (Fig. 24 D – F). Penis papilla long, horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 24 D, F); posterior half of this papilla dilated, tapering to form a very narrow tip. Papilla occupies entire male and female atrium. Total length of papilla equal to seven times its greatest diameter. Papilla covered with a cuboidal, non-ciliated epithelium, pierced by gland cells producing erythrophil granules. A moderate space of 10 µm immediately beneath epithelium of mid-papilla fully packed by necks of gland cells producing cyanophil granules. Subepithelial musculature consisting of a dense layer of circular muscle, tending to decussate distally, 80 µm thick dorsally and 160 µm thick ventrally, followed by a 10 - µm-thick layer of longitudinal muscle. Male atrium ample, not folded; lined with a low, non-ciliated epithelium, and underlain by a 10 - µm-thick layer of circular muscles followed by a 10 - µm-thick layer of longitudinal fibres. Ovaries oval shaped, 500 µm in diameter in anteroposterior axis, and situated at a distance from anterior end equivalent to one-quarter of body length. Ovovitelline ducts emerge from dorsolateral wall of ovaries. Laterally to female atrium, the oviducts curve medially and join to form the common glandular ovovitelline duct. Common glandular ovovitelline duct as long as approximately one-third of the female atrium length, and communicating with posterior portion of female atrium, the latter being 300 µm long, and bent posterodorsally. Female atrium without folds, and being twice the length of the male atrium (Fig. 24 F). Posterior section with 200 µm in length of female atrium lined with a 20 - µm-high, stratified-like, non-ciliated epithelium, which is surrounded by a 30 - µm-thick layer of very thin muscle fibres. Remaining epithelium of female atrium lined with cuboidal epithelium, pierced by gland cells producing erythrophil granules. This epithelium and that of female genital duct surrounded by 5 - to 40 - µm-thick layer of circular-to-decussate muscle fibres, followed by a 40 - to 150 - µm-thick layer of longitudinal fibres.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	description	FIGS 25 AND 26	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	materials_examined	Material examined Holotype MZUSP PL 2070 (field number F 4112): Estação Biológica de Boraceia, Salesópolis / SP, Brazil (23 ° 39 ′ 14.0 ″ S, 045 ° 53 ′ 21.0 ″ W). F. Carbayo et al., coll., 21 December 2009. Cephalic region: horizontal sections on 11 slides; ovarian region: horizontal sections on four slides; pre-pharyngeal region: transverse sections on eight slides; pharynx: sagittal sections on six slides; copulatory apparatus: sagittal sections on 12 slides. Paratype MZUSP PL 2077 (field number F 6575): São Sebastião / SP, Brazil (23 ° 44 ′ 48.7 ″ S, 045 ° 37 ′ 55.7 ″ W). A. L. Almeida et al., coll., 13 February 2015. Ovarian region: horizontal sections on seven slides; pre-pharyngeal region: transverse sections on ten slides; pharynx and copulatory apparatus: sagittal sections on 15 slides. Paratype MZUSP PL 2078 (field number F 6578): São Sebastião / SP, Brazil (23 ° 44 ′ 48.7 ″ S, 045 ° 37 ′ 55.7 ″ W). A. L. Almeida et al., coll., 13 February 2015. Ovarian region: horizontal sections on seven slides; prepharyngeal region: transverse sections on ten slides; pharynx and copulatory apparatus: sagittal sections on nine slides. Paratype MZUSP PL 2073 (field number F 6357): São Sebastião / SP, Brazil (23 ° 44 ′ 48.7 ″ S, 045 ° 37 ′ 55.7 ″ W). A. L. Almeida et al., coll., 29 April 2015. Copulatory apparatus: sagittal sections on 11 slides. Paratype MZUSP PL 2123 (field number F 4389): São Sebastião / SP, Brazil (23 ° 44 ′ 45.6 ″ S, 045 ° 37 ′ 55.2 ″ W). F. Carbayo et al., coll., 26 February 2010. Entire specimen in 80 % ethanol. Paratype MZUSP PL 2095 (field number F 6026): São Sebastião / SP, Brazil (23 ° 44 ′ 39.8 ″ S, 045 ° 38 ′ 13.7 ″ W). Carbayo et al., coll., 11 January 2014. Preserved in 80 % ethanol. Paratype MZUSP PL 2085 (field number F 6994): Estação Biológica de Boraceia, Salesópolis / SP, Brazil. O. Froehlich et al., coll., 14 September 1986. Pharynx and copulatory apparatus: sagittal sections on 48 slides. Paratype MZUSP PL 2124 (field number F 6068): São Sebastião / SP, Brazil (23 ° 44 ′ 48.7 ″ S, 045 ° 37 ′ 55.7 ″ W). Carbayo et al., coll., 12 January 2014. Preserved in 100 % ethanol. Paratype MZUSP PL 2125 (field number F 6055): São Sebastião / SP, Brazil (23 ° 44 ′ 24 ″ S 45 ° 37 ′ 48 ″ W). Carbayo et al., coll., 11 January 2014. Preserved in 80 % ethanol. Paratype MZUSP PL 1086 (field number F 4358): São Sebastião / SP, Brazil (23 ° 45 ′ 0 7.3 ″ S, 045 ° 37 ′ 51.7 ″ W). Carbayo et al., coll., 26 February 2010. Pre-pharyngeal region: transverse sections on five slides; pharynx: sagittal sections on 12 slides; copulatory apparatus: sagittal sections on 12 slides.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	distribution	Distribution Areas covered with Atlantic forest in the municipalities of Salesópolis and São Sebastião, state of São Paulo, Brazil.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	diagnosis	Diagnosis Dorsal colour pattern constituted by a median mustard-coloured band, bounded on either side by a black band; entire dorsum sprinkled with conspicuous whitish specks. Penis bulb extending anteriorly 0.8 mm from penis papilla. Diameter of muscular cylinder around ejaculatory duct equivalent to 1.5 times the diameter of this duct. Female-to-male atrial length ratio, ~ 1.0.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	etymology	Etymology The specific epithet refers to the name of the biological station where the type material was collected.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3454941FCD3FAD79875FC59.taxon	description	External aspect Preserved adults 50 mm long and 4 mm wide. Body elongated, margins nearly parallel; anterior end rounded, posterior pointed. Dorsum strongly convex, ventral side flat. Dorsal body surface constituted by a median mustard-coloured band (one-third of body width) that grades into the black of cephalic region (one-sixth of body length), bounded on either side by a black band (one-third); entire dorsum sprinkled with conspicuous whitish specks (Fig. 25 A). Ventral surface is grey whitish, with blackish margins. In fixed specimens, dorsal colours faded, mostly the median band. Two types of eyes: a conical type, in a single row around anterior tip; and a cup-shaped type, spreading onto dorsum along lateral bands with one-third of body width. Sensory pits simple invaginations, 40 µm deep, located ventromarginally in a single row, from the very anterior end up to at least the first one-eighth of body length. Relative position mouth-to-body length, 57 %. Relative position gonopore-to-body length, 70 %. Internal morphology Creeping sole 85 % of body width. Glandular margin absent. Three typical geoplaninid cutaneous muscle layers present, as thick as 9 % of body height. Muscle fibres of the longitudinal, innermost layer (20 – 25 µm thick) arranged into bundles with nine to 30 fibres each. Three parenchymal muscle layers present: dorsal layer of decussate diagonal fibres, supraintestinal layer of transverse and longitudinal fibres, and subintestinal layer with transverse and longitudinal fibres. Longitudinal fibres around intestine present, constituting a loose tube whose fibres mix with transverse ones. Ventral nerve plate present. Mouth situated at a distance from root of the pharynx equivalent to 34 – 49 % of pharyngeal pocket length. Pharynx cylindrical, dorsal insertion slightly posterior. Oesophagus-to-pharynx ratio, 10 % (Fig. 25 B). Outer pharyngeal musculature consisting of a subepithelial layer (2.5 µm) of longitudinal muscle, followed by a layer (35 µm) of circular fibres, the innermost of which are intermingled with longitudinal ones. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (50 µm) of circular fibres and an innermost layer (15 µm) of longitudinal fibres. Testes dorsally located between supraintestinal parenchymal muscle layer and intestine; anteriormost testes at a distance from anterior end equivalent to 22 % of body length; posteriormost testes slightly anterior to root of pharynx. Penis bulb elongated, extending from 0.8 mm anterior to penis papilla to the level of gonopore region; well developed, dorsally more than ventrally. It consists of tightly packed muscle fibres variously orientated proximally. Part of the muscle fibres from the right dorsal portion of penis bulb detach and, subsequently, run obliquely downwards to embrace ventro-anterior portion of the female atrium (Fig. 25 D, E). Further course of these fibres could not be determined. A lower number of muscle fibres from the dorsoposterior left portion of penis bulb run anteroventrally towards ventral epidermis; further course of these fibres could not be determined. Sperm ducts run dorsolaterally or dorso-internally to the ovovitelline ducts. Laterally to level of the insertion of penis papilla, the sperm ducts bend medially and then ventrolaterally to penetrate the bulb. The ducts subsequently recurve and join to continue posteriorly as an unpaired, curved duct; the unpaired portion continues as the ejaculatory duct. The latter follows a sinuous course to open at the tip of the penis papilla (Figs 25 C, D, 26 A, B). Sperm present in distal portion of sperm ducts, along both their course outside and inside penis bulb. Sperm ducts lined with a cuboidal-to-columnar, ciliated epithelium and surrounded by a 12 - µm-thick layer of circular muscles. Ejaculatory duct lined with tall, ciliated epithelium; extrapapillar section pierced by gland cells producing fine, weakly cyanophil granules, but this cell type may extend along intrapapillar section until the level of the gonopore; intrapapillar section pierced by conspicuous and numerous gland cells producing erythrophil-to-xanthophil granules; surrounded by a 10 - µm-thick layer of circular muscles, and constituting a muscular cylinder with a diameter equivalent of 1.5 times the diameter of this duct. Penis papilla long, horizontal and cylindrical along most of its length; distally conical. Dorsal and ventral insertions of this papilla approximately at the same transverse plane. This papilla occupies the entire male atrium and the first half of the female atrium. Total length of papilla equal to six to seven times its greatest diameter. Penis papilla covered with a squamousto-cuboidal epithelium, pierced by two types of gland cells, producing strongly erythrophil granules and fine cyanophil granules, respectively; former glands pierce entire epithelium except for mid-dorsal; this area pierced by cyanophil glands, the necks of which pack the space immediately beneath this epithelium (Fig. 25 C). Subepithelial musculature consisting of a dense layer of circular muscle, decussates distally, 110 µm thick dorsally, 55 µm ventrally, followed by a 15 - µm-thick layer of longitudinal muscle. Male atrium ample, not folded, and lined with a low, non-ciliated epithelium, pierced by gland cells producing erythrophil granules and underlain by a 10 - µm-thick layer of circular muscles, followed by a 10 - µm-thick layer of longitudinal fibres. Ovaries oval shaped, 500 µm in diameter in anterior – posterior axis, and situated at a distance from anterior end equivalent to one-fifth of body length. Ovovitelline ducts emerge from dorsolateral wall of ovaries. Laterally to female atrium, the oviducts curve medially and join to form the common glandular ovovitelline duct. This duct is as long as 20 – 30 % of female atrium length and communicates with female genital duct; the latter is a canalicular projection of the posterior region of the female atrium, and is lined with a 25 - µm-tall stratified-like, ciliated epithelium. Female atrium cylindrical, becoming conical posteriorly, sometimes here with small folds (Figs 25 C, 26 A, D); a little longer than the male atrium. Conical portion lined with a stratified-like epithelium; cylindrical portion lined with cuboidal epithelium; both epithelia pierced by two types of gland cells, producing amorphous substance; surrounded by 5 - to 20 - µm-thick layer of circular-to-decussate muscle fibres, followed by a layer of longitudinal fibres, 125 µm thick in posterior portion, 20 µm in anterior portion.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
03BFF20FD3494946FCBFF9319F5EFA51.taxon	discussion	A total of five species are distinguished from the remaining congeners in the chromatic pattern of the dorsum, namely G. vaginuloides (median black band, white-yellow para-median stripes externally bordered by black lines, and lateral reddish-iron bands), G. apua (median luminous orange band, bounded on either side by a black stripe, externally to which is a greenishyellow stripe, and this again is bordered externally by a marginal black band), G. mogi (median orange-ochre band, bounded on either side by a thin whitish stripe, and in turn this is bordered externally by a thin black line, externally to which is a grey band with dense white mottling, externally to which is a thin marginal black line), G. iporanga (median melon yellow band, bounded on either side by a black band, externally to which is a white band divided longitudinally in half by a black line), and G. pulchella (anterior third of dorsal body pure orange; remaining dorsal side constituted by a median light grey band, bordered on either side by a black band; entire dorsum sprinkled with conspicuous whitish specks). There are six pairs of species that are indistinguishable by means of only either the external aspect (G. piratininga – G. ibiuna, G. chita – G. boraceia, G. caraguatatuba – G. cananeia and G. cambara – G. paranapiacaba) or the internal organs (G. cambara – G. chita and G. mogi – G. boraceia). Their profound similarity means that we rely on a combination of both external and internal atributes to recognize every species. The dorsum of G. piratininga and G. ibiuna is very alike (median traffic red or orange-ochre band, bounded on either side by a black stripe, externally to which is a white stripe, and in turn this is externally bordered by a black band, externally to which is a thin marginal white line). However, their copulatory apparatus differs in that in G. piratininga the penis papilla is as long as five times its diameter, and the femaleto-male atrial length ratio is 2.0, whereas in G. ibiuna the penis papilla is as long as seven times its diameter, and the female-to-male atrial length ratio is 1.0. The dorsum of G. chita and G. boraceia is also indistinguishable from each other (median mustard-coloured band, bounded on either side by a black band; entire dorsum sprinkled with conspicuous whitish specks). The internal most conspicuous difference between them is that the diameter of the muscular cylinder around the ejaculatory duct is equivalent to ten times the diameter of this duct in G. chita, whereas in G. boraceia it is equivalent to 1.5 times. Geoplana caraguatatuba and G. cananeia present a dorsal colour pattern also very alike (median orangeochre or yellow-orange band, bounded on either side by a black stripe, externally to which is a white stripe, and this in turn is externally bordered by a black stripe that merges into a grey band with white mottling, which fades gradually towards its outer margin, whereas this margin is bordered by a black stripe). However, the penis papilla of G. caraguatatuba is pointed, the female atrium presents smooth walls, and female-to-male atrial length ratio is 2.0, whereas in G. cananeia the penis papilla is blunt, the posterior half of the female atrium has a richly folded wall and the female-to-male atrial length ratio is 1.0. Additionally, in the latter species the stroma of the penis papilla presents necks of glands gathered in gross bundles. The pair of species G. cambara – G. paranapiacaba are externally very alike (median stripe of orange or orange-ochre, bounded on either side by a black line, externally to which is a broader white line, and in turn this is externally bordered by a thin black line, externally to which is a grey band with dense white mottling, bordered by a thin marginal black line). Internally, they can be distinguished in that the longitudinal parenchymal muscle fibres are disposed around intestine as a loose muscular cylinder in G. cambara, whereas in G. paranapiacaba the muscle fibres are positioned only dorsally and ventrally to the intestine. Furthermore, in G. cambara the diameter of the muscular cylinder around the ejaculatory duct is equivalent to 12 times the diameter of this duct, whereas in G. paranapiacaba it is equivalent to 2.5 times. Finally, the species of each of two pairs of species (G. cambara – G. chita and G. mogi – G. boraceia) cannot be differentiate from each other with regard to the internal organs, especially the copulatory apparatus. However, the chromatic pattern of G. cambara is constituted by a median orange stripe plus five pairs of lines (lateralmost lines are difficult to discern) either black, white or grey, whereas the dorsum of G. chita displays a median mustard-coloured band, bounded on either side by a black band, with entire dorsum sprinkled with conspicuous whitish specks. The internal organs of G. mogi and G. boraceia are also very similar to each other, but the dorsum in G. mogi is constituted by a median orange-ochre band plus four pairs of whitish, black or grey lines, whereas in G. boraceia the median mustard-coloured band is bounded on either side by a black band; with the entire dorsum sprinkled with conspicuous whitish specks.	en	Ana Laura Almeida, Fernando P. L. Marques, Fernando Carbayo (2019): ‘ Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65, DOI: 10.1093/zoolinnean/zly022
