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Two subfamilies of land planarians (Geoplanidae) are endemic to the Neotropical region, namely Geoplaninae (with 29 genera and 346 nominal species, most of which are from Brazil) and its sister-group Timyminae, with only two Chilean species. The systematics of these groups through morphology and molecular data (COI and 28S rDNA genes), including nine new Chilean species, is re-assessed in this study. The great morphological diversity of the Chilean species is congruent with the molecular trees and, accordingly, five new genera (Adinoplana, Harana, Myoplana, Sarcoplana and Transandiplana) are proposed, each characterized by putative synapomorphies. Seven new tribes are also erected (Adinoplanini, Gusanini, Haranini, Inakayaliini, Myoplanini, Polycladini and Sarcoplanini), each one monogeneric, except Geoplanini (which includes all genera under the current concept of Geoplaninae plus the Chilean Transandiplana) and Sarcoplanini (with Sarcoplana and the already known Mapuplana, Pichidamas and Wallamapuplana). Re-diagnoses of Geoplaninae, Timymini, Gusana, Inakayalia, Polycladus and Pichidamas are proposed and biogeographic remarks on Transandiplana are provided

The Chilean land planarian genus Gusana (Platyhelminthes, Tricladida, Geoplaninae) currently comprises three species that were described in the 19th century. Four new species of the genus are described herein, namely G. hualpensis Carbayo, sp. nov., G. lujanae Almeida & Carbayo, sp. nov., G. melipeucensis Almeida & Carbayo, sp. nov. and G. purensis Bolonhezi, Almeida & Carbayo, sp. nov. An integrative taxonomy approach was adopted by combining morphological (anatomy, histology) and molecular (COI and 28S genes) information. Additionally, the monophyletic status of Gusana is investigated and the species are delimited by a recently introduced molecular delimitation method based on pairwise genetic distances. All Gusana species are very similar in attributes such as body shape, dorsal colour pattern and internal organs, namely the pharynx and copulatory apparatus and only differ in the details. Gusana is retrieved as monophyletic and the molecular delimitation method recovered the same species recognised morphologically. A re-diagnosis of the genus and replacement of G. lata into Pseudogeoplana are also proposed. A new role of the penis papilla is also suggested based on the spermatophore found in Gusana lujanae.

Why some animals can regenerate while many others cannot remains a fascinating question. Even amongst planarian flatworms, well-known for their ability to regenerate complete animals from small body fragments, species exist that have restricted regeneration abilities or are entirely regeneration incompetent. Towards the goal of probing the evolutionary dynamics of regeneration, we have assembled a diverse live collection of planarian species from around the world. The combined quantification of species-specific head regeneration abilities and comprehensive transcriptome-based phylogeny reconstructions reveals multiple independent transitions between robust whole-body regeneration and restricted regeneration in the freshwater species. Our demonstration that theRNAi-mediated inhibition of canonical Wnt signalling can nevertheless bypass all experimentally tractable head regeneration defects in the current collection indicates that the pathway may represent a hot spot in the evolution of planarian regeneration defects. Combined with our finding that Wnt signalling has multiple roles in the reproductive system of the model speciesS. mediterranea, this raises the possibility of a trade-off between egg-laying and asexual reproduction by fission/regeneration as a driver of regenerative trait evolution. Although initial quantitative comparisons of Wnt signalling levels, reproductive investment, and regenerative abilities across the collection confirm some of the model’s predictions, they also highlight the diversification of molecular mechanisms amongst the divergent planarian lineages. Overall, our study establishes a framework for the mechanistic evolution of regenerative abilities and planarians as model taxon for comparative regeneration research.

Traumatic mating, that is, copulation that involves wounding the partner's body, is a widespread phenomenon that is particularly prevalent in hermaphroditic animals. Traumatic mating is generally a collateral side effect of diverse strategies (from physical anchorage to injection of substances to manipulate the partner), but the trauma could also be adaptive by itself if it delays remating by the injured partner. In the Tricladida (the clade of planarian flatworms), reciprocal sperm transfer is often assumed to occur by means of a 'regular' nontraumatic copulation, that is, insertion of the penis through the partner's gonopore and deposition of the ejaculate into its genital atrium, with subsequent sperm migration to the oviducts. However, while studying the anatomy of Brazilian land planarians for taxonomic purposes, we found foreign bodies, reminiscent of spermatophores, implanted within the parenchyma of Choeradoplana albonigra (Riester, 1938). Herein, we describe and illustrate several lines of morphological evidence indicating that these foreign bodies likely represent a novel case of intragenital copulatory wounding (e.g., structural and histochemical similarity to land planarians spermatophores; implantation at the level of the gonopore; vestiges of rupture of the genital atrium's wall), corroborating that traumatic mating is pervasive and underreported in Metazoa. We also propose two different hypotheses to explain such copulatory wounding, viz., that it concerns (1) a regular mating strategy or (2) an accidental effect of the copulation. In any event, this land planarian may prove useful as a novel, noninsect terrestrial model organism to investigate the evolution of traumatic mating.

Santos, Charles Morphy D., Carbayo, Fernando (2021): Taxonomy as a political statement: the Brazilian case. Zootaxa 5047 (1): 92-94, DOI: https://doi.org/10.11646/zootaxa.5047.1.8

Living representatives of the Neotropical genus Choeradoplana Graff, 1896 ( Geoplaninae , Tricladida , Platyhelminthes ) are easily recognized by the typical shape of the head which is laterally expanded, rolled-up, and ventrally provided with two glandular cushions. In this study, the morphology and phylogeny (cytochrome C oxidase subunit I gene) of several species of land planarians are taxonomically investigated. Four of the six species studied are new to science, namely: Ch. eudoxiae Silva & Carbayo, sp. nov., Ch. claudioi Lago-Barcia & Carbayo, sp. nov., Ch. onae Lago-Barcia & Carbayo, sp. nov., and Ch. riutortae Lago-Barcia & Carbayo, sp. nov. The species Choeradoplana albonigra and Ch. eudoxiae deviate from the usual body shape pattern in that the head does not present lateral expansions nor glandular cushions, becoming indistinguishable from its sister genus Cephaloflexa . Pseudogeoplana tristriata (Schultze & Muller, 1857) is also redescribed from a newly collected specimen and was discovered to be a member of Choeradoplana . Graff (1899) also studied another specimen that was considered to be conspecific with P. tristriata ; however, in this new it is concluded that it is not conspecific but rather a new species. The name Pseudogeoplana aevipandemiae Lago-Barcia & Carbayo, sp. nov. is suggested for Graff’s specimen.

Living representatives of the Neotropical genus Choeradoplana Graff, 1896 (Geoplaninae, Tricladida, Platyhelminthes) are easily recognized by the typical shape of the head which is laterally expanded, rolled-up, and ventrally provided with two glandular cushions. In this study, the morphology and phylogeny (cytochrome C oxidase subunit I gene) of several species of land planarians are taxonomically investigated. Four of the six species studied are new to science, namely: Ch. eudoxiae Silva & Carbayo, sp. nov., Ch. claudioi Lago-Barcia & Carbayo, sp. nov., Ch. onae Lago-Barcia & Carbayo, sp. nov., and Ch. riutortae Lago-Barcia & Carbayo, sp. nov. The species Choeradoplana albonigra and Ch. eudoxiae deviate from the usual body shape pattern in that the head does not present lateral expansions nor glandular cushions, becoming indistinguishable from its sister genus Cephaloflexa. Pseudogeoplana tristriata (Schultze & Müller, 1857) is also redescribed from a newly collected specimen and was discovered to be a member of Choeradoplana. Graff (1899) also studied another specimen that was considered to be conspecific with P. tristriata; however, in this new it is concluded that it is not conspecific but rather a new species. The name Pseudogeoplana aevipandemiae Lago-Barcia & Carbayo, sp. nov. is suggested for Graff’s specimen.

Background Cratera is a genus of land planarians endemic to the Brazilian Atlantic forest. The species of this genus are distinguished from each other by a series of external and internal characters, nonetheless they represent a challenging taxonomic issue due to the extreme alikeness of the species analysed in the present work. To resolve these difficulties, we have performed morphological analyses and used three nuclear markers (ribosomal 18S and 28S, Elongation Factor, a new anonymous marker named Tnuc813) and two mitochondrial fragments (Cytochrome c oxidase subunit I gene, and a fragment encompasing NADH deshydrogenase subunit 4 gene, trnF and the beginning of the Cytochrome c oxidase I gene) in an integrative taxonomic study. Methods To unveil cryptic species, we applied a molecular species delimitation approach based on molecular discovery methods, followed by a validation method. The putative species so delimited were then validated on the basis of diagnostic morphological features. Results We discovered and described four new species, namely Cratera assu, C. tui, C. boja, and C. imbiri. A fifth new species, C. paraitinga was not highly supported by molecular evidence, but was described because its morphological attributes are unique. Our study documents for the genus Cratera the presence of a number of highly similar species, a situation that is present also in other genera of land planarians. The high number of poorly differentiated and presumably recent speciation events might be explained by the recent geological history of the area.

Zoologist Prof. Eudóxia Maria Froehlich (1929*–2015†) kept an uncatalogued personal collection of land planarians in wet medium. We catalogued their contents. Glass jars contained a variable number of small vials, each with one or several labeled specimens. We transcribed the species names and field data, when available, written on card labels to a table. A total of 1772 vials in 80 jars, contained over 2700 specimens. Subsequently, we cross checked field data with that of the type specimens. As a result, we identified, with different degrees of certainty, type specimens of 78 species. These types compose a very significant potential source of additional morphological details of the species to conform with current standards in the taxonomy of the group.

Three new Neotropical land planarians (Platyhelminthes, Tricladida, Geoplaninae) from the Brazilian Atlantic forest are described. Obama apiguara Oliveira, Almeida & Carbayo sp. nov. presents glandular fossae opening through dorsal epithelium, a structure not documented previously among land flatworms. Paraba tata Bolonhezi, Lago-Barcia & Carbayo sp. nov., is characterized by a prostatic vesicle horizontal with an inconspicuous bifurcate portion, a penis papilla as long as male atrium and a female genital duct projected from the postero-dorsal section of the female atrium. The third new species is distinctive in the extraordinarily thick muscle surrounding the prostatic vesicle, a finger-shaped penis papilla, and a dilated female genital cavity. These attributes stands the species apart from all members of Geoplaninae and Piima ata Carbayo gen. et sp. nov. is proposed.

Areas of endemism (AoE) are the main study units in analytical biogeographic methods, and are often defined as an area with two or more endemic species living in them, presenting substantial congruence among their range limits. We explored the distribution of land planarians (Geoplanidae, Platyhelminthes) across the southern region of the Brazilian Atlantic forest (from the state of Rio de Janeiro, to the state of Rio Grande do Sul) utilizing DaSilva's et al. (2015) protocol. We used two methods, Endemicity Analysis (EA), and Geographical Interpolation of Endemism (GIE). We identified nine AoE of terrestrial flatworms in the Southern Atlantic forest. Performance of the methodologies is discussed. These AoE of land planarians can be explained through vicariance events combined with their physiological and ecological own limitations. Interestingly, these AoE are congruent with fine-scale approaches such as that with harvestmen. Most land planarians have revealed to present a very small distributional range evidencing their potential as a good model for fine-scale studies of AoE.

In 2016, the type-material of ten of the 15 Brazilian land planarians (Platyhelminthes, Tricladida, Geoplanidae) described by Schirch (1929) was discovered deposited in the Museu Nacional do Rio de Janeiro (MNRJ). Schirch only described the external morphology of these species, all originally placed in the genus Geoplana. By the 1930s and 1950s Geoplana itatiayana, G. plana, and G. rezendei underwent taxonomic revision based on the study of non-type specimens. The remaining 12 species also underwent a taxonomic revision but only based on the literature. Current names of these species are Geoplana goettei, Pseudogeoplana arpi, Ps. blaseri, Ps. bonita, Ps. bresslaui, Ps. cardosi, Ps. doederleini, Ps. lumbricoides, Ps. obscura, Ps. riedeli, Ps. theresopolitana, and Ps. wetzeli. The species Geoplana maximiliani sensu Schirch (1929) was renamed as Ps. schirchi Ogren & Kawakatsu, 1990. The present study reports a taxonomic revision of seven of Schirch’s species using type material, namely Obama itatiayana, Pasipha plana, Pseudogeoplana arpi, Ps. bresslaui, Ps. doederleini, Ps. schirchi and Ps. wetzeli. Additional specimens of some of these species were also examined. Morphological data from histological preparations and from virtual sections were obtained through a non-destructive technique of X-ray computed microtomography (µCT). This approach resulted in the preservation of the entire body of at least one type-specimen of each species, and the holotype of Ps. bresslaui. Conspecificity of O. itatiayana and P. plana was confirmed, as previously reported in the literature. It is also proposed that Ps. bresslaui belongs to the genus Paraba, while the other species should remain in Pseudogeoplana, since type-specimens are either immature, poorly preserved or simply lost.

Leading taxonomist Prof. Eudóxia Maria Froehlich (1929*–2015†) kept an uncatalogued personal collection of a large set of histological glass slides of land planarians. Many of these slides carry insufficient written information for identification purposes. 2287 slides of her collection were examined under the microscope and compared with relevant literature. We found 1633 slides bearing type material of a total of 119 nominal species, mostly collected in the Neotropical region. Identification of 148 slides (belonging to 16 species) as type material is suspected but not confirmed. Type material represents 110 triclad species. There is also type material of two Polycladida, two Rhabdocoela, three Proseriata, one Prolecitophora, and one Acoelomorpha species. The 119 nominal species were described by Ernest Marcus, Eveline du Bois Reymond-Marcus, Diva Diniz Corrêa, Cláudio Gilberto Froehlich, and Eudóxia Maria Froehlich. While keeping the slides in their original location in cupboards, the catalogue provides a quick means of retrieving desired slides. Slides were deposited in the Museu de Zoologia da Universidade de São Paulo.

Since 2008 there have been many records in Europe (British Isles, Spain, France, Italy) of a large terrestrial planarian morphologically very similar to the Brazilian species Obama marmorata. Sequences of mitochondrial (Cox1) and nuclear (18S, 28S, ITS-1 and EF) genes from European specimens and some from Brazil indicate that they belong to a species different from that of other specimens also collected in Brazil. Moreover, the phylogenetic results show that they are not sister-species. Histological sections of both Brazilian and European specimens reveal subtle morphological differences between the two species. Obama marmorata is confined to Brazil, and the second, herein described new species, Obama nungara sp. nov., is found in Brazil and Europe. These cryptic species may be syntopic in areas in Brazil. The new species occurs in human-modified environments both in Brazil and in Europe. We also conclude that the specimens from Spain and Argentina identified previously as Obama marmorata belong to the new species.

The land planarians in the genus Obama include the largest species of the Neotropical Geoplaninae. Morphological discrimination of Obama species can be difficult, as many species are morphologically similar and some present asymmetric – difficult to interpret – copulatory organs. New techniques are thus welcomed to provide faster species description and identification. Here, we study several specimens of Obama by means of mainly 2D and 3D μCT-based images obtained through X-ray microcomputed tomography (μCT) of a paratype, and, complementarily, histological sections of the holotype and a second paratype, which were digitized into virtual slides. Comparison of these images with traditional histological sections and descriptions of the known species of the genus allows us to conclude that our specimens constitute a new species, which we describe here. We further evaluate the phylogenetic placement of the new species using gene fragments from the mitochondrial cytochrome c oxidase subunit I and the nuclear elongation factor-1α. Raw and derivative μCT data and virtual histological sections were deposited in an open repository (GigaDB) and are freely available. This work leads us to conclude that μCT constitutes a relatively fast, inexpensive non-destructive method that produces results comparable to those of traditional histology, and is thus amenable for describing flatworm species.

A new genus and species of land planarian, Pichidamas piru gen. nov. sp. nov., is described from Pichidamas, province of Osorno in Southern Chile. Pichidamas piru is a sub-cylindrical and slender Geoplaninae characterized by the presence of a creeping sole extending 50% of the body width, a cephalic retractor muscle, five parenchymatic muscle layers, a male atrium with proximal tongue-like fold and distal adenodactyl; and ovovitelline ducts approaching the copulatory apparatus anteroventrally. Details of the organization of the cephalic retractor muscle, the absence of sensory pits, the existence of a parenchymatic muscle layer, located to the inside of the ventral peripheral nervous plexus, and an adenodactyl in the distal portion of the male atrium are unique traits seemingly reflecting an earlier branch of Geoplaninae. Fil: Bulnes, Verónica Natalia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Instituto de Ciencias Biológicas y Biomédicas del Sur. Universidad Nacional del Sur. Departamento de Biología, Bioquímica y Farmacia. Instituto de Ciencias Biológicas y Biomédicas del Sur; Argentina Fil: Grau, José H.. Leibniz-Institut für Evolutions und Biodiversitätsforschung; Alemania Fil: Carbayo, Fernando. Universidade de Sao Paulo; Brasil

The Brazilian land planarians Cratera crioula, C. joia, Geoplana hina, and G. taxiarcha (Platyhelminthes, Tricladida, Geoplanidae) are revised taxonomically from type material and additional specimens. Geoplana hina sensu Carbayo et al. (2013) was found to be an undescribed species and therefore is described and named as Cratera picuia sp. n. A new species of the genus is also described and named as Cratera arucuia sp. n. G. hina and G. taxiarcha are transferred to Cratera. The most remarkable morphological feature of Cratera—a dilated terminal portion of the ejaculatory duct—is either absent, inconspicuous, or variable in C. hina, C. joia, C. picuia sp. n., and C. arucuia sp. n. Based on the monophyletic status of Cratera inferred elsewhere, an emendation of the genus is here proposed to encompass the morphological variation observed in the genus.

The land planarian genus Choeradoplana (Plathelminthes, Tricladida) is currently integrated by 13 species. In previous works, morphological variation in its type species, Choeradoplana iheringi, was reported, but no attempt to test whether it is just a single species has been made yet. In order to disentangle the taxonomy of this species and further members of the genus, we sampled new specimens and combined morphological and molecular data and also have evaluated the performance of diverse methods of molecular species delimitation. Our data point to the presence of two cryptic species named C. iheringi, plus two new species, all hidden under the same general appearance. An in-depth morphological study of the specimens allowed detection of diagnostic morphological traits in each species, for which we also propose a molecular diagnosis. This integrative taxonomic study demonstrates once again the usefulness of molecular tools to weigh minor morphological characteristics and thus reveal the existence of species that would otherwise remain cryptic. However, under certain parameters, the molecular methods may over-split species with a high genetic structure, maybe pointing to incipient speciation. This makes critical the use of these methods combined with a comprehensive morphological approach. We also present a comprehensive phylogenetic tree including most Choeradoplana species. The tree, well supported, allows making some preliminary inferences on the evolution of the group and its historical biogeography.

GEOPLANA VAGINULOIDES (DARWIN, 1844) Planaria vaginuloides Darwin, 1844, p. 244. Geoplana vaginuloides: Riester, 1938, p. 72–75 [part]; Froehlich, 1956a, p. 314–315 [part]; Marcus, 1951, p. 54–56 (misidentification), 1952: p. 76–77 (misidentification). Material examined Neotype MZUSP PL 2074 (field number F6387): 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 N574: 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 N628: 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. Note We could not confirm conspecificity of some specimens studied by Riester (1938) and specimens examined by Froehlich (1955a, 1956a, 1958) (see Supporting Information, Table S1). Distribution Areas covered with Atlantic forest in the municipalities of Rio de Janeiro and Teresópolis, state of Rio de Janeiro. 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. Type locality Rio de Janeiro, Brazil. 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. 4A). 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. 4B). 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. 4C). 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. 4D). 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. 5A). Penis papilla very long, 5 mm, with dorsal and ventral insertions at the same horizontal plane (Fig. 4D). 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. 4D). 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. 5B). 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. 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). Neotype designation satisfies provisions of the ICZN code (Art. 75.3.1.–7.) as follows: (1) the neotype is designated with the express purpose of clarifying the taxonomic status or the type locality of the name Geoplana vaginuloides; (2) the characters differentiating G. vaginuloides from any other taxa are stated in the diagnosis above; (3) these data and the description of the species are sufficient to ensure recognition of the specimen designated; (4) after Dr Hugh Jones [from Natural History Museum London (NHM)], the name-bearing type specimen is lost or even has never been deposited in the NHM; (5) the ‘Remarks’ section shows that neotype is consistent with the former name-bearing type from the original description and from Darwin’s field notes (in Keynes, 2000); (6) the neotype came from the original type locality; and (7) the neotype is the property of the Museu de Zoologia da Universidde de São Paulo (MZUSP), Brazil., Published as part of 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), pp. 1-65 in Zoological Journal of the Linnean Society 185 on pages 23-26, DOI: 10.1093/zoolinnean/zly022, http://zenodo.org/record/2546555, {"references":["Darwin C. 1844. Brief descriptions of several terrestrial planariae, and of some remarkable marine species, with an account of their habits. Annals and Magazine of Natural History, Annales de Sciences Naturelles 14: 241 - 251.","Riester A. 1938. Beitrage zur Geoplaniden-Fauna Brasiliens. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 41: 1 - 88.","Froehlich CG. 1956 a. Tricladida Terricola das regioes de Teresopolis e Ubatuba. Papeis Avulsos de Zoologia 12: 313 - 344.","du Bois-Reymond Marcus ED. 1951. On South American Geoplanids. Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia 16: 217 - 255.","Froehlich CG. 1955 a. Sobre morfologia e taxonomia das Geoplanidae. Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia 19: 195 - 279.","Froehlich CG. 1958. On a collection of Brazilian land planarians. Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia 21: 93 - 121.","Marcus E. 1952. Turbellaria brasileiros. Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia 17: 5 - 187.","Keynes R. 2000. Charles Darwin's zoology notes & specimen list from H. M. S. Beagle. Cambridge: Cambridge University Press."]}

GEOPLANA IPORANGA ALMEIDA & CARBAYO SP.NOV. 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 F3166): 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 F3132) (immature): Cephalic region: horizontal sections on three slides; pharynx and copulatory apparatus: sagittal sections on 12 slides. Distribution Areas covered with Atlantic forest in Parque Estadual de Intervales, Ribeirão Grande, state of São Paulo, Brazil. Etymology The specific epithet is the name of another locality within the conservation area from where the species was collected. 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. 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. 20A). 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. 20B). 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. 20C). Penis papilla horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 21A, 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. 20C). 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. 21B). 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. GEOPLANA PULCHELLA SCHULTZE & MÜLLER, 1857 FIGS 22 AND 23 Geoplana pulchella Schultze & Müller, 1857: p. 25; Froehlich, 1955b, p. 189–190; du Bois-Reymond Marcus, 1951, p. 234–235 (misidentification). Material examined MZUSP PL 1068 (field number F3249): 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 F6528): 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 F6544): 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. Note We could not confirm identification of a specimen studied by Froehlich (1955b); see Supporting Information (Table S1). Distribution Areas covered with Atlantic forest in municipalities of Blumenau, Paulo Lopes, Alfredo Wagner, Eastern part of Santa Catarina state, South Brazil. 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. 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. 22A). 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. 22B). 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. 22C, 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. 23A). 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 ( Female atrium without folds (Fig. 22C, 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.

GEOPLANA BORACEIA ALMEIDA & CARBAYO SP.NOV. FIGS 25 AND 26 Material examined Holotype MZUSP PL 2070 (field number F4112): 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 F6575): 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 F6578): 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 F6357): 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 F4389): 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 F6026): 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 F6994): 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 F6068): 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 F6055): 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 F4358): 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. Distribution Areas covered with Atlantic forest in the municipalities of Salesópolis and São Sebastião, state of São Paulo, Brazil. 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. Etymology The specific epithet refers to the name of the biological station where the type material was collected. 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. 25A). 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. 25B). 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. 25D, 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 25C, D, 26A, 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. 25C). 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 25C, 26A, 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.

GEOPLANA CHITA FROEHLICH, 1956 FIG. 24 Geoplana chita ��� Froehlich, 1956b [in part]. 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 F6993): 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 F6995): Morretes/PR, Brazil. Magno C. Segalla, coll., 1 November 1987. Pharynx and copulatory apparatus: sagittal sections on 37 slides. MZUSP PL 715 (field number F1710) (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 F1730) (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. Note An unlabelled specimen from the original type material is not conspecific (see Discussion below and Supporting information, Table S1). Distribution Areas covered with Atlantic forest in the municipalities of Curitiba and Morretes, State of Paran��, South Brazil. 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. 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. 24A, 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 ��m 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. 24C). Outer pharyngeal musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (35 ��m) of intermingled circular and longitudinal muscle fibres. Inner pharynx musculature consisting of a thin subepithelial layer of longitudinal muscle, followed by a layer (38 ��m) of circular fibres and an innermost layer (10 ��m) 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-��m-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 ��m on the posterior half, and constituting a muscular cylin- der with a diameter equivalent to ten times the diameter of this duct (Fig. 24D���F). Penis papilla long, horizontal and cylindrical along most of its length, with dorsal and ventral insertions at the same transverse plane (Fig. 24D, 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 ��m 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 ��m thick dorsally and 160 ��m thick ventrally, followed by a 10-��m-thick layer of longitudinal muscle. Male atrium ample, not folded; lined with a low, non-ciliated epithelium, and underlain by a 10-��m-thick layer of circular muscles followed by a 10-��m-thick layer of longitudinal fibres. Ovaries oval shaped, 500 ��m 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 ��m long, and bent posterodorsally. Female atrium without folds, and being twice the length of the male atrium (Fig. 24F). Posterior section with 200 ��m in length of female atrium lined with a 20-��m-high, stratified-like, non-ciliated epithelium, which is surrounded by a 30-��m-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-��m-thick layer of circular-to-decussate muscle fibres, followed by a 40- to 150-��m-thick layer of longitudinal fibres., Published as part of 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), pp. 1-65 in Zoological Journal of the Linnean Society 185 on pages 54-56, DOI: 10.1093/zoolinnean/zly022, http://zenodo.org/record/2546555, {"references":["Froehlich CG. 1956 a. Tricladida Terricola das regioes de Teresopolis e Ubatuba. Papeis Avulsos de Zoologia 12: 313 - 344.","Froehlich CG. 1956 b. Planarias terrestres do Parana. Dusenia 7: 173 - 196."]}

A new species of the genus Issoca (Platyhelminthes, Tricladida, Geoplaninae) is described. Issocaassanga sp. n. presents the diagnostic features of the genus, with the exception of the relative position of the subneural parenchymal muscle layer with the cephalic retractor muscle, which are overlapped in the type species of the genus but are intersected in the new species. Rather than a polymorphic character, the relative position of these muscle layers might reflect the polyphyletic status of the genus.

GEOPLANA PARANAPIACABA ALMEIDA & CARBAYO SP . NOV . Material examined Holotype MZUSP PL 2079 (field number F6660): 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 F6998): Cidade Turística de Paranapiacaba, Santo André/ SP, Brazil. E. M. Froehlich et al., coll., 13 May 1990. Copulatory apparatus: sagittal sections on 39 slides. Distribution Only known from the type locality. Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the district where the type material was collected. 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. 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 (1/16th) posteriorly merging with each other, externally to which is a broader white line (one-eighth), and this is externally bordered by a black stripe (1/16th), externally to which is a grey stripe (one-eighth) with dense white mottling, then bordered by a marginal thin black line (Fig. 12A). Ventral body surface is whitish with blackish margins. In preserved specimen F6660, dorsal colour slightly faded. In preserved specimen F6998, 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. 12B, 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. 12D). 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. 13A, 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 12E, 13D). 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. 13A, 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. 13A, B, D). Female atrium long, without folds, as long as 3.5–4 times the male atrium length (Fig. 13A, 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.

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

GEOPLANA CANANEIA ALMEIDA & CARBAYO SP.NOV. FIG. 17 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 F4096): 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 F4097): pharynx: transverse sections on six slides; undeveloped copulatory apparatus: sagittal sections on three slides. Paratype MZUSP PL 2069 (field number F4098): pharynx: sagittal sections on eight slides; copulatory apparatus: sagittal sections on 13 slides. Distribution Area covered with Atlantic forest in Cardoso Island, municipality of Cananeia, state of São Paulo. Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the municipality where the species was collected. 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. 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 (1/13th 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. 17A). 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. 17B). 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. 17C) 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. 17D, 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. 17E), 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. 17E); 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.

GEOPLANA MOGI ALMEIDA & CARBAYO SP.NOV. FIGS 8 AND 9 Geoplana vaginuloides: Marcus, 1951, p. 54���56. Material examined Holotype MZUSP PL 2089 (field number F7000): Parque do Estado, S��o Paulo / SP, Brazil. O. Fran��oso 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 S��o 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 F1022): 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. Distribution Areas covered with secondary Atlantic forest in the municipalities of S��o Paulo and Mogi das Cruzes, state of S��o Paulo, Brazil. Etymology The specific epithet refers to the Tupi-Guarani (indigenous Brazilian tribe) name of the municipality where the species was collected. 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. 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. 8A). 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 F1022, whitish, blackish and greyish colours lost brightness; orange-ochre faded. In specimen F7000, 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 ��m high and 40 ��m in width; and a cup-shaped type, 40 ��m 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 ��m 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 ��m 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. 8B). Oesophagus-topharynx ratio, 15%. Outer pharyngeal musculature consisting of a subepithelial layer (5 ��m) of longitudinal muscle, followed by a layer (70 ��m) 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 ��m) of circular fibres and an innermost layer (30 ��m) 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 8C, 9A). 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-��m-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-��m-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-��m-thick layer of circular muscles, which constitutes a muscular cylinder possessing a diameter equivalent to 1.3 times the diameter of this duct (Fig. 9B). 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. 9C). Subepithelial musculature of penis papilla consisting of dense 80- to 90-��m-thick layer of circular muscle, the fibres of which are decussate distally, followed by a 10- to 20-��m 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-��m-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. 9A) and is lined with a 25-��m-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-��m-thick layer of circular-to-decussate muscle fibres, followed by some longitudinal fibres, and then a layer (50���150 ��m thick) of thicker longitudinal fibres., Published as part of 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), pp. 1-65 in Zoological Journal of the Linnean Society 185 on pages 29-32, DOI: 10.1093/zoolinnean/zly022, http://zenodo.org/record/2546555, {"references":["du Bois-Reymond Marcus ED. 1951. On South American Geoplanids. Boletins da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia 16: 217 - 255."]}