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Sabulirhynchus axi Artois & Schockaert, 2000 Fig. 3A–D Material examined PANAMA • 5 whole mounts; Pacific Ocean, Isla Iguana; 7°37′46″ N, 79°59′49″ W; depth 6 m; 6 Mar. 2016; subtidal, coarse sand; HU XIX.2.01–XIX.2.05 • 2 whole mounts; same collection data as preceding; 7°38′06″ N, 79°59′48″ W; depth 10 m; subtidal, coarse sand mixed with small rocks and covered with organic matter; HU XIX.2.06–XIX.2.07. Distribution Bahia Academy, Santa Cruz, Galapagos Islands (Artois & Schockaert, 2000). Remarks The specimens collected in Panama are morphologically similar to those from the Galapagos. The prostatic stylet type III (Fig. 3A–D) is 46–76 µm long (x = 62 µm; n = 7) and 10–31 µm wide (x = 21 µm; n = 7). A feature of the female system, not mentioned by Artois & Schockaert (2000), is the presence of a proximal, apparently sclerotised cap in the seminal receptacle.

Sabulirhynchus ibarrae sp. nov. urn:lsid:zoobank.org:act: 1EA7110C-D687-42BB-87A7-1259A24E4E0E Figs 3E–F, 4 Diagnosis Species of Sabulirhynchus Artois & Schockaert, 2000 with a very long proboscis, representing 40% of the body length. Prostatic stylet type III plate-shaped, ~53 µm long, proximally tubular and with a ridge at its midlength. Seminal reservoir exhibits a proximal apparently sclerotised cap. Etymology Species named after the late Dr María Elena Ibarra Martín (Marine Research Centre, Havana University, Cuba), head of the MRC for more than 25 years, who indefatigably championed the protection of oceans. Awarded with the National Prize of Environmental Sciences of Cuba in 2007. The Research Group of Marine Ecology of Universidad de Oriente carries her name. Material examined Holotype CUBA • 1 whole mount; Santiago de Cuba, Siboney; 19°57′34″ N, 75°42′07″ W; 7 Feb. 2019; depth 3 m; sublittoral, coarse-grained sand, salinity 34 ‰; https://id.luomus.fi/KV.700; FMNH. Other material CUBA • 1 whole mount; same collection data as for holotype; photographs of live specimens available; HU XIX.2.08. Description The specimens are unpigmented (Fig. 4A), ~ 1.5 mm long. The proboscis (Fig. 4A–B: pr) attains 40% of the body length. Caudally to the proboscis there is a pair of eyes (Fig. 4A–B: e). The pharynx is at the midbody (Fig. 4A–B: ph). A testis (Fig. 4A–B: t) is located beside the pharynx and occupies the middle third of the body. The prostatic stylet type III (Figs 3E–F, 4C; 4A–B, 4D: ps3) is plate-shaped, 49–56 µm long (x = 53 µm; n = 2), and 20–21 µm wide proximally; it is proximally tubular and exhibits a ridge at its midlength. The vitellarium (Fig. 4A–B: vi) runs from the posterior end of the proboscis to the caudal body end. The oval ovary (Fig. 4B, D–E: ov) has the oocytes organised in a row. The female bursa (Fig. 4A–B, D: b) is positioned caudally to the male atrial organs. The spindle-shaped seminal receptacle (Fig. 4B, D–E: sr) is filled with sperm and proximally exhibits a seemingly sclerotised cap (Fig. 4D–E: sc). The gonopore opens at 80% (Fig. 4B: cg). Remarks Until now, the genus Sabulirhynchus included a single species: Sabulirhynchus axi. Sabulirhynchus ibarrae sp. nov. and S. axi share the diagnostic features of the genus: lack of accessory glandular organ in the male system, a plate-shaped prostatic stylet type III, and a pyriform seminal receptacle in the female atrium (Artois & Schockaert 2000). Considering the newly analysed material of both species, we add to the diagnosis of Sabulirhynchus the presence of a proximal, apparently sclerotised cap in the seminal receptacle. This feature is easily recognisable in live specimens; however, we did not distinguish it on the whole mounts. Therefore, the sclerotised nature of this structure is questionable. The prostatic stylet is similar in length in both species, 52–63 µm long in S. axi (population from the Galapagos) and ~53 µm in S. ibarrae sp. nov. However, it is more variable in the population of S. axi from Panama (46–76 µm long). The stylet of S. axi is a simple plate, distally turned backward (Artois & Schockaert 2000), whereas it is straight, proximally tubular, and with a ridge at its midlength in S. ibarrae sp. nov. These differences in stylet morphology warrant the status of a new species., Published as part of Diez, Yander L., Sanjuan, Claudia, Monnens, Marlies & Artois, Tom, 2023, New species of Polycystididae (Platyhelminthes: Kalyptorhynchia) from Cuba and the Pacific coast of Panama, pp. 67-86 in European Journal of Taxonomy 856 on pages 78-80, DOI: 10.5852/ejt.2023.856.2029, http://zenodo.org/record/7554565, {"references":["Artois T. & Schockaert E. 2000. Interstitial fauna of the Galapagos: Typhlopolycystidinae (Platyhelminthes, Polycystididae). Tropical Zoology 13: 141 - 158. https: // doi. org / 10.1080 / 03946975.2000.10531128","Evdonin L. A. 1977. Monograph of the Turbellaria Kalyptorhynchia in the fauna of the USSR and adjacent areas [in Russian]. Fauna of the USSR 115: 1 - 400."]}

Phonorhynchoides lalanai sp. nov. urn:lsid:zoobank.org:act: 94BB22AE-7C11-4ECB-A344-B8ACFB01CF28 Figs 1C, I–J, 2D, G, Table 2 Phonorhynchoides sp. in Leasi et al. 2018: 1–12. Diagnosis Species of Phonorhynchoides Beklemischev, 1927 with a prostatic stylet type IV ~114 µm long, proximally ~9 µm, narrowing to 5 µm wide distally; it is twisted in the middle. Accessory stylet type IV 105–106 µm long, narrowing from 4–5 µm wide proximally to 2 µm wide distally; slightly curved in the proximal half. Etymology Species named after the late Dr Rogelio Lalana (Marine Research Centre, Havana University, Cuba), prominent Cuban marine researcher, awarded with the National Prize of Marine Sciences of Cuba in 2012, for his outstanding contribution to the knowledge of marine invertebrates, mainly crustaceans. Material examined Holotype CUBA • 1 whole mount; Santiago de Cuba, Bueycabón; 19°57′38″ N, 75°57′28″ W; 6 Feb 2018; depth 0.5 m; sublitoral, fine-grained sand rich in organic matter, salinity 33 ‰; https://id.luomus.fi/KV.699; FMNH. Other material PANAMA • 2 whole mounts; Pacific Ocean, Isla Iguana; 07°38′06″ N, 79°59′48″ W; 6 Mar. 2016; depth 10 m; sublittoral, coarse sand with gravel, covered with organic matter; HU XIX.1.49–XIX.1.50. Description The specimens are unpigmented (Fig. 2D), ~ 1 mm long. The proboscis (Figs 1C, 2D: pr) is less than 10% of the body length. A pair of eyes is located just behind the pharynx (Figs 1C, 2D: e). The pharynx (Figs 1C, 2D: ph) is positioned in the anterior body half. A pair of testes (Fig. 1C: t) is located posterior to the pharynx. The prostatic stylet type IV (Figs 1C, 2G: ps4, 1I) is 110–120 µm long (x = 114 µm; n = 3) and narrows from a width of 8–11 µm proximally (x = 9 µm; n = 3) to 5 µm distally (n = 3); the distal end is rounded. At about midway, the prostatic stylet is twisted. The accessory stylet type IV (Figs 1C, 2G: as4, 1J) is 105–106 µm long (n = 3), slightly curved in the proximal half, and narrows from 4–5 µm wide proximally (n = 3) to 2 µm wide distally (n = 3); the distal end is oblique and sharp. The vitellaria (Fig. 1C: vi) run from just behind the pharynx to the caudal body end. The female bursa (Fig. 1C: b) is located beside the stylets; it is muscular but not bipartite. The ovaries (Fig. 1C: ov) are located rostrally to the stylets; they are more or less kidney-shaped, with the oocytes organised in a row; the oocytes distally increase in diameter. Remarks Phonorhynchoides lalanai sp. nov. exhibits the diagnostic features of Phonorhynchoides: the prostatic stylet type IV is larger than the accessory stylet type IV, and the female bursa is muscular but not bipartite (Willems et al. 2017). The prostatic stylet of P. lalanai sp. nov. is twisted at the end of its proximal half, while this structure is never twisted in the other six known species of the genus. For comparison, the prostatic stylet is undulated in P. carinostylis Ax & Armonies, 1987 and P. gondwanae Willems & Artois in Willems et al., 2017, corkscrew-shaped in the middle in P. somaliensis Schockaert, 1971, and almost straight in P. flagellatus Beklemischev, 1927, P. japonicus Ax, 2008, and P. minor Diez, Sanjuan, Reygel & Artois in Diez et al., 2018b (for details see Willems et al. 2017; Diez et al. 2018b). Furthermore, the prostatic stylet carries ornamentations in P. carinostylis (spiral ridge over the whole length) and P. japonicus (thickened proximally and short spiral ridge), which are missing in P. lalanai. In P. lalanai, the prostatic stylet is distally rounded, differing from that in P. minor (terminally oblique) and the other species (sharp tip). The prostatic stylet of P. lalanai. (~114 µm) is considerably longer than that of P. minor (53–58 µm), but shorter than that in the other species (see Table 2). The accessory stylet type IV of P. lalanai sp. nov. ends in an oblique tip, while it ends in a pointed tip in all other species of Phonorhynchoides. The accessory stylet of the new species (105–106 µm) is smaller than that of P. gondwanae and P. japonicus, but larger than that of the other species (see Table 2). The length proportion between the accessory stylet type IV and the prostatic stylet type IV is larger in the Caribbean species (P. lalanai 88–95%, P. minor 78–85%) than in their known congeners (Willems et al. 2017; Diez et al. 2018b) (see Table 2). Indeed, it is characteristic of P. lalanai and P. minor that the prostatic stylet and accessory stylet are similar in length. The distinct morphology of the sclerotised stylets, previously discussed, warrants the status of P. lalanai as a new species.

Djeziraia adriani sp. nov. urn:lsid:zoobank.org:act: 39C06707-C4B5-4640-9B0D-7CAC9784D00E Figs 1B, G–H, 2B–C, F Diagnosis Species of Djeziraia Schockaert, 1971 with paired seminal vesicles. Gonads and atrial organs located caudally in the body. Prostatic stylet type IV 92–93 µm long, curved, S-shaped in live animals. Etymology Species named after Dr Adrian David Trapero Quintana (Havana University, Cuba), specialist in freshwater ecology and the taxonomy of Odonata. Material examined Holotype CUBA • 1 whole mount; Santiago de Cuba, Sardinero; 19°57′42″ N, 75°47′00″ W; 18 Mar. 2021; intertidal, mangrove area surrounding the mouth of Río Sardinero, 80 m from the beach, sample of sediment with rotten leaves and branches, salinity 32 ‰; https://id.luomus.fi/KV.698; FMNH. Other material CUBA • 2 whole mounts; same collection data as for holotype; photographs of live specimens available; HU XIX.1.47–XIX.1.48. Description The live specimens are unpigmented (Fig. 2B–C), 0.8–0.9 mm long, measured on whole mounts. A pair of well-differentiated bristles (Figs 1B, 2B: bt) is located at the sides of the proboscis pore. The proboscis (Figs 1B, 2B: pr) represents less than 10% of the body length. A pair of eyes (Figs 1B, 2B: e) is located just behind the proboscis. The pharynx (Figs 1B, 2B: ph) is positioned in the anterior body half. The gonads and atrial organs occur in the caudal body half (Figs 1B, 2C). The testes are located rostrally to the ovaries. The paired seminal vesicles (Figs 1B, 2F: sv) open through a single ejaculatory duct into the copulatory bulb. The ejaculatory duct is surrounded by the prostate vesicle type IV (Figs 1B, 2F: pv4). The cell bodies of the prostatic glands are extracapsular and enter the copulatory bulb proximally. The prostatic stylet type IV (Figs 1B, 2C, F: ps4; 1G–H) is tubular and 92–93 µm long (n = 3). The proximal funnel-shaped part of the stylet is 9–10 µm wide, and it tapers to a distally rounded tip, subdistally 4 µm wide; the aperture is completely terminal. Oocytes are organised in a row and proximally diminish in diameter. The vitellaria run laterally, from the pharynx almost to the caudal body end. The female bursa is located next to the stylet. The common gonopore opens at 90%. Remarks Djeziraia adriani sp. nov. exhibits most diagnostic features of Djeziraia: the proboscis is very small, less than 10% of the body length; the atrial organs are located in the caudal body half; the prostate vesicle is interposed; and there is a single prostatic stylet type IV in the male atrium (cf. the other representatives of Phonorhynchoidinae Tessens, Janssen & Artois, 2014 with two or three stylets) (see Schockaert 1971; Artois & Schockaert 2001). A striking feature of the new species is the fact that there are two seminal vesicles, while there is only one in its congeners: Djeziraia euxinica (Mack-Fira, 1971) sensu Schockaert 1982, D. incana Artois & Schockaert, 2001, D. longystila Noreña, Damborenea, Faubel & Brusa, 2007, and D. pardii Schockaert, 1971. Furthermore, the presence of a diverticulum at the bursal stalk (Schockaert 1971) was not observed in our live specimens. Despite these differences, the overall morphology of the new species largely corresponds to what is described for Djeziraia, and we therefore provisionally include it in this genus. Molecular phylogenetic work may prove useful to confirm this preliminary classification. The prostatic stylet type IV of all species of Djeziraia is a simple tube: it is slender and very long (D. longystila and D. incana) or short and straight (D. euxinica and D. pardii). In D. euxinica, the stylet is longitudinally striated and ends in a small hook. The general morphology of the prostatic stylet of D. adriani sp. nov. is comparable to that of D. longystila and D. incana. However, it is S-shaped in live specimens of the new species. Furthermore, the prostatic stylet of D. adriani (92–93 µm) is shorter than that of D. longystila (~275 µm; Noreña et al. 2007), D. incana (~179 µm, 295 µm in one specimen; Artois & Schockaert 2001), and D. euxinica (105–120 µm; Mac-Fira 1971; Artois & Schockaert 2001), and larger than that of D. pardii (~84 µm; Schockaert 1971). Both the presence of paired seminal vesicles and the morphology of the stylet support the validity of the new species, Djeziraia adriani sp. nov., Published as part of Diez, Yander L., Sanjuan, Claudia, Monnens, Marlies & Artois, Tom, 2023, New species of Polycystididae (Platyhelminthes: Kalyptorhynchia) from Cuba and the Pacific coast of Panama, pp. 67-86 in European Journal of Taxonomy 856 on pages 72-74, DOI: 10.5852/ejt.2023.856.2029, http://zenodo.org/record/7554565, {"references":["Tessens B., Janssen T. & Artois T. 2014. Molecular phylogeny of Kalyptorhynchia (Rhabdocoela, Platyhelminthes) inferred from ribosomal sequence data. Zoologica Scripta 43 (5): 519 - 530. https: // doi. org / 10.1111 / zsc. 12066","Schockaert E. R. 1971. Turbellaria from Somalia 1. Kalyptorhynchia (part 1). Monitore Zoologico Italiano 4 (5): 101 - 122.","Artois T. & Schockaert E. 2001. Interstitial fauna of the Galapagos: Duplacrorhynchinae, Macrorhynchinae, Polycystidinae, Gyratricinae (Platyhelminthes, Polycystididae). Tropical Zoology 14: 63 - 85. https: // doi. org / 10.1080 / 03946975.2001.10531143","Mack-Fira V. 1971. Deux turbellaries nouveaux de la Mer Noire. Revue romaine de Biologie Serie de Zoologie 16 (4): 233 - 240.","Schockaert E. R. 1982. Turbellaria from Somalia 2. Kalyptorhynchia (part 2). Monitore Zoologico Italiano 18 (2): 81 - 96.","Norena C., Damborenea C., Faubel A. & Brusa F. 2007. Composition of meiobenthonic Platyhelminthes from brackish environments of the Galician and Cantabrian coasts of Spain with the description of a new species of Djeziraia (Polycystididae, Kalyptorhynchia). Journal of Natural History 41 (29 - 32): 1989 - 2005. https: // doi. org / 10.1080 / 00222930701526055"]}

Brachyrhynchoides ortizi sp. nov. urn:lsid:zoobank.org:act: 774F2120-7B24-4F35-82EE-C46D93D9D9A7 Figs 1A, D–F, 2A, E, Table 1 Diagnosis Species of Brachyrhynchoides Artois & Schockaert, 2013 with the prostatic stylet type IV tubular, ~70 µm long, ~15 µm wide proximally, distally slightly narrower; terminal aperture oblique. Accessory stylet I tubular, 70 µm long; accessory stylet II needle-shaped, 59 µm long. Etymology Species named after Dr Manuel Ortiz (Marine Research Centre, Havana University, Cuba), prominent Cuban marine researcher, for his outstanding contribution to the knowledge of marine invertebrates, mainly crustaceans. Material examined Holotype CUBA • 1 whole mount; Santiago de Cuba, Siboney; 19°57′34″ N, 75°42′07″ W; 7 Feb 2019; depth 2 m; sublittoral, coarse-grained sand with fragments of calcareous algae, salinity 34 ‰; https://id.luomus.fi/KV.697; FMNH. Other material CUBA • 1 whole mount; same collection data as for holotype; photographs of live specimens available; HU XIX.1.46. Description The live specimens are unpigmented (Fig. 2A), ~ 1.5 mm long. The proboscis (Figs 1A, 2A: pr) measures less than 10% of the body length. A pair of eyes (Figs 1A, 2A: e) is located caudally to the proboscis. The pharynx (Figs 1A, 2A: ph) is located in the anterior body half. A pair of testes (Figs 1A, 2A: t) is located caudally to the pharynx. The prostatic stylet type IV (Figs 1A, 2A, 2E: ps4; 1D) is tubular, 68–71 µm long (x = 70 µm; n = 2) and 12–17 µm wide proximally (x = 15 µm; n = 2); distally, it is slightly narrower, with the terminal aperture oblique. The accessory stylet I (Figs 1A, 2A, E: as1; 1E) is tubular, 70 µm long (n = 1), 4 µm wide proximally (n = 1), and 2 µm wide distally (n = 1); its terminal aperture is oblique. The accessory stylet II (Figs 1A, 2E: as2; 1F) is needle-shaped, 59 µm long (n = 1), and 3 µm wide proximally (n = 1); it is slightly curved in the distal half and ends in a pointing tip. The proportions between the stylets were calculated following the methods of Artois et al. (2013): α = (length of the accessory stylet I / length of the prostatic stylet) × 100 = 98–103%; β = (length of the accessory stylet II / length of the prostatic stylet) × 100 = 83–87%; γ = (length of the accessory stylet II / length of the accessory stylet I) × 100 = 84%. The vitellaria (Figs 1A, 2A: vi) run at the body sides, from just behind the pharynx to almost the caudal end of the body. The left vitellarium was much smaller than the right one but this may be caused by of squeezing or orientation of the animal. The ovaries (Fig. 1A: ov) are oval-shaped, located rostrally to the stylets; only a single oocyte was observed in each ovary. Remarks The new species, Brachyrhynchoides ortizi sp. nov., shows the diagnostic features of the genus, as listed by Artois et al. (2013): the proboscis is very small (less than 10% of the body length) and three stylets occur in the male atrial organs (a prostatic stylet type IV, an accessory stylet I, and an accessory stylet II). However, these features need revaluation as Tessens et al. (2014) showed the genus to be paraphyletic. Three species of Brachyrhynchoides were known until now: B. triplostylis Artois & Schockaert, 2013, B. acutus Artois & Schockaert, 2013, and B. oosterlyncki Willems, Reygel & Artois, 2013, which can be distinguished by differences in stylet morphology (Artois et al. 2013). The prostatic stylet type IV of B. ortizi sp. nov. is the shortest and widest with respect to that of its congeners (see Table 1). It is more similar in length to that of B. oosterlyncki; however, the prostatic stylet of B. ortizi is straight, slightly wider proximally (15 µm) than distally, and with the distal aperture oblique, whereas in B. oosterlyncki the stylet is 8 µm wide proximally and tapers to a distally-pointing tip. The other two species of Brachyrhynchoides have a much larger prostatic stylet (see Table 1). Furthermore, in B. triplostylis and B. acutus, the prostatic stylet is curved and gradually tapers to a sharp (B. acutus) or rounded (B. triplostylis) tip (Artois et al. 2013). According to Artois et al. (2013), the accessory stylet I in species of Brachyrhynchoides is connected to the larger accessory vesicle. However, no accessory vesicles were observed in B. oosterlyncki and, therefore, the authors considered the shorter stylet to be accessory stylet I. For comparative purposes, we will consider accessory stylet I the larger in all species. In B. ortizi sp. nov., the tubular accessory stylet I is 70 µm long, with the terminal aperture oblique, and more or less of the same width over its entire length. On the other hand, the accessory stylet I in B. oosterlyncki is shorter, needle-shaped, and wider proximally. The accessory stylet I of B. triplostylis and B. acutus is much larger (Artois et al. 2013) (see Table 1). The accessory stylet II in all species of Brachyrhynchoides is needle-shaped. The smallest stylet appears in B. oosterlyncki, followed by B. ortizi sp. nov., and it is much larger in B. triplostylis and B. acutus (see Table 1). The accessory stylet II of B. ortizi is slightly curved and its distal half ends in a pointed tip. It is similar in morphology to what is described for other species, with the exception of B. acutus, where the accessory stylet II is distinctly curved. The proportions between stylets vary greatly interspecifically (Table 1, see Artois et al. 2013).

The origin of introduced Nile tilapia stocks in sub-Saharan Africa is largely unknown. In this study, the potential of monogeneans as a biological tag and magnifying glass is tested to reveal their hosts' stocking history. The monogenean gill community of different Nile tilapia populations in sub-Saharan Africa was explored, and a phylogeographic analysis was performed based on the mitogenomes of four dactylogyrid species (Cichlidogyrus hani, C. sclerosus, C. thurstonae, and Scutogyrus longicornis). Our results encourage the use of dactylogyrids as biological tags. The magnifying glass hypothesis is only confirmed for C. thurstonae, highlighting the importance of the absence of other potential hosts as prerequisites for a parasite to act as a magnifying glass. With the data generated here, we are the first to extract mitogenomes from individual monogeneans and to perform an upscaled survey of the comparative phylogeography of several monogenean species with unprecedented diagnostic resolution. The Special Research Fund of Hasselt University finances MG (7NI02), AJCL (BOF19OWB02) and MPMV (BOF20TT06). Molecular work was funded through research grant 1513419 N of the Research Foundation—Flanders (FWO-Vlaanderen), and part of leading to results presented in this publication was carried out with infrastructure funded by the European Marine Biological Research Centre (EMBRC) Belgium, FWO-Vlaanderen project GOH3817N. The field expedition to the DRC was partly financed by the Stichting ter Bevordering van het Biodiversiteitsonderzoek in Afrika (SBBOA) fund.the research We appreciate the help of the people involved in the field work and sampling procedure. Furthermore, we would like to thank Christopher Laumer (EMBL-European Bioinformatics Institute, UK) and Gontran Sonet (Royal Belgium Institute of Natural Sciences, Belgium) for providing us with the extraction protocol and R scripts, respectively. We could also like to acknowledge Wim Bert, Christoph Hahn, Nikol Kmentova, ´ Jos Snoeks, and Carl Vangestel for suggestions on an earlier version of this manuscript. Finally, we would like to thank Ria Vanderspikken (Hasselt University, Belgium) and Natascha Steffanie (Hasselt University, Belgium) for their technical support in the laboratory

Orostylis gallicus sp. nov. Gobert, Monnens & Artois (Figs. 3F, 4F) Localities. More or less 70 m in front of a cliff at Wimereux beach, Wimereux, France (50��45���36���N; 1��41���31���E) Sand and shell gravel among algae scraped of rocks (Sep. 16, 2015): Type locality. More or less 70 m in front of a cliff at Wimereux beach, Wimereux France (50��45���36���N; 1��41���31���E) Red algae on rocks (Sep. 16, 2015). Material. Two whole mounts, one of which designated holotype (FMNH KV.695), the other one in the HU reference collection (HU XIV.3.10). Etymology. The species is named after the ancient Celtic people that inhabited the region of the type locality (Lat. ���gallicus��� = Gallic). Description. Based on observations of the whole mounts, the general morphology is as described for the genus, with the copulatory organ located anteriorly in the body, lateral to the pharynx. The number of testes is unknown. The stylet (Figs. 3F, 4F) is a strongly curved, ���raptor-claw���-shaped hollow spine, and is 45���51 ��m long. The proximal part is funnel-shaped and has an asymmetric aperture with a rounded flap. The distal third of the stylet is thin and tapers into a sharp tip. Laterally, the stylet shows a narrow, longitudinal slit. Additional details of the male copulatory system and female genital system could not be observed on the available specimens., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on page 37, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911

Orostylis asinaraensis sp. nov. Gobert, Jouk, Revis & Artois (Figs. 3B���C, 4A) Localities. Cala Sant���Andrea, Asinara National Park, Sardinia, Italy (41��00���48���N; 8��14���56���E) Sand from 2 m deep (Sep. 27, 2014): Type locality. Cala dei Ponzesi, Asinara National Park, Sardinia, Italy (41��05���14���N; 08��20���45���E) Sand from the intertidal (Sep. 30, 2014). Material. Observations on live specimens. Two whole mounts, one of which designated holotype (FMNH KV.691), the other deposited in the HU XIV.3.06. Etymology. Species named after the National Park and island of Asinara, Sardinia, where the species was discovered. Description. Specimens are transparent, unpigmented. A pair of round eyes (Fig. 4A: e) is present anteriorly. The overall internal organisation of the specimens corresponds to that of Orostylis dohae sp. nov. The mouth is situated ventrally, near the anterior body end. A horizontally positioned cylindrical pharynx (Fig. 4A: ph) is situated in the anterior third of the body. The anterior edge of the pharynx bears several small papillae, but long tentacles as in O. dohae sp. nov. were not observed. The single testis (Fig. 4A: t) is situated ventrally around the midbody. The male copulatory system lies alongside the pharynx. It consists of a large copulatory bulb containing a seminal vesicle (Fig. 4A: sv) and prostatic vesicle. In live specimens, circular muscles clearly surround the copulatory bulb. The seminal and prostate vesicles enter the stylet separately. The stylet (Figs. 3B,C; 4A: st) is a strongly curved, ���raptor-claw���-shaped, hollow spine, with a ���handle-like��� extension projecting proximally. In one of the specimens (HU XIV.3.06), the stylet is straighter, with the distal third bent to one side (Fig. 3C). The total stylet (incl. the handle) measures 64���76 ��m (x = 70 ��m; n = 2). The handle is 15���19 ��m long (x = 17 ��m; n = 2). The stylet shows a longitudinal slit on one side. Based on observations of live specimens, it is likely that the male system opens into the oral cavity, as is the case in O. dohae sp. nov. An ovary (Fig. 4A: ov) was observed in the midbody. It is large and, together with the intestine, takes up most of the posterior two thirds of the body. A single vitellarium fills the caudal end of the body, with two large branches stretching forwards up to the level of the copulatory organ., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on pages 33-34, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911

Orostylis timucuorum sp. nov. Smith III (Figs 3H, 5, S 1���S 3) Localities. Small beach facing the intracoastal waterway near the boat launch and pier, Whitney Marine Laboratory, FL (29��40���14���N; 81��12���57���W) Muddy sand with oyster shells and shell fragments (May 21, 2019): Type locality. Bear Island, NC (approximately 34��38���51���N; 77��07���02���W) Muddy sand-flat near tidal drainage stream (Jun., 2018). Outer sandy beach on Bogue Banks, Emerald Isle Site, NC (34��38���42���N; 77��05���23���W) Surface sediment halfway between MTL and LTL (Mar. 15, 2013). ���Anne McCrary���s Mudflat��� (muddy sand-flat facing the intracoastal waterway near Wilmington, NC ���approximately 34��13���11���N; 77��48���49���W) (Jan. and Feb., 1970) by R. M. Rieger. Bogue Inlet, NC. Shallow subtidal in ��� Amphioxus sand��� (date unknown) by R. M. Rieger. Material. Five specimens studied alive; two turned into whole mounts; one designated holotype (FMNH KV.696); the other reference material (HU XIV.3.19); three specimens examined by confocal microscopy; three live-drawings made by R. M. Rieger. Etymology. The specific epithet is in honour of the people (called ���Timucua��� by the Spanish) who first inhabited the region near the type locality. Description. Living, sexually mature specimens from the type locality measure 705���895 ��m long; all specimens possess a pair of pigmented eyes, as well as the general characters of the genus (Fig. 5A). The pharynx averages 16% of the body length (Fig. 5A: ph; n = 3). The pharynx bears 6 club-tipped, distal tentacles (Figs. 5A inset, S1). Radial pharyngeal musculature is thickly arranged and pseudostriated (Fig. 5B: rm). The male organ is directed forward and lies just behind or beside the pharynx; testes are paired (Figs. 5A: t, S2). The ovary is V-shaped, with paired germinal zones, sometimes giving the impression of two ovaries (Figs. 5A,C: ov). The vitellarium is either very large, or paired, situated immediately behind the ovary and extending posteriorly (Fig. 5C: vit). A bursal tissue lies in the ventral midline at approximately the level of the ovaries (Fig. 5A,D: bu). Confocal microscopy reveals a musculature associated with this that appears to lead to a mid-ventral pore (Figs. 5D,E: red arrow). The 58-��mlong stylet ranges in length from 47 to 69 ��m (n = 5), possessing a flared base with two distinct areas (Figs. 3H, 5F���G; 5H: sb, st). The stylet appears to have separate pathways for sperm and prostatic secretions. The stylet has a lateral slit that extends at least partway along it (Fig. 5F, arrows). Squeezing apparently flattens a stylet that, at rest, comprises a base with an elongate, semi-circular tip whose point rests in a small ���cup��� (Fig. 5H: green arrow); the base and tip are in nearly orthogonal planes (Fig. 5H: red arrows). Connection of the male canal with the pharynx tube could not be confirmed with the material at hand., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on page 37, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911

Orostylis dohae sp. nov. Gobert, Van Steenkiste & Artois (Figs. 2, 3A, 4D) Neodalyellida n. gen. n. sp. 2 in Van Steenkiste et al. (2013) Neodalyellida sp. 2 in Stephenson et al. (2019) Localities. Doha, Qatar (25��19���04���N; 51��32���18���E) Sand at about the high-water mark (Mar. 19, 2008): Type locality. Material. Observations on live specimens. One whole mount designated holotype (FMNH KV.690), and two serially sectioned specimens (HU XIV.3.04��� XIV.3.05). Etymology. Species named after the capital of Qatar, where the species was discovered. Description. Animals are about 0.6 mm long (measured on whole mount), with two dark brown, kidney-shaped eyes (Figs. 2A, 4D: e). The epidermis is syncytial, strongly ciliated, and 2���5 ��m thick. The cilia measure 2���3 ��m. Nuclei in the epidermis are scarce. A thin basal membrane underlies the epidermis, followed by a circular and a longitudinal muscle layer, respectively. Several basophilic, coarse-grained glands are situated in the anterior end of the body, in the area surrounding the mouth. The caudal end of the body contains eosinophilic cement glands. The mouth is located near the anterior end of the body (Fig. 2A: m) and surrounded by a sphincter. A long, thin-walled oral tube connects the mouth opening to the pharynx. The pharynx (Figs. 2, 4D: ph) is located in the anterior body half and is between 1/4 and 1/5 of the body length long. The pharynx is cylindrical in shape, with a conspicuous musculature, including radial, circular, and longitudinal muscles. This is typical for the pharynx doliiformis in neodalyellids, but the radial muscles are very wide and stacked closely together with little or seemingly no space between them. The anterior end of the pharynx bears a ���collar��� (Fig. 2A: phc) separated by a constriction from the rest of the pharynx. This collar is provided with three pairs of tentacles projecting anteriorly (Figs. 2A, 4D: pht). The intestine (Fig. 2B: i) lies caudally from the ovary (Fig. 2: ov). In live specimens, diatom frustules were observed inside the intestinal tract. The large, paired testes are situated just anterior to the midbody (Figs. 2, 4D: t), behind the pharynx, on either side of the body. In live specimens, two distinct testes were clearly observed. However, in the serial sections, the separation between both testes was less obvious, creating the impression of a single, large testis. The two testes lie against one another in such a way that the spermatogenic distal ends of the testes point laterally.A pair of vasa deferentia extend anteriorly and connect the testes to the seminal vesicle (Figs. 2, 4D: sv) in the proximal half of the male copulatory organ. The copulatory organ is situated in the anterior part of the body, alongside the pharynx. It consists of the round to ovoid seminal vesicle (Figs. 2, 4D: sv), followed distally by a small, globular prostate vesicle (Fig. 2B: pv) filled with eosinophilic gland secretions. The seminal and prostate vesicle are lined with a syncytial, nucleated epithelium and surrounded by circular muscles. In the distal half of the copulatory organ, the seminal vesicle connects directly to a sclerotised stylet (Figs. 2, 4D: st; 3A). The stylet is a simple, thin-walled funnel, curved at an angle of about 90�� at its midpoint. The stylet is 33 ��m long and 9 ��m wide proximally. It connects to a long male duct, which opens in the oral cavity (Fig. 2B: oc). The female genital system consists of a large, curved ovary (Fig. 2: ov), positioned at the midbody, dorsally to and in front of the intestine. A female duct or gonopore was not observed. Instead, the oviduct appears to connect directly to the intestinal lumen. It is assumed that eggs are deposited in the intestinal lumen and expelled from the body via the mouth. Eosinophilic glands are present in the region where the ovary connects to the intestine. In the same area, vesicles with sperm were observed in the intestinal wall (Fig. 2A: isv). Possibly, these vesicles are resorptive and function as a bursa. The posterior half of the body is almost completely taken up by a lobed or branched vitellarium (Fig. 2B: vit)., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on pages 31-33, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911, {"references":["Van Steenkiste, N., Tessens, B., Willems, W., Backeljau, T., Jondelius, U. & Artois, T. (2013) A comprehensive molecular phylogeny of Dalytyphloplanida (Platyhelminthes: Rhabdocoela) reveals multiple escapes from the marine environment and origins of symbiotic relationships. PLoS ONE, 8, e 59917. https: // doi. org / 10.1371 / journal. pone. 0059917","Stephenson, I., Van Steenkiste, N. W. L. & Leander, B. S. (2019) Molecular phylogeny of neodalyellid flatworms (Rhabdocoela), including three new species from British Columbia. Journal of Zoological Systematics and Evolutionary Research, 57, 41 - 57. https: // doi. org / 10.1111 / jzs. 12243"]}

Orostylis gen. nov. Gobert, Armonies, Diez, Jouk, Monnens, Revis, Reygel, Smith III, Van Steenkiste & Artois Etymology. The name Orostylis is derived from the Latin ���os��� (mouth), and ���stylis��� (stylet). The name refers to the fact that the male copulatory apparatus connects to the mouth in the type species. Diagnosis. Anterior-facing, cylindrical pharynx and sclerotised stylet in the anterior part of the body. Male genital opening in the oral cavity. Single ovary and vitellarium (O. timucuorum sp. nov. might have two vitellaria). No other female genital structures. Testes paired or unpaired. No visible connection between the female genital system and the environment, except possibly in O. timucuorum sp. nov. Grammatical gender. masculine Type species. Orostylis dohae, sp. nov., described below., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on page 31, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911

Orostylis caecus sp. nov. Diez & Artois (Figs. 3D, 4B) Localities. Bah��a Larga, Guam�� municipality, Santiago de Cuba province, Cuba (19��58���57���N; 76��18���15���W) Intertidal, very fine sand in the upper 10 cm of the sediment, salinity 34��� (Apr. 5, 2017): Type locality. Bah��a de Gibara, Holgu��n province, Cuba (21��06���00���N; 76��08���09���W) Intertidal, on very fine sand with organic material close to the mouth of the river, salinity 32 ��� (Feb. 17, 2016). Material. Observations on four live specimens, three of which were whole mounted afterwards, one designated holotype (FMNH KV.692), the others deposited in the HU XIV.3.07��� XIV.3.08. Etymology. The specific name refers to the species��� lack of eye spots (Lat. ���caecus��� = blind). Description. General morphology as described for the species above, with the stylet located anteriorly, lateral to the pharynx (Fig. 4B: ph, st). The body is unpigmented, slender, and 0.25���0.42 mm long (measured on two whole-mounted specimens). Eye spots are lacking. The mouth opens subterminally and is surrounded by few tactile hairs. The pharynx (Fig. 4B: ph) is 1/6���1/3 of the body length in live animals, measuring 104���120 ��m in two whole mounts and bearing six slender tentacles anteriorly (Fig. 4B: pht). The paired testes are located at both sides of the pharynx, each one connected to a seminal vesicle. The two seminal vesicles enter a small prostate vesicle. The stylet (Figs. 3D; 4B: st) is formed by a broad, proximal piece (Fig. 3D: bp) and a distal, small, hollow hook (Fig. 3D: h). The entire stylet is 24���31 ��m long (x = 29 ��m; n = 3) and 14���20 ��m wide (x = 16 ��m; n = 3). The hook is 8���15 ��m long (x = 11 ��m; n = 3) and 3���5 ��m wide at its base (x = 4 ��m; n = 3). The broad piece is partially filled by a granular secretion, either originating from the prostate vesicle or part of the prostate vesicle itself. A single ovary (Fig.4B: ov) is situated in the midbody and is surrounded by the gut., Published as part of GOBERT, STEFAN, ARMONIES, WERNER, DIEZ, YANDER L., JOUK, PHILIPPE, MONNENS, MARLIES, REVIS, NATHALIE, REYGEL, PATRICK, SMITH, JULIAN III, STEENKISTE, NIELS VAN & ARTOIS, TOM, 2022, Orostylis gen. nov., a new genus of Dalytyphloplanida with seven new species (Platyhelminthes: Rhabdocoela), pp. 29-46 in Zootaxa 5115 (1) on page 34, DOI: 10.11646/zootaxa.5115.1.2, http://zenodo.org/record/6346911

Krumbachia subterranea Reisinger, 1933 Fig. 8A–C Material examined Neotype GERMANY • 1 spec., studied alive and sagittally sectioned; Hessen; 50°46′30″ N, 09°30′18″ E; 9 Aug. 2011; A.M. Houben and W. Proesmans leg.; submersed meadow at the banks of the Breitenbach creek; neotype no. 823; HU. Reference material GERMANY • 1 spec., live observations; same collection data as for neotype. Description The examined specimens are around 2.5 mm long. The body is slender with a somewhat pointy anterior end and rounded posteriorly (see Fig. 8A). Adenal rhabdite glands occur just behind the brain (Fig. 8A: ar). Paired protonephridiopores (pp) are positioned ventrally, very near to the mouth (Fig. 8A: pp). A rosulate pharynx (Fig. 8A: ph) is situated just behind the centre of the body. The gonopore (Fig. 8A–C: gp) is located at ±75% of the body and connected to the gonoduct (Fig. 8B– C: gd). The gonoduct is surrounded by an inner circular and outer longitudinal muscle layer, and lined with a ciliated, nuclear epithelium resembling the epidermis. The genital atrium (Fig. 8A–C: ga) is surrounded by a similar musculature and lined with a high, nucleated epithelium. On both lateral sides of the genital atrium, a group of proliferating cells (Fig. 8C: pc) is found. A large group of eosinophilic glands is situated posterior to the genital atrium and probably enters this atrium anteriorly (not drawn on the reconstruction). The large, egg-shaped testes (Fig. 8A: t) lie anterior to the pharynx and ventral to the vitellaria (Fig. 8A– B: vi). Paired vasa deferentia unite to form a single vas deferens (Fig. 8A–C: vde) just before entering the copulatory bulbus (Fig. 8A: co). Two layers of spiral muscles surround the long, curved copulatory organ, which bends up to 80° towards its distal end. The copulatory organ bears an intracapsular seminal vesicle (Fig. 8B–C: vs) and a strongly sclerotised ejaculatory duct (Fig. 8B–C: de). Several coarsegrained, extracapsular eosinophilic glands (Fig. 8A–C: gg) enter the copulatory organ at the proximal end. Their secretion is lightly basophilic when entering the copulatory organ, however, it becomes strongly eosinophilic in the ejaculatory duct. A cone-shaped bursa (Fig. 8A–C: bu), surrounded by strong inner circular and outer longitudinal muscles, lies next to the copulatory organ and opens into the genital atrium. The vitellaria extend from the region of the rhabdite glands to the posterior end (Fig. 8A: vi). The vitelloducts (Fig. 8C: vd) fuse just before opening into the female duct (Fig. 8C: fd). Furthermore, this duct is surrounded by a layer of circular muscles and receives the oviduct, a seminal receptacle (Fig. 8A–C: rs), and the female glands (Fig. 8C: fg). The seminal receptacle is provided with a short stalk, which also is surrounded by circular muscles. Discussion See the general discussion on the genus Krumbachia Reisinger, 1924. Previously known distribution Ruhr, Germany (Reisinger 1933), riparian forest near Schlitz, Germany (Schwank 1981). Common species in Europe (Lanfranchi & Papi 1978)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 90-92, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1933. Neues zur vitalen Nervenfarbung. (Gleichzeitig ein Beitrag zur Kenntnis des Protoplanelliden-Nervensystems). Verhandlungen der deutschen Zoologischen Gesellschaft: 155 - 160.","Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Kepner W. A. & Carter J. S. 1931. Olisthanella virginiana nov. sp. Zoologischer Anzeiger 95: 87 - 94.","Ruebush T. K. 1938. Krumbachia minuta n. sp. (Turbellaria Rhabdocoela). Zoologischer Anzeiger 122: 260 - 265.","Schwank P. 1981. Seltene und wenig bekannte Strudelwurmer (Turbellaria) aus Quellen und Bergbachen Mitteleuropas. Beitrage zur Naturkunde in Osthessen 17: 101 - 131.","Lanfranchi A. & Papi F. 1978. Turbellaria (excl. Tricladida). In: Illies J. (ed.) Limnofauna Europaea: 5 - 18. G. Fischer, Stuttgart."]}

Genus Faunulus Houben, Proesmans & Artois gen. nov. urn:lsid:zoobank.org:act: 18E72BA0-D92B-4504-8E17-BD6497247BCA Type species Faunulus nielsi Houben, Proesmans & Artois gen. et sp. nov. Diagnosis ‘Typhloplanidae’ with adenal rhabdites in two anterior tracts, dermal rhabdites could be present distally. Eyes absent. Pharynx slanted obliquely forward and in the central body part. With rod-shaped glands at rostral and caudal body end. Testes differ in size and lie anterior to the pharynx. Copulatory organ with straight, armed cirrus. Seminal receptacle inside of the oviduct. Bursa copulatrix absent. Etymology The genus name refers to the habitat where this taxon was found and is derived from the Latin ‘Faunulus’ (m.), meaning ‘forest dweller’., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on page 75, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040

Hoplopera opaca Reisinger, 1924 Fig. 7D–F Material examined AUSTRIA • 1 spec., studied alive and sagittally sectioned; Hohe Rannach near Graz; 47°09′53″ N, 15°24′50″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; leafy humus of a birch forest; XIV.2.42; HU. Description Our specimen is about 1 mm long and very dark. The anterior end is rounded, with the posterior end showing a distinct tail with pronounced eosinophilic tail glands (Fig. 7D: tg). Small dermal rhabdites occur all over the body. Adenal rhabdites are rather inconspicuous and only visible on the sectioned specimen. Paired protonephridiopores (Fig. 7D: pp) open posterior to the mouth. A rosulate pharynx (Fig. 7D: ph) is positioned at ±75% of the body. The gonopore (Fig. 7E–F: gp) is situated at ±80% of the body and connected to a genital atrium (Fig. 7D– F: ga), which is surrounded by muscles and lined with a high epithelium. The vasa deferentia (Fig. 7F: vde) enter the copulatory organ (Fig. 7D–F: co) separately. This copulatory organ is a 20 µm long, oval structure surrounded by muscles of uncertain orientation. It contains sperm in its proximal part, which probably serves as a seminal vesicle. An ejaculatory duct was not observed. Intracapsular prostate glands (Fig. 7E: gg) were observed on live specimens. The male duct is surrounded by circular muscles. The bipartite bursa (Fig. 7D–F: bu) bears a thin-walled, proximal part containing sperm and a long distal part, the bursal stalk (Fig. 7E–F: bs). This stalk is surrounded by circular muscles and contains a sclerotised structure (Fig. 7E: ss). It seems to consist of several vertical bars, which are fused in the distal part. The vitellaria were still developing in the studied specimen, but already reach up to ±33% of the body. The female duct (Fig. 7F: fd) connects the oviduct (Fig. 7F: od) and vitelloduct (Fig. 7F: vd) to the genital atrium. Discussion See the general discussion on the genus Hoplopera. Previously known distribution Near Graz, Austria, in moist forest soils (Reisinger 1924). General discussion on Hoplopera All species of Hoplopera are characterised by a combination of features: pharynx situated in the caudal body half, presence of dermal rhabdites and adenal rhabdites, the latter organised in rostral tracks (i.e., Stäbchenstassen), presence of tail glands, lack of a stylet, and the presence of a bursa copulatrix (see Van Steenkiste et al. 2010). This combination of features is also shown by Rhomboplanilla bryophila Schwank, 1980. However, species of Hoplopera all have a sclerotised structure in the bursa copulatrix, which is absent in R. bryophila. Species of Hoplopera either occur in limnoterrestrial environments (H. isis Houben, Proesmans & Artois sp. nov.; H. macropharynx Reisinger, 1924; H. maculata Reisinger, 1924; and H. opaca), or in marine to brackish habitats (H. littoralis Karling, 1957 and H. pusilla Ehlers, 1974). However, the marine origin of H. littoralis and H. pusilla has been questioned. Based on its sampling locality, Karling 1957 and Ax 2008 suggested that H. littoralis might as well be a limnoterrestrial species. H. pusilla was found in the supratidal of salty meadows with salinity below 10‰ and was therefore deemed brackish instead of limnoterrestrial (Armonies 1987). All limnoterrestrial species possess a bipartite bursa, with the distal part (probably the bursal stalk) containing a sclerotised structure, while the bursa of the ‘marine’ animals consists of only one part. Furthermore, the vasa deferentia of the limnoterrestrial species enter the copulatory organ separately (Reisinger 1924), while they fuse just before entering the copulatory organ in H. littoralis and H. pusilla (Karling 1957; Ehlers 1974). Both the ecological and morphological differences suggest that Hoplopera may as well be split into two genera. Awaiting a thorough cladistic analysis, however, we prefer to maintain the taxon Hoplopera containing both limnoterrestrial and ‘marine’ animals. The limnoterrestrial species of Hoplopera are most easily distinguished from each other by the shape of the sclerotised structure in the bursa. The fish pot-shaped sclerotic structure as seen in Hoplopera isis Houben, Proesmans & Artois sp. nov. is unique within the genus. In the other three limnoterrestrial species, this structure consists of 6–8 bars proximally connected to a ring (H. macropharynx), 4 (rarely 5) elongated, spoon-shaped structures accompanied with two longitudinal plates (H. maculata) or 6–10 longitudinal bars with broadened distal ends and small protrusions towards the genital atrium (H. opaca) (Reisinger 1924). None of these species possesses the three horizontal bars present in H. isis Houben, Proesmans & Artois sp. nov. Apart from the above-mentioned differences, H. isis Houben, Proesmans & Artois sp. nov. can also be distinguished from H. opaca and H. macropharynx by the presence of a seminal receptacle, a feature it shares with H. maculata. H. opaca has pronounced tail glands, while the tail glands of H. isis Houben, Proesmans & Artois sp. nov. are small and inconspicuous. H. macropharynx has a conspicuous, large, and oval pharynx, which is unique among the species discussed. H. maculata is the largest of all limnoterrestrial species of Hoplopera (1.5 mm vs. at most 1 mm for the other three species) and has typical parenchymal refracting spots (for details, see Reisinger 1924)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 88-89, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Van Steenkiste N., Davison P. & Artois T. 2010. Bryoplana xerophila n. g. n. sp., a new limnoterrestrial microturbellarian (Platyhelminthes, Typhloplanidae, Protoplanellinae) from epilithic mosses, with notes on its ecology. Zoological Science 27 (3): 285 - 291. https: // doi. org / 10.2108 / zsj. 27.285","Karling T. G. 1957. Drei neue Turbellaria Neorhabdocoela aus dem Grundwasser der schwedischen Ostseekuste. Kungliga fysiografiska sallskapets i Lund forhandlingar 27: 25 - 33.","Ehlers U. 1974. Interstitielle Typhloplanoida (Turbellaria) aus dem Litoral der Nordseeinsel Sylt. Mikrofauna des Meeresbodens 49: 427 - 526.","Ax P. 2008. Plathelminthes aus Brackgewassern der Nordhalbkugel. Akademie der Wissenschaften und der Literatur. Franz Steiner Verlag, Stuttgart.","Armonies W. 1987. Freilebende Plathelminthen in supralitoralen Salzwiesen der Nordsee: Okologie einer borealen Brackwasser-Lebensgemeinschaft. Microfauna Marina 3: 81 - 156."]}

Genus Bryoplana Van Steenkiste, Davison & Artois, 2010 Type species Bryoplana xerophila Van Steenkiste, Davison & Artois, 2010. Remark As Bryoplana belgica Houben, Proesmans & Artois sp. nov. is only the second species to be described within the genus, we here provide a new, emended diagnosis for Bryoplana Van Steenkiste, Davison & Artois, 2010, as well as the type species Bryoplana xerophila Van Steenkiste, Davison & Artois, 2010. For further details, please refer to the discussion on B. belgica Houben, Proesmans & Artois sp. nov. below. Diagnoses Bryoplana Diagnosis amended after Van Steenkiste et al. 2010. Genus of ‘Protoplanellinae’ without dermal or adenal rhabdites. Eyes absent. Pharynx somewhat elongate, in anterior part of body and slanting obliquely forward. Paired testes near the caudal body end and ventrocaudal to the vitellaria. Simple muscular tube as copulatory organ. Bursa copulatrix absent. Bryoplana xerophila Species of Bryoplana with the body completely covered by cilia. Testes small, globular. Copulatory organ without any sclerotisation. Without seminal vesicle., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on page 81, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Van Steenkiste N., Davison P. & Artois T. 2010. Bryoplana xerophila n. g. n. sp., a new limnoterrestrial microturbellarian (Platyhelminthes, Typhloplanidae, Protoplanellinae) from epilithic mosses, with notes on its ecology. Zoological Science 27 (3): 285 - 291. https: // doi. org / 10.2108 / zsj. 27.285"]}

Protoplanella simplex Reisinger, 1924 Fig. 11 Material examined Neotype GERMANY • 1 spec., live observations and sagittal sections; Kordel; 49°49′24″ N, 06°38′06″ E; 24 Jul. 2011; A.M. Houben and W. Proesmans leg.; mosses growing on a wall; neotype no. 825; HU. Other material GERMANY • 1 spec., serially sectioned; same collection data as for neotype; XIV.3.11; HU • 2 specs, live observations, one of which serially sectioned; Bavaria, Oberau; 07°33′33″ N, 11°06′57″ E; 13 Jul. 2011; A.M. Houben and W. Proesmans leg.; forest litter; XIV.3.12; HU • 5 specs, live observations, three of which serially sectioned; Lanaken; 50°56′03″ N; 05°39′36″ E; 27 Jul. 2011; A.M. Houben and W. Proesmans leg.; mosses at the forest edge; XIV.3.13–XIV.3.15; HU • 4 specs, live observations, two of which serially sectioned; Lanaken, National Park ‘ Hoge Kempen’; 50°56′02″ N; 05°39′38″ E; 27 Jul. 2011; A.M. Houben and W. Proesmans leg.; moist forest soil; XIV.3.16–XIV.3.17; HU. Description The studied specimens are about 0.8 mm long. The anterior and posterior body ends are rounded (Fig. 11A). Rhabdite glands are present and arranged into two groups behind the brain (Fig. 11A: ar); the rhabdites themselves extend forward in two anastomosing tracts. Dermal rhabdites were not observed. The paired protonephridiopores (Fig. 11A: pp) open laterally and somewhat caudal to the rosulate pharynx (Fig. 11A, C: ph), which is situated just behind the middle of the body. The gonopore (Fig. 11A–C: gp) is located at ±80% of the body and connected to a genital atrium (Fig. 11A–C: ga). The genital atrium is surrounded by an inner circular and outer longitudinal muscle layer. The small, round testes (Fig. 11A–B: t) lie posterior to the pharynx and ventral to the vitellaria (Fig. 11A–C: vi). Both vasa deferentia (Fig. 11B–C: vde) enter the copulatory organ (Fig. 11A: co) separately from the lateral side. This 22 µm long, oval-shaped copulatory organ is surrounded by two layers of spiral muscles and contains an intracapsular seminal vesicle (Fig. 11B–C: vs) with a low, nucleated epithelium and an ejaculatory duct (Fig. 11B–C: de), which is surrounded by two layers of spiral muscles. Extracapsular, eosinophilic prostate glands (Fig. 11A–C: gg) open laterally into the copulatory organ. A male duct (Fig. 11C: md) with the same musculature as the genital atrium connects the copulatory organ to the latter. The paired vitellaria extend from 25% of the body to the posterior end where they fuse via a broad anastomosis. A single ovary (Fig. 11A–C: ov) is closely associated with one of the vitellaria forming an ovovitellarium. The female duct (Fig. 11C: fd) is surrounded by the same musculature as the genital atrium and receives a club-shaped seminal receptacle (Fig. 11A–C: rs) and ovovitelloduct at its proximal end. Fine-grained, eosinophilic glands (Fig. 11C: fg) surround and enter the female duct. Discussion See the general discussion on the genus Protoplanella. Remarks Intensive surveys at the type locality yielded no specimens. The specimens from Oberau, Bavaria, Germany, which is relatively close to the original type locality in Graz, Austria, are of poor quality. Therefore, a specimen from Kordel, Germany was designated neotype, since all diagnostic features are exactly as described by Reisinger (1924). Previously known distribution In the vicinity of Graz, Austria in forest humus (Reisinger 1924, 1954; An der Lan & Franz 1954; An der Lan 1963), on the Faroe Islands (Steinböck 1931); near Poznań, Poland, in moss and litter (Kolasa 1974). General discussion on Protoplanella As mentioned by Van Steenkiste et al. (2011), the identification of Protoplanella simplex is challenging. New material found during several sampling trips showed a need for type material to unravel the morphological differences between all known descriptions of P. simplex. Reisinger (1924) originally described animals with round testes; vasa deferentia that open separately from the lateral side of the copulatory organ; a vitellarium and ovovitellarium connected to each other over a broad anastomosis; and a seminal receptacle that opens into the proximal part of the female duct. Luther (1963) described a specimen with elongated testes; a bursa that could be a seminal receptacle near the male copulatory organ; and, judging from his illustration, most probably paired ovovitellaria. Also, Van Steenkiste et al. (2011) describe a sack-like protrusion at the genital atrium (which was not mentioned by Reisinger 1924); a female bursa (which was called a seminal receptacle by Reisinger 1924) containing remnants of sperm; and their illustration shows the vasa deferentia uniting while entering the copulatory organ at the proximal side. The descriptions of Luther (1963) and Van Steenkiste et al. (2011) differ substantially from that of Reisinger (1924) and, therefore, we consider them not to refer to representatives of P. simplex. Here, we consider these descriptions as referring to a new species: Protoplanella leiae Houben, Proesmans & Artois sp. nov. Moreover, the description of Luther (1963) indicates some, albeit smaller, differences with that of Van Steenkiste et al. (2011). The illustration made by Luther (1963) is not detailed and only made from live specimens. Since Luther’s drawings are in general more detailed, we assume this illustration was based on sub-ideal circumstances, which yielded a quickly made sketch. For now, we consider the specimens described by Luther (1963) and Van Steenkiste et al. (2011) to be members of the same species since differences are small and, in the case of Luther, not based on thoroughly studied material., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 98-99, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Reisinger E. 1954. Edaphische Kleinturbellarien als bodenkundliche Leitformen. Mitteilungen des Naturwissenschaftlichen Vereines fur Karnten 64: 106 - 123.","An der Lan H. & Franz H. 1954. Turbellaria. In: Franz H. (ed.) Die Nordost-Alpen im Spiegel ihrer Landtierwelt: 129 - 134. Universitatsverlag Wagner, Innsbruck.","An der Lan H. 1963. Zur Verbreitung edaphischer Kleinturbellarien in Osterreich. Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 53: 227 - 234.","Steinbock O. 1931. Freshwater Turbellaria. In: Jensen S., Lundbeck W. & Mortensen Th. (eds) Zoology of the Faroes: 1 - 32. Copenhagen.","Kolasa J. 1974. Badania nad wirkami (Turbellaria) zrodel ze szczegolnym uwzglednieniem okolic Poznania (The investigations on spring Turbellaria with a special regard to surroundings of Poznan). Badania Fizjograficzne nad polska zachodnia 26: 7 - 31.","Van Steenkiste N., Tessens B., Krznaric K. & Artois T. 2011. Dalytyphloplanida (Platyhelminthes: Rhabdocoela) from Andalusia, Spain, with the description of four new species. Zootaxa 2791: 1 - 29. https: // doi. org / 10.11646 / zootaxa. 2791.1.1","Luther A. 1963. Die Turbellarien Ostfennoskandiens IV. Neorhabdocoela 2. Typhloplanoida: Typhloplanidae, Solenopharyngidae und Carcharodopharyngidae. Fauna Fennica 16: 1 - 163."]}

Bryoplana belgica Houben, Proesmans & Artois sp. nov. urn:lsid:zoobank.org:act: 1F08C372-CBAE-45E6-BE50-144752F3D44B Fig. 5 Diagnosis Species of Bryoplana with the ciliation reduced or completely lacking dorsally. Testes elongated, in the posterior body part, lateral to the copulatory organ. Copulatory organ with bent, sclerotised ejaculatory duct. Most proximal part of female duct forming a seminal receptacle. Etymology The epithet refers to the fact that the species was found in Belgium. Material examined Holotype BELGIUM • 1 spec., studied alive and serially sectioned; Koksijde, Oostduinkerke, nature reserve ‘De Zeebermduinen’; 51°08′22″ N, 02°41′26″ E; 30 Jul. 2011; A.M. Houben and W. Proesmans leg.; dry moss growing on dunes; KV.687; FMNH. Description The specimen is about 0.8 mm long. Both body ends are rounded (Fig. 5A–B). Free swimming specimens show a small tail. Adenal rhabdite glands (Fig. 5A: ar) are situated in two groups at ±25% of the body. Dermal rhabdites and protonephridiopores were not observed. The epidermis on the ventral body side is 4.5 µm high and entirely covered with locomotory cilia, while on the dorsal body side it is 3.5 µm high and ciliation is strongly reduced, even almost completely lacking. A slightly forward-slanted rosulate pharynx (Fig. 5A–B: ph) is located just rostral to the centre of the body. The gonopore (Fig. 5C–D: gp) is situated at ±80% of the body and connected to a genital atrium (Fig. 5A, C–D: ga), which is lined with a high, nucleated epithelium and surrounded by muscles, the orientation of which could not be observed with certainty. The elongated testes (Fig. 5A, C–D: t) lie at ±70% of the body and ventrally to the paired vitellaria (Fig. 5A–D: vi). They gradually taper into the broad vasa deferentia (Fig. 5C: vde), which laterally enter the copulatory organ (Fig. 5A: co). Circular muscles surround the 34 µm long copulatory organ, which includes an intracapsular seminal vesicle (Fig. 5C–E: vs) and a 22 µm long, strongly sclerotised ejaculatory duct (Fig. 5C–E: de). This ejaculatory duct is more or less straight at its proximal end and bends strongly (±100°) at the distal end. Coarse-grained eosinophilic glands are situated ventrally to the copulatory organ. Although the entrance into the copulatory organ could not be seen, these glands probably represent the prostate glands (Fig. 5D: gg?). The female duct (Fig. 5C–D: fd) is relatively long and lined with a nuclear epithelium. Proximally, it widens into a seminal receptacle (Fig. 5A, C–D: rs), which receives the short oviduct (Fig. 5D: od). The vitelloduct and female glands (Fig. 5D: fg) open into this female duct at the place where it connects to the seminal receptacle. Discussion The new species can readily be placed within ‘Typhloplanidae’ because it possesses all diagnostic features: a pharynx rosulatus, a single ovary, paired testes, and a single genital opening. Furthermore, the ventral position of the testes relative to the vitellaria indicates this species should be placed in ‘Protoplanellinae’, ‘Rhynchomesostominae’ Bresslau, 1933 or ‘Typhloplaninae’ Graff, 1905. However, the species’ general habitus and internal organisation differ markedly from the situation in the latter two subtaxa, and we therefore designate this species to ‘Protoplanellinae’. Most representatives of ‘Protoplanellinae’ have the pharynx situated in the midbody or posterior body half. Only select species have a pharynx in the anterior half of the body, these belong to Achrochordonoposthia Reisinger, 1924; Bockia Reisinger, 1924; Bryoplana; Microcalyptorhynchus Kepner & Ruebush, 1935, Prorhynchella Ruebush, 1939; and Protopharyngiellona Schwank, 1980. Some of these genera show very typical features: presence of a proboscis (Microcalyptorhynchus), presence of ciliated pits in the anterior body half (Prorhynchella), or the fact that the proboscis is of the doliiformis-type (Bockia). Lack of these features in our species indicates that it cannot be allocated to any of these genera. One of the most eye-catching features of B. belgica Houben, Proesmans & Artois sp. nov. is the lack of a bursa copulatrix, which in the remaining taxa is only the case for Bryoplana. We hence allocate our specimens to the latter genus, to which it indeed shows much resemblance. Bryoplana belgica Houben, Proesmans & Artois sp. nov. differs from Bryoplana xerophila by the presence of a sclerotised, curved ejaculatory duct, the presence of a seminal receptacle in the female system, and the fact that the dorsal body ciliation is very much reduced, or even lacking. The latter feature was also mentioned by Kolasa (1977) for Ventrociliella romanae, which he suggested to be an adaptation to limnoterrestrial habitats. However, in all other typical limnoterrestrial taxa, even in B. xerophila, the dorsal ciliation is present. Both species of Bryoplana were recovered from dried out moss – B. xerophila in the USA and B. belgica Houben, Proesmans & Artois sp. nov. in Western Europe – but indeed differ markedly in the extent of the dorsal body ciliation, undermining Kolasa’s (1977) hypothesis. The presence/absence of a seminal receptacle in the female system as a distinguishing character should be used with care at this moment. Indeed, the seminal receptacle in B. belgica Houben, Proesmans & Artois sp. nov. is simply a swelling of the female duct, and not a separate organ. As such, the absence of this structure in B. xerophila may be due to the specimens of Van Steenkiste et al. (2010) not having mated yet. However, as Van Steenkiste et al. (2010) investigated several live specimens and twelve serial sections, this seems unlikely. Regardless, the differences in body ciliation and construction of the copulatory organ proper clearly distinguish both species.

Ventrociliella romanae Kolasa, 1977 Fig. 3 Material examined AUSTRIA • 1 spec., serially sectioned; near Graz, Hohe Rannach; 47°09′53″ N, 15°24′50″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; leafy humus of a birch forest; XIV.2.34; HU • 1 spec., live observations; Graz, Rohrenberg; 47°07′44″ N, 15°24′53″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; between dead leaves • 2 specs, serially sectioned; Graz, Kanzelkügel; 47°06′49″ N, 15°23′11″ E; 26 Aug. 2011; leafy humus; XIV.2.35–XIV.2.36; HU • 1 spec., live observations; Weyer, Kreuzberg; 47°51′36″ N, 14°39′09″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; humus of a mixed forest • 4 specs, live observations, one of which serially sectioned; Weyer, Kreuzberg; 47°51′29″ N, 14°39′14″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; moss growing on a dead trunk; XIV.2.37; HU. Description and discussion The specimens from Austria are about 0.8mm long and very dark due to their gut content (Fig. 3B). Habitus and internal organs are as described by Kolasa (1977), with the exception that protonephridiopores of one specimen (not drawn) are situated just anterior to the pharynx and not lateral to or slightly behind the pharynx as originally described. However, in most specimens, they are situated at ±50% of the body (pp, Fig. 3A). Due to the small size of the specimens, it was not possible to make a thorough reconstruction of the reproductive structures from the sectioned specimens. Although Ventrociliella romanae was originally placed within ’Protoplanellinae’, it was recently listed as a taxon incertae sedis within Dalytyphloplanida (Van Steenkiste et al. 2010). Awaiting the inclusion of this taxon in molecular phylogenetic analyses, we for now keep this genus as incertae sedis, with probably a high affiliation to ‘Protoplanellinae’: Kolasa (1977) considered V. romanae closely related to the protoplanellin Bockia deses Reisinger, 1924 because of their similar habitus, the position of the male reproductive organ, the construction of the pharynx, and their limnoterrestrial lifestyle. Indeed, the only drawing ever published of B. deses (Reisinger 1954: fig. 10) closely resembles the drawing we constructed of a free-swimming specimen of V. romanae (Fig. 3B). Considering their similarity, and especially since the description of B. deses is based on a single specimen, a re-assessment of the validity of these species is warranted. As we found new material of V. romanae on the type locality of B. deses, we here present a detailed morphological comparison between both species. Kolasa (1977) stated that V. romanae and B. deses differ due to the presence of (1) a real pharynx doliiformis, (2) the lack of adenal rhabdites, (3) the absence of a bursa, (4) the occurrence of ciliation all over the body, (5) the lack of a seminal receptacle, and (6) a sclerotised ejaculatory duct. However, the pharynx of V. romanae is strongly tilted forwards, which makes it possible to describe it as a pharynx doliiformis (1). Furthermore, in some of our specimens of V. romanae, adenal rhabdites are barely visible and may easily be overlooked (2). The bursa originally described by Kolasa (1977) was sometimes seen as a large genital atrium, a characteristic also mentioned in the description of B. deses (Reisinger 1924) (3). Reisinger (1924) never made any remarks on body ciliation. Indeed, this is a rather subtle characteristic and may have easily been missed on a single specimen (4). As the seminal vesicle of V. romanae is merely an accumulation of sperm in the oviduct, it could be easily overlooked if it is not filled. Therefore, the absence of this vesicle in B. deses has little diagnostic value (5). The only clear difference between the two species may be that the ejaculatory duct of B. deses is slightly sclerotised, while it is muscular in V. romanae. However, considering their small size and the absence of sectioned material of B. deses, the reliability of this characteristic is also questionable (6). Finally, the typical, conspicuous, vacuolated epidermis of V. romanae was never mentioned for B. deses. As all presumed differences are questionable to some degree, it is highly likely that V. romanae and B. deses belong to the same genus or perhaps are synonyms. In the latter case, V. romanae would become the junior synonym of B. deses. Resolving this possible synonymy is a challenging task, as specimens identified as either species are reported from the same (type) locality. A more exhaustive resampling effort on this sampling locality may provide more clarity: Microscopical study of a large sample may or may not reveal a consistent difference in the above-mentioned morphological traits, possibly demonstrating their diagnostic value. Sequencing morphologically vouchered representatives of both presumed species and subsequent molecular delineation methods may then provide further evidence for the validity of these taxa. However, with the data available today, Bockia deses is better considered a nomen dubium. Previously known distribution Forest soil near the stream Fosso Contesora in Tuscany, Italy (Kolasa 1977)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 77-79, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Kolasa J. 1977. A new representative of terrestrial Typhloplanoida (Turbellaria) - Ventrociliella romanae gen. nov., sp. nov. Bulletin de l'Academie polonaise des sciences. Serie des sciences biologiques 25: 39 - 43.","Van Steenkiste N., Davison P. & Artois T. 2010. Bryoplana xerophila n. g. n. sp., a new limnoterrestrial microturbellarian (Platyhelminthes, Typhloplanidae, Protoplanellinae) from epilithic mosses, with notes on its ecology. Zoological Science 27 (3): 285 - 291. https: // doi. org / 10.2108 / zsj. 27.285","Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Reisinger E. 1954. Edaphische Kleinturbellarien als bodenkundliche Leitformen. Mitteilungen des Naturwissenschaftlichen Vereines fur Karnten 64: 106 - 123."]}

Faunulus nielsi Houben, Proesmans & Artois gen. et sp. nov. urn:lsid:zoobank.org:act: E8B0D042-259A-427D-8F29-5DB6ED8C8411 Fig. 2 Diagnosis Provisionally the same diagnosis as the genus. Etymology The species epithet is dedicated to Niels Houben, son of the first author (A.M. Houben). Material examined Holotype AUSTRIA • 1 spec., live observations and sagittal sections; Kreuzberg near Weyer; 47°51′36″ N, 14°39′09″ E; 29 Aug. 2011; A.M. Houben and W. Proesmans leg.; forest soil of Pinus, Sambucus, Corylus, and Cornus; KV.686; FMNH. Paratypes AUSTRIA • 2 specs, live observations, one of which whole mounted; same collection data as for holotype; paratype no. 821; HU. Description Specimens about 1 mm long. The body shape is pointed anteriorly and blunt to rounded posteriorly (see Fig. 2A). On free-swimming specimens, a small tail can be observed. Both anterior and posterior ends possess very typical, strongly eosinophilic glands that appear rod-shaped, like big rhabdites, and are situated more or less parallel to each other. Rostrally, rhabdite glands (Fig. 2A: ar) are present and arranged in two groups. The distal part of the epidermis is completely filled with coarse, strongly eosinophilic secretions, which probably are dermal rhabdites (Fig. 2B: dr?). The protonephridiopores were not observed with certainty. The slightly forward slanted rosulate pharynx (Fig. 2A: ph) is located centrally. The gonopore (Fig. 2A–B: gp) is situated at ±65% of the body and connected to a genital atrium (Fig. 2A– B: ga) that is surrounded by muscles, although their orientation could not be observed with certainty. The paired, differently sized testes (Fig. 2A: t) lie in front of the pharynx and ventral to the paired vitellaria (Fig. 2A–B: vi). One testis is relatively large, while the other is barely visible. The paired vasa deferentia (Fig. 2B: vde) enter the copulatory organ laterally (Fig. 2A: co). Two layers of spiral muscles surround the 50 µm long copulatory organ, which bears an intracapsular seminal vesicle (Fig. 2B–C: vs) and a cirrus (Fig. 2B–C: cir). Large, coarse-grained, extracapsular eosinophilic glands (Fig. 2B: gg) are associated with the copulatory organ, although the point of entry remains uncertain. The 30 µm long cirrus is more or less straight and covered with spines in its distal part (see Fig. 2B–F: cir). The most distal part of the cirrus bears the larger spines and is enveloped in a conical, sclerotised pouch (Fig. 2C: sp). The female duct (Fig. 2B: fd) is relatively long and lined with a high, nuclear epithelium. Proximally it widens into a seminal receptacle (Fig. 2B: rs) that receives the oviduct and the vitelloduct. Discussion Faunulus nielsi Houben, Proesmans & Artois gen. et sp. nov. strongly resembles species of Adenocerca Reisinger, 1924 because of the combination of following features: centrally positioned pharynx, presence of tail glands and a spiny cirrus, and absence of a bursa (Van Steenkiste et al. 2010). However, F. nielsi Houben, Proesmans & Artois gen. et sp. nov. differs from all species of Adenocerca in having testes that lie rostral to the pharynx. In all species of Adenocerca, they are situated posterior to the pharynx and lie next to the copulatory organ. Moreover, there is a difference in size between the two testes of F. nielsi Houben, Proesmans & Artois gen. et sp. nov., which was never reported for Adenocerca. This size difference is consistent for all observed specimens. All species of Adenocerca either have no dermal rhabdites (Adenocerca teshirogii Kolasa, 1981) or dermal rhabdites occurring throughout the whole epidermis. Conversely, what we assume are dermal rhabdites occur solely in the caudal body part of F. nielsi Houben, Proesmans & Artois gen. et sp. nov. Because of the unique combination of features present in the new species described, we find it justified to erect a new genus within ‘Typhloplanidae’. Remarks Animals seem to feed on rotifers since trophi were found in the gut., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 75-77, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Van Steenkiste N., Davison P. & Artois T. 2010. Bryoplana xerophila n. g. n. sp., a new limnoterrestrial microturbellarian (Platyhelminthes, Typhloplanidae, Protoplanellinae) from epilithic mosses, with notes on its ecology. Zoological Science 27 (3): 285 - 291. https: // doi. org / 10.2108 / zsj. 27.285"]}

Hoplopera isis Houben, Proesmans & Artois sp. nov. urn:lsid:zoobank.org:act: 4DA3359A-D279-4286-B7B0-822382D258F7 Fig. 7A–C Diagnosis Dark species of Hoplopera, about 0.8 mm long with inconspicuous tail glands. Dermal rhabdites small, adenal rhabdites in two tracks. Atrial glands present. Spindle-shaped testes. Copulatory organ 20 µm long, without sclerotised structures. Bipartite bursa with sclerotised structure containing two groups of parallel, vertical rods, separated from each other by vertical bars, as a whole resembling a fish pot. Female duct forming a seminal receptacle. Etymology The species epithet is dedicated to Isis Houben, first daughter of the first author (A.M. Houben). Material examined Holotype AUSTRIA • 1 spec., studied alive and horizontally sectioned; south of Graz, between Glashütten and Trahütten; 46°49′46″ N, 15°06′09″ E; 23 Aug. 2011; A.M. Houben and W. Proesmans leg.; moss growing in a small stream in a pine forest; KV.688; FMNH. Paratypes AUSTRIA • 5 specs, studied alive, one of which whole mounted; same collection data as for holotype; paratype no. 822; HU. Description Specimens are about 0.7–0.9 mm long. The body is rounded anteriorly and possesses a small tail provided with small, inconspicuous tail glands (Fig. 7C: tg). Rostrally, sharply pointed adenal rhabdites (Fig. 7B: ar) are arranged in two groups. Very small dermal rhabdites occur all over the body. The paired protonephridiopores (Fig. 7A: pp) lie posterior and lateral to the mouth. The rosulate pharynx is located just behind the middle of the body (Fig. 7A: ph). The gonopore (Fig. 7B–C: gp) is situated at ±80% of the body and connected to a genital atrium (Fig. 7A–C: ga) that is lined with a high epithelium and surrounded by muscles of uncertain orientation. At its posterior side, the genital atrium receives a large group of eosinophilic glands (Fig. 7A–C: ag). Two small, spindle-shaped testes (Fig. 7A: t) lie in front of the pharynx (Fig. 7A: ph) and ventral to the paired vitellaria (Fig. 7A, C: vi). In some specimens, one testis seems bigger than the other one. The paired vasa deferentia (Fig. 7B–C: vde) enter the egg-shaped copulatory organ laterally. Two layers of spiral muscles surround the 24µm long copulatory organ (Fig. 7A: co), which bears an intracapsular seminal vesicle (Fig. 7B–C: vs) and an ejaculatory duct (Fig. 7B–C: de). Coarse-grained intra- (Fig. 7B– C: gg1) and extracapsular (Fig. 7C: gg2) eosinophilic glands are associated with the copulatory organ. The relatively long male duct (Fig. 7C: md) receives the bursa (Fig. 7A–C: bu) before entering the genital atrium (Fig. 7A–C: ga). This bipartite bursa consists of an 18 µm long, thin-walled proximal part containing sperm and a 22 µm long distal part, which could be considered the bursal stalk (Fig. 7B–C: bs). This stalk is surrounded by circular muscles and contains a sclerotised structure (Fig. 7B: ss) that resembles the structure of a fish pot. It consists of vertical rods that are divided into two groups by a horizontal bar. The proximal and distal part are also delimited by a horizontal bar. As a whole, this bursal stalk is wrapped in a loose matrix surrounded by circular muscles. The vitellaria (Fig. 7A, C: vi) extend from the anterior third of the body to the posterior end where they meet. The oviduct broadens and becomes a seminal receptacle (Fig. 7A–C: rs), which further becomes a long female duct (Fig. 7C: fd), lined with a high, nucleated epithelium and surrounded by muscles. The vitelloduct was not observed with certainty, although it is probably an unpaired one that meets the oviduct. Discussion See the general discussion on the genus Hoplopera. Remarks The hard parts and copulatory structures are not visible on the whole-mounted specimen. Therefore, we designate a serially sectioned specimen as holotype., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 86-88, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143."]}

Adenocerca minima Kolasa, 1981 Fig. 4 Material examined AUSTRIA • 3 specs, studied alive, one of which whole mounted and a second one horizontally sectioned; south of Graz, between Glashütten and Trahütten; 46°49′46″ N, 15°06′09″ E; 23 Aug. 2011; A.M. Houben and W. Proesmans leg.; humid moss from a small stream in a pine forest; XIV.2.38 and XIV.3.18; HU. Description and discussion Specimens are about 0.6 mm long. Habitus and internal organisation correspond to the description of Kolasa (1981b). However, the original description, based on one specimen, mentions a much smaller animal (0.3 mm) (see Kolasa 1981b). Small dermal rhabdites are present all over the body. Protonephridiopores (Fig. 4A: pp) are situated lateral to the pharynx (Fig. 4A–B: ph). The two small testes (Fig. 4A–B: t) are situated caudally from the pharynx, at 65–70% of the body. Adenocerca minima clearly differs from all other representatives of Adenocerca by the orientation of the spines of its copulatory organ (see Fig. 4C–F). Small spines are present in the distal half of the cirrus and at the distal end, where 6 µm long, distally pointing, crescent-shaped spines form a crown (Fig. 4C: cs). Other species only possess small spines (A. clinopharynx Reisinger, 1924 and A. teshirogii) or have a different orientation of these spines, with the proximal spines being the biggest (A. bresslaui Reisinger, 1924). Remarks Kolasa (1981b) mentioned seven crown spines while we only counted six. Both Kolasa’s and our counts were made on a single specimen. Therefore, for the time being, we prefer to designate our specimens to the same species as Kolasa’s (1981b). Previously known distribution In wet mosses on the banks of the stream Fosso Contesora, Italy (Kolasa 1981b)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 79-80, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Kolasa J. 1981 b. New semiterrestrial species of Protoplanellinae (Turbellaria, Typhloplanidae) associated with a submontane stream. Bulletin de l'Academie polonaise des sciences. Serie des sciences biologiques 28 (10 - 11): 611 - 616.","Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143."]}

Prorhynchella minuta Ruebush, 1939 Fig. 10 Material examined USA • 9 specs, live observations, one of which sagittally sectioned and two horizontally sectioned; Alabama, Ponderosa Lake; 32°54′53″ N, 87°19′28″ W; 27 May 2009; Niels Van Steenkiste leg.; submerged vegetation in shallow, clear water in the southeast corner of the lake; XIV.3.01–XIV.3.03; HU. Description and discussion Specimens are about 1 mm long (Fig. 10E) and can be placed within ‘Protoplanellinae’ due to the ventral position of the testes relative to the vitellaria and the separate protonephridiopores (see Reisinger 1924). Additionally, sectioned specimens clearly show ciliated pits (Fig. 10B: cp) at the anterior body end. Within ʻProtoplanellinae’, the only genus containing species with such pits is Prorhynchella (Van Steenkiste et al. 2010). Moreover, the studied specimens also possess the typical proboscis-like organ (Fig. 10B: plo) associated with a glandular complex (Fig. 10B: gl) that is unique for this monospecific genus (described as ‘rüssel-like organ’ by Ruebush 1939). The only difference between our specimens and the original description of P. minuta is the position of the protonephridiopores (Fig. 10A: pp), which are situated somewhat more posteriorly compared to the original description (Ruebush 1939). The studied specimens show some intraspecific variation in the structure of the bursa (Fig. 10A, C–F: bu). In one specimen, this is merely an evagination of the genital atrium while in another it is an oval, thin-walled structure, filled with sperm (compare Fig. 10C–D, F). It is connected to the genital atrium (Fig. 10A, D: ga) by a long bursal stalk (Fig. 10C, F: bs), which is surrounded by very strong sphincters (Fig. 10C, F: sph). Only two sphincters were visible on live specimens (Fig. 10F), but at least six were counted in one sectioned specimen (Fig. 10C). In the original description of P. minuta, Ruebush (1939) describes a bursa with a stalk, similar to the situation in most of our specimens (Fig. 10C, F). Whether or not the specimen drawn in Fig. 10D is an aberration is not clear: The specimen appears fully adult, hence the poorly developed bursal structure is unlikely a result of a different degree of development. With the data available to us, it is difficult to speculate on the biological significance of this reduced organ and we hence refrain from drawing conclusions on the matter. Previously known distribution Freshwater ponds near Pine Orchard, Connecticut, USA (Ruebush 1939)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on pages 94-96, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Ruebush T. K. 1939. A new North American rhabdocoel turbellarian Prorhynchella minuta n. gen., n. sp. Zoologischer Anzeiger 127: 204 - 209.","Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Van Steenkiste N., Davison P. & Artois T. 2010. Bryoplana xerophila n. g. n. sp., a new limnoterrestrial microturbellarian (Platyhelminthes, Typhloplanidae, Protoplanellinae) from epilithic mosses, with notes on its ecology. Zoological Science 27 (3): 285 - 291. https: // doi. org / 10.2108 / zsj. 27.285"]}

Olisthanellinella rotundula Reisinger, 1924 Fig. 9 Material examined AUSTRIA • 3 specs, studied alive, one of which sagittally sectioned and one of which horizontally sectioned; Graz, Kanzelkügel; 47°06′49″ N, 15°23′11″ E; 26 Aug. 2011; A.M. Houben and W. Proesmans leg.; forest soil; XIV.2.49–XIV.2.50; HU. Description and discussion The body length of the studied specimens is about 0.8 mm. Habitus and internal organisation correspond to the original description (Reisinger 1924). The most typical characteristics of this monospecific genus are: adenal rhabdites (Fig. 9A: ar) occurring only in rostral tracks; dermal rhabdites absent; a bipartite bursa (Fig. 9A–D: bu) with a muscular, sclerotised stalk (Fig. 9B, D: bs) and a thin-walled proximal part; and the copulatory organ (Fig. 9A: co) with a muscular ejaculatory duct (Fig. 9B, D: de) and a large seminal receptacle (Fig. 9C: rs) that opens into the oviduct (Fig. 9C: od). The opening of the vitelloduct was never clearly observed. Neither of our sectioned specimens clearly show both male and female reproductive structures. Therefore, and because of (for now) intraspecific differences (see below), we refrain from designating a neotype. Specimens described by Luther (1963) seem to be somewhat different from those described by Reisinger (1924) since they possess an egg-shaped bursa instead of a bipartite bursa and their oviduct functions as the seminal receptacle. Moreover, the specimens collected by Armonies (1987) were identified based on Luther’s (1963) description.As probably is the case for all protoplanellins, a thorough molecular analysis would help to solve which features are apomorphic/diagnostic for the taxon and help identification and delimitation of species. Therefore, we provisionally keep all within Olisthanellinella rotundula. Previously known distribution In the vicinity of Graz, Austria in moist forest soils (Reisinger 1924); North-East Alps, Austria in moist forest soil (An der Lan & Franz 1954); Oldenburg, Germany in humus of oak leaves (Reisinger in Luther 1963); Helsingfors, Finland in moss (Luther 1963); Sylt, Germany, in salt meadows (Armonies 1987)., Published as part of Houben, Albrecht M., Monnens, Marlies, Proesmans, Willem & Artois, Tom J., 2022, Limnoterrestrial ' Typhloplanidae' (Rhabdocoela, Platyhelminthes), with the description of four new species and a new genus, pp. 70-102 in European Journal of Taxonomy 798 on page 94, DOI: 10.5852/ejt.2022.798.1671, http://zenodo.org/record/6323040, {"references":["Reisinger E. 1924. Die terricolen Rhabdocoelen Steiermarks. Zoologischer Anzeiger 59: 33 - 48, 72 - 86, 128 - 143.","Luther A. 1963. Die Turbellarien Ostfennoskandiens IV. Neorhabdocoela 2. Typhloplanoida: Typhloplanidae, Solenopharyngidae und Carcharodopharyngidae. Fauna Fennica 16: 1 - 163.","Armonies W. 1987. Freilebende Plathelminthen in supralitoralen Salzwiesen der Nordsee: Okologie einer borealen Brackwasser-Lebensgemeinschaft. Microfauna Marina 3: 81 - 156.","An der Lan H. & Franz H. 1954. Turbellaria. In: Franz H. (ed.) Die Nordost-Alpen im Spiegel ihrer Landtierwelt: 129 - 134. Universitatsverlag Wagner, Innsbruck."]}