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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