taxonID	type	description	language	source
03C487EEFFD2547751A9F8E42BF00F9F.taxon	description	Tope genus: Hainesia L. Pfeiffer, 1856. Remarks: Tere is one available family-group name for Madagascan Cyclophoroidea: Hainesiinae Tiele, 1929, which was treated as a synonym of the subfamily Megalomastomatinae (family Megalomastomatidae) by Bouchet et al. (2017), based on the phylogeny of Webster et al. (2012). However, the later publication has not included any species from the Caribbean Islands, where Megalomastoma Swainson, 1840, the type species of Megalomastomatidae lives, but only two species of Acroptochia from Madagascar. In the absence of molecular information, the evolutionary relationship of the Caribbean Megalomastomatidae and the Madagascan Hainesia and Acroptochia is unknown. Nevertheless, it is not likely that they would belong to the same monophyletic group. Terefore, we suggest using Hainesiidae for Hainesiinae (including Hainesia and Acroptochia) and Boucardicinae. Egorov (2009) treated Hainesiidae as a family in its own right and included the genera Acroptochia, Hainesia, and, with doubt, Anosocolus and Boucardicus. Based on the characters of the radula and operculum, he included Hainesiidae provisionally in Litorinoidei (= Litorinoidea Children, 1834); however, molecular phylogenetic data (Webster et al. 2012) later confirmed that Acroptochia (and therefore, Hainesiidae) belongs to the Cyclophoroidea. Tere are two additional high-spired operculate genera from Mauritius that might be of relevance to the Madagascan genera in question: Madgeaconcha O. L. Griffiths & Florens, 2004 and Naggsiaconcha O. L. Griffiths & Florens, 2004, but they are much smaller than Acroptochia and Hainesia, and their systematic relationships are highly questionable (Griffiths and Florens 2004). Madgeaconcha, with its minute shell being only slightly larger than 1 mm, is similar to Acmella W. T Blandford, 1869 in size and shape (see Vermeulen et al. 2015, Das et al. 2021), and therefore, it probably belongs to the family Assimineidae. Although Naggsiaconcha is much higher spired than any other terrestrial assimineids, it might also belong to the same family. Te genera Ankoravaratra Emberton, 2002 (described by Emberton 2002 c; type species: Ankoravaratra ambrensis Emberton, 2002, by original designation) and Owengriffithsius Emberton, 2002 (described by Emberton 2002 b; type species: Owengriffithsius capdambrae Emberton, 2002, by original designation), also endemic to Madagascar, are currently classified in the families Maizaniidae and Cyclophoridae, respectively. Nevertheless, their systematic position is highly questionable, and it is possible that they would also belong to the Hainesiidae.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFDF547352CBF9B92A920913.taxon	description	(Fig. 10) h t t p: / / z o o b a n k. o r g / 2 6 9 6 4 1 F 2 - F C 2 5 - 4 1 A E - B C 9 6 - 0632 E 6931 CC 3 Tope genus: Boucardicus Fischer-Piete & Bedoucha, 1965. Content: Anosocolus Fischer-Piete, Ch. P. Blanc, F. Blanc & F. Salvat, 1993 (Fig. 10 A), Boucardicus Fischer-Piete & Bedoucha, 1965 (Fig. 10 B – D), Malarinia Haas, 1961 (Fig. 10 E, F). As a result of a recent intense survey, the number of known Boucardicus species has reached almost 200 species (Emberton and Pearce 1999, Emberton 2002 a, Emberton et al. 2010, Balashov and Griffiths 2015). Anosocolus and Malarinia both have two species. Diagnosis: Sutural tube absent; microtunnels long, formed by folds of the uppermost shell layer, and situated between the peristome and the position of the operculum (= constriction), i. e. anterior to the inner opening; each microtunnel reaches the common opening individually; the single opening is situated immediately behind the operculum. Operculum eccentric. Distribution: Endemic to Madagascar (Fig. 11). Remarks: Te shell sculpture of Boucardicus species, especially the rib morphology of the area between the peristome and the position of the operculum, is crucial for species delimitation and recognition. Tus, it was described in detail for each species (Emberton 1994, 2001, 2002 a, Emberton and Pearce 1999, Emberton et al. 1999, 2010). However, the function of these ribs (i. e. microtunnels) and the presence of a single opening behind the operculum have not been mentioned in the literature.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352E8FEC02ACD0C5D.taxon	description	(Fig. 10 A)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352E8FEC02ACD0C5D.taxon	type_taxon	Tope species: Anosocolus anosocolus Fischer-Piete, Ch. P. Blanc, F. Blanc & F. Salvat, 1993 (by original designation).	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352E8FEC02ACD0C5D.taxon	discussion	Remarks: Anosocolus is included in the Boucardicinae, owing to the presence of microtunnels on the last whorl of the type species. Boucardicus, in the present sense, is an extremely variable genus in terms of shell size, shell and aperture shape, and sculpture, and Anosocolus differs from some large, triangular Boucardicus species (e. g. Fig. 10 B) only in minor characters, such as the keeled body whorl and closed umbilicus. Tis might suggest that Anosocolus and large-shelled Boucardicus are more closely related to each other than are large, triangular-shelled and small, ovoid Boucardicus species (e. g. Fig. 10 D). Clearly, a genus-level revision of Boucardicus is needed in order to provide a phylogenetically meaningful subdivision of this mega-genus.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352CEFB8E28FF0D83.taxon	description	(Fig. 10 B – D)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352CEFB8E28FF0D83.taxon	description	Tope species: Acroptochia notabilis E. A. Smith, 1892 (Fig. 10 B) by original designation.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547352CEFB8E28FF0D83.taxon	discussion	Remarks: Madecataulus Fischer-Piete & Bedoucha, 1965 (type species: Madecataulus goudoti Fischer-Piete & Bedoucha, 1965, by monotypy; Fig. 10 C) is a synonym of Boucardicus according to Emberton and Pearce (1999). See also under Anosocolus and Malarinia.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547351DFFCE72B9C0C99.taxon	description	(Fig. 10 G) Content: Acroptochia Crosse & P. Fischer, 1877 [replacement name for Euptochia Crosse & P. Fischer, 1877, non Hübner, 1816 (Lepidoptera); type species: Euptochia metableta Crosse & P. Fischer, 1873 (Fig. 10 G), by typification of replaced name], Hainesia L. Pfeiffer, 1856 (type species: Coclostoma croceum G. B. Sowerby I, 1843, by subsequent designation; Wenz, 1938: 469). Remarks: Differs from Boucardicinae by the absence of microtunnels.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD85473534CFA512CC30B4D.taxon	description	(Fig. 10 E, F)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD85473534CFA512CC30B4D.taxon	description	Tope species: Malarinia hova Haas, 1961, by original designation. Included species: Malarinia hova Haas, 1961 (Fig. 10 E) and Malarinia calcoperculata Emberton, 1994 (Fig. 10 F, see also Emberton & Pearce, 1999: 364, fig. 22).	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD85473534CFA512CC30B4D.taxon	discussion	Remarks: We examined photographs of the type species of Malarinia (Madagascar, ~ 30 miles south of Moramanga, Chutes de la Mort, leg. E. S. Ross, 10 November 1959, FMNH 106701). Te strong sculpture on the last half whorl suggests that each rib represents a microtunnel, as in the genera Boucardicus and Anosocolus. So far, this genus has been placed in the family Diplommatinidae, which has no other known members in Madagascar or in the entire African continent (Egorov 2013). Tus, its position in the Boucardicinae is more likely, and it is here moved to Boucardicinae. Boucardicus menoi K. C. Emberton, Slapcinsky, C. A. Campbell, Rakotondrazafy, Andriamiarison & J. D. Emberton, 2010 and Boucardicus avo K. C. Emberton, Slapcinsky, C. A. Campbell, Rakotondrazafy, Andriamiarison & J. D. Emberton, 2010 are very similar to two known Malarinia species in shell shape and sculpture (Emberton et al. 2010). It is possible that they are wrongly assigned to Boucardicus and should instead belong to Malarinia. However, it might also mean that there is no clear morphological distinction between Malarinia and Boucardicus. Given that Malarinia is the older name, it has priority over Boucardicus, and the ~ 200 species of the later genus should be transferred to Malarinia. However, we do not formally synonymize Boucardicus with Malarinia here. Instead, a systematic revision of the extremely variable Boucardicus should propose a reasonable subdivision of that genus.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547151FFFA952CCD0E00.taxon	description	(Fig. 12)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFD8547151FFFA952CCD0E00.taxon	description	Tope genus: Alocaeus Baird, 1850. Content: Alocaeus Baird, 1850 (type species: Alocaeus eodouxi Venmans, 1956; Fig. 12 A), Chamalocaeus Möllendorff, 1897 (type species: Alocaeus (Chamalocaeus) fuhstorferi Möllendorff, 1897), Coclorox Godwin-Austen, 1914 (type species: Coclostoma constrictum Benson, 1851), Dicharax Kobelt & Möllendorff, 1900 (type species: Alocaeus hebes Benson, 1857; Fig. 12 D), Diorox Benson, 1859 (type species: Alocaeus amphora Benson, 1856; Fig. 12 C), Metalocaeus Pilsbry, 1900 (type species: Alocaeus (Metalocaeus) melanopoma Pilsbry, 1900, Fig. 12 E), Pincerna Preston, 1907 (type species: Pincerna liratula Preston, 1907), and Stomacosmethis Bollinger, 1918 (type species: Alocaeus (Stomacosmethis) sarasinorum Bollinger, 1918). Currently contains 349 extant species (Páll-Gergely et al. 2020, 2021, Jirapatrasilp et al. 2021, MolluscaBase 2024). Diagnosis: External tube present (visible along the suture externally), microtunnels few (four or five) to many (> 50), long, running from umbilicus to sutural tube; tube and microtunnels rise only in subadults (do not extend beyond the last half whorl), entirely absent in juveniles. Distribution: Western India through the Himalaya to Japan in the east, Korea in the north, and Indonesia to the south (van Benthem Juting 1948, 1959, Azuma 1982, Minato 1988, Páll-Gergely et al. 2020, 2021, Jirapatrasilp et al. 2021) (Fig. 11). Remarks: We do not list below the included genera, because the genus-level classification of the Alycaeinae was published recently (Páll-Gergely et al. 2020). Te only change at the genus level compared with the paper by Páll-Gergely et al. (2020) is that Coclorox has been treated as a synonym of Pincerna following Páll-Gergely (2017), while it is currently accepted as a valid genus (see Gitenberger et al. 2022, Páll-Gergely et al. 2023). In fact, Gitenberger et al. (2022) referred to the unpublished molecular phylogeny published in the present study. However, when we gave information to Edmund Gitenberger, we considered Stomacosmethis balingensis (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) (Tomlin, 1948) a Pincerna species based on shell shape. However, in the present molecular phylogeny no true Pincerna are represented; therefore, we cannot confirm that Coclorox and Pincerna and phylogenetically distinct. Nevertheless, it seems likely that while Coclorox is closely related to Alocaeus and Diorox, Pincerna is a relative of Stomacosmethis. Tus, we maintain Coclorox as a valid genus, but this question has to be addressed by future molecular studies.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFDA546C512CF9D1291F0A43.taxon	description	(Fig. 13 A, B) h t t p: / / z o o b a n k. o r g / E B B 6 5 4 2 E - 6 A 0 0 - 4 1 6 0 - 9 9 E D - 5 B 4373 E 918 D 8 Tope genus: Platorhaphe Möllendorff, 1890. Content: Platorhaphe Möllendorff, 1890 (Fig. 13 A, B), currently contains 76 species (MolluscaBase 2024). Diagnosis: Inner tube present (not visible along the suture externally), situated above the suture and with a single opening, microtunnels short (start near the suture). Distribution: South-eastern China, Taiwan, Okinawa (Japan), the Philippines, Malaysian part of Borneo, Sulawesi, Maluku and North Maluku, Indochina Peninsula (Peninsular Malaysia, Tailand, Laos, Vietnam) (Fig. 11). Remarks: Te genitive of rhaphe is rhaphidis, hence the stem of the genus Platorhaphe is ‘ Platyrhaphid- ’. Te family-group name will thus be Platyrhaphidinae (Philippe Bouchet, pers. comm., September 2019).	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546C519DFEFB2D2A0EB2.taxon	description	(Fig. 13 C)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546C519DFEFB2D2A0EB2.taxon	description	Tope species: Laotia pahiensis Saurin, 1953 (by monotypy). Content: Tree species: Laotia christahemmenae Páll-Gergely, 2014 (Páll-Gergely 2014), Laotia luongi Páll-Gergely & Hunyadi, 2021, and Laotia pahiensis Saurin, 1953 (Saurin 1953; also Fig. 13 C). Distribution: Northern Vietnam and northern Laos. Remarks: Te molecular phylogenetic analysis (Fig. 9) revealed that Laotia forms a strongly supported clade with many species and genera of Diplommatinidae and represents a separate lineage within that clade with an unresolved basal node. Owing to the uncertainty regarding the phylogenetic relationships between Laotia and (other) Diplommatinidae, we erect a new subfamily for Laotia and Messageria within Diplommatinidae, instead of erecting a new family in its own right. Even if the Diplommatinidae comprise two strongly supported subclades (Webster et al. 2012, Köhler 2023; Fig. 9), we lack molecular information on the majority of diplommatinid genera (currently 35 genera, of which 8 are fossil-only; MolluscaBase 2024), and predicting the systematic position of each genus would be very difficult based on morphological characters alone. Terefore, we leave all other Diplommatinidae without classifying them into subfamilies. Te shell shape of Laotia is strikingly similar to that of the extinct (Miocene – Cretaceous) European genus Ferussina Grateloup, 1827 (type species: Ferussina anostomaeformis Grateloup, 1827, by monotypy) (Supporting Information, Fig. S 6 E), which is currently classified in its own subfamily in the Cyclophoridae (Páll-Gergely and Neubauer 2020, Páll-Gergely et al. 2023) [previously in its own family, Ferussinidae Wenz, 1923 (1923 – 1930); see Bouchet et al. 2017]. Te similarity could result either from relatedness or from parallel / convergent evolution. Te former hypothesis can be tested by searching for inner tubes in Ferussina similar to the ones found in Laotia shells. We examined a sample containing ~ 230 shells (O-Oligozän, Landschneckenkalk, Hochheim-Flörsheim, coll. Kinkelin) in the SMF. Two shells had a broken last whorl, which presented the opportunity to investigate the presence or absence of inner gas-exchange chambers. Given that no such chambers were found, we consider the similarity between Laotia and Ferussina to be a result of parallel / convergent evolution.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546C52A9F9B12C6A0969.taxon	description	(Fig. 13 C – F) h t t p: / / z o o b a n k. o r g / 3 D 2 6 7 C 2 2 - E B D 9 - 4 5 B 5 - A 7 5 1 - B 5 BA 463 EC 511 Tope genus: Laotia Saurin, 1953. Content: Laotia Saurin, 1953 (Fig. 13 C) and Messageria Bavay & Dautzenberg, 1904 (Fig. 13 D – F), containing three and two species, respectively. Diagnosis: Inner tube missing, but a series of distinct inner sutural chambers situated below the suture; as the shell grows, old chambers are closed and new ones open, four or five open simultaneously; microtunnels long (start in umbilical area), two or three connected to a single sutural chamber. Distribution: Northern Laos, Northern Vietnam, Southern China (Guangxi) (Fig. 11).	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546C5292FD9129E10DE0.taxon	type_taxon	Tope species: Coclotus pusillus G. B. Sowerby I, 1843 (Fig. 6 B), by original designation.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546C5292FD9129E10DE0.taxon	discussion	Remarks: No recent revision has been published. Tirty-three of the 76 species were described from the Philippines by Bartsch (1919), who is famous for being an extreme spliter (see Waters 2006, and Páll-Gergely et al. 2019 and references therein). Terefore, the real number of species might be lower, although several undescribed species await description from continental Asia. Bartsch (1919) described three subgenera (Platoraphella, Platoraphida, and Platoraphina) based on subtle differences of the operculum, all from the Philippines. Although we have not examined this question in detail, it seems unlikely that four subgenera are known from the edge of the distribution of this genus, whereas the nominotypical subgenus is widely distributed in Southeast Asia. Te subgenera of Platorhaphe are likely to be results of Bartsch’s extreme spliting activity. Tus, in agreement with Egorov (2009), we do not consider them valid, and we handle Platorhaphe as a genus without subgenera. Although Bartsch (1919) clearly indicated the presence of the inner tube of this genus as a characteristic feature, it has not received any atention by later authors. Tus, owing to the simple cyclophoroid shell shape, some species currently classified as Platorhaphe might belong to Coclotus or other similar-looking genera, and vice versa. Tis question will be investigated further in upcoming systematic studies.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546D51E6F8A329030D8F.taxon	description	(Fig. 13 D – F)	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546D51E6F8A329030D8F.taxon	description	Tope species: Helicomorpha (Messageria) scalarioides Bavay & Dautzenberg, 1904, by original designation. Content: Messageria scalarioides (Bavay & Dautzenberg, 1904) (Fig. 13 D – F) and Messageria sinica Z. - Y. Chen & L. - W. Lin, 2021.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546D51E6F8A329030D8F.taxon	distribution	Distribution: Northern Vietnam and Guangxi (China).	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
03C487EEFFC7546D51E6F8A329030D8F.taxon	discussion	Remarks: Two syntypes (an adult and a subadult one) of Helicomorpha scalarioides were examined. One of them is an adult shell, which is hereby selected to be the lectotype (MNHN-IM- 2000 - 32414; fig. 1 in Chen and Lin 2021). In oblique apertural view, several openings on the inner tube are visible, resembling those described in Laotia. In the subadult paralectotype (MNHN-IM- 2000 - 32415; see Fig. 13 E, F), the openings start immediately before the peristome. In the adult lectotype, the openings start more distantly from the peristome. Messageria was described as a subgenus of the diplommatinid Helicomorpha Möllendorff, 1890 (type species: Helicomorpha turricula Möllendorff, 1890 by original designation). Currently, nine species of Helicomorpha inhabit the Philippines (Kobelt 1902). Chen and Lin (2021) elevated Messageria to the genus level and transferred it to the Alycaeidae owing to features of the shell and operculum shared with Laotia. We received photographs of the lectotype of Helicomorpha turricula (Supporting Information, Fig. S 6 A) and borrowed four species from the MNCN (Helicomorpha pilula Quadras & Möllendorff, 1896: Sierra Bullones, isla de Bohol, Filipinas, coll. Azpeitia 2760, MNCN 28808 / 2, see Supporting Information, Fig. S 6 B – D; Helicomorpha quadrasi Möllendorff, 1893: Malitbog, isla de Leyte, Filipinas, coll. Azpeitia, 2761, MNCN 28809 / 2; Helicomorpha turricula, Filipinas, Azpeitia, 2762, MNCN 28812 / 2; and Helicomorpha depressa Möllendorff, 1893: Cantanduanes, isla Cantanduanes, Filipinas, coll. Azpeitia, 2798, MNCN 28801 / 2). Te borrowed shells (Supporting Information, Fig. S 6 B – D) showed no signs of an inner tube or separate sutural chambers, which are visible in Messageria scalarioides and Laotia cf. christahemmenae. Terefore, Helicomorpha is confirmed to belong to the family Diplommatinidae, whereas it does not belong to the subfamily Laotiinae proposed for Laotia and Messageria, both possessing sutural chambers.	en	Páll-Gergely, Barna, Ruthensteiner, Bernhard, Harl, Josef, Magonyi, Nóra M., Asami, Takahiro, Krizsik, Virág, Schwaha, Tomas, Fehér, Zoltán (2024): Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-25, DOI: 10.1093/zoolinnean/zlae158, URL: https://doi.org/10.1093/zoolinnean/zlae158
