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Páll-Gergely, Barna, White, Tom S. (2023): Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae). Journal of Natural History 56 (45-48): 2011-2029, DOI: 10.1080/00222933.2022.2152750, URL: http://dx.doi.org/10.1080/00222933.2022.2152750

Hypselostomatidae Zilch, 1959 Remarks Bouchet et al. (2017) treated Hypselostomatinae as a subfamily of Gastrocoptidae Pilsbry, 1918. The systematic relationships of the genera classified in this (sub)family are largely unclear. Here, to be consistent with our previous papers (i.e. Páll-Gergely et al. 2015, 2020, 2022), we use Hypselostomatidae at the family level. According to Pilsbry (1917), Boysidia Ancey, 1881 possesses a concrescent angular and parietal lamella, whereas they are separate in Bensonella and Paraboysidia. This systematics has been widely used since (i.e. Schileyko 1998; Panha and Burch 2005). However, it was already noted by Pilsbry (1917) that there are species showing intermediate character states. For example, Boysidia strophostoma (Möllendorff, 1885) possesses clearly separated angular and parietal lamellae, but is still classified in Boysidia. Solving the homologies of apertural barriers and the systematics of the related hypselostomatid genera is beyond the scope of the present paper, but should be done as soon as possible., Published as part of Páll-Gergely, Barna & White, Tom S., 2023, Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae), pp. 2011-2029 in Journal of Natural History 56 (45 - 48) on page 2014, DOI: 10.1080/00222933.2022.2152750, http://zenodo.org/record/7560931, {"references":["Zilch A. 1959 - 1960. Gastropoda. Teil 2. Euthyneura. In: Schindewolf OH, editor. Handbuch der Palaozoologie. Berlin: Borntraeger; p. 834.","Bouchet P, Rocroi JP, Hausdorf B, Kaim A, Kano Y, Nutzel A, Parkhaev P, Schrodl M, Strong EE. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia. 61 (1 - 2): 1 - 526.","Pall-Gergely B, Hunyadi A, Jochum A, Asami T. 2015. Seven new hypselostomatid species from China, including some of the world ̾ s smallest land snails (Gastropoda, Pulmonata, Orthurethra). ZooKeys. 523: 31 - 62. doi: 10.3897 / zookeys. 523.6114.","Pall-Gergely B, Hunyadi A, Grego J, Reischutz A, Buczko K, Vermeulen JJ. 2020. Clostophis Benson, 1860, is not a monotypic diplommatinid but a speciose hypselostomatid (Gastropoda: Eupulmonata), with descriptions of six new species. Raffles Bull Zool. 68: 350 - 368.","Pall-Gergely B, Jochum A, Vermeulen JJ, Anker K, Hunyadi A, Orstan A, A S, Danyi L, Schilthuizen M. 2022. The world ̾ s tiniest land snails from Laos and Vietnam (Gastropoda, Pulmonata, Hypselostomatidae). Contrib Zool. 91 (1): 62 - 78. doi: 10.1163 / 18759866 - bja 10025.","Ancey CF. 1881. Description de mollusques terrestres nouveaux. Le Naturaliste. 3 (47): 373 - 374.","Schileyko AA. 1998. Treatise on recent terrestrial pulmonate molluscs. Part 2. Gastrocoptidae, Hypselostomatidae, Vertiginidae, Truncatellinidae, Pachnodidae, Enidae, Sagdidae. Ruthenica. Supplement 2: 129 - 261.","Panha S, Burch JB. 2005. An introduction to the microsnails of Thailand. Malacol Rev. 37 / 38: 1 - 155.","Mollendorff OF von. 1885. Materialien zur Fauna von China. Jahrbucher der Deutschen Malakozoologischen Gesellschaft. 12: 349 - 398."]}

Genus Bensonella Pilsbry and Vanatta, 1900 Bifidaria (Bensonella) Pilsbry and Vanatta 1900: 591. Boysidia (Bensonella) – Pilsbry 1917 (in Pilsbry 1916 –1918): 198. Boysidia (Paraboysidia) Pilsbry, 1917 (in Pilsbry 1916 –1918): 174, 201. Type species Pupa plicidens Benson, 1849 (Fig. 3 (a–e) herein), by original designation. Remarks According to Pilsbry̾s (1917) original description of Paraboysidia, this genus (at that time a subgenus of Boysidia) differs from Bensonella by having normal (not hooked) apertural barriers. However, this claim is incorrect because the type species of Bensonella (Pupa plicidens) has normal barriers. The type species of Paraboysidia (Boysidia paviei Bavay and Dautzenberg, 1912, fig. 3(f–k)) is very similar to that of Bensonella (Pupa plicidens) in general shell shell and aperture shape, and in the arrangement of the apertural barriers. The main difference is the presence of some additional ̍supplementary̾ apertural folds (i.e. tiny plicae between main plicae) in P. plicidens, which does not seem to justify the distinction of two genera. Therefore, herein Paraboysidia is treated as a junior synonym of Bensonella. Gittenberger et al. (2021) also treated Paraboysidia as a synonym of Bensonella but did not explain their decision to do so. Bensonella species with hooked and normal barriers in the Himalaya (i.e. B. hooki sp. nov. and B. plicidens) are very similar to each other in all other shell characters. Moreover, hooked and normal apertural barriers are characteristic of otherwise extremely similar and close-occurring species in the genera Acinolaemus F.G. Thompson and Upatham 1997; Anauchen Pilsbry, 1917; and Hypselostoma Benson, 1856 (Vermeulen et al. 2007, 2019). This suggests that the hooked or normal apertural barriers are subjects of rapid, probably adaptive (see Wada and Chiba 2013) evolutionary changes. Since it is probable that the hooked apertural barriers have been evolved multiple times in different groups within the Hypselostomatidae, it is unlikely that species with apertural barriers would form a monophyletic unit. Therefore, it is not recommended to use this character state to distinguish between genera. We thus classify species with both hooked (B. hooki sp. nov., B. lakainguta Hwang, 2014) and normal apertural barriers (i.e. B. plicidens) in Bensonella. In the last three decades, several Boysidia (Bensonella) species from China have been newly described or transferred into the (sub)genus: Boysidia dilamellaris D.-N. Chen, Y.-H. Liu and W.-X. Wu, 1995 (transferred to Bensonella by Chen et al. 1999); Boysidia (Bensonella) jinpingensis M. Tian, B. Fao and Y.-X. Chen, 2015; Boysidia (Bensonella) nanjiangensis Z.-L. Zhang, W.-H. Zhang and T.-C. Luo, 2011; Boysidia (Bensonella) qingliangfengensis F. Fang, J. Wang and Y. Chen, 2015; Boysidia (Bensonella) shilinensis D.-N. Chen, M. Wu and G.-Q. Zhang, 1999; Boysidia (Bensonella) tianxingqiaoensis T.-C. Luo, D.-N. Chen and G.-Q. Zhang, 2000 (Luo et al. 2000); Boysidia (Bensonella) tongguanensis D.-N. Chen and W.-H. Zhang, 2002; Boysidia (Bensonella) xingyinensis Y.-H. Guo, W.-C. Zhou and T.-C. Luo, 2006 (Guo et al. 2006); and Boysidia (Bensonella) xiuwenensis W.-H. Zhang, T.-C. Luo and W.-C. Zhou, 2010. Of these, B. dilamellaris, B. nanjiangensis, B. shilinensis, B. tongguanensis and B. xiuwenensis were classified as within Boysidia Ancey, 1881 (without subgeneric classification) by Zhang et al. (2014). We agree with this classification because these species are similar in shell shape and apertural dentition (i.e. fused angular and palatal lamella) to the type species, Pupa dorsata Ancey, 1881. We further classify B. jinpingensis in Boysidia for the same reason (i.e. that species is a member of Boysidia and not Bensonella). Boysidia (Bensonella) tianxingqiaoensis (treated as a Boysidia species by Zhang et al. 2014) and Boysidia (Bensonella) xingyinensis are here both transferred to Gyliotrachela Tomlin, 1930 due to the free and keeled last whorl and the separate parietal and angular lamella all being similar to those of the type species of that genus: Hypselostoma hungerfordianum Möllendorff, 1891. Boysidia (Boysidia) fengxianensis D.-N. Chen, Y.-H. Liu and W.-X. Wu, 1995 was transferred to Bensonella by Schileyko (1998) and to Boysidia (Bensonella) by Zhang et al. (2014). Judging as best we can from the poor-quality drawing in the original description (mostly the three parietal lamellae), we agree that this species belongs to Bensonella. Lastly, the Sumatran Bensonella karoensis Maassen, 1999 is retained in Bensonella herein, due to the arrangement of the apertural barriers (separate angular and parietal lamella, two palatal lamellae, a basal and a columellar lamella) that largely match with the situation in Pupa plicidens Benson, 1849 and Boysidia paviei Bavay and Dautzenberg, 1912, the type species of Bensonella and Paraboysidia, respectively., Published as part of Páll-Gergely, Barna & White, Tom S., 2023, Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae), pp. 2011-2029 in Journal of Natural History 56 (45 - 48) on pages 2014-2016, DOI: 10.1080/00222933.2022.2152750, http://zenodo.org/record/7560931, {"references":["Pilsbry HA, Vanatta EG. 1900. A partial revision of the Pupae of the United States. Proc Acad Nat Sci Phila. 52: 582 - 611.","Pilsbry HA. 1916 - 1918. Manual of conchology, structural and systematic, with illustrations of the species. Ser. 2, Pulmonata. Vol. 24: Pupillidae (Gastrocoptinae). pp 1 - 380, pls 1 - 49. Philadelphia, published by the Conchological Section, Academy of Natural Sciences. [pp. 1 - 112, Pl. 1 - 13, 18 Dec 1916; pp. 113 - 176, pl. 14 - 29, 18 July 1917; pp. 177 - 256, pl. 30 - 38, 09 Nov 1917; pp. 257 - 380, pl. 39 - 49, 24 April 1918].","Benson WH. 1849. Descriptions of four new Asiatic species of the genus Pupa of Draparnaud. Ann Mag Nat Hist. 4 (20): 125 - 128. doi: 10.1080 / 03745486009496158.","Bavay A, Dautzenberg P. 1912. Description de coquilles nouvelles de l ̾ Indo-Chine. Journal de Conchyliologie. 60 (1): 1 - 54.","Gittenberger E, Gyeltshen C, Leda P, Sherub S. 2021. The superfamilies Pupilloidea and Enoidea (Gastropoda, Eupulmonata) in Bhutan. Folia Malacol. 29 (2): 69 - 90. doi: 10.12657 / folmal. 029.009.","Thompson FG, Upatham S. 1997. Vertiginid land snails from Thailand (Gastropoda, Pulmonata, Pupilloidea). Bull Fla Mus Nat Hist. 39 (7): 221 - 245.","Benson WH. 1856. Remarks on the genera Tanystoma, Nematura, and Anaulus. Ann Mag Nat Hist. 17 (2): 342 - 343. doi: 10.1080 / 00222935608697520.","Vermeulen JJ, Phung LC, Truong QT. 2007. New species of terrestrial molluscs (Caenogastropoda, Pupinidae & Pulmonata, Vertiginidae) of the Hon Chong - Ha Tien limestone hills, Southern Vietnam. Basteria. 71: 81 - 92.","Vermeulen JJ, Luu HT, Theary K, Anker K. 2019. Land snail fauna of the Mekong Delta Limestone Hills (Cambodia, Vietnam): notharinia Vermeulen, Phung et Truong, 2007, and a note on Plectostoma A. Adams, 1865 (Mollusca: Gastropoda: Caenogastropoda: Diplommatinidae). Folia Malacol. 27 (3): 167 - 177. doi: 10.12657 / folmal. 027.015.","Wada S, Chiba S. 2013. The dual protection of a micro land snail against a micro predatory snail. PLoS ONE. 8 (1): e 54123. doi: 10.1371 / journal. pone. 0054123.","Hwang C-C. 2014. A new subspecies of land snail Bensonella plicidens lakainguta (Gastropoda, Vertiginidae) from southern Taiwan. Bull Malacol Taiwan 貝類學報. 37: 15 - 26.","Chen D-N, Wu M, Zhang G-Q. 1999. Studies on the genus Boysidia Ancey from China with descriptions of six new species (Pulmonata: Stylommatophora: Pupillidae). Acta Zootaxonomica Sinica. 24 (2): 124 - 135.","Luo T-C, Chen D-N, Zhang G-Q. 2000. Two new species of Pupillidae From China (Pulmonata: Stylommatophora: Pupillidae). Acta Zootaxonomica Sinica. 25 (2): 147 - 151.","Guo Y-H, Zhou W-C, Luo T-C. 2006. One new species of the genus Boysidia (Pulmonata: Stylommatophora: Pupillidae). Acta Zootaxonomica Sinica. 31 (3): 541 - 543.","Ancey CF. 1881. Description de mollusques terrestres nouveaux. Le Naturaliste. 3 (47): 373 - 374.","Zhang W-H, Chen D-N, Zhou W-C. 2014. Two new species of the genus Boysidia from China, with a preliminary discussion of its geography (Pulmonata: Stylommatophora: Pupillidae). Zool Syst. 39 (4): 570 - 575.","Tomlin JRB. 1930. Some preoccupied generic names. - II. Proc Malacol Soc Lond. 19: 22 - 24.","Mollendorff OF von. 1891. On the land and freshwater shells of Perak. Proc Zool Soc Lond. 1891: 330 - 348.","Schileyko AA. 1998. Treatise on recent terrestrial pulmonate molluscs. Part 2. Gastrocoptidae, Hypselostomatidae, Vertiginidae, Truncatellinidae, Pachnodidae, Enidae, Sagdidae. Ruthenica. Supplement 2: 129 - 261.","Maassen WJM. 1999. Some new records of Streptaxidae and Hypselostomatidae from Sumatra, Indonesia, with descriptions of three new species. Basteria. 63: 121 - 126."]}

Bensonella plicidens (Benson, 1849) (Figures 3 (a–e), 4, 5, 9(a,b)). Pupa plicidens Benson, 1849: 126 Pupa plicidens – Küster 1841 –1855: 136, pl. 17, figs 23–24 Boysidia plicidens – Pfeiffer 1853: 553 Vertigo (Odontocyclas) plicidens – Adams and Adams 1858: 173 Pupa (Scopelophila) plicidens – Albers and Martens, 1860: 296 Boysidia plicidens – Hanley and Theobald, 1874 (1870–1876): pl. 100, fig. 8 Boysidia plicidens – Sowerby, 1876: pl. 16, fig. 151 Pupa (Odontocyclas) plicidens – Pfeiffer, 1879 (1878–1881): 350 Boysidia plicidens – Ancey 1881: 373 Bifidaria (Bensonella) plicidens – Pilsbry and Vanatta 1900: 591 Boysidia plicidens – Gude 1914: 294 Bifidaria (Bensonella) landurensis [sic] Pilsbry 1915: 73–74 new synonym Boysidia landurensis – Pilsbry, 1917 (in Pilsbry 1916 –1918): 204, pl. 35, fig. 9 Paraboysidia landourensis – Benthem Jutting, 1950: 39 Bensonella plicidens – Hwang 2014: fig. 5 (partim: fig. 5 shows B. plicidens) Bensonella landourensis and plicidens – Budha and Backeljau 2017 (partim: figs 2B, 3, 5 A and F show B. plicidens) Types examined UMZC I.103325.A, lectotype of P. plicidens (selected herein from a series of 15 syntypes in the UMZC), R. McAndrew coll. (ex- W.H. Benson coll.), Rockville, Landour, Himalaya; NHMUK 1954.6.2.937, paralectotype of P. plicidens, J.S. Hawkins coll. (ex- W.H. Benson coll.), W Himalaya (labelled as paratype); ANSP 16721, holotype of B . landourensis (photos examined: see Figure 5). Additional material examined NHMUK 1856.9.15.69/A, 36 shells, Capt. T. Hutton coll., Moussorie; NHMUK 20191111/A, 10 shells, Cherra, India, mixed sample that includes NHMUK 20191111/B, identified as B. hooki sp. nov.; NHMUK 1903.7.1.2882/B, six shells, Godwin-Austen coll., Moussorie, NW Himalaya, mixed sample that includes NHMUK 1903.7.1.2882/C, identified as B. hooki sp. nov.; NHMUK 1906.2.2.190/B, 7 shells, W. T. Blanford coll., Cherra Poonje, Khasi Hills, mixed sample that includes NHMUK 1906.2.2.190/C, identified as B. hooki sp. nov. Extended diagnosis Shell triangular (pyramidal), slightly ovoid, height somewhat variable; shell colour greyishyellowish and corneous; sculpture with inconspicuous growth lines and very fine spiral striation (its strength variable, in some shells nearly absent); aperture with numerous barriers; parietal wall with a parietal and an angular lamella, both being lower at their middles, and a third, short, deeply situated one between parietal lamella and columellarparietal junction; palatal wall with three long main plicae with elevated central parts, usually (but not always) with additional short plicae between them; columellar lamella similar to palatal plicae; between the lowest palatal plicae and the columellar lamella there are two short plicae; an additional, blunt tubercle sits on the edge of the palatal lip, at the position where the palatal lip forms a little sinus; lower part of columellar lip with two very low, blunt denticles (swellings). Differential diagnosis For differences with B. hooki sp. nov., see under that species. Bensonella plicidens differs from B. lakainguta in the following traits: shell shape is similar, but B. plicidens is wider triangular; the apex is less pointed; the sculpture is clearly densely spirally striated, whereas B. lakainguta is smooth; the arrangement of the apertural barriers is similar, but the barriers of B. plicidens are elongated ridges, which are highest at their middle parts, whereas in B. lakainguta the central part of each barrier is a hook pointing anteriorly (outside of the shell), with an anteriorly elongated low ridge before the hook. Remarks The original spelling of B. landourensis was B. landurensis by Pilsbry (1915), although the type locality was spelled Landour (town in Mussoorie, India). Therefore, the spelling ̍ landourensis ̾ by van Benthem Jutting (1950) is a justified emendation, and ̍ landurensis ̾ is an incorrect original spelling (under ICZN Art. 32.5). Pilsbry (1915) distinguished B. landourensis from B. plicidens by the uninterrupted angular lamella in the former. Additional differences were also mentioned, such as the length of the angular lamella, which does not emerge to the edge of the peristome (i.e. it stops before the peristome edge) in B. landourensis but does in B. plicidens, and the parietal lamella, which is shorter in B. plicidens than in B. landourensis. Although the original description of B. landourensis does not mention hooked apertural barriers in B. plicidens, it is clear that Pilsbry̾s (1915) new species was not compared with correctly identified B. plicidens but with the species that is described here as new (B. hooki sp. nov.), because B. hooki sp. nov. has an interrupted angular lamella, whereas that of B. plicidens is not interrupted. Comparison of the holotype of B. landourensis with syntypes of B. plicidens reveal no significant differences, other than the somewhat more slender shell of B. landourensis, which can be explained by intraspecific variability. Pilsbry (1915) mentioned that the single shell of B. landourensis (= B. plicidens in our current understanding) was found amongst shells of B. plicidens (= B. hooki sp. nov. in our current understanding), all of which had been received from Benson. We were unable to examine the shells studied by Pilsbry, other than the holotype of B. landourensis. It is possible that Benson had both B. plicidens (a species with normal barriers) and B. hooki sp. nov. (a species with hooked barriers) when he described B. plicidens. However, neither Benson (1849) nor subsequent authors mentioned hooked barriers as a character of that species, and the syntypes in the original Benson collection in the UMZC all have normal barriers (Preece et al., in litteris). Even the name ̍ plicidens ̾ indicates that the apertural barriers were plica-like. We do not know how Pilsbry received Benson̾s original specimens, or whether it is really true that he did. It is remarkable that all of the syntypes of B. plicidens that remain in England (UMZC and the shell from the J.S. Hawkins collection in the NHM) have normal barriers, but the lot Pilsbry received, according to this analysis, was entirely hooked with only one normal B. plicidens that was redescribed as B. landourensis by Pilsbry (1915). It is possible that the ̍ B. plicidens ̾ specimens that Pilsbry (1915) referred to as originating from Benson were not part of the type series, or were not directly from William Benson. Here we select the syntype in Cambridge (UMZC) as a lectotype to avoid further nomenclatural problems. Consequently, B. plicidens is a species with normal apertural barriers and B. hooki sp. nov. is a species with hooked barriers., Published as part of Páll-Gergely, Barna & White, Tom S., 2023, Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae), pp. 2011-2029 in Journal of Natural History 56 (45 - 48) on pages 2018-2023, DOI: 10.1080/00222933.2022.2152750, http://zenodo.org/record/7560931, {"references":["Benson WH. 1849. Descriptions of four new Asiatic species of the genus Pupa of Draparnaud. Ann Mag Nat Hist. 4 (20): 125 - 128. doi: 10.1080 / 03745486009496158.","Kuster HC. 1841 - 1855. Die Gattungen Pupa, Megaspira, Balea und Tornatellina. In Abbildungen nach der Natur mit Beschreibungen. In: Systematisches ConchylienCabinet von Martini und Chemnitz. Nurnberg: Bauer & Raspe; p. 194.","Pfeiffer L. 1853. Monographia heliceorum viventium. Sistens descriptiones systematicas et criticas omnium huius familiae generum et specierum hodie cognitarum. Vol. 3, Volumen tertium. Lipsiae: Brockhaus.","Adams H, Adams A. 1858. The genera of recent mollusca, arranged according to their organization. Vol. 2. London: John van Voorst.","Albers JC, von Martens E. 1860. Die Heliceen nach naturlicher Verwandtschaft systematisch geordnet von Joh. Christ. Albers. 2 nd ed. Leipzig: Engelman.","Sowerby GB II. 1876. Monograph of the genus Pupinidae. In: Reeve L, editor. Conchologia iconica, or illustrations of the shells of molluscous animals. Vol. 20. London: L. Reeve; p. 1 - 10. unpaginated text.","Ancey CF. 1881. Description de mollusques terrestres nouveaux. Le Naturaliste. 3 (47): 373 - 374.","Pilsbry HA, Vanatta EG. 1900. A partial revision of the Pupae of the United States. Proc Acad Nat Sci Phila. 52: 582 - 611.","Gude GK. 1914. Mollusca-II. Trochomorphidae-Janellidae. In: Shipley AE, Marshall GAK, editors. The Fauna of British India, including Ceylon and Burma. London: Taylor & Francis; p. XII + 520.","Pilsbry HA. 1915. A new Indian species of Pupillidae. Nautilus (Philadelphia). 29 (7): 73 - 74.","Pilsbry HA. 1916 - 1918. Manual of conchology, structural and systematic, with illustrations of the species. Ser. 2, Pulmonata. Vol. 24: Pupillidae (Gastrocoptinae). pp 1 - 380, pls 1 - 49. Philadelphia, published by the Conchological Section, Academy of Natural Sciences. [pp. 1 - 112, Pl. 1 - 13, 18 Dec 1916; pp. 113 - 176, pl. 14 - 29, 18 July 1917; pp. 177 - 256, pl. 30 - 38, 09 Nov 1917; pp. 257 - 380, pl. 39 - 49, 24 April 1918].","van Benthem Jutting WSS. 1950. The Malayan species of Boysidia, Paraboysidia, Hypselostoma and Gyliotrachela (Gastropoda, Pulmonata, Vertiginidae) with a catalogue of all the species hitherto described. Bulletin of the Raffles Museum. 21: 5 - 47.","Hwang C-C. 2014. A new subspecies of land snail Bensonella plicidens lakainguta (Gastropoda, Vertiginidae) from southern Taiwan. Bull Malacol Taiwan 貝類學報. 37: 15 - 26.","Budha PB, Backeljau T. 2017. First report of Paraboysidia landourensis (Pilsbry, 1915) from Nepal, with description of a putative new species and a note on Bensonella plicidens (Benson, 1849) (Stylommatophora: family unclear). In: Budha PB, editor. Taxonomy of terrestrial molluscs of Nepal PhD dissertation. Antwerp (Belgium): Evolutionary Ecology Group, University of Antwerp; p. 189 - 201."]}

Bensonella hooki Páll-Gergely sp. nov. (Figures 6, 9 (c,d)) Boysidia (Bensonella) plicidens – Pilsbry, 1917 (in Pilsbry 1916 –1918): 198 (partim), pl. 34, figs 1, 2, 4 Type material Holotype. NHMUK 20191114.1, 1 shell (SW: 1.72 mm, SH: 2.01 mm, AW: 0.83 mm, AH: 0.86 mm), Cherrapunji, Godwin-Austen coll. Paratypes. NHMUK 20191114.2, 782 shells, same data as for holotype; NHMUK 20191111 /B, 3 shells from a mixed sample identified as B . plicidens (NHMUK 20191111 / A), Cherra, India; NHMUK 1903.7.1.2882/C, 1 shell from a mixed sample with B . plicidens, Moussorie, NW Himalaya, Godwin-Austen coll.; NHMUK 1906.2.2.190C, 6 shells from a mixed sample identified as B. plicidens (NHMUK 1906.2.2.190B), Cherra Poonje, Khasi Hills, W. T. Blanford coll.; NHMUK 20191107, 16 shells, Khasi, R. Beddome coll., no. 5024; NHMUK 1888.12.4.592–8, seven shells, Cherra, W. Theobald coll.; NHMUK 1903.7. 1.2062, three shells, Diyung valley; NHMUK 20191113, 13 shells, Cherrapoonji, Assam, GodwinAusten coll.; NHMUK 20191110, six shells, Teria Ghat, Assam, H.F. Blanford coll.; NHMUK 20191112, five shells, Khasi Hills, A.S. Kennard coll.; NHMUK 20191109, 13 shells, Himalaya, originally glued to two pieces of paper, some detached); NHMW-MO-69532 (2 shells), Khasi Hills, Ind., coll. Oberwimmer, ex coll. Stoliczka; NHMW-MO-113727 (2 shells), Khasi, coll. Stoliczka, 1880.xv.473; NHMW-MO-113725 (1 shell), Cachar, coll. Stoliczka 1870. Additional material. NHMUK 20191114.3, 9 juveniles, same data as for holotype. Diagnosis. A Bensonella species with a triangular-ovoid, reddish brown shell and hooked apertural barriers. Description. Shell triangular-ovoid, height somewhat variable; apex rather blunt, shell colour reddish brown; sculpture with occasionally strong, widely spaced growth lines; spiral striae absent; aperture subrectangular with clearly isolated sinulus due to angular lamella and parietal tubercle; peristome white, strongly widened, expanded, not reflected; aperture with numerous barriers; parietal wall with an angular lamella having two parts: its outer part reaches peristome, its inner part forms an outside-pointing hook; parietal lamella deeply situated, hooked; third lamella between parietal lamella and columellarparietal junction is ca. half as high as parietal lamella, and also forms a hook; palatal wall with three long main plicae (suprapalatal plica, upper palatal plica, lower palatal plica, however it is possible that the uppermost plica is the upper palatal plicae) with hooked central parts, their outer parts very slender, low, reaching peristome; columellar lamella similar to palatal plicae, hooked; between the lowest palatal plicae and the columellar plicae there are two or three short, hooked plicae; there are additional short, low, deeply situated plicae above the suprapalatal plica inside the sinulus; parietal tubercle blunt, knob-like, sits on edge of palatal lip, at the position where the palatal lip forms a little sinus; lower part of columellar lip widened, sometimes a low denticle-like swelling discernible; umbilicus very narrow, rounded. Measurements (in mm). SW = 1.58–1.77, SH = 1.84–2.3, AW = 0.74–0.83, AH = 0.79–0.92 (NHMUK 20191114). Differential diagnosis. This new species differs from B. plicidens in the following traits: Bensonella hooki has a shell that is larger and darker in colour, with more widely spaced and stronger growth lines; spiral striation absent; aperture with wider peristome, and the palatal tubercle on the peristome edge is less pointed, rather knob-like. The most important difference is in the apertural barriers, which are always hooked in the new species, but are never so in B. plicidens. Bensonella lakainguta has a more triangular shell (with straight side lines) and a more pointed apex, the shell colour is yellowish corneous (reddish brown in B. hooki sp. nov.), its parietal callus is adnate to the penultimate whorl (diverges to a greater distance from the penultimate whorl in B. hooki sp. nov.), the peristome is overall thinner and the parietal tooth on the peristome edge is more pointed. The dentition is strikingly similar. Etymology. This new species is named after Captain Hook, a fictional character from J.M. Barrie̾s Peter Pan, referring to the hook-like apertural barriers. It is a Latin singular, masculine noun in the genitive case. Distribution. Shells of this species were collected in the southwest (Mussoorie) and the southeast (Assam) Himalaya. Remarks. Although shells with hooked and normal barriers have sometimes been included in mixed lots from the Himalaya, there are also several lots with only one ̍ type ̾ of apertural barrier. This, and the additional differences between the two types (i.e. B. plicidens and B. hooki sp. nov.) indicates that contrary to the previous view (Budha and Backeljau 2017), these types are not morphological varieties of the same species, but represent two distinct species., Published as part of Páll-Gergely, Barna & White, Tom S., 2023, Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae), pp. 2011-2029 in Journal of Natural History 56 (45 - 48) on pages 2023-2025, DOI: 10.1080/00222933.2022.2152750, http://zenodo.org/record/7560931, {"references":["Pilsbry HA. 1916 - 1918. Manual of conchology, structural and systematic, with illustrations of the species. Ser. 2, Pulmonata. Vol. 24: Pupillidae (Gastrocoptinae). pp 1 - 380, pls 1 - 49. Philadelphia, published by the Conchological Section, Academy of Natural Sciences. [pp. 1 - 112, Pl. 1 - 13, 18 Dec 1916; pp. 113 - 176, pl. 14 - 29, 18 July 1917; pp. 177 - 256, pl. 30 - 38, 09 Nov 1917; pp. 257 - 380, pl. 39 - 49, 24 April 1918].","Budha PB, Backeljau T. 2017. First report of Paraboysidia landourensis (Pilsbry, 1915) from Nepal, with description of a putative new species and a note on Bensonella plicidens (Benson, 1849) (Stylommatophora: family unclear). In: Budha PB, editor. Taxonomy of terrestrial molluscs of Nepal PhD dissertation. Antwerp (Belgium): Evolutionary Ecology Group, University of Antwerp; p. 189 - 201."]}

Bensonella lakainguta Hwang, 2014 (Figures 7–8, 9 (e–h)) Boysidia (Bensonella) plicidens – Pilsbry, 1917 (in Pilsbry 1916 –1918): 198 (partim), pl. 34, figs 3, 9–10 Boysidia (Bensonella) plicidens – Habe 1956: 109, figs 9–11 Bensonella plicidens – Minato 1988: 41 Bensonella plicidens – Schileyko 1998: 139, fig. 159 Bensonella plicidens – Maassen 1999: 126 Bensonella plicidens lakainguta Hwang, 2014: 18, figs 2–4 Boysidia (Bensonella) qingliangfengensis F. Fang, J. Wang and Y. Chen, 2015: 692, fig. 1 new synonym Material examined NHMUK 201901108, 12 shells, Omi, Japan, Hirase coll. 457; NHMUK 20191128, three shells, Rijozen, Omi, Japan; NHMW-MO-38313 (1 shell), Mt. Fujiwara, 300 m, Japan, coll. Blume ex coll. Azuma, 24 September 1951; NHMW-MO-55810 (5 shells), Japan, Mie Pref., Fujiwara-dake, 300 m, coll. Edlauer ex coll. Kuiper; NHMW-MO-38645 (4 shells), Yoro, Mino; NHMW-MO-69480 (2 shells), Riozen, Omi, Japan, coll. Oberwimmer ex coll. Jetschin; NHMW-MO-113726 (3 shells), Omi, Japan, coll. Rušnov; HNHM 105323 (figured shell), Japan, Kochi Prefecture, Kami-shi, Ryugado Prefectural National Park, above the cave, 500 m along a side-road, 290 m, 33.601917°N, 133.746217°E, Leg. Hunyadi, A., Murányi, D. and Páll-Gergely, B., 08 August 2016. Remarks In the original description, Hwang (2014) mentioned three differences between Indian B. plicidens and the subspecies Bensonella plicidens lakainguta. The most important of them was the submarginal elevations of columellar, basal and palatal barriers, i.e. anterior to the hooks those barriers become more elevated. However, this trait can also be seen in Indian B. hooki sp. nov. (Figure 9 (c,d)); see differences between B. lakainguta and B. hooki sp. nov. outlined under the description of the latter. Boysidia (Bensonella) qingliangfengensis (Figure 7 (e,f)) was described from the border region of the Chinese Anhui and Zhejiang Provinces (very close to Pilsbry̾s (1917) record from Hangzhou), and is claimed to be larger than B . plicidens, yet looks identical for all other characters. Therefore, it is treated here as a junior synonym of B . lakainguta Hwang, 2014. Maassen (1999) mentioned that he compared Bensonella karoensis with Japanese shells of Bensonella plicidens (= Bensonella lakainguta). Furthermore, he erroneously stated that Bensonella plicidens inhabited Thailand., Published as part of Páll-Gergely, Barna & White, Tom S., 2023, Solving the mystery of the misunderstood Bensonella plicidens (Benson, 1849) (Gastropoda: Stylommatophora: Hypselostomatidae), pp. 2011-2029 in Journal of Natural History 56 (45 - 48) on pages 2025-2026, DOI: 10.1080/00222933.2022.2152750, http://zenodo.org/record/7560931, {"references":["Hwang C-C. 2014. A new subspecies of land snail Bensonella plicidens lakainguta (Gastropoda, Vertiginidae) from southern Taiwan. Bull Malacol Taiwan 貝類學報. 37: 15 - 26.","Pilsbry HA. 1916 - 1918. Manual of conchology, structural and systematic, with illustrations of the species. Ser. 2, Pulmonata. Vol. 24: Pupillidae (Gastrocoptinae). pp 1 - 380, pls 1 - 49. Philadelphia, published by the Conchological Section, Academy of Natural Sciences. [pp. 1 - 112, Pl. 1 - 13, 18 Dec 1916; pp. 113 - 176, pl. 14 - 29, 18 July 1917; pp. 177 - 256, pl. 30 - 38, 09 Nov 1917; pp. 257 - 380, pl. 39 - 49, 24 April 1918].","Habe T. 1956. Anatomical studies on the Japanese Land Snails (7). Venus (Japanese Journal of Malacology). 19 (2): 109 - 117.","Minato H. 1988. A systematic and bibliographic list of the Japanese land snails. Shirahama: H. Minato.","Schileyko AA. 1998. Treatise on recent terrestrial pulmonate molluscs. Part 2. Gastrocoptidae, Hypselostomatidae, Vertiginidae, Truncatellinidae, Pachnodidae, Enidae, Sagdidae. Ruthenica. Supplement 2: 129 - 261.","Maassen WJM. 1999. Some new records of Streptaxidae and Hypselostomatidae from Sumatra, Indonesia, with descriptions of three new species. Basteria. 63: 121 - 126."]}

Teruel, Rolando, Rodr��guez-Cabrera, Tom��s M. (2020): Revision of the genus Tityopsis Armas 1974 (Scorpiones Buthidae) Part 1 General updates and description of four new species. Euscorpius 304: 1-40, DOI: http://doi.org/10.5281/zenodo.4648293, {"references":["ACOSTA, L. E., D. M. CANDIDO, E. H. BUCKUP & A. D. BRESCOVIT. 2008. Description of Zabius gaucho (Scorpiones, Buthidae), a new species from southern Brazil, with an update about the generic diagnosis. The Journal of Arachnology, 36: 491-501.","ARMAS, L. F. DE. 1973. Tipos de las colecciones escorpiologicas P. Franganillo y Universidad de la Habana (Arachnida, Scorpionida). Poeyana, 101: 1-18.","ARMAS, L. F. DE. 1974. Escorpiones del archipielago cubano. 3. Genero Tityus C. L. Koch, 1836 (Scorpionida: Buthidae). Poeyana, 135: 1-15.","ARMAS, L. F. DE. 1980. Aspectos de la biologia de algunos escorpiones cubanos. Poeyana, 211: 1-28.","ARMAS, L. F. DE. 1982. Algunos aspectos zoogeograficos de la escorpiofauna antillana. Poeyana, 238: 1-17.","ARMAS, L. F. DE. 1984a. Escorpiones del archipielago cubano. VIII. Adiciones y enmiendas (Scorpiones, Buthidae, Diplocentridae). Poeyana, 275: 1-37.","ARMAS, L. F. DE. 1984b. Tipos de Arachnida depositados en el Instituto de Zoologia de la Academia de Ciencias de Cuba. I. Amblypygi, Opiliones, Ricinulei, Scorpiones, Schizomida y Uropygi. Poeyana, 284: 1-11.","ARMAS, L. F. DE. 1988. Sinopsis de los escorpiones antillanos. Editorial Cientifico-Tecnica, Havana, 102 pp.","ARMAS, L. F. DE. 1989a. Cantidad de hijos por parto en escorpiones cubanos (Scorpiones: Buthidae, Diplocentridae). Ciencias Biologicas, 18: 119-122.","ARMAS, L. F. DE. 1989b. Dos casos de anomalia sexual en escorpiones cubanos (Scorpiones: Buthidae, Diplocentridae). Ciencias Biologicas, 21-22: 173-175.","ARMAS, L. F. DE. 2001. Scorpions of the Greater Antilles, with the description of a new troglobitic species (Scorpiones: Diplocentridae). Pp. 245-253 in Fet, V. & P. A. Selden (eds.). Scorpions 2001. In Memoriam Gary A. Polis. Bucks: British Arachnological Society, xi + 690 pp.","ARMAS, L. F. DE. 2003. On two enigmatic organisms associated with some Antillean and Mexican scorpions (Arachnida: Scorpiones). Revista Iberica de Aracnologia, 7: 239-240.","ARMAS, L. F. DE. 2005. Antillean scorpions deposited at the Montana State University (Arachnida: Scorpiones). Euscorpius, 18: 1-4.","ARMAS, L. F. DE. 2006. Name-bearing types of scorpions deposited at the Institute of Ecology and Systematics, Havana, Cuba (Arachnida: Scorpiones). Euscorpius, 33: 1-14.","ARMAS, L. F. DE & G. ALAYON. 1984. Sinopsis de los aracnidos cavernicolas de Cuba (excepto acaros). Poeyana, 276: 1-25.","ARMAS, L. F. DE & E. MARTIN-FRIAS. 1998. Presencia del genero Tityopsis en Mexico y descripcion de una especie nueva (Scorpiones: Buthidae). Anales de la Escuela Nacional de Ciencias Biologicas, Mexico, 43: 45-49.","ARMAS, L. F. DE, R. TERUEL & F. KOVARIK. 2011. Redescription of Centruroides granosus (Thorell, 1876) and identity of Centrurus granosus simplex Thorell, 1876 (Scorpiones: Buthidae). Euscorpius, 127: 1-11.","COMISION INTERNACIONAL DE NOMENCLATURA ZOOLOGICA [CINZ]. 1999. Codigo Internacional de Nomenclatura Zoologica. 4a edicion. The International Trust for Zoological Nomenclature, xxix + 156 pp.","CRUZ, J. DE LA & L. F. DE ARMAS. 1980. Macroquetas digitoterminales en Buthidae (Arachnida: Scorpionida). Poeyana, 199: 1-10.","FET, V. & G. LOWE. 2000. Family Buthidae C. L. Koch, 1837. Pp. 54-286, in \"Catalog of the scorpions of the world (1758-1998)\" (V. Fet, W. D. Sissom, G. Lowe & M. E. Braunwalder, eds.). The New York Entomological Society, v + 690 pp.","FRANCKE, O. F. 1977. Scorpions of the genus Diplocentrus Peters from Oaxaca, Mexico. The Journal of Arachnology, 4: 145-200.","FRANCKE, O. F. 1985. Conspectus genericus scorpionorum 1758-1982 (Arachnida: Scorpiones). Occasional Papers of the Museum, Texas Tech University, 98: 1-32.","FRANCKE, O. F. 2007. A critical review of reports of parthenogenesis in scorpions (Arachnida). Revista Iberica de Aracnologia, 16: 93-104.","FRANCKE, O. F., R. TERUEL & C. E. SANTIBANEZ-LOPEZ. 2014. A new genus and a new species of scorpions (Scorpiones: Buthidae) from southeastern Mexico. The Journal of Arachnology, 42: 220-232.","JAUME, M. L. 1954. Catalogo de la fauna cubana. IV. Catalogo de los Scorpionida de Cuba. Circulares del Museo y Biblioteca de Zoologia de La Habana, pp. 1085- 1092.","KOVARIK, F. 1998. Stiri [Scorpions]. Madagaskar, Jihlava, 175 pp. [In Czech].","KOVARIK, F., R. TERUEL & G. LOWE. 2016. Two new scorpions of the genus Chaneke Francke, Teruel & Santibanez-Lopez, 2014 (Scorpiones: Buthidae) from southern Mexico. Euscorpius, 218: 1-20."]}

Tityopsis pumila sp. n. (Figures 2, 13, 15, 17, 19, 21, 34, 38, Tables 2, 4) http: //zoobank. org/urn: lsid: zoobank. org: act: 806A095B- 3AFE-4D3F-B0B9-5C333A064BBA TYPE LOCALITY AND TYPE DEPOSITORY. Cuba, Pinar del R��o Province, Guane Municipality, Sierra de Paso Real, 1.5 km northwest of Molina, 50 m a. s. l.; RTO. TYPE SPECIMEN. Cuba, Pinar del R��o Province, Guane Municipality, Sierra de Paso Real, 1.5 km northwest of Molina, 50 m a. s. l., 24 August 2016, leg. T. M. Rodr��guez, A. Rodr��guez & R. L��pez, ♂ (holotype), RTO. ETYMOLOGY. The selected epithet is a Latin adjective that literally means ���dwarf���. It alludes to one the most obvious diagnostic character for this species, i.e., the very small adult size. DIAGNOSIS (male only). Adult size small for the genus (21 mm). Coloration: base orange-brown, slightly darker on metasomal segments IV���V, slightly paler on pedipalp chelae and venter, and much paler on legs; pedipalp femur and patella, carapace, tergites and metasoma symmetrically patterned with dark brown; metasoma with carinae conspicuously darker; pedipalp fingers dark brown. Pedipalps short for the genus (length/ width ratio of femur, patella and chela: 2.65, 2.45 and 3.11); chela with manus short oval (length/width ratio: 1.43) and with fingers very short (movable finger/manus length ratio: 1.17), fixed/movable fingers with 11/11 principal rows of denticles. Tegument of carapace, tergites, and metasoma matt, of pedipalp manus, sternites and telson with a subtle satin sheen. Carapace and tergites with many coarse granules scattered. Pectines with 14/14 teeth; basal middle lamella slightly enlarged and teardrop-shaped. Sternite V with median smooth patch large, longer than wide and lanceolate; lateral smooth patches moderately large, oblique and bulky. Metasoma very short and robust (length/width ratio of segments I���V: 1.03, 1.34, 1.45, 1.61 and 2.11), with 10/8/8/8/7 complete to almost complete, coarsely serrate to serrato-denticulate carinae; lateral inframedians on II obsolete on basal half; intercarinal tegument of segment V with many small conical granules. Telson with vesicle globular; subaculear tubercle small. DESCRIPTION (♂ holotype; Figs. 2, 13, 15, 17, 19, 21, 34; Tabs. 2, 4). Coloration (Figs. 13, 34). Base medium orange-brown, slightly darker on metasomal segment V, slightly paler on pedipalp chelae and venter, and much paler on legs (which are pale yellowish brown). Chelicerae manus moderately reticulate with dark brown distally; fingers moderately infuscate. Pedipalps faintly and irregularly infuscate on all segments except chela manus, with carinae darker and fingers dark brown. Carapace symmetrically and densely spotted with dark brown, mostly below coarser granulose areas; all margins black; eyes and ocular tubercles black. Tergites symmetrically and densely spotted with dark brown, mostly below coarser granulose areas and median keel; posterior margin dark brown. Pectines immaculate yellowish, with basal portion and basal plate progressively darker due to heavier sclerotization. Sternites III���VI immaculate, VII only with posterior margin deeply infuscate; V with median and lateral smooth patches bright white. Legs essentially immaculate, only with external surface of patella and tibia very faintly and irregularly infuscate. Metasoma with segments IV���V slightly darker (reddish), with carinae deeply infuscate and with distal portion of each segment faintly infuscate into an irregularly annulated pattern; dorsal surface of I���IV immaculate; lateral surfaces with large but faint, elongate dark brown spots between carinae all over distal third of all segments; ventral surface with a thick but interrupted dark brown stripe between ventral submedian carinae, plus a pair of large but faint, elongate dark brown spots between ventral submedian and ventrolateral carinae over distal half of segments I���IV and all along V. Telson vesicle ventrally with faint, irregular infuscation; aculeus with distal half dark reddish brown. Chelicerae (Fig. 17a). With dentition typical for the genus, teeth standard-sized and sharp. Tegument glossy but with minute granulation scattered, dorsodistal portion of manus with coarse, glossy granules irregularly arranged transversally around dorsodistal depression. Setation very dense ventrally, but essentially lacking dorsally, except for six rigid macrosetae around dorsodistal depression of manus. Pedipalps (Fig. 15a���c). Short for the genus, with chelae shorter and heavier than standard. Femur essentially straight (very slightly bent inwards distally), with all carinae very strong, coarsely denticulate to serrate; intercarinal tegument matt, very densely and irregularly granulose; internal (i) trichobothria surrounding a large, irregularly triangular spur. Patella straight, with all carinae very strong, coarsely granulose to serrato-crenulate; intercarinal tegument matt, very finely and densely granulose, internally with many large, conical denticles. Chela with manus short-oval (1.43 times longer than wide), much wider than patella (ratio 1.27) and with the basal half slightly widest, all carinae strong, coarsely serrate to serrato-crenulate, intercarinal tegument with a subtle satin sheen, very finely and densely granulose on all surfaces; fingers very short for the genus (movable finger 1.17 times longer than underhand) and evenly curved, fixed finger with 11/11 principal rows of denticles, movable finger with 11/11 plus an apical subrow of four denticles and a large internal accessory denticle (large terminal denticle not included), basal lobe/notch combination absent. Carapace (Fig. 17a). Slightly wider than long (length/width ratio = 0.97). Anterior margin roundly bilobed, with two pairs of dark macrosetae. Carinae: anterior medians anteriorly divergent and formed by very coarse, isolated granules and irregularly fused to the superciliaries, which are composed of much smaller and closer granules; posterior medians straight, parallel and formed by coarse granules; other carinae absent or indistinct from general granulation. Furrows: anterior marginal, anterior median, median ocular, lateral oculars, central transverse, central median, posterior median and posterior marginal all fused, wide and moderately deep; posterior laterals very long, oblique, wide and moderately deep, almost fused to posterior marginal; lateral centrals very long, oblique (essentially parallel to posterior laterals), wide and moderately deep, almost fused to central transverse. Tegument matt, very densely and irregularly but finely granulose, with many coarse, rough granules scattered. Median eyes relatively large and separated by about one ocular diameter; lateral eyes noticeably smaller and largely concealed below anterolateral margin. Sternum (Fig. 19a). Standard for the genus, with two pairs of dark macrosetae. Tegument very densely and irregularly granulose. Genital operculum (Fig. 19a). Standard for the genus, with three pairs of dark macrosetae. Valves roundly subtriangular and widely separated medially, each with a median depression; tegument minutely and densely granulose. Genital papillae large, thick and slightly protruding. Pectines (Fig. 19a). Standard-sized for the genus (not reaching coxa-trochanter joint of leg IV). Tooth count 14/14 (right pecten with tooth eight abnormally short), teeth straight and slightly swollen. Fulcra medium-sized to small, bulky and round. Basal middle lamella slightly enlarged and teardropshaped. Pre-pectinal plate well visible, heavily sclerotized, very wide and short; tegument minutely and densely granulose. Basal plate heavily sclerotized, about as long as wide; anterior margin with a very wide, V-shaped anteromedian notch, posterior margin convex; tegument minutely and densely granulose. Legs. Somewhat less slender than standard, with all carinae strong, finely denticulate to granulose. Intercarinal tegument finely and very densely granulose, with abundant slightly coarser granules scattered mostly over external surface, even defining an additional carina (externomedian) on lateral surface of femur of all legs. Claws short and strongly curved. Mesosoma (Figs. 17b, 19). Tergites largely matt, very densely and irregularly but finely granulose, with many medium-sized to coarse, rough granules scattered; carinae: median longitudinal strong, serrato-crenulate to serrate and progressively longer from I���VII, submedians absent on I��� VI, complete, straight and strongly serrate on VII, laterals absent on I���VI, complete, straight and strongly serrate on VII. Sternites with a subtle satin sheen, very densely and irregularly but finely granulose, with some dark macrosetae scattered all over; lateral margins of all sternites serrate to granulose; posterior margin of all sternites smooth to vestigially granulose, on III vestigially bilobed, on IV widely bilobed, on V narrowly bilobed (due to protuberance of lateral smooth patches), on VI widely bilobed, on VII concave; carinae: III���V acarinate, VI-VII with paired laterals and submedians which are long, straight and coarsely crenulate; spiracles small, round to short-oval; sternite V with median smooth patch large, lanceolate (remarkably longer than wide), bulky and glossy, not protruding from posterior margin; lateral smooth patches moderately large, oblique, oval, bulky and glossy, protruding well from posterior margin. Metasoma (Fig. 21). Shorter than standard for the genus (4.08 times longer than carapace), very robust and slightly narrower distally. Segment I with ten complete carinae, II��� IV with eight, V with seven: dorsal laterals strong, coarsely serrate to serrato-denticulate on I���IV, absent on V but subtly indicated by an incomplete row of conical granules; lateral supramedians strong, coarsely serrate to serratodenticulate on I���V; lateral inframedians strong, coarsely serrato-crenulate on I, present only on distal half of II, absent on III���V (but indicated in III���IV by a few coarser, irregularly arranged granules and on V by a well-defined row of coarse granules that almost define true carinae); ventral laterals strong, coarsely serrate to serrato-denticulate on I���V; ventral submedians strong, coarsely serrate to serratodenticulate on I���IV, indicated by an incomplete but raised row of coarse granules on basal half of V; ventral median absent on I���IV, strong, coarsely serrate to serrato-denticulate on V. Intercarinal tegument matt, very finely and densely granulose, with some medium-sized granules scattered on dorsal surface of all segments and on lateral surfaces of V; dorsal furrow poorly defined, wide and shallow, but distally narrower and deeper; setation sparse, with two pairs of dark macrosetae on essentially all carinae. Telson (Fig. 15d). Vesicle globose (1.36 times longer than wide, exactly as wide as deep) and with many dark and whitish setae scattered; tegument coriaceous but with a subtle satin sheen, with vestiges of coarse granules scattered mostly ventrally; ventral median carina weak but coarsely subgranulose, continued into the subaculear tubercle that is small, broadly conical and unarmed (i.e., without any granules). Aculeus standard-sized, sharp, shorter than vesicle and moderately curved. FEMALE. Unknown. VARIATION. Unknown. AFFINITIES (adult males only). This species is very easy to distinguish at first sight from the other three described species of the genus by having the pedipalp movable finger shorter than carapace. Moreover, by the unique combination of a median smooth patch of sternite V lanceolate (longer than wide), coupled to a tiny size (21 mm) and very robust habitus (e.g., pedipalp only 3.58 times longer than carapace and with movable finger shorter than carapace, metasoma only 4.08 times longer than carapace and with segment I essentially as long as wide). These characters compare to the three other Tityopsis as follows, enhanced by additional diagnostic features: T. mulata sp. n: 1) size larger (25���26 mm); 2) coloration much darker (basically olivaceous brown) and very densely spotted all over with blackish brown; 3) pedipalps more elongate (3.95 times longer than carapace), with movable finger longer than carapace; 4) metasoma more elongate (4.27���4.33 times longer than carapace, with segment I 1.15 times longer than wide); 5) tegument of carapace, tergites, sternites, metasoma and telson with a satin sheen; 6) pectines with slightly higher tooth count (14���15); 7) carapace and tergites with granulation finer; 8) metasoma with carinae weaker and finer. T. inaequalis : 1) size much larger (26���29 mm); 2) coloration much paler (basically light to pale yellowish brown) and only with faint infuscation in carapace and metasomal segments II���IV (even absent in many specimens, mostly males); 3) pedipalps much more elongate (4.00���4.02 times longer than carapace), with movable finger longer than carapace; 4) metasoma more elongate (4.26���4.44 times longer than carapace, with segment I 1.21���1.31 times longer than wide); 5) tegument of carapace, tergites, sternites, metasoma and telson with a satin sheen; 6) pectines with higher tooth count (16/16); 7) carapace and tergites with granulation finer; 8) metasoma with carinae weaker and finer. T. inexpectata : 1) size larger (23���25 mm); 2) coloration basically reddish brown, with carapace remarkably darker (heavily spotted with dark brown); 3) pedipalps more elongate (3.74���3.88 times longer than carapace), with movable finger longer than carapace; 4) metasoma more elongate (4.16���4.25 times longer than carapace, with segment I 1.12���1.14 times longer than wide); 5) sternite V with median smooth patch widely cordiform and about as long as wide; 6) pedipalps and metasoma with carinae weaker and finer; 7) telson with subaculear tubercle variable, but almost always markedly larger. ECOLOGICAL NOTES. This species lives in mesophylous semicaducifolious forest on limestone karstic soil (Fig. 38). All specimens were collected in the slopes of the mogote, under rocks semi-buried in leaf litter and organic soil, usually hanging to the underside of the rocks, as well as directly on the soil. It lives syntopically with a very different Tityopsis sp. (much larger, paler and slenderer) and with C. guanensis and H. junceus, both of which also occur sympatrically on the vegetation., Published as part of Teruel, Rolando & Rodr��guez-Cabrera, Tom��s M., 2020, Revision of the genus Tityopsis Armas 1974 (Scorpiones Buthidae) Part 1 General updates and description of four new species, pp. 1-40 in Euscorpius 304 on pages 15-21, DOI: 10.5281/zenodo.4648293

Tityopsis sheylae sp. n. (Figures 2, 23���32, 36, 40, Tables 3���4) http: //zoobank. org/urn: lsid: zoobank. org: act: 7DD2D5D2- 87BC-49B5-A268-D6C2FB94746E TYPE LOCALITY AND TYPE DEPOSITORY. Cuba, Mayabeque Province, border between San Jos�� de Las Lajas and Jaruco Municipalities, Tapaste, Escaleras de Jaruco, Mogote La Jaula, Cueva del Indio, 190 m a. s. l.; RTO. TYPE SPECIMENS. Cuba, Mayabeque Province, border between San Jos�� de Las Lajas and Jaruco Municipalities, Tapaste, Escaleras de Jaruco, Mogote La Jaula, Cueva del Indio, 190 m a. s. l., 3 April 2019, leg. R. Teruel, S. Yong & T. M. Rodr��guez, 1♂ (holotype), RTO, 13 December 2014, leg. T. M. Rodr��guez & A. Longueira, 3♂ 7♀ 2♂ juvs. 3♀ juvs. (paratypes), RTO, 12 May 2017, leg. S. Yong & L. Forcelledo, 1♀1♀ juv. (paratypes), RTO, 5 January 2018, leg. R. Teruel, S. Yong & L. Forcelledo, 5♂ 9♀ 2♂ juvs. 3♀ juvs. (paratypes), RTO, 3 April 2019, leg. R. Teruel, S. Yong & T. M. Rodr��guez, 2♂ 6♀ (paratypes), RTO. OTHER MATERIAL EXAMINED. Cuba, Mayabeque Province, border between San Jos�� de Las Lajas and Jaruco Municipalities, Tapaste, Escaleras de Jaruco, La Jaula, Cueva del Indio, 190 m a. s. l., 23 December 1995, leg. A. P��rez, 1♂ (topotype), IES, 27 September 1999, leg. E. Fonseca, 1♀ (topotype), IES. Note. Both specimens were repeatedly seen by one of us (RT) between 2005 and 2011, but are currently missing from IES collection. As their whereabouts is unknown (there is no official record of a loan, destruction or loss in the collection log), both were excluded from the type series. ETYMOLOGY. The selected epithet is a Latinized matronym honoring the senior author���s wife Sheyla Yong, who enthusiastically assisted us all along the present revision, both in the field (she actually collected several of the type specimens) and in the lab (part of the data gathering and processing, as well as photographic work). DIAGNOSIS. Adult size moderately large for the genus (24��� 27 mm in males, 32���34 mm in females). Adult coloration: base light yellowish brown, slightly darker on metasomal segments IV���V and slightly paler on legs, venter and telson; dark patterns essentially absent, except for faint infuscation in carapace (mostly around eyes) and metasomal segments II���IV (in some specimens only, mostly females). Juvenile coloration: base pale yellowish brown, paler on pedipalp chelae, legs, venter and telson; dark patterns essentially absent, except for diffuse gray to dark brown reticulation in chelicerae, carapace (along anterior margin and around eyes) and metasomal segments II���V (mostly on posterior part and around carinae); tergites with posterior margins translucent whitish; pedipalps fingers infuscate. Pedipalps remarkably elongate and robust for the genus (length/width ratio of femur, patella and chela: 2.94���3.11, 2.58���2.65 and 3.50���3.66 in males, 2.79���2.96, 2.64���2.71 and 3.51���3.57 in females); chelae with manus incrassate oval and very slightly longer in males (length/width ratio: 1.50���1.56 in males, 1.46���1.53 in females) and with fingers very long (movable finger/manus length ratio: 1.33���1.34 in males, 1.33���1.40 in females), fixed/movable fingers with 12���13/12���13 principal rows of denticles (mode 12/12). Tegument of carapace, tergites, sternites, metasoma and telson sexually dimorphic between adults: matt in male vs. with a satin sheen in females. Carapace and tergites with many medium-sized to coarse granules scattered (similar in both sexes). Pectines with 13���16 teeth in males, 13���15 in females (mode 14 and 13���14, respectively); basal middle lamella slightly enlarged and teardrop-shaped in males, greatly enlarged and widely oval in females. Sternite V with median smooth patch widely cordiform and about as long as wide in both sexes, very large in males and moderately large in females; lateral smooth patches oval, oblique and bulky in both sexes, moderately large in males and small in females. Metasoma slightly elongate in both sexes (length/width ratio of segments I���V: 1.12���1.16, 1.42���1.44, 1.54���1.58, 1.76���1.78 and 2.22���2.25 in males vs. 1.14���1.15, 1.43���1.45, 1.54���1.64, 1.70���1.71 and 2.22���2.24 in females), with 10/10/8/8/7 complete to almost complete, coarsely serrate to serrato-crenulate carinae; lateral inframedians on II obsolete only on basal one-sixth (males) to one-fifth (females); intercarinal tegument of segment V with many small (males) to medium-sized (females) conical granules. Telson with vesicle less globose than standard for the genus, subaculear tubercle small. DESCRIPTION (adult male holotype; Figs. 2, 23, 25, 27, 29, 31, 36a; Tabs. 3���4). Coloration (Figs. 23, 36a) base light yellowish brown, slightly darker on metasomal segments IV���V (medium brown) and slightly paler on legs and venter. Chelicerae manus very sparsely reticulate with brown (more densely distally); fingers very faintly and irregularly infuscate. Pedipalps immaculate, with carinae and fingers slightly darker due to heavier sclerotization; fingers infuscate. Carapace largely immaculate, with faint symmetrical infuscation around all three eye groups; eyes and ocular tubercles black. Tergites immaculate, with carinae of VII slightly darker due to heavier sclerotization. Pectines immaculate pale yellowish brown, with basal portion and basal plate progressively darker due to heavier sclerotization. Sternites immaculate; V with median and lateral smooth patches bright white. Legs immaculate, with carinae slightly darker due to heavier sclerotization. Metasoma with segments IV���V slightly darker (medium brown), with carinae darker due to heavier sclerotization and without any annulated pattern; all surfaces immaculate. Telson immaculate, with distal half of aculeus dark reddish brown. Chelicerae (Fig. 27a). With dentition typical for the genus, teeth standard-sized and sharp. Tegument glossy but with minute granulation scattered, dorsodistal portion of manus with coarse, glossy granules irregularly arranged transversally around dorsodistal depression. Setation very dense ventrally, but essentially lacking dorsally, except for five rigid macrosetae around dorsodistal depression of manus. Pedipalps (Fig. 25a���c). Remarkably enlarged for the genus, i.e., all segments longer and chelae also heavier than standard. Femur essentially straight (very slightly bent inwards distally), with all carinae very strong, coarsely denticulate to serrate; intercarinal tegument matt, very densely and irregularly granulose; internal (i) trichobothria surrounding a large, irregularly triangular spur. Patella straight, with all carinae very strong, coarsely granulose to serrato-crenulate; intercarinal tegument matt, very finely and densely granulose, internally with many conical denticles. Chela with manus incrassate oval (1.56 times longer than wide), much wider than patella (ratio 1.26) and with the basal half slightly widest, all carinae strong, coarsely serrate to serrato-crenulate, intercarinal tegument matt, very finely and densely granulose on all surfaces; fingers very long for the genus (movable finger 1.34 times longer than underhand) and evenly curved, fixed finger with 12/12 principal rows of denticles, movable finger with 12/12 plus an apical subrow of four denticles and a large internal accessory denticle (large terminal denticle not included), basal lobe/notch combination absent. Carapace (Fig. 27a). Wider than long (length/width ratio = 0.93). Anterior margin widely V-shaped, with two pairs of dark macrosetae. Carinae: anterior medians anteriorly divergent and formed by coarse, isolated granules and irregularly fused to the superciliaries, which are composed of smaller and much closer granules; posterior medians straight, parallel and formed by medium-sized granules; other carinae absent or indistinct from general granulation. Furrows: anterior marginal, anterior median, median ocular, lateral oculars, central transverse, central median, posterior median and posterior marginal all fused, wide and moderately deep; posterior laterals long, oblique, wide and moderately deep, almost fused to posterior marginal; posterior transverse narrow and moderately deep; other furrows absent or indistinct. Tegument matt, very densely and irregularly but finely granulose, with many medium-sized, rough granules scattered. Median eyes relatively large and separated by about one ocular diameter; lateral eyes noticeably smaller and moderately exposed beyond anterolateral margin. Sternum (Fig. 29a). Standard for the genus, although more pentagonal in shape, with three pairs of dark macrosetae. Tegument very densely and irregularly granulose. Genital operculum (Fig. 29a). Standard for the genus, with three pairs of dark macrosetae. Valves roundly subtriangular and not separated medially; tegument minutely and densely granulose. Genital papillae large, thick and not protruding. Pectines (Fig. 29a). Moderately large for the genus (but not reaching coxa-trochanter joint of leg IV). Tooth count 14/15, teeth straight and slightly swollen. Fulcra large and bulky, paraboloid to round. Basal middle lamella slightly enlarged and teardrop-shaped. Pre-pectinal plate well visible, heavily sclerotized, very wide and short; tegument coriaceous to smooth. Basal plate normally sclerotized, about as long as wide; anterior margin with a very wide, V-shaped anteromedian notch, posterior margin convex; tegument smooth. Legs. Slender, with all carinae strong, finely denticulate to granulose. Intercarinal tegument minutely and densely granulose, with a row of slightly coarser granules defining an additional carina (externomedian) on lateral surface of femur of all legs. Claws short and strongly curved. Mesosoma (Figs. 27b, 29). Tergites matt, very densely and irregularly but finely granulose, with many medium-sized to coarse, rough granules scattered; carinae: median longitudinal strong, serrato-crenulate to serrate and progressively longer from I���VII, submedians absent on I���VI, complete, straight and strongly serrato-denticulate on VII, laterals absent on I���VI, complete, straight and strongly serrato-denticulate on VII. Sternites matt, very densely and irregularly but finely granulose, with some dark macrosetae scattered all over; lateral margins of all sternites subgranulose to granulose; posterior margin of all sternites smooth to vestigially granulose, on III���IV vestigially bilobed, on V bilobed (due to protuberance of lateral smooth patches), on VI essentially straight, on VII widely concave; carinae: III���V acarinate, VI-VII with paired laterals and submedians which are long, straight and finely crenulate; spiracles small, narrowly oval to short slit-like; sternite V with median smooth patch very large, widely cordiform (about as long as wide), bulky and glossy, not protruding from posterior margin; lateral smooth patches moderately large, oblique, oval, bulky and glossy, moderately protruding from posterior margin. Metasoma (Fig. 31). Slightly longer than standard for the genus (4.23 times longer than carapace) and slightly narrower distally. Segments I���II with ten complete to almost complete carinae, III���IV with eight, V with seven: dorsal laterals strong, coarsely serrate to serrato-denticulate on I���III, coarsely crenulate-denticulate on IV, absent on V but subtly indicated by a few granules; lateral supramedians strong, coarsely serrate to serrato-crenulate on I���V; lateral inframedians strong, coarsely serrato-crenulate on I���II (obsolete on basal one-sixth of the latter), absent on III���V (but indicated in each segment by an incomplete row of granules that never form a true, raised carina); ventral laterals strong, coarsely crenulate to serrato-crenulate on I���V; ventral submedians strong, coarsely serrate to serrato-crenulate on I���IV, indicated by an incomplete but raised row of coarse granules on less than basal half of V; ventral median absent on I���IV, strong, coarsely serrato-crenulate on V. Intercarinal tegument matt, very finely and densely granulose, with many medium-sized granules scattered on dorsal surface of all segments and many small, comical granules on lateral surfaces of V; dorsal furrow absent or indistinct on segments I���III, complete, narrow and moderately deep on IV���V; setation sparse, with two pairs of dark macrosetae on essentially all carinae. Telson (Fig. 25d). Vesicle less globose than standard for the genus (1.48 times longer than wide, 1.02 times wider than deep) and with several dark and whitish setae scattered; tegument coriaceous and matt, smooth even ventrally; ventral median carina very weak but coarsely subgranulose, continued into the subaculear tubercle that is small, broadly conical and unarmed (i.e., without any granules). Aculeus standard-sized, sharp, shorter than vesicle and moderately curved. FEMALE (adult paratopotypes; Figs. 24, 26, 28, 30, 32, 36c���d; Tabs. 3���4). Sexual dimorphism well-marked, differing from described male by: 1) size conspicuously larger (32.1���34.0 mm); 2) tegument of carapace, tergites, sternites, metasoma and telson with a satin sheen; 3) pedipalps relatively shorter, slightly more robust (especially chela manus) and with carinae finer; 4) carapace slightly wider posteriorly; 5) mesosoma relatively wider, with lateral sides markedly convex; 6) pre-pectinal plate absent; 7) genital papillae absent; 8) pectines relatively smaller, with teeth shorter, narrower and slightly fewer in number (13���15, mode 13���14), and with basal middle lamella greatly enlarged, widely oval in shape and whitish; 9) sternite V with median and lateral smooth patches much smaller, less bulky and translucent; 10) metasoma with carinae slightly stronger; 11) telson vesicle with many medium-sized granules scattered ventrally and ventrolaterally, and ventral median carina stronger. VARIATION.The 44 paratopotypes are remarkably homogeneous for most morphological characters such as coloration, tegument sculpture, carination, setation and morphometric ratios, where variation was only minor and mostly due to eval differences (Fig. 36), e. g., older adults have slightly darker coloration and setation, granulation and claws worn-out to different degrees. The only noteworthy exceptions are discussed as follows. Adult size (Tab. 3) varied from 23.6���26.8 mm in males and 32.1���34.0 mm in females. There are two size classes even distinguishable to unaided eye in adult males (separated by a maximum gap of 3.14 mm, i.e., a 12% size difference), but only one in females (5% maximum size difference). Number of principal rows of denticles varied from 12���13 on both fixed and movable fingers, with a clearly defined mode of 12 in each case. Fixed, symmetrical counts for both pedipalps (i.e., same number of rows in all four fingers of the same individual), were the dominant occurrence by far: of 35 specimens analyzed (10 paratopotypes excluded due to having at least one finger broken or too damaged for reliable counting), 25 had 12/12 rows (including the holotype) and one had 13/13 rows (an adult female). The cases of asymmetry did not show any bias towards a defined finger, i.e., they occur in similar frequencies on fixed and movable fingers, and on left and right pedipalps. Pectinal tooth counts (Tab. 4) varied from 13���16 in males and 13���15 in females, with clearly defined modes of 14 and 13��� 14, respectively. Single-tooth asymmetry was a very common occurrence: 7 out of 15 males and 11 out of 25 females (two adults of this sex had a pecten either missing or too damaged for counting); there were no specimens with two or more teeth asymmetry. AFFINITIES. By adults exhibiting the unusual combination of a size large for the genus (over 23 mm in males and 32 mm in females) with a light yellowish, essentially immaculate coloration, this species resembles only T. inaequalis. All other congeners (either described or not) are combined conversely, i.e., if coloration is pale, then size is small and vice versa (actually, most of them are medium-sized and reddish-colored with well-defined darker patterns). Adults of T. inaequalis can be easily separated from those of T. sheylae sp. n. by the following characters: 1) size slightly but consistently larger (26���29 mm in males, 33���36 mm in females); 2) female pedipalp chela slightly shorter and more robust (length/width ratio: 3.43���3.51 in females); 3) female pectines with basal middle lamella slightly smaller and more oval in shape; 4) male sternite V with median smooth patch narrower, longer than wide and lanceolate in shape. Besides, both species are widely allopatric in isolated orographic systems: T. inaequalis in Sierra de Los ��rganos (Pinar del R��o Province) and T. sheylae sp. n. in Escaleras de Jaruco (Mayabeque Province), with their nearest occurrences separated by 170 km air-distance. ECOLOGICAL NOTES. T. sheylae sp. n. has been found so far only inside a single cave (Fig. 40). It occurs scattered all along it, from the first chamber right beyond the entrance (not completely dark, relatively cold, less humid and essentially devoid of bat guano), through the deepest corridors (in absolute darkness, very hot, with water-saturated air and densely covered by bat guano). We always used the UV-detection technique to find it and were able to notice that apparently because of the different microclimatic conditions, the scorpions changed their behavior accordingly: those from the first chamber were mostly found hidden under rocks (only one adult of each sex was found walking between the rocks, in separate collecting events), while all those living in the deep were totally or partially exposed on the walls and floor, either walking slowly or sit-and-wait hunting. It lives syntopically with the buthid Centruroides gracilis (Latreille, 1804), but one species always outnumbers the other depending on the cave site. In the hottest chamber T. sheylae sp. n. is rare (only three specimens found so far), while hundreds of C. gracilis virtually cover all available surfaces. Conversely, in the colder chambers and corridors we have found less than 15 individuals of C. gracilis in total, against 42 of T. sheylae sp. n. Interestingly, no cases of mutual predation have been observed yet, in spite of the coexistence and abundance of both scorpion species all along the cave. On 3 April 2019, we found a small-sized adult male that was being predated upon by an adult female of the spider Ciboneya nuriae Huber & P��rez, 2001 (Pholcidae). The scorpion had just molted (its exoskeleton was still soft and not completely stretched yet, see Fig. 36f) and was already dead about 1 cm away from its exuvium, while the spider sucked it somewhere between the chelicerae and pedipalp coxae. We deduce the scorpion anchored to molt besi, Published as part of Teruel, Rolando & Rodr��guez-Cabrera, Tom��s M., 2020, Revision of the genus Tityopsis Armas 1974 (Scorpiones Buthidae) Part 1 General updates and description of four new species, pp. 1-40 in Euscorpius 304 on pages 26-38, DOI: 10.5281/zenodo.4648293, {"references":["ARMAS, L. F. DE. 1974. Escorpiones del archipielago cubano. 3. Genero Tityus C. L. Koch, 1836 (Scorpionida: Buthidae). Poeyana, 135: 1 - 15."]}

Tityopsis canizaresorum sp. n. (Figures 2, 14, 16, 18, 20, 22, 35, 39, Tables 2, 4) http: //zoobank. org/urn: lsid: zoobank. org: act: AC88BB73- DE45-4045-890D-74C4D6E2A98B TYPE LOCALITY AND TYPE DEPOSITORY. Cuba, Matanzas Province, Ci��naga de Zapata Municipality, Los Hondones, 2���3 m a. s. l., RTO. TYPE SPECIMENS. Cuba, Matanzas Province, Ci��naga de Zapata Municipality, Los Hondones, 2���3 m a. s. l., 22���25/ January/2013, leg. R. Teruel & T. M. Rodr��guez, 1♀ (holotype), RTO; Matanzas Province, Ci��naga de Zapata Municipality, Playa Larga, 1 m a. s. l., 20 June 2011, leg. L. A. Casanella, F. Cala, 1♀ juv. (paratype), RTO; 3.5 km southeast of Soplillar, 3 m a. s. l., 25 January 2013, leg. R. Teruel & T. M. Rodr��guez, 2♀ (paratypes), RTO; Los Hondones, 2���3 m a. s. l., June 1985, leg. J. Novo & L. V. Moreno, 1♀ juv. (paratype), IES, 1 February 2001, leg. R. Teruel, 2♀1♀ juv. (paratypes), RTO, 28 November���4 December 2012, leg. T. M. Rodr��guez, 5♀1♀ juv. (paratypes), RTO, 22���25/January 2013, leg. R. Teruel & T. M. Rodr��guez, 12♀9♀ juvs. (paratypes), RTO, 16 August 2014, leg. T. M. Rodr��guez, 4♀2♀ juvs. (paratypes), RTO. ETYMOLOGY. The selected epithet is a Latinized patronym honoring two good friends of us: the Ca��izares brothers (Maikel and Maydiel), who greatly helped us during our repeated collecting trips to the type locality. DIAGNOSIS (females only). Adult size medium for the genus (25���29 mm). Adult coloration: base yellowish brown, slightly darker on metasomal segments IV���V and slightly paler on legs, venter and telson; dark patterns essentially absent, except for faint brown reticulation in chelicera manus and faint infuscation in carapace (along anterior margin and around eyes) and metasomal segments II���IV (on ventral surface). Juvenile coloration: base pale orange to yellowish brown, slightly paler on pedipalps, legs, venter and telson; dark patterns essentially absent, except for faint gray reticulation in chelicera manus and faint infuscation in carapace (along anterior margin and around eyes) and metasomal segments II���V (mostly on posterior part and around carinae); tergites with posterior margins translucent whitish; pedipalps fingers not infuscate. Pedipalps remarkably elongate for the genus (length/width ratio of femur, patella and chela: 2.70���2.80, 2.54���2.61 and 3.65���3.74); chela with manus elongate-oval (length/width ratio: 1.53���1.61) and with fingers long (movable finger/manus length ratio: 1.31���1.39), fixed/movable fingers with 11���12/11���12 principal rows of denticles (mode 12/12). Tegument of carapace, tergites, sternites, metasoma and telson with a satin sheen. Carapace and tergites with many coarse granules scattered. Pectines with 11���14 teeth (mode 12���13); basal middle lamella greatly enlarged and oval. Sternite V with median smooth medium-sized, as long as wide and cordiform; lateral smooth patches small, oblique and slightly bulky. Metasoma slightly elongate (length/width ratio of segments I���V: 1.09���1.16, 1.40���1.47, 1.52���1.58, 1.71���1.88 and 2.26��� 2.32), with 10/8/8/8/7 complete to almost complete, coarsely denticulate to serrato-denticulate carinae; lateral inframedians on II obsolete on basal half; intercarinal tegument of segment V with many large, conical granules. Telson with vesicle less globose than standard for the genus; subaculear tubercle small. DESCRIPTION (♀ holotype; Figs. 2, 14, 16, 18, 20, 22, 35a; Tabs. 2, 4). Coloration (Figs. 14, 35a). Base light yellowish brown, slightly darker on metasomal segments IV���V (medium brown) and slightly paler on legs, venter and telson. Chelicerae manus very sparsely reticulate with brown (more densely distally); fingers faintly and irregularly infuscate. Pedipalps immaculate, with carinae and fingers slightly darker due to heavier sclerotization. Carapace largely immaculate, with faint symmetrical infuscation along anterior margin and around all three eye groups; eyes and ocular tubercles black. Tergites immaculate, with carinae of VII slightly darker due to heavier sclerotization. Pectines immaculate pale yellowish brown, with basal portion and basal plate progressively darker due to heavier sclerotization. Sternites immaculate; V with median and lateral smooth patches translucent. Legs immaculate, with carinae slightly darker due to heavier sclerotization. Metasoma with segments IV���V slightly darker (medium brown), with carinae darker due to heavier sclerotization and without any annulated pattern; all surfaces immaculate, except for ventral surface with a pair of very faint, elongate brownish spots between ventral submedian and ventrolateral carinae over distal half of segments II���IV. Telson immaculate, with distal half of aculeus dark reddish brown. Chelicerae (Fig. 18a). With dentition typical for the genus, teeth standard-sized and sharp. Tegument glossy but with minute granulation scattered, dorsodistal portion of manus with coarse, glossy granules irregularly arranged transversally around dorsodistal depression. Setation very dense ventrally, but essentially lacking dorsally, except for five rigid macrosetae around dorsodistal depression of manus. Pedipalps (Fig. 16a���c). Remarkably elongate for the genus, i.e., moderately longer and with chelae slenderer than standard. Femur essentially straight (very slightly bent inwards distally), with all carinae very strong, coarsely denticulate to serrate; intercarinal tegument with a satin sheen, very densely and irregularly granulose; internal (i) trichobothria surrounding a large, irregularly triangular spur. Patella straight, with all carinae strong, coarsely granulose to serrato-crenulate; intercarinal tegument with a satin sheen, very finely and densely granulose, internally with many conical denticles. Chela with manus elongate-oval (1.53 times longer than wide), wider than patella (ratio 1.19) and with the basal half slightly widest, all carinae strong, coarsely serrate to serratocrenulate, intercarinal tegument with a satin sheen, very finely and densely granulose on all surfaces; fingers long for the genus (movable finger 1.39 times longer than underhand) and evenly curved, fixed finger with 12/12 principal rows of denticles, movable finger with 12/12 plus an apical subrow of four denticles and a large internal accessory denticle (large terminal denticle not included), basal lobe/notch combination absent. Carapace (Fig. 18a). Wider than long (length/width ratio = 0.93). Anterior margin widely V-shaped, with two pairs of dark macrosetae. Carinae: anterior medians almost parallel (very slightly divergent anteriorly) and formed by very coarse, isolated granules and irregularly fused to the superciliaries, which are composed of much smaller and closer granules; posterior medians almost parallel (very slightly divergent posteriorly) and formed by coarse, isolated granules and irregularly fused to the central laterals and lateral oculars which are equally formed by coarse, isolated granules; other carinae absent or indistinct from general granulation. Furrows: anterior marginal, anterior median, median ocular, lateral oculars, central median, posterior median and posterior marginal all fused, wide and moderately deep; posterior laterals very long, oblique, wide and moderately deep, almost fused to posterior marginal; other furrows absent or indistinct. Tegument with a satin sheen, very densely and irregularly but finely granulose, with many coarse, rough granules scattered. Median eyes relatively large and separated by about one ocular diameter; lateral eyes noticeably smaller and largely concealed below anterolateral margin. Sternum (Fig. 20a). Standard for the genus, although more pentagonal in shape, with two pairs of dark macrosetae. Tegument minutely and densely granulose. Genital operculum (Fig. 20a). Standard for the genus, with three pairs of dark macrosetae. Valves roundly subtriangular and not separated medially; tegument coriaceous. Genital papillae absent. Pectines (Fig. 20a). Smaller than standard for the genus (barely reaching sternite III spiracles). Tooth count 13/12, teeth very short, straight and not swollen. Fulcra large and bulky, paraboloid to round. Basal middle lamella greatly enlarged and oval in shape. Pre-pectinal plate vestigial, poorly sclerotized, very wide and short; tegument smooth. Basal plate normally sclerotized, about as long as wide; anterior margin with only a subtle indication of anteromedian notch, posterior margin paraboloid; tegument smooth. Legs. Slenderer than standard for the genus, with all carinae strong, finely denticulate to granulose. Intercarinal tegument finely and very densely granulose, with abundant slightly coarser granules scattered mostly over external surface, even defining an additional carina (externomedian) on lateral surface of femur of all legs. Claws short and strongly curved. Mesosoma (Figs. 18b, 20). Tergites with a satin sheen, very densely and irregularly but finely granulose, with many coarse, rough granules scattered; carinae: median longitudinal strong, serrato-crenulate to serrate and progressively longer from I��� VII, submedians absent on I���VI, complete, straight and strongly serrate on VII, laterals absent on I���VI, complete, straight and strongly serrate on VII. Sternites with a satin sheen, very densely and irregularly but finely granulose, with some dark macrosetae scattered all over; lateral margins of all sternites subserrate to subgranulose; posterior margin of all sternites essentially smooth, on III���IV widely bilobed, on V vestigially and widely bilobed (lacking any protuberance of lateral smooth patches), on VI essentially straight, on VII shallowly concave; carinae: III���V acarinate, VI-VII with paired laterals and submedians which are long, straight and finely crenulate; spiracles small, narrowly oval to short-oval; sternite V with median smooth patch medium-sized, cordiform (as long as wide), moderately bulky and glossy, not protruding from posterior margin; lateral smooth patches small, oblique, oval, slightly bulky and glossy, not protruding from posterior margin. Metasoma (Fig. 22). Length standard for the genus (3.97 times longer than carapace) and slightly narrower distally. Segment I with ten complete to almost complete carinae, II���IV with eight, V with seven: dorsal laterals very strong, coarsely denticulate to serrato-denticulate on I���IV, absent on V but subtly indicated by an incomplete row of conical granules; lateral supramedians very strong, coarsely denticulate to serrato-denticulate on I���V; lateral inframedians very strong, coarsely serrato-denticulate on I, present only on distal half of II, absent on III���V (but indicated in III���IV by a few coarser, irregularly arranged granules and on V by a well-defined row of very coarse granules that almost define true carinae); ventral laterals very strong, coarsely serrate to serrato-denticulate on I���V; ventral submedians very strong, coarsely serrate to serrato-denticulate on I���IV, indicated by an incomplete but raised row of coarse granules on more than basal half of V; ventral median absent on I���IV, very strong, coarsely denticulate to serrato-denticulate on V. Intercarinal tegument with a satin sheen, very finely and densely granulose, with some medium-sized granules scattered on dorsal surface of all segments and with many large, conical granules on lateral surfaces of V; dorsal furrow complete, narrow and moderately deep on all segments; setation sparse, with two pairs of macrosetae on essentially all carinae. Telson (Fig. 16d). Vesicle less globose than standard for the genus (1.59 times longer than wide, exactly as wide as deep) and with several dark and whitish setae scattered; tegument coriaceous but with a satin sheen, with many medium-sized granules scattered ventrally and ventrolaterally; ventral median carina weak but coarsely subgranulose, continued into the subaculear tubercle that is obsolete. Aculeus standardsized, sharp, shorter than vesicle and moderately curved. MALE. Unknown. This is apparently a female-only species that reproduces asexually by thelytokous parthenogenesis (see below, in Comments section). VARIATION. The 40 paratypes are remarkably homogeneous for most morphological characters such as coloration, tegument sculpture, carination, setation and morphometric ratios, where variation was only minor and mostly due to eval differences, e.g., older adults have slightly darker coloration and setation, granulation and claws worn-out to different degrees. The only noteworthy exceptions are discussed as follows. Adult size (Tab. 2) varied from 24.9���29.0 mm. The maximum gap observed between two specimens was 4.07 mm (a 14% absolute difference), which suggests the existence of at least two distinct size-classes; on the contrary, within each putative size class specimens differed by not more than 1.2 mm (4%). Number of principal rows of denticles varied from 11���12 on both fixed and movable fingers, with a clearly defined mode of 12 in each case. Fixed finger was slightly more variable, with five fingers out of 66 (7.6%) having 11 rows; on movable finger, only two fingers out of 65 (3.1%) had 11 rows. All cases of fixed and movable fingers with 11 rows were singlefinger occurrences (most frequently on right pedipalp), i.e., the specimen had the three other fingers with 12 rows. An adult female paratopotype had right movable finger with seven rows only, but it was a teratology: about half of the denticles were irregularly aligned into a single, poorly defined row. Pectinal tooth counts (Tab. 4) varied from 11���14, with a clearly defined mode of 12���13. Occurrence of symmetric counts between left/right pectines was essentially as common as one-tooth asymmetry (20 vs. 16 individuals, respectively, two specimens excluded from analysis due to having one pecten either missing or too damaged for reliable counting); the only two naturally deviant specimens (i.e., not showing any evidences of teratology), had 11/13 and 14/12. AFFINITIES (adult females only). This species is very easy to distinguish at first sight from the three other described species of the genus where this sex is known, by two unique characters: the metasomal segment V with intercarinal tegument very coarsely denticulate (especially on lateral surfaces) and the pedipalp chela remarkably more elongate and slender (3.65��� 3.83 times longer than wide). In the remaining species, metasomal segment V always has much weaker and sparser intercarinal granulation. Besides, pedipalp chela length/width ratio is much lower and compares as follows, enhanced by additional diagnostic features: T. mulata sp. n.: 1) size larger (32���34 mm); 2) coloration much darker (basically olivaceous brown) and very densely spotted all over with blackish brown; 3) pedipalp chela shorter and more robust (length/width ratio: 3.24���3.30); 4) tegument of carapace, tergites, sternites, metasoma and telson matt; 5) pectines with slightly but consistently higher tooth counts (12���15) and with basal middle lamella slightly smaller and more oval in shape; 6) sternite V with median smooth patch markedly narrower; 7) metasoma more robust, especially segment V (length/width ratio: 2.16���2.17); 8) telson vesicle more globose. T. inaequalis : 1) size much larger (33���36 mm); 2) pedipalp chela shorter and more robust (length/width ratio: 3.43���3.51); 3) carapace and tergites with granulation finer; 4) pectines larger, with higher tooth counts (14���16) and with basal middle lamella slightly smaller and more oval in shape; 5) sternite V with median smooth patch smaller, longer than wide and narrowly cordiform in shape; 6) metasoma with carinae weaker and finer; 7) telson vesicle more globose. T. inexpectata : 1) coloration basically light reddish brown, with carapace darker and pedipalp fingers deeply infuscate; 2) pedipalp chela shorter and more robust (length/width ratio: 3.31���3.48); 3) sternite V with median smooth patch shorter and wider than long; 4) metasoma markedly more robust (length/width ratio of segments I���V: 1.01���1.08, 1.31���1.35, 1.49���1.53, 1.63���1.70 and 2.01���2.06); 5) telson vesicle more globose. The females of T. pumila sp. n. are still unknown, but it can be distinguished from T. canizaresorum sp. n. based on a nonsexually dimorphic character: the coloration remarkably darker (base orange-brown) and symmetrically patterned with dark brown on pedipalps (femur and patella), carapace, tergites and metasoma. Moreover, T. pumila sp. n. and T. canizaresorum sp. n. are two of the most widely allopatric members of Tityopsis, by occurring near the opposite longitudinal tips of the genus range (Fig. 2). ECOLOGICAL NOTES. This species lives exclusively in mesophylous semicaducifolious forest on karstic soil (Fig. 39). Almost all specimens were collected during diurnal searches, under rocks of various sizes semi-buried in the leaf litter, usually hanging to the underside of the rock, as well as directly on the soil. In the type locality, T. canizaresorum sp. n. is more common inside the small collapsed depressions (dolines) that are locally called ���hondones��� and give the place its name. Repeated nocturnal searches by the authors with UV light at Los Hondones, Playa Larga and other sites scattered all over Zapata Swamp, were completely unsuccessful to yield any specimens of this elusive species. Nevertheless, two juveniles were unintentionally found at night inside human-inhabited rooms: one walking on the floor at the type locality at about 22:30 hrs, the other on a hotel bed at Playa Larga at about 2:15 hrs. The latter stung one of the collectors in his back while sleeping and caused him excruciating pain for about two hours (Luis A. Casanella, personal communication). In the three known localities, T. canizaresorum sp. n. lives syntopically with the buthids C. guanensis and H. junceus (which are also commonly found on the vegetation, especially the former), as well as with the diplocentrid Heteronebo morenoi (Armas, 1973), which is largely restricted to the seashore. All collecting events listed above included pregnant females, thus, this species reproduces all year round. Five captive females gave birth to litters of 6���17 newborn, which underwent the first ecdysis 4���6 days later. COMMENTS. Although not proved yet, there is strong factual evidence that T. canizaresorum sp. n. reproduces by obligate thelytokous parthenogenesis. Most adult females were collected pregnant, but not a single male is present in the entire sample (41 specimens, 26 of them adults) and the five females that gave birth in captivity, produced all-female broods. We tried to get a direct, positiv, Published as part of Teruel, Rolando & Rodr��guez-Cabrera, Tom��s M., 2020, Revision of the genus Tityopsis Armas 1974 (Scorpiones Buthidae) Part 1 General updates and description of four new species, pp. 1-40 in Euscorpius 304 on pages 23-26, DOI: 10.5281/zenodo.4648293, {"references":["ARMAS, L. F. DE. 1973. Tipos de las colecciones escorpiologicas P. Franganillo y Universidad de la Habana (Arachnida, Scorpionida). Poeyana, 101: 1 - 18.","FRANCKE, O. F. 2007. A critical review of reports of parthenogenesis in scorpions (Arachnida). Revista Iberica de Aracnologia, 16: 93 - 104."]}

Helicopsyche alajuela Johanson and Holzenthal, 2010 PANAMA, Chiriqu�� Province, Cuenca 108, R��o Majagua, Banquito de Palmira, 8.68083��N and 82.53250��W, 840 m, Malaise trap, T. R��os and Y. Aguirre, 28 February���14 March 2019, 1 male. Distribution. Costa Rica, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 6, DOI: 10.5281/zenodo.3688655

Marilia kingsolveri Bueno-Soria and Rojas-Ascencio, 2004 PANAMA, Chiriqu�� Province, Cuenca 108, R��o Majagua, Banquito de Palmira, 8.68083˚N and 82.53250˚W, 840 m, Malaise trap, T. R��os and Y. Aguirre, 28 February-14 March 2019, 1 male. Distribution. Costa Rica, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 5, DOI: 10.5281/zenodo.3688655

Ochrotrichia unica Bueno-Soria and Santiago, 1992 PANAMA, Veraguas Province, Cuenca 132, Santa Fe National Park, R��o Mulaba, Isleta, PSPSCB- PNSF-C 132-2017-015, 8.54513��E and 81.11970��W, 412 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 22 April 2017, 1 male. Distribution. Colombia, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 5, DOI: 10.5281/zenodo.3688655

Brysopteryx esparta Harris and Holzenthal, 1994 PANAMA, Veraguas Province, Cuenca 097, Santa Fe National Park, afluente R��o Calovebora, PSPSCB-PNSF-C097-2017-005, 8.54318��E and 81.16398��W, 536 m, Malaise trap, T. R��os, E. Alvarez, and C. Nieto, 19���23 April 2017, 1 male. Distribution. Costa Rica, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 4, DOI: 10.5281/zenodo.3688655

Mayatrichia illobia Harris and Holzenthal, 1990 PANAMA, Veraguas Province, Cuenca 097, Santa Fe National Park, R��o Llanito, PSPSCB-PNSF- C097-2017-012, 8.56553��E and 81.18817��W, 340 m, UV light trap, A. Cornejo, T. R��os, E. Alvarez, and C. Nieto, 20 April 2017, 8 males. Distribution. Costa Rica, Ecuador, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 4, DOI: 10.5281/zenodo.3688655

Wormaldia bolivari Mu��oz-Quesada and Holzenthal, 2015 PANAMA, Chiriqu�� Province, Cuenca 108, Quebrada Grande, Boquete, Valle Escondido, 8.78365˚N and 82.44429˚W, 1147 m, T. Arefina-Armitage, 15 September 2018, 1 male. Distribution. Panama, Venezuela., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 3, DOI: 10.5281/zenodo.3688655, {"references":["Munoz-Quesada, F., and R. W. Holzenthal. 2015. Revision of the Neotropical species of the caddisfly genus Wormaldia McLachlan (Trichoptera: Philopotamidae). Zootaxa 3998: 1 - 138."]}

Wormaldia zunigae Mu��oz-Quesada and Holzenthal, 2015 PANAMA, Chiriqu�� Province, Cuenca 108, tributary of Quebrada Grande, Boquete, Valle Escondido, 8.78173��N and 82.44395��W, 1194 m, UV light trap, B. Armitage and T. Arefina-Armitage, 11 March 2018, 1 male; ibid., Quebrada Grande, 8.78365˚N and 82.44429˚W, 1147 m, T. Arefina-Armitage, 15 September 2018, 1 male. Distribution. Colombia, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 3, DOI: 10.5281/zenodo.3688655, {"references":["Munoz-Quesada, F., and R. W. Holzenthal. 2015. Revision of the Neotropical species of the caddisfly genus Wormaldia McLachlan (Trichoptera: Philopotamidae). Zootaxa 3998: 1 - 138."]}

Costatrichia falsa Santos, Takiya, and Nessimian, 2013 PANAMA, Veraguas Province, Cuenca 132, Santa Fe National Park, R��o Mulaba, afluente 1er Brazo, PSPSCB-PNSF-C132-2017-008, 8.54318��E and 81.16398��W, 770 m, Malaise trap, T. R��os, E. Alvarez, and C. Nieto, 19-23 April 2017, 2 males. Distribution. Costa Rica, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 4, DOI: 10.5281/zenodo.3688655

Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A., Aguirre, Yusseff (2020): The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama. Insecta Mundi 744 (744): 1-8, DOI: http://doi.org/10.5281/zenodo.3688655, {"references":["Aguila, Y. 1992. Systematic catalogue of the caddisflies of Panama (Trichoptera). p. 532-548. In: D. Quintero and A. Aiello (eds.). Insects of Panama and Mesoamerica: Selected Studies. Oxford University Press; Oxford, England. 714 p.","Armitage, B. J., R. J. Blahnik, S. C. Harris, A. Cornejo, and T. I. Arefina-Armitage. 2018. The Trichoptera of Panama. VII. Additional new country records for caddisflies from the Republic of Panama. Insecta Mundi 0614: 1-7.","Armitage, B. J., and S. C. Harris. 2018a. The Trichoptera of Panama. V. Descriptions of new species, new country records, and a synonymy. Insecta Mundi 0604: 1-11.","Armitage, B. J., and S. C. Harris. 2018b. The Trichoptera of Panama. VIII. The Hydroptilidae of Panama: current status, biodiversity comparisons, projections, and needs. Aquatic Insects 39: 95-115.","Armitage, B. J., S. C. Harris, R. J. Blahnik, and R. E. Thomson. 2016. The Trichoptera of Panama IV. New records for caddisflies (Insecta: Trichoptera) from the Republic of Panama. Insecta Mundi 0511: 1-13.","Armitage, B. J., S. C. Harris, and R. W. Holzenthal. 2015. The Trichoptera of Panama. I. New records for caddisflies (Insecta: Trichoptera) from the Republic of Panama. Insecta Mundi 0435: 1-10.","Blahnik, R. J., and B. J. Armitage. 2019. The Trichoptera of Panama XII. Contributions to our knowledge of the caddisfly family Glossosomatidae (Trichoptera) in Panama. Insecta Mundi 0740: 1-17.","Blahnik, R. J., and R. W. Holzenthal. 2004. Collection and curation of Trichoptera, with an emphasis on pinned material. Nectopsyche, Neotropical Trichoptera Newsletter 1: 8-20.","Bowles, D. E., S. G. Tiemann, and G. W. Easley. 2007. Caddisfly (Insecta: Trichoptera) assemblages of large springs and spring-runs in central Texas, U.S.A. p. 15-29. In: J. Bueno-Soria, R. Barba- Alvarez, and B. J. Armitage (eds.). Proceedings of the 12th International Symposium on Trichoptera. Caddis Press; Columbus, Ohio. 378 p.","Bueno-Soria, J., and R. Barba-Alvarez. 2015. New species of Plectropsyche Ross 1947 (Trichoptera: Hydropsychidae: Hydropsychinae). Zootaxa 4040: 421-432.","Bueno-Soria, J., J. J. Morrone, and R. Barba-Alvarez. 2007. Trichoptera of the Sierra Tarahumara, Chihuahua, Mexico. p. 31-35. In: J. Bueno-Soria, R. Barba-Alvarez, and B. J. Armitage (eds.). Proceedings of the 12th International Symposium on Trichoptera. Caddis Press; Columbus, Ohio. 378 p.","Calor, A. R., and R. Mariano. 2012. UV light pan traps for collecting aquatic insects. EntomoBrasilis 5: 164-166.","Harris, S. C., and B. J. Armitage. 2015. The Trichoptera of Panama. II. Ten new species of microcaddisflies (Trichoptera: Hydroptilidae). Insecta Mundi 0437: 1-17.","Harris, S. C., and B. J. Armitage. 2019. The Trichoptera of Panama. X. The Quebrada Rambala drainage, with description of 19 new species of microcaddisfies (Trichoptera: Hydroptilidae). Insecta Mundi 0707: 1-54.","Holzenthal, R. W., R. E. Thomson, and B. Rios-Touma. 2015. Order Trichoptera. p. 965-1002. In: J. Thorp and D. C. Rogers (eds.). Ecology and general biology: Thorp and Covich's freshwater invertebrates. Volume I. Fourth edition. Academic Press; New York, NY. 1148 p.","Moulton, S. R., II, and S. C. Harris. 1997. New species of southwestern Nearctic microcaddisflies (Trichoptera: Hydroptilidae). Proceedings of the Entomological Society of Washington 99: 494-501.","Munoz-Quesada, F., and R. W. Holzenthal. 2015. Revision of the Neotropical species of the caddisfly genus Wormaldia McLachlan (Trichoptera: Philopotamidae). Zootaxa 3998: 1-138.","Razuri-Gonzalez, E., and B. J. Armitage. 2019. The Trichoptera of Panama XI. Three new species of caddisfies in the genus Smicridea McLachlan (Trichoptera: Hydropsychidae) from Omar Torrijos and Santa Fe National Parks. Insecta Mundi 0710: 1-13.","Thomson, R. E., and B. J. Armitage. 2018. The Trichoptera of Panama. VI. Seven new species of microcaddisflies (Insecta: Trichoptera: Hydroptilidae) from Mount Totumas Cloud Forest and Bio- logical Reserve. Insecta Mundi 0613: 1-15."]}

Helicopsyche breviterga Flint, 1991 PANAMA, Chiriqu�� Province, Cuenca 108, R��o Majagua, Banquito de Palmira, 8.68083��N and 82.53250��W, 840 m, Malaise trap, T. R��os and Y. Aguirre, 28 February���14 March 2019, 2 males; ibid., 14 March���4 April, 2019, 1 male. Distribution. Colombia, Ecuador, Panama, Venezuela., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 6, DOI: 10.5281/zenodo.3688655

Ochrotrichia quinealensis Bueno-Soria and Holzenthal, 1998 PANAMA, Veraguas Province, Cuenca 097, Santa Fe National Park, afluente R��o Calovebora, PSPSCB-PNSF-C097-2017-005, 8.54318��E and 81.16398��W, 536 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 21 April 2017, 2 males; ibid., Cuenca 132, Santa Fe National Park, R��o Mulaba, 2do Brazo, PSPSCB-PNSF-C132-2017-007, 8.52577��E and 81.13045��W, 623 m, Malaise trap, A. Cornejo, T. R��os, E. Alvarez, and C. Nieto, 19-23 April 2017. 4 males; ibid., R��o Mulaba, afluente 1er Brazo, PSPSCB-PNSF-C132-2017-008, 8.51706��E and 81.12140��W, 770 m, Malaise trap, T. R��os, E. Alvarez, and C. Nieto, 19���23 April 2017, 4 males; ibid., R��o Mulaba, 1er Brazo, PSPSCB-PNSF-C132-2017-009, 8.52560��E and 81.12956��W, 623 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 19 April 2017, 6 males; ibid., R��o Mulaba, 3er Brazo, PSPSCB-PNSF-C132-2017-010, 8.52906��E and 81.13943��W, 662 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 19 April 2017, 6 males; ibid., R��o Mulaba, afluente, antes de caseta MiAmbiente, PSPSCB-PNSF-C132-2017-014, 8.53143��E and 81.14975��W, 746 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 21 April 2017, 20 males; ibid., R��o Mulaba, Isleta, PSPSCB- PNSF-C132-2017 - 015, 8.54513��E and 81.11970��W, 412 m, UV light trap, T. R��os, E. Alvarez, and C. Nieto, 22 April 2017, 6 males. Distribution. Costa Rica, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 5, DOI: 10.5281/zenodo.3688655

Triaenodes morai Holzenthal and Andersen, 2004 PANAMA, Cocl�� Province, Cuenca 103, Omar Torrijos National Park, Quebrada Corazones, PSPSCB- PNGDOTH-C103-2017-001, 8.67760��N and 80.60007��W, 728 m, UV light trap, A. Cornejo, E. P��rez, T. R��os, E. Alvarez, and C. Nieto, 24 March 2017, 1 male. Distribution. Costa Rica, Nicaragua, Panama., Published as part of Armitage, Brian J., Harris, Steven C., Blahnik, Roger J., Thomson, Robin E., R��os Gonz��lez, Tom��s A. & Aguirre, Yusseff, 2020, The Trichoptera of Panama XIII. Further new country records for caddisflies (Insecta: Trichoptera) from the Republic of Panama, pp. 1-8 in Insecta Mundi 744 (744) on page 5, DOI: 10.5281/zenodo.3688655

Anacroneuria planicollis Klap��lek, 1923 (Figs. 2���12) Material examined: Panam��: Chiriqu��, Jaramillo Abajo, Caldera River, 1065 masl, 8��44���46.97���N and 82��25���8.77���O. 1 n, 04.ii.2015, #17; ibid., 1 n, 04.ii.2015, #19; ibid., 1 n, 04.ii.2015, # 22; ibid., 1 n, 21.ii.2015, #23;1 n, 21.ii.2015, #24;1 n, 21.ii.2015, #25;1 n, 21.ii.2015, #26;1 n, 21.ii.2015, #27;1 n, 21.ii.2015, #28;1 n, 21.iii.2015, #48; ibid., 1 n, 21.iii.2015, #49; ibid., 1 n, 21.iii.2015, #50; ibid., 1 n, 21.iii.2015, #51; ibid., 1 n, 21.iii.2015, #52; ibid., 1 n, 21.iii.2015, #53; ibid., 1 n, 21.iii.2015, #54; ibid., 1 n, 25.iv.2015, #99; ibid., 1 n, 25.iv.2015, #100; ibid., 1 n, 24.viii.2015, #190;1 n, 17. ix.2015, #196; ibid., K. Castillo, MUPADI (nymphs: m # 1, f # 1, adults: m # 2, f # 16). Adult: Description by Stark (1998). Description of the last nymphal instar. Male (n=3): Body brown and stout, body length 12.3 �� 0.23 mm (not including cerci) (Fig. 2). Head (length = 1.9 �� 0.17 mm, width = 2.9 �� 0.26 mm): mostly brown, with a dark spot in dorsal view that extends from the back of the ocelli to the middle part of the head. Light colored ocelli with dark margins and a clear area extended laterally, front with a clear and discontinuous M-shape spot, brown labrum (Fig. 3), filamentous antennae (length: 7.5 �� 1.10 mm) with dark hairs in the proximal region of the scape (Fig. 4). Thorax (n=3): Pronotum (length = 1.6 �� 0.11 mm, width = 2.9 �� 0.21 mm) with band pattern (dark, light, dark, light) from the midline to the lateral margin of the pronotum, pronotal margin clear with spines (Fig. 3). Mesonotum (length = 2.6 �� 0.73 mm, width = 32 �� 0.21 mm) and metanotum (length = 2.6 �� 0.17 mm, width = 2.7 �� 0.23 mm) with pattern of clear, diffuse drawings, wing pads dark in nymphs closed to emerge, with spines in the margins (Fig. 9). Thoracic gills: sc1 present, sc2 and sc3 absent; pl1, pl2 and pl3 present (Fig. 10). Legs mostly brown, covered with dark hairs and outer side margins with few fringes of bristles. Foreleg (Figs 5���6): coxa with numerous dorsal spines in the distal region and without ventral spines (Fig. 8). Femur covered with dark hairs, with a transverse row of spines (7 to 8) in the middle part and a patch of small spines in the proximal lower part, presence of a dark spot at the distal end, without ventral distal spines (Fig. 7). Abdomen: Brown tergites with long, strong spines, presence of a circular brown spot in the ninth sternite (Fig. 11), anal gills absent, cerci with few fine bristles at the distal end. Variations Female (n=11): Body characteristics similar to those of the male. Body length 16.1 �� 0.82 mm (does not include cerci, n = 3), antennas: n = 10, long = 7.55 �� 2.69 mm, head: long = 2.42 �� 0.17 mm, width = 3.71 �� 0.21 mm, pronotum: long = 2.15 �� 0.08 mm, width = 3.77 �� 0.28 mm, mesonotum: long = 3.23 �� 0.24 mm, width = 4.08 �� 0.32 mm, metanotum: long = 3.31 �� 0.16 mm, width = 3.68 �� 0.25 mm, inverted V-shaped notch in the eighth sternite (Fig. 12). Diagnosis: The nymphs of A. planicollis can be distinguished from A. quetzali, by the presence of a dark spot on the head in dorsal view that extends from the back of the ocelli to the middle part of the head, presence of pattern of bands from the midline towards the lateral margin of the pronotum, coxa without ventral spines, and femur with a transversal row of spines (7 to 8) medially. Distribution: Belize, Costa Rica, Guatemala, M��xico, Nicaragua, and Panam��. Ecology: The nymphs of A. planicollis were collected in a shallow area of the Caldera River with bedrock substrate, submerged leaf litter, and slow current. The nymphs of A. planicollis were positively correlated with temperature and conductivity (Castillo S��nchez et al., 2018)., Published as part of Castillo S��nchez, Kayla N., Aguirre, Yusseff P., R��os Gonz��lez, Tom��s A. & Bernal Vega, Juan A., 2019, The Plecoptera of Panama. IV. Description of the last nymphal instar for two species of Anacroneuria Klap��lek (Plecoptera: Perlidae) from the Caldera River, Chiriqu��, Panam��, pp. 446-458 in Zootaxa 4711 (3) on pages 447-450, DOI: 10.11646/zootaxa.4711.3.2, http://zenodo.org/record/3586528, {"references":["Klapalek, F. (1923) Plecopteres nouveaux. Cinquieme partie. Annales de la Societe Entomologique de Belgique, 63, 21 - 29.","Stark, B. P. (1998). The Anacroneuria of Costa Rica and Panama (Insecta: Plecoptera: Perlidae). Proceedings of the Biological Society of Washington, 11, 551 - 603.","Castillo Sanchez, K. N., Aguirre, E. Y. P., Rios Gonzalez, T. A. & Bernal Vega, J. A. (2018) Anacroneuria (Plecoptera: Perlidae) from Caldera River, Chiriqui, Panama: new distribution records, and comments on altitudinal distribution and seasonal variation. Biologia Tropical, 66 (1), 164 - 177. https: // doi. org / 10.15517 / rbt. v 66 i 1.28924"]}

Petroscolex centenarius Csuzdi, Szederjesi & Sherlock, sp. nov. (Figures 2, 3) Holotype. NHMUK 1997.1594 clitellate adult. DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 20.02.1973. Paratypes. HNHM AF/3555 1 ex. clitellate adult, NHMUK 1997.1595 1 ex. clitellate adult broken in two pieces. Locality and date same as of the Holotype. NHMUK 1997.1548 ��� 1549 1 ex. clitellate adult, 1 ex. juvenile, HNHM AF/3539 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 08.03.1973. NHMUK 1997.1526 ��� 1529 1 ex. clitellate adult, 1 ex. aclitellate adult, 2 ex. juvenile, NHMUK ANEA 2019.7368 1 ex. clitellate adult, HNHM AF/5754 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 09.03.1973. NHMUK 1997.1530 1 ex. aclitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 07.06.1973. Etymology. The specific epithet refers to the 100th birthday of Prof. Pietro Omodeo. Description. Holotype: Preserved length 160 mm, diameter after clitellum 5 mm, segment number 196. Paratypes: 150���190 mm in length, 4���5 mm in diameter. Segment number 153���206, anteclitellar segments frequently multiannulate, segments ii���xi carinate. Colour preserved brownish, alive unknown. Head epilobous 1/2 open, dorsal pores lacking. Setae: ab distant cd paired, setal ratio after clitellum aa:ab:bc:cd:dd = 4:3:3.6:1:12. Nephridial pores begin on segment ii, aligned somewhat dorsal to setal line c. Clitellum annular on segments ��xiii�����xvii. Prostatic pore midventral on a spherical papilla in xiii, near to the 12/13 intersegmental furrow which sometimes emerges as a penis-like organ. Female pores small dots on xiv, close to the intersegmental furrow 14/15 just below setal line d. Spermathecal pore single midventral on xviii (Fig. 2 A���C). Internal characters: Muscular gizzard in vi, large, cylindrical. Septa 7/8���10/11 moderately, 11/12 slightly thickened. Calciferous glands paired in xiv, large mouflon-horn shaped. Small chylus-sacs in ix, x, xi. Dorsal blood vessel simple throughout. Hearts in x���xi, moniliform. Nephridial system holoic with thin biramous nephridial bladders. Typhlosole lacking. Proandric. Testes and sperm funnels in x enclosed in a coiled sperm-reservoir. Seminal vesicles large in xi. Ovaries in xiii enclosed in an ovarian capsule pendant from septum 12/13. This ovarian capsule laterally continuing in a short, hardly recognizable ovarian duct joining the fertilisation chamber and bearing a pendant ovisac. From the fertilisation chamber a straight oviduct leads to the female pore in xiv. The ovarian capsule medially communicates on both sides with a delicate ovo-spermathecal duct running alongside the bursa copulatrix and joining apically to the pear-shaped, highly muscular spermathecal atrium (Fig. 3A). This atrium opens midventrally on segment xviii. The left and right ovarian apparatus connected by a small intercommunicating duct running above the intestine (Fig. 3A). Prostates a pair of long spiral tubes, joining the bursa copulatrix (copulation pouch) via a common Y-shaped duct. Bursa copulatrix situated above the spermathecal atrium and completely covers it dorsally. Bursa copulatrix highly muscular, somewhat spindle-shaped, oriented headwards and opening (through the prostatic pore) on a penislike protuberance on xiii (usually retracted). The two male ducts run backward along both sides of this copulatory pouch, and join to it subapically near each other (Fig. 3B). Penial setae lacking. Discussion. Eudrilidae is the only earthworm family with internal fertilisation (Clausen 1965; Jamieson 1967; Sims 1969), which requires intercommunication of the ovarial and spermathecal systems resulting in a common ovo-spermathecal apparatus. This ovo-spermathecal apparatus is highly variable among different taxa both in its structure and external opening (Sims 1987). The external opening of the spermatheca, usually located near the front of the clitellum, is frequently shifted back: for example, in Beddardiella Michaelsen, 1910 (spermathecal pore at 16/17), Keffia Clausen, 1963 (in xix), or in Parapolytoreutus Segun, 1980 (as far back as segment 22/23). Sometimes it shows wide variation even inside a genus: for example, in Buettneriodrilus aequatorialis Michaelsen, 1935 the spermathecal pore is in 12/13, while in Buettneriodrilus armatus (Michaelsen, 1913) it is in segment xix (Zicsi & Csuzdi 1986; Sims 1987). The position of the prostate pores is less variable; in the genera within the family described to date, it can be found in xvii, 17/18 or in xviii. Petroscolex gen. nov. is the first genus in Eudrilidae with prostatic pore in xiii, anterior to the clitellum. The new genus is also unique in the position of calciferous glands in xiv, not xii or xiii as in other species of the subfamily. Sims (1987) distinguished two groups of genera based on this character, one group with calciferous glands in xii, distributed in West Africa, and another one with calciferous glands in xiii, distributed across tropical Africa. Petroscolex gen. nov. with its calciferous glands in segment xiv is more similar to the latter group because all the genera with backward shifted spermathecal pores possess calciferous glands in xiii. The distribution of the new genus supports this as well, the type locality Irangi, Kivu is at the eastern border of the Congo-basin. A third peculiarity of the new species is the proandric condition of the testes. In Eudrilidae the male apparatus is mostly holandric, although a few genera show metandric reduction: Metascolex Michaelsen, 1903, Polytoreutus Michaelsen, 1890 or Okudrilus Csuzdi & Sherlock, 2015. Petroscolex gen. nov. is the first eudrilid genus with proandric reduction., Published as part of Csuzdi, Csaba, Rota, Emilia, Szederjesi, T��mea, Sherlock, Emma, Brown, George G., Chang, Chih-Han, Cosin, Dario Diaz, Fragoso, Carlos, Jamieson, Barrie G. M., Hong, Yong, James, Samuel W., Paoletti, Maurizio G., Pavl��cek, Tom��s, Plisko, Danuta, Pop, Victor V. & Shen, Huei-Ping, 2019, Description of a new Central African earthworm, Petroscolex centenarius gen et sp. nov. (Crassiclitellata, Eudrilidae), celebrating the 100 th birthday of Pietro Omodeo, pp. 501-508 in Zootaxa 4674 (5) on pages 504-507, DOI: 10.11646/zootaxa.4674.5.1, http://zenodo.org/record/3468756, {"references":["Clausen, M. W. (1965) Description of two new species of Libyodrilus Beddard, 1891, with remarks on the genus and on the female system in the family Eudrilidae. Videnskabelige Meddelelser Dansk Naturhistorisk Forening, 128, 273 - 292.","Jamieson, B. G. M. (1967) A taxonomic review of the African megadrile genus Stuhlmannia (Eudrilidae, Oligochaeta). Journal of Zoology London, 152, 79 - 126. https: // doi. org / 10.1111 / j. 1469 - 7998.1967. tb 01640. x","Sims, R. W. (1969) Internal fertilization in Eudrilid earthworms with the description of a new Pareudriline genus and species (Oligochaeta) from Ghana. Journal of Zoology, 157, 437 - 447. https: // doi. org / 10.1111 / j. 1469 - 7998.1969. tb 01714. x","Sims, R. W. (1987) A review of the Central African earthworm family Eudrilidae (Oligochaeta). In: Bonvicini Pagliai, A. M. & Omodeo, P. (Eds.), On earthworms. Selected symposia and monographs, UZI, 2. Mucchi, Modena, pp. 359 - 388.","Zicsi, A. & Csuzdi, Cs. (1986) Weitere Angaben zur Regenwurmfauna des Kongo-Gebietes (Oligochaeta: Eudrilidae und Glossoscolecidae). Acta Zoologica Hungarica, 32, 385 - 412.","Csuzdi, Cs., Sherlock, E., Talla Kouete, M. & Dohertty-Bone, T. M. (2015) Four new earthworm species from the highlands of Cameroon with description of a new genus Okudrilus gen. n. (Oligochaeta: Eudrilidae & Acanthodrilidae). African Invertebrates, 56, 25 - 38. https: // doi. org / 10.5733 / afin. 056.0103"]}

Petroscolex Csuzdi, Szederjesi & Sherlock, gen. nov. Type species. Petroscolex centenarius sp. nov. Diagnosis. Eudrilinae with single midventral prostatic pore in xiii and single midventral spermathecal pore in xviii. Female pores paired in xiv close to 14/15 near d. Oesophageal gizzard in vi, intestinal gizzards absent. Dorsal blood vessel simple throughout. Paired calciferous glands in xiv and unpaired chylus-sacs in ix, x, xi. Male genital apparatus proandric with sperm reservoirs in x and vesicles in xi. Excretory system holoic, vesiculate. Ovo-spermathecal apparatus paired with a dorsal interconnecting duct. Penial setae lacking. Etymology. The new genus is named in honour of Prof. Pietro Omodeo. Remarks. The new genus is unique among Eudrilidae due to the forward shift of the prostatic pore and the position of calciferous glands in segment xiv., Published as part of Csuzdi, Csaba, Rota, Emilia, Szederjesi, T��mea, Sherlock, Emma, Brown, George G., Chang, Chih-Han, Cosin, Dario Diaz, Fragoso, Carlos, Jamieson, Barrie G. M., Hong, Yong, James, Samuel W., Paoletti, Maurizio G., Pavl��cek, Tom��s, Plisko, Danuta, Pop, Victor V. & Shen, Huei-Ping, 2019, Description of a new Central African earthworm, Petroscolex centenarius gen et sp. nov. (Crassiclitellata, Eudrilidae), celebrating the 100 th birthday of Pietro Omodeo, pp. 501-508 in Zootaxa 4674 (5) on page 504, DOI: 10.11646/zootaxa.4674.5.1, http://zenodo.org/record/3468756

Dicranomyia (Neoglochina) curraniana (Alexander, 1944) Original Genus: Limonia; Holotype: male; Type Locality: Panama, Panama Province, Canal Zone, Barro Colorado Island; Description: (Alexander, 1944q); Wing Reference: Alexander, 1945y; Hypopygium Reference: Alexander, 1945y; Panamanian Provinces: Panama; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 22, DOI: 10.5281/zenodo.3727017

Shannonomyia (Shannonomyia) atroapicalis Alexander, 1934 Original Genus: Shannonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Finca Parada; Description: Alexander, 1934d; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 20, DOI: 10.5281/zenodo.3727017

Atarba (Ischnothrix) patens (Alexander, 1940) Original Genus: Ischnothrix; Holotype: female; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: (Alexander, 1940l); Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 12, DOI: 10.5281/zenodo.3727017

Gonomyia (Paralipophleps) gladiator Alexander, 1919 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Panama Province, Real de Santa Maria; Description: Alexander, 1919k; Wing Reference: Alexander, 1919k; Hypopygium Reference: none; Panamanian Provinces: Darien, Panama; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 16, DOI: 10.5281/zenodo.3727017

Teucholabis (Teucholabis) subleridensis Alexander, 1938 Original Genus: Teucholabis; Holotype: male; Type Locality: Panama, Chiriqui Province, Finca Lerida; Description: Alexander, 1938r; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 18, DOI: 10.5281/zenodo.3727017

Rhipidia (Rhipidia) variicosta (Alexander, 1934) Original Genus: Limonia; Holotype: female; Type Locality: Panama, Chiriqui Province, Finca Parada; Description: (Alexander, 1934d); Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 28, DOI: 10.5281/zenodo.3727017

Orimarga (Orimarga) melampodia Alexander, 1940 Original Genus: Orimarga; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: Alexander, 1940l; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 27, DOI: 10.5281/zenodo.3727017

Gnophomyia banksiana Alexander, 1945 Original Genus: Gnophomyia; Holotype: male; Type Locality: Panama, Panama Province, Canal Zone, Barro Colorado Island; Description: Alexander, 1945y; Wing Reference: none; Hypopygium Reference: Alexander, 1945y; Panamanian Provinces: Panama; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 14, DOI: 10.5281/zenodo.3727017

Dicranomyia (Caenolimonia) translucida (Alexander, 1912) Original Genus: Furcomyia; Holotype: male; Type Locality: Panama, Bocas del Toro Province; Description: (Alexander, 1912g); Wing Reference: Alexander, 1912g; Gelhaus 2009; Hypopygium Reference: none; Panamanian Provinces: Bocas del Toro; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 22, DOI: 10.5281/zenodo.3727017

Gonomyia (Gonomyia) remota obtusistyla Alexander, 1940 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: Alexander, 1940l; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 14, DOI: 10.5281/zenodo.3727017

Teucholabis (Teucholabis) trifasciata trifasciata Enderlein, 1912 Original Genus: Teucholabis; Holotype: female; Type Locality: Colombia, Rio Magdalena; Description: Enderlein, 1912a; Wing Reference: Enderlein, 1912a; Hypopygium Reference: none; Panamanian Provinces: undetermined; Regional Distribution: Colombia, Ecuador, Guyana, Mexico, Panama, Peru, Venezuela., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 18, DOI: 10.5281/zenodo.3727017

Elephantomyia (Elephantomyia) arcuaria Alexander, 1945 Original Genus: Elephantomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: Alexander, 1945y; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 23, DOI: 10.5281/zenodo.3727017

Styringomyia maya Ribeiro, 2003 Original Genus: Styringomyia; Holotype: male; Type Locality: Ecuador, Azuay, 18 km SE Jesus Maria; Description: Ribeiro, 2003; Wing Reference: Alexander, 1914g; Hypopygium Reference: Ribeiro, 2003; Panamanian Provinces: Darien; Regional Distribution: Belize, Bolivia, Colombia, Costa Rica, Ecuador, Guyana, Honduras, Panama, Suriname, Venezuela., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 17, DOI: 10.5281/zenodo.3727017

Ellipteroides (Progonomyia) subcostatus (Alexander, 1919) Original Genus: Gonomyia; Holotype: female; Type Locality: Panama, Panama Province, Rio Chico near mouth of Rio Procona; Description: (Alexander, 1919k); Wing Reference: Alexander, 1919k; Hypopygium Reference: none; Panamanian Provinces: Panama; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 13, DOI: 10.5281/zenodo.3727017

Gonomyia (Leiponeura) macswaini Alexander, 1940 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: Alexander, 1940l; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Costa Rica, Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 15, DOI: 10.5281/zenodo.3727017

Gonomyia (Gonomyia) aitkeniana Alexander, 1979 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Fortuna Dam Site; Description: Alexander, 1979a; Wing Reference: none; Hypopygium Reference: Alexander, 1979a; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 14, DOI: 10.5281/zenodo.3727017

Dicranomyia (Neoglochina) fumosa (Alexander, 1912) Original Genus: Furcomyia; Holotype: female; Type Locality: Guyana, Amatuk; Description: (Alexander, 1912h); Wing Reference: Alexander, 1912h; Alexander 1927o; Hypopygium Reference: none; Panamanian Provinces: Panama; Regional Distribution: Guyana, Panama, Peru, Venezuela., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 22, DOI: 10.5281/zenodo.3727017

Geranomyia recondita recondita Alexander, 1921 Original Genus: Geranomyia; Holotype: male; Type Locality: Peru, Iquitos; Description: Alexander, 1921r; Wing Reference: none; Hypopygium Reference: Alexander, 1954h; Panamanian Provinces: undetermined; Regional Distribution: Colombia, Panama, Puerot Rico., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 25, DOI: 10.5281/zenodo.3727017

Dicranomyia (Neoglochina) trialbocincta (Alexander, 1941) Original Genus: Limonia; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: (Alexander, 1941q); Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 23, DOI: 10.5281/zenodo.3727017

Cheilotrichia (Empeda) brevifida Alexander, 1970 Original Genus: Cheilotrichia; Holotype: male; Type Locality: Panama, Chiriqui Province, El Volc��n; Description: Alexander, 1970c; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 12, DOI: 10.5281/zenodo.3727017

Atarba (Atarba) serena Alexander, 1969 Original Genus: Atarba; Holotype: male; Type Locality: Panama, Chiriqui Province, El Volc��n; Description: Alexan- der, 1969n; Wing Reference: none; Hypopygium Reference: Alexander, 1969n; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 12, DOI: 10.5281/zenodo.3727017

Trentepohlia (Paramongoma) chiriquiana Alexander, 1934 Original Genus: Trentepohlia; Holotype: female; Type Locality: Panama, Chiriqui Province, Finca Parada; Description: Alexander, 1934d; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 28, DOI: 10.5281/zenodo.3727017

Polymera (Polymera) superba Alexander, 1913 Original Genus: Polymera; Holotype: male; Type Locality: Panama, Panama Province, Canal Zone, Tabernilla; Description: Alexander, 1913d; Wing Reference: Alexander, 1913d; Hypopygium Reference: none; Panamanian Provinces: Panama; Regional Distribution: Brazil, Ecuador, Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 20, DOI: 10.5281/zenodo.3727017

Gonomyia (Paralipophleps) peracuta conifera Alexander, 1941 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, Potrerillos; Description: Alexander, 1941q; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Costa Rica, Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 16, DOI: 10.5281/zenodo.3727017

Gonomyia (Gonomyia) sejuncta Alexander, 1970 Original Genus: Gonomyia; Holotype: male; Type Locality: Panama, Chiriqui Province, El Volc��n; Description: Alexander, 1970c; Wing Reference: none; Hypopygium Reference: none; Panamanian Provinces: Chiriqu��; Regional Distribution: Panama., Published as part of Armitage, Brian J. & R��os Gonz��lez, Tom��s A., 2019, The Diptera of Panama. I. Annotated catalog of the Tipulomorpha of Panama, pp. 1-30 in Insecta Mundi 712 on page 14, DOI: 10.5281/zenodo.3727017