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Moravec, Jiří, Razanajaonarivalona, Elisé Hugo, Hori, Michio (2020): New or rare Madagascar tiger beetles-19. Description of Pogonostoma (Pogonostoma) natsuae sp. nov. and elaborated redescription of P. (M.) meridionale Fleutiaux (Coleoptera: Cicindelidae). Zootaxa 4790 (2): 261-276, DOI: https://doi.org/10.11646/zootaxa.4790.2.4

Laterality has been studied in several vertebrates, mainly in terms of brain lateralization and behavioral laterality, but morphological asymmetry has not been extensively investigated. Asymmetry in fishes was first described in scale-eating cichlids from Lake Tanganyika, in the form of bilateral dimorphism in which some individuals, when opening their mouths, twist them to the right and others to the left. This asymmetry has a genetic basis, and is correlated with lateralized attack behaviors. This has subsequently been found in fishes from numerous taxa with various feeding habits. The generality of such morphological laterality should thus be investigated in as wide a range of fishes as possible. Using specific indicators of lateral differences in mandibles and head inclination, we find that representative species from all 60 orders of extant gnathostome fishes (both bony and cartilaginous) possess morphological laterality. Furthermore, we identify the same laterality in agnathans (hagfish and lamprey), suggesting that this trait appeared early in fish evolution and has been maintained across fish lineages. However, a comparison of asymmetry among groups of bony fishes reveals, unexpectedly, that phylogenetically more recent-groups possess less asymmetry in body structures. The universality of laterality in fishes indicates a monophyletic origin, and may have been present in the ancestors of vertebrates. Ecological factors, predator—prey interactions in particular, may be key drivers in the evolution and maintenance of dimorphism, and may also be responsible for the cryptic trend of asymmetry in derived groups. Because lungfish and coelacanths share this trait, it is likely that tetrapods also inherited it. We believe that study of this morphological laterality will provide insights into the behavioral and sensory lateralization of vertebrates.

Calochroa harmandi(Fleutiaux, 1893) (Figs 6C, 10A) Cicindela harmandi Fleutiaux, 1893:494. Calochroa harmandi: Rivalier 1961:126. Specimens examined. (N = 1) [Laos] 1♂ (MHJ), Attapeu Prov. Bolavens Plateau, E Tad Katamtok, 11-12. V. 2010, V. Ryj��ček leg. Description. Internal sac everted dorso-apically in lateral view. DA largest in bladders; VA same as large as VB; B, DLL and VLR slightly swollen; BLR not swollen. VA roundly everted dorso-apically, vested with short spines. VLR roundly swollen dorsally, vested numerously with short and weakly chitinous spines. DA swollen dorso-basally. DLL roundly swollen, vested with fine spines. VB widely and oval everted left latero-ventrally, vested numerously with long spines, which are forming a band. Flagellum very long, curved and formed S-shaped. Medial tooth short and flattened, extending apically. Remarks. This species shares the characteristics of the position of each bladder with Calochroa whithillii but is distinguished by the following characteristics: VA everted dorso-apically; medial tooth flattened., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on page 294, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528, {"references":["Fleutiaux, E. (1893) Remarques sur quelques Cicindelidae et descriptions d'especes nouvelles. Annales de la Societe Entomologique de France, 62, 483 - 502.","Rivalier, E. (1961) Demembrement du genre Cicindela Linne, IV. Faune indomalaise. Revue francaise d'Entomologie, 28, 121 - 149."]}

Cicindela(Cicindela) gemmata ainoLewis, 1891 (Figs 8G, 15E) Cicindela gemmata aino Lewis, 1891:20. Cicindela gemmata shikokuana Oike, 1939:15 [synonymy by Cassola & Nidek 1984:9]. Cicindela (Cicindela) gemmata Faldermann, 1835: Rivalier 1950:224 (only nominotypical subspecies). Specimens examined. (N = 2) [Japan] 1♂ (YFJ), Tottori, Sugasaki, K��fu-ch��, 4. V. 2013, Y. Fukuda leg; 1♂ (YFJ), Tottori, Sugasaki, K��fu-ch��, 6. V. 2013, Y. Fukuda leg. Description. Internal sac widely everted dorso-apically in lateral view. VA largest in bladders; DA and VLR the second largest; B and VB the third largest; DLL and VLL smallest in bladders. BLR not exist. DLL roundly swollen, vested with fine spines. DA well swollen basally. VA long and cylindrically swollen, everted apically, vested numerously with short and weakly chitinous spines. VB roundly swollen left laterally, vested numerously with short spines. VLR cylindrically swollen left laterally, with weakly chitinous spines. VLL roundly swollen right laterally, vested with short and weakly chitinous spines. B oval and roundly swollen, everted right laterally, numerously with short spines, which are forming a band. Flagellum stout and well short, curved. Medial tooth flattened, extending ventrally. Remarks. This species is clearly distinguished from other members of the subgenus Cicindela by sharing the characteristics of the position of each bladder but differs by the following characteristics: VA long and cylindrically swollen; VLR everted left laterally., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on pages 300-301, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528, {"references":["Lewis, G. (1891) Note on a new Cicindela from north Japan. The Entomologist's Monthly Magazine, 27, 20.","Oike, K. (1939) Cicindela gemmata shikokuana subsp. nov. Transactions of the Kansai Entomological Society, 8, 15 - 16.","Cassola, F. & Brouerius van Nidek, C. M. C (1984) Checklist of Cicindela (s. auct.) of the Palaearctic region (Coleoptera: Cicindelidae). Cicindela, 16, 7 - 17.","Faldermann, F. (1835) Coleopterorum ab illustrissimo Bungio in China boreali, Mongolia, et montibus Altaicis collectorum, nec non ab ill. Turczaninoffio et Stchukino e provincia Irkutzk missorum illustrationes. Memoires presentes a l'Academie Imperiale des Sciences de Saint-Petersbourg, 2, 337 - 464., 5 pls.","Rivalier, E. (1950) Demembrement du genre Cicindela Linne (Travail preliminaire limite a la faune palearctique). Revue francaise d'Entomologie, 17, 249 - 268."]}

Cosmodelasp. cf. separata (Figs 8A, 13A) Specimens examined. (N = 1) [Vietnam] 1♂ (MHJ), Lao Cai Prov., Sapa, 20-23. V. 2012, M. Hori leg. Description. Internal sac everted widely and dorsally in lateral view. DA largest in bladders; AL the second largest; B same as large as VA, A, DLL, VLL and VLR. A thick and cylindrically everted apically. AL roundly right ventro-laterally swollen, vested with fine spines. Medial tooth short, extending ventrally, weakly curved at apex. DLL roundly swollen, vested with fine spines. VA short triangular swollen, everted dorsa-apically. VLR oval and weakly swollen left laterally at base of VA. VLL widely swollen right laterally, with well sclerotized spines, which are forming a narrow band. B slightly and roundly swollen, everted right laterally, numerously with long spines. Flagellum very long, curved and formed S-shaped. Medial tooth short, extending ventrally, weakly curved at apex. Remarks. This species is clearly distinguished from other members of Cosmodela by the following characteristics: A and VA short; VB non-existent; AL everted ventro-laterally., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on page 287, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528

Calochroa lacrymans(Schaum, 1863) (Figs 6H, 10D) Cicindela lacrymans Schaum, 1863:57. Calochroa discrepans lacrymans: Fowler 1912:390. Calochroa lacrymans: Naviaux 1986:61. Cicindela (Calochroa) lacrymans: Acciavatti & Pearson 1989:129. Calochroa lacrymans: Wiesner 1992:108. Specimens examined. (N = 1) [Sri Lanka] 1♂ (MHJ), Kandy, VIII. 1990, unknown leg. Description. Internal sac everted dorso-apically in lateral view. DA largest in bladders; VA the second largest; VLR, VB, DLL slightly swollen; BLR and B not swollen. VA shortly and roundly everted right dorso-apically, vested with short and weakly chitinous spines. VLR roundly everted dorsally. DA semicircular swollen right latero-basally. DLL roundly swollen, vested numerously with spines, which are weakly chitinous. VB slightly and roundly swollen left latero-ventrally, vested numerously with long spines, which are forming a spot. Flagellum very long and thin, S-shaped. Medial tooth extending apically, weakly curved at apical 1/3. Remarks. This species is clearly distinguished from other members of Calochroa by the characteristics lacking B., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on page 292, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528, {"references":["Schaum, H. R., (1863) Contributions to the knowledge of the Cicindelidae of tropical Asia, containing descriptions of new species, a list of those hitherto described, and synonymical notes. The Journal of Entomology. Descriptive and Geographical, 2, 57 - 73.","Fowler, W. W. (1912) The fauna of British India including Ceylon and Burma. Coleoptera. General introduction and Cicindelidae and Paussidae, Taylor and Francis, London, 529 pp. https: // doi. org / 10.5962 / bhl. title. 83566","Naviaux, R. (1986) Coleoptera, Cicindelidae. Premier complement aux Cicindeles du Sri Lanka. Revue Scientifique Du Bourbonnais, 58 - 69.","Acciavatti, R. E. & Pearson, D. L. (1989) The tiger beetle Genus Cicindela (Coleoptera, Insecta) from the indian subcontinent. Annals of Carnegie Museum, 58, 77 - 353.","Wiesner, J. (1992) Checklist of the Tiger Beetles of the World. Verlag Erna Bauer, Keltern, 364 pp."]}

Cosmodela aurulenta(Fabricius, 1801) (Figs 8C, 13B) Cicindela aurulenta Fabricius, 1801:239. Cosmodela aurulenta aurulenta: Rivalier 1961:128. Cosmodela aurulenta: Lopez-Lopez et al. 2015:8. Specimens examined. (N = 4) [Malaysia] 2♂♂ (YFJ), Is. Penang, Penang National Park, 2. III. 2012, Y. Fukuda leg; [Indonesia] 2♂♂ (EUMJ), Yogyakarta, Mt. Merapi, Kaliurang, Java Is., 1. III. 2013, R. Ogawa leg. Description. Internal sac everted widely and dorsally in lateral view. DA largest in bladders; B, VLL and A the second largest; DLL and VA the third largest; BLR, VLR, AL and VB smallest in bladders. A cylindrically everted apically. AL and VB slightly bulging at base of A, vested with fine spines, which are forming oblong spots. DA well swollen basally. DLL roundly swollen, vested numerously with spines, which are weakly chitinous. VA triangular swollen at base of DA, everted dorso-apically, tapered. BLR roundly swollen left ventrally, vested numerously with well sclerotized spines, which are forming an oblong spot. VLR swollen left laterally at base of VA. VLL oval and widely swollen right laterally, with well sclerotized spines, which are forming a narrow band. B oval and widely swollen right laterally, with well sclerotized spines, which are forming an oblong spot. Flagellum very long, curved and formed S-shaped. Medial tooth short, extending apically. Remarks. This species mostly shares the characteristics of the position of each bladder with C. batesi but is distinguished from the latter by VA and A not long., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on pages 286-287, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528, {"references":["Fabricius J. C. (1801) Systema Eleutheratorum secundum ordines, genera, species; adiectis synonymis, locis, observationibus, descriptionibus. Tomus I & II. Bibliopolii Academici Novi, Kiliae, 226 pp.","Rivalier, E. (1961) Demembrement du genre Cicindela Linne, IV. Faune indomalaise. Revue francaise d'Entomologie, 28, 121 - 149.","Lopez-Lopez, A., Abdul, A. & Galian, J. (2015) Molecular phylogeny and divergence time estimation of Cosmodela (Coleoptera: Carabidae: Cicindelinae) tiger beetle species from Southeast Asia. Zoologica Scripta, 44 (4), 437 - 445. https: // doi. org / 10.1111 / zsc. 12113"]}

Sophiodela chinensis chinensis(DeGeer, 1774), comb. nov. (Figs 7A, 12D) Cicindela chinensis DeGeer, 1774: 119. Cicindela sinensis [sic]: Wood 1874: 16 [incorrect subsequent spelling]. Cicindela (Sophiodela) chinensis: Nakane 1955: 25. Cicindela (Sericina) chinensis: Rivalier 1961:123 [synonymy by Nakane 1976: 3]. Specimens examined. (N = 1) [China] 1♂ (MHJ), Hunan, Huaishua, VII. 1996, unknown leg. Description. Internal sac everted dorsally in lateral view. DA largest in the bladders; DLL the second largest; VA as large as B. VLR, BLR and VB not exist. VA short triangular everted apically. DA everted basally. B roundly swollen ventrally, with well sclerotized spines, which are forming a narrow band. DLL swollen dorsally, covered numerously with well sclerotized spines, which are a forming band. Flagellum well short, curved and formed S shape. Medial tooth short, gently curved right ventro-laterally at apical 1/3. Remarks. Sophiodela chinensis flammifera (Horn, 1920) distributed in South Korea and North Korea is related to this species. The structures of the male internal sac of this subspecies indicated possible synonymy with S. c. chinensis, but the former has a strong reddish body color and is slightly larger than the latter in body size. Sophiodela chinensis chinensis is almost the same size as Sophiodela japonica and apparently different from S. c. flammifera. Further research would be required to test the validity of this possibility. Sophiodela chinensis flammifera (Horn, 1920) , comb. nov. (Figs 7E, 12E) Cicindela chinensis flammifera Horn, 1920: 19. Cicindela (Sophiodela) chinensis flammifera: Cassola & Nidek 1984:11. Specimens examined. (N = 2) [North Korea] 1♂ (MHJ), Hamgyeongbuk-do, Kwangmo-pong, 3–6. VII. 1994, un- known leg; [Korea] 1♂ (ATJ), Gyeonggi-do, Gwangju-si, Docheok-myeon, Sangnim-ri, Seoul National University Experiment forest, Mt. Taehwasan, 23–25. VI. 2006, A. Tamura leg. Description. Internal sac everted dorsally in lateral view. DA largest in the bladders; DLL the second largest; VA as large as B. VLR, BLR and VB not exist. VA short triangular with slightly dimple, everted apically. DA everted basally. B roundly swollen ventrally, with well sclerotized spines, which are forming a narrow band. DLL swollen dorsally, covered numerously with well sclerotized spines, which are a forming band. Flagellum well short, curved and formed S shape. Medial tooth short, gently curved right ventro-laterally at apical 1/3., Published as part of Fukuda, Yuki, Ogawa, Ryo & Hori, Michio, 2019, The reclassification of Sophiodela and other tiger beetles (Coleoptera, Cicindelidae) based on the structure of the everted internal sac of the male genitalia, pp. 271-308 in Zootaxa 4661 (2) on pages 282-284, DOI: 10.11646/zootaxa.4661.2.3, http://zenodo.org/record/3998528, {"references":["DeGeer, C. (1774) Memoires pour servir a l'histoire des insectes and quadrieme by quatrieme. Tome quadrieme. Hesselberg, Stockholm, xii + 457 pp.","Wood, J. G. (1874) Insects abroad, being a popular account of foreign insects, their structure, habits, and transformations. Longmans, Green, and Co., London, xiii + 648 pp. https: // doi. org / 10.5962 / bhl. title. 7906","Nakane, T. (1955) New or little-known Coleoptera from Japan and its adjacent regions, XII. The Scientific Report of the Saikyo University, 2 (1), 24 - 40.","Rivalier, E. (1961) Demembrement du genre Cicindela Linne, IV. Faune indomalaise. Revue francaise d'Entomologie, 28, 121 - 149.","Nakane, T. (1976) Check-list of Coleoptera of Japan, Cicindelidae. The Coleopterists' Association of Japan, 3, 1 - 7.","Horn, W. (1920) Cicindelinen-Studien aus dem schwedischen Reichsmuseum nebst einigen Bemerkungen neuer Arten etc. Arkiv for Zoologi, 13 (11), 1 - 21.","Cassola, F. & Brouerius van Nidek, C. M. C (1984) Checklist of Cicindela (s. auct.) of the Palaearctic region (Coleoptera: Cicindelidae). Cicindela, 16, 7 - 17."]}