Abstract
Two new members of Tridactylidae Brullé, 1835 are described based on specimens found in mid-Cretaceous Kachin amber from Northern Myanmar. Duappendactylus melanocephalus gen. et sp. nov. is assigned to the family based on its two-segmented cerci and inflated mesotibia. It is similar to Birmitoxya Gorochov, 2010 by its lack of (visible) paraproctal processes and the presence of an elongated, upward-facing subgenital plate, but differs from that genus in several other aspects of its morphology. Additionally, we describe Hiemalidactylus eileenae gen. et sp. nov. based on a well preserved, presumably nymphal specimen. It is assigned to Tridactylidae based on the two-segmented cerci and the inflated mesotibia. Like the two aforementioned genera, Hiemalidactylus does not have cerciform paraproctal processes. However, this genus differs from Birmitoxya and Duappendactylus by not having an elongated subgenital plate. Instead, the paraproct is elongated and reaches almost the length of the basal segment of the cercus. These new species of Tridactylidae raise the diversity of this family known from Kachin amber to nine genera and eleven species.
References
- Ander, K. (1936) Orthoptera Saltatorias fylogeni pa grundval av amforande anatomiska studier. In: Kemner, N.A. (Ed.), Det femte Nordiska Entomologmetet i Lund 3–6 augusti 1936. Opuscula Entomologica. Edidit Societas Entomologica Lundensis, 1, pp. 93–94.
- Ander, K. (1939) Vergleichend-anatomische und phylogenetische Studien über die Ensifera (Saltatoria). Opuscula Entomologica. Edidit Societas Entomologica Lundensis, 2 (Supplement), 306, 172 figs.
- Baena-Bejarano, N., Heads, S.W. & Taylor, S.J. (2018) Comments on the neglected nymphs of mud crickets in the genus Mirhipipteryx (Caelifera: Tridactyloidea: Ripipterygidae). Zootaxa, 4486 (2), 180–188. https://doi.org/10.11646/zootaxa.4486.2.7
- Brullé, G.A. (1835) Cinquième Ordre. Orthoptères. In: Histoire naturelle des insectes: Vol. 9 [1] (5). Société bibliophile, Paris, pp. 1–225. [225–416 in 1836]
- Cao, C.Q., Chen, S.Z. & Yin, Z. (2019) A new genus and a new species of pygmy mole cricket in Cretaceous amber from Burma (Orthoptera: Tridactylidae). Zootaxa, 4559 (1), 193–195. https://doi.org/10.11646/zootaxa.4559.1.12
- Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2025) Orthoptera Species File. Available from: http://orthoptera.speciesfile.org/
- de Saussure, H. (1896) Orthoptera. Fam. Gryllidae. Biologia Centrali-Americana, 1, 201–216.
- Deyrup, M. (2005) A new species of flightless pygmy mole cricket from a Florida sand ridge (Orthoptera: Tridactylidae). Florida Entomologist, 88 (2), 141–145. https://doi.org/10.1653/0015-4040(2005)088[0141:ANSOFP]2.0.CO;2
- Du, X.H., Xu, C.P. & Zhang, L.Y. (2022) Two new pygmy mole crickets from mid-Cretaceous Kachin amber of northern Myanmar (Orthoptera: Tridactyloidea: Tridactylidae). Cretaceous Research, 129, 105002. https://doi.org/10.1016/j.cretres.2021.105002
- Engel, M.S. (2020) Myanmar: paleontologists must stop buying conflict amber. Nature, 584 (7822), 525. https://doi.org/10.1038/d41586-020-02432-z
- Fan, S., Gu, J.J. & Cao, C.Q. (2023) A new species of the genus Burmadactylus Heads, 2009 from mid-Cretaceous amber in north Myanmar (Orthoptera: Caelifera: Tridactyloidea). Zootaxa, 5306 (5), 595–598. https://doi.org/10.11646/zootaxa.5306.5.7
- Giglio-Tos, E. (1914) Sulla posizione sistematica del gen. Cylindracheta Kirby. Annali del Museo Civico di Storia Naturale di Genova, 46, 81–101. https://doi.org/10.5962/bhl.part.14781
- The GIMP Team (2025) GIMP. Version 3.0.2-1. Available from: https://www.gimp.org/ (accessed 15 July 2025)
- Gorochov, A.V. (1992) New fossil Orthoptera and Phasmoptera from the Mesozoic and Cainozoic of Mongolia [in: New taxa of fossil invertebrates from Mongolia]. Trudy - Sovmestnaya Sovetsko-Mongol’skaya Paleontologicheskaya Ekspeditsiya [Transaction - Joint Russian-Mongolian Paleontological Expedition], 41, 117–121.
- Gorochov, A.V. (1994) New data on Triassic Orthoptera from Middle Asia. Zoosystematica Rossica, 3 (1), 53–54, illustr.
- Gorochov, A.V. (2005) Review of Triassic Orthoptera with descriptions of new and little known taxa: Part 2. Paleontologicheskiy Zhurnal, 3, 46–54.
- Gorochov, A.V., Jarzembowski, E.A. & Coram, R.A. (2006) Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England. Cretaceous Research, 27, 641–662. https://doi.org/10.1016/j.cretres.2006.03.007
- Gorochov, A.V. (2010) New and little-known orthopteroid insects (Polyneoptera) from fossil resins: Communication 4. Paleontological Journal, 44, 657–671. https://doi.org/10.1134/S0031030110060080
- Gu, J.J., Zheng, C.J., Ren, D., Cao, C.Q. & Yue, Y.L. (2022) Two New Species of Ripipterygidae (Orthoptera, Tridactyloidea) from Mid-Cretaceous of Myanmar with a Key to the Genera of Tridactyloidea in Amber. Insects, 13 (11), 979. https://doi.org/10.3390/insects13110979
- Günther, K.K. (1979) Einige Bemerkungen über die Gattungen der Familie Tridactylidae Brunner und zur Klassifikation der Tridactylodea (Orthopteroida, Caelifera). Deutsche Entomologische Zeitschrift, 26 (4–5), 255–264. https://doi.org/10.1002/mmnd.19790260408
- Haug, J.T., Azar, D., Ross, A.J., Szwedo, J., Wang, B., Arillo, A., Baranov, V.A., Bechteler, J., Beutel, R.G., Blagoderov, V.A., Delclòs, X., Dunlop, J.A., Feldberg, K., Feldmann, R.M., Foth, C., Gehler, A., Harms, D., Hedenäs, L., Hyžný, M., Jagt, J.W.M., Jagt-Yazykova, E.A., Jarzembowski, E.A., Kerp, H., Khine, P.K., Kirejtshuk, A.G., Klug, C., Kopylov, D.S., Kotthoff, U., Kriwet, J., McKellar, R.C., Nel, A.N., Neumann, C., Nützel, A., Peñalver, E., Perrichot, V., Pint, A., Ragazzi, E., Regalado, L., Reich, M., Rikkinen, J.K., Sadowski, E.M., Schmidt, A.R., Schneider, H., Schram, F.R., Schweigert, G., Selden, P.A., Seyfullah, L.J., Solórzano-Kraemer, M.M., Stilwell, J., van Bakel, B.W.M., Vega-Vera, F.J., Wang, Y., Xing, L., & Haug, C. (2020) Comment on the letter of the Society of Vertebrate Paleontology (SVP) dated April 21, 2020 regarding “Fossils from conflict zones and reproducibility of fossil-based scientific data”: Myanmar amber. Paläontologische Zeitschrift, 94, 431–437. https://doi.org/10.1007/s12542-020-00524-9
- Heads, S.W. (2009) A new pygmy mole cricket in cretaceous amber from Burma (Orthoptera: Tridactylidae). Denisia, 26 (2 April 2009), 75–82.
- Heads, S.W. (2010) New Tridactyloidea in Miocene amber from the Dominican Republic (Orthoptera: Caelifera). Annales de la Société Entomologique de France, 46 (1–2), 204–210. https://doi.org/10.1080/00379271.2010.10697659
- Houston, T.F. (2007) Observations of the biology and immature stages of the sandgroper Cylindraustralia kochii (Saussure), with notes on some congeners (Orthoptera: Cylindrachetidae). Records-Western Australian Museum, 23 (3), 219. https://doi.org/10.18195/issn.0312-3162.23(3).2007.219-234
- The Inkscape Team (2025) Inkscape. Version 1.3.2. Available from: https://inkscape.org/de/ (accessed 15 July 2025)
- Latreille, P.A. (1809) Histoire Naturelle, genérale et particuliere, des Crustacés et des Insectes, 4. F. Dufart, F., Paris, 394 pp.
- Martins-Neto, R.G. (1990) Um novo genero e duas novas especies de Tridactylidae (Insecta, Orthopteridea) na Formacao Santana (Cretaceo Interior do nordeste do Brasil). Anais da Academia Brasileira de Ciências, 62 (1), 51–59.
- Olivier, A.G. (1789) Encyclopedie methodique, dictionnaire des insectes, Vol. 4. Paris: Pankouke, 373, 331.
- Peretti, A. (2021) An alternative perspective for acquisitions of amber from Myanmar including recommendations of the United Nations Human Rights Council. Journal of International Humanitarian Action, 6 (1), 12. https://doi.org/10.1186/s41018-021-00101-y
- Poinar, G. (2020) Pygmy mole crickets (Orthoptera: Tridactylidae) in Dominican and Burmese amber. Historical Biology. https://doi.org/10.1080/08912963.2018.1483362
- Schall, O.K.O., Cumming, R., Lian, Z. & Husemann, M. (2025) A new species of Burmecaelidae (Orthoptera: Caelifera) sheds light on the family’s possible taxonomic placement. Zootaxa, 5636 (1), 174–182. https://doi.org/10.11646/zootaxa.5636.1.8
- Scudder, S.H. (1902) On the United States Orthoptera which have been referred to the genus Tridactylus. Psyche, a Journal of Entomology, 9 (310), 308–310. https://doi.org/10.1155/1902/519583
- Sharov, A.G. (1968) Filogniya orthopteroidnykh nasekomykh [1971 English translation: Phylogeny of the Orthopteroidea]. Trudy Paleontologieskogo Instituta Akademii Nauk SSSR, 118, 1–216.
- Shi, G.G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M.C., Lei, W.Y., Li, Q. & Li, X.H. (2012) Age constraint on Burmese amber based on U-Pb dating of zircons. Cretaceous Research, 37, 155–163. https://doi.org/10.1016/j.cretres.2012.03.014
- Song, H., Amédégnato, C., Cigliano, M.M., Desutter‐Grandcolas, L., Heads, S.W., Huang, Y., Otte, D. & Whiting, M.F. (2015) 300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling. Cladistics, 31 (6), 621–651. https://doi.org/10.1111/cla.12116
- Targioni-Tozzetti, A. (1882) In Ortotteri agrari cioè dei diversi insetti dell’ordine degli ortotteri nocivi o vantaggiosi all’agricoltura o all’economia domestica e principalmente delle cavallette. Annali di Agricoltura, Firenze, Bencini, 238 pp., 68 figs.
- Uchida, K., Husemann, M. & Kotthoff, U. (2024) A new Cretaceous orthopteran family of the Caelifera (order: Orthoptera) from Burmese amber of northern Myanmar. PalZ, 98 (1), 117–125. https://doi.org/10.1007/s12542-023-00670-w
- Woo, B. (2021) A new species of pygmy mole cricket (Orthoptera: Tridactylidae) from the Lake Wales ridge of Florida and new records of Ellipes eisneri from the northern Brooksville ridge. Journal of Orthoptera Research, 30 (2), 131–143. https://doi.org/10.3897/jor.30.65603
- Xing, L. & Qiu, L. (2020) Zircon UPb age constraints on the mid-Cretaceous Hkamti amber biota in northern Myanmar. Palaeogeography, Palaeoclimatology, Palaeoecology, 558, 109960. https://doi.org/10.1016/j.palaeo.2020.109960
- Xu, C.P., Fang, Y. & Jarzembowski, E.A. (2020) A new pygmy mole cricket (Orthoptera: Tridactyloidea: Tridactylidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 111, 104371. https://doi.org/10.1016/j.cretres.2020.104371
- Xu, C.P., Wang, B., Fan, L., Jarzembowski, E.A., Fang, Y., Wang, H., Li, T., Zhuo, D., Ding, M. & Engel, M.S. (2022) Widespread mimicry and camouflage among mid-Cretaceous insects. Gondwana Research, 101, 94–102. https://doi.org/10.1016/j.gr.2021.07.025
- Zhao, J.Y., Xu, C.P., Cao, C.Q., Jarzembowski, E.A., Fang, Y. & Xiao, C.T. (2024) A new genus and species of mud cricket (Orthoptera: Ripipterygidae) from mid-Cretaceous Kachin amber of northern Myanmar. Cretaceous Research, 160, 105880. https://doi.org/10.1016/j.cretres.2024.105880
- Zheng, C.J., Cao, C.Q. & Gu, J.J. (2023) A new genus and a new species of pygmy mole cricket from mid-Cretaceous amber in North Myanmar (Orthoptera: Caelifera: Tridactyloidea). Zootaxa, 5339 (3), 296–300. https://doi.org/10.11646/zootaxa.5339.3.7