Elcanidae Explained

Elcanidae are an extinct family of Mesozoic and early Cenozoic orthopterans. Members of the family are distinguished by the presence of spurs on the distal part of the metatibia, unique among orthopterans, these have been suggested to have been used for controlling gliding,^[1] swimming aids,^[2] or for jumping on water.^[3] The group combines characteristics from both major groups of orthopterans, with long antennae and nymphal morphology similar to Ensifera, but with wing venation and adult morphology more similar to Caelifera. Elcanidae is part of Elcanoidea, which is thought to have diverged from living orthopterans by the beginning of the Permian, around 300 million years ago.^[4] The family also includes Permelcanidae, known from the Early-Late Permian. The relationship of Elcanoidea to Ensifera and Caelifera is currently unresolved. Elcanids are known from the Late Triassic to Paleocene of Eurasia, North and South America. Some members of the group exhibited aposematic coloration. They are thought to have been herbivorous.

thumb|right|upright|Sinoelcana minuta
Jurassic, Chinathumb|right|upright|Elcanonympha diana nymphthumb|right|upright|Wings of Parelcana pulchmacula

Taxonomy

• subfamily Archelcaninae Gorochov, 2011^[5]
  • Archelcana Sharov, 1968^[6]
    • Archelcana durnovaria Whalley, 1985^[7] – Charmouth Mudstone Formation, England, Early Jurassic (Sinemurian)
    • Archelcana ornata Zherikhin, 1985^[8] – Cheremkhovskaya Formation, Russia, Early Jurassic (Toarcian)
    • Archelcana shurabica Sharov, 1968 – Sagul Formation, Kyrgyzstan, Toarcian
    • Indeterminate remains attributed to the genus are also known from the latest Triassic of Germany^[9]
  • Jeholelcana Fang et al. 2018b^[10]
    • Jeholelcana yanensis Fang et al. 2018b – Yixian Formation, China, Early Cretaceous (Aptian)
  • Parelcana Handlirsch, 1906^[11]
    • Parelcana anglicana Handlirsch, 1939^[12] – Blue Lias, United Kingdom, Early Jurassic (Hettangian)
    • Parelcana dubia Handlirsch, 1939 – United Kingdom, Late Triassic (Rhaetian)
    • Parelcana pulchmacula Tian et al., 2019^[13] – Daohugou, China, Middle Jurassic (Callovian)
    • Parelcana tenuis Handlirsch, 1906 – Green Series, Germany, Toarcian
  • Sibelcana Gorochov, 1990^[14]
    • Sibelcana rossica Gorochov, 1990 – Glushkovo Formation, Russia, Late Jurassic (Tithonian)
    • Sibelcana transbaicalica Gorochov, 1990 – Ukurei Formation, Russia, Tithonian
  • Synelcana Zessin, 1988^[15]
    • Synelcana muelleri Zessin, 1988 – Green Series, Germany, Toarcian
• subfamily Elcaninae Handlirsch, 1906
  • Cratoelcana Martins-Neto, 1991^[16] – Crato Formation, Brazil, Aptian
    • Cratoelcana damianii Martins-Neto, 1991
    • Cratoelcana zessini Martins-Neto, 1991
  • Ellca Kočárek, 2020
    • Ellca nevelka Kočárek, 2020 – Burmese amber, Myanmar, Late Cretaceous (Cenomanian)
  • Eubaisselcana Gorochov, 1986^[17] – Gurvan-Eren Formation, Mongolia, Aptian
    • Eubaisselcana mongolica Gorochov, 1986
    • Eubaisselcana sharovi Gorochov, 1986
  • Minelcana Gorochov et al., 2006^[18]
    • Minelcana dubia (Giebel 1856)^[19] – Lulworth Formation, United Kingdom, Early Cretaceous (Berriasian)
    • Minelcana membranacea Gorochov et al., 2006 – Durlston Formation, United Kingdom, Berriasian
  • Panorpidium Westwood, 1854^[20]
    • Panorpidium bimaculatum Gorochov et al., 2006 – Weald Clay, United Kingdom, Early Cretaceous (Barremian)
    • Panorpidium maculosum Zhou et al. 2022^[21] Burmese amber, Myanmar, Cenomanian
    • Panorpidium proximum Gorochov et al., 2006 – Durlston Formation, United Kingdom, Berriasian
    • Panorpidium parvum Gorochov et al., 2006 – Weald Clay, United Kingdom, Early Cretaceous (Hauterivian)
    • Panorpidium sibirica Sharov, 1968 – Zaza Formation, Russia, Aptian
    • Panorpidium spica Kim et al., 2021 – Jinju Formation, South Korea, Early Cretaceous (Albian)
    • Panorpidium tessellatum Westwood, 1854 – Durlston Formation, United Kingdom, Berriasian
    • Panorpidium yixianensis Fang et al., 2015^[22] – Yixian Formation, China, Aptian
  • Probaisselcana Gorochov, 1989^[23]
    • Probaisselcana cretacea Gorochov et al., 2006 – Weald Clay, United Kingdom, Hauterivian
    • Probaisselcana euryptera Tian et al., 2019 – Yixian Formation, China, Aptian
    • Probaisselcana karatavica Sharov, 1968 – Karabastau Formation, Kazakhstan, Late Jurassic (Oxfordian)
    • Probaisselcana oculata Hu & He, 2023^[24] - Burmese amber, Myanmar, Cenomanian
• Incertae sedis
  • Burmelcana Peñalver & Grimaldi, 2010^[25]
    • Burmelcana longirostris Peñalver & Grimaldi, 2010 – Burmese amber, Myanmar, Cenomanian
  • Cascadelcana Fang et al., 2018a^[26]
    • Cascadelcana virginiana Fang et al., 2018a – Cow Branch Formation, North Carolina, Late Triassic (Norian)
  • Cenoelcanus Schubnel et al., 2020^[27]
    • Cenoelcanus menatensis Schubnel et al., 2020 – Menat Formation, France, Paleocene
  • Elcanonympha Heads et al., 2018^[28]
    • Elcanonympha diana Heads et al., 2018 – Burmese amber, Myanmar, Cenomanian
  • Hispanelcana Peñalver & Grimaldi, 2010 – Álava amber, Escucha Formation, Spain, Albian
    • Hispanelcana alavensis Peñalver & Grimaldi, 2010
    • Hispanelcana arilloi Peñalver & Grimaldi, 2010
    • Hispanelcana lopezvallei Peñalver & Grimaldi, 2010
  • Longioculus Poinar et al., 2007^[29]
    • Longioculus burmensis Poinar et al., 2007 – Burmese amber, Myanmar, Cenomanian
  • Hukawnelca Uchida, 2021^[30]
    • Hukawnelca gracile Uchida, 2021 – Burmese amber, Myanmar, Cenomanian
  • Caelielca Uchida, 2021
    • Caelielca spinocrus Uchida, 2021 – Burmese amber, Myanmar, Cenomanian
  • Monitelcana Xu et al., 2022^[31]
    • Monitelcana penalveri Xu et al., 2022 – Burmese amber, Myanmar, Cenomanian

Notes and References

1. Kočárek P . December 2020 . A diminutive elcanid from mid-Cretaceous Burmese amber, Ellca nevelka gen. et sp. nov., and the function of metatibial spurs in Elcanidae (Orthoptera) . Cretaceous Research . 116 . 104574 . 10.1016/j.cretres.2020.104574 . 2020CrRes.11604574K . 225028485 .
2. Tian H, Gu JJ, Huang F, Zhang H, Ren D . July 2019 . A new species of Elcaninae (Orthoptera, Elcanidae) from the Lower Cretaceous Yixian Formation at Liutiaogou, Inner Mongolia, NE China, and its morphological implications . Cretaceous Research . 99. 275–280. 10.1016/j.cretres.2019.03.010. 2019CrRes..99..275T . 134966211 .
3. Kim. Do-Yoon. Lee. Mirinae. Nam. Gi-Soo. Park. Tae-Yoon S.. April 2021. The first orthopteran fossils from the Lower Cretaceous (Albian) Jinju Formation of Korea: ethological implications for elcanids. Cretaceous Research. 125. en. 104843. 10.1016/j.cretres.2021.104843. 2021CrRes.12504843K . 0195-6671 .
4. Poschmann . Markus J. . Nel . André . Raisch . Manfred . 2023-05-31 . A new archaeorthopteran insect from the Early Permian of the Saar-Nahe Basin, SW Germany, and the earliest record of Elcanoidea (Orthoptera) . Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen . en . 308 . 2 . 181–187 . 10.1127/njgpa/2023/1136 . 0077-7749.
5. Gorochov AV . 2011 . New family-group taxa in the infraclass Polyneoptera (Insecta): making the names, proposed early, available. . Zoosystematica Rossica . 20 . 159–160 . 10.31610/zsr/2011.20.1.159 .
6. Sharov AG . 1968 . Filogeniya ortopteroidnykh nasekomykh . Phylogeny of orthopteroid insects . ru. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR . 118 . 1–216 .
7. Whalley PE . 1985 . The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. . Bulletin of the British Museum (Natural History) Geology . 39 . 3 . 107–189 .
8. Zherikhin VV . 1985 . Yurskie pryamokrylye yuzhnoy Sibiri i zapadnoy Mongolii (Gryllida=Orthoptera). . Jurassic Orthoptera of southern Siberia and western Mongolia (Grillida = Orthnoptera). . ru. Yurskie Nasekomye Sibiri I Mongolii . 171–184 .
9. Barth G, Ansorge J, Brauckmann C . December 2011 . First record of the genus Ipsvicia (Hemiptera: Ipsviciidae) outside Gondwana - an Australian genus from the Upper Triassic of Germany . Polish Journal of Entomology / Polskie Pismo Entomologiczne. 80. 4. 645–657. 10.2478/v10200-011-0050-z. 84850244. 0032-3780. free.
10. Fang Y, Heads SW, Wang H, Zhang H, Wang B . August 2018 . The first Archelcaninae (Orthoptera, Elcanidae) from the Cretaceous Jehol Biota of Liaoning, China . Cretaceous Research. en. 86. 129–134. 10.1016/j.cretres.2018.02.008. 2018CrRes..86..129F . 134206028 .
11. Book: Handlirsch A . 1906 . Die Fossilen Insekten und die Phylogenie der Rezenten Formen, parts I-IV. . The fossil insects and the phylogeny of the recent forms . Ein Handbuch fur Palaontologen und Zoologen . A manual for palaontologists and zoologists . de . 1–640 .
12. Handlirsch A . 1939 . Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. II Teil. . New investigations into the fossil insects with additions and supplements as well as perspectives on phylogenetic, palaeogeographical and general biological problems. II part. . de. Annalen des Naturhistorischen Museums in Wien . Annals of the Natural History Museum in Vienna . 49 . 1–240 .
13. Tian H, Gu JJ, Yin XC, Ren D . The first Elcanidae (Orthoptera, Elcanoidea) from the Daohugou fossil bed of northeastern China . ZooKeys . 897 . 19–28 . 2019-12-09 . 31857785 . 6914707 . 10.3897/zookeys.897.37608 . free . 2019ZooK..897...19T .
14. Gorochov V . 1990 . Orthopterans, Gryllida, in Pozdne-Mezozoyskie Nasekomye Vostochnogo Zabaykal'ya. . Orthnopterans, Grillida, in Late Mesozoic Insects of Eastern Transbaikalia . ru . Akademiya Nauk SSSR, Trudy Paleontologicheskogo Instituta . USSR Academy of Sciences, Proceedings of the Paleontological Institute . 239 . 210–214 .
15. Zessin W . 1988 . Neue Saltatoria (Insecta) aus dem Oberlias Mitteleuropas . New Saltatoria (Insecta) from the Upper Liaison of Central Europe . German . Freiberger Forschungshefte C . Freiberg Research Books C . 419 . 107–121 .
16. Martins-Neto RG . 1991 . Sistemática dos Ensifera (Insecta, Orthopteroida) da formação Santana, Cretáceo Inferior do Nordeste do Brasil. . Systematics of the Ensifera (Insecta, Orthopteroida) from the Santana, Lower Cretaceous formation in Northeastern Brazil. . pt. Acta Geologica Leopoldensia . 14 . 32 . 5–160 . 10.11606/D.44.1990.tde-26082015-144345 . free .
17. Gorochov AV . 1986 . Grasshoppers. Gryllida (=Orthoptera), in Nasekomye v rannemelovykh ekosistemakh zapadnoy Mongolii. . The Joint Soviet-Mongolian Palaeontological Expedition . 28 . 171–174 .
18. Gorochov AV, Jarzembowski EA, Coram RA . 2006-10-01. Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England . Cretaceous Research. en. 27. 5. 641–662. 10.1016/j.cretres.2006.03.007. 2006CrRes..27..641G . 0195-6671.
19. Giebel CG . 1856 . Die Insecten und Spinnen der Vorwelt mit steter Berücksichtigung der lebenden Insekten und Spinnen. . Die Fauna der Vorwelt . 2 . 1–511 .
20. Westwood JO . 1854 . Contributions to fossil entomology. . Quarterly Journal of the Geological Society of London . 10 . 1–2 . 378–396 . 10.1144/GSL.JGS.1854.010.01-02.43 . 129712238 .
21. Zhou . Qian . Xu . Chunpeng . Jarzembowski . Edmund A. . Xiao . Chuantao . 2022-08-01 . A new species of Elcanidae (Insecta: Orthoptera) from mid-Cretaceous Kachin amber . Cretaceous Research . en . 136 . 105226 . 10.1016/j.cretres.2022.105226 . 2022CrRes.13605226Z . 248105782 . 0195-6671.
22. Fang Y, Wang B, Zhang H, Wang H, Jarzembowski EA, Zheng D, Zhang Q, Li S, Liu Q . January 2015 . New Cretaceous Elcanidae from China and Myanmar (Insecta, Orthoptera) . Cretaceous Research. en. 52. 323–328. 10.1016/j.cretres.2014.05.004. 2015CrRes..52..323F .
23. Gorochov AB . 1989 . Novye taksony prjamokrylyx semejstv Bintoniellidae, Xenopteridae, Permelcanidae, Elcanidae, i Vitimiidae (Orthoptera, Ensifera) iz mezozoja azii . New taxa of the Orthopteran families Bintoniellidae, Xenopteridae, Permelcanidae, Elcanidae, and Vitimiidae (Orthoptera, Ensifera) from the Mesozoic Asia . ru. Vestnik Zoologii . 1989 . 4 . 20–27 .
24. Hu . T.-H. . He . Z.-Q. . 2023 . A new species of genus Probaisselcana (Orthoptera: Elcanidae) from mid-Cretaceous Burmese amber . Cretaceous Research . 151 . en . 105671 . 10.1016/j.cretres.2023.105671. 2023CrRes.15105671H .
25. Peñalver E, Grimaldi DA . 2010-01-01. Latest occurrences of the Mesozoic family Elcanidae (Insecta: Orthoptera), in Cretaceous amber from Myanmar and Spain . Annales de la Société Entomologique de France . N.S.. 46. 1–2. 88–99. 10.1080/00379271.2010.10697641. 84115545. 0037-9271.
26. Fang Y, Muscente AD, Heads SW, Wang B, Xiao S . 2018-05-25. The earliest Elcanidae (Insecta, Orthoptera) from the Upper Triassic of North America . Journal of Paleontology. 92. 6. 1028–1034. 10.1017/jpa.2018.20. 2018JPal...92.1028F . 133821611. 0022-3360.
27. Schubnel. Thomas. Desutter. Laure. Garrouste. Romain. hervé. sophie. Nel. André. 2020. Paleocene of Menat Formation, France, reveals an extraordinary diversity of orthopterans and the last known survivor of a Mesozoic Elcanidae. Acta Palaeontologica Polonica. 65. 10.4202/app.00676.2019. 0567-7920. free.
28. Heads SW, Thomas MJ, Wang Y . A new genus and species of Elcanidae (Insecta: Orthoptera) from Cretaceous Burmese amber . Zootaxa . 4527 . 4 . 575–580 . December 2018 . 30651491 . 10.11646/zootaxa.4527.4.8 . 58668337 .
29. Poinar G, Gorochov AV, Buckley R . 2007 . Longioculus burmensis, n. gen., n. sp. (Orthoptera: Elcanidae) in Burmese amber. . Proceedings of the Entomological Society of Washington . 109 . 649–655 .
30. Uchida. Kei. 2021-11-17. Two new genera of the Elcanidae (order: Orthoptera) from middle Cretaceous Burmese amber of northern Myanmar. Cretaceous Research. 131 . en. 105092. 10.1016/j.cretres.2021.105092. 244377365 . 0195-6671. free.
31. Xu. Chunpeng. Luo. Cihang. Jarzembowski. Edmund A.. Fang. Yan. Wang. Bo. 2022-03-28. Aposematic coloration from Mid-Cretaceous Kachin amber. Philosophical Transactions of the Royal Society B: Biological Sciences. 377. 1847. 20210039. 10.1098/rstb.2021.0039. 35124999 . 8819367 . 246608514 .