Abstract

Ghost lineages have always challenged the understanding of organism evolution. They participate in misinterpretations in phylogenetic, clade dating, biogeographic, and paleoecologic studies. They directly result from fossilization biases and organism biology. The Cylindrotomidae are a perfect example of an unexplained ghost lineage during the Mesozoic, as its sister family Tipulidae is already well diversified during the Cretaceous, while the oldest Cylindrotomidae are Paleogene representatives of the extant genus Cylindrotoma and of the enigmatic fossil genus Cyttaromyia. Here we clarify the phylogenetic position of Cyttaromyia in the stem group of the whole family, suggesting that the crown group of the Cylindrotomidae began to diversify during the Cenozoic, unlike their sister group Tipulidae. We make a comparative analysis of all species in Cyttaromyia, together with the descriptions of the two new species, C. gelhausi sp. nov. and C. freiwaldi sp. nov., and the revision of C. obdurescens. The cylindrotomid biogeography seems to be incongruent with the phylogenetic analysis, the apparently most derived subfamily Stibadocerinae having apparently a ‘Gondwanan’ distribution, with some genera only known from Australia or Chile, while the most inclusive Cylindrotominae are Holarctic.

Highlights

  • Cylindrotomidae Schinner, ­18631 together with Limoniidae Speiser, ­19092, Pediciidae Osten-Sacken, ­18603, and Tipulidae Latreille, 1­ 8024 sensu stricto are classified within Tipuloidea Latreille, 1­ 8024, group of insects present in the fossil record since at least 220 Ma (Triassic)[5]

  • Three species of Cylindrotoma are known from impressions of the Ypresian Fur Formation; two species from the Ølst Formation of Denmark were described within Cyttaromyia[29] (Supplementary Table S1)

  • Four species of Cyttaromyia were described from the Eocene Green River Formation USA, the other were described from Florissant Formation and Kishenehn Formation in USA, Middle Salt Formation in Alsace (France), Biamo Formation in Russia, and from Baltic ­amber[24,27,28,30,31,40]

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Summary

Introduction

Cylindrotomidae Schinner, ­18631 together with Limoniidae Speiser, ­19092, Pediciidae Osten-Sacken, ­18603, and Tipulidae Latreille, 1­ 8024 sensu stricto are classified within Tipuloidea Latreille, 1­ 8024, group of insects present in the fossil record since at least 220 Ma (Triassic)[5]. According to phylogenetic synthesis based on combined morphological characters of adult, larvae and pupae, together with nuclear gene sequence data as 28S rDNA or CAD, the Cylindrotomidae are found as a sister group of Tipulidae This group of insects is generally indicated as a sister group or being closely related to the Tipulidae by other a­ uthors[10,11,12,13,14,15,16,17,18,19,20]. We describe two new species of Cyttaromyia on the basis of new fossils from the same Formation, and new technics of research give us possibility to redescribe Cyttaromyia obdurescens Cockerell, ­192427

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