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      The presumed oldest flying insect: more likely a myriapod?

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      PeerJ
      PeerJ Inc.
      Chilopoda, Rhyniognatha hirsti, Rhynie chert, Crussolum, Devonian insect

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          Abstract

          The early fossil record of insects is scarce, with only few finds in the Devonian. All these finds appear problematic and controversial, partly due to incomplete preservation and challenging interpretation of many structures. We provide details of one of these important forms, Rhyniognatha hirsti from the famous Rhynie Chert Lagerstätte with up-to-date 3D imaging techniques. The fossil has been interpreted as the remains of one of the earliest flying insects. The specimen mainly preserves the remains of the head. The structures of the mandibles have been used as a main argument for an interpretation as an insect, but these are in fact less easy to interpret. New observed structures include the remains of a head capsule and an additional pair of mouth parts. Structures formerly suggested to represent remains of the head capsule or apodemes are more likely to be representing glands of ectodermal origin. The newly observed structures do not support an interpretation as an insect. Instead they make the interpretation as a myriapod more likely, possibly as a centipede. Centipede remains from the Rhynie Chert are known from scutigeromorphs. We therefore point out that R. hirsti could be interpreted as an early centipede.

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          New light shed on the oldest insect.

          Insects are the most diverse lineage of all life in numbers of species, and ecologically they dominate terrestrial ecosystems. However, how and when this immense radiation of animals originated is unclear. Only a few fossils provide insight into the earliest stages of insect evolution, and among them are specimens in chert from Rhynie, Scotland's Old Red Sandstone (Pragian; about 396-407 million years ago), which is only slightly younger than formations harbouring the earliest terrestrial faunas. The most well-known animal from Rhynie is the springtail Rhyniella praecursor (Entognatha; Collembola), long considered to be the oldest hexapod. For true insects (Ectognatha), the oldest records are two apparent wingless insects from later in the Devonian period of North America. Here we show, however, that a fragmentary fossil from Rhynie, Rhyniognatha hirsti, is not only the earliest true insect but may be relatively derived within basal Ectognatha. In fact, Rhyniognatha has derived characters shared with winged insects, suggesting that the origin of wings may have been earlier than previously believed. Regardless, Rhyniognatha indicates that insects originated in the Silurian period and were members of some of the earliest terrestrial faunas.
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            Fossil calibrations for the arthropod Tree of Life

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              Phylogenetic Distribution of Extant Richness Suggests Metamorphosis Is a Key Innovation Driving Diversification in Insects

              Insects and their six-legged relatives (Hexapoda) comprise more than half of all described species and dominate terrestrial and freshwater ecosystems. Understanding the macroevolutionary processes generating this richness requires a historical perspective, but the fossil record of hexapods is patchy and incomplete. Dated molecular phylogenies provide an alternative perspective on divergence times and have been combined with birth-death models to infer patterns of diversification across a range of taxonomic groups. Here we generate a dated phylogeny of hexapod families, based on previously published sequence data and literature derived constraints, in order to identify the broad pattern of macroevolutionary changes responsible for the composition of the extant hexapod fauna. The most prominent increase in diversification identified is associated with the origin of complete metamorphosis, confirming this as a key innovation in promoting insect diversity. Subsequent reductions are recovered for several groups previously identified as having a higher fossil diversity during the Mesozoic. In addition, a number of recently derived taxa are found to have radiated following the development of flowering plant (angiosperm) floras during the mid-Cretaceous. These results reveal that the composition of the modern hexapod fauna is a product of a key developmental innovation, combined with multiple and varied evolutionary responses to environmental changes from the mid Cretaceous floral transition onward.
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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                peerj
                peerj
                PeerJ
                PeerJ Inc. (San Francisco, USA )
                2167-8359
                30 May 2017
                2017
                : 5
                : e3402
                Affiliations
                [-1]Department of Biology, Ludwig-Maximilians-Universität München , Planegg-Martinsried, Germany
                Article
                3402
                10.7717/peerj.3402
                5452959
                1f40df11-b427-401f-95d9-cf8aed1bd4b4
                ©2017 Haug and Haug

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

                History
                : 17 October 2016
                : 10 May 2017
                Funding
                Funded by: European Community Research Infrastructure Action
                This research received support from the SYNTHESYS Project ( http://www.synthesys.info/) which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Entomology
                Paleontology
                Zoology

                chilopoda,rhyniognatha hirsti,rhynie chert,crussolum,devonian insect

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