Abstract
The relic dragonfly family Epiophlebiidae is recovered as sistergroup of Anisoptera (= Epiprocta) by most molecular and morphological analyses. However, in a recent study it was placed within Anisoptera as sister group of Cordulegastridae. In another study, several affinities to Zygoptera in the morphology of the ovipositor and the egg-laying behaviour were pointed out. Here, we present a detailed study of the outer, as well as the inner, head morphology of Epiophlebia superstes. Compared with the last detailed literature account, three additional mandibular muscles were discovered, as well as additional buccal and pharyngeal muscles. The results are compared with the anatomic features of Zygoptera and Anisoptera. A formal character evaluation focused on head characters confirms the sistergroup relationship of Epiophlebiidae and Anisoptera.
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Acknowledgment
Fabian Wilde (HZG, Geesthacht), as well as Sebastian Thieß and Wolfgang Drube (DESY, Hamburg), provided excellent support at the DESY synchrotron facilities. The scanning was done in the course of proposals no. I-20080169 and I-20090211. Marco Stampanoni and Peter Modregger are sincerely thanked for their support at the PSI for SLS TOMCAT beamline analysis (proposals 20100137 and 20110069). Irina Ruf (Steinmann Institute, Bonn, Germany) made microCT investigations at the device of the Steinmann Institute possible. We would like to express our gratitude to Professor Ryuichiro Machida (University of Tsukuba, Japan) who kindly provided specimens of E. superstes. Benjamin Wipfler and Rolf Beutel (University of Jena, Germany) are thanked for useful advice. Furthermore, we thank two anonymous reviewers for their valuable comments. The laboratory members of the ZFMK are sincerely thanked for their support.
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Appendix 1. Character discussion (Fig. 4; ESM 3)
Appendix 1. Character discussion (Fig. 4; ESM 3)
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0.
Composition of mandibular abductor: (0) undivided (0md3); (1) divided into two bundles (0md2 and 0md3). Ephemeroptera possess two mandibular abuctors (0md2 and 0md3). In Odonata only the 0md3 is present. The muscle is divided distally into two bundles. The origin of the two bundles closely resembles the origin of 0md2 and 0md3 in Ephemeroptera.
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1.
M. frontobuccalis lateralis: (0) absent; (1) present. In Odonata M. frontobuccalis lateralis (0hy2), M. tentoriohypopharyngalis (0hy3) and M. praementosalivarialis posterior (0hy8) are absent. Ephemeroptera possess these muscles (Staniczek 2000, 2001).
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2.
M. tentoriohypopharyngalis (0hy3): (0) absent; (1) present.
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3.
M. praementosalivarialis posterior (0hy8): (0) absent; (1) present.
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4.
M. tentoriopraementalis inferior (0la5): (0) absent; (1) present. The labial muscles M. tentoriopraementalis inferior (0la5) and M. praementoglossalis (0la11) are absent in Odonata, but present in Ephemeroptera.
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5.
M. praementoglossalis (0la11): (0) absent; (1) present.
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6.
Galea: (0) absent; (1) present. Ando (1962) discovered that no galea is formed during embryonic development in Odonata. In Ephemeroptera the galea is fused with the lacinia but still discernible in certain taxa (Staniczek 2000, 2001).
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7.
Salivary system: (0) absent; (1) present. A salivary system is present in Odonata but absent in Ephemeroptera.
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8.
Head: (0) transversely elongate; (1) globular. The head of Zygoptera is transversely elongated with the dome-shaped eyes located at the distal ends.
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9.
M. tentoriomandibularis medialis superior (0md7): (0) absent; (1) present. The M. tentoriomandibularis medialis superior (0md7) is present in all studied Zygoptera and Ephemeroptera. It is also present in the anisopteran taxa Austropetaliidae, Cordulegastridae, Petaluridae and Gomphidae but absent in Neopetaliidae, Aeshnidae, Libelullidae, Macromiidae and Corduliidae. Owing to its absence in E. superstes the primary loss of 0md7 is regarded as a groundplan feature of Epiprocta.
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10.
Eyes: (0) seperated by less than their own width; (1) separated by more than their own width. In all adult Anisoptera and Anisozygoptera the eyes are separated by less than their own width, all Zygoptera and Ephemeroptera show the opposite state.
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11.
Anteclypeus and postclypeus: (0) not facing anteriorly; (1) facing anteriorly. In Zygoptera the anteclypeus and postclypeus can form distinct anterior and dorsal faces respectively. It can also be greatly swollen (e.g., Chlorocyphidae). Anteclypeus and postclypeus face anteriorly in all studied Anisoptera. In contrast to Rehn (2003) this state is also present in E. superstes.
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12.
Labrum transversely enlarged: (0) absent; (1) present. The lateral sides of the labrum are strongly protruding transversely in E. superstes. This is a potential autapomorphy of Epiophlebiidae.
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13.
Vertex: (0) flat; (1) grossly enlarged and shield like; (2) formed as a moderate protuberance. A flat vertex characterizes all Zygoptera. All Anisoptera and Anisozygoptera exhibit a more or less prominent elevation of the vertex. The unique shape of the vertex in E. superstes is a potential autapomorphy of Anisozygoptera.
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14.
Pedicellus: (0) flattened with a lateral vestiture of long setae; (1) not as above. The shape and structure of the pedicellus is another potential autapomorphy of Epiophlebiidae.
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15.
M. frontobuccalis posterior (0bu3): (0) absent; (1) present. The 0bu3 is present in E. superstes but absent in all Anisoptera and Zygoptera. It is also present in Ephemeroptera.
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16.
Internal part of the interantennal ridge (interantennal apodeme): (0) absent; (1) present. An interantennal apodeme originating at the interantennal ridge is present in all studied Anisoptera and absent in E. superstes and Zygoptera.
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17.
Internal part of the epistomal ridge (epistomal apodeme): (0) absent; (1) present. An epistomal apodeme originating at the epistomal ridge is present in all studied Anisoptera and absent in E. superstes and Zygoptera.
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Blanke, A., Beckmann, F. & Misof, B. The head anatomy of Epiophlebia superstes (Odonata: Epiophlebiidae). Org Divers Evol 13, 55–66 (2013). https://doi.org/10.1007/s13127-012-0097-z
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DOI: https://doi.org/10.1007/s13127-012-0097-z