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Journal of Natural History Ovipositor Structure and Relationships Within See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/236146132 Ovipositor structure and relationships within the Hymenoptera, with special reference to the Ichneumonoidea Article in Journal of Natural History · June 1994 DOI: 10.1080/00222939400770301 CITATIONS READS 120 278 5 authors, including: Donald L J Quicke Simon Nicholas Ingram Chulalongkorn University University of Plymouth 575 PUBLICATIONS 10,653 CITATIONS 61 PUBLICATIONS 1,629 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Aspilota-group (Braconidae: Alysiinae) View project Hymenoptera Phylogeny View project All content following this page was uploaded by Simon Nicholas Ingram on 22 August 2014. The user has requested enhancement of the downloaded file. This article was downloaded by: [Univ of Plymouth] On: 22 August 2014, At: 06:48 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnah20 Ovipositor structure and relationships within the Hymenoptera, with special reference to the Ichneumonoidea D.L.J. Quicke b d , M.G. Fitton a , J.R. Tunstead b , S.N. Ingram b & P.V. Gaitens c a Department of Entomology , The Natural History Museum , London, SW7 5BD, UK b Department of Animal and Plant Sciences , University of Sheffield , Sheffield, S10 2TN, UK c Electron Microscope Unit , University of Sheffield , Sheffield, S10 2TN, UK d Department of Biology , Imperial College , Silwood Park, Ascot, SL5 7PY, UK Published online: 17 Feb 2007. To cite this article: D.L.J. Quicke , M.G. Fitton , J.R. Tunstead , S.N. Ingram & P.V. Gaitens (1994) Ovipositor structure and relationships within the Hymenoptera, with special reference to the Ichneumonoidea, Journal of Natural History, 28:3, 635-682, DOI: 10.1080/00222939400770301 To link to this article: http://dx.doi.org/10.1080/00222939400770301 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/ terms-and-conditions Downloaded by [Univ of Plymouth] at 06:48 22 August 2014 JOURNAL OF NATURAL HISTORY, 1994, 28, 635-682 Ovipositor structure and relationships within the Hymenoptera, with special reference to the Ichneumonoidea D. L. J. QUICKE*t, M. G. FITTON$, J. R. TUNSTEAD*, S. N. INGRAM* and P. V. GAITENS§ Department of Entomology, The Natural History Museum, London SW7 5BD, UK * Department of Animal and Plant Sciences and § Electron Microscope Unit, University of Sheffield, Sheffield $10 2TN, UK t Present address: Department of Biology, Imperial College, Silwood Park, Ascot SL5 7PY, UK (Accepted 30 August 1993) The morphology, function and phylogenetic significance of the hymenopterous ovipositor, as revealed by transverse sections, is explored. Ovipositors (including stings) of > 240 species belonging to some 69 families (representing all super- families) have been prepared and examined, and almost 180 are illustrated. Particular attention is paid to Ichneumonoidea. Sections show many new, phylogenetically informative characters as well as providing further insight into ovipositor function. Examples of synapomorphies are given which suggest various groupings at the family or subfamily levels. Functional interpretations are given for several characters, with particular reference to substrate penetration, passage of the egg along the ovipositor and stinging. KEYWORDS: Hymenoptera, ovipositor, morphology, phylogeny, Ichneumonoidea. Introduction The morphology of the ovipositor and its related structures has provided many useful characters for the systematics of Hymenoptera (see, for example, Fergusson, 1988). This is not surprising as the ovipositor and oviposition behaviour seem to have Downloaded by [Univ of Plymouth] at 06:48 22 August 2014 played crucial roles in the evolution of the order (Gauld and Bolton, 1988). However, of all the many studies on ovipositors, including the derived stings of aculeate wasps, bees and ants, few have examined the detailed structure of the ovipositor valves themselves, having instead relied principally on simple observations of gross external appearance. A few exceptions to this are worthy of note: the works of Scudder (1961), Oeser (1961), Brothers (1975) and Smith (1968-1970, 1972). Several textbooks of entomology and papers on ovipositor structure provide diagrams of cross sections of hymenopteran ovipositors, but most such illustrations were not based on original work, many are highly stylized, and some even misleading (for example, Weber, 1933; Riek, 1970; Askew, 1971; Richards and Davies, 1977; Gauld and Bolton, 1988). However, from examination of the publications that give more accurate diagrams or descriptions of cross sections through ovipositors and stings (for example, Abbott, 1934, 1935; Soliman, 1941; Venkatraman and Subba Rao, 1954; King, 1962; Delanoue and Arambourg, 1965; Hermann and Blum, 1967; Rogers, 1972; Smith, 1968-1970, 1972; Bronner, 1985; Billen, 1990) it was apparent 0022-2933194 $10.00 © 1994 Taylor & Francis Ltd. 636 D.L.J. Quicke et al. that there is considerable variation in structure between taxa. Our interest in the phylogenetic relationships of the families of Hymenoptera and, in particular, the subfamilies of Ichneumonoidea led us to conduct a wide-ranging survey of ovipositor/ sting structure using the technique of semi-thin sectioning, following resin embedding, in order to identify characters of phylogenetic or biological importance. In a com- panion paper (Quicke et al. 1992b) we consider the functional and phylogenetic significance of ovipositor valvilli in Hymenoptera. Together these two studies examine characters which should add significantly to our understanding of the evolution of the groups of Hymenoptera and pose many interesting questions about ovipositor function. General ovipositor morphology and terminology There have been several different, widely-used nomenclatures for the parts com- prising the hymenopteran ovipositor or sting system. Here we largely follow Smith (1968-1970, 1972), except that we refer to the gonapophyses of the ninth and eighth abdominal segments as the upper and lower valves of the ovipositor, respectively. Our reason for employing this more descriptive terminology is that we hope to increase the accessibility of our results to workers outside the field of hymenopteran morphology and systematics. The terms, and the consequent references to 'dorsal' and 'ventral' parts, assume the ovipositor is directed posteriorly from the apex of the metasoma, though in some taxa the rotation of the metasomal apex may cause the ovipositor to protrude anteriorly, even in repose, for example in Vanhornia and some chalcidoids. Also, we use 'ovipositor' to include the sting, the term used to describe it in aculeate Hymenoptera in which it retains only that function. Smith (1969, 1970) gave detailed glossaries of equivalent terms and Fergusson (1988) provided a small comparative table for the main parts of the ovipositor system. Smith (1969, 1970) provided good evidence that the paired appendages forming the upper and lower valves were originally segmented (meristic) structures, com- prising a basal segment termed the radix, followed by the multisegmented lanmium. Segments of the lamnium are modified to form the tooth-like processes involved in substrate penetration. In most Symphyta the lamnia of both the upper and lower valves are serrated (Fig. 1), in ichneumonoids usually only the lower valves have teeth (Fig. 2), whereas in most other groups, such as Megalyroidea and Chalcidoidea, it is the upper valve that is serrated (Fig. 3). Frequently, the apex of the ovipositor valve(s) Downloaded by [Univ of Plymouth] at 06:48 22 August 2014 opposite the one with serrations is furnished with a single protuberance, referred to as a node or nodus, (Fig. 2) or occasionally it has a notch. Smith (1969, 1970) concluded that in those groups in which the teeth are restricted to the apex of the ovipositor valve, the smooth basal region represents the elongated radix. However, this may not be so (see Results). The upper and lower valves are hollow structures, the lumen of each (Figs
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