Morphology of the Female Terminalia of Nymphes Myrmeleonoides Leach, 1814 (Neuroptera Nymphidae) and Phylogenetic Implications
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Ann. Mus. civ. St. nat. Ferrara Vol. 8 2005 [2007] pp.55-58 ISSN 1127-4476 Morphology of the female terminalia of Nymphes myrmeleonoides Leach, 1814 (Neuroptera Nymphidae) and phylogenetic implications Verena Feuerstein Georg-August-University Gottingen. Johann-Friedrich-Blumenbach-Institute of Zoology and Anthropology, De partment of Morphology and Systematics, Berliner Str. 28, 0-37073 Gottingen, Germany, e-mail: [email protected] The female terminal structures of Nymphes myrme/eonoides Leach, 1814 (Nymphidae) are de scribed in detail. Within Neuroptera the genitalia of Nymphidae are considered to be highly derived. However, all elements of the "orthopteroid" ovipositor are obviously present in N. myrme/eonoides. The ovipositor is short and is composed basically of the gonocoxites IX (+ valvulae III). A pair of scierites (valvulae II) is present along the inner membranous sides. This pair of sclerites interlocks with a structure often called "subgenitale" (fused valvulae I). Similar structures exist in Chrysopidae and the arrangement may even be homologous with the olistheter of Raphidioptera. The valvulae I and II are missing in some other Neuroptera (e.g. Osmylidae). The interpretation of the subgenitale and the corresponding sclerites of Nymphes (and chrysopids) as ovipositor derivates sheds new light on the evolution of the ovipositor in Neuroptera and provides new views of their phylogeny. Key words - Neuroptera, Nymphidae, Nymphes myrme/eonoides, ovipositor, morphology, phylo geny. Introduction the internal and external genitalia of Neu ropterida was performed by Stitz (1909). Nymphidae is a small taxon consisting Principi (1949) and Philippe (1972) de of approximately 35 extant species restric scribed the morphology of the female ge ted to Australia, New Guinea and adjacent nitalia of Chrysopa, that is characterized islands. It is considered to be the sister by a very short ovipositor. Investigations group of Myrmeleontidae + Ascalaphidae of taxa with a long ovipositor similar to that + Nemopteridae (New, 1984; Oswald, of the Raphidioptera are restricted to the 1998) or of Myrmeleontidae + Ascalaphi external morphology. Ferris (1940) exami dae (Mansell, 1992; Aspock et al., 2001) ned the ovipositor of Plega signata (Ha within the Myrmeleontiformia (Myrmeleon gen, 1877) (Mantispidae) and Tjeder toidea sensu Mansell, 1992), one of the (1937) that of Dilaridae. more species rich subtaxa within Neuro The overall similarity of the female ge ptera. nital structures of Neuroptera, Megalopte Contradictory interpretations of the ra and Raphidioptera was demonstrated neuropteran ovipositor structures exist in by Mickoleit (1973a). He hypothesized a the literature and much controversy sur short ovipositor for the ground plan of rounds the homology of female genitalia. Neuropterida and complete loss of the go Morphological investigations of the ovipo napophyses IX (valvulae II). The gonapo sitor have been carried out predominantly physes VIII (valvulae I) are assumed to for taxonomic reasons. Detailed morpho have been lost in the ground plan of Neu logical examinations exist only for a few roptera as well as that of Megaloptera. Al neuropteran taxa. The first examination of ternatively, Mickoleit (1973a) discussed a 55 Proceedings of the IX International Symposium on Neuropter%gy long raphidiopteran-like ovipositor with present along the inner membranous si valvulae I and valvulae II for the ground des of gonocoxites IX. This pair of scleri plan of Neuropterida. Although Mickoleit tes interlocks with an unpaired sclerotized abandoned this theory for functional rea projection of the segment VIII, the so sons, Achtelig (1978) supported the hypo called "subgenitale" (Figs 1 C and 2 A) - a theses after further investigations of Ra structure invariably present in all nym phidioptera. phids (New, 1984). The short ovipositor of Nymphidae and Comparable structures exist in Heme other Myrmeleontiformia was assumed to robiidae (e.g. Wesmaelius, Aspock et al., be highly derived and for this reason con 1980) and Chrysopidae (e.g, Chrysopa, sidered to be less important for the under Principi, 1949; Philippe, 1972; Chrysoper standing of the ground pattern of female la: Fig. 2 B). In Chrysopidae the "subgeni genitalia in Neuroptera. However, exami tale" is a pair of lobes which articulate with nation of Nymphes myrmeleonoides Le a pair of sclerotized pockets like a snap ach, 1814 (Nymphidae) showed that obvi press button ("button-pression", Philippe. ously all elements of an "orthopteroid" ovi 1972). positor are present and that it is not as de Achtelig (1978) compared the arrange rived as previously assumed. ment of the sclerites found in Chrysopidae with the interlock of the long ovipositor of Material and methods Raphidioptera (Fig. 3 C). In contrast to Smith (1969) and Mickoleit (1973a) he The ovipositor morphology of N. myrmeleo noides was examined based on alcohol fixed material (70%) and macerated specimens that were studied in glycerine. Histological sections of N. myrmeleonoides and Chrysoper/a carnea (Stephens, 1836) were made. For this purpose specimens were fixed in Bouin's solution, sec tioned serially at 10l-lm and stained with "Nu clear Fast Red" and "Aniline Blue-Orange G" (Azan stain). Females of N. myrmeleonoides were col lected at Rocksberg near Brisbane (Queens land, Australia) in October 1997 by Ulrike Nol te. Specimens of C. carnea were collected at s VII B G6ttingen (Germany) in February 2004 by Ve rena Feuerstein. Results and discussion The female external genitalia of N. myrmeleonoides (Figs 1 A and 1 B) con sist primarily of a pair of so-called "gona pophyses laterales", a structure of segment IX present in all Neuropterida. It is inter preted as the gonocoxites IX + gonostyli Fig. 1 - Female of Nymphes myrmeleonoides: A. IX (valvulae III) (Mickoleit, 1973a; Achte lateral view of terminalia; B, ventral view of termina lig, 1978). In N. myrme/eonoides the go lia; C, fused valvulae I ("subgenitale") and valvulae II. nocoxites IX + valvulae III are relatively a ::: anus; cat::: cataprocessus; epr = ectoproct; gx = gonocoxite; i = interlock; vv = valvula; s ::: sternite; t = short and joined to the sternum along al tergite. most the entire length. A pair of sclerites is 56 V. Feuerstein - Female terminalia of Nymphes myrmeleonoides Leach, 1814 fused unpaired valvulae I. Extrapolating the situation of the oviposi tor found in Raphidioptera to that of Chrysopidae Achtelig came to the conclu sion that the lobes (subgenitale) of Chryso pidae are the reduced valvulae I and the snappress button interlock is a reduced oli stheter (Fig. 3 8). As a consequence the subgenitale of Nymphidae and Hemerobiidae may also be interpreted as the fused valvulae I. The sclerites along the inner sides of the gono coxites IX are the valvulae II (gonapo physes IX) with the primary olistheter as the interlock between them and the valvu lae I (Fig. 3). This leads to the conclusion that all elements of the orthopteroid ovipo sitor are not only present in the ground plan of Neuropterida, but also in the ground plan of Neuroptera. The question as to the characteristic of the ovipositor in the neuropteran ground plan still remains unanswered. Assuming a primary olistheter a long raphidiopteran-like ovipositor seems probable for the ground plan of Neuropterida. Taxa with a long ovi Fig. 2 - Cross section of the ovipositor through the pOSitor must be examined in order to deter region of the fused valvulae I: A, Nymphes myrme leonoides; B, Chrysoperla carnea. mine whether this also applies to Neuropte ra. Tjeder (1937) described an unpaired considered the interlock (= olistheter) of the subgenitale and a pair of rod-like appenda raphidiopteran ovipositor to be primary. His ges for Dilaridae which he homologized with examinations of prothetelic larvae showed the valvulae I and valvulae II. Mickoleit that the dorsal part of the ovipositor of Ra (1973a), who pointed out the similarity of the phidioptera, consisting mainly of the gono ovipositor of Dilaridae and Raphidioptera, coxites IX + valvulae III, also contains the discussed the rod-like appendages of Dilari valvulae II along their inner sides (Fig. 3 C). dae as the valvulae I. Similar structures of The valvulae II, which have the function of the segment VIII have not been described a rail, interlock on each side with the long for the long ovipositor of Plega signata 8 c vv I 200 Ilm Fig. 3 - Schematic cross section of the ovipositor of (A) Nymphes myrmeleonoides, (B) Chrysoperla carnea, (C) Raphidioptera (after Mickoleit, 1973b). Emphasized structures show sci erotized part. 57 Proceedings of the IX International Symposium on Neuropterology (Mantispidae) (Ferris, 1940). In accor Aspock U., Plant, J. D. & Nemeschkal H., 2001 - Cla dance with Achtelig (1978) the presence distic analysis of Neuroptera and their systematic position within the Neuropterida (Insecta: Holometa of valvulae I and valvulae II in Neuropte bola: Neuropterida: Neuroptera). Systematic Entomo ra is considered to be plesiomorphic and logy, 26: 73-86. can be found at least in Nymphidae and Ferris G.F., 1940 - The morphology of Plega signata within Hemerobiiformia in Hemerobiidae (Hagen) (Neuroptera: Mantispidae). Microentomo logy, 5: 33-56. and Chrysopidae. The structures dis Mansell M.W., 1992 - The systematic position of the cussed are apparently lacking in other Nemopteridae (Insecta: Neuroptera: Myrmeleontoi Neuroptera, for example in Myrmeleon