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The Phylogeny and Geographical Distribution of the Aleuropteryginae (Neuroptera, Coniopterygidae)

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The Phylogeny and Geographical Distribution of the Aleuropteryginae (Neuroptera, Coniopterygidae) The phylogeny and geographical distribution of the Aleuropteryginae (Neuroptera, Coniopterygidae) MEINANDER,11. 1979. The phylogeny and geographical distribution of the Aleuropteryginae (Neuroptera, Coniopterygidae) . - Ann. Ent. Fenn. 45, 16-23. An attempt is made to apply cladistic analysis to the subfamily. The division into three tribes is accepted, Fontenelleini being considered to be a sister group to Coniocompsini and Aleuropterygini. Fontenelleini is divided into three different evolutionary lines. The fossil species are reviewed and their systematic position discussed. Coniopterygidae existed before Pangaea broke up into different continents and an attempt is made to correlate the phylogeny with the theory of con- tinental drift. M. Meinander, Division of Entomology, Zoological Museum, N. Jarnviigs- gatan 13, SF-00100 Helsingfors 10, Finland. Index words: Aleuropteryginae, phylogeny, distribution. In my revision of the family Coniopterygidae easy task to decide the evolutionary status of its (MEINANDER1972), all the species then described characters from the few fossils available. The were arranged phenetically in systematic groups establishment of synapomorphy on the basis of of different categories, which were supposed to recent material generally requires that the char- be monophyletic or at least paraphyletic in the acters are unique and complex, and in the sense of HENNIG(1966), but the phylogeny re- Coniopterygidae such characters are found only ceived only a superficial analysis. in the genitalia. However, although the genital The vast progress made by the earth sciences characters are generally quite sufficient to during the last few decades has improved our establish monophyletic genera, the great spe- knowledge of continental drift and, despite the cialization of the genitalia makes it hard to trace inevitable controversy characterizing the theory homolo~ousorgans. and it is thus often difficult " ', 2 and data of plate tectonics, consensus seems to to group the genera into monophyletic lines. have been reached regarding the sequence of That the family Coniopterygidae is a mono- continental movements. Continental drift, re- phyletic group is certain and is supported by sulting in the break-up of the continents and many synapomorphic characters (MEINANDER the establishment of new contacts with other 1972). The families Coniopterygidae and Itho- continents, evidently played a great role in the nididae apparently form two separate mono- evolution of groups like Coniopterygidae exist- ohvletic1, lines. which branched off early from ing from the Mesozoic. the other Planipennia, probably forming inde- An attempt is made here to apply cladistic pendent sister groups to the bulk of the Plani- analysis to Aleuropteryginae, one of the two pennia (MEINANDER1972). The two subfamilies subfamilies of Coniopterygidae, and to discuss into which the Coniopteryginae are divided. the phylogeny and history of the subfamily in Aleuropteryginae and - Coniopteryginae, are the light of its present geographical distribution. clearly sister groups, since they possess several synapomorphic characters (MEINANDER1972). Cladistic analysis of the Aleuropteryginae In MEINANDER1972, the subfamily Aleurop- teryginae was divided into three phenetically Cladism is a logical framework for recon- well-characterized tribes, which could be proved structing phylogeny, in which the basic principle to be monophyletic (Fig. 1). Two of these, Al- is HENNIG'Scriterion of monophyly, that only europterygini and Coniocompsini, were found synapomorphic (shared derived) relationships to form a sister group to the third, Fontenelleini. can aid in the establishment of monophyletic In Aleuropterygini, Heteroconis and Aleu- lines. Although apparently rather old, and with ropteryx are sister groups. TJEDER( 1972) divid- a narrowr morphological spectrum, it is no ed Heteroconis into two subgenera, creating Ann. Ent. Fenn. 45: 1. 1979 17 Fig. 1. Cladogram of the Aleuropteryginae with the six main evolutionary lines. Drepanoconis for an aberrant species from New Guinea, for which only the female is known. I refer to consider Drepanoconis a synonym of Heteroconis until the male is described. Dreba- tloconis can hardly be a sister group to the rest of the widely distributed Heteroconis, six species of which were described by TJEDERfrom the same mountain range in New Guinea, but it has some highly specialized characters, and the creation of a new monotypic generic group for it would make Heteroconis paraphyletic. Al- though it is sometimes necessary to have phene- tically homogeneous paraphyletic groups, I think they should be avoided when possible. In MEINANDER1972, Aleuropteryx was divid- ed into three species groups, the A. loewii Kla- Fig. 2. Cladogram of the Fontenelleini. Apomorphic palek group ( 11 spp.), the A, minuta Meinan- characters marked with black and synapomorphic in- der group (6 spp.) and the monotypic A. ar- dicated by hatching. Further explanation in the text. gentata Tjeder group. The A. loewii group is a sister group to the other two. Apomorphic char- ly represent three different evolutionary lines, acters in the A. loewi group are the large and group A with Paraconis, Cryptoscenea, Neoconis con~plexprocess of sternite IX and the complex and Pampoconis, group B with Spiloconis and dructure of this sternite; its dorsal edges are group C with Bidesmia, Vartiana, Pseudoconis, fused so that it forms a ring, through which the Capoconis, Helicoconis, Ohmopteryx and Fonte- penis protrudes. Synapomorphic characters in nella. It has not been possible to arrange the the A, minuta and A. argentata groups are the groups into a system of sister groups. inaculate wings and the strong prolongation Synapomorphic characters for group A are that caudad of the penis sclerite. Whether the A. the ectoprocts are completely merged with seg- tnilzuta group is a sister group to A. argentata ment 9 and are equally strongly sclerotized (A). or a paraphyletic group from which the highly The ectourocts of the aberrant monotvuic, L Var- hpecialized A. argentata is derived cannot be tiana in group C are also incorporated in seg- determined. ment 9 but this has obviously occurred inde- The hypothetical evolution of the Fontenel- pendently of the development in group A. The leini is shown in the cladogram in Fig. 2. The styli in group A (A2) are placed on separate smbols used in the discussion of the characters ~late~s. below refer to the cladogram. Group B is characterized by very prolonged The Fontenelleini can be divided into three scapes and pedicels (Bi); in their plcsio~norphic phenetically well-defined groups, which obvious- condition in the other groups these structures 18 Meinnnder, The phylogeny and geographical distribution . are thus found only on segments 3-6, and (22) a synapomorphic character is probably also the development of strong spines on the ectoprocts. Cryptoscenea and Paraconis have hairy paired lobes caudad of segment 9, which are called coxo~odites in MEINANDER1972. If thev are coxopodites, their presence is plesiomorphic and their reduction in line 2 could be considered a synapomorphic character. Whether Cryptoscenea and the monotypic Pa- raconis can be considered sister groups is diffi- cult to decide, due to their narrow morphologi- cal spectrum. An apomorphic character of Para- conis (3) is the structure of the penis sclerite: which has lost all traces of the two ~arallelscle- rites found in all the other genera of the tribe. The genitalia of the four species of Crytoscenea for which the male is described show a funda- mental similarity, which may be synapomorphic. but on the other hand the genitalia of Para- conis could well be derived from those of Cry- ptoscenea, in which case Cryptoscenea is a para- phyletic genus. When the family was revised (MEINANDER Fig. 3. The position of the three southern continents 1972), the genera Neoconis and Pampoconis remained almost the same until the Eocene (45 m.y,). were readily separated phenetically. A key char- (After MCKENZIE& SCLATER1971). acter used was the presence of a distal cross- vein M-Cui in the hind wing in Pampoconis and its absence in Neoconis. but additional are about one and a half times as long as broad. characters could be found in the male genitalia, Sternite IX is incorporated in the internal geni- e.g. the shape of the penis sclerite and the pres- talia (B2); in the plesiomorphic condition in the ence of an extra rod articulating with the para- other groups it is external although sometimes mere in Neoconis. At the time of the revision somewhat telescoped into segment 8. Sternite (MEINANDER1972), seven species of Neoconis VIII is prolonged and forms the floor of the had been recorded, while Pampoconis was mo- abdominal tip (B3), whereas in the other groups notypic. Since then five new species have been it is not prolonged. As a synapomorphic char- described ( ADARIS 1973, MEINANDER1973. racter of group C I recognize the wide separ- 1974a, 1974b) and using the key character ation of veins M and Cui in the hind wing. In given in MEINANDER1972 one species has been the plesiomorphic condition, represented in all assigned to Neoconis and four to Parnpoconis. the species of Aleuropterygini, Coniocompsini Of these species Pampoconis insulana Meinander and groups A and B, they run very close to each has an interesting intermediate position be- other. The separate sclerite called the hypand- tween the two genera. Besides the wing charac- rium, which exists in most genera of group C, ter, other
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