Russian Entomol. J. 14(2): 87–100 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2005

Metamorphosis and homology of mouthparts in Neuropteroidea (Hexapoda: Metabola), with remarks on systematics and nomenclature

Ìåòàìîðôîç è ãîìîëîãèÿ ðîòîâûõ ÷àñòåé ó Neuropteroidea (Hexapoda: Metabola), ñ çàìå÷àíèÿìè ïî ñèñòåìàòèêå è íîìåíêëàòóðå

Nikita J. Kluge Í.Þ. Êëþãå

Department of Entomology, Saint-Petersburg State University, Universitetskaya nab., 7/9, Saint-Petersburg, 199034, Russia; e-mail: [email protected] Êàôåäðà ýíòîìîëîãèè, áèîëîãî-ïî÷âåííûé ôàêóëüòåò, Ñ-Ïåòåðáóðãñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Óíèâåðñèòåòñêàÿ íàá., 7/9, Ñ-Ïåòåðáóðã, 199034, Ðîññèÿ.

KEY WORDS: Neuroptera, Planipennia, Neuropteroidea, Euneuropteroidea, Birostrata, Chrysopidae, Myrme- leontidae, Megaloptera, Rhaphidioptera, Raphidia, Meganeuroptera, Sialis, Eumegaloptera, maxilla, mandible, metamorphosis, larva, pupa.

ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Neuroptera, Planipennia, Neuropteroidea, Euneuropteroidea, Birostrata, Chrysopi- dae, Myrmeleontidae, Megaloptera, Rhaphidioptera, Raphidia, Meganeuroptera, Sialis, Eumegaloptera, ìàêñèëëà, ìàíäèáóëà, ìåòàìîðôîç, ëè÷èíêà, êóêîëêà.

ABSTRACT. Examination of larval/pupal transfor- ìàêñèëëÿðíûì ùóïèêîì, ñëèòûì ñ ãàëååé, ëàöèíè- mation in Chrysoperla carnea and Chrysotropia ciliata åé è äèñòàëüíîé ÷àñòüþ ñòèïåñà; åãî êîëþùàÿ âåð- allows to understand homology of the larval paired øèíà îáðàçîâàíà íå ëàöèíèåé èëè ãàëååé, à ìàêñèë- piercing-sucking stylets, whose presence is the single ëÿðíûì ùóïèêîì. Êîëþùàÿ âåðøèíà ìàíäèáóëÿð- known autapomorphy of Birostrata (= Euneuropteroi- íîãî êîìïîíåíòà ýòîãî ñòèëåòà — íå ìàíäèáóëÿð- dea s.str.). The maxillary component of the piercing- íûé èíöèçîð, à âûðîñò ëàòåðàëüíåå èíöèçîðà. Îïè- sucking stylet is formed by maxillary palp fused with ñàíà òðàíñôîðìàöèÿ ìàêñèëë è ìàíäèáóë ïðè ïðå- galea, lacinia and a distal portion of stipes; its piercing âðàùåíèè ëè÷èíêè â êóêîëêó ó Raphidia è Sialis — apex is formed not by lacinia or galea, but by the Neuropteroidea ñ èñõîäíûì ãðûçóùèì ëè÷èíî÷íûì maxillary palp. Piercing apex of the mandibular compo- ðîòîâûì àïïàðàòîì. Ââîäèòñÿ íîâîå îáú¸ìíîå íà- nent of the stylet is not the mandibular incisor, but an çâàíèå Birostrata, taxon nov. Îáñóæäàþòñÿ îáú¸ìíûå outgrowth laterad of the incisor. Larval/pupal transfor- íàçâàíèÿ Neuropteroidea, Neuropterida, Euneuroptero- mation of maxillae and mandibles in Raphidia and idea, Planipennes, Planipennia, Neuroptera, Rhaphidio- Sialis (as Neuropteroidea with primitively biting larval ptera, Raphidioptera, Meganeuroptera è Megaloptera. mouth apparatus) is described. A new circumscriptional Âåðîÿòíî, Neuropteroidea — ãîëîôèëåòè÷åñêèé òàê- name Birostrata, taxon nov. is introduced. Circum- ñîí, õàðàêòåðèçóþùèéñÿ íàëè÷èåì èìàãèíàëüíîãî scriptional names Neuropteroidea, Neuropterida, Eu- äîðñàëüíîãî äèâåðòèêóëà ñòîìîäåóìà è äðóãèìè neuropteroidea, Planipennes, Planipennia, Neuroptera, àóòàïîìîðôèÿìè. Rhaphidioptera, Raphidioptera, Meganeuroptera and Megaloptera are discussed. Neuropteroidea is probably a holophyletic taxon characterized by presence of imag- Introduction inal dorsal stomodeal diverticulum and other autapo- morphies. Since strict rules on circumscriptional names are elaborated [Kluge, 1999a, 1999b, 2000, 2004], it be- ÐÅÇÞÌÅ. Èçó÷åíèå ëè÷èíî÷íî-êóêîëî÷íîé comes possible to bring in order existent not-typified òðàíñôîðìàöèè ó Chrysoperla carnea è Chrysotropia names which formerly were used differently by differ- ciliata ïîçâîëèëî ïîíÿòü ãîìîëîãèþ ëè÷èíî÷íûõ ent authors. The most complete catalogue of these names ïàðíûõ êîëþùå-ñîñóùèõ ñòèëåòîâ, íàëè÷èå êîòîðûõ is available from Internet [http://www.bio.pu.ru/win/ ÿâëÿåòñÿ åäèíñòâåííîé èçâåñòíîé àóòàïîìîðôèåé entomol/KLUGE/nom/~~~Cont.htm]. However, a large Birostrata (= Euneuropteroidea s.str.). Ìàêñèëëÿðíûé generally accepted holophyletic taxon known under êîìïîíåíò êîëþùå-ñîñóùåãî ñòèëåòà îáðàçîâàí names “Neuroptera”, “Planipennia”, “Megaloptera” and 88 Nikita J. Kluge others, appears to have no correct circumscriptional among secondary biting denticles. In many carnivorous name, which could be accepted; here for this taxon is insects mola is lost. Apex of incisor usually represents proposed a new name Birostrata. apex of the whole mandible. Initially, mandible has The only recently known reliable autapomorphy of muscles going from its base to cranium (anterior abduc- Birostrata is a highly specialized structure of larval tors and posterior adductors) and to endoskeleton (in mouth apparatus, which represents a pair of composite Amyocerata — to tentorium). In Metapterygota man- piercing-sucking stylets. Mandibles and maxillae, which dibular base is diminished, and mandibular-tentorial constitute these stylets, are so modified, that till now muscles are reduced, often completely lost. homology of their parts was unclear. In larvae of other Neuropteroidea — Rhaphidioptera, Sialis/fg1 and Eu- 1. Neuropteroidea megaloptera — mouthparts are not so modified, but homology of their maxillary parts also was a subject of discussion. For insects which undergo metamorphosis, Circumscriptional names: Neuropteroidea Handlirsch, 1903; homology of external parts of the body can be discovered Neuropterida Whiting & Carpenter & Wheeler & if this homology is known for another stage of ontogene- Wheeler, 1977. sis of the same insect: in this case it is possible to trace Hierarchical name: Myrmeleon/f1=Hemerobius/g1 (incl. development of this organ during metamorphosis. For Raphidia, Chauliodes) [f: Myrmeleonides Latreille, 1802: example, this method allowed to clarify homology of 287 (page priority upon Hemerobiini Latreille, 1802: parts of imaginal maxilla in Aphaniptera [Kluge, 2002]. 288; priority by first reviser: Myrmeleonina MacLeay, In all Neuropteroidea including Birostrata, homolo- 1821); g: Hemerobius Linnaeus, 1758: 549, type-species gy of mandibular and maxillary parts is quite clear in H. humulinus Linnaeus, 1758 (designated by the Com- pupa and imago, as pupal-imaginal mandibles and max- mission, Opinion 211, 1954)]. illae have structure close to the ancestral for Hexapoda. Typified name in use: Myrmeleontidea SENSU Rohdendorf The aim of this work was to trace development of 1977. maxillae and mandibles from larva to pupa, in order to NOMENCLATURAL COMMENTS. The name clarify homology of their parts in larva. Neuropteroidea Handlirsch, 1903 is well known, and Below, some comments are given about general the name Neuropterida Whiting et al., 1977 is its structure of maxillae and mandibles. evident junior circumscriptional synonym. In spite of MAXILLA. One of the good autapomorphies of this, some authors prefer to use Neuropterida instead of Hexapoda is structure of maxilla [Kluge, 1999b, 2000]. Neuropteroidea. Probably they think that the name Neu- Basal portion of maxilla normally has only one wall — ropteroidea should be rejected, because it contains the ventral (originally posterior), which is divided into two suffix with ending “-oidea”, which belongs to superfam- sclerites — cardo and stipes; from the stipes arise three ily, but not to taxa of other ranks (ICZN 29.2) [Interna- projecting parts of maxilla — lacinia, cardo and palp. tional Code of Zoological Nomenclature, 1999]. How- The palp is initially 5-segmented (but in some Hexapo- ever, this rule can be applied only to typified names of da can have more or less than five segments). Maxilla the family-group, but not to other names (ICZN 29.2.1). has following muscles (Figs 18–20): stipital-cranial; The name Neuropteroidea does not belong to family- muscles going from cardo and stipes to head endoskel- group in any sense (either strict sense of the ICZN, or eton (in Amyocerata — cardinal-tentorial and stipital- wide sense formulated by Kluge [1999]), because it is tentorial); lacinial-cranial, passing dorsad (originally not formed from any generic name; thus, its ending “- anteriad) of all other muscles; lacinial-stipital; galeal- oidea” does not contradict ICZN. Unlike Neuropteroi- stipital and palpal-stipital, both passing ventrad (origi- dea, another superordinal name from the same Handlir- nally posteriad) of the lacinial-stipital muscle; intrinsic sch’s classification — Panorpoidea — really should be muscles of palp segments. Among Hexapoda muscula- rejected, because it is a typified family-group name ture of maxilla is very conservative, and in many cases whose rank is not superfamily. allows to clarify homology of parts of maxillae [Kluge, STATUS OF NEUROPTEROIDEA. While the tax- 2002]. During metamorphosis maxillary musculature on Neuropteroidea is widely accepted, its autapomor- can disappear completely (in Birostrata) or partly (in phies are not well known, so some authors regard it to be Sialis and Raphidia), but in all cases larva and imago a plesiomorphon1. have the same set of muscles peculiar for Hexapoda. Some authors regard Neuropteroidea to be para- MANDIBLE. The initial mandible of Mandibulata phyletic, because assume that Meganeuroptera and Co- has incisor (or apical canine), kinetodontium (or subap- leoptera have a sister relationship. The only reason for ical canine) and mola [Kluge, 2000, 2004]. Some au- this is an assumption that a proximal widening of hind thors confused kinetodontium with prostheca (fused wing, which is characteristic for Meganeuroptera, is a setae between kinetodontium and mola in some insects). first step toward a hind-motor wing specialization, which In many insects kinetodontium is indistinguishable is characteristic for Coleoptera [Brodsky, 1994]. How-

1 The term “plesiomorphon” was introduced by Kluge [2004] to indicate a taxon which has no autapomorphies, being possibly paraphyletic, but whose paraphyly (as well as holophyly) is not proven [http://www.bio.pu.ru/win/entomol/KLUGE/ plesiomorphon.htm]. Metamorphosis and homology of mouthparts in Neuropteroidea 89

Fig. 1. Imago of Chrysoperla carnea: a — longitudinal section with alimentary canal; b — proventriculus and base of dorsal diverticulum. Ðèñ. 1. Èìàãî Chrysoperla carnea: à — ïðîäîëüíûé ñðåç ñ ïèùåâàðèòåëüíûì êàíàëîì; á — ïðîâåíòðèêóëþñ è îñíîâàíèå äîðñàëüíîãî äèâåðòèêóëà. ever, the same hind wing widening occurs in Ithone, In many representatives some of this characters are which belongs to Birostrata. lost: in most myrmeleontids wings are narrowed, so the Another idea about paraphyly of Neuropteroidea is widening of costal field is not expressed; in Rhaphid- connected with an assumed synapomorphy of all Me- ioptera and Dictyneuroptera Crampton 1916 (= Mantis- tabola except for Meganeuroptera. Sehnal [1985] re- pa/fg1) distal parts of Sc is transformed to a pterostig- ported: “To stress the similarity between larvae, pupae ma, so Sc looks falling into the fore margin of wing and adults of Megaloptera, this order and occasionally before the pterostigma; Rhaphidioptera, Sialis/fg1 and also Raphidioptera were by some authors separated some Birostrata have a small number of RS branches, so from other endopterygotes (called Euholometabola) as its pectinate branching is non-expressed. In Coniop- Eoholometabola”. Like F. Sehnal, I was failure to find teryx/fg1 all these characters of wing venation are lost. the original paper where the taxa names Euholometabo- • Ovipositor lost gonapophyses of the 9th abdominal la and Eoholometabola were proposed. Meganeuroptera segment (which initially for Amyocerata couple with really have an outstanding type of larval/pupal transfor- gonapophyses of 8th segment to form the working part of mation, where the phase of leg diminishing is absent, ovipositor); working part of ovipositor is constituted by and some of leg muscles are retained, while Rhaphid- coxites of the 9th abdominal segment, which are fused ioptera in this respect are similar to Birostrata and other dorsally, filled with intrinsic muscles (new formations), Metabola; however, some facts indicate that metamor- largely desclerotized and bear a pair of longitudinal phosis of Meganeuroptera is a result of secondary sim- lateral sclerites articulated with lateral sides of the 9th plification, but not a plesiomorphy [Kluge, in press]. abdominal tergite. If the ovipositor is long (in Rhaphid- Most probably, Neuropteroidea is a holophyletic ioptera, Dilar/fg1 and Dictyneuroptera–Trichoscelia/ taxon, as it has the following autapomorphies. fg1) it is actively flexible and in rest arched dorsally. In AUTAPOMORPHIES OF NEUROPTEROIDEA. most representatives ovipositor is short or reduced. • Imago (but not larva) has a peculiar dorsal divertic- • In imago the 10th abdominal tergite bears a pair of ulum of stomodaeum [Grinfeld, 1962]. This diverticu- groups of trichobothria. In some taxa they are lost lum arises from dorsal wall at hind part of foregut just [Aspöck et al., 2001]. before proventriculus, is directed backward and lies • In larva a pair of lateral cervical sclerites are fused above midgut (Fig. 1a). Proventriculus has a peculiar ventrally to form an integral unpaired semicircular scler- structure: it is armed by 8 arched sclerites which form a ite (Fig. 15). Imaginal cervical sclerites either retain the flower-like figure directed by its petals anteriorly (Fig. plesiomorphic paired structure, or have the larval struc- 1b). This proventricular armature is found in Raphidia, ture (in Eumegaloptera–Corydalus/f3=g1), or have a Chrysoperla, Osmylus and Mantispa; but not found in modification of their own (in Rhaphidioptera). Sialis and Sisyra. MOUTH APPARATUS. Imaginal maxilla has seto- • On fore wing, fore margin is convex in proximal se lacinia (Fig. 16) and probably is initially specialized part so that costal field is widened in this area, gradually for pallinophagy (while Rhaphidioptera and some Biros- narrowing toward apex; the costal field has many ob- trata are carnivorous, and Meganeuroptera and some lique cross-veins; Sc goes parallel to RA up to wing Birostrata are aphagous). Other features of imaginal apex, where it arches backward together with RA (or mouth apparatus are either evidently plesiomorphic, or fused with RA), repeating shape of fore margin of the variable. In all Neuropteroidea maxilla has structure wing. RS usually has pectinate branching. close to the initial for Hexapoda (see above), with 90 Nikita J. Kluge distinctly developed cardo, stipes, galea, lacinia and 5- ily Planipennes matched in circumscription Plecoptera segmented palp. Imaginal mandible is biting, either + Isoptera + Copeognatha + Birostrata + Meganeuroptera functional, or non-functional. + Rhaphidioptera + Mecaptera. Planipennes is a junior Structure of larval mouth apparatus is quite different circumscriptional synonym of Stegoptera Dumeril, in Birostrata, Rhaphidioptera, Sialis/fg1 and Eumega- 1806. Burmeister [1839] was the first to use the spelling loptera (see below). “Planipennia”. The taxon Planipennia Burmeister, 1839 CLASSIFICATION. Traditionally, Neuropteroidea matches in circumscription Birostrata + Meganeuroptera are divided into three subordinate taxa — Birostrata, + Rhaphidioptera + Mecaptera. Rhaphidioptera and Meganeuroptera. Some authors do The only name whose original circumscription pos- not recognize the taxon Meganeuroptera and accept sibly matches the taxon under consideration is Eu- four directly subordinate taxa — Birostrata, Rhaphid- neuropteroidea Krausse & Wolff, 1919. In our paper, ioptera, Sialis/fg1 and Eumegaloptera (see below). where this name is discussed [Kluge & Krivokhatsky, 1998] an error takes place; there is written: “The taxon 1.1. Birostrata Euneuropteroidea was proposed as a volumetrical tax- Circumscriptional names: on, but not as a ranking one (for explanation — see Euneuropteroidea Krausse & Wolff, 1919 (see com- above). It means, that in further classifications the name ments); Euneuropteroidea should be used for taxa of the same Birostrata Kluge, taxon nov. allowed volume (i.e. including at least type species of In circumscription also matches: Megaloptera SENSU Bur- all families listed above as included, and excluding at meister, 1839 (non Latreille, 1802, nec 1807); Neu- least the taxa of Neuropteroidea listed above as exclud- roptera SENSU Handlirsch, 1903 (non Linnaeus, 1758); ed ones) ...”. This wrong sentence came from the previ- Stegoptera SENSU Banks, 1907 (non Dumeril, 1806); ous version of the text, where we proposed a new name Planipennia SENSU Heymons, 1915 (non Planipennes Latreille, 1817, nec Planipennia Burmeister, 1839). for this taxon, and unfortunately was not corrected. The Hierarchical name: Myrmeleon/f2=Hemerobius/g2 (sine correct text should be: “In further classifications the Raphidia, Chauliodes; incl. Chrysopa, Osmylus, Sisyra, name Euneuropteroidea should be used for taxa of the Coniopteryx, Dilar, Mantispa, Berotha, Ithone, Polysto- same allowed volume (i.e. including at least type spe- echotes, Nevrorthus). cies of all orders listed in Krausse & Wolff [1919] as Typified names in use: Myrmeleonina, Myrmeleontida, included, and excluding all taxa listed in the same paper Hemerobiina, Hemerobiites, Hemerobida, Hemero- as excluded ones) ...”. The ordinal names listed by bidae, Hemerobiida, Hemerobiidae, Hemerobiidi, Krausse & Wolff [1919] as belonging to Euneuropteroi- Hemerobiiformes, Hemerobiodea. dea, are Myrmeleonida, Ascalaphida, Nemopteridida NOMENCLATURAL COMMENTS. Nowadays this (misprint “Neuropteridida”), Sisyrida, Osmylida, Kalli- taxon is often called either Neuroptera or Planipennia, grammida, Nymphitida, Mesochrysopida, Prohemero- but neither name in its original circumscription match this biida, Solenoptilida, Dilarida, Polystoechotidida, taxon. Nymphesida, Hemerobiida, Chrysopida, Mantispida and The name Neuroptera Linnaeus, 1758 was origi- Coniopterygida. nally applied to an order which united genera Libellula Thus, among the taxa originally included into Eu- (matching Odonata), Ephemera (matching Ephemero- neuropteroidea, there are extinct taxa Prohemerobius/ ptera), Phryganea (matching Trichoptera + Plecoptera fg, Solenoptilon/fg, Kalligramma/fg, Nymphites/fg and + part of Meganeuroptera), Hemerobius (matching Biros- Mesochrysopa/fg, known only as fossil adults. The only trata + part of Meganeuroptera + Isoptera + Copeo- reason to unite these taxa with Birostrata, is that their gnatha), Panorpa (matching Mecaptera) and Raphidia wing venation have features of Neuropteroidea (see (matching Rhaphidioptera). Now Neuroptera SENSU Lin- above), but have no special features of Rhaphidioptera, naeus, 1758 is regarded to be a paraphyletic taxon Sialis/fg1 or Eumegaloptera; at the same time, no one of ancestral to other orders of winged insects in the Lin- these taxa has distinct diagnosis basing on wing vena- naean classification. Subsequently, the paraphyletic or- tion. Probably, the taxon Euneuropteroidea is poorly a der Neuroptera was broken down to a number of smaller plesiomorphon, ancestral for Rhaphidioptera and Mega- natural orders, one of which was given the name Neu- neuroptera and existing since Permian (or at least since roptera. MacLeay [1821] designated the genus Libellu- Lias, from which the type-species of Prohemerobius la as the type for the name Neuroptera, so some workers Handlirsch, 1906 was described). splitting Linnaean Neuroptera into smaller orders ap- Unlike Euneuropteroidea, the taxon Birostrata, tax- plied the name Neuroptera to an order including odo- on nov. is holophyletic and characterized by an autapo- nates (Libellula). Others, however, used, and still do, morphy in larval structure (see below); its original listed the name Neuroptera to refer to orders of various cir- membership includes only Myrmeleon/fg, Hemerobius/ cumscriptions not including Libellula. fg, Chrysopa/fg, Osmylus/fg, Sisyra/fg, Coniopteryx/ The name Planipennes Latreille, 1817 is often fg, Dilar/fg, Mantispa/fg, Berotha/fg, Ithone/fg, Polys- spelled out as “Planipennia”, because some people mis- toechotes/fg and Nevrorthus/fg — taxa, for all of which take “Planipennes” for a French spelling; however, this apomorphy is known. If necessary, one can use the elsewhere Latreille [1825] spells “Planipennes” both in name Birostrata as “Neuroptera-Birostrata”, because Latin and French. Originally [Latreille, 1817] the fam- all representatives of Birostrata were originally includ- Metamorphosis and homology of mouthparts in Neuropteroidea 91 ed into the artificial order Neuroptera Linnaeus, 1758. only lacinia and cardo have muscles going to cranium. Unlike Euneuropteroidea, which are reported from Per- Thus, base of mala includes at least lacinia and a distal mian and Lias, the taxon Birostrata possibly is no older portion of stipes movably articulated with the proximal than Late Jurassic. portion of stipes, which retains its normal position, AUTAPOMORPHY OF BIROSTRATA. The only structure and function. known autapomorphy of Birostrata is a unique paired The Rousett’s conclusion, based on musculature, is sucking-piercing mouth apparatus of larva, which in- correct. A question, which was not answered, is which variably exists in all representatives (with this the name maxillary parts attached to stipes and lacinial base “Birostrata” is connected). (lacinia, galea, or palp) constitute the long piercing- LARVAL MOUTH APPARATUS. Besides labrum sucking portion of the maxillary mala, and which are and labium with labial palps, mouth apparatus includes reduced. a pair of unusual piercing-sucking stylets composed by I examined larval/pupal transformation in represen- a pair of modified mandibles and a pair of modified tatives of two species of Chrysopidae and one species of distal portions of maxillae. These movable distal por- Myrmeleontidae. Larvae of Chrysoperla carnea tions of maxillae have complex origin, and I suggest to (Stephens, 1836 [Chrysopa]) were collected in July call them maxillary malae. The mandible and the max- 2004 in Borisovka of Belgorod Province (Russia), and illary mala of each side are coupled together by a kept in cages; 8 pronymphs taken from cocoons were peculiar lock [Gaumont, 1976] and function as an inte- fixed at various phases of development. One pronymph gral movable stylet. Homology of the maxillary malae of Chrysotropia ciliata (Wesmael, 1841 [Chrysopa]) was the subject of discussion and is clarified now (see was got from larva collected in August 2004 in Saint- below). Petersburg (Russia). Larvae of Myrmeleon bore (Tjed- MAXILLARY COMPONENT OF STYLET. Max- er, 1941 [Grocus]) were collected in August 2004 on illa of larval Birostrata consists of a basal portion and a southern shore of Gulf of Finland in Izhora near Saint- distal portion. The basal portion is formed by two Petersburg and kept in cages; 25 pronymphs were fixed sclerites (often called cardo and “stipes”), which consti- at various phases of development. All pronymphs were tute a part of ventral wall of the head. The distal portion dissected and mounted on slides in Canadian balsam. (maxillary mala) is a movably articulated process di- The examined species of Chrysopidae have such a rected anteriorly; it couples with mandible and consti- phase of pronymphal development, when musculature tutes a ventral half of the piercing-sucking stylet. Max- of mouthparts is already lost, hypoderm of mouthparts illa has the following musculature (Fig. 2) [Rousett, is strongly diminished and starts to get the pupal shape, 1966]: (1) intrinsic muscles inside proximal part of the but retains slender stretched tips, which are still inserted maxillary mala (dilators of sucking channel); (2) into the larval cuticular appendages (Figs 4, 6, 11, 12). muscle(s) going from base of the mala into the “stipes” This phase corresponds to the arbitrary 2nd and 3rd (present not in all representatives); (3) strong muscle(s) phases of leg transformation, when the hypodermal going from base of the mala to cranial wall; (4) muscle(s) parts of legs also retain vestigial apical processes [Kluge, going from base of the mala to tentorium; (5) muscle(s) in press: Figs 33a–b, 34, 38, 41]. Presence of the stretched going from the “stipes” to tentorium; (6) muscle(s) vestigial tips of future pupal appendages inserted into going from the cardo to tentorium. Unlike imago and the larval cuticular appendages allows to understand pupa (Fig. 3), larva has no muscle going from the cardo homology between larval and pupal appendages (but to cranium. not homology between their segments, because at least Basing on musculature, Rousett [1966] discussed all in legs ontogenetic homology of segments is broken). possible hypotheses about origin of these parts of max- The examined species of Myrmeleon has no such phase illa. Hypothesis 1: maxillary mala is either lacinia, or of development; when its mouthparts transform from galea; the basal portion of maxilla consists of the whole larval to pupal, their tissues at first degenerate and move stipes and the cardo. Hypothesis 2a: maxillary mala out from the corresponding larval cuticle, and only after includes the whole stipes (possibly with lacinia, galea this get pupal shape. So conclusions about homology of and palp); the basal portion of maxilla represents only maxillary parts are based mainly on development of the cardo divided into two pieces. Hypothesis 2b: max- Chrysopidae. illary mala includes a part of stipes (possibly with Among the examined pronymphs of Chrysopidae, 4 lacinia, galea and palp); the basal portion of maxilla specimens of Chrysoperla carnea and one specimen of consists of the rest part of stipes and the cardo. Presence Chrysotropia ciliata are at the phase when appendages of the muscle going from mala to tentorium excludes the retain hypodermal projections inserted into the larval Hypothesis 1, because only stipes and cardo (but not cuticular appendages. In all these specimens hypoderm lacinia, galea or palp) have muscles going to tentorium. of the future pupal maxilla already has distinguishable Presence of the muscle going from mala to stipes (i.e. to lacinia, galea and palp, and tip of the palp continues to the distal sclerite of basal portion of maxilla) excludes a slender process which is inserted into the larval cutic- the Hypothesis 2a, because only stipes (but not cardo) ular stylet-like distal portion of maxillary mala (Figs 4, has muscles going to lacinia, galea and palp. Presence of 6, 11, 12). At the same time, the future pupal galea the muscle going from mala to cranium excludes as- locates inside the proximal swollen part of the larval sumptions that mala is the galea or the palp, because cuticular maxillary mala, and the future pupal lacinia 92 Nikita J. Kluge

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Figs 2–4. Maxilla of Chrysoperla carnea, ventral view: 2 — left half of head of active larva (only maxillary sclerites dotted; tentorium and maxillary muscles shown by interrupted lines; other muscles not drawn); 3 — maxilla of pupa (anlages of pupal- imaginal muscles shown by interrupted and dotted lines); 4 — maxilla of pronymph at that phase, when under larval cuticle (shown by integral line) hypoderm is detached and starts to get pupal shape (shown by interrupted line). Abbreviations: crd — larval cardo; gal — future pupal galea; lac — future pupal lacinia; mal — larval maxillary mala (distal portion of stipes + lacinia + galea + palp); p.mx — future pupal maxillary palp; stp — larval proximal portion of stipes. Ðèñ. 2–4. Ìàêñèëëà Chrysoperla carnea, âåíòðàëüíî: 2 — ëåâàÿ ïîëîâèíà ãîëîâû àêòèâíîé ëè÷èíêè (òîëüêî ìàêñèëëÿðíûå ñêëåðèòû ïóíêòèðîâàíû; òåíòîðèóì è ìàêñèëëÿðíûå ìûøöû ïîêàçàíû ïðåðûâèñòûìè ëèíèÿìè; äðóãèå ìûøöû íå íàðèñîâàíû); 3 — ìàêñèëëà êóêîëêè (çà÷àòêè êóêîëî÷íî-èìàãèíàëüíûõ ìûøö ïîêàçàíû ïðåðûâèñòûìè è ïóíêòèðíûìè ëèíèÿìè); 4 — ìàêñèëëà ïðîíèìôû íà òîé ôàçå, êîãäà ïîä ëè÷èíî÷íîé êóòèêóëîé (ïîêàçàííîé ñïëîøíîé ëèíèåé) ãèïîäåðìà îòñëîèëàñü è íà÷èíàåò ïðèîáðåòàòü êóêîëî÷íóþ ôîðìó (ïîêàçàíà ïðåðûâèñòîé ëèíèåé). Ñîêðàùåíèÿ: crd — ëè÷èíî÷íîå êàðäî; gal — áóäóùàÿ êóêîëî÷íàÿ ãàëåà; lac — áóäóùàÿ êóêîëî÷íàÿ ëàöèíèÿ; mal — ëè÷èíî÷íàÿ ìàêñèëëÿðíàÿ ìàëà (äèñòàëüíàÿ ÷àñòü ñòèïåñà + ëàöèíèÿ + ãàëåà + ùóïèê); p.mx — áóäóùèé êóêîëî÷íûé ìàêñèëëÿðíûé ùóïèê; stp — ëè÷èíî÷íàÿ ïðîêñèìàëüíàÿ ÷àñòü ñòèïåñà. locates more proximally. In some specimens bound- ed into the larval cuticular mala (in other specimens aries between segments 3, 4 and 5 of the future pupal fixed at the same phases of development, parts of the maxillary palp are already visible (Fig. 6). Among the future pupal maxilla are non-distinguishable). examined pronymphs of Myrmeleon bore, about 8 spec- Such development of pupal maxilla allows to con- imens have more or less distinguishable lacinia, galea clude that in larva of Birostrata maxillary palp has not and palp of the future pupal maxilla, and at the same been reduced, but gave rise to the long, heavily sclero- time retain some connection between it and the larval tized, pointed, non-segmented, stylet-like, distal por- cuticular maxillary mala; in all of them only the future tion of maxillary mala; galea, lacinia and apical portion pupal maxillary palp is connected with, or partly insert- of stipes are completely fused with base of the palp. Metamorphosis and homology of mouthparts in Neuropteroidea 93

7

6

9 8

5 10

Figs 5–10. Head of Chrysoperla carnea: 5–9 — pronymphs: 5 — head of pronymph (in left — dorsal view; in right — ventral view; integral lines — larval cuticle; interrupted lines — hypoderm getting shape of future pupal head and appendages; imaginal eyes dotted); 6 — optic section of maxilla (black filled — optic section of larval cuticle; thin line — outline of hypoderm getting shape of future pupal maxilla); 7 — optic section of mandible (black filled — optic section of larval cuticle; thin line — outline of hypoderm getting shape of future pupal mandible); 8 — hypoderm getting shape of future pupal mandible at next phase of pronymphal development; 9 — the same, at next phase (the same specimen as in Fig. 5); 10 — mandible of pupa. Ðèñ. 5–10. Ãîëîâà Chrysoperla carnea: 5–9 — ïðîíèìôû: 5 — ãîëîâà ïðîíèìôû (ñëåâà — äîðñàëüíî; ñïðàâà — âåíòðàëüíî; ñïëîøíûå ëèíèè — ëè÷èíî÷íàÿ êóòèêóëà; ïðåðûâèñòûå ëèíèè — ãèïîäåðìà, ïðèîáðåòàþùàÿ ôîðìó áóäóùåé êóêîëî÷íîé ãîëîâû è åå ïðèäàòêîâ; èìàãèíàëüíûå ãëàçà ïóíêòèðîâàíû); 6 — îïòè÷åñêèé ñðåç ìàêñèëëû (÷åðíàÿ çàëèâêà — îïòè÷åñêèé ñðåç ëè÷èíî÷íîé êóòèêóëû; òîíêèå ëèíèè — î÷åðòàíèÿ ãèïîäåðìû, ïðèîáðåòàþùåé ôîðìó áóäóùåé êóêîëî÷íîé ìàêñèëëû); 7 — îïòè÷åñêèé ñðåç ìàíäèáóëû (÷åðíàÿ çàëèâêà — îïòè÷åñêèé ñðåç ëè÷èíî÷íîé êóòèêóëû; òîíêèå ëèíèè — î÷åðòàíèÿ ãèïîäåðìû, ïðèîáðåòàþùåé ôîðìó áóäóùåé êóêîëî÷íîé ìàíäèáóëû); 8 — ãèïîäåðìà, ïðèîáðåòàþùàÿ ôîðìó áóäóùåé êóêîëî÷íîé ìàíäèáóëû â ñëåäóþùåé ôàçå ïðîíèìôàëüíîãî ðàçâèòèÿ; 9 — òî æå, â ñëåäóþùåé ôàçå (òîò æå ýêçåìïëÿð, ÷òî íà ðèñ. 5); 10 — ìàíäèáóëà êóêîëêè. 94 Nikita J. Kluge

11

12

Figs 11–12. Chrysopidae, head of pronymph: 11 — Chrysoperla carnea, left half of head, view from median section; 12 — Chrysotropia ciliata, head (in left — dorsal view; in right — ventral view). Larval cuticle shown as colourless; surface of hypoderm (which gets shape of future pupal parts) dotted, hidden parts of hypoderm shown by interrupted line. Abbreviations: ant — future pupal antenna; gal — future pupal galea; lac — future pupal lacinia; md — future pupal mandible; p.lb — future pupal labial palp; p.mx — future pupal maxillary palp. Ðèñ. 11–12. Chrysopidae, ãîëîâà ïðîíèìôû: 11 — Chrysoperla carnea, ëåâàÿ ïîëîâèíà ãîëîâû, âèä ñ ìåäèàëüíîãî ñðåçà; 12 — Chrysotropia ciliata, ãîëîâà (ñëåâà — äîðñàëüíî; ñïðàâà — âåíòðàëüíî). Ëè÷èíî÷íàÿ êóòèêóëà ïîêàçàíà êàê áåñöâåòíàÿ; ïîâåðõíîñòü ãèïîäåðìû ïóíêòèðîâàíà, ñêðûòûå ÷àñòè ãèïîäåðìû ïîêàçàíû ïðåðûâèñòîé ëèíèåé. Ñîêðàùåíèÿ: ant — áóäóùàÿ êóêîëî÷íàÿ àíòåííà; gal — áóäóùàÿ êóêîëî÷íàÿ ãàëåà; lac — áóäóùàÿ êóêîëî÷íàÿ ëàöèíèÿ; md — áóäóùàÿ êóêîëî÷íàÿ ìàíäèáóëà; p.lb — áóäóùèé êóêîëî÷íûé ëàáèàëüíûé ùóïèê; p.mx — áóäóùèé êóêîëî÷íûé ìàêñèëëÿðíûé ùóïèê. Metamorphosis and homology of mouthparts in Neuropteroidea 95

MANDIBULAR COMPONENT OF STYLET. The [1916] decided that the correct spelling of the generic mandibular component of larval piercing-sucking stylet name was not Raphidia, but Rhaphidia; because of this, undoubtedly represents the whole mandible: it has a he used the family name Rhaphididae and created the usual for Metapterygota articulation with head capsule new ordinal name Rhaphidioptera. According the rules and strong muscles going from its base to cranium. of the actual International Code of Zoological Nomen- Pupal and imaginal mandible of Birostrata has a clature [1999], generic name should be used in its distinct pointed apical incisor and a medial biting edge original spelling independently if it corresponds to gram- formed by fused kinetodontium and mola (this edge can matical rules or not (ICZN 32.5.1); subsequent spelling be smooth or dentate, but has kinetodontium and mola different from the original, i.e. an unjustified emenda- indistinguishable). Pupal mandibles are functional; in tion, is regarded to be a junior objective synonym, with course of larval/pupal transformation in pronymphal its own author and date (ICZN 33.2.3); family-group phase, all larval mandibular musculature at first com- name should be formed from the original spelling of pletely disappears, and than new pupal-imaginal man- generic name (ICZN 35.4.1). Because of this, the gener- dibular musculature appears. ic name should be Raphidia Linnaeus, 1758, but not When muscle-less primordium of pupal mandible Rhaphidia Billberg, 1820; family-group names should develops inside the larval cuticular mandible of pro- be formed from Raphidia, but not from Rhaphidia. nymph (at the same phase, as maxilla — see above) its However, these rules can not be spread to the name incisor develops not from the point of larval mandible, Rhaphidioptera: this name does not belong to the family- but as a blunt outgrow of its median side; slender group, because it is formed not from generic name, suffix pointed process, which is a remainder of the larval and ending only, but includes also a base “-pter-”. Unlike mandibular stylet, locates on lateral side of the future typified names (i.e. family-group names in widest sense) pupal mandible (Fig 7). During subsequent develop- the name Rhaphidioptera Navas, 1916 should be used ment, this process becomes shorter and shorter (Figs 8, in its original spelling with its own author and date, and 9), and finally disappears (Fig. 10). the name Raphidioptera Martynov, 1938 should be Such development allows to conclude that the larval regarded as an unjustified emendation, i.e. junior objec- mandibular portion of piercing stylet proceeds not from tive synonym, and used with its own author and date. the incisor, but from a peculiar lateral outgrow of man- AUTAPOMORPHIES OF RHAPHIDIOPTERA. dible. The lateral mandibular outgrowth of larval Biros- This is a small group of insects with rather uniform trata resembles mandibular task of certain larval structure, so it has many autapomorphies [Aspöck & Ephemeroptera [Kluge, 2004]. Aspöck, 2004]. One of autapomorphies is elongate, ASSUMED HOMOLOGY. Possibly, the mandibu- prognathous head with long gula, so bases of all mouth- lar and the maxillary components of stylet originated parts are translocated to the extreme anterior margin of from serially homologous parts of mandible and maxilla the head. — their telopodites. Maxillary telopodite is a maxillary MOUTH APPARATUS. In all stages of develop- palp; in most Hexapoda (including pupa and imago of ment (larva, pupa and imago) head has the same struc- Birostrata) it is distinctly developed and 5-segmented. ture (Figs. 13, 15, 17): it is sharply prognathous, elon- Mandibular telopodite was lost in the common ancestor gate, with long gula; tentorium is vestigial, and posteri- of Hexapoda or earlier, in the common ancestor of or tentorial pits locate at the middle of the long sclero- Atelocerata [Kluge, 2000]. However, it is not excluded tized ventral side of the head far from mouthparts and that the place on mandible where the mandibular te- far from occipit. Maxillae and labium are located ante- lopodite was lost, retains ability to produce a non- riorly; their bases are hidden by anterior margin of the segmented process. Such process could independently gula (Fig. 13). Muscles going from maxillae and labium appear in some non-related taxa of Ephemeroptera- to cranium and tentorium (cardinal-cranial, cardinal- Furcatergaliae, in larval Birostrata and possibly in some tentorial, stipital-tentorial, lacinial-cranial and labial- other insects. tentorial muscles) are strongly elongated, longitudinal, nearly parallel and can somewhat retract the maxillae 1.2. Rhaphidioptera and the labium into the head (Fig. 13). These muscles Circumscriptional names: are equally developed in larva, pupa and imago and Tetramera Billberg, 1820 (non Tertamera Dumeril, 1806); seem to retain without lysis during metamorphosis. Aponeuroptera Crampton, 1916; MAXILLA. Unlike the maxilla initial for Hexapo- Rhaphidioptera Navas, 1916; Raphidioptera Martynov, 1938. da, larval and imaginal maxilla of Rhaphidioptera has Hierarchical name: Raphidia/fg1 (incl. Inocellia) [f: Raphid- stipes cylindrically projected, with developed median inae Latreille, 1810; g: Raphidia Linnaeus, 1758; type- sclerotized wall proximad of lacinia (Fig. 14: stp), so species R. ophiopsis Linnaeus, 1758]. the lacinial-cranial muscle passes trough the stipes (Fig. Typified names in use: Raphidia, Raphidiacea, Raphidia- 13) (such structure is not found in other Neuropteroi- des, Raphididae, Raphidides, Raphidiidae, Raphidi- dea). At all stages, maxillary palp is 5-segmented and idea, Raphidiina, Raphidiides, Raphidina, Raphidi- attached on lateral side of the stipes. nae, Raphidiodea, Raphidioidea, Raphidites. Larval maxilla has no separate lacinia; galea is re- NOMENCLATURAL COMMENTS. Basing on tra- tractable and locates on apex of cylindrical stipes (Figs ditions of transliteration from Greek to Latin, Navás 13, 14). Imaginal maxilla has well-separated lacinia (se- 96 Nikita J. Kluge

14 16

13 15 17

Figs 13–17. Raphidia spp.: 13–15 — larva: 13 — part of head of active larva, ventral view; galea partly retracted into stipes, cardo and base of stipes retracted into head (maxillary muscles and hidden parts of maxilla shown by interrupted lines; other muscles not drawn); 14 — optic section of pronymphal maxilla (black filled — optic section of larval cuticle; thin line — outline of hypoderm getting shape of future pupal maxilla); 15 — larval head and prothorax, lateral view; 16–17 — pupa: 16 — optic section of maxilla (black filled — optic section of pupal cuticle; thin line — outline of imaginal maxilla developing inside); 17 — pupal head and prothorax, lateral view. Abbreviations: crd — cardo; gal — galea; lac — lacinia; stp — median wall of stipes. Ðèñ. 13–17. Raphidia spp.: 13–15 — ëè÷èíêà: 13 — ÷àñòü ãîëîâû àêòèâíîé ëè÷èíêè, âåíòðàëüíî; ãàëåà ÷àñòè÷íî âòÿíóòà â ñòèïåñ, êàðäî è îñíîâàíèå ñòèïåñà âòÿíóòû â ãîëîâó (ìàêñèëëÿðíûå ìûøöû è ñêðûòûå ÷àñòè ìàêñèëëû ïîêàçàíû ïðåðûâèñòûìè ëèíèÿìè; äðóãèå ìûøöû íå íàðèñîâàíû); 14 — îïòè÷åñêèé ñðåç ïðîíèìôàëüíîé ìàêñèëëû (÷åðíàÿ çàëèâêà — îïòè÷åñêèé ñðåç ëè÷èíî÷íîé êóòèêóëû; òîíêàÿ ëèíèÿ — î÷åðòàíèÿ ãèïîäåðìû, ïðèîáðåòàþùåé ôîðìó áóäóùåé êóêîëî÷íîé ìàêñèëëû); 15 — ëè÷èíî÷íàÿ ãîëîâà è ïåðåäíåãðóäü, ëàòåðàëüíî; 16–17 — êóêîëêà: 16 — îïòè÷åñêèé ñðåç ìàêñèëëû (÷åðíàÿ çàëèâêà — îïòè÷åñêèé ñðåç êóêîëî÷íîé êóòèêóëû; òîíêàÿ ëèíèÿ — î÷åðòàíèÿ èìàãèíàëüíîé ìàêñèëëû, ðàçâèâàþùåéñÿ âíóòðè); 17 — êóêîëî÷íàÿ ãîëîâà è ïåðåäíåãðóäü, ëàòåðàëüíî. Ñîêðàùåíèÿ: crd — êàðäî; gal — ãàëåà; lac — ëàöèíèÿ; stp — ìåäèàëüíàÿ ñòåíêà ñòèïåñà. tose, as in other Neuropteroidea); galea is non-retractable MANDIBLE. Pupal mandibles are functional; prob- (Fig. 16). During the larval/pupal and pupal/imaginal trans- ably, mandibles retain functional musculature during all formations imaginal lacinia develops from a median-distal metamorphosis (unlike Birostrata). At pronymphal part of the integral larval stipes, which actually represents phase, the future pupal mandible locates inside the a result of fusion of stipes and lacinia (Figs 14, 15). larval cuticular mandible in such a way, that apex of the Metamorphosis and homology of mouthparts in Neuropteroidea 97 pupal incisor lies inside apex of the larval incisor (as in 1.3.1. Sialis/fg1 Fig. 21) (unlike Birostrata, whose apices of larval and Hierarchical name: Sialis/fg1 [f: Sialida Leach, 1815; g: pupal mandibles are different — see above). Sialis Latreille, 1802, type-species Hemerobius lutarius 1.3. Meganeuroptera Linnaeus, 1758]. Typified names in use: Sialis, Sialida, Sialidae, Sialinae, Circumscriptional names: Sialoidea. Semblides Latreille, 1825 (non Semblis/fg); AUTAPOMORPHIES OF SIALIS/FG1. Known rep- Meganeuroptera Crampton, 1916; Eumegalopterida Krausse & Wolff, 1919. resentatives of this taxon have rather uniform structure, In circumscription also matches: Megaloptera SENSU La- so the taxon has many autapomorphies. Larva has seg- treille, 1807 (non 1802). mentation on the abdominal gill appendages and a unique Hierarchical name: Corydalus/f1=Chauliodes/g1 (incl. Sia- ventral tail on the 10th abdominal segment. lis) [f: Corydalides, Leach 1815; g: Chauliodes Latreille, MOUTH APPARATUS. In most respects mouth 1796, type-species Hemerobius pectinicornis Linnaeus, apparatus of larva, pupa and imago is similar and close 1763 (designated by Latreille, 1802 & 1810)]. to ancestral for Hexapoda; only larval maxillae have Typified names in use: Corydalides, Corydalida. significant difference from the pupal-imaginal ones. NOMENCLATURAL COMMENTS. The oldest MAXILLA. In larva galea and palp arise from a correct circumscriptional name of the taxon under con- common basal segment, and palp looks as 4-segmented sideration is Meganeuroptera Crampton, 1916. The (Fig. 18). In pupa (Fig. 20) and imago labial palp is name Meganeuroptera Crampton, 1916 (formed as separate from galea and distinctly 5-segmented. Röber “mega” + “Neuroptera”) should not be confused with its [1941] wrongly described musculature of larval maxilla junior homonym Meganeuroptera Tillyard, 1918 (formed and confused lacinia with galea. as “Meganeura” + “ptera”). Larvae of Sialis sordida Klingstedt, 1932 ready to Usually this taxon is called Megaloptera; however, in pupation were collected 11 May 2004 at the bank of modern literature the name Megaloptera is used in two Peterhoff Canal (Saint-Petersburg, Russia) in the upper different senses: for the taxon uniting Rhaphidioptera + layer of ground and in rotten wood. Their development Meganeuroptera [Hennig, 1953; Hinton, 1981] or for continued in laboratory. 12 specimens were fixed at, or Meganeuroptera without Rhaphidioptera [Hennig, 1969 near, the moment of larval/pupal ecdysis — 5 larvae and others]. In the past the name Megaloptera was used ready to moult, 4 moulting specimens and 3 pupae just also in a quite different sense — for Birostrata, i.e. for the after moult. They were dissected and mounted on slides taxon which does not include neither Rhaphidioptera, nor in Canadian balsam. Meganeuroptera [Burmeister, 1839 and others]. Origi- Homology between parts of larval and pupal maxil- nally, the name Megaloptera was introduced by Latreille; lae can be clarified (1) by comparison of musculature, in the original publication [Latreille, 1802–1805] this (2) by comparison of setation and (3) by examination of name was used for the taxon uniting genera Chauliodes, development of pupal tissues inside larval cuticle. Corydalus, Sialis and Raphidia — i.e. for Rhaphid- (1) Maxillary musculature is identical in larva, pupa ioptera + Meganeuroptera. In his next publications [La- and imago, and only partly degenerates during larval/ treille, 1806–1809 et al.] this author used the name pupal transformation. Labial palp has only one intrinsic Megaloptera for Meganeuroptera only. Thus, the formal muscle in the 1st segment; other maxillary musculature original circumscription of the name Megaloptera La- is typical for Hexapoda. The cardinal-tentorial, stipital- treille 1802 is Rhaphidioptera + Meganeuroptera. In tentorial, lacinial-cranial, and lacinial-stipital muscles order to avoid confusion, it is better not to use the name are visible at all stages of development. The galeal- Megaloptera, and if accept the taxon uniting Rhaphid- stipital, the both palpal-stipital and the single palpal ioptera + Meganeuroptera, use for it the junior circum- muscles are not seen when larva transforms to pupa scriptional synonym Emmenognatha Borner, 1904. (Fig. 19), but they have identical arrangement in larval STATUS OF MEGANEUROPTERA. Some authors and pupal-imaginal maxilla (Figs 18, 20). In larva the do not recognize this taxon and regard Sialis/fg1 to be a galeal-stipital muscle passes through the common base sister group to Rhaphidioptera, but not to Eumega- of galea and palp, and the single palpal muscle locates in loptera [Štys & Bilinski, 1990]. This conclusion is the same common base (Fig. 18). In pupa and imago the based on the fact that examined species of Sialis and galeal-stipital muscle passes through a separate galeal Rhaphidioptera have telotrophic ovaria of a peculiar pedestal (Fig. 20). This means that the galeal pedestal, structure, while examined species of Eumegaloptera which in pupa and imago is a separate projection of have panoistic ovaria [Büning, 1996]. stipes, in larva is completely fused with the 1st segment There are only a few non-unique derived characters, of palp to form something like a segment. which can be regarded as autapomorphies of Mega- (2) Maxillary setation is identical in larva and pupa neuroptera: larvae are aquatic; larval abdomen bears (Figs 18, 20), but imago has numerous additional setae paired lateral gill appendages (whose structure differs which mask the primary ones. There are following primary in Sialis/fg1 and Eumegaloptera); wings have setae on (larval-pupal) setae: 1 long seta on ventral surface of cardo; membrane. 6 long setae on lateral side of stipes; 1 seta on base of Meganeuroptera are divided into Sialis/fg1 and Eu- lacinia; 5 setae on biting edge of lacinia (3 dorsal, 1 ventral megaloptera. and 1 apical); 1 seta on midlength of galea; some minute 98 Nikita J. Kluge

Fig. 18. Active larva of Sialis sordida, part of head, ventral view. Tentorium and maxillary muscles shown by interrupted lines; other muscles not drawn. Abbreviations as in Figs 19–21. Ðèñ. 18. Àêòèâíàÿ ëè÷èíêà Sialis sordida, ÷àñòü ãîëîâû, âåíòðàëüíî. Òåíòîðèóì è ìàêñèëëÿðíûå ìûøöû ïîêàçàíû ïðåðûâèñòûìè ëèíèÿìè; äðóãèå ìûøöû íå ïîêàçàíû. Ñîêðàùåíèÿ êàê íà ðèñ. 19–21. apical galeal setae; 3 setae on 1st palpal segment; 1 seta (3) At early stage of development of the pupal on lateral side of 3rd palpal segment; 2 setae on medial maxilla under larval cuticle, tips of pupal lacinia, galea side of 4th palpal segment; some minute apical setae on and palp remain inside the homologous larval append- apex of 5th palpal segment. In pronymphal phase pupal ages (Fig. 19); this helps to find correct correspondence setae are visible through larval cuticle, and the setation between the larval and the pupal-imaginal parts. At later allows to see how pupal parts are shifted proximally stage, the pupal lacinia, galea and palp are shifted more relatively to the homologous larval parts. proximally and become crumpled inside the larval sti- Metamorphosis and homology of mouthparts in Neuropteroidea 99

19 20 21

Figs 19–21. Sialis sordida: 19 — maxilla of pronymph, optic section (larval cuticle shown by black as optic section; future pupal maxilla shown as ventral view; retained and partly degenerated muscles shown by interrupted and dotted lines correspondingly); 20 — labium and maxilla of pupa, ventral view (maxillary muscles shown by interrupted lines, other muscles not drawn); 21 — mandible of late pronymph, optic section (larval cuticle shown by black as optic section; outline of future pupal mandible shown by thin line). Abbreviations: crd — cardo; gal — galea; gp — galeal pedestal (distal outgrowth of stipes) adhered to first segment of maxillary palp; lac — lacinia; p1, p2, p3, p4, p5 — 1st–5th segments of maxillary palp; stp — stipes. Ðèñ. 19–21. Sialis sordida: 19 — ìàêñèëëà ïðîíèìôû, îïòè÷åñêèé ñðåç (ëè÷èíî÷íàÿ êóòèêóëà ïîêàçàíà ÷åðíûì êàê îïòè÷åñêèé ñðåç; áóäóùàÿ êóêîëî÷íàÿ ìàêñèëëà ïîêàçàíà ñ âåíòðàëüíîé ñòîðîíû; ñîõðàíèâøèåñÿ è ÷àñòè÷íî äåãåíåðèðîâàâøèå ìûøöû ïîêàçàíû ïðåðûâèñòûìè è ïóíêòèðíûìè ëèíèÿìè ñîîòâåòñòâåííî); 20 — íèæíÿÿ ãóáà è ìàêñèëëà êóêîëêè, âåíòðàëüíî (ìàêñèëëÿðíûå ìûøöû ïîêàçàíû ïðåðûâèñòûìè ëèíèÿìè; ïðî÷èå ìûøöû íå íàðèñîâàíû); 21 — ìàíäèáóëà ïîçäíåé ïðîíèìôû, îïòè÷åñêèé ñðåç (ëè÷èíî÷íàÿ êóòèêóëà ïîêàçàíà ÷åðíûì êàê îïòè÷åñêèé ñðåç; î÷åðòàíèå áóäóùåé êóêîëî÷íîé ìàíäèáóëû ïîêàçàíî òîíêîé ëèíèåé). Ñîêðàùåíèÿ: crd — êàðäî; gal — ãàëåà; gp — ïîñòàìåíò ãàëåè (äèñòàëüíûé âûðîñò ñòèïåñà), ïðîðîñøèé ê ïåðâîìó ÷ëåíèêó ìàêñèëëÿðíîãî ùóïèêà; lac — ëàöèíèÿ; p1, p2, p3, p4, p5 — 1–5-é ÷ëåíèêè ìàêñèëëÿðíîãî ùóïèêà; stp — ñòèïåñ. pes, thus the larval cuticular lacinia, galea and palp of palp is free, and the palp has normal 5-segmented appear to be empty. structure. As can be seen, the larval segment, which looks as a MANDIBLE. Pupal mandibles are functional; man- common base for galea and palp, is actually a result of dibles retain functional musculature during all metamor- fusion of the first segment of palp with a galeal pedestal phosis (unlike Birostrata). At pronymphal phase, the (a projected portion of stipes, to which galea is at- future pupal mandible locates inside the larval cuticular tached). Thus, 4-segmented larval palp consists of ini- mandible in such a way, that apex of pupal incisor lies tial palp segments 2–5. In pupa and imago first segment inside apex of larval incisor (Fig. 21) (unlike Birostrata, 100 Nikita J. Kluge whose apices of larval and pupal mandibles are differ- // International Trust for Zoological Nomenclature, 1999. ent — see above). 306 pp. Kluge N.J. 1999a. A system of alternative nomenclatures of the 1.3.2. Eumegaloptera supraspecific taxa // Entomol. Obozrenie. Vol.78. No.1. P.224–243 [in Russian]. English translation: A system of Circumscriptional name: Eumegaloptera Riek, 1974. alternative nomenclatures of supra-species taxa. Linnaean Hierarchical name: Corydalus/f2=Chauliodes/g2 (sine Sialis). and post-Linnaean principles of systematics // Entomolog- Typified names in use: Corydalida, Corydalidae, Corydali- ical Review. 1999. Vol.79. No.2. P.106–110. nae, Chauliodidae. Kluge N.J. 1999b. Mitos en sistemática y principos de nomencla- AUTAPOMORPHIES OF EUMEGALOPTERA. tura zoológica (Myths in systematics and principles of zoological nomenclature) // A. Melic, J.J. de Haro, M. Mendes Larval pygopod is transformed to a pair of leg-like & I.Ribera (eds). Evolution y Filogenia de Arthropoda / projections, each with two apical claws and a lateral gill Boletin de la Sociedad Entomologica Aragonesa. Vol.26. appendage. At all stages (larva, pupa and imago) head P.347–377. has a large gula (but unlike Rhaphidioptera, retains Kluge N.J. 2000. Sovremennaya sistematika nasekomyh [Modern strongly developed tentorium). Maxilla is modified: systematics of insects. Part I. Principles of systematics of living organisms and general system of insects with classifi- larval, pupal and imaginal stipes is strongly elongate cation of primary wingless and paleopterous insects]. St.- and cylindrically projected (but, unlike Rhaphidioptera, Petersburg: Lan’. 336 pp. [in Russian] without a median wall proximad of lacinia); larval (but Kluge N.J. 2002. Gomologiya rotovyh chastey bloh (Aphaniptera) not pupal and imaginal) lacinia is lost, being completely [Homology of the mouthparts in fleas (Aphaniptera)] // fused with stipes (similar to Rhaphidioptera). Entomol. Obozrenie. Vol.81. No.4. P.808–816. [in Russian] Kluge N.J. 2004. The phylogenetic system of Ephemeroptera. Kluwer External structure of mouthparts of larva, pupa and Academic Publishers. 442 pp. imago, and musculature of larva and imago of Cory- Kluge N.J. 2005. Larval/pupal leg transformation and a new dalus cornutus are adequately described and figured by diagnosis for the taxon Metabola Burmeister, 1832 = Kelsey [1954]. Oligoneoptera Martynov, 1923 // Russian Entomological Journal. Vol.13 (for 2004). No.4. P.189–229. Kluge N.J. & Krivokhatsky V.A. 1998. Neuroptera, Megaloptera, ACKNOWLEDGEMENTS. The work was support- Stegoptera, Planipennia, etc. — where is a true name for the ed by the Federal Russian program “Universities of lacewings (Insecta)? // Kharkov Entomological Society Russia” (grant ÓÐ.07.01.332) and the federal program Gazette. Vol.6. No.1. P.5–15. on supporting of leading scientific schools (grant ÍØ Krausse A. & Wolff M. 1919. Eine Uebersicht über die bisher 2232.2003.4). aufgestellten fossilen und rezenten Insectenordungen // Arch. Naturg. (A). Bd.85. Nr.3. S.151–171. Latreille P.A. 1802–1805. Histoire naturelle, generale et parti- References culiere des Crustaces et des Insectes. T.1–14. Paris. Latreille P.A. 1806–1809. 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