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A d \ a n ~ e \ In Ncuropterology Proceedrng\ ot the Thrrd Internat~onal Syrnposlum on Ncuropterology Bcrg en DaI, Kruger Y a t ~ o n ~ ~ l Park, R S A 1988 Manwll, M W & A\pock, H (Eds) Pretoria, R S A 1990 Pp 151 169

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Classification of the Myrmeleontidae based on larvae (Insecta: Neuroptera) l

Lionel A. STANGE and Robert B. MILLER Florida Dept. of Agriculture and Corisu~ner Services, Gainesville, U.S.A. &

Project City. California, U.S.A.

ARSTRACT

A classification of the ktriiily Myr~neleontidac is pre\ented, based on studies of 223 specres representing 58 genera known in the larval \tage. This clasbiiication is in close agreerrlerlt with [he currerlt classification of the family hased on adult morphology. Only one tribe (Maulini) is unknown in the larval stage and its relationships are not clear. The larval characters sugge\t that the Bmchynernurini should be classified with the Myrmecaeluri- ni, Gepini and Isoleontini. However, there arc a few gcncra of Bsachqnemurini which are structurally discor- dant and lnay thrrn a separate group. The largest tribe. Nernoleontini, should include genera classified in the Dimarellini and Cilenurini.

Separation of subfanr~lies based on larvae is not clearly supported by larval characters. However, most of the tribes can be defined by synapomorphic characters. The Dendroleontini may not be a natural group since only four o l the 31 described genera arc known In the larval stage, Tribal drfferences are expressed in a key to tribes. Important characters that separate tribes appear to be the relative length of the labial palp (coniparcd to basal mandibular width), presence of hlatlc-like digging setae on sternite IX, mandibular dentition, development of dolichastera, loss of the subrrredial tooth on stcrnite V111 and the development of a mesothoracic spiracular tubercle. A tliicussion of the characters that delimrt genera IS also presented.

Key words: Myrmeleontidae, larvae, ~norphology, classification, t;txonorny, subfamilies, tribes, genera.

INTRODUCTION

Our present classification of the family Myrmeleontidae into subfamilies, tribes and genera is still quite problematic with many questions to settle. It is, of course, based mainly on adult morphology. One problem has been that most workers have dealt only with a limit- ed fauna. On a world-wide scale the definition of higher categories is less clear. It is appar- ent that no one character is completely reliable. The lack of tibial spurs, for example, can vary within a genus as is quite clcar in the genera Bruchyrzemurus, Ahatoleon and Dimur-ellti. Larvae offer another completely different set of characters. However, since

( I ) Contr rbutron N o 678, Bureau of Fntornology. Florrd,~ Depdrtment o f Agrrculture and Conqurner Services, Ga~rle \ \ rlle, FL 32602

L. A. Stnnge & R. B. Miller

larvae live in a diffcrcnt environnlcnt froin ~~du l t s , the selective pressul-ks have probably been quite dissimilar. ant1 the rate of change Inay be faster in adults than larvae in some lines of evolution but Just the opposite in othcrs. Furthermore, as with adults, basing genera or tribes on one character can be misleatiing.

It is also important to know the nmicroenvirvnnlcnt of the larva because. as we will show later, this can profoundly affect the structure of the larva without necessarily being an indication of phylogenetic separation. Of course, the more species we know in the larval stage, the better our understanding of the relationships in the family.

At present we know about 60 gcncra in the larval stage representing all the currently recog- nized subfamilies, tribcs (except Maulini) and about 30 percent of the genera. The reader is referred to Gepp (1984) for most of thc published accounts and pertinent literature. Important references since Gepp have been a study of the acanthaclisine larvae (Stange & Miller 1985), P~rrrgI~nurus (Tanaka 1979). Dictyoleon (New 1982a), Nnvusoleon (Miller & Stange 1985), Nophis (Sinmon 1985), Gymnocnemia (Insom et al. 1985), Cymothules (Mansell 1987), European larvae (Friheden 1973) and Florida larvae (Stange 1980; Lucas & Stange 1981). Mansell, Miller and Stange have unpublished notes on many other African and Anmerican genera which have been used in this review. The diversity of larval types is better understood in some tribes than in othcrs. For example, we know eight of the 15 genera of Acanthaclisini, 12 of the 22 genera of the new world Brachynemurina, but we know only six of the 15 genera of the Palparinae, and only one genus (Dendroleon) of the highly diverse Australian fauna of Dendroleontini. The classification of the Aus- tralian Dendroleontini is probably the least understood of all groups of antlions. Thc great diversity in fenlale terminalia of this Australian group suggests that the larvae might like- wise be diverse in structure and in microenvironnmental requirements.

We should recognize that the related family Ascalaphidae is not clearly distinct from the Myrnmeleontidae in all larval characters. We know that scoli occur in both families, ap- parently associated with a nmore free-living existence (not in sand or soil, but uncovered) that demands better camouflage and protection. The scoli allow the larva to cling better to the substrate, especially during predatory or defensive Inaneuvers. The cordate head found in some Ascalaphidae is not found in others (Pieper & Willman 1980). The most significant difference between the two familics appears to be the enlarged hind pretarsal claws of the Myrrneleontidae. However, a few species of antlions (i.e. Nu~~(~.s(>leon) have the hind pretarsal claws somewhat reduced, apparently a secondary condition due to the niche that these larvae occupy. Probably other rock-face inhabiting nmyrrneleontid larvae will be found and may show even more extreme reduction of the pretarsal claws. As WC

will discuss later, ant-lion larvae that have adapted to living on bare rock-faces have some interesting convergences in larval structure.

VALUE OF LARVAE IN DEFINING SUBFAMILIES

We recognize three subfarnilies, Stilbopteryginae, Palparinae and Myrmeleontinae. This is in agreement with most contemporary workers although New (1985) also recognizes the Acanthaclisinae. We know relatively few larvae of the Palparinae and Stilbopterygi- nae which limits our discussion. Nearly all of the Palparinae appear to have one synapo- morphic character which helps to characterize the group. This is the dcvclopmcnt of a pair of blade-like digging setae on the posterior margin of sternite IX (Fig. I ) . This prob- ably evolved in relation to the large size of most of the species which burrow into the sand. The larvae of the Stilbopteryginae also attain large size, but instead of blade-like setae they have at the posterior margin a pair of large submedial processes which bear several digging setae. This is also found in the Peruvian Dirnmres and also apparently in one un-

CIussij7cation 5 f Mymeleontidae

determined palparine larva recently discovered in Israel by Sirnon (Pers. Cnmrn.). Also, !his srructurc occurs in the Myrmccaelurini. This structure may be the precursor of thc btadc-like sctac. and in the Asgenrine Dimnres (Fig. 1) there is a double blade-like struc-

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L.A. Stungr & R.B. Miller

turc. The lisual number of teeth in the Myrmeleontinae and also of the one known species of Stilbopteryginac is three. with only a few exceptions. G1ctzlirli.s and one species of Ccv- troc.1isi.t have two, whereas Pnrtrrirhuc~1i.si.s (Fig. 5 ) has only one tooth. In the Palparinae the number of teeth varies from three to six. The Myrmeleontinac have some synapo- rnorphic characters which unfortunately occur sporadically among the tribes. Thcsc arc the development of a tubercle which bears the mesothoracic spiraclc, and the loss of the submedial teeth o n sternite VIII. At present we can only hypothesize that the subfalllilies are monophyletic and hope that the characters used here for definition will hold up as further species arc discovered.

VALUE OF LARVAL CHARACTERS FOR DEFINING TRIBES

Based on our studies of larvae representing about 30 percent of the described genera, we have modified the tribal classification to a considerable extent. mainly by combining rnany of the tribe\ recognized by Markl (1954) and others.

The largest tribe in terms ofnurnbcr of genera is the Ncmoleontini, which now combines the Glenurini, Protoplcctrini. Obini, Nyutini, Dirnarellini and Distolcontini. This tribe is fairly uniform in larval characters except for a very few genera (i.e. Nu~.usolrotz) which do not live in sandy habitats. Also, we now place the Brachynemurini, Gepini, Isoleontini and Nesoleontini in the tribe Myrmecaelurini. Both larval and adult characters support this taxonomic treatment. The Dendroleontini are the rnost difficult to define, since few genera are known in the larval stage. There is an obvious close relationship to the Neniolcon- tini and further study especially of the rich Australian fauna will no doubt modify our view on the content of the tribe. The tribes Dimarini, Acanthaclisini and Myrmeleontini appear to be well defined in the larval stzge. The only tribe u,~known in the larval stage is the Maulini with only two known genera both occurin5 in southern Africa. Adult struc- tures (i.e. structure of labial palps. lack of fen~vral sense hair) suggest closer relat~onship with the Palparinae than with thc Myrmeleontinae. but the discovery of the larval stage will probably decide the question.

Characters which serve to distinguish tribes are the following: (1) length of the labial pal- pus compared to basal width of mandible; (2) whether the rncsothoracic spiracle is borne on a tubercle or nl>t; (3) presence or absence of submedial teeth on sternitc VIII; (4) dis- tribution and configuration of teeth on mandible; ( 5 ) presence or absence of specialized setae (clolichastcrs; hair tufts: modified digging setae).

We recognize nine tribcs at present: Stilboptcrygini, Palparini, Dirnarini, Maulini, Nemoleontini. Dendr~~lcontini, Myrrric;contini, Acanthaclisini and Myrn~ccaclurini. The small tribe MAULINI is poorly known taxonomically, and the larval stagc is completely unknown. so this is a doubtful tribal taxon. Of the other eight, all can be fairly well de- fined in the larval stagc, except for the Dendroleontini. There are, of course, some dift'icult genera of ant-lions. such as Lpn~o1~!rir:.s. Gtio/)holro!l and others, which especially in the larval stage, are not ciza~,ly assignable to any cxisting tribe. 'The PALPAZINI (Fig. 4) have mandibles longer than the head capsule, three to six mandibular teeth and one or two pairs of blade-like setae on posterior margin of sternite IX. Larvae of P~11p~1ridiu.s havc been found. The elongate mandible with three sharp tecth agrees with thc characters of Palparini rather than Dimarini.

MY RMELEONTINI, DIMARINI and ACANTHACLISINI share the short labial palpus (shorter than mandibular width) and lack the lnesothoracic spiracular tubercle. Myrmeleon- tini larvae have the submedian teeth on stcrnitc VIII; those of Acanthaclisini (Fig. 7) lack them, whereas Dimarini is variable in this character. Dimarini (Figs 1 , 2) and many Acan- thaclisini (Figs S, 6) havc relatively short. thick mandibles, often with reduction in teeth

Classification of Myrmeleonridae

or crowding together of the teeth. Dinfarini has the bluntest teeth known in the family. MYRMECAELURINT (Figs 12, 13, 15) are characterized by having the distal tooth equal to or shorter than the middle tooth, distal tooth often set at different angle. and middle roorh usualIy closcr to distal tooth than basal one. Certain New World Brachynemurina (Fig. 15) are quite bizarre in having the bases of the mandibles close together. in having abdominal scoli, and in othcr fcnturcs, Gnopholeon, Menkeleon and Tytrholeon represent these stranse genera which probably descsve at least subtribal recognition. The largest tribe, NEMOLEONTINI, is typically a uniform group with three parallel teeth (Fig. 10). on the rnnndihlc which gradually increase in length toward the apex. However, thcre are a few hizarrc genera such as GI~~~rrnrs which has only two teeth and a two-segmented labial palpus, and which lives in leaf litter. Also, the larvae of hrovnsol~on and at lcast cmc species of Ercrnol~on live on hare rock and have well dcvcloped abdominal scoii. elongate mandibles, with teeth distally placed, reduced digging sctae, etc.

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L.A. Stange & R.B. Miller

GENERA AS DEFINED BY LARVAE

As one might expect, some genera are easily defined by larval characters whereas in other cases several genera have structurally identical larvae. 'The NEMOLEONTINI offer the best contrast in this regard. Adult structure is fairly siniilar between New World genera Glenur~ls and Eremoleon less so between Na~wsoleon and Araucaleotz and these genera, and with significant adult structural differences between Dimarellu and Psammoleotl.

We now know the larvae of the aforementioned genera. Glenurus larvae are the most dis- tinctive, having only two mandibular teeth and only two segments in the labial palpus. Although the five species of Glenurus known in the larval stage are uniform in these charac- ters, they differ in many others, such as relative head width and chaetotaxy. All of these larvae live in litter, possibly associated with animal dung, and are slow-moving. Yet, adult structure is quite similar to Erernoleorz. The larvae have clearly shown Glenurus to be a valid genus.

Dimarellu has been classified in a different tribe from Nemoleontini since Markl's 1954 basic wing venation classification. Studies of Dtmarellu and the related Pachyleon re- vealed other differences in the head, terminalia and other features which suggest that this group is quite isolated from other Nemoleontini. However, the larvae are very similar to Psarnmoleon larvae and no significant structural differences can be found between these genera. The larval stage in this case is of value in showing that Dimarellu and allies are not distantly removed from other genera. However, the larva does not provide good generic separation. This is comn~onplace in the Nelnoleontini since Old World larvae that we have studied, such as Neuroleon, Creoleon, Obus, Disroleon, Gymnocnernia and Megistopus are very difficult to separate on larval characters. The adults have good structural differ- ences and some of these genera have been placed in different tribes.

Recently we have found some unusual larvae belonging to the Nemoleontini that live on bare rock faces. Nearly all the larvae of the Nemoleontini to date have been found in sand or loose soil in the open, or often in protected areas such as rock overhangs or cave mouths. All of ihese larvae that basically do the same thing are quite sirnilar in structure. Some of the rock overhang species, such as many Erenzoleon, have evolved longer mandibles and creep rather than run and can climb rock faces easily. Eremoleon contains about 30 species in the New World. We have reared most of them. All but three species live under rock overhangs or cave mouths. Many anchor themselves to the substrate covered by only fine dust. Apparently, on the Island of Hispaniola in the Caribbean, where five species of Ercwzolcon occur, one species has evolved to living on bare rock on the ceiling rather than being covered by fine dust in hidden places. These larvae are quite sessile, flattened, with abdominal scoli. When we first found them we thought they were a species of Navusoleotz whose larvae (two species known) live on bare rock. 'The adults of this Hispanio- lan Eremoleon and E. rerverinus Navis are almost identical in every respect and have been confused in the past by taxonomists. The female terlninalia do differ, however, and once the situation was revealed, several other minor characters were found to distinguish these two species. What we have is a striking larval evolution in structure (scoli, mandi- ble, digging setae) due to a new niche but almost no character differentiation in the adult. Eremoleon is of further interest because a second species (E. fernoralis Banks) has adapt- cd to living in tree holes, not large. gaping ones, but any small burrow that the s~uall larva can squeeze into. However, this tree hole larva does not show any striking structur- al differences from most other Eremoleotz. Another Eremoleon, E. pullens Banks, is a true cave inhabitant. This ant-lion apparently stays in the same cave generation after gener- ation without leaving. Both the larvae and adults can live in permanently dark parts of the cave, larvae feeding on dermestid larvae and other insects in bat guano, the adults feeding on miscellaneous insects, including, apparently, bed bugs of the bats.

Classification of M,vrmeleontidae

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L. A. S t a n g ~ & R. B. Miller

The larva of Eremoleon palletzs (Fig. 9) differs from all other Eretnolcotz larvae by having very short mandibles. In the relatcd genus Pstlnzmol~on a similar situation occurs where nearly all species have relatively long mandibles, but one specics, P. tninor Banks (Fig. I I), has unusually short ones. Although this larva lives in a specialized niche, organic material and sand under fallen palm fronds, the habitat would not seem to offer any clear explanation as to the modification. The larva probably spends its whole life in a very restricted, dark area (similar perhaps to E. pullens) with abundant food, and shorter man- dibles have evolved.

Gcncric characters in the Ncrnoleontini are as follows: ( l ) number and configuration of mandible tecth; (2) abdominal spiracles borne on tubercles or not; and (3) presence or abscnce of cubmedial teeth on stcrnite VIII.

ACANTHAC1,ISINI have some excellent generic characters (Stange & Miller 1985). Eight genera are known in the larval stagc. Gencra differ in the niandibular teeth and setae, chactotaxy of sternite VIII and shape of the anterior margin of the clypeal-labrum. At least two genera have larvae which can only rnove backwards (Vella, Ph~lnoclisis). Two gcncra, Parunthac1i.sis and Coritroc.lisis, have a reduced number of mandibular teeth, short antennae and short, powerful jaws.

Five genera are known in the tribe MYRMELEONTTNI: Bnligu, Myrmrleon, Dictyoleon. E~iroleon and Porrerils. Larvae in these genera :ire all very similar without apparent diffcr- ences at the gcncric levcl. They arc all highly specialized, in only rnoving backwards and constructing pitfall traps. Porrcrus was considered to belong to a different tribc until the larvae were discovered. The adults have a number of structural differences, but the larvae of Myrmeleon and Porrerus arc very similar in structure and behavior.

The genera of the MYRMECAELURINI are often quite distinctive. A few such as CIIP- ta, Isoleon, Myrmccaelurus and Scotoleon have species which make pitfall traps. As far as is known, all can move forwards. We have studied larvae of Nophis, Cuetu, Gepus, Solter, M~lmt.undu. Isolron, LOI)PZLIS, M p r m ~ c ~ ~ e l ~ i r u s and most of the New World gencra of Brachynernurina.

Most of these genera are structurally sirnilar with the same kind of mandibular dentition with the distal tooth srnall and set off at a diffcrcnt angle. However, there are a few New World genera (Mettkeleotz, Tyttholcon, Gnopholeon) which arc very diffcrent. In these genera the mandibular bases arc close together, there are sometimes abdominal scoli, and certain other differences in head structure.

The Old World gencra of Myrmecaclurini can be divided into two subtribcs, Myrmecae- lurina (A~ l zuno l~o t z , A.spoeckiatza, Gepellu, Gepus, Irtmolron, I.soleorl, Lopezus, Myr- rnecuelurus, Murtlcundu, Mongolean, Furgella, Napa, Nophis, Soltcr) and Nesoleontina (Cirefcr, Nudus, Nesoleon) based on wing venation. These subtribes cannot be separated at this time nor complctcly from the New World subtribe Brachynernurina based on lar- vae. However, many gcncric characters are present in the tribe, such as: (1) presence or absence of submedial tecth on sternite VIII (absent in Nol,his, Scotoleon, present in the rest); (2) dentition and length of mandible; and (3) modification of abdominal spiracles.

As we have stated earlier, only a few gcncra of DENDROL,EONTINI are known. We have reared Derldroleon and Pieholeon. Mansell (1987) has described the larvae of Cymofhales. Significant data on other genera of the tribc are not available. These larvae have one characteristic in common which separates them from others; the presence of a setal tuft medially on the mesoscutum. A similar type of tuft occurs in I~molcrnus from Chile. sug- gesting some relationship. However, in the case of Lxnzolemus the tufts are serial on several abdominal segments and the middle mandibular tooth is closer to the distal one. We are

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L. A. Stange & R. B. Miller

still studying the relationships of Lemolemus: it might belong in the Myrmecaelurini or the Dendroleontini, or its own tribe.

It is likely, as additional genera of this large tribe are discovered, that the tuft of setae will not hold up as a tribal character. This is in some ways one of the most primitive groups of antlions so that i t may lack synapomorphies which help define other groups.

Genera of the PALPARIN1 appear to have good generic differences in the number of man- dibular teeth (three to six) and the shape of specialized digging setae on sternite IX. However, few species are known to us.

The DIMARINI have two genera in the New World and one in the Old World. Only the New World genera are known in the larval stage. The larvae are sluggish creatures, liv- ing deep in sand, can live for many years (at least six years) before pupating and have very short mandibles with extremely blunt teeth. The two genera are quite different. One has two nlandibular teeth, no specialized digging setae on sternite IX and no submedial teeth on sternite VIII. Dirrlczrcs itself has three mandibular teeth, two pairs of specialized digging setae and has prominent submedial teeth.

CLASSIFICATION OF THE MYRMELEONTIDAE. BASED ON LARVAE (223 SPECIES IN 58 GENERA)

Subfamily Stilbopteryginae van der Weele, 1908 (2 genera - Australia). Larvae known: Stilbopteryx (1).

Subfamily Palparinae Banks, 191 1 (16 genera - worldwide except Australia, North America). Tribe Dimarini Navas, 1914 (2 genera - South America, l genus - Palaearctic). =Echthromyrmicini Markl, 1954. Larvae known: Dimres (3). Tribe Palparini Banks, 191 1 (1 1 genera - Old World except Australia). = Palparidiini Markl, 1954. Larvae known: Crumhornorphus (1); Go1ufiu.s ( ? 1 ); Nosu (2); Palpares (3); Pa1paridiu.s (1 ) . Tribe Maulini Markl, 1954 (2 genera - southern Africa). Larvae unknown (may be referable to Myrmeleontinae).

Subfamily Myrmeleontinae Latreille, 1802 (1 65 genera - worldwide). Tribe Acanthaclisini Navas, 1912 (15 genera - worldwide). Larvae known: Acunthuc1i.si.s ( 1 ) ; Centroclisis (3); Faclrinu (2) ; Heocli.sis (2); Purunthaclisis (4) ; Synclisis (1); Syngenes (1); Vellu (3).

Tribe Dendroleontini Banks, 1899 (32 genera - worldwide except South America). Larvae known: Cymotha1e.c (2); Denclroleon (4); Tricholeon (2).

Tribe Myrmecaelurini Esben-Petersen, 1918 (35 genera - worldwide except Australia). Subtribe Brachynemurina Banks, 1927 (22 genera - New World). = Austroleonina Banks, 1943. Larvae known: Abatoleon (8); Ameromyiu (2); Austmleon (2); Brachynemurus (9); Chuetoleon (1); Chzopholeon ( l ) ; Lenzolemus ( 1 ) ; Menkeleon ( l ) ; Scotoleon (17); Tyttholeon (l) . Subtribe Myrmecaelurina Esben-Petersen, 191 8 (9 genera - Old World except Australia). = Lopezini Esben-Petersen, 19 18.

Classification of Myrrneleontidae

=Gepini Markl, 1954. =Isoleonini Holzel, 1972. Larvae known: Furgellu (1); Gepus (2); Isoleon (1); Lopezus ( 1); Maracanda (I); Myrrnecaelurus (2 ) ; Nophis (2); Solter (1). Subtribe Nesoleontina Markl, 1954 (3 genera - southern Africa). Larvae known: Cuetu (7).

Tribe Myrmeleontini Latreille, 1802 (7 genera - Cosmopolitan). = Porrerini Navas, 1913. =Grapini Navas, 1932.

Larvae known: Baligu (2); Dictyoleon (1); Euroleon ( l ) ; Myrmeleon (40+); Porrerus (3).

Tribe Nemoleontini Banks, 191 1 (76 genera - Cosmopolitan). = Forn~icaleonini Navas, 19 12. = Neuroleini Navlis, 19 12. =Creagrini Navlis, 1912. = Megistopini Navas, 1912. =Gymnocnemini Navas, 19 12. = Protoplectrini Tillyard, 1916. =Glenurini Banks, 1927. = Macronemurini Esben-Petersen, 19 18. =Nyutini Markl, 1954. = Dimarellini Markl, 1954. = Distoleonini Holzel, 1972.

Larvae known: Araucaleon (I); Creoleon (4); Dimarella (10); Distoleon (5); Eremoleon (20); Glenurus (5); Gymnocnemiu (1); Mucronernurus (1); Megistopus ( I ) ; Navusoleon (2); Neuroleon (8); Obus (2); Puruglenurus ( I ) ; Psummoleon (1 7); Xuntholeon ( I ) .

KEY TO TFUBES OF MYRMELEONTIDAE BASED ON LARVAE I . Abdominal sternite IX with enlarged, blade-like setae submedially at posterior mar-

gin (Figs 1, 4) or mandible with 2 teeth with vestigial apical seta (Fig. 3); mesothorac- ic spiracle not borne on tubercle (Palparinae) ........... . . . . .. . . . . . . ............ . . ... 2

Abdominal sternite IX without blade-like digging setae (Fig. 7) although sometimes several large digging setae borne on common base; mandible with 3 teeth or if reduced to 1 or 2 teeth without trace of apical seta; mesothoracic spiracle sometimes borne on tubercle .................................................................................. 3

2. Mandible elongate (Fig. 4), usually longer than head capsule, sometimes with 4 or more sharply-pointed teeth; labial palpus and antenna longer than basal width of mandible; modified digging seta single-bladed; Old World except Australia . . . . . ........................................................................................ Palparini

Mandible shorter than head capsule, with 2 or 3 teeth thakire blunt apically or tipped with vestigial seta; labial palpus and antenna shorter than basal width of mandible (Figs 2, 3); modified digging seta at posterior margin of sternite IX absent or double- bladed (Fig. 1); South America ... ......... ...... .. . . ..... . . . .... . . . . . . . . . . Dimarini

3 . Ocular tubercle longer than medial width with short, thick apical dolichasters; ab- dominal sternitc 1X with submedial process at posterior rnargin bearing several well- developed digging sctae; antennal tubercle with several dolichasters longer than basal antennal segment; mandible with dolichasters on median margin; Australia (Stilbopteryginae) .......................................................... Stilboptergini

L.A. Stunge & R. B. Miller

Without above cornbination of characters (Myrmeleontinae) ...................... 4

4. Labial palpus shorter than basal width of mandible; mesothoracic spiracle not borne on tubercle; head without dolichasters; backward movement only (many genera) or backward and forward movement ....................................................... 5

Labial palpus longer than basal width of mandible (Fig. 15) or mesothoracic spira- cle borne on tubercle; head often with dolichasters (Fig. l l ) ; backward and for-

............................................................................. ward movement 6

5 . Mandible with some setae on exterior margin as long or longer than greatest man- dibular width (Fig. 14); sternite V111 with pair of inconspicuous submedian teeth near posterior margin; make pitfall traps ............................. Myrmeleontini

Mandible with longest setae on exterior margin less than one-half greatest mandibu- lar width (Fig. 6); sternite V111 without teeth on subapical margin (Fig. 7); pitfall

.................................................................. traps absent Acanthaclisini

6. Mandible with distal tooth equal to or shorter than middle tooth, distal tooth often set at different angle (acute projecting anteriorly) (Figs 12, 13, 15); usually middle tooth closer to distal tooth than to basal tooth; (Fig. 15) sternite V111 often with pair of inconspicuous submedial teeth near posterior margin (absent in Nophis, Scotoleon); scoli usually absent on abdomen and much reduced on thorax (known exceptions

......................................... are Gnopholeon and Jafielia) Myrmecaelurini

Mandible with distal tooth longer than middle tooth (Fig. 10) (Glenurus with 2 teeth), teeth more or less parallel and usually equidistant (Fig. 10); sternite VIII with or without submedian teeth; scoli often developed on thorax, less common on abdo- men ...................................................................................... 7

7 . Mesoscutum with tuft of setae at middle (Fig. 8); abdominal sternite IX at least as ................................................................ long as wide Dendroleontini

Mesoscutum without tuft of setae at middle; abdominal sternite IX wider than long ............................................................................. Nemoleontini

APPENDIX A:

PUBLISHED DESCRIPTIVE ACCOUNTS OF ANT-LION LARVAE P

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SPECIES pp P P--

LOCALITY REFERENCES p - P - - -

PALPARINAE PALPARINI

Palpares Rambur

1. cephulotes (Klug) E ~ Y pt

2. hispanus Hagen Europe

3. latipennis Rambur Congo

4. libelluloides (Linnaeus) Europe

Redtenbacher 1884b

McLachlan 1873; Navas 1936

McLachlan 1873; Redtenbacher 1884b

Hagen 1873; Withycombe 1925; Willmann 1977

Clussij?cution of Myrrneleontidue

STILBOPTERYGINAE Stilboptrvx Newman

5. 1itzeuri.s Navas Australia McFarland 1968; New 1982b

MYRMELEONTINAE ACANTHACLISINI Acanthuc~1isi.s Ralnbur

6. occitutlicci (Villers) Europe Redtenbacher 1884b; Steffan 1975

7. pulliclu McLachlan

8. species

Centrocli,\is Nav6s

9. pwzctuluta Navlis

Fadrrna Navas

10. species

Heoclisis Navas

11. filnduta (Walker)

Russia

Egypt

Luppova 1969

Stange & Miller 1985

Tunisia Stange & Miller 1985

South Africa Stange & Miller 1985

Australia Mathews 1950; Tillyard 1926; Adams 1936

Stange & Miller 1985 12. species

Phanoclisis Banks

13. longicollis (Rambur)

Purunthuclisis Banks

14. congener (Hagen)

15. hugeni Banks

16. specics

Australia

Tunisia Stange & Miller 1985

USA

USA

Florida (USA)

Stange & Miller 1985

Stange' & Miller 1985

Stange 1980; Hagen 1887; Stange & Miller 1985

Synclisis Navas

17. hueticu (Rambur) Europe

Tunisia

Redtenbacher 1884b; Principi 1947; Richard 1952; Stange & Miller 1985

Syngenes Kolbe

18. Iongicornis (Rambur)

Vellu Navas

19. urncricurza (Drury)

South Africa Stange & Miller 1985

Florida (USA) Hagen 1873; 1887; Redtenbacher 1884b; Stange & Miller 1985

20. j&llr~x (Rambur) USA; Mexico Stange & Miller 1985

L.A. Stange & R.B. Miller

DENDROLEONTINI Cyrnothules Gerstacckcr

2 1 . ec.c.et1tro.t (Walker)

Dendroleotz Brauer

22. ohsoletus (Say)

23. puntherinus (Fabricius)

24. jezoetzsis Okamoto

MYRMECAELURINI Hrachynemurina Brachynemurus Hagen

25. jkrox (Walker)

26. nebulosus Brach

27. signutus (Hagen)

28. ahdominulis (Say)

Scotol~on Banks

29. pallidus (Banks)

30. peregrinus (Hagen)

3 1. quadripunctatus (Currie)

Myrmecaelurina

Myrmecuelurus Costa

32. trigrammus (Pallas)

Nophis Navas

33. rrilhardi Navas

Nesoleontina

Cuetu Navas

34. beieri Hiilzel

35. lineosa (Rambur)

South Africa Mansell 1987

Florida (USA) Redtenbacher 1884b; Stange 1980

Europe Brauer 1867; Redtenbacher 1884b; Hagen 1873; Roubal 1936; Steffan 1975

Japan Baba & Edashige 1955

California (USA) Stange 1970

Florida (USA) Brach 1978; Stange 1970

New Jersey (USA) Stange 1970

South Dakota (USA) Stange 1970

California (USA) Wheeler 1930; Stange 1970

Cal i f~miz (USA) Stange 1970

California (USA) Stange 1970

Europe

Israel

Redtenbacher 1883; 1884b Steffan 1975; Willmann 1977

Simon 1985

Europe Willmann 1977

Lackinger 1973

Classi$cation of My rmeleontidae

MYRMELEONTINI

Dictyoleon Esben-Petersen

36. rzervosus Esben-Petersen Fiji

Euroleon Esben-Petersen

37. rlostrus (Fourcroy) Europe

Myrmeleon Linnaeus

38. hore (Tjeder)

39. carolirzus Banks

40. crudelis Hagen

41 . fusciarus (Navis)

42. formicurius Linnaeus

43. fronrulis Burrneister

44. gerlindae Holzel

45. hyalinus Olivier

46. irnmacu1utu.s DeGeer

47. inconspicuus (Rambur)

48. insertus Hagen

49. junuuriu~ (Navzis)

50. rrlobili~ Hagen

5 1 . rusrit u~ Hagen

52. ".,ingrilur~.\ " Westwood

53. tcwrnlu Bankj

Sweden

Florida (USA)

Florida (USA)

Dodecanese Islands

Europe

Java

Europe

Dodecanese Islands

eastern USA

Europe

Florida (USA)

Brazil

Florida (USA)

Texas (USA)

Indla

Tcxas (USA)

New 1982a

Redtenbacher 1884a, b; Eglin 1939; Principi 1943; Friheden 1973; Steffan 1975; Bernard 1963

Dewitz 1882: Ohnl 1965; Friheden 1973

Lucas & Stange 1981

Lucas & Stange 1981

Willmann 1977

Redtenbacher 1884b; Doflcin 1916; Eglin 1939; Friheden 1973; Steffan 1975

van der Weele 1909; Jacobson 1912

Hiilzel 1974

Willmann 1977

Hagen 1873; Rcdtenbacher 1884b; Lucas & Stange 1981

Redtenbacher 1883; Principi 1943; Steffan 1975

Lucas & Stange 198 1

Jurberg 1963

Lucas & Stange 198 l

McClendon 1902

Ch~tnis 1960

McClendon 1902

L.A. Stangc. & R. B. Miller

NEMOLEONTINI

Creolron Tillyard

54. lugclunensis (Villers)

55. plumbeus (Olivier)

Distoleon Banks

56. mr~nulatus (Klug)

57. c.clftu.\ (Fabricius)

58. contub~rtzali~ (McLachlan)

59. retrczgrclrnmicus (Fabricius)

Glenurus Hagen

60. gratus (Say)

Gyrnnocnerniu Schneider

6 1 . vclriegata

Mrgistoj)us Rambur

62. flflaviconlis (Rossi)

Navclsolcon Banks

63. boliviuna Banks

Nel~roleon Navis

64. arcnarius (Navas)

65. distic-hu~ (Navis)

66. ocrecltus (Navas)

67. rr~ic.rc>sterlus (McLachlan)

68. n~rrlnusierzsi.~ (Borkhausen)

Purciglctzurus van der Wccle

69. jczpotric~r.\ (McLachlan)

70. c>kinalz9ensis Okamoto

BIBLIOGRAPHY

Algeria, France

Dodecanese Islands

Dodecanese Islands

Madeira

Ryukus Island

Europe

Florida (USA)

Italy

France

Peru

France

France

France

Japan

Ryukus

Redtenbachcr 1883; Steffan 1964, 1975; Willmann 1977

Willmann 1977

Redtenbacher 1884b;

Brauer 1854, 1855; Redtenbacher 1884; Steffan 1975

Miller & Stange 1983

Insom et ul. 1985

Steffan 1964, 1969, 1975

Miller & Stange 1985

Steffan 1971

Steffan 1971

Steffan 1971

Steffan 197 1: Gepp 1974

Stefan 1971; Auber 1956a

Redtenbacher 1884b

Tanaka 1979

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Classi$cation of M y rmeleontidae

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L.A. Stunge & R.B. Miller

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TILLYARD, R.J. 1926. The Insects of.4u.stralia and New ZPuland. Angus & Robertson Ltd., Sydney. 560 pp. VAN DER WEELE, H.W. 1909. Mecoptera and Planipennia of Insulinde. With biological notes from Edw.

Jacobson. Notes from the Leyden Museutn 3 l : 1-98. WHEELER, W.M. 1930. Demons of the Dust. W.W. Norton & Co., New York. 378 pp. WILLMANN, R. 1977. Die Myrnieleontidae (Insecta, Neuroptera) der DodekanesIAgais. Z~ologisches

Jahrbuch (Systematik) (Jena) 104: 98-136. WITHYCOMBE, C.L. 1925. Some Aspects of the Biology and Morphology of the Neuroptera. With special

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Classification of Myrrneleontidae

Addresses of authors:

Dr Lionel A. Stange Taxonomic Entomologist Florida State Collection of Arthropods Division of Plant Industry P.O.Box 1269 Gainesville, Florida, 32602 U.S.A.

Mr Robcrt B. Miller P.O.Box 1092 Project City California, 96079 U.S.A.

Bibliography of the Neuropterida Bibliography of the Neuropterida Reference number (r#): 7265 Reference Citation: Stange, L. A.; Miller, R. B. 1990 [1990.??.??]. Classification of the Myrmeleontidae based on larvae (Insecta: Neuroptera). Pp. 151-169 in Mansell, M. W.; Aspöck, H. (eds.). Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology (3-4 February 1988, Berg en Dal, Kruger National Park, South Africa). South African Department of Agricultural Development, Pretoria. 298 pp. Copyrights: Any/all applicable copyrights reside with, and are reserved by, the publisher(s), the author(s) and/or other entities as allowed by law. No copyrights belong to the Bibliography of the Neuropterida. Notes: File: File produced for the Bibliography of the Neuropterida (BotN) component of the Lacewing Digital Library (LDL) Project, 2010.