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Vol. 8, No.1 - 2, March - June, 1994 67

Reclassification of tile New Vior Id antlioll gellera formerly included in the tribe (: Myrmeleontidae)

Lionel A. Stange Flolida State Collection of ArUllopods Department of Agriculture & Consumer Services P () Box 147100 Gainesville, FL. 32614-7100 U.S.A.

Abstract A cladistic analysis of the New World tribe Brachynemurini has resulted in se\'eral new taxonomic designations. The tribe is divided into 3 tribes, 2 of which are newly described. The Brachynemurini s.s. now contains 12 genera of which Argentoleon, Atricholeon, Mexoleon and Venezueleon are newly described. The Gnopholeontini (NEVI TRIBE) includes 4 North American genera whereas the Lemolemini (NE'N TRIBE) contains 6 South American genera of which &ualeon and Galapagoleon are newly described. Descriptions of genera in the 3 tribes, based on adults and kno'JlIl larvae, are given. Keys to the genera in each tribe are provided, as well as a key to the tribes of Myrmeleontidae.

KEY WORDS: Neuroptera, Myrmeleontidae, New World, keys, cladistics, phylogeny, classification.

Introduction lowing autapomorphies: 1) post\'entrallobe ofmale Since the revision of the North Ameriean ectoproct; 2) a narrow gonapophyseal plate; and 3) BrachynemurinibyStange (1970), additionaladult a short distal tooth ofthe larval mandible, which is and larvalcharacters (Stange & Miller, 1990) have ~een fOIl;;Whi~ h::I";; Jecljo the present reclassi- enee Of phylogenetie relationships, t"'lO analyses lcatlOn 0 ew or raChYnemurme genera mto three tribes. Twenty three genera are hereby ree were made: one of all genera and the other using ognized. Representative larvaeofnineteenofthese only genera in which the larvae are known. genera are known. A cladistic analysis of these genera has led to significant changes in their clas­ HIgher level relationships Within the sification. Two gro'lpS Of genera fOrmerly placed m Myrrneleontoidea the Braehynemurini are given tribal status Relationships among higher taxa within the (Gnopholeontini and Lemolemini); the remaining Myrmeleontoideahave notpreviously been investi­ genera are maintained as the Braehynemurini. gated eladistieally. MacLeod (197Q) has indicated The Gnopholeontini are restricted to the Sonoran several autapomorphies of the superfamily based Region; they are defined by the autapomorphy of on the larval head The apomorpbic larval bead the close approximation of the larval mandibles. characters unite the , , The Lemolemmlare most diverse m Chile; they are Nemoptendae, AScalaphldae and Myrmeleontidae defined by the autapomorphies ofthe development into a group. Mansell (1992) has provided the most ofspecialized setaeon the median areaofthelarval recent discussion of the relationships within the abdominal tergites and the lack ufdigging setae 011 indicating apomorphies for the the female ectoproct. Both the Gnopholeontini and various families and providing a . Lemolemini have highly modified posterior Oswald (1993) has eompleted a eIadistiC analYsis of gonopophyses ofthe female terminalia. The more the Psychopsidae, a hypothesized as the restricted Brachynemurini are defined by the fol- sister-group of the rest of the Myrmeleontoidea 68 Insecta Mundi

(Withycombe, 1925, IIeIllY, 1978). The following Lachlathetes) but these exceptions are considered analysis of the tribes of the Myrmeleontidae uses secondary modifications. these four families as outgroups whelp determine outgroups for the generic analysis. 3. Labial palp, sensory area - (0) palpimacula present The Psychopsidae (26 extant in 5 gen­ but without pit; (1) o...al shaped pit; (2) elongate, slit-shaped pit era, found in Afriea, Australia and the Orient) and Comments: The apomorphic condition (2) is found in the Nymphidae (about25 speciesin 8 genera, restricted I\canthaclisini, , Palparidiini and some to Australia and New Guinea) are small relict genera of the Palparml. Whether the latter trIbe families. The and are should be considered plesiomorphic or apomorphic much larger and more diverse. Since many of the In regards to thIS character IS not completely clear characters used in the tribal analysis of the but results scoring this character both ways do not MyrmeleontIdae vary consIderably among the two change the results except for a minor node differ­ subfamiliesofNemopteridae, bothsubfamilieswere ence. The outgroups have small palpimacula and the evolution of enlarged palpimacula seems to be entered indehendent~for a clearer resolution of found only in the Myrmeleontidae. 'fhe the relations ips. Th Ascalaphidae appears to be Nemopteridae may have lostthe pit-likepalpimacula even mo~~ dive;:ce with at ~eas~ 3 s;~ilie~ ait when they adapted to feeding.

(THORAX) acters are entered for the family as a whole. 4. Pronotum· (0) wider than long; (1) as long as wide . The tribal classification of the world or longer Myrmeleontidae is based on Markl.(l956) with Comments: Mansell (1992b) considers without much modifications indicated by Stange & Miller (1985; justification the broadpronotum asanautapomorphy 1990) Since larvae are known for all the major ofthe . It is treated here as plesiomorphic mainly because it is found in the Psychopsidae groups, it was decided to utilize both larval and which is considered to be the sister group ofthe rest adult characters in the analysis. The only tribes of of the Myrmeleontoidea The polarity is subjective Myrmeleontidae unknown in the larval stage are fundamentally but scored either way would not thePseudimarini(1 rarespecies) andMaulini(about change the . 5 species in 2 genera) Although the larvae in general remforce the trIbalclassifIcatIOn, oneprob­ 5. Pronotal artIculatIOn (taken from Adams, 1958); (0) lem in using laNae was recognized. The laNae of pronotum articulates with mesot.horax the Dimarini are highly diverse (Stange, 1989) so (psychopsidae; NymphIdae); (1) pronotum artICU­ lates with mesothoracic s iracle' 2 ronotum ar- rather than the tribe as a whole Preliminary rac1e analysis led to the beliefthat three distmct groups we~e present inthe Brachynemuri~.To attemptto 6. Pretarsal arolium - (0) present, (1) absent Comments: The presence ofan arolium is considered the plesiomorphic condition in the Myrmeleontoidea. The following informative characters were used. There is anevident aroliuminPsychopsidaewhereas The data matrix is given in Appendix I in the Ny mphidae the arolium is deeply bifid, for m- ing two pulvilliform lobes. In the Ascalaphidae and M)rmeleontidae the auxiliae are fused forming a List of myrmeleontid tribal apomorphies bristly median lobe co-adapted to a strong (HEAD) unguitraeter plate which strengthens the flexing 1. Mouthparts - (0) short mouthparts; (1) head pro­ mechanism ofthe claws. The arolium is completely longed with long mouthparts absent in these families Comments: The elongated head and mouthparts is associated with pollen feeding in the Nemopteridae. 7. Femoral sense haIr - (0) absent; (1) present on proleg and mesoJeg (Fig 9) 2. Antennal shape· (0) filiform, (1) clavate (Fig. 36), Comments: The femoral sense haIr IS an autapomorphy (2) Clubbed of the subfamily Myrmeleontjnae In a few genera Comments. The shape of the antennae is fairly consis­ (I.e., Atncholeon, Venezueleon) the haIr has been tent but there are a few exceptions. Filiform-like secondarily lost as evidenced by the strong reduc- antennae are found both in the Ascalaphidae tion of all leg setae in these genera. In the (Tmesibasis) and Myrmeleontidae (Anteonoleon, there is also a femoral sense hair (sometimes two) on the metaleg. Vol. 8, No.1 - 2, March - June, 1994 69

8. Hindwing - (0) similar ro forewing, (1) greatly elon Mymeleontidae is a derived state, then the distal gated crossveins in Pseudimares iris Kimmins may repre- sent a derived condition rather than a vestigial (WING VENATION) condition. 9. Forewing, origin vein CuP (~at or before crossvein; (1) distad ofM-Cu or CuP obsolete; (2) M­ 14. Pilula axillaris (#12) - (0) absent; (1) present Cu obsolete Comments: The pilula axillaris or Eltringham's organ is Comments: The apomorphic condition is found in found only in the Myrmeleontidae and is an GnophoIeontmI, Lemolemini and Brachynemurmi. autapomorphy of the family. The absence of the Elsewhere, the apomorphic condition has been ob- structure in several groups (especially ) served only m the SOuth AfrIcan genus Nanrwleon IS conSIdered a reversal. and and in some .

10. Forewing, condition ofvein lA - (0) Free course from (ABDOMEN) base to hind margin (see Markl 1954. Fig. 62), (1) 15. First abdominal spiracle (data taken from Adams, uniood with CuP near base (Figs. 10-11) 1958) - (0) eonneeood to mesepimeron; (l) incorpo- Comments: The genus Palparidius is variable in this rated with the metepimeron character. Comments. The migration of the first abdominal spi- racle onto the metepimeron appears associated with 11. IIindwing, OrIgm of radial seeror - (0) near wing the diminution of the first tergite. Probably this base, well before origin ofmedial fork; (1) originates relocation of the spiracle improves the air supply to distal ro medial fork (Fig. 15), (2) greatly reduced the flight muscles. Comments: This is a progressive character where the origin of the radial seeWr has apparently migrated 16. Female terminalia, posterior gonapophysis (#16) distally according to Tillyard (1915). Usually there (0) absent; (1) plate-like; (2) elongate, digitiform are only 1 or 2 presectoral cross"eins in the (Figs. 17 22); (3) inflated or greatly enlarged (Figs. plesiomorphic condition but some palparine genera 23-27, 30-32) have more due to the extensive crossveins fOund in Comments' There is confUsion about terminology ofthe wing areas. gonapophyses. Tjeder (1977) decided to use other terms. However, I conSIder that the proper names 12 Both wings, hypostigmatic cell - (0) absent; (1) are still to be resolved and use my terminology Short, less than 3x longer than hIgh; (2) elongate, (Stange, 19/0) here. The posterIOr gonapophysIS more than fix longer than high (Fig 15) appears to have evolved from an inconspicuous Comments: The shape ofthe hypostIgmatIc cell appears swollen plate m the Ascalaphldae and Palparmae to to be a very constant character in the Myrme- huge structures in Nellwtlls (Fig 27) Miller (1991) leontOldea. However, there are a few genera of has suggested that the Inflated posterIor Myrmeleontidae Uvlaracandula; Nuglerus) where gonapophyses function as handlers whereas the thehypostigmaticcell isshortas in the Ascalaphidae. digitiform gonapophyses are used for soil testing. This is probably parallel evolution since these gen- era in other characters appear to be relatively 17. Female terminalia, gonapophyseal plate - (0) ab­ apomorphic in their tribes. The short and long sent, (1) narrower than width of posterior hypostigmatic cell conditions are entered as non­ gonapophysis; (2) wider than width of posterior additive since it is not clear whether the asealaphid gonapophysis type or myrmeleonidtype represents the apomorphic Comments: The gonapophyseal plate is an asetose condition. selerite which forms part of the floor of the genital chamber and apparently articulates with the poste- 13. Both wings, subcostal area (0) with at least basal rior gonapophyses. Usually the larger the posterior crossvein; (1) without crossveins (Fig. 15) gonapophysis, the larger the gonapophyseal plate Comments: Oswald (1993) tentatively asserts that the although with some exceptions (f.canthaclisini; presence of the basal subcostal crossvein is Nemoleontini). This might be a synapomorphy of pleslOmorphic m Psychopsldae and the prolIfera­ the except it appears to be absent tion of additional crossveins as apomorphic This in the (a sQcondary loss?). Some may be true m the Nymphldae as well smce New parallel evolutIon m the reductIOn of thIS plate is (1981) clearly shows some genera of Nymphidae seen in the Brachynemurini It can be moderately WIth only I basal crossvem and others WIth many large (Dejuna) to very small ( and some crossveins The basal crossvein is absent in the Dendroleontini such as Glenoleon which is variable, Ascalaphldae and MyrmeleontIdae whICh IS taken but Glerwleon IS probably paraphyletlC. as a synapomophy of these two families. If the development of distal crossveins in the 70 Insecta Mundi

18. Lateral gonapophysis, ehaetetaxy (Figs. 17 32) (0) ) additional teeth (4 to 6) are developed. digging setae weaklyto moderatelydeveloped, never The shortening of the mandible has led to a reduc­ longer than gonapophyseal width (Figs. 17 27); (1) tien ofteeth in only a few eases. Two teeth aro found digging setae large, down curved, longer than in (Nemoleontini), gonapophyseal width (Figs. 28,29). (.A.eanthaelisini) and Millerloon (Dimarini). The lat ter genus is the most enigmatic since the short teeth 19 Male ectoproct (#20 ) - (0) without elongate stilI show remnants of the seta (Inly 1 tooth is postventrallobe (Figs 50,51, 53); (1) with elongate present in Paron thaclisis (Acanthaclisini) IIsu- postventral lobe (F'lgs. 47, 49, 55). ally, the teeth are parallel WIth an mcreasmg length Comments' The Ascalaphidae is about as variable as the distally although several gronps have the middle Myrmeleonhdae m thIS character so that the polar­ tooth the longest. These are all probably derIved ity can not be establisbed for tbe family as a whole conditions and are not tbe foclls of the present Some variatIOn occurs m the Brachynemunm; analYSIs. In the present analysIs, the prImary Myrmecaelurini and Nemoleontini, but the ground apomorphy is the modification of the distal tooth pIau is considered apomorphic for the Brachy- which is much shorter than the middle tooth but, nemurini and plesiomorphic for the other two tribes. more important, situated at a different angle. This apomorphy is found in Brachynemurini, (LARVAL CIIARACTERS) Myrmecaelurini, and Nesoleontini. Larval characters of the Psychopsidae and Nymphidae are diseussed by MaeLeod, 1970, Ne w 1982, 1983, 22. Labial palpus, length - (0) elongate, longer than 1989. basal width of mandible; (1) shorter than basal width of mandible 20. Mandibular distance - (0) widely separated; (1) elosely approximated 23. Oeular tubereles (0) absent; (1) present Comments: The approximation ofthe mandibular bases is found only in the Gnopholeontini amoung the 24. Mesothoracie spiraele (0) sessile; (1) peduneulate Myrmeleontidae. The most extreme case is Unopholeon barner! Wig 69) basewhere the closer 25 Hindtarsus and tibia - (0) articulated; (1) fused approximation of the mandible is found only in the Ascaiaphidae. ThIS IS eVIdently a derIved condItIOn 26. Hmd pretarsal claws - (0) not larger than fore and is probably associated with sessile larvae in pretarsal claws; (1) much larger than fore pretarsal whIch the trap structure IS more advantageous. claws Comments' The enlarged hind pretarsal claws are 21. Mandible, dentition - (0) without teeth; (1) WIth 2- eVIdently an adaptatIon for burrowmg backwards 3 parallel teeth (Figs. 64·70); (2) distal tooth at into the sand The only known exception in the different angle than middle tooth (Fig. 63). Myrmeleonudae IS Gnopholeon barberi. The reduc­ Comments. The development of mandibular teeth is tion in the size of the hind pretarsal claws is prob- clearlysynapomorphicfor Nymphidae, Ascalaphidae ably a reversal in this species, which does not and Myrmeleontidae. All other families of burrow in the sand and is ecologically similar to the Neuroptera including the Myrmeleontoid families Ascalaphidae. It is also possible that the Nemopteridae and Psychopsidae lack true man- Ascalaphidae are derived from tile Myr meleontidae, dibular teeth. Mandibular teeth are apparently andhave secondarily lostthe enlargedhind pretarsal derived from enlarged setal bases (Eaba, 1954) and, claws as an adaptation to li\'ing on plants. In some based on known evidence, the development of at of the Nemopterinae which burrow into the sand least 3 teeth appears to he the next step without head first, the anterior pretarsal daws are en intermediate stages of having 1 or 2 teeth. All larged. known ascalaphid larvae and the great majority of - larvae have 3 teeth. Some extant nymphid 27. Abdominal fossoria - (0) absent; (1) present larvae have only 1 large tooth, but this is probably Comments: The fossoria are enlarged digging setae on due to reduction since a nymphid (Pronymphes) sternite IX and sometimes found only on the third has 3 parallel teeth and at least 1 undescribed instar . extant species has several teeth Related to larval strategIes, the mandIble has become longer 28. Abdommal stermte VIII - (0) WIthout submedIal in some groups shorter in ot.hers Probahly there teeth; (1) with submedial teeth (Fig 16) has been repeated lengthenmg and shortemng of Comments: The development of submedial teeth may be mandibles as the different types appear in different a synapomorphy of the Ascalaphidae and evolutionary lines. The lengthemng ofthe mandIble Myrmeleontidae often is accompanied by having the teeth situated more distally, or in a few palparine genera (Gola/rus; Vol. 8, No.1· 2, March· June, 1994 71

The characters and taxa were analyzed using the extlerne diffelencesin thelalvaeofthe 2 geneta the Hennig86 cladistics program (version 1.5) of Goss ofthefossoria & submedialteeth inMillerleon), Farris (1989). Using the ie option, 6 equally parsi­ the analysis supports the two genera forming a monious were found (length 71; con. monophyletic group sIstency mdex (CI) 0.53; retentIOn mdex (rI) The of the NemoleontmI appears to 0.70). The successive approximations character be weakly supported only by the reversal ofchar weighting feature (Farris 1969; Carpenter, 1988) acter #14 (secondary loss of the pilula axillaris). resulted in 2 cladograms (ci 0.83, ri 0.92). This apomOlphy is also found in the Nesoleontilli Character weights are given in Appendix 3 The and in the Brachynemurini (Scotoleori) Since the cladograms are shown in FIgs. 1·2. trIbe forms a trIchotomy WIth the groups treated The suite of characters used in this analysis here in one cladogram and as a sister group to the support the recognition offive families. As can be Brachynemurini+ (Lemolemini + (Gnopholeontini appreciatedfrom theeladograms, the Ascalaphidae I Dendroleontini» in the other cladoglam, it was is so close that itcouldbe considered as a subfamily decided to use the Nemoleontini as one of the ofMylIueleolltidae. All the autapolllOlphies for the outgroups. Ascalaphidae appear somewhere in the MyrmeleontIdae. However, two otherapomorphles Relationships Within the Gnopholeontlnl, exist which may strengthen the case for family Lemolemini, and Brachynemurini recognition ofthe Ascalaphidae. The antennae are In earlier papers on the classification of the significantly longer than in the Myrmeleontidae Brachynemurini (Stange & Miller 1985; ~992), the ex e tfor h iz rr nu A r Th ea 0 e arva may e apomorp IC e Myrmecaelurini. This was based on the remark· Ascalaphidae. able convergence in the dentitiOn of the larval The 2 weightedcladograms arethe sameexcept mandible. However, the present analysis suggests for the placement of the Nemoleontini. According a closer relationship of the My rmecaelurini to the tocurrentclassification, the , Palparidiini, Acanthaclisini basedon female terminalia charac· ter #18). The relationship of the Acanthaclisini with the Myrmecaelurini + Nesoleontini is sup- portedby anotherapomorphy notusedin the analy­ sis, the presence of abdominal hair pencils. Hair analysis indicate that the subfamily Palparinae in pencils arefound in allgroups ofthe Acanthaclisini. the traditional sense is paraphyletic. The Elsewhere, they ale fuund unly in SOllie genera of the Myrmecae)urini The present tribal analysis supports the split· the Palparinae if one wishes to recognize ting of the Brachynemurini into 3 tribes paraphyletic taxa. Another treatment supported (Gnopholeontini; Lemolemini; andBrachy!!emurini} by this cladistic analysis is to recognize three sub· andindicatesa close relationship ofGnopholeontini families; thePalparinae (palparidiiniandPalparini), with the Dendroleontini. The mono~hYIY of the the StiJbopteryginae, and the Dimarinae with the Gnopholeontini is supportedby the au apomorphy of the close approximation Of the larval mandibles Imarmae. owever, more c aracter ana ySIS 0 (#20). The monophyly of the Lemolemini is sup­ theStilbopteryginaeandassociatedgroupsis needed ported by one larval autapomorphy not entered before such a change is recommended. The into the anal~sis: the presence of::cialized setae Stilbopteryginae have always been a eontroversial along the lIle ian line of the abdo en. The mono· group, sometimes classified with the Ascalaphidae phyly of the Bracbynemurini is supported by the based on the clubbed and short elongate postventral of the ectoproct (# 19) and hypostigmatic cells which are key characters ofthe larval dentition (#21). Finally, the monophyly of Ascalaphidae. Ithas also been treatedasa separate the Dendroleontini is sUI>I>orted by one larval family and, more recently (New 1982), as a sabfam· autapomolphic character, namely tlu;setae which ily of the Myrmeleontidae. New's concept is sup· hold debris on the mesonotum (not entered in data ported by this analysis matrix). Although the genera Dimares and Millerleon were entered independently in the analysis due to 72 Insecta Mundi

Using theDemholeontini and Nemoloontini as 8. Pletalsal claws - (0) small to modelately large, less outgroups, thefollowing phylogenetic analysis was than one-half length of distal tarsomere; (1) en­ made on genera of the BrachynemUrIOl, lalged, mOle than one-halflength ofdistal talsomete Gnopholeontini and Lemolemini. The following mformatlve characters were used, WIth the WING CH.ARACTERS 9. Forewing, origin of vein CuP (#9) - (0) at or before plesiomorphic condition and comments where reI crossvein m cu; (1) distad of m cu or CuP obsolete evant. The data matrix is given in Apendix 2. Comments' The distal origin of vein CuP is a Characters #4, #22, #25, and #31 were treated as synapomorphy of the tribes Brachynemurml, non-additive Numbersin bracketsindicateeqlliva- Gnopholeontini and Lemolemini Sometimes vein lence WIth hst of trIbal apomorphIes. CuP IS obsolete m these tribes. The character IS very stahle, the most hasal position of Cnp was observed In Janueha. In a few groups of the Dendroleonbm List of gelleric apOIIIOl pllies (adult) the ;:gin ~fvein ~P=ay h~dis~a~ of:ss~ei~ m- cu. sew ere meyrme eon i ae,lserlved HE~ CHA&A.CTERS state is found in ZVannoloon Esben-Petersen and in the tribe Maulini.

10. Forewing, vein 2A - (0) runs at even curve toward 3A, (1) luns along 1A fOI shOlt distance, then back at a sharp angle towards 3A (Nemoleontini)

. and Scotoleon. 11. Hindwing, vein CuA - (0) bends to hind margin at or 3. Ocular rim setae - (0) absent; (I) present before medial fork, (1) runs along posterior branch Comments: The development of ocular rim setae ap­ of medial fork for some distance pears correlated with the developmentofprofemoral Comments: The derived state is found in most genera of clavate setae. Usually there are two or more setae Brachynemurini albeit with some variation in ex­ on the dorsal surface ofthe rim which vary m length pression. The most extreme expression of tho de from genus to genus. The derived state is a rived state is found in the genus Ameromyia in the synapomorphy ofAbatoleon, Chaetoleon, Gnopholeon New World Brachynemurini. (2 of 3 species) and Menkeleon. Also, ocular rim setae ale found in some genelaofthe Acanthaclisini, 12. Hmdwmg, pIlula axIlIarls (#14) . (0) present; (1) Dendroleontini and Myrmecaelurini. absent Comments: The pllula aXlllarls IS sometImes call THORACIC CHARACTERS "Eltringham's Organ" This structure, odoriferous (0) 4. Profemoral elavate setao (Fig. 9) absent, (1) m function, IS found only In the male With a few present, ;ran~ed in a FeW; (2) elustered exceptions (e.g. B. longiCaudu8) Although a unique Comments:heerivod stato is a synapomorphy of character for the family, it appears to have been lost Abatoleon and Cha€toleon. In these genora thore in some groups of the MyrmeleontInae are at least four such sotae In a row and in most (Nemoleontini; Scotoleon). Abatoleon there' are secondary rows. In 2 of tho 3 speCies of GnophOleon theso setae are clustered; ABDOMINAL CHARACTERS this state is considerod of independent evolution 13. Female telminalia, ectoproct - (0) with digging setae; (1) without digging setao O. Fomoral sense hair (Fig 7) (#7) • (10 present; (I) absent 14. Female terminalia, lateral gonapophyses - (0) sopa­ Comments: The derived state is a synapomorphy of rated (Figs. 17 20, 22 30); (1) partially fused (Fig. Atricholeon and Venezueloon. Tho derived state is 21); (2) completely fused (Fig. 31) evidently a reversal to the primitive state found in Comments: The derived states are found in }Jara Palparinae. candula, Jaffuelia, Venezueleon and Ameromyia.

6. Profemoral sense hair - (0) much shorter than 15. Female terminalia, latoral gonapophyses (0) elon plofemUI, (1) as long as or longer than femur gate (Figs. 17-19, 23-29); (1) transverse (Figs. 20·21, 3032) 7. Tibial SpUIS - (0) present, (1) absent Commonts: The dorived state is found in the Comments: The derivod state is a synapomorphy of Gnopholeontini. Maraeandula, Menkeloon, and Tyttholeon. Tibial spurs are sporadically absent in Brachynemurus (2 spocios) and Abatoleon (1 specios). Vol. 8, No.1 - 2, March - June, 1994 73

16. Female terminalia, posterior gonapophyses (#16} Comments. The derived state is found in most genera of (0) digitiform (Figs. 17-22); (1) inflated or greatly the Brachynemurini. enlarged (Figs. 20 22) Comments: The derived state is found in the 24. Male genitalia, paramere . (0) plate·like; (1) re- Gnopholeontini and the Lemolemini. Extreme ex duood to separate rods (Fig. 48) pressions are found in Maracandula and Neulatus. Comments: the apomorphic condition is found in Miller (1991) stated that the djgitiform type is Ameromyia and venezlleteOn mostly used for soil depth t.esting whereas the mflated gonapophysls IS assoCiated With egg han­ 25. Male gemtaha, sculpture of medlUncus - (0) smooth dling. (noD.spinose, not. cross striat.e); (1) micro spinose (Austroleon); (2) cross-stnate (Figs. 54, 59) 17. Female terminalia. posterior gonapophyses . (0) (Gnopholeontini) without scraping setae (Figs. 11-24), (1) with scrap' Comments: The derived characters are non-additive. ing setae (Fig. 25) Comments. The derived state is found in Chaetoleon and LARVAL CHARACTERS Jaffuelia and is associated with sCiaping the sub- 26. Mandibles, (#20) - (0) widely separated (Figs. 60- strate prior to egg deposition. 67); (1) closely approximated (Figs. 68-71) Comments. The derived state is found in Gnopholeon, 18. Female terminalia, gonapophyseal plate - (0) nar· Menkeleon and Tyttholeon. Similar approximation rower than posterior gonapophysis (Figs. 17 21); (1) oeeurs in many asealaphid larvae. Wider than posterior gonapophysis (Figs. 22-27, 30­ 32) 27. Mandibular teeth (#21) (0) 2 parallel teeth, usually Comments: The derived state is found in Gnopholeontini increasing in length distally (Figs. 64-71); (1) distal and Ilendroleontini Although there is considerable tooth shorter than middle tooth, set at different variation in the size of the plate, I am dividing the angle (Figs. 60-63) st.at.e into 2 majOr groups usmg the wjdth of the Comments' The derived state is found in the plate. Some interpretation is needed since although Brachynemurini. This type of mandible is also the plate is somewhat narrower t.han the found m the (lid World Myrmecaelurini and gonapophysis in Maracandula, it is still very large Nesoleontini. See comments on tribal relationships and the discrepancy IS due to the extremely mflated (#21) posterior gonapophysis in that genus The Lemolemini is split into two groups. lv'eulatus and 28. Hmd pretarsal claw - (0) Without basal tooth; (1) Jaffuelia have large gonapophyseal plates whereas with basal tooth ElicUia, Lemolemu8 and Sical have narrower plates. Comments. The derived state is a synapomorphy of Neulatus and Jaffuelia. 19. Female ter minalia, pregenital plate . (0) with tooth (Figs. 24-26, 30.32), (1) without tooth (Figs. 17-18, 29. Mesothorax, chaetotaxy . (0) no specialized median. 20) setae; (1) with specialized setae medially on mesonotum, 20. Male ectoproct (# 19) - (0) without elongate sometimes borne on common stalk (Bankisus; many postventral lobe (Fig. 52); (1) with posv.'entrallobe Australian genera of Dendroleontini) (Fig. 41-43, 47, 49) Comments: The development of a medial group of elongate bristles on the mesonotum appeals to be 21. Male ectoproct· (0) not produced ventrally (Figs. 47, the best apomophic characterization of the 49); (1) produced ventrally (Figs 51, 53) Dendroleontini. The funotion of the bristles is to Comments: The apomorphic condition is best expressed hold a debris ball which acts as a lure to prey and in Mexoleon represents a unique predatorstrategy iathe . In Bankisus and many Australian groups 22. Male genitaha, genital sac· (OJ without cluhhed (Periclystrina) this structure is very long with the setae or secondary sclerites; (1) with clubbed setae bristles fused as a common stalk. (Fig. 48); (2) with lateral lobe·like sclerite; (3) With median setose sclerite near mediuncus (Fig. 44) 30 Abdominal scali - (0) absent; (1) present Comments. The derived state (#1) is a synapomorphy of Comments: The development of abdommal scoh prob· Ameromyia and Venezueleon whereas the extra ably is related to living on rock surfaces or plants sclerites are found in Aigentoleon and Atrieholeon. (Gnopholeon; Jaffuelia; Neulatus), parallehng the These derivatives are non-additive. ascalapbid conditjon

23. Male genitalia, parameres - (0) not·hinged (Figs. 57, 59); (1) with hinge (Figs. 46, 54, 56) 74 Insecta Mundi

31. Abdominal tergites, chaetotaxy - (0) without spe­ female ectoplOct (# 13) and the latvalautapomOIphy cialized setae medially; (1) with dolichasters; (2) of the development of specialized setae on the with blaek, tufted setae along midline dorsal mesal margin of abdomen (#31-2). Elicura Comments: The derived stateisfound in the Lemolemini. and LemolBmus have autapomorphiQs which sup There are two groups. Roseate, squat doliehasters are found in Neulatus and Jaffuelia whereas other port their monophyly: the sexual dimorphism of Iemolemine genera have tufted black setae the tibial spurs in Lemolemus and the forefemoral setal comb and expanded forewing costal area in Based on the character matrix given in Appen­ dix 2, 3 separate analyses were made. The first analysis was madeofthe generainwhich thelarvae are known The other 2 analyses included the 5 generain which the larvae are unknown in order to and Ensorra are the sister group to the rest of the dQtQrminQ phylOgQnQtic rQlahonshij)s Of thQse 5 brachynemutine genera although this telationship genera. is found in only 1ofthe 2 weighted cladograms and The first analysis was made of 19 genera with is based on character# 19, a very homoplastic char­ laIval infollnation. Characters #4. 22. 25, and 30 ac~r.~mBromyia+ \4mezuBlBon are defined as a were run as non-additive. The mhennig*/bb* op­ cla e oy 3 autapomorphies. These two genera also tion ofHennig86 resultedin 112 cladograms (length are restrictedto SouthAmerica. Theother5 genera =62; ci =O. 59; ri=O. 71). Thesuccessive approxima- are all North American. Although both Chaetoleon tions characterweightingfeature reducedthe num­ and lv/exoleon have the male ectopIOct produced berofeladograms to 2 (ci 0.87; ri 0.93), character ::~t~:l~~:I~::~1RZ:~:t~~~:~~~~.!Jc~;t~nf~: weights are given in Appendix 3. The 2 dadograms (Figs 3, 4) are nearly identl- genera treated in the analysis, only the original cal, differing only in the treatment ofthe loss ofthe genus Brachynemurus has no apomorphies to de­ mQdial tooth on thQ prQgQnital plate (#19). This is fend it and will need furthet study. a homoplastic characterappearingin severalplaces To determine the relationships of the 5 genera in the cladogram. One cladogram (Fig. 3) has a of Brachynemurini for which larval data are mISS­ tIichotomy involving Austroleon, Abatoleon + ing, two additio~l analyses were done. The first Argentoleon + Ensorra, and the rest of bracbynemurine genera The other cladOgram (Fig. 4) eliminates the trichotomy. mhennig*/bb* option of Hennig86 program was Examination ofthe 2 cladograms clearly shows run and resulted in 3,248 eladograms (length 67; that larval characters are most important for the discrimination of the three of the Brachynemurini s I ,now recognized as threetribes The Gnopholeontini include Gnopholeon, appendix 5. A Nelson consensus tree of these MBUkBlBOU, and 1'Yttholeou, and are defined by weighted cladogIams is given in Fig. 5. characters#15, 25-2 & 26. The Lemolemini are the Maracandula comes out in the Gnopholeontini, sistet glOup oftheGnopholeontini+Dendroleontini, ECllaleon and Galapagoleon in the LemolQmlnI, defined hyeharacters#13 & 31-2 Finally, the t.bird whereasDejuuaand Clathroueuriaare shown to be large group of genera now comprising the related to genera comprising the tribe Brachynemurini, arQ dQIinQd by charactQr #27. Brachynemmini 15.15. The genera of Gnopholeontini are well defined The second analysis was to run the 25 genera in both cladograms. Tyttholeolt has no listed char- usingonly adult characters This analysis was less acters in this analysis butis abundantly character- informative regarding the relationships of the five ized by autapomorphies such as the shape of the genera. Usmg the mhenning*/bb* option of illal e ectoprocts (Fig 50), enlargedpregenitalplatQ Hennig86, 3,503 eladograms were produced(length (Fig. 30), and lanceolate setae on the larval - 59; ci - O. 52; ri- 0.65). The successive approxima­ meosternum. tions characterweightingfeature reduced the num- ber of cladograms to 353 (ci = 0.80; ri = 0.90); The Lemolemini ate defined by the adult charactQr WQights arQ gWQn in appendix 6. A autapomorphy of the Joss of digging setae of the Nelson consensus tree (Fig. 6) was produced. The consensus tree exhibits considerably less resolu- Vol. 8, No.1 - 2, March - June, 1994 75 tion than the plevious cladoglams. The BRACIIYNEMURINI Gnopholeontini are resolved, but the genera of Abatoleon- (1) Mediuncus and the parameresofthe Lemolemini form a polytomy with the male genitalia highly modifIed (FIg. 58) Dendxoleontini and Gnopholeontini The genera Marcwandula, Ecualeon, and Galapagoleon fall AmeromYla - Enlarged pretarsal claws. within the polytomy and their relationships are thus uncertain. The Brachynemurini s.s. are only Argentoleon - Secondary. lobe-like sclerites later- paItly Iesolved with Abatoleon, Algentoleon, ally on genital sac. Austroleon and Ensorra forming part of a basal polytomy for the entIre tree. DeJuna and Atncholeon-SecondarysclerotIzedandsetoseplates Clathroneuria fall out in this region. anterior to gonarcus (Fig. 44).

Generic autapomorphies Austrokon - Male mediuncus miclO-spinose.

LEMOLEMINI Brachynemurus - The apically narrowed male Ecualeon (1) Female tergite VIII with egg han ectoproct is autapomorphic for 8 of the 21 species dling setae (Fig. 33); (2) presenceoflarge, scale-like (FIg. 42). setae on male abdominal segments III-V; (3) male ectoproct ventrally emarginate (Fig. 35). Chaetoleon 0 Posterior gonapophysis with scraping setae. Galapagolron - Female posterior gonapophysis strongly curved (Fig. 23). Ctathroneuria -(I) Postventrallobeofmaleectoproct with median lobe (Fig. 43). (2) mediuncus ofmale Elicura - Profemur with elongate white bristles on genitalia narrow and elongate (Fig. 46). exterior face Dejuna - Male genitalia with gonarcus elongate, Jattuelw - Larva WIth most of body covered with rnediuncus with spiniform process, and parameres scale like setae. abbreviated (Fig. 45). iVeulatus - Apex of st.ernit.e VIII wit.h median pm- Ensorra -Male stemite IX extIemely elongate

hair absent, that of profemuI small, southem 15), metafemur without sense hair (except Africa Maulini Sericoleon and Epacanthaclisis); tibial spurs Labial palps shorter than head width, area not andpretarsalela..s various, usuall) notstrongl) extending to apex; mesofemoral sense hair arched or bent; male abdomen without hair pFesent, sometimes very small or absent in pencils (exeept Myrmeeaelurini) 7 some genera (Atricholeon; Venezueleon) in which all leg setae are greatly reduced 3 7. Forewing vein 2A runs in fairly even eurve toward 3..A• (Fig. 11) 8 3. SOuthern Mrica; female terminalia with lateral Forewing vein 2A runs close to 1A for short gonapophysis elongate and with long down distance, then bends at sharp angle toward 3A ward curved setae whiCh are longer than Wig. 10) (modIfied In Pachyleon and Xantholeon gonapophyseal width (Fig 28); antennae close of the Nemoleontini) 9 together, antennal fossae separated by less than pedicel width...... 8 Hindwing posterior broad, nearly twice as high ...... Myrmecaelurini (Nannoleon) rIght before medIal fork than correspondmg New World; female terminalia with lateral area of forewing (similar to Fig. 14)' origin of gonapophysis small to moderately long, some­ hindwing radial sector often near or beyond times transverse, with or without digging se- medial fork, often 3 or more presectoral tae, but when present not longer than crossveins; female anterior gonapophysis plate­ gonapophyseal width (Figs. 17-22), antennal like or membranous; lateral gonapophysis elon- fossae usually separated by more than pedicel gate with long, downward curved digging setae width 4 (Fig. 29) .... Myrmecaelurini (Myrmecaelurina) Hindwing posterior area less than 1.5 x higher 4. Male eetoproet with postventral lobe at least than that offorewing before fOlks (Figs. 12, 13), longer than diameter (Figs. 41-43, 47, 49) or origin of hindwing radial sector before medial profemur with row ofclav 8te setae (Chaetoloon) fork, usually only 1 OF 2 preseetoral eFossveins; (Fig. 9) or mesofemoral sense hair longer than female anterior gonapophysis often lobe-like; remur CAllstroZeon); female terminalia with pos female lateral gonapophysis moderately long at terior gonapophysis subcylindrical (Figs. 17- most, digging setae if present shorter than 22), sometImes greatly reduced .. gonapophyseal width Dendroleontini Brachynemurini Male ectoproct WIthout postventral lobe (rarely 9. Hmdwmg WIth 1 (rarely 2, 3) presectoral produced but then shorter than greatest diam- crossveins; male pilula axillaris always absent eter of lobe); profemur WIthout row of clavate ...... NemoleontInI setae (clustered in Gnopholeon); mesofemoral Hindwing with 4 or more presectoral crossveins; sense haIr much shorter than femur except m male pIlula axIllarls present or absent...... 10 North American GnopMleon; female posterior gonapophysis swollen thumb-like (Figs. 24-26, 10. Basitarsusofhindleglongerthan apical tarsomere; 30-32), distally swollen (Fig. 2'1), or strongly procoxa with elongate white brIstles; female curved (Fig. 23, 34 ) 5 lateral gonapophysis with many long, down- curved digging setae as in Fig. 29 . 5. Tibial spurs absent (Fig. 40) or (in Gnopholeon) ...... Nesoleontini mesoremoral sense hair as long as or longer Basitarsus of hindleg shorter than apical than femur; Sonoran Subregion (southwest tarsomere; procoxa without white bristles; lat- United States, Mexico) Gnopholeontil1i elal gonapophysis without elongate, down- Tibial spurs present (Figs. 38, 39); mesofemoral curved bristles Myrmeleontini sense hair no more than 112 as long as femur; South America (Galapagos Islands; Ecuador; Larvae Chile; Argentina; south Brazil) .... Lemolemini (Larvae ofMaulim and Namwleon unknown) T,abia! paJpus shorter than basal width of man- 6. dible; mesothoracic spiracle not borne on tu­ bercle' head without dolichasters; backward movement only (many genera) or backward and orwar movemen ...... Labial palpus longer than basal width of man­ dible or mesothoracic spiracle borne on tu- bercle; head often with dolichasters; backward and forward movement.. 3 Vol. 8, No.1· 2, March· June, 1994 77

2. Mandible with some setae on outer lateral margin erossveins to posterior fork of MP2 or not reaching as long as or longer than greatest mandibular fork, Rs originates near or beyond medial fork, width; sternite VIII with pair of inoonspi6uoUS usually 3 or more presectoral crossveins, posterior submedian teeth near posterior margin; make pitfall traps MYrmeleontini Mandible with longest setae on outer lateral mar· gin less t.han one.half greatest mandibular width; sternite VIII without teeth on subapical gonapophysis membranous or plate·like; margm; do not make pItfall traps .. gonapophyseal plate mode18tely lalge to vely Ie· ...... Acanthaclisini duced; pregenital plate with or without median tooth; postenor gonapophyslS cylindncal, often reo 3. Mandible bases separated by less than width of duced; ffimale ectoproct with digging setae (except one mandibular base (Figs. 68·'11), Sonoran Ensorra). subregion Gnopholeontini Mandible bases separated more than width ofone mandibular base (Figs. 60 67) 4 Larval Description: Labial palpus longer than basal width of mandible, mandibles not upcurved, 4. Mesoseutum ·..... ith medial tuft of setae at middle; separated by much more than hasal width ofman. abdominal sternum IX usually at least as long dible, WIth mIddle tooth closer to distal tOOth than as wide; mandibles up turned (best viewed in to basal tooth; distal tooth shorter than middle profile) Dendroleontini tooth, set at different angle; head often with Mesos6utum "llithout medial tuft of setae; ab dolichastels, mesothOiacic spitacle pedunculate or dominal sternum IX wider than long; man- dibles straight .. .. 5 su~edian ~h 5 Abdominal terga I-I V with specialized setae sag- near posterior margin; abdomen ittally consisting of incurved setae or squat witout speciaized setae along median line; with· dolichasters; western and southern Sout.h out fossoria. America (especially Chile; Galapagos Islands) ...... Lemolemlnl Discussion: Thephylogenetic relationships ofthe Abdominal terga I-IV without specialized setae Brachynemllrini are difficult to understand On sagittally 6 the baSIS of the cladistic analYSIS of the hIgher categories (Fig. 1) and of genera previously eon 6. Mandible with distal tooth shorter than middle tained in the Brachynemurini (Fig. 3), I am defin­ tooth and usually set at different angle than othel teeth (with 3 teeth only) .. ing the HI achynem utini by the following

Nesoleontini WIt Mandible with distal tooth as long as or longel Gnopholeontini, Lemolemini, Maulini, some than middle tooth, teeth parallel (few genera DendroleontiniandNannoleon); (2) cylindricalpos­ with 2 teeth) Nemoleontini terior gonapophyses (found in some Old '!Iodd groups); (#3) short, deflected distal tooth of larval mandible (shared with Myrmecaelllrini and Tribe Brachynemurini Banks, 1927

Austroleonini Banks, 1927 from the Gnopholeontini and Lemolemini are (1) Adult Description: Labial palps short to moder the elongate postventrallohe ofthe male ect.oproct ately long, palpimacula not extending to apex; (found in the majority of the genera) and (2) the metafemur without sensory hair; pretarsal claws hmgedmaleparamere (foundm 50% ofthe genera). not toothed, rarely strongly bent near base; tibial spurs present or rarely absent (l species of Key to genera of Brachynemurini flbatoleou; 2 speCIes of Brachjtuefflurus) fOrewing vein CuP originates at or near crossvein M-Cu; 2A Adults runsin a failly even CUI ve tow ald 3A, pilulaaxillalis 1. Hindwing vein CuA bends to posterior margin at present or absent: hindwing CuA connected by or before medial fork, only 1 or 2 crossveins between CuA and posterior fork of MP2 (Fig. 78 Insecta Mundi

12), postelior alea of hindwing less than twice digging setae (Fig. 20), apex of hindwing in as high as corresponding area of forewing ..... 2 repose coincides with apex of forewing; Argen- Hindwing '1ein CuA runs aleng posterior fork of tina EnsoTM Navas MP2for a long distance, with 3 ormore crossveins Male sternite IX much smaller than sternite VIII; between them (Fig. 13); or posterior area of female eetoproet with digging setae (Fig. 22); hindwing broad (Fig. 14), at least twice as high apex of hindwing in repose clearly extends before medial furk as corresponding area of beyond apex of forewing; Mexico to Costa Rica forewing 8 D€juna Navas

2 Profemur wit.h clavate setae (Fig 9); ocular rim 8 Profemoral sense hair absent 9 usually With several short setae that project Protemoral sense hall' present 10 over eye. 3 Profemur without clavate setae; ocular rim with- 9. Antennal fossae separated by about Width ot out setae 4 pedicel; frons without setae; forewing costal area with costal cells interconnected; male 3. Male ectoplOct with postventral lobe at least ectoproct with postventrallobe more than ex as twice as long as middle diameter; female long as diameter at midlength (Fig. 41); North terminalia with a memLI anous plegenital plate, America Atlicholeon n. genus (Fig. 19); posterior gonapophysis more than 4X Antennal fossae separated by about 2X width of as long as diameter at midlength, "ithout scrap pedicel, flons with setae, fmewing costal alea ing setae; Neotropics Abatoleon Banks with cells not interconnected; male ectoproct Male eetopreet with post entrallone never longer "ith postventrallobe about as long as wide (Fig. than diameter at midlength, sometimes absent 47); South America ...... Venezueleon n. genus (Fig. 53); female terminalia with selerotized pregenital plate bearing median tooth; poste­ 10. Pretarsal claws as long as distal tarsomere; frons rior gonapophysis at most 3X as long as diam with numerous setae (Fig. 37); forowing CuP + eter at midlength, with scraping setae; North lA parallels posterior branch of CuA for a America Chaetaleon Banks distanoo, about equal to that between hindwing CuA and posterior branch of MP2; profemoral 4. Profemoral sense hall' at least three-fourths as sense hall' as long as temur; Venezuela to Ar­ long as femur, or pret.arsal claws longer than gentina Ameromyia Banks hind basltarsus (Austroleon hzen) 5 Pretarsal claws much shorter than distal Profemoral sense hair (if present) no more t.han t.arsomere (Fig 38), or frons witbout setae; one-halflength offemur; pretarsal claws longer or shorter than hind basitarsus 7 sense hair shorter than femur;2 5. Male ectoproct with conspicuous median lobe in ...... II addition to postventral lobe (Fig. 43); female abdomen usually longer than wings; profemur 11. Pretarsal claws shorter than hind basitarsus (Fig. and procoxa without row oflong white bristles; 38); tibial spurs longer than greatest tarsomere tempelate Nolth America . diameter; male ectoproct with postventrallobe ...... ,' Clathroneuria Banks about as long as wide, ventrally with a short Male eetoproet without median lobe, sometimes plocess (Fig. 51, 52) lJexoleon n. genus without postventrallobe; profemur and procoxa Pretarsal claws longer than hind basitarsus, or sometimes with 1'0'''' of long white bristles; tibial spurs absent or vestigial, male ectoploet female abdomen usually shorter than wings; witb postventral lobe longer than wide, not SOuth America .. . 6 direeted ventrally 12

6 Male ectoproct with postventral lobe at loast 8x 12. Posterior area of hindwing broad (Fig. 14), at longer than middle diameter; profemur without least twice as broad at medial fork as corre- row of long white bnstles .. sponding area of forewing; 4-6 crossveins be­ Argentoleon n genus tween hindwing euA and hind margin; male Male ectoproct Without postventral lobe or if pllula axillaris absent; setal bases on male short lohe developed, Jess than 2x longer than tergite IV often spined; vertex with dark pig Wide; profemur With row of long white setae .. ment, when present, restncted to scars ...... Austroleon Banks Seotoleon Banks PosterlOr area of hmdwmg narrow (Fig. 13), less 7. Male sternite IX enlarged, nearly as wide as t.han t.wice as wide at medial fork as corre- sternite VIII (Fig. 49); female ectuproct without sponding region of forewing, more than 10 Vol. 8, No.1· 2, March· June, 1994 79

cross.eins between euA and hind margin of 8. Ventral surface of head capsule with dolichasters hindwing or (in B. sackeni) vertex scars yellow- ...... 9 ish with dark pigment in intersear area; setal Ventral surfaee of head eapsule without bases on male tergite IV not spined; pilula dolichasters 10 axillaris present or absent ...... Brachynemurus Hagen 9. Head capsule longer than wide (measured ven­ trally); abdominal tergites with numerous Larvae thread-like setae between regular setae . 1. Sternite VIn without a pair of well defined ...... Ameromy~a (modesta Group) submedian teeth 2 Head capsule about as wide as long (measured Sternite VIII with a pair ofsclerotized submedian ventrally); abdommal tergites Without thread­ teeth (Fig. 16) g like set.ae VenezlIeleon

2. Mesothoracic spiracle not borne on tubercle ...... 10 Mesothoracic spiracle borne on tubercle that is ...... Scotoleon longer than its basal Width Austroleon Mesothoracic spiracle borne on tubercle MesothoraclC spIracle borne on tubercle that is ...... Atricholeon and Dejuna shorter than its basal width 11

3. Abdominal spiracles enlarged and/or borne on 11. Submedian teeth of sternite VIn weakly devel- tubercles . 4 oped, shorter than basal widths; head capsule Abdominal spiracles not enlarged or borne on (dorsal view) longer along dorsal midline than tubercles 6 across ocular tubercles, distance between basal and distal teeth less than distance between 4. Mandible longer than head capsule, with distal base and first tooth Abatoleon tooth separated from middle tooth by at least Submedian teeth of sternite VIn well developed length of distal tooth, lllesothoracic spiracle longer than basal widths; head capsule shorter borne on tubercle that is shorter than basal along dorsal midline than across ocular tu-

Included species: bruneli (Alavo) , camposi =FoyaNavas 1914. Broteria 12:53 ;afterEsben-Petersen 1920.19o). Type species: Fo a tmpezia Navas, by (Banks), deprivatus Banks, dolichogaster (Navas), original designation. dorsalis (Banks), frontalis (Banks), garciana (Banks); indiges (Walker). Nemotolus Banks, 1943. Bal. Ent. "enezolana 2.163 (after Stange 1967:45). Type species: Distribution: Nearctic and Neotropical Region prownsis Gel'staeeker, by original designation. (Florida Keys; Caribbean Islands; Southwestern Texas to Atgentina). Adult Description: FlOns with setae; antenna of same length on both sexes, moderately short with DISCUSSIon: The presence of clavate setae on the about 30 fiagellomeres; antennal fossae separated profemur distinguishes this genus from all others by about 2X pedicel width; profemur without clav- in the subtribe except for Chaetoleon. However, ate setae; ocular rim without setae; femoral sense sometimes the setae ale only weakly thickened hair of proleg as long as femur and much longel apically. The elongate male postventral ectoproct than thatofmesoleg; mesonotum withoutbladelike lobe and the alltapomorpbic medillnCIJS and set.ae; t.ibial spurs present; pret.arsal claws nearly parameres of the male genitalia distinguish this as long as distal tarsomere; pilula axillaris large; sex. In several characteristics Abatokon parallels h1Odw1Og as long as forew1Og or shorter; anterior Dejuna; both have highly specialized male genita banksian line present in both '.vings; forewing cos lia, similar wing venation, and are restricted to talareasimple; posterior area ofhindwing aboutas tropical climates broad as that offorewing, fmewing vein CuP + lA runs along posterior fork of CuA for long distance; The present center of dlVerslty of Abatoleon IS h1Odw1Og euA runs along posterior fork ofmedIal South ...A.merica, especially in the Peruvian coastal fork for a long distance; posPlentrallobe of male desert and Argentina. Many undescribed species ectoproct developed, usually less than 3X longer exist hi PelU. The two aberrant Atgentinian spe· than middle diameter; without median m second- cies, A. frontalis (Banks) andA. deprivatus Banks, ary lobes; male genitalia highly modified, gonarcus may belong to a different genus These two species reduced, medillncllS smooth, greatly expanded cov- have short abdomens and very elongate male ering modified rod·like parameres; genitalic sac postventral ectoproct lobes, and deprivatus lacks WIth clavate setae; female terminalia with pregeni. Vol. 8, No.1 - 2, March - June, 1994 81 tal plate membrano\:U;; gonapophy seal plate short; one-halflength ofdistal tarsomere; pilula axillaris lateral gonapophysis fused, with well developed large: hindwing shorter than forewing. in repose digging setae; ectoproct with well developed dig­ apicesofwings nearlycoincide; costalareaofforew­ ging setae ingwith one seriesofcells; posteriorareaofhindwing narrower than presectoral area, euAbends to hmd Larval Description (based on modesta and margin at or before origin of medial fork; anterior nigriventris): Head capsule with dolichasters; banksianline weakly developed; postventrallobe of mandible with distance between base and basal male ectopIOct more than 8 times langei than tooth less than between teeth; head capsule a little middle diameter, without secondary lobes; male longer than wide (measured ventrally) with or paramere plate-hke, WIthout hmge; gemtal sac without dolichasters; mesothoracic spiracle borne with lateral lobe like sclerite; mediuncus smooth; on tubercle; abdominal tergites sometimes with female terminalia with pregenital plate membra­ thread like setae; abdominal spiracle not enlarged nous, without median tooth; gonapophyseal plate norborneon tubercles; submedian tooth on sternite narrow, short, nottransverse; lateralgonapophyses VIII longer than basal width. separated, with strong digging setae; ectoproct with digging setae; posterior gonapophyses with- Included species: d~m~dwtaNavas, longwentns out scrappmg setae. (Navas), hirsuta Navas, modesta (Banks), muralli Navas, nigriventris (Walker), pentheri Navas, Larval Description (BasedonA. irrigatus): Head pleuralis Navas, pubiventris (Walker), pI otensus capsule with dolichastels, mandible with distance (Gerstaecker), stevensi Navas, strigosa (Banks), between base and basal tooth about equal to t.hat. and tendinosa (Gerstaecker). between teeth; mesothoracic spIracle borne on tu­ bercle that is longer than basal width; abdominal Geographic Range: South America east of the spiracles weakly raised; submedian tooth on stern­ l\ndes. ite VIII longer than basal width.

Discussion: This is a distinctive genus in which Included species: irngaws (Gerstaecker) & the pretarsal claws are greatly enlarged and the longitudinalis (Navas). male gemtalia hIghly modihed. The same modihed male genitalia are found in Veneweteen. The Distribution: South America east of the Andes. setose frons is shared only with Venezueleon. The PIOfemOIal sense hait is extIemely long, but the Discussion: Algentoleon has one autapoIDorphy, mesofemoral sense hair is quite short There are the development. of secondary, lobe-like sclerites two species groups. Larvae of the A. modesta from the gemtal sac. The enlargedpretarsal claws', Group have thread-like setae on the abdominal and elongate femoral sense hairs relate this genus tergites whereas the nigriventris group lack these to Austroleon and Ameromyia. The latter genus setae. Larvae have been found living in sand has many apomorphic chatactels (clavate setae of dunes. The larval head capsule is relatively wide, the male genitalia; form of the male parameres; but. less so than in t.he relat.ed Veuezueleou shape offorewingvein CliP + IN whiCh relate it to V€n€ZU€l€on and l~ss to Argentoleon. One variable 3. Argento/eon Stange, new genus species, A. irrigatus, ranges from Venezuela to Patagonia with some striking geographicfOims. In Type species: Myrmeleon irrigatu8 Gerstaecker, 1893, Patagonia, the individuals are much smaller than by present designation. elsewhere in its range A. longitudinalis (Navas) may only be a geographic variant of A. irrigatus, AdultDescription: Frons wIthoutsetae; antenna but evaluatIon of the geographIc VariatIOn present short, same length in both sexes, about 30 in A. irrigatus is needed to determine the validity flagellomeres; antennal fossae separated by about of longitudinalis. The larva was found in sand on width of pedicel; profemul without clavate setae, a glassy slope and resembles closely the larva of ocular rim without setae; profemoral sense hair as Austroleon. long as profumur and as long as mesOfemoral sense hair; mesonotum without blade-like setae; tibial spurs present; pretarsal claws large, longer than 82 Insecta Mundi

4. Atricholeon Stange, new genus development of sclelOtized and seLose plates ante­ (Figs. 41, 44) rior~gonarcusis an autB;P0;hy e two known speCIes ~r by numerous Type species: BrachjJnemurusparkeriStange, by present designation.

Adult Description. Frons without setae, anten- ofthis ~ateau. A. tUberculatusJlP~ears to be more nal fossae separated by about width of pedicel; profemur withoutclavate setae; ocularrim without mesoscu~rium setae; femoral sense hair absent; mesonotum with­ erances.e s;'ongyumped of out bladelike setae; tibial spurs present; pretarsal parkeri is apomorphic for that species. claws shorter than hind basitarsus and less than one halflength ofdistal tarsomere; pilula axillaris 5 Aus/m/eon BarIks smallwithsetae I eshicted to apicalmargin ofknob; hindwing shorter than or nearly equal to forewing; Austroleon Banks, 1909. J. New York Ent. Soc. 17:3. anterior hankgian line weakly developed; costal Type speeies: Aust7n dispar Banks, by subse areaofforewing mostly biareolate; posterior areaof hindwing narrower than presectoral area at radial -'Moza Navas, 1912. BroterIa 10:34 (after Stange sectoralthough wider than prefork area; CuA runs 1967'47) Type species: Mom nubilis Navas, by along posterior fork of MP2 for a short distance; original designation. antenna relatively long, usually with more than 40 flagellomeres; postventral lobe of male ectoproct =Correa Navas, 1914. Broteria 12:217 (after Stange lOnger than abdOminal segment VIII, without a 1967:47) Type species: Correa expansus Navas, median lobe; gonarcus simple; smooth mediuncus by original designation. Preoccupied by Correa which is small hooded process, paramere hinged, Fauvel, 1878 (Coleoptera). with elongate hook; secondary sclerotized and se- Rev. Acad. Madrid 19.251 (after tose plates present anterior to gonarcus; female T e s ecies: Clotus atomicus outa median tooth; medianfloor with sclerotization mediallyandsubapically; lateralgonapophysissepa =Calleo Navas, 1926. Brotelia 23.5. New name for rate, with very strongly developed digging setae; Correa Navas, 1914. ectoplOct with stlOngly developed digging setae, gonapophyseal plate relatively short and with fjne AdultDesctiption. FIOns withoutsetae; antenna striae. short, usually about 30 flagellomeres; antennal fossae separated by about widthofpedicel; pro£'emur Larval Description (based on parkeri): Head without clavate setae, but comb of setae present; capsule with dolichasters; mandible with distance ocular rim without setae; profemoral sense hair as between base and basal tooth less than between long as or shorter than profemur (except male A. teeth, mesothoracic spiracle borne on tubercle; ab­ lizen) and as long as mesofemUIal sense han, dominal spiracles not enlarged; submedian tooth mesonotum without blade-like setae; tibial spurs on sternite VIII reduced to bulge. present; pretarsalclaws longer than one-halfdistal tarsomere; pilula axillaris large; hindwing shorter Included species: parkeri (Stange) and than forewing, in repose apices of wings nearly tabel calatas (Banks). coincide, costal area offOIewing with one series of cells; posterior area of hindwing narrower than Distribution: Mexico, Arizona, and New Mexico. presectoral area, CuA bends to hind margin at or before origin of medial fOrk; anteriOr bankslan hne Discussion: The reduction in leg setae, particu- weakly developed or absent; postventral lobe of larry the loss of the femolal sense hair, is an male ectoproct absent or weakly de\' eloped (less apomorphy of this genus. This condition is dupli- than 2X longer than wide); without median or catedonly inDejuna mimicaand Venezueleon. The secondary lobes, maleparamereplate-like, without hinge; mediuncus micro spinose; female terminalia Vol. 8, No.1- 2, March - June, 1994 83 with pregenital plate membranous, without me­ implexa Navas ( Brachynemurus californicus dian tooth; gonapophysealplate narrow, short, not Banks), by monotypy. transverse; lateral gonapophyses separate, with strong digging setae; ectoproct with digging setae =Netroneurus Banks, 1927. Bull. Mus. Compo Zool. Harv. 68: 42. Type speeies: Braehynemul'us carolinus Banks, by original designation. LarvalDescription (based on immittt8; lizeri): Head capsule with dolichasters; mandible with Bollenga Navas, 1932. Bull. Mus. Zool. Torino (Ser. 3) distance between base and basal tOOttl about equal 42: 13. Type species: Bollenga dinamitensis Navas to that hetween teeth; mesothoracic spiracle borne (-BrachiYnemllrlls aachen; Hagen) on tubercle that IS longerthan basalwuith; abdomI­ nal spiracles weakly raised; submedian tooth on AdultDescription: Fronswithoutsetae; antenna stemite VIII longer than basal width. at least moderately long with more than 30 flagellomeres, that ofmale longer and less clavate Included species: dispar Banks, immitus than in female; antennal fossae separated by less (Walker), lizeri (Nav~s). than width of pedicel; profemur without clavate setae; ocular rim without setae; femoral sense hair DistributIon: South America east of the Andes. elongate, that of profemur longer than that of mesofemur but shorter than length of femur; Discussion: Austroleonis apomorphically defined mesonotum without blade-like setae; tibial spurs by the microspinose male mediuncus. The elongate present or absent (nebulosus and signatus); femoral sensehairofboth the prolegand mesoleg is pret.arsal claws equal to or longer than hind found m all speCIes except for male A. hzen whIch basitarsus but less than one-half length of distal has a shorter femoral sense hair. Larvae of two tarsomere; pHula axilaris present or absent; speCIes have been found in sand dunes in Venezu­ hindwing equal to or shorter than forewing; ante­ ela (A. immitus) and Argentina (A. lizen). Basedon rior banksian line usually well developed; costal larval exuviae these 2 larvae differ morefrom each area of forewing sometimes biareolate, narrower other than does A. immitus from the larvae of above radial sector than greatest presectoral area; Argentoleon irrigatus post~rior area of hindwing moderately broadened (exceptB. sacken~); much broadernearbase than at 6. Brachynemurus Hagen medial fork, euA runs along posterior fork of MP2 (Figs. 11, 13,16,17,42 for a longdistance; postventrallobeofmaleectoproct

urus a en ana ian n. Type species: Myrmeleon longicaudus Burmeister, by subsequent designation of Banks 1899.

Cal;nemUlU8 Banks, 1899. Canadian Ent. 31.70 (after Stange 1970:51). Type species: Brachynemurus ealifomicus Banks, by original designation.

Hespereleon Banks, 1913: Entomol. News 24:64 (after Stange 1970:51). Type species: Myrmeleon ferox Walker, by original designation. Larval Description (based on carolinus, Mastega NllyaS, 1914 Bull. Brooklyn Ent. SOG. 9: 13 divf.sus, exigua,terox, longicaudu8, nebulosu8, (after Stange 1970:51). Type species: Mastega ramburi, sackeni, seminolae): Head capsule texanus Navas (-8. lrregutans Currie). by monotypy. WIth dohchasters; mandIble with distance between base and basal tooth less than that between teeth; Comptesa Navas, 1915. Mem. R. Acad. Clen. Barcelona mesothoracic spiracleborneon tubercle; abdominal

Included species. abdominalis (Say), blandus lobe of male ectoproct simple or weakly produced, (Hagen). californicus Banks, carolinus Banks, about as broad as long. without median lobe but divisus Navas, elongatus Banks, exigua (Navas), sometimesproducedventrally; malegenitaliarather ferox (Walker), fuscus (Banks), hubbard; Currie, ordinary, paramere with hinge and weakly devel- trregulans Curne, longwaudus (BurmeIster), oped hook; female termmalia with pregenitalplate mexwanus Banks, henshawi (Hagen), nebuwsus selerotized with a small median tooth; (Olivier), pulchellus Banks, ramburi Banks, B. gonapophyseal plate large; posterior gonapophysis sackeni Hagen, B. seminome Stange, B. signatus with scraping setae, lateral gonapophyses sepa­ (Hagen), and B versutll.'i (Walker) rate, with digging setae

Distribution· Nearctic region to about 50" N Larval Description (based on pusillus): Head latItude and south to southern MexIco. capsule with many dolichasters; mandible with distance between base and basal tooth longer than Discussion: There is a great deal of structural between teeth; mesothoracic spiracle borneonweak diversity in this gwup, making it difficult to char­ tubercle, abdominalspiraclesnotenlargednorborne acterize as a whole No synapomorphies areknown on tubercles; submedian tooth on sternite VIII for the genus, consequently It may prove to be a shorter than basal WIdth. paraphyletic assemblage. The modified male ectoproct (Fig. 42) is common to about one-third of Included species: pumilis (Burmeister),pusillus the component species. Tibial spurs may be pwmi- (Currie), ttipunctatus (Banks) and variabilis nent, minute, or absent Thepilula axillarismayhe (Banks) well developed, weakly developed, or absent. The parallel tracksofthe Cu Aand posteriorfork ofMP2 Distribution: North America. m the hmdwing are a unifying generic characteris­ tic, except for B. saekeni. The genus shares with Discussion: The presence of profemoral clavate Scotoleon the synapomorphy of antennal sexual setae distinguishes Chaetoleon from all other dimorphism. Larvaehavebeenfound inopen sand, Brachynemurine genera except Abatoleon. The e g sand dunes presence of scraping setae on the female posterior gonapophysis is an autapomorphy of Chaetoleon 7. Chaeto/eon Banks and diagnostic within the Brachynemurini. Rela- (Figs. 7, 53, 54) tionship with Mexoleon is indicated by a weak ventral lobe oft.he male ectoproct (Compared with a Chaetoleon Banks, 1920. Bull. Mus. Compo Zoot. 64:328. strong ventral lobe in Mexol€Qu). There are two Type species: Myrmecoleon pumilis Burmeister, very distmct speCIes groups. The C. pumilis group by original designation. has a simple maleeetoproet, more elongatelegs and antennae, blade-like mesoscutal bristles, andmore AdultDescription: Fronswithoutsetae; antenna weakly swollen posterior gonapophyses relative to moderately long to very long; antennalfossae sepa­ the C. tripunctatus Group. One species, C. pusillus, rated by less than width of pedicel; prowmur with bas a bag-like structure on the male postmentum row of clavate setae; ocular rim ·Nith several short The larva of C. pusillus has been found in sand setae that project over eye; femoral sense hair of underplants. Chaetoleon variabilis has the female proleg much shorter than tarsus, longer than that scraping setae less de" eloped than in other species. ofmidfemur; mesonotum withconspicuous, weakly tlat.t.ened or bJadelike bristles; tibial spurs present; pretarsal claws longer or shorter than hind aSItarsus, not onger t an one- a length ofdistal tarsomel'e; pilula axillaris small with setae re- Clathroneuria Banks, 1913 Entomol News 24:65. stricted to distal margin ofknob; hindwing shorter Type Species: Brochynemurus schwarzi Currie, than forewing, in repose apices coincide; banksian by original designation lines absent; forewing costalareawith single series Of celIS; posteriOr area of hindwing narrower than AdultDescription: Fronswithoutsetae; antenna presectoral area before radial sector, euA bends to moderately long with over 30 flagellomeres; anten- hind margin at or near medial fork; postventral nal fossae separated by less than width ofpedicel; Vol. 8, No.1· 2, March· June, 1994 85 profemurwithoutclavate setae; Deular rim without AdultDescription: Fronswithoutsetae; antenna setae; femoral sense hair ofproleg unusually long short, usually with less than 30 flagellomeres; (as long as feIllUl), ove! twice as long as that of antennalfossae separatedby about widthofpedicel midfemur; mesonotum without bladelike setae; or more (D mimica); profemur without clavate tibial spurs present; pretarsal claws much longer setae; ocular nm WIthout setae; femoral sense haIr than hind basitarsus and more than one half as various (absent in mimica), that ofprofemurlonger long as distaltarsomere. Pilulaaxillarissmallwith than that of midfemur but shorter than length of setae restricted to apical margin ofknob, hindwing femur, mesonotuIll without blade-like setae, tibial shorter thanforewing, in repose wing apices nearly spurs present; pretarsal claws longer than hind coincide; anteriorbanksian line usually well devel­ basitarsus, no longer than one-halflength of dIstal oped; posterior area of hindwing narrower than tarsomere; pilula axillaris large with setae densely presectoral area Just before radial sector (except clustered on dorsal face of knob; hindwing as long coquilletti), euA bends to hind margin before me as forewing, in repose apex of hindwing extends dial fork or runs along posterior fork of MP2 for a beyond that of forewing; anterior banksian line short distance, forewing costal area with single weakly developed, costal area offOI ewi:ng with one series of cells; postventrallobe of male ectoproct series ofcells for mostofdistance; posterior area of with a median lobe in addition to postventrallobe hmdwmg narrower than presectOral area, euA (Fig 43); mediuncus smooth, long and narrow; bends to hind margin at or before medial fork; paramere hmged without hooks (Fig. 46); female postventrallobe ofmale ectoproct longer than terg­ terminalia with pregenital plate selerotized with a ite VIII, without a median lobe, gonarcus of male small median tooth; gonapophyseal plate expan- genitalia elongate, mediuncus with a spiniform sive, about as in Dejuna (Fig. 22), lateral process, parameres abbreviated (FIgS. 59, 60); fe­ gonapophyses separate, with digging setae; male terminaliawith pregenitalplatemembranous ectoproct WIth dIggmg setae. orsclerotIzed, WIthout median tooth; gonapophyseal plate large (Fig. 30); lateral gonapophyses sepa Larval Description: Larvae unknown. rate, with digging setae; female ectoproct with digging setae. Included Species: arapahoe Banks, coquilletti (Currie), navajo Banks, schwarzi (Currie), westcotti Larval DeSCription: Presumed larvae of D. (Stange) mimica have recently been discovered in Honduras and are being reared. They agree closely with Distribution: Southwestern USA and Northern larvae ofAtricholeon. Mexico. Included SpecIes: j'enestrata (Banks) ( angusta Discussion: The unusual male ectoproct (Fig. 43), Navas), mimica (Stange),persimila(Stange), setosa m whIch a median lobe IS developed m addItion to (Stange), and straminea (Stange). the postventral lobe is a diagnostic and autapomorphic character for this genus. Male Distribution: Arid tropical areas of North and genitalia are also distinctive in having an unusu- Central America. ally narrowed and elongate mediuncus A longer femoral sense hair helps to Identify the female sex. DISCUSSIOn: The male gemtaha (FIgS. 59, 60) are B. cgquilkttiis thQ only spQcies in the group haying autapomorphic for the genus. D. mimica which a broadhindwing posteriorarea, one similarto that lacksfemoral sensehairs, hasabbreviatedpretarsal found in Scatakan, and a short male eetoproct lobe. claws and tibial SpUlS. exhibits reduced leg setae, Although large populations of adults of both C. and has a small distal palpamere with reduced coquilletti and C. schwarzi have been discovered palplIllacula parallels m many ways speCIes of many times, all attempts to find larvae have failed. Atrichgl€gn.

9. Dejuna Navas 10. Ensorra Navas (Figs. 12, 22, 45) (Figs. 20, 49)

Dejuna Navas, 1924. Broteria 21:57. Type Species: Ensorra Navas, 1915. Broteria 13:10. Type Species: Dejuna angusta Navas, by original designation. Ensorra modestus Navas, by original designation. 86 Insecta Mundi

AdnltDescIiption: FIOns withoutsetae, antenna AdnltDescIiption. FIOns withoutsetae, antenna short with about 28 flagellomeres: antennal fossae with more than 35 flagellomeres; antennal fossae separatedby about widthofpedicel, profemurwith­ separated by less than width of pedicel; profemur out clavate setae; ocular rim without setae; femoral without clavate setae; ocular rim without setae; sensehall'shorton prolegand mesoleg; mesonotum femoral sense haIr moderately long (subequal m without bristles; tibial spurs present; pretarsal length to distaltarsomereofproleg),thatofprofemur claws about equal to basitarsus and much shorter about equal in length to that of midfemur; than one-haJflength ofdistal talsOInele, hindwing mesonotum without blade-like setae, tibial SpUIS shorter than forewing, in repose apices of wings and pretarsal claws shorter t.han hind basit.arsus, coincide; pilula axillaris present; costal area of lattershorterthanone-halfdistal tarsomere; pilula forewing simple; banksian lines absent; posterior axillaris with setae restricted to distal margin of area of hindwing narrower than presectoral area knob; hindwing shorter than forewing, in repose justbefore radialsector, hindwing\' ein CuA1 eaches apices coincide; anteliol banksian line poody de- hindmargin aboutat levelofmedialfork; banksian veloped: posterior area ofhindwing narrower than lines absent, male ectoploct with elongate greatest width of presectoral area, about as broad post.vent.ral lobe; male sternit.e IX unusually long; at base as at medial fork, euA runs along posterior male gemtaha WIth SImple gonarcus, smooth branch of medIal fork a short dIstance; forewmg mediuncus and plate like parameres; pregenital costal area at radial sector much narrower than plate of female sclerotized, without median tooth; greatest presectoral width, with one series ofcells; gonapophyseal plate nallOW, elongate, not tIans- postventlallobe ofmale ectoproct barely produced, verse; lateral gonapophyses separate, with digging about. as wide as long, ectoproct producedventrally setae; ectoproct without dIggmg setae: mtoa process (FIg. 112); male gemtaliaWIth hmged paramere and smooth mediuncus; genital sacwith Larval Description (based on verticalis): out secondary sclerites, and clavate setae; female Mandible with distance between base and basal teuninalia with plegenital plate scleIOtized, with a tooth less than that between teeth; mesothoracic median tooth anteriorly; gonapophysealplate mod- Spilacle bmne on tubercle, abdominal spiracles oot erately large (about as in Fig. 30); lateral modified; submedian tooth on sternite VIII shorter gonapophyses separate, with digging setae; than basal WIdth. ectoproct WIth strong dIgging setae.

Included Species: verticalis (Banks) (=modestus Larval Description (based on papago): Head Navas). capsule depressed posteriorly, with dolichasters, manyverysbort; mandible elongate, notbroadened Distribution: Argentma. subbasally, WIth dIstance between base and basal tooth shorter than that between basal and distal Discussion: This genus contains a single small teeth; mesothoracic spiracle borne on tubercle that species restricted to Argentina. The outstanding is aboutas long as basal width, abdominalspitacles autapomorphy ofthis species is the enlargement of enla'-.:;ed, raised; submedian teeth on sternite VIII male sternite IX (Fig. 49). The lack of clavate longer t.han basal widt.bs profemoralsetae andsetalcomb, absenceofdigging setae on the female ectopIOct, the plesenceofShm t Included Species: mixtecus (Stange) andpapago pretarsal claws and profemoral sense hair distin- (Currie).

Distribution: Arid t.ropical regions of sout.hern CO aeon pal i us b1.1 he on y arva avalla e for Mexico andmontaneregionsofsouthwesternUnited examination is damaged and therefore not keyed. States and northern MeXICO. The larva makes a pitfall trap. Discussion: Themaleectoproctwhich isproduced 11. M~~~~n ~rnge, new genus ventrally (Fig. 52) is diagnostic aodsyoapomorphic (FIgs. II 5 for the genus, although it is approached in struc­ ture by species ofChaetoleoll Mexoleoll, however, Type Species: Brachynemurus mixtecus Stange, lacks the clavate profemoral setae of Chaetoleon. by present deSIgnatIOn. Vol. 8, No.1 - 2, March· June, 1994 87

12. Seato/eon Barlks quadl ipunctatus (Currie), singulm is (Currie), and (Figs. 8, 14, 18, 60 63) yavapai (Currie)

Scotoleon Banks, 1913. Ent. News 24:64. Type Spe- Distribution: West oftheMississippiRiver, north cies: Brachynemurus longipalpis Hagen, by origi­ to about 45° North latItude and south to temperate nal designation. Mexico.

AdultDescription: Frons withoutsetae; antenna Discussion: The blOad posteIior area of the shortor elongate, usually longer andless clavate in hindwing (Fig 14) is the key character for the male; antennal fossae separated by about WIdth of group, whIch IS duplicated only m Clathroneuna pedieel; profemur without clavate setae; ocular rim coquilletti and Brachynemurus sackeni in the New without setae; femoral sense hair of proleg much World. Clathroneuria coquilletti is easily distin· shorter than distal tarsomere. not much longer guished by the long femoral sense hair, mediallobe than that of mesoleg; mesonotum without hlade· ofthe male ectoproct, andmale genitalia Scotoleon like seta; tIbIal spurs eVIdent; pretarsalclaws longer expansus tends to have this area narrower, espe· than hind basitarsus but not longer than one-half cialJy in smalJer specimens There is considerable length of distal tarsomere; pI1ula axilians absent; differentIatIOn of the labIal palps m thIS genus and hindwing shorter than forewing, in repose apices these differences are usually constant at the spe nearly coincide; anterior banksian line present; cific level. S. expansus and S. yavapai are rather forewing costal area with only one series of cells; aberrant species, especially in the modification of hindwing posterior area greatly broadened, about the male genitalia as broad near medIal fork as basally, usually WIth The loss ot the pilula aXIllans, broadened 6 or fev..er cross'leins between Cu}~ and posterior hindwing posterior area, andthe presence ofmodi margin; CuA diverges to hind margin along poste­ fied tergal setae on the male abdomen are generic rior fOrk of MP2 (Figs. 37, 38); postventrallobe of synapomorphies. This is probabh one of the most male ectoproct short to elongate, without second- actively evolving genera in the Brachyneumurini ary lobes; male geOltaba ordinary WIth slffiple judgingby the difficulty ofseparating some speCIes archedgonarcus andsmooth mediuncus; paramere and the presence of polytypic species The wing WIth hmge and hook (except S. expansus and S. venatIOn, espeCIally the broadenedhmdwmgposte· ,yavapai); genital sac without secondary sclerites rior area, is similar to that found in the Old World (except carrizonus and nigrescens); female Myrmecaelurini. Larvae have been identified for terminalia with pregenital plate scI emtized , with. most species and are different from other out a median tooth; gonapophysealplate small(Fig. Bracbynemurini (except Atricholeon and Dejuna) 32); lateral gonapophyses separate, WIth dlggmg m lackmg submedIal teeth on stermte VIII. setae; eetoproct with digging setae. 13. Venezueleon Stange, new genus Larval Description (based on all species ex- (Figs. 21, 47, 48) cept marshi, nigrescens, yavapai): Head cap- sule WIthout dolichasters; mandible with distance Type Species: Venezueleon guaricus Stange, by present designafjon asal an ista teeth; mesothoracic spiracle not borne on tubercle; abdominal spiracles not en- Adult Description: Frons with setae; antenna larged; sternite VIII without submedial teeth. same length in both sexes, about 30 flagellomeres; antennalfossaeseparatedby mOl e than twice width Included Species: carrizonus (Hagen), deflexus of pedicel; pronotum about twice as wide as long; (Adams), dtsstmths (Banks), eiseni (Banks), profemurwithoutclavatesetae orsetalcomb; femo· expansus (Navas), [idelitas (Adams), in[uscatus ralsense hairabsent; tibialspurspresent; pretarsal (Adams), intermedius (Currie), longipalpis (Hagen), claws shorter than hind basitarsus and one·half marshi (Stange), minusculus (Banks), miuutw: distal tarsomere, aboutsame length as tibial spurs, (Adams), niger (Currie), nigrescens (Stange), mesonotum i hi' . , , we eve ope w 1 e se ae pos enor y; m wmgas (Banks),peninsulanis (Banks), peregrinus (Hagen), long as forewing, in repose wing apices nearly 88 Insecta Mundi coincide, pilula axillatis well developed, weak ante­ capsule. The extreme width of the head capsule is rior banksian line in both wings; forewing costal autapomorphic for the genus and probably is asso- area slIl1ple; postenor area of hmdwmg narrower clated WIth Its mIcrohabItat. The absence ofthread­ than prQsQctoral arQa bQforQ radialsQctor; forQwing likQ SQtaQ on thQ abdominal tQrgitQs may bQ a larval vem CuP+ lAonlyruns along posteriorfork of CuA character from the A. modesta Group. There are for a short distance (in contrast to f.meromyia), four undescribed species, one ofwhich is described interconnected by about 4 crossveins; hindwing below. euA runs along posterior fork of medial fork for a short distance; radial sectororiginatesa littlebasad VenezueJeon guaricus Stange, new species of medial fork m hmdwmg, about 3-4 presectoral crossveins; post>t'entrallobe ofmale ectoproct mod Holotype male: Rio Orituco. 15 km S Calabozo, erately developed, lessthantwice aslong as median Guarico, Venezuela, February27,1988, R. B. Miller diameter; sternite IX much wider than long, not & L. A. StangQ (F'SCA). produced medially; male genit.alia highly modified, gonarcusbroadly expanded, plate-likelaterallyWIth AdultDescription: Length ofbody 32mm; length lat;al tooth; mQdiun~us arched, bifurcate yen offorewing 20 mm, greatest width 5l1lIn. General coloration pale brown with considerable yellow orange wing suffusion Head mostly pale brown with pregenitalplate membranous; gonapophyseal with darker brown vertex, anterior row of vertex plate elongate, posterior gonapophysis at least 3X scars most promment; frons WIth transverse light longer than median diameter; ectoproct and lateral brown stripe near middle; antenna nearly all pale gonapophysIS WIth strongly flattened diggmg se­ brown with darker brown on pedicel, extreme base tae; latter partially fused. of flagellomeres (especially dorsally) and apical spine; pronotum pale brown with median whitish Larval Description (basedongual icus). Man- strij)Q latQrad Of whiCh somQwhat darkQr brown dible as long as head capsule, distance between than laterally, setal bases darker brown; teeth more than that between base ofmandIble and mesoscutum and metascutum pale brown with basal t:r;,th;£Qad~JaP~IQas wtdQ ~ lonlL~measured darker brown sublaterally, scutelli with faint sublateral dark stripe; pleura pale brown, dark brown stripe on upper episternum; legs pale brown on tubercle thatis longerthan basalwidth; abdomi­ except darker brown ventrally; femora and tibiae nal tergites without thickened setae or thread-like with darkbrown stripe on mesal margin; tarsidark brown ventrally, wings with veins mostly pale brown, membrane suffused with yellow orange; basal v,idths. forewing with rbegmal mark along posteriorfotk of CuA near hind margin, several crossveins espe­ Distribution: Venezuela. cially in radial sector, near cubital fork, and crossveins at mar ins alon costal area and poste- rQlatQd to Amerom~pia, especially in regards to the sinlilar male genitalia, female terminalia (lateral Frons with setae; pronotum with outstanding gonapophy ses atleastpartially fused), widely sepa- white bristles along anterior, posterior, and lateral ratedantennalfossae, andfrontal setae. Leg struc­ margins; a few medially; scutelli with outstanding ture and wing venation offer significant differ- ....,hite bristles posteriorly; mesoseutum v. ith nu· ences. The pretarsal claws are weakly developed merous white bristles; legs without femoral sense andthefemoral senseharr IS absentin Venezueleon. hairand nearlywithoutoutstanding setae, a few on Fore;ving vein CuP .. Ii'. only parallels posterior femora and tibiae are sborter than diameter at fork of CuA for a short distance in contrast to point of origin; mesocoxa and metacoxa with a few Ameromyia The larva resembles those of the white bristles posteriorly; male abdomen with long AmeromyiamodestaGroup inhaving mesothoracic ~ale brown setae' those on sternites I-V twice as spiraclQ bornQ on an elongate tubercle, the rela (Jng as those 011 c~IIesPolldillg tergites, posteriorly tively wider head capsule, and the presence of setae become of eqllal lengths; postventral lobe of dolichasters on the ventral surface of the head Vol. 8, No.1· 2, March· June, 1994 89 male eetoproct with about 4 dark brown setae, GnopholeorItirIi Starlge, New Tr ibe mostly apically. Greatest ocular width about two-thirds Adult Description: Labial palps short, ~t~r~uI~rdistance: ~n~~nna with 30~~gellomeres; palpimacula not extending to apex; metafemUl IS a pa pornere of a mm not swo en, sensory without sensory hair; pretarsal claws not toothed opening nearly circular, located near middle and nor strongly bent near base; tibial spurs present. or occupying most of dorsal surface; pronotum twice absent; forewing vein CliP originates distad of as wide as long, proleg and mesoleg about equal in crossvem m-cu; forewmg vem 2A runs m a fmrly length, metaleg the longest; basitarsomere of even curve toward 3A; pilula axillaris present, metaleg about ax longer than diameter, somewhat usually well developed; hindwing vein CuA runs longer than that ofmes;l:g; dis~l tarsomerep~ut dose to hind margin, ends near medial fOlk; as long as tarsomeres . toget er; pretarsa c aw hindwing radial sector originates before medial about as long as hind basitarsus, about as long as fork, 2 or 3 presectoral crossveins; male abdomen tibial spurs; wings narrow, somewhat sinuate near without hair pencils or postventral lobe; male base, apexasymmetrical,forewingcostalcellslonger paramere plate-like; female anterior gonapophysis ~:; :j~ :~~:w~~o~;~~s~e~~::~~~r~~c~~~s~~:i plate like or membranous, posterior gonapophysis inflated; pregenitalplatewith tooth; gonapophyseal plate large; female ectoproct with digging setae

Larval DescriptIon: Mandibular bases close to- area narrower than prefork area; CuA runs along gether, separatedby aboutbasalwidth ofmandible; posterior fork of MP2 for short distance, intercon­ mandible with 3 teeth, middle tooth closer to distal n~ctedby4 cross~m' :dom;~uch W;:ger t~an tooth than basal tooth, distal tooth not shmter than middle tooth; abdomen without dolichasters or tufted setae along median area, sometimes with as greatest diameter; male genitalia as in Figure lateral scoli. 48. Discussion. This tribe is characteristic of the Sonoran Region. Larvae are either free living (Tyttholeott), live in prot.ect.ed areas (rock over- hangs) (Menkeleon) or live on rocks or tree trunks (Guopholsou). The larvae are unknown for

ciated with rocks. The principle synapomorphy of the genera in this tribe is the dose approximation of the larval mandibles. The inflated posterior gonapophyses (BIgS 31, 32) functiOn as egg han Paratypes: 9 males, 9 females, same data as types dlers rather than egg coaters according to Miller (CAS, IISNM, FSCA, Miller collection); 3 males, I (1990). E\' enmore modifiedposteriOi gonapophyses are found in the Lemolemini. Some species (espe- e. -1 , 1 ,. pang er (; U NM). cially Maracandula) are afternoon fliers. Discussion: There is some variation in this spe­ cies at the type locality. The degree ofwing suffu- Key to the genera of Gnopholeontini sion varies andsmaller specimens appear tohave a Adults relatIvely shorter and broader wmg. There are 1. Tibial spurs present (Fig. 39) . evidently 3 additional undescribed species, all Onopholeon Stange knownfrom single specimensfrom Venezuela. They Tibial spurs absent (Fig. 40) 2 differ in wing shape and coloration. Larvae were discovered at the type locality. Theylive in shallow 2. Wings narrow with sparsevenation; male ectoproct sand, anchored to bedrock Two adults were reared produced mesally below (Fig 50)' posterIOr to confirm the association. gonapophysis of female subcylindrical; prege· 90 Insecta Mundi

nital plate very large (Fig. 30) . Adult Description: Legs short and lelatively . Tyttholeon Adams stocky, profemurless than 1.5Xlongerthanprocoxa; "lings broad -".;ith denser venation; male eetoproct hindbasitarsusmuchshorterthan distaltarsomere; not produced mesally; posterior gonapophysis fQmoral SQnSQ hair QlongatQ, onQ on prolQg and offemale greatly swollen, pregenital plate small (Figs. 31, 32) 3 mesoleg; pretarsal claws and tibial spurs well de­ veloped, wings narrow with sparse venation, 3 Antennal tlagellomere 3 at least 1.5X longer than banksian lines absent; hypostigmatic cell without wide (F'lg. 36); cubItal tork of forewmg and crossveins, posteriorareaofhindwingnarrow, euA medial fork of hindwing arise near middle of bends to hind margin near medial fork; pilula wmg; pretarsal claws more than 2X longer than aXInans wendeveloped WIth setae covermg mostof greatest diameter of apical tarsomere; female dorsal surface of knob; male genitalia with simple lateral gonapophyses fused (FIg. 31) . gonarcus, free parameres; male ectoproct simple, An·w·~~~·i·fi~~~ii~·~~·~~· 3··~b~~r:a~o~:laasC~~~~ postventlallobe weakly developed at most; female termin Ii wi h ri r n h i w II n I a Ia or 0 hindwing arise well before middle of wing; urn e; pregeni a p a e narrow WI pretarsal claws not much longer than greatest tooth; gonapophysQal plate Qxpansive. diameter of apical tarsomere (Fig. 40); female lateral gonapophyses not fused .. Lanai Description (based on hal he' i): Head ...... Menkeleon Stange capsule greatly modified, much wider than long, greatly produced anterolaterally into a pair of ro- Larvae bust stemmatal tubercles which bear long setae 1. Mesothoracic sternum with specialized patch of WhICh camouflage mandibles m trap pOSitIOn; man­ lanceolate setae at middle (Fig. 71); labial pal- dible elongate, narrow, ...... ith three closely spaced, pus very long, distal palpomere beyond sensory equidistant teeth far removed from base; mesotho­ area more than 5X lon~er than wide' head capsule subquadrate, p;lterolateral c~rners lacic spiracle not borne on tubercle; meso- and roduced laterall i. 71 . live in 0 en sand metathoracic as well as abdominal segments I-VIII ...... Tyttholeon Adams eac WI a I a era y symme rIca parr 0 arge MetathOlacic stemum without specialized patch scoli; pretarsal claws of hindleg no longer than ofsetae; labial palpus short to moderately long, those of mesoleg; submedian teeth on sternite VIII distal palpomere less than 3X longer than wide longel than basal widths. beyond sensory area; head capsule various, but not mueh produeed at posterolateral corners .. Included Species· barbed (Currie), dPUcatulus ...... 2 (Currie), and zapotecus Stange. 2. Abdomen with scoli (except 1st instar); head greatly produced at anterolateral areas which Distribution: Deserts of southwestern United bearstemmatal tubercles, with long setae which States to southern Mexico. camouflage mandible in trap position (Fig 68); maximum jaw gape ca. 260°; mandibular teeth Discussion: The female genitalia provide funda­ about equidist.ant.· on t.ree bark or bare rock mental diIIQrences from all other gQnera in the faces GnopholPfln Stange tribe except for Menkeleon. The small pregenital Abdomen WIthout scoh; head not greatly pro- plate is a notable difference from Tyttholeon; and f:~:;~~:O~:~n~~~tit~a::::~~~~:e~::~~: the sepalate late!al gonapophyses, smalie! poste- tooth much closer to distal tooth than basal rior gonapophysis, stockier legs with one elongate tooth, under rock overhangs .. femoral sense hair, sparserwingvenation, and well ...... Menkeleun Stange developed tibial spurs are important differences from Maracandula. Thewelldevelopedtibialspurs 13. GnophoJeon Stange ale a key diagnostic ttait I eadily diffelentiating (Figs 15, 32, 39, 68-69) Gnopholeon from all other genera in the tribe. This may be a paraphyltic genus since the two species groups differ in characters that are of ge- Gnopheleen Stange, 1970. Uni.... Calif. Pub. Ent. 55:148. Type Species: Gnopholeon zapotecu8 Stange, by nerie importanee in the related Braehynemurini. original designation. The G. barberi Group have forefemoral clavate setae (absent in G. delicatulus Group) whereas the Vol. 8, No.1- 2, March - June, 1994 91 lack of digging setae on the female ectopIOct of the truncate pIOcess near origin of weakly developed G. delicatulus Group is an apomorphy of that postventral lobe, somewhat produced ventrally; group. The pretarsal claws are also larger than in female terminalia with posterIOr gonapophysiS ex­ the G barberi Group This taxonomic decision is tremelyenlargedandswollen; lateralgonapophyses bemgdeferreduntilthelarvalstageofG. delu:atUlus fused; pregemtal plate narrow WIth medIan tooth; is known. The larva ofthe G. barberi Group is one gonapophyseal plate expansive. ofthe most bizarre larvaltypes known in the family MYllueleontidae and resembles in some ways the Larval Description: Unknown. larvae ofAscalaphidae The hind pretarsal claws are not enlarged as in other antlions and the Included Species: apu:al£s (Banks) andpygmaeus mandibular bases are the most approximated of (Hagen). any known larva. The head capsule IS highly modified, with the mandibles capable of a Distribution: Central and Southern Mexican gape more than 260" which is a strong apomorphy. highlands. The abdominal scoli are well developed in later instal'S Thelarva was reared through the 3 instars Discussion: This genus exhibits a number of from an egg laId by a captIve female. It IS suspected unusualcharacters. Thefemale gemtaliaarehighly that the larva lives on tree trunks due to itscamou specialized with the posterior gonapophysis ex flage and structure. tremely enlarged and swollen and the lateral gonapophyses fused. The presence of several 14 Maracandula Currie crossveins in the hypostigmatic cell in the de- (Figs. 31, 36, 59) scribed species IS a feature not found elsewhere m the Myrmeleontidae of the Western Hemisphere, Maracandula Currie, 1901. Proc. ent. Soc. Wash. 4:436. although some individuals of Menkeleon bellula Type species: Mjrmeloon pygmORUs Hagen by approach thiscondition. Also of unusual interest is original designation. Banks 1927. Bull. Mus. Compo the extreme reduction ofthe eyes and highly modi- Zoo1. 68:58 (further description). Stange 1970. Umv. fled antenna in the described species Calif. Publ. Ent. 55:154 (further description). (autapomorphies) Theclosest relative is the genus Menkeleon. The female gemtaha ofMaracandula -Microleon Banks, 1901. Trans. Amer. ent. Soc. 27:365. appeal' more specialized than in lJenkeleon with Type species: Microleon apicalis Banks, by origi­ the lateral gonapophyses fused and the posterior nal designation. Preoccupied by Microloon Butler, 1885. gonapophysis greatlyenlarged Also, t.he suhapicaJ1y broadened wings of Maracandula are distinctive =Mimoleon, Banks 1913. Trans. Amer. ent. Soc. 39:226. However, m 2 undeSCrIbed speCIes ofMaracandulii; Ne w name for ..Yicroloon Banks, 1901. leg and wing characters are more similar to Menkeleon. From the spurless genus Tyttholeon, Adult Description. Antennae withflagellomeres the small pregenital plate, differently fonned male 2-4 much longer than wide; legs moderately long ectoproct, andwingvenationoffer importantdiffer­ and slender, pro£emur more than 1 5X longer than ences There are t.wo described species and several procoxa; hind basitarsus shorter or longer than undescribed ones. The adults in the afternoon. diStal tarsomere; femoral sense hair small and hardly distinguishable flOm stllrounmng bristles, one each on proleg and mesoleg; pretarsal claws with dense venation; banksian lines absent; Menkeloon Stange, 1970. Univ. Calif. Pub. Ent. 55' hypostIgmatIc cell WIth or WIthout several Type Species: Maracandula bellula Banks, by crossveins; posteriorareaofhindwingnarrow, CuA 01 iginal designation. bends to hind margin near medial fork; pilula axillaIis well developed with setae covering most of Adult Description: Ant.ennae with tlagellomeres dorsal surface of knob; male genitalia with rather 2-4 about as long as wide; legs moderately long and simple gonarcns, free parameres; eversible sac slender, protemur more than 1.5X longor than present laterally and ventral to male genitalia; procoxa; hind basitarsus longer than distal mesal margin of male ectoproct with short, rather tarsomere; twosubequal, rather longfemoral sense 92 Insecta Mundi hairs on pIoleg and mesoleg, wings bIoad with shoItel than hind basitaIsus which is eitheI IongeI rather dense venation, hypostigmatic cell usually or shorter than distal tarsomere; wings slender, without crossveins; banksian lines absent; poste­ with sparse venation; hypostigmatic cell without rior area of hindwing narrow, euA bends to hind crossveins; banksian lines absent; hindwing poste- marglO near medmIfork; pIlula axIllarIs smallwIth rIor area narrow, CuA bends to hindmargin before setae restricted to apical margin of knob; male medialfork; pilula axillaris moderately well devel genitalia with rather simple gonarcus, free oped with setae concentrated toward distal margin paIameIeS, male ectopIOct lathel simple, some­ of knob, male genitalia with simple gonarcus, free what produced ventrally, postventral lobe weakly parameres; male ectoproct produced mesally be- developed; female termlOaha WIth posterior low, no postventral lobe; female posterior gonapophysis swolhm thumblike; lateral gonapophysis subcylindrical, lateral gonapophysis gonapophyses separate; pregenital plate narrow separate; pregenital plate much enlarged with a with median tooth; gonapophysealplateexpanswe. prominent median proeess postelioIly, gonapophyseal plate large. LaIval Description (based on bellalas): Head capsule wider than long, weakly produced Larval Description (based on puerilis)- Head posterolateraIIy; labmlpalpusmoderately long, dis­ capsule WIder thanlong, producedposterolateraIIy, talpalpomere less than 3X longerthan wide beyond with doliehasters; labialpalpusextraordinarilyeIon sensory area; mesothoracic spiracle borne on tu­ gate; distal palpomere beyond sensory area more belcle, mesostelOum withoutspecializedsetalpatch than 5x lungel than wide, mesothoracic spiracle at middle; abdomen without scoli horne on elongate tubercle; mesotboracic sternum WIth speCIaliZed lanceolatesetaeatmIddle; pretarsaI Included Species- Only b€llulus (Banks). clawsofhindlegabout 2x longerthanthatofmesoleg; abdomen withoutscoli; submedian tooth ofsternite Distribution: SouthwesternUnitedStates, north- VIII longer than basal width. ern Mexico. Included Species: puerilis Adams. Discussion- The female terminaJia ofMenkeleon are qUIte SlIDlIar to Gnopholeon but Menketeon Distribution: MOJave-Colorado Desert; Great lacks tibial spurs, Superficially, the species of Basin; Baja California. Menkeleon resembles those of Marcreandula but genelic distinctions ale detailed in the key. The Discussion: The highly modified female pregeni- presence of 2 short femoral sense hairs is tal plate (Fig 30) and shape of the male ectoproct autapomorphlC. The larva was found under an (F'Ig. 7I) are autapomorphic. The larva (FIg. 65) IS overhanging rock. also very distinctive with the development of Ian ceolate setaeon themetasternum (autapomorphic). 16. Tyttho,'eon Adams Other distinctive charactels are the loss of tibial (Figs. 30, 50, 57, 71 spurs (present in Gnopholeon) and narrow wings

liv~s th~ On~ TyttholeonAdams, 1957. Psyche 63:106. Type Species: The larva in sand. undescribed Tyttholeon puerilis Adams, by original designation. species is found in southern Baja California. . Stange 1970.157. Lemolemini Stange, new tribe Adult Description: Antenna with 20-25 flagellomeres, flagellomere 1 longer than wide, Adult Description: Labial palps short, sensory about one-half length of pedicel and subequal to area oval; metafemur without sensory hair; flagellomere 2; greatest ocular width about one- pretarsal claws not toothed nor strongly bent near half interocular distance; legs rather short and base; tibial spurs present, sometimes shorter in stocky. profemur about 1 5 times longer than male; fOrewing vein CuP originat~s distad Of bas~; procoxa; hindlega littlelongerthanproleg; profemur forewing vein 2A runs a fairly even curve toward WIthout clavat~ s~ta~; prot~moraI sens~ hair much 3A; pilula axillaris present; hindwing as long as less than one-halflength ofprofemur, equal to that forewing, in repose apex of hindwing extends be- of midfemur; tibial spurs absent; pretarsal claws yond that of forewing (except Galapagoleon); Vol. 8, No.1 - 2, March - June, 1994 93 hindwing'\lein CuAIuns close to hind maIgin, ends Pretarsal clav;s longer than length of proleg nearmedialfork; hindwing radialsectororiginates basitarsus; profemoral sense hair at least 1/2 well before medial fork, usually 2 presectoral length of femur; antennal fossae separated by crossveins; abdomen without hair pencils, shorter not more than width ofpedicel; female posterior gonapophysis shorter than height of elltoproet than wmgs, about same length In both sexes; male or strongly curved mesally (Figs. 23, 24, 26, 34) ectoproct without postventrallobe; male paramere 3 plate-like; female anterior gonapophysis plate-like Ol memblanous, posteliUl gonapophysis highly 2. Forewmg costal area biareolate m part; hmdwmg modified, either swollen or strongly curved; shorter than forewi ng, in repose apices of wings gonapophyseal plate usually very large (except nearly comcide; male ectoproct With large ven­ Elieura and Lemolemus); female ectoproct without tral process' female gonapopbysis evenly swol- digging setae. len for complete length, with apiCal scrapmg setae (Fig. 25) Jaffuelia Forewing costal area with single series of L-ells, LarvalDescription: Mandibularbasesfar a~art, hindwing as long as forewing, in repose apex of sepalated by much mOl e than basal width ofnan­ hindwing extends beyond that offorewing; male dible; mandible with 3 parallel teeth, middle tooth ectoproct without • entral process, female closer to distal tooth than basal tooth, dIstal tooth gonapophysis swollen toward apex, without is longer than middle tooth; basal tooth distal to seraping setae (Fig. 27) I·,reulatu8 midpoint of mandible; abdominal segments with dolichasters OJ tufted setae along midline, some- 3. Male eGtoproet with indieation ofpostventrallobe; timeswithlateralscoli; sterniteVIIIwith submedian female posterior gonapophysis longer than teeth. height of elltoprollt, strongly lluFVed mesally (Fig. 23, 34); Galapagos Islands; southwestern Ecuador 4 Discussion: ThIS trIbe IS apomorphlCally defined Male ectoproct without indication of postventral by the specialized setae found medially on the lobe' CAm ale post.enorgonapophySIS shorterthan abdominal tergites of the larva. The adults have height ofectoproct, not strongly curved mesally one apomOlphic chalacteI, the lack ufdigging setae (FigS. 24, 26); Central Chile; Argentma; Uru­ on the female ectoproct This is evidently a relict guay; southern Brazil 5 group WIthout close relatIves. Oddly, although some larvae (Lemolemus, Sieal, Elieura) live in 4. Forewing posterior area highest near cubital fork; loose soil under rock overhangs or near tree bases, female tergite VII with subapical row ofpromi­ none has invaded the sand dune habitats which nent setae (Fig. 33) ; male ectoproct mesally emarginate ventrally (Fig. 35), Western Ecua- abound in Central Chile This tribe is most diverse dor Ecualeon in Chile where 5 oC the 7 genera occur. Three Forewing posterior area highest near end of vein g~er~ ~lieura, Lemol~us, and Sieal are closely 2A, female tergite VII without subapical row of prominent setae; male ectoproct not emargin- ate mesally; Galapagos Islands . Ecualeon, found in western Ecuador, is assigned to ...... Galapagoleon the tribe based 011 a presumed larva which agrees with Lemolemioi and adult characters 5 Profemur "'ith long white bristles along exterior Galapagoleon IS unknown III the larval stage and IS face; forewing costal area broad, area above tentatively referred to the Lemolemini since the cubital fork nearly ax as high as subcostal area, female tenninalia are similar to Ecualeon. often triareolate; mesofemoral sense hair about 1/2 length of femur; female terminalia "'ith Key to genera of Lemolemini anterior gonapophysis plate-like (Fig. 24); pre- Adults 1. Pretalsal claws shorter than length of proleg basitarsus; profemoral sense hair about 1/4 length of femur; antennal fossae separated by Profemur without long bristles along exterior more than width of pedicel; female posterior face, except sometimes 1 or 2 subapiCal ones; gonapophysis longer than height of eetoproet, forewing costal area narrower, area above not curved (Figs. 25, 27) 2 cubital fork less than 2X as high as subcostal area, biareolate at most; mesofemoral sense hairabout 1I4lengthoffemur;female terminalia 94 Insecta Mundi

with anterior gonapophysis lobe-like (Fig. 26), AdultDesetiption. Antennalfossae sepalatedby pregenital plate wider than long; Central Chile about width of scape; distal palpomere of labium ...... 6 moderately swollen; profemurwithoutwhitebristles on exteriorface except lor2 subapicalones; procoxa 6. Basitarsus of metaleg less than 3X longer than WIth 2 whIte bnsties anterIOrly and posterIOrly, wide; forewing costal area with few, if any, interconnected cross"eins in basal half; prege setaemOl'e thanone halfeoxaldiameter; profemOl'al nital plate large, without median tooth Sical sensehairaboutone-halflengthofprofemur, about Basitarsus of metaleg at least 4X longer than as long as that ofmesofemUl, pretarsal claws mod­ Wide; forewing costal area with most crossveins erately developed, longer than pro-basitarsus but mterconnected; pregemtal plate small, WIth shorter than tIbIal spurs whIch are longer than median tooth T,emolemu.tl meta basitarsus; tibialspurs ofsame length inboth sexes; forewing costal area biareolate in about distal one-foUlth to one-thild; fmewing ladialsec- (CU on and Galapagoleon unknown) tor arises well basad ofcubital fork, in contrast to 1. Abdominal segments I VII with 'oVell developed position ofhindwmg radialsectorwhich ansesonly scoli, at least longer than wide (Figs. 62, 63); little basad of medialfork; bindwing usually witb 4 abdomen without median row of tufted b1aek setae but with roseate, squat dolichasters; presectoralcrossveins;forewingposteriorareahigh­ pretarsal claws with basal tooth 2 estneal'cubitalfOl'k; anterior banksian line absent; Abdominal segments I-VII without scoli, or if hindwing nearly as long as forewing, in repose short scob present, then WIder than long; abdo­ hindwing apex extends somewhat beyond that of men with median row of tufted black setae forewing; male sternite IX large, much narrowed (except first mstar), WIthout roseate postenorly; maleabdomen much longer than wmgs, dolichasters; pretarsal claws wit.hollt. hasal t.oot.h segment VIII at least 4X longer than high; seale ...... 3 like setaeon segmentsIII-V; male ectoproct bilobed 2. Abdomjna~ scoli at I:st 3X lo~ge~ th; w~' ventIally on mesal malgin (Fig. 35), postventral tapered 0 apex, wi out dolic as rsig~ ; head longer than wide; live in Puya plants ..... lobes indicated; gonarcus nearly transverse, ...... Neulatus Navas mediuncus large, smooth, except doubly ndged

broadened apically, with dolichasters (Fig. 63); cess ig. 35); ema e wit large, transverse prege­ head wider than long, live on boulders .. nitalplatewith minute medialtooth, gonapophyseal ...... Jaffuelia Navas plate large and expansive; posterior gonapophysis swollen and curved subapically; anterior 3. Abdominal segments I-VII with weak scoli; head gonapophysis swollen, transverse, witb many elan- eapsule and seoli '.vith abundant silky ';;hite, gate setae; ectoproct WIth digging setae, those on hairlike setae Sical Navas l'..bdominal segments I VII without seoli; head lateral gonapophysis moderately developed; stem- capsule and scoli without silky, white, hairlike ite VII not produced medially; tergite IX offemale setae 4 not produced ventrally; tergite VII of female with subapical row of large, curved setae (Fig. 33). 4 Mandible pale with numerous dark brown spots; anterior scolns of mesothorax 2X longer than Distribution: Southwest Ecuador

Discussion: This genus is plOvisionallv assigned ase; an erlor scolus 0 meso orax. onger to the Lemolemini. It appears related to than wide ...... T,emolemu.s Navas GalapagoTMn especially in regards to the curved posterior gonapophysis of the female terminalia. The structure ofthe female abdomen suggests that 17. Ecualeon Stange, New Genus this genus may throw similar to that obser ved (Figs. 33-35) in Psammoleonwhichhas similarfemale terminalia. The presence of the prominent row of setae on Type species: Ecualeon ovispargus Stange, by tergite VII (Fig. 33) may be a significant sign ofsuch present designation behaviOr. Another significant character of thiS genus is the broadening of the posterior area near thecubitalfork. Quiteunusualandunique, atleast Vol. 8, No.1- 2, March - June, 1994 95 among New 'vVorld antlions, is the development of Flons without setae, clypeus with transvelse large, scale-like setae on male abdominal segments row of 4 long, black setae; scattered decumbent III-V. The male ectoproct (FIg. 35) IS emargmate setae on vertex; labmm with many long black setae ventrally which is unique in the tribe decreasing in length distally; pronotum with long (some one-thIrd length of pronotal WIdth) setae on margins, shorter ones on middle, blackones mostly Ecualeon ovispargus Stange, New Species lateral, white ones medial; prescutum with 3-4 black OI white setae in a lOW, mesonotum with 3-5 Holotype male. 7 km. S. Piiias, PlOvince EIOIO, w~e !'l~~a~ in a ~o: np.Rr middle' m~so!'lcute)]u; Ecuador, July7, 1989, R. Miller & L. Stange (Florida State Conection Of Aithropods:) bristles anteriorly and posteriorly, posterior ones AdultD.escription: Length of~ody84mm; length somewhat longel than coxal width, antelior ones shorter' rofemur wi h u ri I n x ri r coloration yellowish brown, wing membrane trans­ gm excep a ew su apica ones; c osmg ace 0 a parent except small brownish suffusion at nearly femora and tibt! and ~xterior face of t~biae ~ith everyveinjuncture. Head colorvariable, from pale manyelongatenstiest atare mostlytWIce asong to dark brown; anteriOr row of vertex scars most as femur or tibia, black bristles on exterior faee' prominent, dark brown lateral scar and smaller profemoral sense hair about as long as that of submedian dark brown scar; middle row reduced to mesofemur, about one-half length of femur; wing submedian dark brown area; posterior row with ;:~;:r;~?n V~~ ~:~e~~;:~~~~:Cl~:r:.t1~:; a~~ prominent dark brown lateral scar, middle scar lighterbrown; clypeu8 andlabiUm mostly yellOwiSh ubiquitous setae; tergites II V and sternites III V brown, gena mostly dark brown, prominent dark with numerous brown, scale-like setae, some of brown spotbeneathantennalfossa; postgenanearly which on stelnites ale 3X as long as on tergites; completely shiny dark brown; palpi dark brown setae mostly at middle of tergites and sternites. except pale apically except distal palpomeres; an­ Greatest ocular WIdth nearly as great as tenna neRrly RJ) brown, seRpe pale brown posteri- ~teroculardistance; antennawith 30flagellomeres; orly and less so anteriorly; pedicel and base of agellomere 1-13 about 1.5X longer than wide, flageIIomere 1 dark brown; pronotum mostlY pale abruptly broadening beyond so that flagellomele 20 about 3X broader than long; distal palpomere of IRbium somewhRt swollen, circular sensory areR a and metanotum similar to pronotum with pale brown medially, darkbrown laterally; scutellimostly pale brown; mesopleuron dark brown anteriOI'Iy, mostly palebrown posteriorly; metapleuron mostly mesoleg about 3X longer than diameter, aboutone­ palebrown with scatteled dalkblown aleas, coxae half length of distal tarsomere; hind hasitarsus andmera mostl ale brown meso- andme a-coxa about 4X longer than diameter; tibial spurs reach e own pos erior y; emora an i iae pa e brown with dark brown spots at setal bases and tarsus, and somewhat beyond apex ofmetatarsus; apically, especially profemur and mesofemur; tarsi pretarsal claws shorter than tibial spurs, about as pale brown with dark bro...m mostly on tarsomeres long as pIO-basitalsus; hindwing nearly as long as 4 and 5 and aJ!ex of distal tarsomere' tibial sj!urs forewing, in repose apex of hindwing projects be­ yond that offurewing; wings all about same shape and claws I eddish brown, lighter dist~lly, forewing and hindwing nearly equaHy colored; wing veins but hindwing narrower; forewing costal area pale brown with extensive dark brown sections biareolate in distalone-half, cells higher than wide near vein and crossvein junctures; stigma pale in basal one-half, fOlewing with radial sector aris- ing well before cubital fork, 5-6 presectoral crossveins; CuP + 1A runs along posterior fork of euA for short distance, interconnected by 3 crossveins; posterior area highest near distal end' hindwing radial sector areas somewhat before me~ dial fork, 4-5 presectoral crossveins; hindwing pos- 96 Insecta Mundi terror area nallower than prefOlk area, highest AdultDesctiption: Antennalfossae separatedby near distal end; euA runs to hind margin just about width ofpedicel; profemur with row ofwhite belore medial lork; piluia aXIllarIs large; abdomen brIstles on exterIor lace; prolemoral and much longer than wings; segment VIII about 4X mesofemoral sense hair more than one halflength longer than high; male ectoproct with postventral offemur; pretarsalclawslonger thanprobasitarsus; lobe weakly indieated, bifurcate mesally (Fig. 35); tibial spurs of same length in both sexes; forewing male genitalia (Fig. 35) with gonarcus arched but costal area broad, about 3X higher than subcostal smaller than mediuncus which is a concave plate area above cubital fork, usually triareolate, ante­ doubly ridged apically; paramere flat plate, later- rior banksian line weakly developed; hindwing any produced. about same length as forewmg, m repose apex of hindwing extends beyond that of fore.....ing; male Female: About as described for male. Abdomen ectoproct simple, without lobes or processes; male shorter than wings; scale-like setae absent; tergite sternite IX much wider than long, weakly produced VII with posterior row oflong, posteriorly directed medially; male genitalia with arched gonarcus, bristles; terminalia (Fig. 33) with posterior mediuncus short, notridged; paramere longerthan gonapophysis swollen, strongly curved mesally, wide; much narrower at mediuncus; female about 2X longer than median diameter; terminalia with sternite VII not produced; tergite gonapophyseal plate expansive; anterior IX not produced ventrally; pregenital plate sclero tized, longer than wide, with medial tubercle; pos­ teriOl gmmpophysis swollen, much shorter than as' width; ectoproct with row of4 dorsally directed height of ectoproct; ectoproct an d lateral brIstles; pregemtal plate sclerotized, transverse, gonapophysls WIthout dlggmg setae; anterIOr with small median tubercle. gonapophysis plate like.

Paratypes: 3 males, 20 females, same date as Larval Description (based on litigatOt). Head holotype (MC, USNM, Quito, SC, FSCA). capsule longer than wide, with dolichasters; man- dlble pale WIth dark brown areas; anterIor me­ Biology: Adults were found flying among 5 foot sothoracic scolus 2X longer than wide, with pale tall grasses or bushes, usually in the shade. When dolichasters; abdominalscoliabsent; abdomen with disturbed they fell to the base of the grass whele median lOW of tufted black setae; pletarsal claws detection was difficult. None was attracted to withoutbasaltooth; mesothoracicspiracle borneon lights but Olle was found by jack lighting An tubercle that is longer than wide; abdominal spi- unidentified larva collected nnder a rock overhang racles enlarged, borne on tubercles. m Ecuador may be thIS speCIes. It has speclahzed setae along middle ofabdomenandabdominalseoli. Included Species: uuquus (Navas), justus (Navas), litigator Navas.

18. Elicura Navas Distribution: Southern Chile, Argentina; Uru­ (Fig. 24) guay; southQrn Brazil. Discussion: Stange (1967) placed some of the Elicura Navas, 1911. Rev. chilena Rist. nat. 15:126. Type Species. Elicura litigator Navas, b~ original species ofElicwa in Lemolemus which this genus designation. closely resembles. One notable difference between Elicura and T,emolemus is that the anterior =Plater Navas, 1919. Rev. Acad. civ. Madrid 17:297. gonapophysis is lobe-like in Lemolemus. Theforew- New synonym. Type Species: Plater iniquus ingcostalareaisverybroadin Elicura andprovides Navas, by original designation. one character to separate it from all othels in the tribe. Larvae of E. litigator were found in many Bridarollus Navas, 1933. Rev. Aead. Ciene. Zaragoza habitat.s such as rock overhangs, tree holes, and at 16:90. New synonym. Type Species: Bridarollus bases of trees and resemble Nemoleontini and sollirs Navas, by original designation. DendiOleontini in mandibular dentition. The dark spotting of the mandible is distinctive in the tribe. Vol. 8, No.1- 2, March - June, 1994 97

19. Jaffuefia Navas

a ue £a av so. oc. r one e Type species: Jaffuelia chilensis Navas, by origi­ nal designatioIl.

AdultDescription: Antennalfossae separatedby more than width of pedicel; distal palpomere of labmm slender; profemoral sense haIr short, less than one fourth length of femur, equal to that of mesofemur; pretarsal claws shorter than meta­ basitarsus, about same length as tibial SPUlS; tibial areamos y lareo a e; orewmgra la sec oranses WQIl basad of cubital fork about QquivalQut to thQ distance between radial sector and medial fork in T~pe hmdwmg; anterior banksian line weakly devel- Species: Brach;nemurus darwini Stange 1 69, by present desigl tion. oped; hindwing shorter than forewing, in repose wingapices nearlycoincide, male sterniteIX trans- AdultDescription: Antennalfossae separatedby Jess than width ofscape; distalpalpomere Of labIUm swollen; profemur without white bristles on exte- mediuncus, parameresplate like; female terminalia with posterior gonapophysis evenly swollen for riorface except 3 subapical ones; procoxa with only ~osterior~ complete length; with scraping setae; ectoproct and 2 white bristles that are less than one- half coxal diam1;er, profrnoral sense hair about lateral gonapophysis without digging setae; prege- t.hree-fourt.hs lengt.h offemur, over twice as long as mesofemoralsensehair; pretarsalclaws well devel­ oped, longer than basitarsus butshorter than tibial spurs; tibial spurs of same length in both sexes; forewing costal alea simple, except 1 UI 2 intercon- LarvalDescription (based on chilensis): Head . . nec e I' in' .. I I U arises we asa 0 cu I a or, m con ras 0 ~o:~::~l :~ta:~;~:i~a:lt~~~~s:~~r~:~~~~~:~)l~~ positiOn or hindwmg radial SQctor which arisQs a little basad of medial fork; hindwing with 2 though less than 2X longer than wide, with many dolichasters; abdomen with median row ofclumped presectoralcross"ems;for ew ingposterior ar eahigh­ dulichasters; pretalsal claws with basal tooth, me- est near end of vein 2A anterior banksian line developed, hindwing sho;ter than forewing, in re- sothoracic spiracle borne on tubercle that is wider than long; abdominal spiracles enlarged, but not much raisQd. arge, somew at WI er t an ong, earmg ong setae; ectoproet simple but postventral lobe indi Included Species. chilensis Navas andporterina cated, without ;rocesses' male ;enitalia with gonarcusa simple Ich, medluncus s newhatlonger Navas. than wide with ridges; paramere with hook; female Distribution: Central Chile. termmaha With large sclerotIzed pregemtal plate which lacks mQdial tooth; gonapophysealplatelarge and expansive; posteriorgonapophysis swollen and Discussion: Thisgenushasseveralautapomorphic CUI ved subapieally, ectoplOct without digging se­ characters in the tribe. The shape of the male . . tae those on lateral ona 0 h si onl m I' eve ope ;s erni e no pro uce me la y; erg- distinctive and the long, evenly swollen female fe~alQ n~t postQrlOr gonapophysis WIth scraping SQtaQ (Fig. ite IX of pr1uCQd v\utrally; antQrior 25) is diagnostic for that sex. The biareolate forew- ing costal area is duplicated only in Elicura and in 98 Insecta Mundi

Larval Description. Unknown. Lively shott, ectoplOct and lateial gonapophysis without digging setae Included Species: darwini (Stange). Larval Description (based on modestus and Distribution: Galapagos Islands. several unnamed species): Headcapsule longer than wide, with dolich8sters; mandible nearly uni Discussion: This genuscontainsonespeciesfound form color exceptdarker basally; anterior mesotho- ill the Galapagos Islands. Its taxonomic position is racic scolus 1.OX longer than wide, with pale unclear but is referred to the Lemolemini based on dolichasters; other scoli wider than long; abdomen overall SImIlarity. The curved posterior WIthout scoh; abdomen WIth medIan row of tufted gonapophysis is similartoEcualeon. The discovery black setae; pretarsal claws without basal tooth; ofthe larval stage will probably clarify its relation­ mesothoracic spiracle borne on tubercle that is ships. longel than wide; abdominal spilacles enialged, borne on tubercles. 21. Lema/emus Navas (Fig. 26) Included Species' modestus (Blanchard)

LemolemusNavas, 1911 Rev chilenaHist nat 15'123 Distribution: Central Chile. Type species: Lemolemus nectator Navas, by origi- nal deSIgnatIOn (- L. modestus (Blanchard) 1851). Discussion: This genus is in needofrevision. One taxonomic problem is that the males and females of -Puren Navas, 1911. Rev. chIlena RISt. nat. 15:125. the same speCIes are vanable ill the length of the New synonym. Type species: Puren bellator tibial spurs. The lobe like anterior gonapophysis Navas, by original designation ( L. modestus found in this genus is remarkable and points to a (Blanchard) 1851). possible relationship with Dendtoleontini. An -Licura Navas, 1918. BoI. soc. aragonese 17:330. New autapomorphy is the sexual dimorphism in the synonym. Type species: Licura {erus Navas, by length of tibial spurs. I have been able only to original designation ( L. modestus (Blanchard) identify by name one species but several other 1851). species are present.

AdultDescription: Antennalfossae separatedby 22. Neulatus Navas about width of pedicel; distal paJpomere o£labium (Figs. 27, 64 65) slender; profemur without white bristles on exte nor face except subapIcally; profemur sense hair =Lincoya Navas, 1911. Rev. chilena Hist. nat. 15:121. moderately long, about one half length of femur, Type species: Lincoya dealbatus Navas by origi- about equal to that of mesofemur; pretarsal claws nal designation. Preoccupied by Lincoya Kirby, hmger than pro-hasitarsus, shorter or longer than 1871 (Lepidoptera) tibial spurs; tibial spurs usually shorter in males; fOrewmg costal area simple or biareolate; forewing Neulatus Navas, 1912b. Arxivs lust. Cieucies 1.108. radialsector arises wellbasadofcubitalfork, about Type species: Glyptobasis porteri Brethes. by origi- equivalent to the distance between radial sector nal designation. and medialfOlk in hindwing; hindwingaboutsame Nikoya Navas, 1913. Rev. chilena !list. nat. 17.243. length as forewing, in repose hindwing extends New Synonym. New name for Lincoya Navas beyond fOrewing; anteriOr banksian line of fOrew- ing weakly developed or absent; male sternite IX AdultDescription: Antennalfossae separatedby much wider than long, weakly produced medially; more than width of pedicel; distal palpomere of male eetoproct simple, without lobes or plocesses; labium slender; profemnrwithout white bristles on male genitalia with arched gonarcus, mediuncus exterior face; profemoral hair short, less than one- present or absent, not ridged; female ferminalia fourth length offemur, equal to that ofmesofemur; with sternite VII not produced; tergite IX not pro- retarsalclawsshorterthan meta-basitarsus about , thanlong, with median tooth; anteriorgonapophysis lobe-like, posterior gonapophysis swollen, butrela- Vol. 8, No.1 - 2, March - June, 1994 99 forewing radial sector arises well basad of cubital !'kteta Navas, 1934. Rev. aBilena Hist. Nat. 38:10. fork, about equivalent to the distance between New synonym. Type species: Neteta barrosi radial sector andmedialfork in hindwing, anterior Navas, by original designation. develope:~~i~cl~inga~lo~g banksian line weakly Chiloleon Navas, 1934 Rev chilena Hist Nat 38'10. asfOrewmg, m repose apex 0 m wmg ex en s New synonym. Type species: ChiZoleon stuardinus beyond that of forewing; male sternite IX trans Navas, by orlgmal deSIgnation verse; male ectoproct simple, without lobes or pro- cesses, malegenitalia with simple archedgonarcns, Adult Description: Aritennalfossae separatedby mediuncus not developed, paramere short, plate- about width ofpedicel; distal palpomere oflabium slender; profemur without white bristles on exte­ Iior face except subapically; profemur sense hair moderately long, about one-half length of femur, ahout equal to that of mesofemur; pretarsal claws len; ectoproct without digging setae; lateral longer than pro basitarsus, shorter or longer than tibial spurs; tibial spurs usually shorter in males; forewing costal area simple; {mewing radial sector ariseswellbasadofcubitalfork, aboutequivalentto the distance hetween radial sector and medial fork in hindwing; hindwing aboutsame length asforew- LanaI Description (based on ported): Head ing, in repose hindwing extends beyond forewing; I n er than wide with dolichasters on dorsal sur- . .. . oped; male sternite IX much wider than long, weakly produced medially; male eetopnx:t simple, t an WI e, wIt60ut 0 c asters; a omen WIt without lobes or processes; male genitalia with "veak row of clumped dolichasters; pretarsal claw arched gonarchus, mediuncns present; female with basal tooth; mesothoracic spiracle borne on terminalia with sternite VII not produced; tergite tubetcle that is lunge! than wide, abdominal spi- IX not produced ventrally; pregenital plate sclero­ racles small, clark, but somewhat elevated tized, withontmediantooth; anterior gonapophvsis lobe-like; posterior gonapophysis swollen, butrela­ Included species: Only porteri (Brethes) ( tively short; ectoproet and lateral gonapophysis dealbatus (Navas). without digging setae.

Distribution: Central Chile LarvalDescription(basedonperalinus). Head capsule wider than long with many dolichasters and fme white hair; mandible nearly uniform dark brown; first 3 pairs of thoracic scoli longer than wide; abdominal scOli present but WIder than long; in the tribe. Also, the median processofsterniteVII all scoli with fine, white hair; abdomen with row of is unique The very pale larvae live in Puya plants tufted black setae; pretarsal claw without basal tooth; mesothmacic spiracle borne on tubercle that is about as long as wide; abdominal spiracles en­ larvae move fairly fast and have elongate abdomi larged, horne on tubercles nal scoli (Brethes 1908:15, larva). The type species is based on a larva which was misidentified as an Included species: barrosi (Navas), peralinus ascalaphid. The pale adults rest on Puya plants Navas, and stuardinus (Navas). durmg the day and are attracted to lights. Distribution: Central Chjle 23. Sical Navas (Fig. 67) Discussion: This genus is sustainedmostly by the larval stage which has moderately developed ab­ Sical Navas, 1928. Rev. chilena Risto Nat. 31.311. Type dominalscoli incontrastto ElicumandLemolemus. species: Sical peralinus Navas, by original desig- The female terminalia are most similar to nation. 100 Insecta Mundi

Lemolemus but the pregenital plate lacks the me­ Banks,Nathan. 1913b. The genusBrachynemurus dian tooth. The larva lives in soil andcarries debris (Neuroptera). Entomological News 24:63-65. held by long white setae and scoli. At least one additional species is found in this genus Banks, Nathan 1924 Descriptions of new Autapomorphles for thIS genus are the long, whIte neuropterOldinsects. BuIIetm Museumof Com­ setae on the scoli and the lack of a median tooth on parative Zoology 65:421 455. the female pregenital plate. AcknO\vledgements The author is grateful to Robert Bruce Miller para Ive 00 ogy 6 : - . fur the photographs of the larvae and for many constructive ideas. John Oswald made a vigorous Banks, Nathan. 1939. On some new and previ­ critique ofthe manuscript. Although the selection ously known Neuropterain the collection ofthe of the characters used in the cladistic anal!sis was AcademTof Natural Sciences of Philadelphia. done solely by the author, special thanks are due to Notulae Ifaturae 32.1-5. DavidWahl who spent manyhours analyzingdata Thanks are also due to the typists of the manu­ Banks, Nathan. 1943. Neuroptera of northern scriPt, MiChelle M. F'aniOla andBnmda J. Lovelace. SouthAmerica. Part2. Myrmeleonidae. Boletin The illustrations are original except for Figures 9- de entomologia venezolana 2:161-173. 1.9, 22, 30-32, 36, 38-46, and 50-59 which are taken frOID Stange (1970), 23from Stange, (1969) andFig. 8 (Adams, 1957). entomologia 12: 15 17, pI. II.

Carpenter, J. M. 1988. Choosing among multiple Aams, P lip A. 1957. New antlions from the equally parsimonious cladograms. Cladistics southwestern United States. Psyche 63:82 4:291-296. 108. Currie, Rollo. 1901. Adwarfant-lion fly. Proceed­ ings EntomologicalSoeietyofWashingron 4:435 437. evolution in the Osmyloidea. Ph. D. Thesis, Harvard University, 105 pp. , 79 Figs. (unpub- Esben-Petersen, Peter. 1928 Neue und wenig lished). bekannte Neuropteren des Hamburger muse­ ums. Deutsche entomologische Zeitschrift Banks,Nathan. 1899. AclassIDcatlon oltheNorth (1928):73-77. American Myrmeleonidae. Canadian Ento mologist 31:67-71. Farris, J. S. 1969. A successive approximations approach to character weighting Systematic Banks, Nathan. 1901a. A new genus of Myrme- Zoology 18:374-385. leonidae. Canadian Entomologist 32:329-330. Farris, J. s. 1989. Hennig86 referenee: doeumen Banks, Nathan. 19016. A list of neuropterOld tation for version 1. 5. Port Jefferson Station, insectsfrom Mexieo. TransactionsoftheAmen New ylJrk can Entomological Society 27:361-371. Fraser, F. C. 1952. New speCIes of Neuroptera ill Banks, Nathan. ]909 New genera and species of the Museum National d'Histoire naturelle, tropIcal MyrmeleoOldae. Journal of the New Paris. Revue Franl;aise d'Entomologie 19:55­ York Entomological Socwty 17: 1 4. 64.

Banks, Nathan. 1913a. Synopses and deserip- Hagen, Hermann 1888 Stray notes on tions ofexotic Neuroptera. Transactions ofthe Myrmeleonidae. Part 4. Canadian Entomolo- American Entomological Society 39:201-242. gist 20:34-38. Vol. 8, No.1· 2, March· June, 1994 101

Henry, Charles S. 1978. An unusual ascalaphid Navas, Longinos. 1919. Algunos insectos larva (Neuroptera' Ascalaphidae) from south· neuropteros de Republica Argentina Revista ern Mrica, with comments on larval evolution de la realAcademiade ClenCIaS exactas, FiSlcas within the Myrmeleontoidea Psyche 85'265· y natnrales de Madrid 17'288·305 2'14. Navas, Longinos. 1920. Sur des Nevropteres MacLeod, Ellis G. 1970. The Neuroptera of the nouveauxou critiques (Serie 2). AnnalesSociete Baltic amber. I. Ascalaphidae, Nymphidae, Scientifique de Bruxelles 39.189·203. and Psychopsidae Psyche 77' 147·}80 Navas, Longmos. 1922. Insectos Sudamerlcanos Miller, RobertB. 1990. Reproductive characteris (Series 4). Revista de la real Academia de tics of some western hemisphere ant· ciencias exactas, Fisicas y naturales de Madrid (Insecta. NemopteIa. Myrmeleontidae). PIO' 19.255·267. ceedings ofthe Third international symposium on Neuropterology (Berg en Dat Kruger Na· Navas, Longinos. 1924. Insectos de la Amenca tional Park), pp 17}.}79 central Broteria (Serie Zool6gica) 21'55·86

Navas, Longinos. 1911. Neuropteroschilenos (Se Navas, Longinos. 1926a. Inseetanova (SeriesXI). ries 2). Revista chilena de Historia Natural Memorie dell'Accademia pontifica dei Nuovi 15.121·128. Lincei, Rome 9.101·110.

Navas, Longlnos. 1912a. Notas sobre Navas, Longinos. 1926b. Algunos msectos del Mirmeleonidos. Broteria (Serie Zool6gica) 10:29 Brasil (SentS 3). Broteria (Serie Zoo16giea) 97. 23:5·15.

Navas, Longinos. 1912b. Sinopsis de los Navas, Longinos. 1927. Insectanova (Series XII). Ascalafidos. Arxius de l'Institut de Ciencies, Memone dell'Accadem18 pontiflca del NUOVI Barcelona 1(3)'45·143, 11 Figs, 2 pis Lincei, Rome 10: 1·10.

Navas, Longinos. 1913. Mirmeleonido e Chile. Navas, Longinos. 1928. Insectos neotIOpieos (Se· RevistachilenadeHistoria natural 17:243·244. ries 3). Revista chilena de historia natural 31.316·325 Navas, Longinos. 1914a Neuropteros sudamencanos (Sene 1). Brotena (Sene Navas, Longmos. 1932. Alcum msettI del museo Zool6gica) 12:45 56. di zoologia della R. Universita di Torino. Bollettino dei Musei di Zoologia e di Anatomia Navas, Longinos. 1914b. New NeUIoptera flom Compamtadella R. UniversitadiTorino (3)42: 1· the United States. Bulletin of the Brooklyn 38. Entomological Society 9"13·211 Navas, Longinos. 1934a. Algunosinsectosde Chile Navas, Longinos. 1915a. Neuropteros nuevos 0 (6a Serie). Revista chilena de historia natural poeo eonoeidos. Memoria della R. Academia 38.9·12. Ciencias y Artes Barcelona (3) 11:373·398, 455· 480 Navas,I.onginos 1934b Tnsectos suramericanos (8 Serie). Revista de la real Academia de Navas, Longinos. 1915b. Neuropteros Clenciasexactas, Fisicasy naturalesde Madrid Sudamerieanos (Serie 2). Broteria (Serie 31:9·28. Zool6gica) 13:5·13. New, T. R. 1~R1 A revision of the Australian Navas,Longinos. 1918. Insectoschilenos. Boletin Nymphidae (Insecta: Neuroptera). Australian de la Sociedadaragonesade Ciencias Naturales Journal Of ZoolOgy 29:707 750, Figs. 1 188. 17:212·230. 102 Insecta Mundi

New, T. R. 1982. A leappraisalof the status of the on Neuloptelology. (Belg en Dal, 1(1 ugel Na- Stilbopterygidae (Neuroptera' tional Park), pp 151-169 MyrmeleontOldea). Journal of the Austrahan Entomological Society 21:71 75. Tillyard, R. J. 1915. Studies in Australian Neuroptera. No. 1. The wing-venation of the Nixon, K. C. 1992. Clados l'efel'enee: doeumenta MYl'meleonidae. Proceedings of the Linnean tion for version 1. 1. Trumansburg, New York. Society of New South Wales 40(4):734·751.

Oswald, J o. 1993 Phylogeny, taxonomy, and Tjeder, Ro. 1977 Distal abdominal segments and bIOgeographyofextantsilkyIacewmgs (Insecta: scIerotIzed parts of gemtaha m AScalaphIdae Neuoptera: Psychopsidae). Memoirs of the (Neuroptera).•A..nnales Entomologici Fennici American Entomological Society 40: 1·65. 43:61-65.

Stange, Lionel A. 1969 Myrmeleontidae of the Withycombe, C. L. 1925 Some aspects of the Galapagos Islands. Acta zoologIca Lilloana bIology and morphology of the Neuroptera, 25: 187·198. with special reference to the immature stages and their possible phylogenetic significance. Stange, Lionel A. 1970. Revision of the ant lion Transaetions of the Entomologieal 80eiety of tribe Brachynemurini ofNorth America. Uni­ London (1924):303-411. versity of California Publications in Entomol- ogy 55'1.192

Stange, L. A. , & R. B. Miller 1990. ClassifICation of the Myrmeleontidae based on larvae. Pro­ ceedings ofthe thitd Intelnational SymposiuIll Vol. 8, No.1· 2, March· June, 1994 103

12 n

3 4 1221

1 1 2 1 ~. .lVemoptenaae ~ ~ ~ 1.1 I ~ •••• A 0 U,,? ~ ~ n irn·"-

6 1315 210112024

1 1 1 2 1 1 1 1

'~alpanm : ~ ~ 1.62! 2.32.528 I92;; 0 n 32224 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ n n n ... "" 121426 2 12 ., 2 1 1 2 1

'Ulmares 1.6 ~ ~ 1" ~ ~2! ~ ~ v 10 1122 .. "~ 1 1 1

.; ~ ~ Acanthaclisini 4 '0 ~ " ~ ~ ~ . . . n -n I ~ ~ I 1428 .. • n • u o '1 v '4

U Nemoleontini 0 1921 ji n n ~ w : 1 2 928 .. 1 1 16 9 28 --t r-lH}- Dendroleontini n n 2 O '-11 n . i

Figure 1. One of' 2 cladograms showmg group relatIOnships within the Myrmeleontoidea based on adult and larval characters Black bar apomorphy; shaded bar parallelism; white bar reversal. This cladogram and others were made with the Clados program by Nixon (1992). 104 Insecta Mundi

1:2 (] : :~-~ Nymphidae

28 - " NemoDteridae

,~ ,~ 2 ~ tI • El ~ 1 2 1 2 ~ .~,~ ~

Paloarini ~ ~ ; '.bL';L.,U:L:; '::L~ I tI B ~ oooA .... If-U IJ , 2 1 1

': .-. n ." Stilbopteryginae u •• tI U L '" 2! Dimares 1 }.L +l -~ ~ tI U IVlill,"II<:UI 1 0

.~ --n Myrmeleontini + tI U 14 1m 0 :! ~

1 1 tr-UUU ~cantnaclISlm .- 2 1 1 0 Ii-. Myrmecaelurini 172428 1 1 0< • n n -:: ~ ~ g "Y 1428 u ~ 0 1 rz

tI ~ Brachynemurim 1 2 a ~J> • n ~ ~ Lemolemini '1. 928 .. J "tI U L1J o 0 -= ~ Gnopholeontini

Figure 2 One of 2 cladograms showing group relationships within the Myrmeleontoidea based on adult and larval characters, Vol. 8, No.1 - 2, March - June, 1994 105

Ie 12 lel2 - ~ Nemo/eontini . " Nemo/eontini

:! 2,,9 .. ~ 2; .. v v !t 3 4 6 830 1: 3 4 6 830 " u uno o eon 1 1 , 1 1 1 1~2~2.6 ' 2 1.52;2.6 ' 2 3 ... 3 £ ~ I 1 7 I '1 --., T rl- Tyttho/eon r-+. 'I Tyttho/eon - 1 Elicura Elicura

,~~, LAma/emus ,~~ I

YfT 19 lft '0 ~ ~'v ' _""0 30 30 '1 . JaTTUeila 1 1~2.83j ~ 1 182831 rr-t""JaffUelia 9 I 1 '~ -.. 1 1 ,. , ~ " .. " .. '0 I Neu/atus LIJ-- Neu/atus 1 ---r ~ ~ 2~ 6 825 ~ n • " ~ " " . 3 4 19 3 4 r-11- alOleo 1 , 1 1 1 20 6 822 20. 6 822

'~ .. 't ..'t 't i '1 ., '1 i .- n Ensorra Ensorra Lf!- ~ 27 ~ ~ ~ 1.4 ~ ~ 1.4 Ameromyia i Ameromyia , 'An?A ".. 1. 'O??? " " . i,i i ~ il~iil~ Vpnpzue/enn

" .? '" 1117 3 4 1117 r -.:tJ'1~'1 r r4-l i '101 , f .MexOIeon .lVIexoleon 1923 23 -r 5 r.q, --r- ,; r,q:):) o 1 A tricho/eon 1 Atricho/eon 1 '1 3 '1 '1 .3 YJ- L+- 2 1 '11 ! '11'.2

Figure 3. One of 2 weighted c1adograms of the genera F'lgure 4. One of 2 weighted cladograms of'the genera formerly included in the Brachynemurini, based on formerly included in the Brachynemurini, based on adult and larval characters. adult and larvae characters. 106 Insecta Mundi

Nemo/eontini Nemo/eontini

Dendro/eontini Abato/eon

Maracandu/a Argento/eon -

. ~n~M'~ ---l

Icura

'LemOlemus .Ameromy,a

Sica/ venezueteon

Ecua/eon f-- Chaeto/eon - Ga/apago/eon Mexo/eon

'--- Jaffue/ia Atricho/eon I - Neu/atus ---, Brachvnemurus

n .. . .

ct:.

vOuucuo

Ensorra Lemo/emus

C/athroneuria Neu/atus

Ameromyia Sica/

~~,,~/~nn

- ri, lVIexoeon

'--- Atricho/eon unopllo'f'UII ~ Brachynemurus U . lVIenKe eon ~ Scoto/eon '----- Tyttho/eon

F'lgure 5. StrIct consensus tree of 341 cladograms ofthe Figure 6. Strict consensus tree of 353 trees showing genera of the Brachynemurini & Myrmecaelurini, relationships ofthe gemlra formerly included in the based on adult and larval characters, Brachynemurini, based on adult characters. Vol. 8, No.1· 2, March· June, 1994 107

7 - Chaetoleon pusillus (Currie)

8 - Scotoleon deflexus (Adams)

Figure 7 8 7) Adult habitus drawing of Chaetoleon pusillus (Currie) (male). 8) Adult habitus dl'u.....ing of &otokon deflexus (Adams) (male). 108 Insecta Mundi

mavU. Seeae

... ~ " CJ.08 l'ac8

..-- - Tibia1 Spurs

real Claws

9 Hypost;gmatic Ce"

PiiulQ Allilloris

15 • Gnopholeon delicatulus 2 2 2 !! 7

rus divisus

14· Scotoleon nigrilabris

Figures 9-16. 9) Proleg; 10·11) Base of forewing; 12-14) Base of hindwing; 15) Wings; 16) Ventral apex oflarva Vol. 8, No.1· 2, March· June, 1994 109

lateral gonapophyslS

9 osterior

targite

pa

a eflor gonapophysis

17 - Bracnynemurus sp. 18 - Scotoleon .Is.ni

19 - Abatoleon dorsalis 20 - Ensorra vertlcalls

21 - Venezueleon guarlcus 22 - Dejuna persimllis

Figures 17-22. Ventral view of female terminalia. 110 Insecta Mundi

24 - Ellcura litigator 23 - Galapagoleon darwlnl

26 - Lemolemus nectator 25 Jatfllelia GhileRSe

27 • Neulatus porterl

Figures 23-27. Ventral view of female terminalia. Vol. 8, No.1· 2, March· June, 1994 111

\ ill \\\! 1/ ////1// ------\-\\ \~ I \ /

\\1\\1 // 7 \~\ J /)

\.....,. >;;t I ,,\1//11/1 IA/

28 Nannoleon michaelseni /

\ I '<\\ if

\ II' 'I 1// III)}, f.I I, N r .' '~ 30 - Tyttholeon puerilis

32 • Gnopholeon delicatulus 31 - Maracalldula apicalis

Figures 28-32. Ventral view of female terminalia. 112 Insecta Mundi

33 - Ecualeon ovisparg'ls 34 - Ecualeon ovispargus

36 - Maraeandula apicalis

35 - Ecualeon ovispargus ~~~~

38 • Mexoleon papago

39 - Gnopholeon

37 - Ameromyia nigriventris

Figures 33-40 33) Lateral view offemale terminalia; 34) Ventral view offumale terminalia; 35) Ventral vi€l'" ofmal€l terminalia and genitalia; 36-37) Frontal view of head; 38-40) Hind tarsus. Vol. 8, No. 1 - 2, March· June, 1994 113

/ / / ,\ \i:;; 0/ '-.: \ ~ It'll \ ,!~0~ \;~/ -~ V ~\ \ \ ,\ M .~ 7/; n r~(f: ~~(~

, ~\ ., '-J~r~~,\ '/.J,--:::- \ ,'\ '7 X"; /:/, \{I~ ~\~

~~ 1"( {\. 'f,<:o ~\i'T \ • ~.J/ ~( ~ ~\ ~~ " "1~,/\-- ': \(q-- '~tr ';: J( ~\~~YJ::: ;C•• (/# ':1\,'..\' IIiff ,lor- ~ "\~\l ,~ AJfJ!E \J:, '/,/ ~.JY/ ~,J'~ ~~/!I.~ :t~ -;~f \Ut. "- lif! ' J,",:oJ , ~ 'Ir ,"F '\: 'I": 3:'!1'/ ~1'l ;S \l / v 43 - Clathroneuria schwarzi 41 - Atricholeon tuberculatus 42 - Brachynemurus seminolae

44 - Atricholeon tuberculatus 45 - Dejuna persimilis 46 - Clathroneurla schwarzi

48 'Jenezueleon guarieus

49 - Ensorra verticalis

Figures 41 50. 41 43) Ventral view of male ectoproct; 44 46) Ventral view ofmale genitalia; 47) Lateral view ofmale terminalia; 48) Male genitalia; 49) Lateral view of male terminalia; 50) Posterior view of male ectoprocts. 114 Insecta Mundi

,i/ /

S1 Mexoleon papago

53 • Chaetaleon pusilllls 54 • Chaetoleon pus/llus

56 • Brachynemurus pulchellus

59 • Maraeandllia apiealis 57 •IYttholeon puerills 58 • Abatoleon dorsalis

Figures 51-59.51,53,55) Lateral view ofmale terminalia. 52, 54, 56-59) Posterior view of male genitalia. Vol. 8, No.1· 2, March· June, 1994 115

60 • Scotoleon fidelitas 61 • Scotoleon fidelitas

62 • Scotoleon pallidus 63 • Scotoleon eiseni

Figures 60-63. Third instar larvae ofBrachynemurini. 116 Insecta Mundi

64 - Neulatus porteri 65 - Neulatus porteri

)

67 - SicaI peralinus

66 - Jaffuelia chilense

Figures 64-67. Third inst.ar larvae of Lemolemini. Vol. 8, No.1· 2, March - June, 1994 117

68 - Gnopholeon barberi 69 - Gnopholeon barberi

70 - Menkeleon bellula 71 - Tyttholeon puerilis

Figures 68-71. Third instar larvae ofGnopholeontini. 118 Insecta Mundi

APPENDIX 1. Data Mattix fOI Tlibal Relationships

Character I~umber I I I I I IIIII 2 2 2 2 2 2 2 2 2 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 456 7 8

Psychopsidae o 0 0 0 0 0 000 o 0 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 0 0 Nymphidae (l (l I 1 (l (l (l (l (l (l (l 2 (l (l (l (I (l (I (I (I I (I (I (I (I (I (I (I Nemopterinae 1 o 0 0 2 1 0 1 0 o 2 1 1 0 1 o 0 0 0 0000000 0 1 Crocinae 1 o 0 1 2 1 0 1 2 1 2 2 1 0 1 o 0 0 0 00000 0 0 0 0 Ascalaphidae o 2 1 0 1 1 000 1 1 1 1 o 1 1 o 0 ? 1 1 0 1 1 1 o 0 1 Stilbopterygilli o 2 1 0 1 1 o 0 0 o 0 1 1 1 1 :1 000 0 1 (l 1 (l 1 1 (l 1 Palparini 0 1 1 0 1 1 000 002 1 1 1 1 o 0 1 0 1 0 1 0 1 1 1 1 Palparidiini Q 1 2 0 1 1 000 002 1 1 1 1 o 0 1 0 1 1 1 1 1 1 1 1 Dimares (I I 2 (I II (I (I (I I (I 2 1 1 1 2 0 0 0 0 1 0 1 0 1 1 1 1 Millerleon 0 1 2 0 1 1 000 1 o 2 1 1 1 2 0 0 0 0 1 1 1 0 1 1 o 0 DendroleoRtiRi 0 1 1 1 1 1 1 o 0 1 o 2 1 1 1 3 2 0 0 0 1 0 1 1 1 1 o 0 Nemoleonlini 0 1 1 1 1 1 1 o 0 1 o 2 1 0 1 2 1 o 0 0 1 0 1 1 1 1 o 0 BlachyllelllUlini (l I 1 I 1 I I (l II (l 2 I II 2 I (l I o 2 0 II II 0 I Gnopholeontini 0 1 1 1 1 1 1 0 1 1 o 2 1 1 1 3 2 0 0 1 1 0 1 1 1 1 0 1 Lemolemini 0 1 1 1 1 1 1 0 1 1 o 2 1 1 1 3 1 o 0 0 1 0 1 1 1 1 0 1 Myrmecaelurini 0 1 1 1 1 1 1 o 0 1 1 2 1 1 1 2 1 1 0 020 1 1 1 1 o 0 Nesoleolllilli 0 1 1 1 1 1 1 (l (l 1 1 2 1 (l I 2 1 I (I (12(1 III I 0 I Myrmeleontini 0 1 1 1 1 1 1 o 0 1 1 2 1 1 1 2 0 0 0 0 1 1 1 0 1 1 0 1 Acanlbaclisini 0 1 2 1 1 1 1 o 0 1 1 2 1 1 1 2 1 1 1 0 1 1 1 0 1 1 o 0

Maulini 0 2 1 1 0 1 1 1 2 1 1 3 0 0 0 'I '1'1'1 'I 'I 'I 'I 'I Pseudimarini 0 1 1 o 0 0 o 0 2 0 1 1 000 ?????????

APPENDIX 2: Data Matrix for Brachynemurini, Lemolemini and Gnopholeontini

Character Number 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1

Nemoleontini 0 0 0 0 0 0 o 0 0 1 0 1 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 0 0 0 0 Dendroleontini 0 0 0 0 o 0 0 0 0 0 0 o 0 o 0 1 0 1 0 0 o 0 0 0 0 0 0 0 1 0 0 BrachyllelllUlilli Abaloleon 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 o 0 0 0 0 0 0 0 0 0 Ameromyia 1 0 0 0 0 1 0 1 0 1 0 0 2 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 Argentoleon 0 0 0 0 0 1 0 1 1 0 0 0 0 o 0 0 0 0 1 1 0 2 o 0 0 0 1 0 0 0 0 Alrieholeon 0 0 0 0 1 0 o 0 1 0 1 0 0 o 0 o 0 0 1 1 0 3 1 0 0 0 1 (l (l (l (l Austroleon 0 0 0 0 0 1 0 1 1 0 0 0 0 o 0 o 0 0 1 0 0 0 o 0 1 0 1 0 0 0 0 Brachynemuws () 1 () () () () () () 1 () 1 () 0 o 0 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 0 Chaetoleon 0 0 1 1 0 0 0 0 1 0 0 0 0 o 0 0 1 0 0 0 1 0 1 0 0 0 1 0 0 0 0 Ensorra 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 0 (l (l (l (l 0 1 (l (l (l (l Mexoleon 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 o 0 0 1 0 1 0 0 0 1 0 0 0 0 Scololeon 0 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 1 o 0 1 0 0 0 1 0 0 0 0 Vellezueleoll 1 (l (l (l 1 (l (l (l I (l 1 (l (l I (l (l (l (l I (l 0 I 0 I 0 0 I 0 0 0 0 Gnepholeonlini Gnopholeon 0 0 1 2 0 1 0 1 0 0 o 0 0 1 0 1 0 0 0 0 0 0 2 1 0 0 0 1 0 Menkeleon 0 0 1 0 0 0 1 0 1 0 0 0 0 0 1 0 1 0 0 o 0 0 0 2 1 0 0 0 0 0 Tylllioleoli 0 (l (l (l (l (l 1 (l 1 (l (l (l (l (l I (l I 0 (l 0 0 o (l 2 I 0 0 0 0 0 Lemolemini Flicllra 0 0 0 0 0 0 0 0 1 0 0 0 1 o 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Jaffuelia 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 1 0 1 1 Lemolemus 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 (l (l (l (l 0 (l 0 0 0 0 0 2 Neulalus 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 1 1 0 o 0 o 0 0 0 0 1 0 1 1 S,cal () () () () () 0 0 0 1 0 0 0 1 o 0 1 0 0 1 0 0 0 o 0 0 0 0 0 0 1 2 Unknown Larvae: Clathroneuria 0 0 0 0 0 1 0 0 1 0 0 (I (I (I (I (l (I 1 (l 1 0 0 o (l 0 ( ( ( ( ( 'I Dejuna 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 ? ? ? ? ? ? Ecualeon 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 0 0 0 o 0 0 7 7 7 7 7 7 Galapagoleon (l 0 0 0 0 0 0 0 I 0 0 0 1 0 0 1 0 0 1 0 0 0 o 0 0 III '! ! Maracandula 0 0 0 0 0 0 1 0 1 0 0 0 0 2 1 1 0 0 0 0 o 0 0 0 0 ? 7 ?? 7 7 Vol. 8, No.1 - 2, March - June, 1994 119

APPENDIX 3. Successive Approximation JvVeights APPENDIX 5. Successive Approximation \Veights for Tribal Analysis for New World Brachynemurini. Generic Analysis wIth genera added with missing larval data. WEIGHT CHARACTER:# WEIGHT CHARACTER # ° 12,20,22,28 1 3,19,24 a 3,4.5,6,8,12,17 2 4, 10, 14, 21, 27 1 14, 19 3 9, 11 2 18 5 2 3 30 10 1,5-8, 13, 15-18,23,25,26 4 11, 13, 20 5 25 APPENDIX 4. Successive Approximation Weights 10 1, 2, 7, 9, 10, 15, 16, 21-24, 26-29, 31 for New World Brachynemurini. Generic Analysis with 5 genera with missing larval data deleted. APPENDIX 6. Successive Approximation Weights for New World Brachynemnrini Generic Analysis WEIGHT CHARACTER # of all genera but only adult characters. o 3,4,5,6,8,9,12,17 WEIGHT CHARACTER # 1 .19 3 14.30 4 11, 13, 18, 20 o 3-6, 8, 9, 12, 17 5 25 1 14, 19 10 1, 2, 7,10, 15, 16,21·24,26-29,31 4 11, 13,20 5 25 10 1, 2, 7, 15, 16, 18, 21·24

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