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March 1994

Reclassification of the New orldW antlion genera formerly included in the tribe Brachynemurini (Neuroptera: Myrmeleontidae

Lionel A. Stange Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, FL

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Stange, Lionel A., "Reclassification of the New orldW antlion genera formerly included in the tribe Brachynemurini (Neuroptera: Myrmeleontidae" (1994). Insecta Mundi. 295. https://digitalcommons.unl.edu/insectamundi/295

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Vol. 8, No. 1 - 2, March - June, 1994 67

Reclassification of the New World antlion genera formerly included in the tribe Brachynemurini (Neuroptera: Myrmeleontidae)

Lionel A. Stange Florida State Collection of Arthropods Department of Agriculture & Consumer Services P.O. Box 147100 Gainesville, FL, 32614-7100 U.S.A.

Abstract A cladistic analysis of the New World tribe Brachynemurini has resulted in several 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, Mmleon and Venezueleon are newly described. The Gnopholeontini (NEW TRIBE) includes 4 North American genera whereas the Lemolemini (NEW TRIBE) contains 6 South American genera of which Ecualwn and Galapagolwn are newly described. Descriptions of genera in the 3 tribes, based on adults and known 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) postventral lobe ofmale Since the revision of the North American ectoproct; 2) a narrow gonapophyseal plate; and 3) Brachynemurini by Stange (1970), additional adult a short distal tooth of the larval mandible, which is and larval characters (Stange & Miller, 1990) have not parallel with other teeth. Since larval traits are been found which have led to the present reclassi- here considered to be very important for the infer- fication of New World brachynemurine genera into ence of phylogenetic relationships, two analyses three tribes. Twenty-three genera are hereby rec- were made: one of all genera and the other using ognized. Representative larvae of nineteen of these 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 groups of genera formerly placed in Myrmeleontoidea the Brachynemurini are given tribal status Relationships among higher taxa within the (Gnopholeontini and Lemolemini); the remaining Myrmeleontoidea have notpreviously been investi- genera are maintained as the Brachynernurini. gated cladistically. MacLeod (1970) 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 apomorphic larval head the close approximation of the larval mandibles. characters unite the Psychopsidae, Nymphidae, The Lemolemini are most diverse in Chile; they are Nemopteridae, Ascalaphidae and Myrmeleontidae defined by the autapomorphies of the development into a group. Manse11 (1992) has provided the most of specialized setae on the median area of the larval recent discussion of the relationships within the abdominal tergites and the lack of digging setae on Myrmeleontoidea indicating apomorphies for the the female ectoproct. Both the Gnopholeontini and various families and providing a phylogenetic tree. Lemolemini have highly modified posterior Oswald (1993) has completed a cladistic analysis of gonopophyses of the female terminalia. The more the Psychopsidae, a family hypothesized as the restricted Brachynemurini are defined by the fol- sister-group of the rest of the Myrmeleontoidea Insecta Mundi

(Withycombe, 1925; Henry, 1978). The following Lachlathetes) but these exceptions are considered analysis of the tribes of the Myrmeleontidae uses secondary modifications. these four families as outgroups to help determine outgroups for the generic analysis. 3. Labial palp, sensory area - (0)palpimacula present but without pit; (1) oval-shaped pit; (2) elongate, The Psychopsidae (26 extant species in 5 gen- slit-shaped pit era, found in Africa, Australia and the Orient) and Comments: The apomorphic condition (2) is found in the Nymphidae (about 25 species in €$genera,restricted Acanthaclisini, Dimarini, Palparidiini and some to Australia and New Guinea) are small relict genera of the Palparini. Whether the latter tribe families. The Nemopteridae and Ascalaphidae 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- subfamilies of Nemopteridae, both subfamilies were ence. The outgroups have small palpimacula and the evolution of enlarged palpimacula seems to be entered independently for a clearer resolution of found only in the Myrmeleontidae. The the relationships. The Ascalaphidae appears to be Nemopteridae may have lost the pit-like palpimacula even more diverse with at least 3 subfamilies and when they adapted to pollen feeding. many tribes. However, the classification of the Ascalaphidae is so poorly established that the char- (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 Mark1 .(1956) with Comments: Mansell (1992b) considers without much modifications indicated by Stange & Miller (1985; justification the broad pronotum as an autapomorphy of the Palparinae. It is treated here as plesiomorphic 1990). Since larvae are known for all the major 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 of the 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 the Pseudimarini (1 rare species) and Maulini (about change the cladogram. 5 species in 2 genera). Although the larvae in general reinforce the tribal classification, one prob- 5. Pronotal articulation (taken from Adams, 1958); (0) lem in using larvae was recognized. The larvae of pronotum articulates with mesothorax the Dimarini are highly diverse (Stange, 1989) so (Psychopsidae; Nymphidae); (1) pronotum articu- that 2 of the 3 constituent genera were entered lates with mesothoracic spiracle; (2) pronotum ar- ticulates with greatly enlarged mesothoracic spi- rather than the tribe as a whole. Preliminary racle analysis led to the belief that three distinct groups were present in the Brachynemurini. To attempt to 6. Pretarsal arolium - (0) present; (1) absent substantiate a separate treatment of these 3 groups, Comments: The presence of an arolium is considered the they were entered independently in the analysis. plesiomorphic condition in the Myrmeleontoidea. The following informative characters were used. There is an evident arolium in Psychopsidae whereas The data matrix is given in Appendix 1. in the Nymphidae the arolium is deeply bifid, form- ing two pulvilliform lobes. In the Ascalaphidae and Myrmeleontidae the auxiliae are fused forming a List of myrmeleontid tribal apomorphies bristly median lobe co-adapted to a strong (HEAD) unguitractor plate which strengthens the flexing 1. Mouthparts - (0) short mouthparts; (1) head pro- mechanism of the 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 mesoleg (Fig. 9) 2. Antenna1 shape - (0) filiform; (1) clavate (Fig. 36); Comments: The femoral sense hair is an autapomorphy (2) Clubbed of the subfamily Myrmeleontinae. In a few genera Comments: The shape of the antennae is fairly consis- (i.e., Atricholeon, 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, Acanthaclisini 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 to forewing; (I) 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 - (0) at or before basal crossvein; (I) distad of M-Cu or CUPobsolete; (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 Gnopholeontini, Lemolemini and Brachynemurini. autapomorphy of the family. The absence of the Elsewhere, the apomorphic condition has been ob- structure in several groups (especially Nemoleontini) sewed only in the South African genus Nannoleon is considered a reversal. and Maulini and in some Dendroleontini.

10. Forewing, condition of vein 1A - (0) Free course from (ABDOMEN) base to hind margin (see Mark1 1954: Fig. 62); (1) 15. First abdominal spiracle (data taken from Adams, united with CUP near base (Figs. 10-11) 1958) - (0) connected to mesepimeron; (1) 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. Hindwing, origin of radial sector - (0) near wing the diminution of the first tergite. Probably this base, well before origin of medial fork; (I) originates relocation of the spiracle improves the air supply to distal to medial fork (Fig. 15); (2) greatly reduced the flight muscles. Comments: This is a progressive character where the origin of the radial sector 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 crossveins 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 of the 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, 1970) here. The posterior gonapophysis more than 5x longer than high (Fig. 15) appears to have evolved from an inconspicuous Comments: The shape of the hypostigmatic cell appears swollen plate in the Ascalaphidae and Palparinae to to be a very constant character in the Myrme- huge structures in Neulrrtus (Fig. 27). Miller (1991) leontoidea. However, there are a few genera of has suggested that the inflated posterior Myrmeleontidae (Maracandula; Nuglerus) where gonapophyses function as egg handlers whereas the the hypostigmaticcell is short as 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 ascalaphid gonapophy sis type or myrmeleonid type represents the apomorphic Comments: The gonapophyseal plate is an asetose condition. sclerite 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 (Acanthaclisini; presence of the basal subcostal crossvein is Nemoleontini). This might be a synapomorphy of plesiomorphic in Psychopsidae and the prolifera- the Myrmeleontinae except it appears to be absent tion of additional crossveins as apomorphic. This in the Myrmeleontini (a secondary loss?). Some may be true in the Nymphidae as well since New parallel evolution in the reduction of this plate is (1981) clearly shows some genera of Nymphidae seen in the Brachynemurini. It can be moderately with only 1 basal crossvein and others with many large (Dejuna) to very small (Scotoleon and some crossveins. The basal crossvein is absent in the Dendroleontini such as Glenoleon which is variable, Ascalaphidae and Myrmeleontidae which is taken but Glenoleon is probably paraphyletic. as a synapomophy of these two families. If the development of distal crossveins in the Insecta Mundi

18. Lateral gonapophysis, chaetotaxy (Figs. 17-32) - (0) Palpares) additional teeth (4 to 6) are developed. digging setae weakly to moderately developed, never The shortening of the mandible has led to a reduc- longer than gonapophyseal width (Figs. 17-27); (1) tion of teeth in only a few cases. Two teeth are found digging setae large, down curved, longer than in Glenurus (Nemoleontini), Centroclisis gonapophyseal width (Figs. 28,29). (Acanthaclisini) and Millerleon (Dimarini). The lat- ter genus is the most enigmatic since the short teeth 19. Male ectoproct (#20) - (0) without elongate still show remnants of the seta. Only 1 tooth is postventral lobe (Figs. 50, 51, 53); (1) with elongate present in Paranthaclisis (Acanthaclisini). Usu- postventral lobe (Figs. 47, 49, 55). ally, the teeth are parallel with an increasing length Comments: The Ascalaphidae is about as variable as the distally although several groups have the middle Myrmeleontidae in this character so that the polar- tooth the longest. These are all probably derived ity can not be established for the family as a whole. conditions and are not the focus of the present Some variation occurs in the Brachynemurini; analysis. In the present analysis, the primary Myrmecaelurini and Nemoleontini, but the ground apomorphy is the modification of the distal tooth plan 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 CHARACTERS) Myrmecaelurini, and Nesoleontini. Larval characters of the Psychopsidae and Nymphidae are discussed by MacLeod, 1970; New 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) closely approximated 23. Ocular tubercles - (0) absent; (1) present Comments: The approximation of the mandibular bases is found only in the Gnopholeontini amoung the 24. Mesothoracic spiracle - (0) sessile; (1) pedunculate Myrmeleontidae. The most extreme case is Gnopholeon barberi (Fig. 69). Elsewhere the closer 25. Hindtarsus and tibia - (0) articulated; (1) fused approximation of the mandible is found only in the Ascalaphidae. This is evidently a derived condition 26. Hind 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 burrowing 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). Myrmeleontidae is Gnopholeon barberi. The reduc- Comments: The development of mandibular teeth is tion in the size of the hind pretarsal claws is prob- clearly synapomorphicfor 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 the Myrmeleontidae, dibular teeth. Mandibular teeth are apparently and have secondarily lost the enlarged hind pretarsal derived from enlarged setnl bases (Baba, 1954) and, claws as an adaptation to living 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 be the next step without head-first, the anterior pretarsal claws are en- intermediate stages of having 1 or 2 teeth. All larged. known ascalaphid larvae and the great majority of ant-lion 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 fossil nymphid (Pronyrnphes) sternite IX and sometimes found only on the third has 3 parallel teeth and at least 1 undescribed instar larva. extant species has several teeth. Related to larval predation strategies, the mandible has become longer 28. Abdominal sternite VIII - (0) without submedial in some groups, shorter in others. Probably there teeth; (1) with submedial teeth (Fig. 16). has been repeated lengthening and shortening 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 lengthening of the mandible Myrmeleontidae often is accompanied by having the teeth situated more distally, or in a few palparine genera (Golafrus; Vol. 8. No. 1 - 2. March - June. 1994

The characters and taxa were analyzed using the extreme differences in the larvae of the 2 genera the Hennig86 cladistics program (version 1.5) of (loss of the fossoria & submedial teeth in Millerleon), Farris (1989). Using the ie option, 6 equally parsi- the analysis supports the two genera forming a monious cladograms were found (length = 7 1; con- monophyletic group. sistency index (ci) = 0.53; retention index (ri) = The monophyly of the Nemoleontini appears to 0.70). The successive approximations character be weakly supported only by the reversal of char- 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 apomorphy is also found in the Nesoleontini Character weights are given in Appendix 3. The and in the Brachynemurini (Scotoleon). 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 of five families. As can be Brachynemurini + (Lemolemini + (Gnopholeontini appreciated from the cladograms, the Ascalaphidae +Dendroleontini)) in the other cladogram, it was is so close that it could be considered as a subfamily decided to use the Nemoleontini as one of the of Myrmeleontidae. All the autapomorphies for the outgroups. Ascalaphidae appear somewhere in the Myrmeleontidae. However, two other apomorphies Relationships within the Gnopholeontini, exist which may strengthen the case for family Lemolemini, and Brachynemurini recognition of the Ascalaphidae. The antennae are In earlier papers on the classification of the significantly longer than in the Myrmeleontidae Brachynemurini (Stange & Miller 1985; 1992), the except for the bizarre genus Albardia. The caudate Brachynemurini was reduced to a subtribe of the head of the larva may be apomorphic for the Myrmecaelurini. This was based on the remark- Ascalaphidae. able convergence in the dentition of the larval The 2 weightedcladograms are the same except mandible. However, the present analysis suggests for the placement of the Nemoleontini. According a closer relationship of the Myrmecaelurini to the tocurrent classification, the Palparini, Palparidiini, Acanthaclisini based on female terminalia (charac- and Dimarini (Dimares, Millerleon), together with ter #18). The relationship of the Acanthaclisini the Pseudimarini (not included in the analysis with the Myrmecaelurini + Nesoleontini is sup- because the larval stage unknown) constitute the portedby another apomorphy not used in the analy- subfamily Palparinae. The results of the present sis, the presence of abdominal hair pencils. Hair analysis indicate that the subfamily Palparinae in pencils are found in all groups of the Acanthaclisini. the traditional sense is paraphyletic. The Elsewhere, they are found only in some genera of Stilbopteryginae (considered currently as a sepa- the Myrmecaelurini. rate subfamily) could also be a constituent tribe of 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; andBrachynemurini) by this cladistic analysis is to recognize three sub- and indicates aclose relationship of Gnopholeontini families; the Palparinae (Palparidiini and Palparini), with the Dendroleontini. The monophyly of the the Stilbopteryginae, and the Dirnarinae with the Gnopholeontini is supported by the autapomorphy Pseudimarini as incertae sedis, but possibly in the of the close approximation of the larval mandibles Dimarinae. However, more character analysis of (#20). The monophyly of the Lemolemini is sup- the Stilbopteryginae and associatedgroups is needed ported by one larval autapomorphy not entered before such a change is recommended. The into the analysis: the presence of specialized setae Stilbopteryginae have always been a controversial along the median line of the abdomen. The mono- group, sometimes classified with the Ascalaphidae phyly of the Brachynemurini is supported by the based on the clubbed antenna and short elongate postventral of the ectoproct (#19) and hypostigmatic cells which are key characters of the larval dentition (#2 1). Finally, the monophyly of Ascalaphidae. It has also been treatedas a separate the Dendroleontini is supported by one larval family and, more recently (New 1982), as a subfam- autapomorphic character, namely the 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 the Dendroleontini and Nemoleontini as 8. Pretarsal claws - (0) small to moderately large, less outgroups, the following phylogenetic analysis was than one-half length of distal tarsomere; (1) en- made on genera of the Brachynemurini, larged, more than one-half length of distal tarsomere Gnopholeontini and Lemolemini. The following WING CHARACTERS informative characters were used, with the 9. Forewing, origin of vein CUP (#9) - (0) at or before plesiomorphic condition and comments where rel- 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 Brachynemurini, non-additive. Numbers in brackets indicate equiva- Gnopholeontini and Lemolemini. Sometimes vein lence with list of tribal apomorphies. CUPis obsolete in these tribes. The character is very stable, the most basal position of CUPwas observed in Jaffuelia. In a few groups of the Dendroleontini the origin of vein CUPmay be distad of crossvein m- List of generic apomorphies (adult) cu. Elsewhere in the Myrmeleontidae, this derived HEAD CHARACTERS state is found in Nannoleon Esben-Petersen and in 1. Frons - (0) glabrous; (1) setose (Fig. 37) the tribe Maulini. Comments: The derived state is a synapomorphy of Ameromyia and Venezueleon. 10. Forewing, vein 2A - (0) runs at even curve toward 2. Antennae - (0) similar in both sexes; (1) male an- 3A; (1) runs along 1A for short distance, then back tenna longer (more flagellomeres) than female at a sharp angle towards 3A (Nemoleontini) Comments: The derived state is a synapomorphy of Brachynemurus and Scotoleon. 11. Hindwing, vein CuA - (0) bends to hind margin at or 3. Ocular rim setae - (0) absent; (1) 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 development of profemoral 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 of the rim which vary in length pression. The most extreme expression of the de- from genus to genus. The derived state is a rived state is found in the genus Ameromyia in the synapomorphy of Abatoleon, Chaetoleon, Gnopholeon New World Brachynemurini. (2 of 3 species) and Menkeleon. Also, ocular rim setae are found in some generaof the Acanthaclisini, 12. Hindwing, pilula axillaris (#14) - (0) present; (1) Dendroleontini and Myrmecaelurini. absent Comments: The pilula axillaris is sometimes call THORACIC CHARACTERS "Eltringham's Organ". This structure, odoriferous 4. Profemoral clavate setae (Fig. 9) - (0) absent; (1) in function, is found only in the male with a few present, arranged in a row; (2) clustered exceptions (e.g. B. longicaudus). Although a unique Comments: The derived state is a synapomorphy of character for the family, it appears to have been lost Abatoleon and Chaetoleon. In these genera there in some groups of the Myrmeleontinae are at least four such setae in a row and in most (Nemoleontini; Scotoleon). Abatoleon there are secondary rows. In 2 of the 3 species of Gnopholeon these setae are clustered; ABDOMINAL CHARACTERS this state is considered of independent evolution. 13. Female terminalia, ectoproct - (0) with digging setae; (1) without digging setae 5. Femoral sense hair (Fig. 7) (#7) - (0) present; (1) absent 14. Female terminalia, lateral gonapophyses - (0) sepa- Comments: The derived state is a synapomorphy of rated (Figs. 17-20, 22-30); (1) partially fused (Fig. Atricholeon and Venezueleon. The 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 Mara- Palparinae. candula, Jaffuelia, Venezueleon and Ameromyia.

6. Profemoral sense hair - (0) much shorter than 15. Female terminalia, lateral gonapophyses - (0) elon- profemur; (1) as long as or longer than femur gate (Figs. 17-19, 23-29); (1) transverse (Figs. 20-21, 30-32) 7. Tibial spurs - (0) present; (1) absent Comments: The derived state is found in the Comments: The derived state is a synapomorphy of Gnopholeontini. Maracandula, Menkeleon, and Tyttholeon. Tibial spurs are sporadically absent in Brachynemurus (2 species) and Abatoleon (1 species). Vol. 8, No. 1 - 2, March - June, 1994

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. 30-32) Comments: The derived state is found in the 24. Male genitalia, paramere - (0) plate-like; (1) re- Gnopholeontini and the Lemolemini. Extreme ex- duced to separate rods (Fig. 48) pressions are found in Maracandula and Neulatus. Comments: the apornorphic condition is found in Miller (1991) stated that the digitiform type is Ameromyia and Venezueleon. mostly used for soil depth testing whereas the inflated gonapophysis is associated with egg han- 25. Male genitalia, sculpture of mediuncus - (0) smooth dling. (non-spinose, not cross striate); (1) micro spinose (Austroleon); (2) cross-striate (Figs. 54, 59) 17. Female terminalia, posterior gonapophyses - (0) (Gnopholeontini) without scraping setae (Figs. 17-24); (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 scraping 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) occurs in many ascalaphid larvae. wider than posterior gonapophysis (Figs. 22-27, 30- 32) 27. Mandibular teeth (#21) - (0) 3 parallel teeth, usually Comments: The derived state is found in Gnopholeontini increasing in length distally (Figs. 64-71); (1) distal and Dendroleontini. 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) state into 2 major groups using the width 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 than the found in the Old 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 inflated (#2 1) posterior gonapophysis in that genus. The Lemolemini is split into two groups. Neulatus and 28. Hind pretarsal claw - (0) without basal tooth; (1) Jaffuelia have large gonapophyseal plates whereas with basal tooth Elicura, Lemolemus and Sical have narrower plates. Comments: The derived state is a synapomorphy of Neulatus and Jaffuelia. 19. Female terminalia, 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 (#IS) - (0) without elongate sometimes borne on common stalk (Bankisus; many postventral lobe (Fig. 53); (1) with postventral lobe Australian genera of Dendroleontini) (Fig. 41-43, 47, 49) Comments: The development of a medial group of elongate bristles on the mesonotum appears 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 function 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 predator strategy in the antlions. In Bankisus and many Australian groups 22. Male genitalia, genital sac - (0) without clubbed (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 scoli - (0) absent; (1) present Comments: The derived state (#I) is a synapomorphy of Comments: The development of abdominal scoli prob- Ameromyia and Venezueleon whereas the extra ably is related to living on rock surfaces or plants sclerites are found in Argentoleon and Atricholeon. (Gnopholeon; Jaffuelia; Neuhtus), paralleling the These derivatives are non-additive. ascalaphid condition.

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

31. Abdominal tergites, chaetotaxy - (0) without spe- female ectoproct (# 13) and the larval autapomorphy cialized setae medially; (1) with dolichasters; (2) of the development of specialized setae on the with black, tufted setae along midline dorsal mesa1 margin of abdomen (#31-2). Elicura Comments: The derived state is found in the Lemolemini. and Lemolemus have autapomorphies which sup- There are two groups. Roseate, squat dolichasters are found in Neulatus and Jaffuelia whereas other port their monophyly: the sexual dimorphism of lemolemine genera have tufted black setae. the tibia1 spurs in Lemolemus and the forefemoral setal comb and expanded forewing costal area in Based on the character matrix given in Appen- Elicura. dix 2, 3 separate analyses were made. The first The Brachynemurini as newly restricted have analysis was made of the genera in which the larvae several well defined subgroups. The South Ameri- are known. The other 2 analyses included the 5 can genera Austroleon, Abatoleon, Argentoleon, genera in which the larvae are unknown in order to and Ensorra are the sister group to the rest of the determine phylogenetic relationships of these 5 brachynemurine genera although this relationship genera. is found in only 1 of the 2 weighted cladograms and The first analysis was made of 19 genera with is based on character # 19, a very homoplastic char- larval information. Characters #4, 22,25, and 30 acter. Ameromyia + Venezueleon are defined as a were run as non-additive. The mhennig*/bb* op- clade by 3 autapomorphies. These two genera also tion ofHennig86 resulted in 112 cladograms (length are restricted to South America. The other 5 genera = 62; ci = 0.59; ri = 0.7 1). The successive approxima- are all North American. Although both Chaetoleon tions character weightingfeature reduced the num- and Mexoleon have the male ectoproct produced ber of cladograms to 2 (ci = 0.87; ri= 0.93); character ventrally, in Mexoleon the process is much stronger weights are given in Appendix 3. and is considered an autapomorphy. Of all the The 2 cladograms (Figs. 3,4) are nearly identi- genera treated in the analysis, only the original cal, differing only in the treatment of the loss of the genus Brachynemurus has no apomorphies to de- medial tooth on the pregenital plate (#19). This is fend it and will need further study. a homoplastic character appearingin several places 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- trichotomy involving Austroleon, Abatoleon + ing, two additional analyses were done. The first Argentoleon + Ensorra, and the rest of was to add the five genera to the previous data brachynemurine genera. The other cladogram matrix, coding the missing data as unknown. The (Fig. 4) eliminates the trichotomy. mhennig*/bbe option of Hennig86 program was Examination of the 2 cladograms clearly shows run and resulted in 3,248 cladograms (length =67; that larval characters are most important for the ci = 0.56; ri = 0.71) before weighting. Successive discrimination of the three clades of the approximation weighting produced 341 cladograms Brachynemurini s.l., now recognized as three tribes. (ci = 0.84; ri= 0.92) with character weights given in The Gnopholeontini include Gnopholeon, appendix 5. A Nelson consensus tree of these Menkeleon, and Tyttholeon, and are defined by weighted cladograms is given in Fig. 5. characters#15,25-2 & 26. The Lemolemini are the Maracandula comes out in the Gnopholeontini, sister group of the Gnopholeontini + Dendroleontini, Ecualeon and Galapugoleon in the Lemolemini, defined by characters#13 & 31-2. Finally, the third whereas Dejuna and Clathroneuria are shown to be large group of genera now comprising the related to genera comprising the tribe Brachynemurini, are defined by character #27. Brachynemurini s.s. The genera of Gnopholeontini are well defined The second analysis was to run the 25 genera in both cladograms. Tyttholeon has no listed char- using only adult characters. This analysis was less acters in this analysis but is abundantly character- informative regarding the relationships of the five ized by autapomorphies such as the shape of the genera.Using the mhenning*/bb* option of male ectoprocts (Fig. 50), enlarged pregenital plate Hennig86,3,503cladograms were produced (length (Fig. 30), and lanceolate setae on the larval = 59; ci= 0.52; ri= 0.65). The successive approxima- meosternum. tions character weighting feature reduced the num- ber of cladograms to 353 (ci = 0.80; ri = 0.90); The Lemolemini are defined by the adult character weights are given in appendix 6. A autapomorphy of the loss 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 tion than the previous cladograms. The BRACHYNEMURINI Gnopholeontini are resolved, but the genera of Abatoleon - (1) Mediuncus and the parameres of the Lemolemini form a polytomy with the male genitalia highly modified (Fig. 58) Dendroleontini and Gnopholeontini. The genera Maracandula, Ecualeon, and Galapagoleon fall Ameromyia - Enlarged pretarsal claws. within the polytomy and their relationships are thus uncertain. The Brachynemurini S.S. are only Argentoleon - Secondary, lobe-like sclerites later- partly resolved with Abatoleon, Argentoleon, ally on genital sac. Austroleon and Ensorra forming part of a basal polytomy for the entire tree. Dejuna and Atricholeon - Secondary sclerotized and setose plates Clathroneuria fall out in this region. anterior to gonarcus (Fig. 44). Generic autapomorphies Austroleon - Male mediuncus micro-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) presence of large, scale-like (Fig. 42). setae on male abdominal segments 111-V; (3) male ectoproct ventrally emarginate (Fig. 35). Chaetoleon - Posterior gonapophysis with scraping setae. Galapagoleon - Female posterior gonapophysis strongly curved (Fig. 23). Clathroneuria - (1) Postventrallobe of male ectoproct with median lobe (Fig. 43). (2) mediuncus of male Elicura - Profemur with elongate white bristles on genitalia narrow and elongate (Fig. 46). exterior face. Dejuna - Male genitalia with gonarcus elongate, Jaffuelia - Larva with most of body covered with mediuncus with spiniform process, and parameres scale-like setae. abbreviated (Fig. 45).

Neulatus - Apex of sternite VIII with median pro- Ensorra - Male sternite IX extremely elongate (Fig. jection (Fig. 27); (2) pregenital plate modified as 49). elongate, scoop-like structure (Fig. 27). Mexoleon - Ventral process of male ectoproct Sical - Female pregenital plate without median spiniform or truncated (Figs. 5 1, 52). tooth. Scotoleon - Modified tergal setae on male tergite IV (most species). GNOPHOLEONTINI Gnopholeon - (1) Larval head greatly produced at Venezueleon - Larval head capsule about as wide as anterolateral area, with long setae which camou- long. flage mandible in trap position (Figs. 68, 69); (2) profemur with clavate setae clustered. Key to tribes of Myrmeleontidae Adults Maracandula - (1) Eye extremely reduced (Fig. 36); 1. Forewing vein CUPoriginates well distad of basal (2) antenna with flagellomeres 2-4 much longer crossvein (Fig. 11) or obsolete; forewing vein 2A than wide. runs in fairly even curve to hind margin (Fig. 11) ...... 2 Menkeleon Profemur with 2 sensory hairs. Forewing vein CUP originates at or very near - (about cubital vein diameter distad) basal crossvein (Fig. 10); forewing vein 2A variable Tyttholeon - (1) Female pregenital plate very ex- pansive (Fig. 30); (2) male ectoproct produced mesally below (Fig. 50); (3) larva with specialized 2. Labial palps longer than head width, sensory area patch of lanceolate setae on mesothoracic sternum. extends at least to apex; mesofemoral sense 76 Insecta Mundi

hair absent, that of profemur small; southern 15); metafemur without sense hair (except Africa ...... Maulini Sericoleon and Epacanthaclisis); tibial spurs Labial palps shorter than head width, area not andpretarsal claws various, usually not strongly extending to apex; mesofemoral sense hair arched or bent; male abdomen without hair present, sometimes very small or absent in pencils (except Myrmecaelurini) ...... 7 some genera (Atricholeon; Venezuelwn) in which all leg setae are greatly reduced ...... 3 7. Forewing vein 2A runs in fairly even curve toward 3A (Fig. 11) ...... 8 3. Southern Africa; 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 (Fig. 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 corresponding 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 ectoproct with postventral lobe at least than that of forewing before forks (Figs. 12, 13); longer than diameter (Figs. 41-43, 47, 49) or origin of hindwing radial sector before medial profemur with row of clavate setae (Chaetoleon) fork, usually only 1 or 2 presectoral crossveins; (Fig. 9) or mesofemoral sense hair longer than female anterior gonapophysis often lobe-like; femur (Austroleon); 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. Hindwing 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 in male pilula axillaris present or absent ...... 10 North American Gnopholwn; female posterior gonapophysis swollen thumb-like (Figs. 24-26, 10. Basitarsusof hindleg longer than apical tarsomere; 30-32), distally swollen (Fig. 27), 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 Gnopholwn) ...... Nesoleontini mesofemoral 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) ...... Gnopholeontini eral 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 narvae of Maulini and Nannoleon unknown) 1. Labial palpus shorter than basal width of man- 6. Hindwing vein CuA unites with posterior fork of dible; mesothoracic spiracle not borne on tu- MP, shortly after fork or (Paranthaclisis) bercle; head without dolichasters; backward metafemur with elongate sense hair; pretarsal movement only (many genera) or backward and claws strongly arched near base; tibial spurs forward movement ...... 2 strongly curved, sometimes bent at right angle; Labial palpus longer than basal width of man- male abdomen with hair pencils (except dible or mesothoracic spiracle borne on tu- Arcuaplectron) ...... Acanthaclisini bercle; head often with dolichasters; backward Hindwing vein CuA connected by crossveins to and forward movement ...... 3 posterior fork of MP, or not reaching fork (Fig. Vol. 8, No. 1 - 2, March - June, 1994

2. Mandible with some setae on outer lateral margin crossveins to posterior fork of MP, 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 inconspicuous usually 3 or more presectoral crossveins, posterior submedian teeth near posterior margin; make area often broadened; male abdomen without hair pitfall traps ...... Myrmeleontini Mandible with longest setae on outer lateral mar- pencils; male ectoproct usually with postventral gin less than one-half greatest mandibular lobe; male paramere often hinged; female anterior width; sternite VIII without teeth on subapical gonapophysis membranous or plate-like; margin; do not make pitfall traps ...... gonapophyseal plate moderately large to very re- ...... Acanthaclisini duced; pregenital plate with or without median tooth; posterior gonapophysis cylindrical, often re- 3. Mandible bases separated by less than width of duced; female ectoproct with digging setae (except one mandibular base (Figs. 68-71); Sonoran Ensorra) . subregion ...... Gnopholeontini Mandible bases separated more than width of one mandibular base (Figs. 60-67) ...... 4 Larval Description: Labial palpus longer than basal width of mandible; mandibles not upcurved, 4. Mesoscutum with medial tuft of setae at middle; separated by much more than basal width of man- 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 Mesoscutum without medial tuft of setae; ab- dolichasters; mesothoracic spiracle pedunculate or dominal sternum IX wider than long; man- not; mesonotum without medial bunch of debris- dibles straight ...... 5 holding setae; sternite VIII with or without 5. Abdominal terga I-IV with specialized setae sag- submedian tooth near posterior margin; abdomen ittally consisting of incurved setae or squat without specialized setae along median line; with- dolichasters; western and southern South out fossoria. America (especially Chile; Galapagos Islands) ...... Lemolemini Discussion: The phylogenetic relationships of the Abdominal terga I-IV without specialized setae Brachynemurini are difficult to understand. On sagittally ...... 6 the basis of the cladistic analysis of the higher categories (Fig. 1) and of genera previously con- 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 ing the Brachynemurini by the following other teeth (with 3 teeth only) ...... Brachynemurini, Myrmecaelurini & apomorphies: (1) migration of forewing CUPwell, Nesoleontini distad of crossvein m-cu (shared with Mandible with distal tooth as long as or longer Gnopholeontini, Lemolemini, Maulini, some than middle tooth, teeth parallel (few genera Dendroleontini and Nanmleon); (2) cylindrical pos- with 2 teeth) ...... Nemoleontini terior gonapophyses (found in some Old World groups); (#3) short, deflected distal tooth of larval mandible (shared with Myrmecaelurini and Tribe Brachynemurini Banks, 1927 Nesoleontini). Additional apomorphies found in the = Austroleonini Banks, 1927 Brachynemurini which help distinguish the tribe from the Gnopholeontini and Lemolemini are (1) Adult Description: Labial palps short to moder- the elongate postventral lobe of the male ectoproct ately long, palpimacula not extending to apex; (found in the majority of the genera) and (2) the metafemur without sensory hair; pretarsal claws hingedmale paramere (found in 50%of the genera). not toothed, rarely strongly bent near base; tibia1 spurs present or rarely absent (1 species of Key to genera of Brachynemurini Abutoleon; 2 species of Brachynemurus) forewing vein CUPoriginates at or near crossvein M-Cu; 2A Adults runs in a fairly even curve toward 3A, pilula axillaris 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 MP, (Fig. Insects Mundi

12); posterior area 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 vein CuA runs along posterior fork of tina ...... Ensorra Nav4s MP,for a long distance, with 3 or more crossveins Male sternite IX much smaller than sternite VIII; between them (Fig. 13); or posterior area of female ectoproct with digging setae (Fig. 22); hindwing broad (Fig. 14), at least twice as high apex of hindwing in repose clearly extends before medial fork as corresponding area of beyond apex of forewing; Mexico to Costa Rica forewing ...... 8 ...... Dejuna Nav4s

Profemur with clavate setae (Fig. 9); ocular rim 8. Profemoral sense hair absent ...... 9 usually with several short setae that project Profemoral sense hair present ...... 10 over eye ...... 3 Profemur without clavate setae; ocular rim with- 9. Antennal fossae separated by about width of out setae ...... 4 pedicel; frons without setae; forewing costal area with costal cells interconnected; male Male ectoproct with postventral lobe at least ectoproct with postventral lobe more than 6X as twice as long as middle diameter; female long as diameter at midlength (Fig. 41); North terminalia with a membranouspregenital plate; America ...... Atricholeon n. genus (Fig. 19); posterior gonapophysis more than 4X Antennal fossae separated by about 2X width of as long as diameter at midlength, without scrap- pedicel; frons with setae; forewing costal area ing setae; Neotropics ...... Abatoleon Banks with cells not interconnected; male ectoproct Male ectoproct with postventral lobe never longer with postventral lobe about as long as wide (Fig. than diameter at midlength, sometimes absent 47); South America ...... Venezueleon n. genus (Fig. 53); female terminalia with sclerotized 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); forewing CUP+ eter at midlength, with scraping setae; North 1A parallels posterior branch of CuA for a America ...... Chaetokon Banks distance, about equal to that between hindwing CuA and posterior branch of MP,; profemoral Profemoral sense hair at least three-fourths as sense hair as long as femur; Venezuela to Ar- long as femur, or pretarsal claws longer than gentina ...... Ameomyia Banks hind basitarsus (Austroleon lizeri) ...... 5 Pretarsal claws much shorter than distal Profemoral sense hair (if present) no more than tarsomere (Fig. 38), or frons without setae; one-half length of femur; pretarsal claws longer forewing CUP+ 1A runs to posterior margin at or shorter than hind basitarsus ...... 7 or before posterior fork of MP,, or profemoral sense hair shorter than femur; North America Male ectoproct with conspicuous median lobe in 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 of long white bristles; 38); tibial spurs longer than greatest tarsomere temperate North America ...... diameter; male ectoproct with postventral lobe ...... Clathroneuria Banks about as long as wide, ventrally with a short Male ectoprdct without median lobe, sometimes process (Fig. 51, 52) ...... Mexokon n. genus without postventral lobe; profemur and procoxa Pretarsal claws longer than hind basitarsus, or sometimes with row of long white bristles; tibial spurs absent or vestigial; male ectoproct female abdomen usually shorter than wings; with postventral lobe longer than wide, not South America ...... 6 directed ventrally ...... 12

Male ectoproct with postventral lobe at least 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 bristles ...... sponding area of forewing; 4-6 crossveins be- ...... Argentoleon n. genus tween hindwing CuA and hind margin; male Male ectoproct without postventral lobe or if pilula axillaris absent; setal bases on male short lobe developed, less than 2x longer than tergite IV often spined; vertex with dark pig- wide; profemur with row of long white setae .. ment, when present, restricted to scars ...... Austrolwn Banks ...... Scotoleon Banks Posterior area of hindwing narrow (Fig. 13), less Male sternite IX enlarged, nearly as wide as than twice as wide at medial fork as corre- sternite VIII (Fig. 49); female ectoproct without sponding region of forewing, more than 10 Vol. 8,No. 1 - 2. March - June, 1994

crossveins between CuA and hind margin of 8. Ventral surface of head capsule with dolichasters hindwing or (in B. sacken9 vertex scars yellow- ...... 9 ish with dark pigment in interscar area; setal Ventral surface of head capsule without bases on male tergite IV not spined; pilula dolichasters ...... 10 axillaris present or absent ...... Brachynernurus Hagen 9. Head capsule longer than wide (measured ven- trally); abdominal tergites with numerous Larvae thread-like setae between regular setae ...... 1. Sternite VIII without a pair of well defined ...... Arneromyia (modesta Group) submedian teeth ...... 2 Head capsule about as wide as long (measured Sternite VIII with a pair of sclerotized submedian ventrally); abdominal tergites without thread- teeth (Fig. 16) ...... 3 like setae ...... Venezueleon

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 ...... Mesothoracic spiracle borne on tubercle that is ...... Atricholeon and Dejuna shorter than its basal width ...... 11

3. Abdominal spiracles enlarged andlor borne on 11. Submedian teeth of sternite VIII 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 VIII well developed, length of distal tooth; mesothoracic spiracle longer than basal widths; head capsule shorter borne on tubercle that is shorter than basal along dorsal midline than across ocular tu- width ...... Mexoleon bercles; distance between basal and distal teeth Mandible as long as head capsule, with distal longer than that between base and first man- tooth separated from middle tooth by less than dibular tooth ...... 112 length of distal tooth; mesothoracic spiracle ...... Arneromyia (nigriventris Group) borne on tubercle that is longer than basal width ...... 5 1. A batoleon Banks 5. Mandible with distance between base and basal tooth longer than between basal and distal (Figs. 19, 58) tooth; abdominal spiracles often with associ- ated secondary tube-like structures; submedian Abatoleon Banks, 1924. Bull. Mus. Comp. Zool. 65:436. teeth on sternite VIII shorter than basal widths Type species: Abatoleon deprivatum Banks, by ...... Abatoleon monotypy . Mandible with distance between base and basal tooth equal to that between basal and distal Adult Description: Frons without setae; anten- tooths; abdominal spiracles without associated nal fossae separated by about width of pedicel; tube-like structures; submedian tooth of stern- profemur with clavate setae (sometimes weakly ite VIII longer than basal widths ...... thickened subapically); ocular rim with or without ...... Argentoleon and Austroleon setae that project over eye; femoral sense hair of 6. North America ...... 7 foreleg about as long as femur and much longer South America ...... 8 than that of midleg; mesonotum without bladelike setae; tibia1 spurs present (except A. deprivatus 7. Mandible elongate with distance between base Banks in Argentina); pretarsal claws longer than and basal tooth much longer than between hind basitarsus but less than Yz length of distal basal and distal teeth ...... Chaetoleon tarsomere;pilula axillaris present;hindwing shorter Mandible shorter with distance between base and than forewing, in repose apex of wings nearly basal tooth less than that between basal and coincide;banksian lines absent; costal area offorew- distal teeth ...... Brachynemurus ing with one series of cells for most of distance; posterior area of hindwing narrower than presectoral area, CuA bends to hind margin at or 80 Insecta Mundi before origin of medial fork; antenna short, usually tibial spurs. Also, these two species lack digging with less than 30 flagellomeres; postven tral lobe of setae on the female ectoproct. Abatoleon is the only male ectoproct well developed but less than one- Neogeic element of the Brachynemurina present in half length of eighth abdominal segment (except A. continental North America. One species,A. indiges, frontalis; A. deprivatus), without median or sec- has reached the Florida Keys, having apparently ondary lobes; gonarcus reduced, mediuncus of male dispersed from South America via the Caribbean genitalia smooth, conspicuously expanded cover- islands. The other North American representative, ing modified parameres; genital sac without sec- A. dorsalis, is found throughout most of Mexico and ondary sclerites or clavate setae; female terminalia is established in southern Texas. Known North with pregenitalplate membranous; gonapophyseal American larvae have enlargedabdominalspiracles, plate long and slender; lateral gonapophysis sepa- but this character is known to be variable among rate, with weak digging setae; ectoproct with dig- South American species. ging setae; posterior gonapophysis elongate, some- times somewhat swollen. 2. Ameromyia Banks (Fig. 37) Larval Description (based on dorsalis, indiges, and several undescribed Peruvian species): AmeromyiaBanks, 1913. Trans. Amer. Ent. Soc. 39:227. Head capsule with or without dolichasters; basal Type species: Brachynemurus strigosus Banks, by tooth is closer to distal tooth than to mandibular original designation. #base;mesothoracic spiracle borne on tubercle that is longer than wide; abdominalspiracles sometimes =AmazoleonBanks, 1913. Trans. Amer. Ent. Soc. 39:229 borne on tubercles; submedian teeth on sternite (after Stange 1967:45). Type species: Myrmeleon VIII present, shorter than basal widths. pubiventris Walker, by original designation.

Included species: bruneri (Alayo), camposi =FoyaNavhs, 1914. Broteria 12:53 (after Esben-Petersen 1920:190). Type species: Foya trapezia Navls, by (Banks), dep rivatus Banks, dolichogaster (NavAs), original designation. dorsalis (Banks), frontalis (Banks), garciana (Banks); indiges (Walker). =Nemotolus Banks, 1943. Bol. Ent. venezolana 2:163 (after Stange 1967:45). Type species: Myrmeleon Distribution: Nearctic and Neotropical Region protensis Gerstaecker, by original designation. (Florida Keys; Caribbean Islands; Southwestern Texas to Argentina). Adult Description: Frons with setae; antenna of same length on both sexes, moderately short with Discussion: The presence of clavate setae on the about 30 flagellomeres; antenna1 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 are only weakly thickened hair of proleg as long as femur and much longer apically. The elongate male postventral ectoproct than that of mesoleg; mesonotum without bladelike lobe and the autapomorphic mediuncus and setae; tibial spurs present; pretarsal claws nearly parameres of the male genitalia distinguish this as long as distal tarsomere; pilula axillaris large; sex. In several characteristics Abatoleon parallels hindwing as long as forewing or shorter; anterior Dejuna; both have highly specialized male genita- banksian line present in both wings; forewing cos- lia, similar wing venation, and are restricted to tal area simple; posterior area of hindwing about as tropical climates. broad as that of forewing; forewing vein CUP+ 1A runs along posterior fork of CuA for long distance; The present center of diversity of Abatoleon is hindwing CuA runs along posterior fork of medial South America, especially in the Peruvian coastal fork for a long distance; postventral lobe of male desert and Argentina. Many undescribed species ectoproct developed, usually less than 3X longer exist in Peru. The two aberrant Argentinian spe- than middle diameter; without median or second- cies, A. frontalis (Banks) and A. deprivatus Banks, ary lobes; male genitalia highly modified, gonarcus may belong to a different genus. These two species reduced, mediuncus smooth, greatly expanded cov- have short abdomens and very elongate male ering modified rod-like parameres; genitalic sac pos tven tral ectoproct lobes, and deprivatus lacks with clavate setae; female terminalia with pregeni- Vol. 8, No. 1 - 2, March - June, 1994 tal plate membranous; gonapophyseal plate short; one-half length of distal tarsomere; pilula axillaris lateral gonapophysis fused, with well developed large; hindwing shorter than forewing, in repose digging setae; ectoproct with well developed dig- apices ofwings nearly coincide; costalarea offorew- ging setae. ing with one series ofcells; posterior area ofhindwing narrower than presectoral area, CuA bends to hind Larval Description (based on modesta and margin at or before origin of medial fork; anterior nigriventris): Head capsule with dolichasters; banksian line weakly developed; postventral lobe of mandible with distance between base and basal male ectoproct more than 8 times longer 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-like, without hinge; genital 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 nor borne on tubercles; submedian tooth on sternite narrow, short, not transverse; lateral gonapophyses VIII longer than basal width. separated, with strong digging setae; ectoproct with digging setae; posterior gonapophyses with- Included species: dimidiata NavAs, longiventris out scrapping setae. (NavAs), hirsuta NavAs, modesta (Banks), muralli NavAs, nigriventris (Walker), pentheri NavAs, Larval Description (Based on A. irrigatus): Head pleuralis NavAs, pubiventris (Walker), protensus capsule with dolichasters; mandible with distance (Gerstaecker), stevensi NavAs, strigosa (Banks), between base and basal tooth about equal to that 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- Andes. ite VIII longer than basal width.

Discussion: This is a distinctive genus in which Included species: irrigatus (Gerstaecker) & the pretarsal claws are greatly enlarged and the longitudinalis (Navas). male genitalia highly modified. The same modified male genitalia are found in Venezueleon. The Distribution: South America east of the Andes. setose frons is shared only with Venezueleon. The profemoral sense hair is extremely long, but the Discussion: Argentoleon has one autapomorphy, 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 genital 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 characters (clavate setae of dunes. The larval head capsule is relatively wide, the male genitalia; form of the male parameres; but less so than in the related Venezueleon. shape of forewing vein CUP+ 1A) which relate it to Venezueleon and less to Argentoleon. One variable 3. Argentoleon Stange, new genus species, A. irrigatus, ranges from Venezuela to Patagonia with some striking geographicforms. In Type species: Myrmeleon irrigatus 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, Adult Description: Frons without setae; 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; antenna1 fossae separated by about of longitudinalis. The larva was found in sand on width of pedicel; profemur without clavate setae; a grassy slope and resembles closely the larva of ocular rim without setae; profemoral sense hair as Austroleon, long as profemur and as long as mesofemoral sense hair; mesonotum without blade-like setae; tibia1 spurs present; pretarsal claws large, longer than Insecta Mundi

4. Atricholeon Stange, new genus development of sclerotized and setose plates ante- (Figs. 41, 44) rior to gonarcus is an autapomorphy. The two known species differ by numerous Type species: Brachynemurusparkeri Stange, by structural details and are completely allopatric. A. present designation. tuberculatus is found only north of the Neovolcanic Plateau of Mexico whereas A. parkeri occurs south Adult Description: Frons without setae; anten- of this plateau. A. tuberculatus appears to be more nal fossae separated by about width of pedicel; highly derived than A. parkeri in characters of the profemur without clavate setae; ocular rim without male genitalia, male ectoproct, and vertex protu- setae; femoral sense hair absent; mesonotum with- berances. The strongly humped mesoscutellum of out bladelike setae; tibial spurs present; pretarsal parkeri is apomorphic for that species. claws shorter than hind basitarsus and less than one-half length of distal tarsomere; pilula axillaris 5. Austroleon Banks small with setae restricted to apical margin of knob; hindwing shorter than or nearly equal to forewing; Austroleon Banks, 1909. J. New York Ent. Soc. 17:3. anterior banksian line weakly developed; costal Type species: Austroleon dispar Banks, by subse- area of forewing mostly biareolate; posterior area of quent designation by Banks 1927:53. hindwing narrower than presectoral area at radial =Mom NavBs, 1912. Broteria 10:34 (after Stange sector although wider than prefork area; CuAruns 1967:47). Type species: Moza nubilis NavBs, by along posterior fork of MP, for a short distance; original designation. antenna relatively long, usually with more than 40 flagellomeres; postventral lobe of male ectoproct =Correa NavBs, 1914. Broteria 12:217 (after Stange longer than abdominal segment VIII, without a 1967:47). Type species: Correa expansus NavBs, 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- =Clotus NavBs, 1922. Rev. Acad. Madrid 19:257 (after tose plates present anterior to gonarcus; female Stange 1967:48). Type species: Clotus atomicus terminalia with pregenitalplate membranous, with- NavBs, by original designation. out a median tooth; median floor with sclerotization medially and subapically ;lateral gonapophysis sepa- =Cameo NavBs, 1926. Broteria 23:5. New name for rate, with very strongly developed digging setae; Correa NavBs, 1914. ectoproct with strongly developed digging setae, gonapophyseal plate relatively short and with fine Adult Description: Frons without setae; antenna striae. short, usually about 30 flagellomeres; antenna1 fossae separated by abou t width of pedicel; profemur 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- Eizeri) and as long as mesofemoral sense hair; dominal spiracles not enlarged; submedian tooth mesonotum without blade-like setae; tibial spurs on sternite VIII reduced to bulge. present; pretarsal claws longer than one-half distal tarsomere; pilula axillaris large; hindwing shorter Included species: parkeri (Stange) and than forewing, in repose apices of wings nearly tuberculatus (Banks). coincide; costal area of forewing 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 banksian line Discussion: The reduction in leg setae, particu- weakly developed or absent; postventral lobe of larly the loss of the femoral sense hair, is an male ectoproct absent or weakly developed (less apomorphy of this genus. This condition is dupli- than 2X longer than wide); without median or catedonly in Dejuna mimica and Venezueleon. The secondary lobes; male paramere plate-like, without hinge; mediuncus micro spinose; female terminalia Vol. 8. No. 1 - 2. March - June. 1994 with pregenital plate membranous, without me- implexci Navbs (=Brachynemurus californicus dian tooth; gonapophyseal plate narrow, short, not Banks), by monotypy. transverse; lateral gonapophyses separate, with strong digging setae; ectoproct with digging setae. =Netroneurus Banks, 1927. Bull. Mus. Comp. Zool. Harv. 68: 42. Type species: Brachynemurus carolinus Banks, by original designation. Larval Description (based on immitus; lizeri): Head capsule with dolichasters; mandible with =Bollenga NavBs, 1932. Bull. Mus. Zool. Torino (Ser. 3) distance between base and basal tooth about equal 42: 13. Type species: Bollenga dinamitensis Navls to that between teeth; mesothoracic spiracle borne (=Brmhynemurus sackeni Hagen). on tubercle that is longer than basal width; abdomi- nal spiracles weakly raised; submedian tooth on Adult Description: Frons without setae; antenna sternite VIII longer than basal width. at least moderately long with more than 30 flagellomeres, that of male longer and less clavate Included species: dispar Banks, immitus than in female; antenna1 fossae separated by less (Walker), lizeri (NavBs). 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: Austroleon is apomorphically defined mesonotum without blade-like setae; tibia1 spurs by the microspinose male mediuncus. The elongate present or absent (nebulosus and signatus); femoral sense hair of both the proleg and mesoleg is pretarsal claws equal to or longer than hind found in all species except for male A. lizeri which basitarsus but less than one-half length of distal has a shorter femoral sense hair. Larvae of two tarsomere; pilula 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. lizeri). Based on rior banksian line usually well developed; costal larval exuviae these 2 larvae differ more from 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. posterior area of hindwing moderately broadened (except B. sackeni); much broader near base than at 6. Brachynemurus Hagen medial fork, CuA runs along posterior fork of MP, (Figs. 11, 13, 16, 17, 42) for a long distance; postventrallobe ofmale ectoproct well developed but less than one-half length of Brmhynemurus Hagen, 1888. Canadian Ent. 20:34. abdominal segment VIII (except B. sackeni), with- Type species: Myrmeleon longicaudus Burmeister, out secondary lobes but narrowed preapically $ by subsequent designation of Banks 1899. many species; male genitalia ordinary with simple gonarcus and smooth mediuncus, paramere with =Calinemurus Banks, 1899. Canadian Ent. 31:70 (after hinge and hook; female terminalia with pregenital Stange 1970:51). Type species: Brachynemurus plate large, sclerotized, with small median tooth californicus Banks, by original designation. anteriorly; gonapophyseal plate moderately expan- sive; lateral gonapophyses separate, with digging =Hesperoleon Banks, 1913: Entomol. News 24:64 (after Stange 1970:51). Type species: Myrmeleon ferox setae; ectoproct with digging setae. Walker, by original designation. Larval Description (based on carolinus, =Mastega Navls, 1914. Bull. Brooklyn Ent. Soc. 9:13 divisus, exigua, ferox, longicaudus, nebulosus, (after Stange 1970:51). Type species: Mastega ramburi, sackeni, seminolae): Head capsule texanus Navls (=B.irregularis Currie), by monotypy. with dolichasters; mandible with distance between base and basal tooth less than that between teeth; =Comptesa Navls, 1915. Mem. R. Acad. Cien. Barcelona mesothoracic spiracle borne on tubercle; abdominal (Ser. 3) 11:464 (after Banks 1942:139). Type spe- spiracles not enlarged nor borne on tubercles; cies: Comptesa ambitiosa Navls, (=B.californicus Banks), by monotypy. submedian tooth on sternite VIII shorter than basal width. =Belluga Navbs, 1920. Ann. Soc. Sci. Bruxelles 39:194 (after Banks 1942:139). Type species: Belluga 84 Insecta Mundi

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 NavBs, elongatus Banks, exigua (NavAs), sometimes producedventrally ;male genitalia rather ferox (Walker), fuscus (Banks), hubbardi Currie, ordinary, paramere with hinge and weakly devel- irregularis Currie, longicaudus (Burmeister), oped hook; female terminalia with pregenital plate mexicanus Banks, henshawi (Hagen), nebulosus sclerotized with a small median tooth; (Olivier), pulchellus Banks, ramburi Banks, B. gonapophyseal plate large; posterior gonapophysis sackeni Hagen, B. seminolae Stange, B. signatus with scraping setae; lateral gonapophyses sepa- (Hagen), and B. versutus (Walker). rate, with digging setae.

Distribution: Nearctic region to about 500 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 borne on weak diversity in this group, making it difficult to char- tubercle; abdominal spiracles not enlargednor borne acterize as a whole. No synapomorphies are known 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. Tibia1 spurs may be promi- (Currie), tripunctatus (Banks) and variabilis nent, minute, or absent. Thepilula axillaris may be (Banks). well developed, weakly developed, or absent. The parallel tracks of the CuA and posterior fork of MP, Distribution: North America. in the hindwing are a unifying generic characteris- tic, except for B. sackeni. The genus shares with Discussion: The presence of profemoral clavate Scotoleon the synapomorphy of antennal sexual setae distinguishes Chaetoleon from all other dimorphism. Larvae have been found in open 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. Chaetoleon Banks and diagnostic within the Brachynemurini. Rela- (Figs. 7, 53, 54) tionship with Mexoleon is indicated by a weak ventral lobe of the male ectoproct (compared with a Chaetoleon Banks, 1920. Bull. Mus. Comp. Zool. 64:328. strong ventral lobe in Mexoleon). There are two Type species: Myrmecoleon pumilis Burmeister, very distinct species groups. The C. pumilis group by original designation. has a simple male ectoproct, more elongate legs and antennae, blade-like mesoscutal bristles, and more Adult Description: Frons without setae; antenna weakly swollen posterior gonapophyses relative to moderately long to very long; antennal fossae sepa- the C. tripunctutus Group. One species, C. pusillus, rated by less than width of pedicel; profemur with has a bag-like structure on the male postmentum. row of clavate setae; ocular rim with several short The larva of C. pusillus has been found in sand setae that project over eye; femoral sense hair of under plants. Chuetoleon variabilis has the female proleg much shorter than tarsus, longer than that scraping setae less developed than in other species. of midfemur; mesonotum with conspicuous, weakly flattened or bladelike bristles; tibia1 spurs present; 8. Clathroneuria Banks pretarsal claws longer or shorter than hind (Figs. 43, 46) basitarsus, not longer than one-half length of distal tarsomere; pilula axillaris small with setae re- Clathroneuria Banks, 1913. Entomol. News 24:65. stricted to distal margin of knob; hindwing shorter Type Species: Brachynemurus schwarzi Currie, than forewing, in repose apices coincide; banksian by original designation. lines absent; forewing costal area with single series of cells; posterior area of hindwing narrower than Adult Description: Frons without setae; antenna presectoral area before radial sector, CuA bends to moderately long with over 30 flagellomeres; anten- hind margin at or near medial fork; postventral nal fossae separated by less than width of pedicel; Vol. 8, No. 1 - 2, March - June, 1994 profemur without clavate setae; ocular rim without Adult Description: Frons without setae; antenna setae; femoral sense hair of proleg unusually long short, usually with less than 30 flagellomeres; (as long as femur), over twice as long as that of antenna1fossae separated by about width of pedicel midfemur; mesonotum without bladelike setae; or more (D. mimica); profemur without clavate tibial spurs present; pretarsal claws much longer setae; ocular rim without setae; femoral sense hair than hind basitarsus and more than one-half as various (absent in mimica), that ofprofemur longer long as distal tarsomere. Pilula axillaris small with than that of midfemur but shorter than length of setae restricted to apical margin of knob; hindwing femur; mesonotum without blade-like setae; tibial shorter than forewing, in repose wing apices nearly spurs present; pretarsal claws longer than hind coincide; anterior banksian line usually well devel- basitarsus, no longer than one-half length 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), CuA bends to hind margin before me- as forewing, in repose apex of hindwing extends dial fork or runs along posterior fork of MP, for a beyond that of forewing; anterior banksian line short distance; forewing costal area with single weakly developed; costal area of forewing with one series of cells; postventral lobe of male ectoproct series of cells for most of distance; posterior area of with a median lobe in addition to postventral lobe hindwing narrower than presectoral area, CuA (Fig. 43); mediuncus smooth, long and narrow; bends to hind margin at or before medial fork; paramere hinged without hooks (Fig. 46); female postventral lobe of male ectoproct longer than terg- terminalia with pregenital plate sclerotized 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 terminalia with pregenitalplate membranous ectoproct with digging setae. or sclerotized, 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 of Atricholeon. Mexico. Included Species: fenestrata (Banks) (=angusta Discussion: The unusual male ectoproct (Fig. 43), Navas), mimica (Stange),persimila (Stange), setosa in which a median lobe is developed in 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 genitalia (Figs. 59, 60) are B. coquilletti is the only species in the group having autapomorphic for the genus. D. mimica which a broad hindwing posterior area, one similar to that lacksfemoral sense hairs, has abbreviatedpretarsal found in Scotoleon, and a short male ectoproct lobe. claws and tibial spurs, 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 palpimacula parallels in many ways species of many times, all attempts to find larvae have failed. Atricholeon.

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

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

Adult Description: Frons without setae; antenna Adult Description: Frons without setae; antenna short with about 28 flagellomeres; antennal fossae with more than 35 flagellomeres; antennal fossae separatedby about width ofpedicel; profemur with- separated by less than width of pedicel; profemur out clavate setae; ocular rim without setae; femoral without clavate setae; ocular rim without setae; sense hair short on proleg and mesoleg; mesonotum femoral sense hair moderately long (subequal in without bristles; tibial spurs present; pretarsal length to distal tarsomere of proleg), that of profemur claws about equal to basitarsus and much shorter about equal in length to that of midfemur; than one-half length of distal tarsomere; hindwing mesonotum without blade-like setae; tibial spurs shorter than forewing, in repose apices of wings and pretarsal claws shorter than hind basitarsus, coincide; pilula axillaris present; costal area of latter shorter than one-half distal 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 just before radial sector; hindwing vein CuAreaches apices coincide; anterior banksian line poorly de- hind margin about at level of medial fork; banksian veloped; posterior area of hindwing narrower than lines absent; male ectoproct with elongate greatest width of presectoral area, about as broad postventral lobe; male sternite IX unusually long; at base as at medial fork, CuA runs along posterior male genitalia with simple gonarcus, smooth branch of medial fork a short distance; forewing 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 of cells; gonapophyseal plate narrow, elongate, not trans- postventral lobe of male ectoproct barely produced, verse; lateral gonapophyses separate, with digging about as wide as long, ectoproct producedventrally setae; ectoproct without digging setae. intoa process (Fig. 112);male genitalia with hinged paramere and smooth mediuncus; genital sac with- Larval Description (based on verticalis): out secondary sclerites, and clavate setae; female Mandible with distance between base and basal terminalia with pregenital plate sclerotized, with a tooth less than that between teeth; mesothoracic median tooth anteriorly; gonapophysealplate mod- spiracle borne on tubercle; abdominal spiracles not 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 Nav As). capsule depressed posteriorly, with dolichasters, many very short; mandible elongate, not broadened Distribution: Argentina. 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 about as long as basal width; abdominal spiracles autapomorphy of this species is the enlargement of enla.- ;ed, raised; submedian teeth on sternite VIII male sternite IX (Fig. 49). The lack of clavate longer than basal widths. profemoral setae and setal comb, absence of hgging setae on the female ectoproct, the presence of short Included Species: mixtecus (Stange) andpapago pretarsal claws and profemoral sense hair distin- (Currie). guish this species from other South American Brachynemurini. The larva resembles that of Distribution: Arid tropical regions of southern Scotoleon pallidus but the only larva available for Mexico and montane regions of southwestern United examination is damaged and therefore not keyed. States and northern Mexico. The larva makes a pitfall trap. Discussion: The male ectoproct which is produced 11. Mexoleon Stange, new genus ventrally (Fig. 52) is diagnostic and synapomorphic (Figs. 38, 51, 52) for the genus, although it is approached in struc- ture by species of Chaetoleon. Mexoleon, however, Type Species: Brachynemurus mixtecus Stange, lacks the clavate profemoral setae of Chaetoleon. by present designation. Vol. 8. No. 1 - 2. March - June. 1994

12. Scotoleon Banks quadripunctatus (Currie), singularis (Currie), and (Figs. 8, 14, 18, 60-63) yavapai (Currie).

Scotoleon Banks, 1913. Ent. News 24:64. Type Spe- Distribution: West of the Mississippi River, north cies: Brachynemurus longipalpis Hagen, by origi- to about 45" North latitude and south to temperate nal designation. Mexico.

Adult Description: Frons without setae; antenna Discussion: The broad posterior area of the short or elongate, usually longer and less clavate in hindwing (Fig. 14) is the key character for the male; antennal fossae separated by about width of group, which is duplicated only in Clathroneuria pedicel; 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, medial lobe than that of mesoleg; mesonotum without blade- of the 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 cially in smaller specimens. There is considerable length of distal tarsomere; pilula axillaris absent; differentiation of the labial palps in 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 of the pilula axillaris, broadened 6 or fewer crossveins between CuA and posterior hindwing posterior area, and the presence of modi- margin; CuA diverges to hind margin along poste- fied tergal setae on the male abdomen are generic rior fork of MP, (Figs. 37, 38); postventral lobe of synapomorphies. This is probably one of the most male ectoproct short to elongate, without second- actively evolving genera in the Brachyneumurini ary lobes; male genitalia ordinary with simple judging by the Wiculty of separating some species arched gonarcus and smooth mediuncus; paramere and the presence of polytypic species. The wing with hinge and hook (except S. expansus and S. venation, especially the broadened hindwingposte- 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 sclerotized, with- most species and are different from other out a median tooth; gonapophysealplate small (Fig. Brachynemurini (except Atricholeon and Dejuna) 32); lateral gonapophyses separate, with digging in lacking submedial teeth on sternite VIII. setae; ectoproct 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 between base and basal tooth less than between present designation. basal and distal 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; antennal fossae separated by more than twice width Included Species: carrizonus (Hagen), def2ems of pedicel; pronotum about twice as wide as long; (Adams), dissimilis (Banks), eiseni (Banks), profemur without clavate setae or setalcomb; femo- expansus (NavBs), fidelitas (Adams), infuscatus ral sense hair absent; tibial spurs present; pretarsal (Adams), intermedius (Currie), longipalpis (Hagen), claws shorter than hind basitarsus and one-half marshi (Stange), minusculus (Banks), minutus distal tarsomere, about same length as tibial spurs; (Adams), niger (Currie), nigrescens (Stange), mesonotum without blade-like setae; scutelli with nigrihbris (Hagen), nivatensis (NavAs), pallidus welldeveloped white setae posteriorly; hindwing as (Banks),peninsulanis (Banks), peregrinus (Hagen), long as forewing, in repose wing apices nearly 88 Insecta Mundi coincide; pilula axillaris 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 simple; posterior area of hindwing narrower ciated with its microhabitat. The absence of thread- than presectoral area before radial sector; forewing like setae on the abdominal tergites may be a larval vein CUP+ 1Aonly runs along posterior fork of CuA character from the A. modesta Group. There are for a short distance (in contrast to Ameromyia), four undescribed species, one of which is described interconnected by about 4 crossveins; hindwing below. CuA runs along posterior fork of medial fork for a short distance; radial sector originates a little basad Venezueleon guaricus Stange, new species of medial fork in hindwing, about 3-4 presectoral crossveins; postventral lobe of male ectoproct mod- Holotype male: Rio Orituco, 15 km S Calabozo, erately developed, less than twice as long as median Guarico, Venezuela, February 27,1988, R. B. Miller diameter; sternite IX much wider than long, not & L. A. Stange (FSCA). produced medially; male genitalia highly moddied, gonarcus broadly expanded, plate-likelaterally with Adult Description: Length ofbody 32 mm; length lateral tooth; mediuncus arched, bifurcate ven- of forewing 20 mm, greatest width 5 mm. General trally; parameres simple, elongate rods; genital sac coloration pale brown with considerable yellow with many strongly clavate setae; female genitalia orange wing suffusion. Head mostly pale brown with pregenital plate membranous; gonapophyseal with darker brown vertex, anterior row of vertex plate elongate; posterior gonapophysis at least 3X scars most prominent; frons with transverse light longer than median diameter; ectoproct and lateral brown stripe near middle; antenna nearly all pale gonapophysis with strongly flattened digging 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 (based onguaricus): Man- stripe laterad of which somewhat darker brown dible as long as head capsule, distance between than laterally, setal bases darker brown; teeth more than that between base of mandible and mesoscutum and metascutum pale brown with basal tooth; head capsule as wide as long (measured darker brown sublaterally, scutelli with faint ventrally); dorsal and ventral surface of head cap- sublateral dark stripe; pleura pale brown, dark sule with dolichasters; mesothoracic spiracle borne brown stripe on upper episternum; legs pale brown on tubercle that is longer than basal width; abdomi- except darker brown ventrally; femora and tibiae nal tergites without thickened setae or thread-like with dark brown stripe on mesa1 margin; tarsi dark setae; abdominal spiracles not enlarged; sternite brown ventrally; wings with veins mostly pale VIII with submedian teeth that are longer than brown, membrane suffused with yellow orange; basal widths. forewing with rhegmal mark along posterior fork of CuA near hind margin, several crossveins espe- Distribution: Venezuela. cially in radial sector, near cubital fork, and crossveins at margins along costal area and poste- Discussion: This genus appears to be closely rior area; posterior vein white. related to Ameromyia, especially in regards to the similar male genitalia, female terminalia (lateral Frons with setae; pronotum with outstanding gonapophyses at least partially fused), widely sepa- white bristles along anterior, posterior, and lateral rated an tennal fossae, and fron tal setae. Leg struc- margins; a few medially; scutelli with outstanding ture and wing venation offer significant differ- white bristles posteriorly; mesoscutum with nu- ences. The pretarsal claws are weakly developed merous white bristles; legs without femoral sense and the femoral sense hair is absent in Venezueleon. hair and nearly without outstanding setae, a few on Forewing vein CUP + 1A only parallels posterior femora and tibiae are shorter 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 Ameromyia modesta Group in having mesothoracic pale brown setae; those on sternites I-V twice as spiracle borne on an elongate tubercle, the rela- long as those on corresponding tergites, posteriorly tively wider head capsule, and the presence of setae become of equal lengths; postventral lobe of dolichasters on the ventral surface of the head Vol. 8, No. 1 - 2, March - June, 1994 89 male ectoproct with about 4 dark brown setae, Gnopholeontini Stange, New Tribe mostly apically. Greatest ocular width about two-thirds Adult Description: Labial palps short, interocular distance; antenna with 30flagellomeres; palpimacula not extending to apex; metafemur distal palpomere of labium not swollen, 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 CUP originates distad of as wide as long; proleg and mesoleg about equal in crossvein m-cu; forewing vein 2A runs in a fairly length, metaleg the longest; basitarsomere of even curve toward 3A; pilula axillaris present, metaleg about 3X longer than diameter, somewhat usually well developed; hindwing vein CuA runs longer than that of mesoleg; distal tarsomere about close to hind margin, ends near medial fork; as long as tarsomeres 2-4 together; pretarsal claw 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, apex asymmetrical; forewing costal cellslonger paramere plate-like; female anterior gonapophysis than high; forewing with 3 presectoral crossveins; plate-like or membranous; posterior gonapophysis CUP + 1A runs along posterior fork of CuA short inflated; pregenitalplate with tooth; gonapophyseal distance, interconnected by 3 crossveins; hindwing plate large; female ectoproct with digging setae. radial sector originates at about level of medial fork, 4 presectoral crossveins; hindwing posterior Larval Description: Mandibular bases close to- area narrower than prefork area; CuA runs along gether, separated by about basal width of mandible; posterior fork of MP, for short distance, intercon- mandible with 3 teeth, middle tooth closer to distal nected by 4 crossveins. Abdomen much longer than tooth than basal tooth; distal tooth not shorter than wings; segment VIII about 3X longer than high; middle tooth; abdomen without dolichasters or male ectoproct with postventral lobe about as long 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 Female: About as described for male. Abdomen Sonoran Region. Larvae are either free living shorter than wings; setae on basal abdominal ster- (Tyttholeon), live in protected areas (rock over- nites about equal to those on basal abdominal hangs) (Menkeleon) or live on rocks or tree trunks tergites; pregenital plate (Fig. 21) membranous but (Gnopholeon). The larvae are unknown for median tubercle present; posterior gonapophysis Maracandula, but evidence suggests they are asso- about 3X longer than wide, without digging setae; ciated with rocks. The principle synapomorphy of digging setae on ectoproct andlateralgonapophysis the genera in this tribe is the close approximation well developed; gonapophyseal plate small. of the larval mandibles. The inflated posterior gonapophyses (Figs. 31, 32) function as egg han- Paratypes: 9 males, 9 females, same data as types dlers rather than egg coaters according to Miller (CAS, USNM, FSCA, Miller collection); 3 males, 1 (1990). Even more modifiedposterior gonapophyses female, 12 km S Calabozo, Guarico, Venezuela, are found in the Lemolemini. Some species (espe- Feb. 6- 12, 1969, P. Spangler (FSCA; USNM). cially Maracandula) are afternoon fliers. Discussion: There is some variation in this spe- cies at the type locality. The degree of wing suffu- Key to the genera of Gnopholeontini sion varies andsmaller specimens appear to have a Adults relatively shorter and broader wing. There are 1. Tibial spurs present (Fig. 39) ...... evidently 3 additional undescribed species, all ...... Gnopholeon Stange known from single specimens from Venezuela. They Tibia1 spurs absent (Fig. 40) ...... 2 differ in wing shape and coloration. Larvae were discovered at the type locality. They live in shallow 2. Wings narrow with sparse venation; male ectoproct sand, anchoredto bedrock. Two adults were reared produced mesally below (Fig. 50); posterior to confirm the association. gonapophysis of female subcylindrical; prege- Insecta Mundi

nital plate very large (Fig. 30) ...... Adult Description: Legs short and relatively ...... Tyttholeon Adams stocky, profemur less than 1.5X longer than procoxa; Wings broad with denser venation; male ectoproct hindbasitarsus much shorter than distal tarsomere; not produced mesally; posterior gonapophysis femoral sense hair elongate, one on proleg and of female 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 flagellomere 3 at least 1.5X longer than banksian lines absent; hypostigmatic cell without wide (Fig. 36); cubital fork of forewing and crossveins; posterior area of hindwing narrow, CuA medial fork of hindwing arise near middle of bends to hind margin near medial fork; pilula wing; pretarsal claws more than 2X longer than axillaris well developed with setae covering most of greatest diameter of apical tarsomere; female dorsal surface of knob; male genitalia with simple lateral gonapophyses fused (Fig. 3 1)...... gonarcus, free parameres; male ectoproct simple, ...... Maracandula Currie postventral lobe weakly developed at most; female Antennal flagellomere 3 about as long as wide; cubital fork of forewing and medial fork of terminalia with posterior gonapophysis swollen hindwing arise well before middle of wing; thumblike; pregenital plate narrow with median pretarsal claws not much longer than greatest tooth; gonapophyseal plate expansive. diameter of apical tarsomere (Fig. 40); female lateral gonapophyses not fused ...... Larval Description (based on barberi): Head ...... Menkeleon Stange capsule greatly modified, much wider than long, greatly produced anterolaterally into a pair of ro- ~arvae bust stemmatal tubercles which bear long setae 1. Mesothoracic sternum with specialized patch of which camouflage mandibles in trap position; man- lanceolate setae at middle (Fig. 71); labial pal- dible elongate, narrow, with three closely spaced, pus very long, distal palpomere beyond sensory equidistant teeth far removed from base; mesotho- area more than 5X longer than wide; head capsule subquadrate, posterolateral corners racic spiracle not borne on tubercle; meso- and produced laterally (Fig. 71); live in open sand metathoracic as well as abdominal segments I-VIII ...... Tyttholeon Adams each with a bilaterally symmetrical pair of large Metathoracic sternum without specialized patch scoli; pretarsal claws of hindleg no longer than of setae; labial palpus short to moderately long, those of mesoleg; submedian teeth on sternite VIII distal palpomere less than 3X longer than wide longer than basal widths. beyond sensory area; head capsule various, but not much produced at posterolateral corners .. Included Species: barberi (Currie), delicatulus (Currie), and zapotecus Stange. 2. Abdomen with scoli (except 1st instar); head greatly produced at anterolateral areas which Distribution: Deserts of southwestern United bear stemmatal 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 equidistant; on tree bark or bare rock mental differences from all other genera in the faces ...... Gnopkoleon Stange tribe except for Menkeleon. The small pregenital Abdomen without scoli; head not greatly pro- plate is a notable difference from Tyttholeon; and duced and without long lateral setae; maximum the separate lateral gonapophyses, smaller poste- jaw gape 200°; mandibular dentition with middle tooth much closer to distal tooth than basal rior gonapophysis, stockier legs with one elongate tooth; under rock overhangs ...... femoral sense hair, sparser wingvenation, andwell ...... Menkeleon Stange developed tibial spurs are important differences from Maracandula. The well developed tibial spurs 13. Gnopholeon Stange are a key diagnostic trait readily differentiating (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- Gnopholeon Stange, 1970. Univ. Calif. Pub. Ent. 55:148. neric importance in the related Brachynemurini. Type Species: Gnopholeon zapotecus Stange, by 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 lack of bging setae on the female ectoproct of the truncate process near origin of weakly developed G. delkatulus Group is an apomorphy of that postven tral 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 tremely enlarged and swollen; lateral gonapophyses being deferred until the larval stage of G. delicatulus fused; pregenital plate narrow with median tooth; is known. The larva of the G. barberi Group is one gonapophyseal plate expansive. of the most bizarre larval types known in the family Myrmeleontidae and resembles in some ways the Larval Description: Unknown. larvae of Ascalaphidae. The hind pretarsal claws are not enlarged as in other antlions and the Included Species: apicalis (Banks) andpygmueus mandibular bases are the most approximated of (Hagen). any known antlion 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 instars. The larva 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. The female genitalia arehighly that the larva lives on tree trunks due to its camou- 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 in the Myrmeleontidae of the Western Hemisphere, Maracandula Currie, 1901. Proc. ent. Soc. Wash. 4:436. although some individuals of Menkeleon bellula Type species: Myrmelwn pygmaeus Hagen, by approach this condition. Also of unusual interest is original designation. Banks 1927. Bull. Mus. Comp. the extreme reduction of the eyes and highly modi- Zool. 68:58 (further description). Stange 1970. Univ. fied antenna in the described species Calif. Publ. Ent. 55:154 (further description). (autapomorphies). The closest relative is the genus Menkeleon. The female genitalia of Maracandula =Microleon Banks, 1901. Trans. Amer. ent. Soc. 27:365. appear more specialized than in Menkeleon with Type species: Microleon apicalis Banks, by origi- the lateral gonapophyses fused and the posterior nal designation. Preoccupied by Microlean Butler, 1885. gonapophysis greatly enlarged. Also, the subapically broadened wings of Maracandula are distinctive. =Mimoleon, Banks 1913. Trans. Amer. ent. Soc. 39:226. However, in 2 undescribedspecies ofMaracandula; New name for Microlwn Banks, 1901. leg and wing characters are more similar to Menkeleon. From the spurless genus Tyttholeon, Adult Description: Antennae with flagellomeres the small pregenital plate, differently formed male 2-4 much longer than wide; legs moderately long ectoproct, and wing venation offer important differ- and slender, profemur more than 1.5X longer than ences. There are two described species and several procoxa; hind basitarsus shorter or longer than undescribed ones. The adults fly in the afternoon. distal tarsomere; femoral sense hair small and hardly distinguishable from surrounding bristles, 1 5. Menkeleon Stange 1970:154 one each on proleg and mesoleg; pretarsal claws (Figs. 40, 70) weakly developed; tibia1 spurs absent; wings broad with dense venation; banksian lines absent; Menkeleon Stange, 1970. Univ. Calif. Pub. Ent. 55: hypostigmatic cell with or without several Type Species: Maracandula bellula Banks, by crossveins; posterior area of hindwing narrow, CuA original designation. bends to hind margin near medial fork; pilula axillaris well developed with setae covering most of Adult Description: Antennae with flagellomeres dorsal surface of knob; male genitalia with rather 2-4 about as long as wide; legs moderately long and simple gonarcus, free parameres; eversible sac slender, profemur more than 1.5X longer than present laterally and ventral to male genitalia; procoxa; hind basitarsus longer than distal mesa1 margin of male ectoproct with short, rather tarsomere; two subequal, rather long femoral sense Insecta Mundi hairs on proleg and mesoleg; wings broad with shorter than hind basitarsus which is either longer 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, CuA bends to hind crossveins; banksian lines absent; hindwing poste- margin near medial fork; pilula axillaris small with rior area narrow, CuA bends to hind margin before setae restricted to apical margin of knob; male medial fork; pilula axillaris moderately well devel- genitalia with rather simple gonarcus, free oped with setae concentrated toward distal margin parameres; male ectoproct rather simple, some- of knob; male genitalia with simple gonarcus, free what produced ventrally, postventral lobe weakly parameres; male ectoproct produced mesally be- developed; female terminalia with posterior low, no postventral lobe; female posterior gonapophysis swollen thumblike; lateral gonapophysis subcylindrical, lateral gonapophysis gonapophyses separate; pregenital plate narrow separate; pregenital plate much enlarged with a with median tooth; gonapophyseal plate expansive. prominent median process posteriorly; gonapophyseal plate large. Larval Description (based on bellulus): Head capsule wider than long, weakly produced Larval Description (based on puerilis): Head posterolaterally; labial palpus moderately long, dis- capsule wider than long, producedposterolaterally, talpalpomere less than 3X longer than wide beyond with dolichasters;labial palpus extraordinarilyelon- sensory area; mesothoracic spiracle borne on tu- gate; distal palpomere beyond sensory area more bercle; mesosternum without specialized setalpatch than 5x longer than wide; mesothoracic spiracle at middle; abdomen without scoli. borne on elongate tubercle; mesothoracic sternum with specializedlanceolatesetae at middle; pretarsal Included Species: Only bellulus (Banks). claws of hindleg about 2x longer than that of mesoleg; abdomen without scoli; submedian tooth of sternite Distribution: Southwestern Unitedstates, north- VIII longer than basal width. ern Mexico. Included Species: puerilis Adams. Discussion: The female terminalia of Menkeleon are quite similar to Gnopholeon but Menkeleon Distribution: Mojave-Colorado Desert; Great lacks tibial spurs. Superficially, the species of Basin; Baja California. Menkeleon resembles those of Maracandula but generic distinctions are 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 autapomorphic. The larva was found under an (Fig. 7 1) are autapomorphic. The larva (Fig. 65) is overhanging rock. also very distinctive with the development of lan- ceolate setae on the metasternum (autapomorphic). 1 6. Tyttholeon Adams Other distinctive characters are the loss of tibial (Figs. 30,50, 57, 71) spurs (present in Gnopholeon) and narrow wings (broad wings in Maracandula and Menkeleon). Qttholeon Adams, 1957. Psyche 63: 106. Type Species: The larva lives in the sand. One undescribed Tyttholeon puerilis Adams, by original designation. species is found in southern Baja California. Taxonomy: 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 CUPoriginates distad of base; procoxa; hindleg a little longer than proleg; profemur forewing vein 2A runs a fairly even curve toward without clavate setae; profemoral sense hair much 3A; pilula axillaris present; hindwing as long as less than one-half length of profemur, 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 hindwing vein CuA runs close to hind margin, ends Pretarsal claws longer than length of proleg near medial fork; hindwing radial sector originates basitarsus; profemoral sense hair at least 112 well before medial fork, usually 2 presectoral length of femur; antennal fossae separated by crossveins; abdomen without hair pencils, shorter not more than width of pedicel; female posterior gonapophysis shorter than height of ectoproct than wings, about same length in both sexes; male or strongly curved mesally (Figs. 23, 24, 26, 34) ectoproct without postventral lobe; male paramere plate-like; female anterior gonapophysis plate-like or membranous, posterior gonapophysis highly 2. Forewing costal area biareolate in part; hindwing modified, either swollen or strongly curved; shorter than forewing, in repose apices of wings gonapophyseal plate usually very large (except nearly coincide; male ectoproct with large ven- Elicura and Lemolemus); female ectoproct without tral process; female gonapophysis evenly swol- digging setae. len for complete length, with apical scraping setae (Fig. 25) ...... Jaffuelia Forewing costal area with single series of cells; Larval Description: Mandibular bases far apart, hindwing as long as forewing, in repose apex of separated by much more than basal width of man- hindwing extends beyond that of forewing; male dible; mandible with 3 parallel teeth, middle tooth ectoproct without ventral 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 scraping setae (Fig. 27) ...... Neulatus midpoint of mandible; abdominal segments with dolichasters or tufted setae along midline, some- 3. Male ectoproct with indication of postventral lobe; times with lateral scoli; sternite VIII with submedian female posterior gonapophysis longer than teeth. height of ectoproct, strongly curved mesally (Fig. 23, 34); Galapagos Islands; southwestern Ecuador ...... 4 Discussion: This tribe is apomorphically defined Male ectoproct without indication of postventral by the specialized setae found medially on the lobe; female posterior gonapophysis shorterthan abdominal tergites of the larva. The adults have height of ectoproct, not strongly curved mesally one apomorphic character, the lack of digging setae (Figs. 24, 26); Central Chile; Argentina; Uru- on the female ectoproct. This is evidently a relict guay; southern Brazil ...... 5 group without close relatives. Oddly, although some larvae (Lemolemus, Sical, Elicura) 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 of promi- nent setae (Fig. 33) ; male ectoproct mesally none has invaded the sand dune habitats which emarginate ventrally (Fig. 35); Western Ecua- abound in Central Chile. This tribe is most diverse dor ...... Ecualeon in Chile where 5 of the 7 genera occur. Three Forewing posterior area highest near end of vein genera, Elicura, Lemolemus, and Sical are closely 2A; female tergite VII without subapical row of related, but the other four genera are isolated. prominent setae; male ectoproct not emargin- Elicura reaches Argentina and southern Brazil. ate mesally; Galapagos Islands ...... Ecualeon, found in western Ecuador, is assigned to ...... Galapagoleon the tribe based on a presumed larva which agrees with Lemolemini and adult characters. 5. Profemur with long white bristles along exterior Galapagoleon is unknown in the larval stage and is face; forewing costal area broad, area above tentatively referred to the Lemolemini since the cubital fork nearly 3X as high as subcostal area, female terminalia are similar to Ecualeon. often triareolate; mesofemoral sense hair about 112 length of femur; female terminalia with Key to genera of Lemolemini anterior gonapophysis plate-like (Fig. 24); pre- Adults genital plate much longer than wide; southern 1. Pretarsal claws shorter than length of proleg Chile, Argentina; Uruguay; southern Brazil ... basitarsus; profemoral sense hair about 114 ...... Elicura 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 ectoproct, 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 hair about 114 length offemur; female terminalia Insecta Mundi

with anterior gonapophysis lobe-like (Fig. 26); Adult Description: Antennalfossae separatedby pregenital plate wider than long; Central Chile about width of scape; distal palpomere of labium moderately swollen;profemur without white bristles on exterior face except 1or 2 subapicalones; procoxa 6. Basitarsus of metaleg less than 3X longer than with 2 white bristles anteriorly and posteriorly, wide; forewing costal area with few, if any, interconnected crossveins in basal halt prege- setae more than one-halfcoxal diameter; profemoral nital plate large, without median tooth ..Sical sense hair about one-halflength of profemur, about Basitarsus of metaleg at least 4X longer than as long as that of mesofemur; pretarsal claws mod- wide; forewing costal area with most crossveins erately developed, longer than pro-basitarsus but interconnected; pregenital plate small, with shorter than tibial spurs which are longer than median tooth ...... Lemolemus meta-basitarsus; tibial spurs of same length in both sexes; forewing costal area biareolate in about Larvae distal one-fourth to one-third; forewing radial sec- (Ecualeon and Galapagoleon unknown) tor arises well basad of cubital fork, in contrast to 1. Abdominal segments I-VII with well developed position of hindwing radial sector which arises only scoli, at least longer than wide (Figs. 62, 63); little basad of medial fork; hindwing usually with 4 abdomen without median row of tufted black setae but with roseate, squat dolichasters; presectoralcrossveins; forewing posterior area high- pretarsal claws with basal tooth ...... 2 est near cubital fork; anterior banksian line absent; Abdominal segments I-VII without scoli, or if hindwing nearly as long as forewing, in repose short scoli 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 instar), without roseate posteriorly; male abdomen much longer than wings, dolichasters; pretarsal claws without basal tooth segment VIII at least 4X longer than high; scale- ...... 3 like setae on segments 111-V; male ectoproct bilobed 2. Abdominal scoli at least 3X longer than wide, ventrally on mesa1 margin (Fig. 35), postventral tapered to apex, without dolichasters (Fig. 62); head longer than wide; live in Puya plants ..... lobes indicated; gonarcus nearly transverse, ...... Neulatus Navls mediuncus large, smooth, except doubly ridged Abdominal scoli less than 2X longer than wide, apically, paramere a flat sclerite, with lateral pro- broadened apically, with dolichasters (Fig. 63); cess (Fig. 35); female with large, transverse prege- head wider than long; live on boulders ...... nitalplate with minute medial tooth; gonapophyseal ...... Jaffuelia Nav4s plate large and expansive; posterior gonapophysis swollen and curved subapically; anterior 3. Abdominal segments I-VII with weak scoli; head gonapophysis swollen, transverse, with many elon- capsule and scoli with abundant silky white, gate setae; ectoproct with digging setae, those on hairlike setae ...... Sical Nav4s Abdominal segments I-VII without scoli; head lateral gonapophysis moderately developed; stern- capsule and scoli without silky, white, hairlike ite VII not produced medially; tergite IX of female 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 scolus of mesothorax 2X longer than Distribution: Southwest Ecuador wide ...... Elicura Navhs Mandible pale with dark brown areas mostly at Discussion: This genus is provisionally assigned base; anterior scolus of mesothorax 1.5X longer to the Lemolemini. It appears related to than wide ...... Lemolemus Navls Galapagoleon especially in regards to the curved posterior gonapophysis of the female terminalia. The structure of the female abdomen suggests that 17. Ecualeon Stange, New Genus this genus may throw eggs similar to that observed (Figs. 33-35) in Psammoleon which has similar female 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 of such present designation. behavior. Another significant character of this genus is the broadening of the posterior area near the cubital fork. Quite unusual and unique, at least Vol. 8, No. 1 - 2, March - June, 1994 among New World antlions, is the development of Frons without setae; clypeus with transverse large, scale-like setae on male abdominal segments row of 4 long, black setae; scattered decumbent III-V. The male ectoproct (Fig. 35) is emarginate setae on vertex; labium 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, black ones mostly Ecualeon ovispargus Stange, New Species lateral, white ones medial; prescutum with 3-4 black or white setae in a row, mesonotum with 3-5 Holotype male: 7 km. S. Piiias, Province El Oro, white setae in a row near middle; mesoscutellum Ecuador, July 7,1989, R. Miller & L. Stange (Florida with sublateral white seta near posterior margin (2 State Collection of Arthropods). pairs in some paratypes); procoxa with 2 white bristles anteriorly and posteriorly, posterior ones Adult Description: Length ofbody 34mm; length somewhat longer than coxal width, anterior ones of forewing 22 mm, greatest width 6 mm; general shorter; profemur without bristles on exterior mar- coloration yellowish brown, wing membrane trans- gin except a few subapical ones; closing face of all parent except small brownish suffusion at nearly femora and tibiae and exterior face of tibiae with every vein juncture. Head color variable, from pale many elongate bristles that are mostly twice as long to dark brown; anterior row of vertex scars most as femur or tibia, black bristles on exterior face; 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 setae very long and ubiquitous, more than 6X prominent dark brown lateral scar, middle scar longer than vein diameter; abdomen with long and lighter brown; clypeus and labium 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 spot beneath antennalfossa; postgena nearly which on sternites are 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 nearly all brown, scape pale brown posteri- interocular distance; antenna with 30flagellomeres; orly and less so anteriorly; pedicel and base of flagellomere 1-13 about 1.5X longer than wide, flagellomere 1 dark brown; pronotum mostly pale abruptly broadening beyond so that flagellomere brown medially, darker brown laterally with nu- 20 about 3X broader than long; distal palpomere of merous dark brown spots at setal bases; mesonotum labium somewhat swollen, circular sensory area a and metanotum similar to pronotum with pale little beyond midpoint of palpomere; legs moder- brown medially, darkbrown laterally; scutellimostly ately long, mesoleg about 0.5X longer than proleg pale brown; mesopleuron dark brown anteriorly, but shorter than hindleg; basitarsus of proleg and mostly pale brown posteriorly; metapleuron mostly mesoleg about 3X longer than diameter, about one- pale brown with scattered dark brown areas; coxae half length of distal tarsomere; hind basitarsus and mera mostly pale brown, meso- andmeta-coxae about 4X longer than diameter; tibial spurs reach darker brown posteriorly; femora and tibiae pale nearly to apex of tarsomere 2 of pro- and meso- brown with dark brown spots at setal bases and tarsus, and somewhat beyond apex of metatarsus; apically, especially profemur and mesofemur; tarsi pretarsal claws shorter than tibial spurs, about as pale brown with dark brown mostly on tarsomeres long as pro-basitarsus; hindwing nearly as long as 4 and 5 and apex of distal tarsomere; tibial spurs forewing, in repose apex of hindwing projects be- and claws reddish brown, lighter distally; forewing yond that of forewing; wings all about same shape and hindwing nearly equally colored; wing veins but hindwing narrower; forewing costal area pale brown with extensive dark brown sections biareolate in distal one-half, cells higher than wide near vein and crossvein junctures; stigma pale in basal one-half; forewing with radial sector aris- white preceded by relatively large dark brown spot; ing well before cubital fork, 5-6 presectoral small dark brown rhegmal spot; abdomen brown, crossveins; CUP+ 1A runs along posterior fork of pale brown areas mostly at base and less at middle CuA for short distance, interconnected by 3 of both tergites and sternites. 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 terior area narrower than prefork area, highest Adult Description: Antennalfossae separated by near distal end; CuA runs to hind margin just about width of pedicel; profemur with row of white before medial fork; pilula axillaris large; abdomen bristles on exterior face; profemoral and much longer than wings; segment VIII about 4X mesofemoral sense hair more than one-half length longer than high; male ectoproct with postventral of femur; pretarsalclaws longer than pro basitarsus; lobe weakly indicated, bifurcate mesally (Fig. 35); tibia1 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 ally produced. about same length as forewing, in repose apex of hindwing extends beyond that of forewing; 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 of long, posteriorly directed medially; male genitalia with arched gonarcus, bristles; terminalia (Fig. 33) with posterior mediuncus short, not ridged; paramere longer than 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- gonapophysis swollen, beset with setae; lateral tized, longer than wide, with medial tubercle; pos- gonapophysis elongate, digging setae about as long terior gonapophysis swollen, much shorter than as width; ectoproct with row of 4 dorsally directed height of ectoproct; ectoproct and lateral bristles; pregenital plate sclerotized, transverse, gonapophysis without digging setae; anterior with small median tubercle. gonapophysis plate-like.

Paratypes: 3 males, 20 females, same date as Larval Description (based on litigator): Head holotype (MC, USNM, Quito, SC, FSCA). capsule longer than wide, with dolichasters; man- dible 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; abdominal scoli absent; abdomen with disturbed they fell to the base of the grass where median row of tufted black setae; pretarsal claws detection was difficult. None was attracted to without basal tooth; mesothoracic spiracle borne on lights but one was found by jack lighting. An tubercle that is longer than wide; abdominal spi- unidentified larva collected under a rock overhang racles enlarged, borne on tubercles. in Ecuador may be this species. It has specialized setae along middle of abdomen and abdominal scoli. Included Species: iniquus (NavAs), justus (NavAs), litigator NavAs.

18. Elicura Navas Distribution: Southern Chile, Argentina; Uru- (Fig. 24) guay; southern Brazil. Discussion: Stange (1967) placed some of the Elicura NavPs, 1911. Rev. chilena Hist. nat. 15:126. Type Speciea: Elicura litigator NavBs, by original species of Elicura in Lemolemus which this genus designation. closely resembles. One notable difference between Elicura and Lemolemus is that the anterior =Plater Navls, 1919. Rev. Acad. civ. Madrid 17:297. gonapophysis is lobe-likein Lemolemus. Theforew- New synonym. Type Speciea: Plater iniquus ing costal area is very broadin Elicura and provides Navls, by original designation. one character to separate it from all others in the tribe. Larvae of E. litigator were found in many =Bridarollus Navls, 1933. Rev. Acad. Cienc. Zaragoza habitats such as rock overhangs, tree holes, and at 16:90. New synonym. Type Species: Bridarollus bases of trees and resemble Nemoleontini and solers NavPs, by original designation. Dendroleontini in mandibular dentition. The dark spotting of the mandible is distinctive in the tribe. Vol. 8, No. 1 - 2, March - June, 1994

19. Jaffuelia Navas a few species of Lemolemus. The posterior (Fig. 25, 63) gonapophysis has scraping setae which are associ- ated with oviposition on rocks. Forewing vein CUP Jaffuelia Navhs, 1918. Bol. Soc. aragonese 17:217. originates close to basal crossvein which is a Type species: Jaffuelia chilensis Navis, by origi- plesiomorphic condition approaching the Old World nal designation. Myrmecaelurini. Structurally, it appears most closely related to Neulatus. Larvae of this genus Adult Description: Antennalfossae separated by were found on boulders. They have abdominal scoli more than width of pedicel; distal palpomere of and odd medial processes bearing dolichasters on labium slender; profemoral sense hair short, less abdominal segments I-IV. Most of the body is than one-fourth length of femur, equal to that of covered with scale-like setae. mesofemur; pretarsal claws shorter than meta- basitarsus, about same length as tibial spurs; tibial spurs the same length in both sexes; forewing costal 20. Galapagoleon Stange, New Genus area mostly biareolate; forewing radial sector arises (Fig. 23) well basad of cubital fork about equivalent to the distance between radial sector and medial fork in Type Species: Brachynemurus darwini Stange hindwing; anterior banksian line weakly devel- 1969, by present designation. oped; hindwing shorter than forewing, in repose wing apices nearly coincide, male sternite IX trans- Adult Description: Antennalfossae separated by verse; male ectoproct with medial lobe ventrally; less than width ofscape; distal palpomere of labium male genitalia with arched gonarcus, short swollen; profemur without white bristles on exte- mediuncus, parameres plate-like; female terminalia rior face except 3 subapical ones; procoxa with only with posterior gonapophysis evenly swollen for 2 white bristles posteriorly that are less than one- complete length; with scraping setae; ectoproct and half coxal diameter; profemoral sense hair about lateral gonapophysis without digging setae; prege- three-fourths length of femur, over twice as long as nital plate sclerotized with median ridge; sternite mesofemoral sense hair; pretarsal claws well devel- VII not produced medially; tergite IX not produced oped, longer than basitarsus but shorter than tibial ventrally; anterior gonapophysis plate-like. spurs; tibial spurs of same length in both sexes; forewing costal area simple, except 1 or 2 in tercon- Larval Description (based on chilensis): Head nec ted crossveins distally; forewing radial sector wider than long, with many squat, dark dolichasters arises well basad of cubital fork, in contrast to on dorsal surface similar to those on rest of dorsum; position of hindwing radial sector which arises a thoracic and abdominal scoli well developed al- little basad of medial fork; hindwing with 2 though less than 2X longer than wide, with many presectoralcrossveins; forewing posterior area high- dolichasters; abdomen with median row of clumped est near end of vein 2A; anterior banksian line dolichasters; pretarsal claws with basal tooth; me- developed; hindwing shorter than forewing, in re- sothoracic spiracle borne on tubercle that is wider pose wing apices nearly coincide; male sternite IX than long; abdominal spiracles enlarged, but not large, somewhat wider than long, bearing long much raised. setae; ectoproct simple but postventral lobe indi- cated, without processes; male genitalia with Included Species: chilensis NavBs andporterina gonarcus a simple arch, mediuncus somewhat longer NavBs. than wide with ridges; paramere with hook; female terminalia with large sclerotized pregenital plate Distribution: Central Chile. which lacks medial tooth; gonapophysealplate large and expansive; posterior gonapophysis swollen and Discussion: This genus has several autapomorphic curved subapically ; ectoproct without digging se- characters in the tribe. The shape of the male tae, those on lateral gonapophysis only moderately ectoproct which has a mesa1 process ventrally is developed; sternite VII not produced medially; terg- distinctive and the long, evenly swollen female ite IX of female not produced ventrally; anterior posterior gonapophysis with scraping setae (Fig. gonapophysis swollen; female abdominal tergite 25) is diagnostic for that sex. The biareolate forew- VII without posterior row of bristles. ing costal area is duplicated only in Elicura and in Insecta Mundi

Larval Description: Unknown. tively short; ectoproct and lateral gonapophysis without digging setae. Included Species: darwini (Stange). Larval Description (based on modestus and Distribution: Galapagos Islands. several unnamed species): Head capsule longer than wide, with dolichasters; mandible nearly uni- Discussion: This genuscontains one species found form color except darker basally; anterior mesotho- in the Galapagos Islands. Its taxonomic position is racic scolus 1.5X 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 scoli; abdomen with median row of tufted gonapophysis is similar to Ecuakon. The discovery black setae; pretarsal claws without basal tooth; of the larval stage will probably clarify its relation- mesothoracic spiracle borne on tubercle that is ships. longer than wide; abdominal spiracles enlarged, borne on tubercles. 2 1. Lemolemus Navas (Fig. 26) Included Species: modestus (Blanchard)

Lemolemus NavBs, 1911. Rev. chilena Hist. nat. 15123. Distribution: Central Chile. Type species: Lemolemus nectator NavBs, by origi- nal designation (= L. modestus (Blanchard) 1851). Discussion: This genus is in need of revision. One taxonomic problem is that the males and females of =Puren Navls, 1911. Rev. chilena Hist. nat. 15:125. the same species are variable in the length of the New synonym. Type species: Puren bellator tibial spurs. The lobe-like anterior gonapophysis Navls, by original designation (= L. modestus found in this genus is remarkable and points to a (Blanchard) 1851). possible relationship with Dendroleontini. An autapomorphy is the sexual dimorphism in the =Licura Navls, 1918. Bol. soc. aragonese 17:330. New synonym. Type species: Licura ferus NavBs, 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.

Adult Description: Antennalfossae separated by 22. Neulafus Navas about width of pedicel; distal palpomere of labium (Figs. 27, 64-65) slender; profemur without white bristles on exte- rior face except subapically; profemur sense hair =Lincoya Navls, 1911. Rev. chilena Hist. nat. 15:121. moderately long, about one-half length of femur, Type species: Lincoya dealbatus Navls, by origi- about equal to that of mesofemur; pretarsal claws nal designation. Preoccupied by Lincoya Kirby, longer than pro-basitarsus, shorter or longer than 1871 (Lepidoptera) tibial spurs; tibial spurs usually shorter in males; forewing costal area simple or biareolate; forewing Neulatus Navls, 1912b. Arxivs Inst. Ciencies 1:108. radial sector arises well basad of cubital fork, about Type species: Glyptobasisporteri Brethes, by origi- nal designation. equivalent to the distance between radial sector and medial fork in hindwing; hindwing about same =Nilcoya NavBs, 1913. Rev. chilena Hist. nat. 17:243. length as forewing, in repose hindwing extends New Synonym. New name for Lincoya Navhs beyond forewing; anterior banksian line of forew- ing weakly developed or absent; male sternite IX Adult Description: Antenna1fossae separated by much wider than long, weakly produced medially; more than width of pedicel; distal palpomere of male ectoproct simple, without lobes or processes; labium slender; profemur without white bristles on male genitalia with arched gonarcus, mediuncus exterior face; profemoral hair short, less than one- present or absent, not ridged; female terminalia fourth length of femur, equal to that of mesofemur; with sternite VII not produced; tergite IX not pro- pretarsalclaws shorter than meta-basitarsus, about ducedventrally; pregenitalplate sclerotized, wider equal to length of tibial spurs; tibial spurs of equal than long, with median tooth; anterior gonapophysis lengths in both sexes; forewing costal area simple; lobe-like, posterior gonapophysis swollen, but rela- Vol. 8, No. 1 - 2, March - June, 1994 forewing radial sector arises well basad of cubital =Neteta NavPs, 1934. Rev. chilena Hist. Nat. 38:lO. fork, about equivalent to the distance between New synonym. Type species: Neteta barrosi radial sector and medial fork in hindwing; anterior NavBs, by original designation. banksian line weakly developed; hindwing as long =Chibleon NavPs, 1934. Rev. chilena Hist. Nat. 38:lO. as forewing, in repose apex of hindwing extends New synonym. Type species: Chibleonstuardinus beyond that of forewing; male sternite IX trans- NavPs, by original designation. verse; male ectoproct simple, without lobes or pro- cesses; male genitalia with simple arched gonarcus, Adult Description: Antennalfossae separated by mediuncus not developed, paramere short, plate- about width of pedicel; distal palpomere of labium like, ventrally produced; female terminalia (Fig. slender; profemur without white bristles on exte- 27) with pregenital plate large, narrow, posterior rior face except subapically; profemur sense hair end hood-like; sternite VII with narrow median moderately long, about one-half length of femur, lobe; posterior gonapophysis large, apically swol- about equal to that of mesofemur; pretarsal claws len; ectoproct without digging setae; lateral longer than pro-basitarsus, shorter or longer than gonapophysis with numerous short digging setae; tibial spurs; tibial spurs usually shorter in males; tergite IX with median lobe ventrally; anterior forewing costal area simple; forewing radial sector gonapophysis plate-like, gonapophyseal plate ex- arises wellbasadofcubitalfork, about equivalent to pansive. the distance between radial sector and medial fork in hindwing; hindwing about same length as forew- Larval Description (based on porteri): Head ing, in repose hindwing extends beyond forewing; longer than wide, with dolichasters on dorsal sur- anterior banksian line of forewing weakly devel- face similar to those on rest of dorsum; thoracic and oped; male sternite IX much wider than long, abdominal scoli well developed, at least 3X longer weakly produced medially; male ectoproct simple, than wide, without dolichasters; abdomen with without lobes or processes; male genitalia with weak row of clumped dolichasters; pretarsal claw arched gonarchus, mediuncus present; female with basal tooth; mesothoracic spiracle borne on terminalia with sternite VII not produced; tergite tubercle that is longer than wide; abdominal spi- IX not produced ventrally; pregenital plate sclero- racles small, dark, but somewhat elevated. tized, without median tooth; anterior gonapophysis lobe-like; posterior gonapophysis swollen, but rela- Included species: Only porteri (Brethes) (= tively short; ectoproct and lateral gonapophysis dealbatus (Navas) . without digging setae.

Distribution: Central Chile. Larval Description (based onperalinus): Head capsule wider than long with many dolichasters Discussion: The female terminalia (Fig. 27) of this and fine white hair; mandible nearly uniform dark genus are apomorphic and highly modified. Tergite brown; first 3 pairs of thoracic scoli longer than IX has a largeventral lobe not duplicated elsewhere wide; abdominal scoli present but wider than long; in the tribe. Also, the median process of sternite VII 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 and were collectedby Miller, Solervicens, and Stange tooth; mesothoracic spiracle borne on tubercle that (1989) by beating the dead leaves and bracts. The is about as long as wide; abdominal spiracles en- larvae move fairly fast and have elongate abdomi- larged, borne on tubercles. nal scoli (Brethes 1908:15, larva). The type species is based on a larva which was misidentified as an Included species: barrosi (Ntlvas), peralinus ascalaphid. The pale adults rest on Puya plants Navtls, and stuardinus (Nhvas). during the day and are attracted to lights. Distribution: Central Chile. 23. Sical Navas (Fig. 67) Discussion: This genus is sustainedmostly by the larval stage which has moderately developed ab- Sical NavPs, 1928. Rev. chilena Hist. Nat. 3 1:317. Type dominal scoli in contrast to Elicura and Lemolemus. species: Sical peralinus NavBs, by original desig- The female terminalia are most similar to nation. Insecta Mundi

Lemolemus but the pregenital plate lacks the me- Banks, Nathan. 1913b. The genus Brachynemurus dian tooth. The larva lives in soil and carries 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 Autapomorphies for this genus are the long, white neuropteroidinsects. Bulletin Museum of Com- setae on the scoli and the lack of a median tooth on parative Zoology 65:421-455. the female pregenital plate. Banks, Nathan. 1927. Revision of the Nearctic Acknowledgements Myrmeleonidae. Bulletin Museum of Com- The author is grateful to Robert Bruce Miller parative Zoology 68: 1-81. for the photographs of the larvae and for many constructive ideas. John Oswald made a vigorous Banks, Nathan. 1939. On some new and previ- critique of the manuscript. Although the selection ously-knownNeuroptera in the collection of the of the characters used in the cladistic analysis was Academy of Natural Sciences of Philadelphia. done solely by the author, special thanks are due to Notulae Naturae 32: 1-5. David Wahl who spent many hours analyzing data. Thanks are also due to the typists of the manu- Banks, Nathan. 1943. Neuroptera of northern script, Michelle M. Faniola andBrenda J. Lovelace. SouthAmerica. Parts. 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 from Stange (1970), 23 from Stange, (1969) and Fig. Brethes, Juan. 1908. Descripci6n de una larva de 8 (Adams, 1957). Glyptobasis? de Chile. Revista chilena de entomologia 12: 15-17, pl. 11.

References Carpenter, J. M. 1988. Choosing among multiple Adams, Phillip A. 1957. New antlions from the equally parsimonious cladograms. Cladistics southwestern United States. Psyche 6332- 4291-296. 108. Currie, Rollo. 1901. Adwarf ant-lion fly. Proceed- Adams, Phillip A. 1958. Studies in the Neuroptera, ings Entomological Society of Washington 4:435- with special reference to wing structure and 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.Aclassification of the North (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 En tomologist 32: 329-330. Farris, J. S. 1989. Hennig86 reference: documen- Banks, Nathan. 1901b. A list of neuropteroid tation for version 1. 5. Port Jefferson Station, insects from Mexico. Transactions of the Ameri- New York. can Entomological Society 27:361-371. Fraser, F. C. 1952. New species of Neuroptera in Banks, Nathan. 1909. New genera and species of the Museum National &Histoire naturelle, tropical Myrmeleonidae. Journal of the New Paris. Revue Fran~aise8Entomologie 19:55- York Entomological Society 17: 1-4. 64.

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

Henry, Charles S. 1978. An unusual ascalaphid Navas, Longinos. 1919. Algunos insectos larva (Neuroptera: Ascalaphidae) from south- neuropteros de R6publica Argentina. Revista ern Africa, with comments on larval evolution de la real Academia de ciencias exactas, Fisicas within the Myrmeleontoidea. Psyche 85:265- y naturales de Madrid 17:288-305. 274. Navas, Longinos. 1920. Sur des Nevropteres MacLeod, Ellis G. 1970. The Neuroptera of the nouveaux ou critiques (Serie 2). Annales Societe Baltic amber. I. Ascalaphidae, Nymphidae, Scientifique de Bruxelles 39: 189-203. and Psychopsidae. Psyche 77: 147-180. Navis, Longinos. 1922. Insectos Sudamericanos Miller, Robert B. 1990. Reproductive characteris- (Series 4). Revista de la real Academia de tics of some western hemisphere ant-lions ciencias exactas, Fisicas y naturales de Madrid (Insecta: Neuroptera: Myrmeleontidae). Pro- 19:255-267. ceedings of the Third international symposium on Neuropterology (Berg en Dal, Kruger Na- Navas, Longinos. 1924. Insectos de la America tional Park), pp. 171-179. central. Broteria (Serie Zool6gica) 2 1: 55-86.

Navas, Longinos. 1911. Neuropteros chilenos (Se- Navas, Longinos. 1926a. Insecta nova (Series XI). ries 2). Revista chilena de Historia Natural Memorie dell'Accademia pontifica dei Nuovi 15:121-128. Lincei, Rome 9: 101-110.

Navas, Longinos. 1912a. Notas sobre Navas, Longinos. 1926b. Algunos insectos del Mirmeleonidos. Broteria (Serie Zool6gica) 10:29- Brasil (Serib; 3). Broteria (Serie Zool6gica) 97. 2315-15.

Navas, Longinos. 1912b. Sinopsis de 10s Navas, Longinos. 1927. Insecta nova (Series XII). Ascalafidos. Arxius de YInstitut de Ciencies, Memorie dell'Accademia pontifica dei Nuovi Barcelona 1(3):45-143, 11 Figs. , 2 pls. Lincei, Rome 10: 1-10.

Navas, Longinos. 1913. Mirmeleonido e Chile. Navas, Longinos. 1928. Insectos neotropicos (Se- Revista chilena de Historia natural 17:243-244. ries 3). Revista chilena de historia natural 31:316-325. Navas, Longinos. 1914a. Neuropteros sudamericanos (Serie 1). Broteria (Serie Navis, Longinos. 1932. Alcuni insetti 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 Neuroptera from Comparata della R. Universita di Torino (3)42: 1- the United States. Bulletin of the Brooklyn 38. Entomological Society 9: 13-20. Navas, Longinos. 1934a.Algunos insectos de Chile Navas, Longinos. 1915a. Neuropteros nuevos o (6a Serie). Revista chilena de historia natural poco conocidos. Memoria della R. Academia 38:9-12. Ciencias y Artes Barcelona (3) 11:373-398,455- 480. NavLs, Longinos. 1934b. Insectos suramericanos (8 Serie). Revista de la real Academia de Navas, Longinos. 1915b. Neuropteros Ciencias exactas, Fisicas y naturales de Madrid Sudamericanos (Serie 2). Broteria (Serie 31:9-28. Zool6gica) 13:5-13. New, T. R. 1981. A revision of the Australian Navis, Longinos. 1918. Insectos chilenos. Boletin Nymphidae (Insecta: Neuroptera). Australian de la Sociedad aragonesa de Ciencias Naturales Journal of Zoology 29: 707-750, Figs. 1-188. 17:212-230. Insecta Mundi

New, T. R. 1982. A reappraisal of the status of the on Neuropterology. (Berg en Dal, Kruger Na- Stilbopterygidae (Neuroptera: tional Park), pp. 151-169. Myrmeleontoidea). Journal of the Australian 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 reference: documenta- Myrmeleonidae. Proceedings of the Linnean tion for version 1. 1. Trumansburg, New York. Society of New South Wales 40(4):734-751.

Oswald, J. 0. 1993. Phylogeny, taxonomy, and Tjeder, Bo, 1977. Distal abdominal segments and biogeography of extant silky lacewings (Insecta: sclerotized parts of genitalia in Ascalaphidae Neuoptera: Psychopsidae). Memoirs of the (Neuroptera). Annales 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 Transactions of the Entomological Society of tribe Brachynemurini of North 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 of the third International Symposium Vol. 8, No. 1 - 2, March - June, 1994 103

3 4 1221 nnnn Nymphidae 1121

Dimares

Millerleon 10 1122 Myrmeleontini

3 1922 Acanthaclisini 4 7 1118

11 2124 M yrmecaclurini 1428 n n 1728 u u Nesoleontini mn, 0 1 10 14 n u Nemoleontini 0

Brach ynemurini

9 28

16 Dendroleontini

Gnopholeontini 1

Figure 1. One of 2 cladograms showing 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

3 4 1221 Nyrnphidae 1121 28 - -A Nernopteridae ....15811 7 4 91012 7 :7 : Crocinae 1212 6 13 15 2 10112024 Ascalaphidae 111 Z111I

Nernoleontini

3 192224 Acanthaclisini 1118 lo Ll I 172428 21 M yrrnecaelurini

110 Nesoleontini 0 1 I 2; Brach ynernurini

Qf- Dendroleontini 20 Gnopholeontin; 1

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

10 12 10 12 I n I n Nemoleontini I U Nemoleontini 1 I 11 9 29 9 29 n I u I Dendroleontini I Dendroleontini 0 1 0 1 18 34 6 830 18 34 6830 Gnopholeon nln 1 1 ,j C # Gnopholeon 152526 ' ' 152526 ' ' . I. 3 3 111 111 121 Menkeleon 121 Menkeleon A 1Tyttholeon 16 I Tyttholeon Elicura

1331 Lemolemus 1331 Lemolemus n I I 12 19 n 12 19 u Sical Sical 30 ' 30 1417 Jaffuelia

Neulatus 1 1 6 8 25 n nI n n I UUI Austroleon u u I Austroleon 111 I I11 3 4 19 3 4 Abatoleon 4 t3 Abatoleon I 1 I 20 6 8 22 20 6 8 22 n nnm q- nnm Argentoleon UYUI Argentoleon 1112 1112 13 Ensorra 1 6 8 14 Amerom yia Amerom yia

Venezueleon Venezueleon

3 4 11 17 Chaetoleon

Mexoleon

5 1922 A tricholeon

Brachynernurus Brachynemurus

12 Scotoleon 1 Scotoleon 1 G Figure 3. One of 2 weighted cladograms of the genera Figure 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

Nernoleontini INernoleontini I YDendroleontini YAbatoleon

Gnopholeon Austroleon

Menkeleon +Ensorra Tyttholeon IClathroneuria IDejuna I ILernolernus Amerorn yia Venezueleon

Chaetoleon I IGalapagoleon Mexoleon Jaffuelia A tricholeon

Neulatus Brachynemurus IAustroleon Scotoleon 7Dejuna Dendroleontini A batoleon Elicura

Argen toleon Jaffuelia

Lernolernus

aClathroneuria Neulatus - Amerom yia Sical Venezueleon IEcualeon TChaetoleon IGalapagoleon Y Mexoleon IMaracandula Atricholeon Gnopho/eon

Brachynernurus Menkeleon

Scotoleon Tyttholeon

Figure 5. Strict consensus tree of 341 cladograms of the Figure 6. Strict consensus tree of 353 trees showing genera of the Brachynemurini & Myrmecaelurini, relationships of the genera 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 drawing of Scotoleon deflexus (Adams) (male). 108 Insecta Mundi

c1avate Setae

10 - Myrmeleon sp. 11 - Brachynemurus sp.

HYPosfigmolic. .. Cell

C I rodotes

12 - Dejuna mimica

15 - Gnopholeon delicatulus

13 - Brachynemurus divisus

16 - Brachynemurus ferox

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

,* gonapophysis

gonapophysis I '. pregenital plate

17 - Brachynemurus sp. 18 - Scotoleon eiseni

19 - Abatoleon dorsalis 20 - Ensorra verticalis

21 - Venezueleon guaricus 22 - Dejuna persirnilis Figures 17-22. Ventral view of female terminalia. 110 , Insecta Mundi

24 - Elicura litigator 23 - Galapagoleon darwini

<\ ' 26 - Lemolemus nectator 25 - Jaffuelia chilense

27 - Neulatus porteri

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

28 - Nannoleon michaelseni

29 - Myrmecaelurus sp.

32 - Gnopholeon delicatulus 31 - Maracandula apicalis

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

33 - Ecualeon ovispargus 34 - Ecualeon ovispargus

36 - Maracandula apicalis

35 - Ecualeon ovispargus

38 - Mexoleon papago

40 - Menkeleon bellula 37 - Ameromyia nigriventris

Figures 33-40.33) Lateral view of female terminalia; 34) Ventral view of female terminalia; 35) Ventral view of male terminalia and genitalia; 36-37) Frontal view of head; 38-40) Hind tarsus. Vol. 8. No. 1 - 2. March - June. 1994 113

" .. 43 Clathroneuria schwarzi 41 - Atricholeon tuberculatus 42 - Brachynernurus seminolae -

44 - Atricholeon tuberculatus 45 - Dejuna persirnilis 46 - Clathroneuria schwarzi

47 - Venezueleon guaricus 48 - Venezueleon guaricus

49 - Ensorra verticalis 50 - Tyttholwn puerilis

Figures 41-60.41-43)Ventral view of male ectoproct; 44-46)Ventral view of male genitalia; 47) Lateral view of male terminalia; 48) Male genitalia; 49) Lateral view of male terminalia; 50) Posterior view of male ectoprocts. 114 Insecta Mundi

51 - Mexoleon papago 52 - Mexoleon papago

53 Chaetoleon pusillus - 54 - Chaetoleon pusillus

55 - Brachynemurus pulchellus

I 56 - Brachynemurus pulchellus

59 - Maracandula apicalis 57 - Tyttholeon puerilis 58 - Abatoleon dorsalis

Figures 51-59. 51, 53, 55) Lateral view of male terrninalia. 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 6063. Third instar larvae of Brachynemurini. 116 Insccta Mundi

64 - Neulatus porteri 65 - Neulatus porteri

67 - Sical peralinus 66 - Jaffuelia chilense

Figures 64-67. Third inntnr Inwan nf T.nmnlnmini. 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 of Gnopholeontini. 118 Insecta Mundi

APPENDIX 1. Data Matrix for Tribal Relationships

Character Number 1111111111 222222222 123456789 0123456789 012345678

Psychopsidae Nymphidae Nemopterinae Crocinae Ascalaphidae Stilbopterygini Palparini Palparidiini Dimares Millerleon Dendroleontini Nemoleontini Brachynemurini Gncpholeontini Lemolemini Myrmecaelurini Nesdeontini Myrmeleontini Acanthaclisini

Maulini 012111101 1121113000 ????????? Pseudimarini 01 11 11000 0020111000 ?????????

APPENDIX 2: Data Matrix for Brachynemurini, Lemolemini and Gnopholeontini

Character Number 1111111112 2222222223 33 123456789 0423456789 0123456789 01

Nemoleontini 000000000 1010000000 0000000000 00 Dendroleontini 000000000 000000101 0 000000000 1 00 Brachynemurini Abatoleon 001 100001 0000000001 1000000100 00 Ameromyia 10000101 1 0100200001 0010100100 00 Argentoleon 00000101 1 0000000001 1020000100 00 Atricholeon 000010001 0100000001 1031000100 00 Auslroleon 000001011 0000000001 0000010100 00 Brachynemurus 01 OOOOOOI 0100000000 10010001 00 00 Chaetoleon 001100001 0000000100 0101000100 00 Ensorra 000000001 0001000001 1000000100 00 Mexoleon 000000001 0100000000 0101000100 00 Scotoleon 010000001 0110000000 1001000100 00 Venezueleon 100010001 0100100001 0010100100 00 Gnopholeontini Gnopholeon 00120101 1 000001 1010 0000021000 10 Menkeleon 001000101 0000011010 0000021000 00 Tyttholeon 000000101 000001 1010 0000021000 00 Lemolemini Elicura 000000001 0001001000 0000000000 02 Jaffuelia 000000001 0001101110 0000000010 11 Lemolemus 000000001 0001001000 0000000000 02 Neulatus 000000001 0001001011 0000000010 11 Sical 000000001 0001001001 0000000000 12 Unknown Lawae: Clathroneuria 000001001 0000000010 IOOOOO???? ?7 Dejuna 000000001 0000000011 000000???7 ?? Ecualeon 000000001 0001001000 OOOOOO???? ?? Galapagoleon 000000001 0001 001 001 OOOOOO7~7? ?? Maracandula 000000101 000021 1000 000000??77 ?? Vol. 8, No. 1 - 2, March - June, 1994 119

APPENDIX 3. Successive Approximation Weights APPENDIX 5. Successive Approximation Weights for Tribal Analysis for New World Brachynemurini. Generic Analysis with genera added with missing larval data.

APPENDIX 4. Successive Approximation Weights for New World Brachynemurini. Generic Analysis with 5 genera with missing larval data deleted. APPENDIX 6. Successive Approximation Weights for New World Brachynemurini. Generic Analysis of all genera but only adult characters.