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Reclassification of the New World Antlion Genera Formerly Included in the Tribe Brachynemurini (Neuroptera: Myrmeleontidae)

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Reclassification of the New World Antlion Genera Formerly Included in the Tribe Brachynemurini (Neuroptera: Myrmeleontidae) Center for Systematic Entomology, Gainesville, Florida Insecta Mundi University of Nebraska - Lincoln Year Reclassification of the New World 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 This paper is posted at DigitalCommons@University of Nebraska - Lincoln. http://digitalcommons.unl.edu/insectamundi/295 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.
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