33 NEUROPTERA (NEUROPTERIDA)

John D. Oswald', Atilano Contreras-Ramos" & Norman D. Penny

RESUMEN. En este capitulo se presenta un panorama difficult to encounter. They probably attain their sobre la sistematica, biologia y distribuci6n geografi­ greatest abundance (but not diversity) in desert ca de los Neuroptera (Planipennia) de Mexico, con communities and in a variety of temperate habi­ una orientaci6nhacia la literatura taxon6mica.Se con­ tats, such as forests, grasslands, and urban back­ sideran las familias actualmente conocidas en Mexi­ yards. On warm, early fall evenings in north tem­ co,las cuales estan en orden descendente por riqueza perate towns and cities, storefront and home win­ de especies registradas (entre parentesis): Myrme­ dows are often covered with hundreds of adult leontidae (97), Chrysopidae (81), Hemerobiidae (44), lacewings attracted to the lights. Coniopterygidae (36), Mantispidae (22), Ascalaphidae Neuroptera have two distinctive characteristics (21), Sisyridae (4), Ithonidae (2), Berothidae (2), Dila­ that make them fascinating creatures. First, they ridae (1) y Polystoechotidae (1). Lafauna total de Neu­ are predators, especially as larvae, giving them the roptera actualmente registrada en el pais suma 311 es­ distinction of helping protect us from a wide vari­ pecies. Como en otroscasos,elorden ha sido estudiado ety of agricultural and horticultural pests (Tauber s610 superficialmente en Mexico, por 10 que se consi­ et al., 2000) as well as disease carriers. Secondly, dera importante que se realicen estudios sistematicos they have developed broad, membranous wings y faunisticos en las diferentes regiones del pais. for flight, which are strengthened by an elaborate network of crossveins, and hence the name lacew­ ings. A few of the larger, diurnal species of Myrme­ INTRODUCTION leontidae, Nemopteridae, and Ascalaphidae can be quite colorful, and at a distance can be mis­ Neuroptera, or lacewings, antlions, and their al­ taken with large butterflies. lies, is one of the smaller orders of insects, with approximately 6000 described species, of which only about 4000 species are estimated to be valid SYSTEMATICS AND DIVERSITY taxa (Aspock et al., 1980). They are found in all temperate and tropical parts of the world, but Documenting biodiversity for insects is a difficult rarely constitute a major component of the insect task. Insects are often small and elusive, and pass fauna. Neuropterans probably reach their great­ many months of the year in immature and resting est diversity in tropical forest communities, but stages. Even when abundant, they are often cryp­ seldom are very abundant there, and often are very tically colored to avoid being eaten by visual predators, like birds and lizards. Thus, few com­ parative statements have been made about the diversity of insect groups for any region. For some of the more conspicuous groups, such as butter­ 1 Department of Entomology, Texas A & M University, College Station, Texas 77845-2475,USA. e-mail: [email protected] flies (DeVries, 1986) and fulgorid bugs (Porion, 2 Centro de Investigaciones Biol6gicas, Universidad Aut6no­ 1994), however, the tropical parts of the Neotropi­ rna del Estado de Hidalgo, Apdo. Postal 1-69, Plaza Juarez, cal region appear to contain the richest taxonomic Pachuca, Hidalgo 42001, MEXICO. assemblage in the world. Penny (1978) listed 1100 e-mail: [email protected] species and subspecies of Neuroptera for America 3 Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, south of the United States, or more than 25% of USA. e-mail: NPennyOCalAcademy.org the known world fauna, including 171 taxa from 560I JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

Mexico. Our taxonomic knowledge of the orderhas Neuropterans however, remains unidentifiable at increased considerably in the past 24 years, but the this time. proportions remain roughly the same; Mexico ap­ Collections and studies in Mexico. Although pears to contain approximately 4% of the world's Neuropterans have been studied to some extent in known Neuroptera fauna (Appendix 33.1). Mexico with respect to their potential use as biologi­ Several families of Neuroptera are absent from cal control agents, at this time there are no formal Mexico. Members of Nymphidae are restricted to specialists in the country. Also, we are not aware of Australia. Species of Nevrorthidae, Psychopsidae, graduate students being trained in systematics of Rapismatidae and Rhachiberothidae are found only Neuroptera within or out of the country that in the in the Old World. Representatives of Nemopteridae near future might engage in the study of the Mexi­ and Osmylidae are found in northern South can fauna. There has been, however, considerable America, but have never been collected in Central work on the Mexican fauna, mostly by non-Mexi­ America or Mexico. All eleven other families can can specialists describing species as part of revision­ be found in some part of the country, including all ary work (e.g., Victor J. Monserrat from Spain on major diverse and speciose families of the order. Hemerobiidae, etc.).Itis evident, therefore, thatthere is room for entomologists within the country to fo­ cus on the study of the different taxonomic and bio­ TAXONOMIC KNOWLEDGE logical aspects of the group. Regarding collections, in Mexico Neuropterans There has never been a taxonomic treatment of the tend to be underrepresented. Perhaps the largest Neuroptera of Mexico. Part of the reason for this number of specimens is at the collection of the are the numerous poor descriptions by pioneer Instituto de Biologia of the National University, taxonomists. For instance, Longinos Navas de­ Mexico City, but even there only several drawers scribed 582 species and subspecies from the Neo­ hold Neuropterans and the specimens are prob­ tropical region between 1908 and 1936 (Penny, ably sorted only to family level. Evidently, much 1978) with short, undiagnostic descriptions, few field work is required. Collections with important comparisons with other taxa known at the time, holdings of specimens out of Mexico are mostly and fewer illustrations. Many of the type speci­ in the United States, mainly in California, for in­ mens used in his descriptions have subsequently stance the California Academy of Sciences (CAS) been destroyed (Monserrat, 1985, 1986). This cre­ in San Francisco, Los Angeles County Museum ated a long period of cautious reluctance to make (LACM) in Los Angeles, the San Diego Natural taxonomic decisions about the tropical American History Museum (SDNHM), and others, such as fauna, and only slowly we are emerging from this the Smithsonian Institution (NMNH) in Washing­ period. Good taxonomic treatments exist for spe­ ton and Texas A & M University (TAMU) in Col­ cific taxa (Tauber, 1969; Stange, 1970a, b; Meinan­ lege Station. Other USA collections at universities der, 1972,1990; Oswald, 1988; Brooks & Barnard, near the border with Mexico might hold Mexican 1990; Brooks, 1994; Monserrat, 1996, 1997, 2000) Neuropterans, but also institutions where special­ found in Mexico. With the help of newer mono­ ists have settled (e.g., YaleUniversity, Florida State graphs it is now possible to identify almost all Collection of Arthropods). One of us (ACR) has adult Mexican Neuropterans to genus (e.g., Henry started collecting efforts in Central Mexico, par­ et al., 1992), but identification of species for many ticularly in Hidalgo State, and it is hoped that new genera remains extremely difficult. students might become involved in formal train­ Even less is known of immature stages. Most ing to study the group in the country. larval Neuropterans cannot be identified below family level, although Stange (1994) provides keys to tribes of larval Myrmeleontidae and gen­ PHYLOGENY era of larval Brachynemurini. Tauber & De Leon (2001) have published a key to larval Ceraeo­ The evolution of Mexican Neuroptera must be chrysa for Mexico. The vast majority of larval viewed in a global context. Neuroptera have a rich NEUROPTERA (NEUROPTERIDA) I 561 fossil history, with records dating back to early better. This state is found in the Chrysopidae and Permian, more than 250 million years ago. By the Hemerobiidae. The final step in larval mandibu­ middle Triassic there was a rich assemblage of now lar evolution was the development of teeth on the extinct lineages present (Kukalova-Peck, 1991). curved mandibles for yet better grasping of prey. Thus, extant fauna must be viewed as relicts of a This state is found in the lineage containing Psy­ much more diverse past. chopsidae, Nymphidae, Nemopteridae, Myrme­ We are in great need of a better phylogeny for leontidae, and Ascalaphidae. extant lineages of Neuroptera. Much of the basis Elements of subfamilial and tribal phylogeny for currentrelationships is based on larval morphol­ have been worked out for most families. A phylo­ ogy, as originally presented by Withycombe (1925); genetic structure for subfamilies and genera of however, Aspock (1992)has noted some problems Hemerobiidae (Oswald, 1994a) and for Psychopsi­ with this phylogeny. UnderWithycombe's scenario, dae (Oswald, 1993) was developed. The subfam­ the oldest lineages are the Ithonidae-Rapisma­ ily and tribal relationships of Chrysopidae are dis­ tidae-Polystoechotidae, whose larvae appear to cussed in Brooks & Barnard (1990). Stange (1994) have more than three ins tars (the number found has done much the same for Myrmeleontidae. The in all other families) and stout mandibles which tribal relationships of Coniopterygidae (Meinan­ may not be grooved for consumption of fluids. der, 1972) and Mantispidae (Lambkin, 1986) have These are also the only larvae in the order which been analyzed. are subterranean. According to Withycombe, the most inefficient type of grooved larval mandibles are straight and BIOGEOGRAPHY without teeth, making grasping prey more diffi­ cult. Groups in this lineage are Coniopterygidae, , Mexico is one of the few countries containing two Osmylidae, Sisyridae, Nevrorthidae, Dilaridae, biogeographical regions -the Nearctic region in Berothidae, Rhachiberothidae and Mantispidae. the north and higher elevations and the Neotro­ Within this lineage Coniopterygidae have an un­ pical region in the south and along the coasts. This certain placement. They are very small, with very regional confluence coupled with considerable to­ reduced wing venation. Larvae have only six pographical relief has generated a tremendous Malpighian tubules, while all other Neuropterans number of biomes and habitats for a single coun­ have eight. Adult coniopterygids do have one very try. The close proximity of mountains and oceans distinctive apomorphy, they have hypodermal has allowed species to survive over long periods glands on the body which secrete a white wax of time and changing environmental conditions which covers the entire body. Sisyridae and by simply migrated short distances up and down Nevrorthidae both have aquatic larvae, but the a mountainside, around a mountain slope, or Sisyridae have several morphological adaptations nearer or farther away from the coast. The sur­ for aquatic life not found in larvae of the vival of many species over time leads to 'species Nevrorthidae. The lineage containing Berothidae­ packing' in an area, with more diversity and nar­ Rhachiberothidae-Mantispidae has adapted to a rower niches. The various isolated mountain sedentary larval existence with hypermetamor­ chains and tropical enclaves along the coast gen­ phosis. The components of this lineage merge and it erate an island effect of small insect populations could be argued that they belong in a single family evolving in isolation and eventually developing (see Aspock & Mansell, 1994;Aspock & Nemesch­ endemic species with small geographical distri­ kal, 1998;Tjeder, 1968). Dilaridae appear to be the butions (see McArthur & Wilson, 1967). sister-group of this lineage based on components The influence of northern temperate faunal ele­ of the male genitalia (such as an extremely long ments may be seen in such groups as the chrysopid penisfilum) and wing venation. According to Meleoma and the hemerobiid Micromus. The tropi­ Withycombe (1925); the next step in larval man­ cal influence is present in the chrysopid Gonzaga dibular evolution was the development of curved, and the hemerobiid Nusalala. More problematical but untoothed, larval mandibles to grasp prey is the presence of the ithonid genera Narodona and 562/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS &: NORMAN D. PENNY

Oliarces. Ithonidae has a bimodal distribution in TAXA IN MEXICO Australia and the region from southwestern U.s.A. to Honduras. The simplestexplanation for Coniopterygidae such a distribution would be a former Gondwa­ The family Coniopterygidae, dustywings, is a naland distribution with the South American el­ moderate-sized group of minute Neuropterans. ement eventually moving northward into Mexi­ The common name 'dustywings' refers to the co. The absence of thorough revisionary works white or gray waxy powder that covers the body of many of the Mexican groups, together with and wings of adults. The wax is produced by cu­ solid species level phylogenies, strongly limit ticular glands on the head, thorax and abdomen inference on the historical biogeography of the and is spread over the rest of the body by the adult Mexican area based on modern trends such as insect, primarily using its hind legs. Coniopte­ cladistic biogeography. rygids are notable within the Neuroptera for their small size. Although a few species belonging to other Neuropteran families (e.g., Hemerobiidae CONSERVATION AND IMPORTANCE and Sisyridae) fall into the upper size range of coniopterygids (which have forewing lengths of Because of their role as biocontrol agents on agri­ 1-5 mm), the Coniopterygidae constitute the most cultural crops, a few Neuropterans are clearly rec­ diverse group of minute Neuropterans and con­ ognized as beneficial insects. However, since it is tain the order's smallest members. Other charac­ likely that all Neuropterans are predators, their ters of dustywings that are unusual within the role in the larger ecosystem could be potentially order Neuroptera are their highly modified (re­ much greater. A few species tend to thrive in an­ duced) wing venation and distinctive male termi­ thropogenic habitats, such as Myrmeleon larval pits nalic structures. Because of their small size and under houses and bridges, Ceraeochrysa in or­ waxy body covering, coniopterygids often super­ chards, and Chrysoperla in grasslands, pastures, ficially ressemble adult whiteflies. and backyards. When life histories of these insects Although uncommonly collected by non­ are studied, almost invariably they are found to neuropterists, coniopterygids can be extremely be beneficial to mankind. abundant locally where favorable preyconditions As for many other groups of plants and ani­ occur. Specimens can be collected by net but are mals, the rapid depletion of natural habitats is a generally more easily collected at lights. Histori­ great concern for preservation of overall Neu­ cally,most specimens of adultconiopterygids have ropteran diversity. The forests, grasslands, deserts been preserved by air drying on point mounts. and mountainsides act as reservoirs of genetic di­ However, air-dried specimens shrivel badly over versity and once gone cannot be replaced. Be­ time because the adult body is only weakly scle­ cause of the species richness and potential for rotized. Specimens prepared in this way require many locally endemic species, the Neuropteran time-consuming dissections to obtain confident fauna is vulnerable to local destruction and ex­ identifications. A better method of preserving co­ ploitation. The most positive aspect is the corol­ niopterygids (and other small Neuropterans) is to lary -each tract of land that is placed off limits collect them into alcohol and point mount them to development holds the promise of preserving after critical point drying. With this technique, Neuropteran diversity along with the often more reliable identifications of male specimens can be charismatic plants and animals that it was de­ made without dissection after a workingknowled­ signed to protect. In order to understand what ge of a local fauna has been established. we may be potentially losing, it is vitally impor­ Biology. Coniopterygid larvae are predaceous. tant that the Neuropteran diversity be docu­ Adultsare also generally predaceous,butmay also mented along with geographical distributions. feed on honeydew. Adults and larvae have been Further collections are clearly needed, as well as reported to feed on a wide variety of slow-mov­ trained specialists and detailed publications to ing arthropod prey, including mites, scale insects, help develop this foundation. insect eggs, coccids, aphids and phylloxerans. NEUROPTERA (NEUROITERIDA) I 563

Most coniopterygid species are associated with ever, given the under-collected nature of this woody vegetation (trees or shrubs) rather than group, many additional species (both undescribed herbs. Plant 'host' records demonstrate that at least and simply unreported) are to be expected in this some coniopterygid species show preferences for region. The total number of species present in particular plant species, suggesting that such spe­ Mexico will verylikely be more than50.The Mexi­ cies may exhibit arthropod prey preferences that can fauna contains numerous species in the gen­ are linked to the host plant preferences of the prey era Coniopteryx and Semidalis, and fewer species species. Many coniopterygid species have multi­ in the genera Conwentzia and Neoconis, as is typi­ voltine life cycles, even in temperate regions. Be­ cal of many New World faunas. The number of cause of their small size, rapid generation time and Mexican species reported here in the genus feeding preferences, some coniopterygid species Aleuropteryx (4) is certainly low. Fourteen species may be valuable for the biological control of mites of this genus are known from southwestern United and sessile homopterans. States, and most of these are probably also present Taxonomy and geographic distribution. The fam­ in northern Mexico. It should be noted that in­ ily Coniopterygidae currently contains ca. 490 de­ traspecific variation in important morphological scribed, valid, species worldwide and is the fourth traits such as the male terminalia is still being most speciose family in the order Neuroptera. worked out in some species complexes, which Coniopterygids are known from all continents (ex­ could lead to future nomenclatural changes (par­ cept Antarctica) and from many oceanic islands. ticularly in the genus Semidalis). The Mexican fau­ The known New World fauna contains ca. 100spe­ na appears to contain few endemic and few wide­ cies placed in 12 genera. Within the New World spread species. Only five of the 32 reported spe­ fauna, 70 percent of the species diversity is con­ cies are presently known only from Mexico, but tained in the three genera Aleuropteryx, Coniopteryx, most of these are known from very few records and Semidalis. Because of the small size of its spe­ and may prove to be more Widespread with addi­ cies, the family Coniopterygidae has historically tional collecting. Only four Mexican species are been under-collected, poorly-studied and pre­ known to occur both to the north and south of sumed to constitute a small group of rather rare Mexico. The vast majority of Mexican species ex­ Neuropterans. The modem era of study of this hibit clear Nearctic or Neotropical affinities and group, dating from the groundbreaking world re­ reach either the southern or northern limits of their vision of Meinander (1972), has revealed quite a range within Mexico. different picture. The known species diversity of this family has more than doubled from 231in 1972 Ithonidae to ca. 490 in 2001, and it has become more well The family Ithonidae, moth lacewings, is an un­ known that coniopterygids, far from being rare, usual relic family of lacewings, whose species are are common and often locally abundant. Given rarelycollected because of their remotehabitats and the rapid rate of description of new species in this unusual lifehistories. Adults are robust-bodied and group over the past 30 years, which shows little hold their brownish or pale wings relatively flat of sign of slowing, additional new species can be over the back, in contrast to most other lacewings expected in many regions of the world, particu­ in which the wings are typically held at a steeper larly in the relatively poorly studied regions of the 'roof-like' angle over the body. The common name New and Old World tropics including Mexico. In for this group denotes the superficial resemblance addition to Meinander's world revision, other of adults to certain moth species. Ithonids are gen­ works relevant to the Mexican fauna include erally regarded as 'primitive' Neuropterans related Meinander (1974a,b, 1975,1981,1983,1990,1995), to polystoechotids and rapismatids. Monserrat (1984a) and Monserrat & Bayo (1995). Biology. The only ithonid species for which sig­ Mexican fauna. Seven genera and 36 species of nificant biological information is available are the coniopterygids are currently known from Mexico. Australian species Ithone fusca (see Tillyard, 1922) This represents approximately one-third of the and the American Oliarces clara (see Faulkner, described New World fauna of this family; how­ 1990a, b), the biologies of which are exceedingly 564/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY unusual for the Neuroptera. Larvae are robust and undescribed ithonid species are also known from C-shaped, resembling scarab beetle larvae, and are Mexico (penny, 1996). found in the soil. Initially thought to be preda­ Mexican fauna. The named Mexican fauna of ceous on soil-dwelling scarabeoid Coleoptera (e.g., Ithonidae consists of the single very poorly known Tillyard, 1922), it now seems likely (though not species Narodona mexicana which has been reported yet conclusively demonstrated) that the larvae of from a single collection from Colima, Mexico. As these species are saprophagous (Gallard, 1932; noted above, at least two undescribed species of Faulkner, 1990a),feeding onthe fluids of plantroot ithonids are also known from Mexico. In addition, tissues. If confirmed, these species would be the apterous females similar to those described for only known plant-feeding Neuropterans. Adults Adamsiana curoei have been collected in Guatemala of these species, and other Australian species and it is possible that this species, or unknown known from live adults, also exhibit unusual bio­ related species, may also occur in southern Mexico. logical characteristics. Adults appear suddenly in synchronized mass emergences of short temporal Sisyridae duration (ca. 1-2 weeks). Males form large aggre­ The family Sisyridae, spongillaflies, is a small group gations, generally associated with a local elevated of Neuropterans that is closely associated with prominence, to which females fly for mating. Be­ water. Adults strongly resemble small brown lacew­ cause these species are relatively poor fliers, ag­ ings (Hemerobiidae), with which they are com­ gregations probably form from only locally emerg­ monly confused. Sisyrids, however, can easily be ing individuals. Females deposit eggs on, and per­ distinguished from hemerobiids by their lack of haps in, the soil. Young larvae actively burrow into parapsidal furrows on the mesonotum (in heme­ the soil. Larvae of OIiarces clara appear to be asso­ robiids and other Neuropterans these furrows di­ ciated with the roots of the creosote bush (Larrea vide the prescutum from the scutum) and the pres­ tridentata), a Widespread desert shrub. At least in ence of a pectinately branched Rs (rather than the this species, spring emergences may be cued by pectinately branched R in hemerobiids). Sisyrids earlier rainfall events that promote the subsequent are typically collected at lights or by sweeping wa­ growth of desert shrubs (Wiesenborn, 1998). Life ter-side vegetation and can be very abundant lo­ cycles take 2-3 years to complete, most of which is cally where favorable aquatic habitats occur. spent in the larval stages. Unlike most Neuropter­ Biology. The families Sisyridae and Nevrorthidae ans, which have three larval instars, five instars (the latter strictly Old World) are the only Neu­ have been reported for !thane fusca. The biologies ropterans with true aquatic larvae. Sisyrid larvae of the New World genera Narodona and Adamsiana are highly specialized and feed on the intracellu­ are entirely unknown. Based on the known col­ lar contents of freshwater sponges. Larvae appar­ lecting sites of Narodona and Adamsiana adults, the ently exhibit little or no host specificity and may microhabitats occupied by species in these gen­ assume the color of the host sponge through feed­ era are almost certainly to be more cool and moist ing. The extremely slender, straight, larval mouth­ than those known for !thane and OIiarces. parts facilitate the specialized feeding regimen of Taxonomy and geographic distribution. The world this group. Second and third instar larvae (but not ithonid fauna currently consists of 17 species first instars in examined species) possess a series placed in six genera. Three genera and 14 species of moveable ventral abdominal processes which are endemic to Australia (see Riek, 1974). The re­ apparently aid in respiration by creating a flow of maining three species are placed in three mono­ water near the body. Late third instar larvae leave typic genera with restricted North and Central the water to pupate on objects, typically vegeta­ American distributions as follows: OIiarces clara tion, along the margins of the water. The cocoon (desert regions of southern California, southern is bilayered with a loose outer mesh enclosing a Nevada and western Arizona; Adams, 1950; Bel­ denser inner envelope. Adults feed on pollen and kin, 1954), Narodona mexicanus (Mexico: Colima), small arthropods gleaned from waterside vegeta­ and Adamsiana curoei (Honduras: Cordillera del tion. Eggs are laid over or near the water and are Merend6n, Cusuco). At least two additional generally covered by the female with a fine silken NEUROPTERA (NEUROPTERIDA) 1565 mat. For detailed biological accounts of several (Adams, 1970), for which they also share the nu­ sisyrid species see Brown (1952), Pupedis (1987), merous long setae and female ovipositor. and Weissmair (1999). Biology and geographical distribution. Most of Taxonomy and geographic distribution. The fam­ what we know about the biology of Dilaridae ily Sisyridaecontains ca. 60 described species that comes from the studies of Nallachius americanus are collectively distributed throughout the New (McLachlan) by MacLeod & Spiegler (1961). Lar­ and Old Worlds. The New World fauna consists vae were collected under the tightly adherent bark of ca. 30 known species placed in two genera. The of erect, recently dead trees. They were success­ cosmopolitan genus Sisyra contains ca. 10 New fully fed on soft, disabled insect larvae and eggs. World (and ca. 30 Old World) species with a cen­ Adults are attracted to lights and are most often ter of diversity in northern South America. The collected in mature forests. Penny & Arias (1982) endemic New World genus Climacia contains ca. found that in the Amazon Basin adults were more 20 species and also exhibits a center of diversity abundant in the forest canopy than at ground level, in northern South America. Discoveries of addi­ and were more frequently collected during the tional new species are to be expected in the New drier part of the year. However, individuals of this World sisyrid fauna. Parfin & Gurney (1958) and family are not often collected, even under ideal Poirrier & Arceneaux (1972) provide starting conditions, and they must be considered one of points for the identification of Mexican sisyrids. the more uncommon families of Neuroptera. Mexican fauna. The confirmed Mexicansisyrid fauna Dilaridae are found in both the Old and New currently consists of four species: Climacia areolaris, C. World. The Old World distribution is in tropical chapini, C. californica, and Sisyra apicalis. The first of Asia as far south as Borneo and in Europe. They these, C. areolaris, is widely distributed in the eastern appear to be absent from Australia and tropical and southernU.s.A. and extends south into northeast­ Africa. In the New World they are found from ern Mexico. Thesecond, C.chapini, is known only from northeastern U.s.A. south to northern Argentina, the south-central U.s.A. and Mexico. The third, C. but appear to be absent west of the Andes. califarnica, has been collected in Sonora and was for­ Taxonomy. New World Dilaridae all belong to a merly known from the western U.s.A. The fourth, S. single genus, Nallachius, in the subfamily Nalla­ apicalis, is known from Cuba, southeastern U.s.A., chiinae. There are seventeen described species, CentralAmericaand northernSouth America,and has most from South America. Only four species are been collectedon the YucatanPeninsula (D.E.Bowles, known from North and Central America. Adams personal communication). Several other Climacia spe­ (1970)divided Nallachius into three species groups: cies are known from central America and it is likely the pu1chellus, limai, and americanus groups that at least some of these will subsequently be found based on wing characters, color of the labrum, and in southern Mexico. Another species of the genus structures of the male genitalia. Sisyra, Sisyra nociurna, is known from Belize and Hon­ Mexican fauna. There are no published records of duras and might be later found within the country. Mexican Dilaridae. Two species of Nallachius (N. americanus and N. pulchellus) are known from Costa Dilaridae Rica and U.s.A. and can be presumed to exist in The Dilaridae, or pleasing lacewings, are a small Mexico. In fact, collected specimens of N. puldiellus distinctive family of Neuropterans. They are rela­ (Banks) existfrom 16milesnorth ofMazatlan (Sinaloa) tively small in size; the body is covered with nu­ and Madre Maria Island off the coast of Nayarit. Two merous long hairs; the wings are relatively broad, other Central American species of Nallachius (N. with transverse dark bands; males have pectinate championi [Navas] from Guatemala and N. parkeri antennae; and this is one of the few families of Penny from Costa Rica) might be found in Mexico, Neuroptera in which the females bear ovipositors. particularly in the southernmost states. In many respects these insects resemble small moths (especially the males) for which they are Polystoechotidae often mistaken. The male genital armature most The family Polystoechotidae, giant lacewings, is closely resembles that of the family Osmylidae a small group of rarely-collected Neuropterans. 5661 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS &: NORMAN D. PENNY

Adults are large (forewing length ca. 15-35 mm) the swollen, raptorial forelegs which give them the with dark, rather robust, bodies andhyaline wings appearance of small preying mantids. The family with variable dark markings. The phylogenetic is cosmopolitan in tropical and temperate parts of affinities of this group are uncertain. They are gen­ the world, but much more diverse and abundant erally regarded as 'primitive' Neuropterans whose in the tropics. Adults are most frequently seen at affinities may lie with the Ithonidae and/or lights, butonespecies, Climaciella brunnea (Say, 1824) Rapismatidae. The monophyly of the group has is a very good vespid wasp mimic (see Opler, 1981) not been demonstrated. and waits at flowers for prey. Biology.The biology of this family remains essen­ Biology and geographical distribution. In those tially unknown. Eggs have been obtained from few species where it is known, the number of eggs captive females by several workers (e.g., Welch, deposited is quite large. Redborg & MacLeod 1914; Hungerford, 1931), from which first instar (1985) found for the North American Mantispa larvae have been reared. The feeding and micro­ uhleri Banks that individual females laid an aver­ habitat requirements of these presumably preda­ age of 12.6 clutches of individually placed eggs ceous larvae however, have not been discovered on long stalks. The number of eggs per clutch var­ and no species of the group has ever been suc­ ied from 614 to 2976. Mantispid larvae are hyper­ cessfully reared beyond the first instar. The field metamorphic. That is, the first instar larva is elon­ biology of the group is entirely unknown, though gate and actively seeks quiescent prey. The subse­ it is suspected that the larvae will be soil dwell­ quent two instars are scarabaeiform, sluggish, and ing. Apart from the late-instar larva partially de­ feed on the prey that they have located during the scribed by MacLeod (1964), no mature larvae of first instar. The type of larval food varies with the this group are known. The Widespread North subfamily. Larvae of Symphrasinae have been American species Polystoechotes punctatus, which found in subterranean scarab pupae, in Polybiinae was once a common insect in many parts of its wasp nests and in individual solitary wasp cells. range, has apparently gone extinctoverlarge parts Larvae of Mantispinae appear to obligately feed on of its former range since the 1930's or 1940's (as a wide variety of spider eggs (Hoffman & Brush­ indicated by the absence of recent collecting wein, 1989, 1990. They enter the spider egg sacs records). The cause of this decline is unknown. using two different strategies: a) boarding an adult Taxonomy and geographic distribution. The spider and then entering the egg sac as it is being Polystoechotidae are exclusively New World in produced, or b) penetrating directly into the egg distribution, with two species occurring in North sac after formation (Redborg & MacLeod, 1984). America and two in South America. The four Mantispids of the genus Climaciella appear to re­ known extant species are as follows: Fontecilla quire a blood meal from an adult spider before graphicus (Chile), Platystoechotes lineatus (U.S.A.: entering the egg sac (Redborg & Macleod, 1983). California), Polystoechotes gazullai (Chile). and P. Redborg and MacLeod (1985) have perfected a punctatus (southernCanada south to Panama). The technique for raising large quantities of larval Nearctic species were revised by Carpenter (1940). Mantispinae in the laboratory on artificial spider Mexican fauna. The Mexican fauna consists of a egg sacs. single species, Polystoechotes punctata, which is Geographically, the Mexican mantispid fauna is Widespread in North America. The few known transitional between species found more frequently Mexican records for this species are from north­ to the north and south. As can be seen from the spe­ ern and central Mexico. None of the other species cies list below, until now there have been few Mexi­ in this family are likely to occur in Mexico, and no can records of the fauna commonly collected in the undescribed species are known. USA. At this time little can be said about the geo­ graphical distribution within Mexican due to the Mantispidae paucity of collection records from many areas. The Mantispidae (Fig. 33.1), or mantid-flies, are a Taxonomy. The family has been divided into four medium-sized group of Neuropterans (300species; subfamilies (Lambkin, 1986). Two African genera, Aspock et al., 1980) immediately recognizable by Mucroberotha Tjeder and Rhachiberotha Tjeder, vari­ NEUROPTERA (NEUROPTERIDA) I 567

Figs. 33.1-3. Habitus of adult Neuroptera: 1) Plega signata (Hagen), Mantispidae (redrawn from Henry et al.,1992); 2) Chaetoleon pusillus (Currie), Myrmeleontidae (redrawn from Stange, 1994); 3) Ululodes mexicanus (McLachlan),Ascalaphidae (redrawn from Henry et al., 1992). ously been have placed in the Berothidae (Tjeder, unpublished PhD. thesis). This thesis contains much 1959)and Mantispidae (Tjeder, 1968),butcurrently new morphological, taxonomic, and geographical are placed in their own family, Rhachiberothidae information on the Latin American fauna. Even so, (Aspock & Mansell, 1994). Of the four recognized identification of mantispids from most parts of the subfamilies, Symphrasinae are found in the south­ tropical world remains problematical. western part of the Nearctic region and through­ Mexican fauna. Penny (1977) listed 14 species of out the Neotropical region. Drepanicinae are found Mantispidae from Mexico; however, only two of in Australia and southern South America. Calo­ the 13 species recognized from the southern USA mantispinae are found in Australia and tropical have previously been recorded from Mexico and is America as far north as the southwestern U.s.A. a reflection on the lack of work on the Mexican Finally, Mantispinae are found throughout the fauna. In this work, the authors record 22 species world in temperate and tropical climates. The fau­ for the country. The fauna is almost certainly much na of only a few regions have been treated in re­ more diverse than currently known, and much cent years: Japan (Kuwayama, 1962), western Eu­ more collecting and study is needed. rope (Aspock etal., 1980),Brazil (penny& da Costa, 1985), and Australia (Lambkin 1986a, b). The New Berothidae World Mantispinae were treated by Hoffman (1992, The family Berothidae, beaded lacewings, is a 568/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS &: NORMAN D. PENNY small group of rather uncommon Neuropterans. ior, prey specificity and habitat partitioning of The fore wings of North American species (but sympatric species are currently outstanding. not most species from other regions) are slightly Taxonomyand geographic distribution.The Bero­ elongate and more or less falcate (i.e., with a more thidae are a small family of ca. 105 described ex­ or less concave area distally on the posterior mar­ tant species placed in ca. 25 genera and 6-7 sub­ gin of the wing). Females possess a pair of elon­ families. The group is widespread in both the New gate processes that arise from the end of the abdo­ and Old Worlds, but the New World fauna is rela­ men and fold under the abdomen at rest. Females tively depauperate with only ca. 20 species in four of several species also possess distinctive patterns genera (Cyrenoberotha, 1 sp., Chile; Naizema, 2 spp., of scale-like setae on the wings and/or thorax. Argentina; Spiroberotha, 2 spp., Costa Rica and Many species are very similar and can be confi­ Venezuela; Lomamyia, 12 spp., southern Canada dently identified to species only by detailed ex­ south to Brazil). The group as a whole (and the amination of the terminalic segments. Most New World fauna too) appears to be most speciose berothids are collected at lights. On a light sheet, in desert regions, with few species in the moist specimens superficially resemble the elongate tropics. Several undescribed species of the genus body form of caddisflies, from which they can Lomamyia are known from north of the Mexico­ usually be distinguished by their behaviors of U.s.A. border, and additional new taxa should be holding their wings off the sheetat an angle (rather expected in this group from farther south. The than flush with the sheet) and their constantly primary taxonomic references on the New World moving antennae (rather than still antennae). berothid fauna are Carpenter (1940), MacLeod & Berothids are believed to be most closely related Adams (1967), Penny (1983), and Adams (1989). to mantispids and dilarids. Gurney (1947) and Tauber & Tauber (1968) treat Biology. The biology of berothids is poorly known. the morphology of the immature stages of Loma­ Even though some species can be locally abun­ myia. Aspock (1986) reviewed the world berothid dant in desert regions around the world, the only fauna, Aspock & Nemeschkal (1998) provided an two detailed life history studies of species of this initial phylogenetic hypothesis of intergeneric re­ family are those of Tauber & Tauber (1968), for lationships within the family. Lomamyia laiipennis, and Brushwein (1987),for Lo­ Mexican fauna. The confirmed berothid fauna of mamyia longicollis. Larvae of both of these species Mexico currently consists of a single species, Lama­ are apparently obligate termitophiles in the gal­ muia squamosa, whose range is known to extend leries of termites of the genus Reticulitermes (Isop­ from southern Texas south to Honduras. During tera: Rhinotermitidae). Adults lay stalked, pen­ the preparation of this work the authors have ex­ dent, egg clusters in microhabitats occupied by amined material of at least one additional uniden­ Reticulitermes species. First instar larvae locate and tified Lomamyia species from northern Mexico penetrate the gallery system of termite colonies, (Tamaulipas). Six described and several unde­ where they immobilize and feed on individual scribed Lomamuia species are known to occur in the termites. Prey are apparently immobilized by a desert regions the southwestern United States. Itis poison injected with a quick thrust of the straight almost certain that future sampling will reveal at larval jaws. The reportbyJohnson & Hagen (1981) least some of these in northern Mexico. It is also that 1. latipennis uses an aggressive gaseous possible that the Costa Rican species Spiroberotha allomone to immobilize prey requires additional sanctarosae may be present in southern Mexico. confirmation. Feeding occurs only after immobi­ lization. Interestingly, only first and third instar Hemerobiidae larvae are active and feed. Second instar larvae The Hemerobiidae, or brown lacewings, is a spe­ are inactive and exhibit morphologically reduced cies rich family of virtually cosmopolitan Neu­ legs and mouthparts. It is currently unknown if ropterans. Individuals of this group are among the the termitophilous habit is a general biological most commonly encountered Neuropterans dur­ characteristic of the entire family. Many interest­ ing survey work. They have also attracted some ing questions regarding female ovipositionbehav­ research due to their economic importance as po­ NEUROPTERA (NEUROPTERIDA) /569 tential biocontrol agents of small phytophagous species and several more have been described af­ insects. The world fauna of brown lacewings con­ terwards. Most of the latter have received little at­ tains ca. 546 valid species, third in diversity after tention since their original descriptions (Oswald, Myrmeleontidae and Chrysopidae (Monserrat, 1990), so the actual number of valid Neotropical 1990b).Adults are typically brown, less frequently species may be reduced somewhat due to syn­ yellowish, black, or green. Most species are rather onymy although considerable clarification has al­ small, with forewing lengths ranging from 4 to 12 ready taken place (e.g., Monserrat, 1996,1997,2000). nun (New, 1989). They are easily distinguished Taxonomy. The family is currently divided in 10 from almost all other Neuropterans by the Rs and subfamilies based on cladistic analyses of intra­ MA being partially fused, with forewing R appear­ generic relationships by Oswald (1993a, b, 1994), ing to have multiple radial sectors (Henry et al., although an eleventh Chilean subfamilyhas been 1992[fig.28.17]). Adults can be collected by sweep­ recently proposed by Monserrat & Deretsky ing vegetation and at lights, and confident spe­ (1999). Both the Nearctic and Mexican faunas are cies identification often requires clearing and ob­ composed of taxa placed in the subfamilies He­ servation of the male genitalia. merobiinae, Megalominae, Microminae, Notio­ Biology and geographical distribution. Adult biellinae, and Sympherobiinae. habits are poorly known, in part due to their cryp­ Mexican Fauna. The known fauna of Mexican he­ tic nocturnal or crepuscular behavior. They are merobiids corresponds to nearly 67% of the Ne­ generally found on branches and foliage of trees arctic fauna (U.S.A. and Canada). Assuming that and bushes, but some species (e.g., several Micro­ the Nearctic fauna is well known, it may be specu­ mus spp.) tend to be found in open, grassy areas lated that after formal sampling in the different (penny et al., 1997). Some adults of this group are physiographic regions of Mexico the fauna of He­ noted for their well-developed brachyptery (e.g., merobiidae of the country might increase consid­ Nusalala andina from Colombia [Penny & Sturm, erably. The two most diverse genera in the coun­ 1984]),which may be related to the isolated habi­ try to date are Hemerobius (17 spp.) and Symphero­ tats occupiedby these species. In general, hemero­ bius (13 spp.), which are also diverse in other New biids appear to be more abundant in temperate World faunas. Species-level identifications are still than tropical climates, however it remains an open somewhat difficult, often requiring the dissection question whether actual species richness is higher of male genitalia. Good starting points to sort out in tropical ecosystems. Some genera, such as Notio­ genera are the works by Penny & Monserrat (1985), biella,are characteristic of the tropics (Henry et al., Henry et al. (1992), and Oswald (1993a). Species­ 1992) and are absent from temperate regions. The level identifications are possible in many cases due physiographic complexity in Mexico may provide to several recent revisionary works, e.g. Monserrat opportunities for making interesting comparisons (1984, 1990a, 1996, 1997, 1998, 2000), Oswald of hemerobiid faunas at different elevations and (1988), and Monserrat & Deretsky (1999). The in diverse vegetation types. Hemerobiid larvae are world checklist by Monserrat (1990b) is a basic predatory and feed on soft-bodied insect preys, guide to the taxonomic literature. typically sessile homopterans (e.g., aphids, white­ flies, and scales). Some larvae appear to have spe­ Chrysopidae cific preference for prey items. Adults appear to The Chrysopidae, or green lacewings, is one of the be primarily predators, but some may take non­ largest and most beneficial families of Neuroptera. animal food as well. Theyare usually medium-sized (wing length about Examples of cosmopolitan genera are Hemero­ 15 nun), pale green wing venation and body, with btu» and Micromus, but many genera are small and goldeneyes. The general greencoloration and trans­ geographically restricted (e.g., Biramus from Ven­ parentwings makes them very difficult to see when ezuela [Oswald, 1993b], Adelphohemerobius from resting under leaves. Adults of some species, such Chile [Oswald, 1994]). In the Nearctic, Penny et al. as those in the genus Chrysopa, emit a foul smell (1997) have recorded 60 species in the family, when annoyed. The voracious appetite of larvae whereas Penny (1978) recorded 110 Neotropical has made them one of the most favored agents of 570 I JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS &: NORMAN D. PENNY biological control on agricultural crops. The elon­ ings, whose larvae appear to be ant-nest preda­ gate, fusiform shape of larvae, with curved, un­ tors. They are medium to large-sized species (wing toothed mandibles indicates a close relationship length 15to more than30 mm) and tend to be more with the brown lacewings (family Hemerobiidae). abundant in drier habitats. Leucochrysini are Biology and geographical distribution. Eggs are found only in the New World, mostly in tropical laid on the ends of long filaments. The eggs canbe forests, although one species is found throughout laid singly, in groups, or together with filamentous much of eastern North America. There are seven stalks clumped. Some larvae, such as Chrysoperla, genera and approximately 180 described species, have short setae on the body and bear no debris, all but 15 of them in the genus Leucochrysa. Mem­ while other larvae, such as Ceraeochrysa and Leuco­ bers of this tribe are easily recognized by their long chrysa, have long thoracic and abdominal setae on antennae, dark pterostigmal spot, and low angle tuberacles and carrybits of debris and skins of their of juncture of the outer gradate series with the prey covering their bodies. The second type of lar­ pseudomedia vein of the forewing. Chrysopini is vae are called trash-bearers, or in Portuguese 'bicho the largest tribe, with 31 genera and about 1000 de lixo'. Larvae go through three instars before described species. They are found world-wide, in forming small, round, whitish, silken pupal cells deserts, forests, grasslands and cultivated areas. on leaf surfaces. Larvae feed on soft-bodied larval Mexican fauna. With its rich variety of biomes, and adult insects, as well as on insect eggs. There Mexico has an attendant rich diversity of chryso­ maybe one to several generationsperyear. In colder pid species. They are abundant in all areas of the regions most chrysopids overwinter in the imma­ country, with different faunal elements in the dif­ ture stages; however, Chrysoperla overwinter as ferent biomes. No apochrysines have yet been col­ adults. Adult chrysopids feed on nectar and pol­ lected in Mexico, but two species are known from len, but apparently do not feed on other insects. Costa Rica and another from the Dominican Re­ Henry (1980) has discovered that male courtship public, so that it is likely that apochrysines occur behavior involves tapping the leaf substrate with in the most tropical parts of the country. Only a his abdomen, which is transmitted to the female as single species of Nothochrysinae is known from vibrations. Without the proper tapping behavior, the northwestern part of the country, but two other females will not respond. species have been collected in the U.S.A near the Chrysopids are found from boreal to tropical border with Baja California and Sonora. There are climates and deserts to rainforests in all parts of 15 genera, five subgenera, and 81 species currently the world, except that they appear to be absent known from Mexico. It is possible that a hundred from New Zealand. They are quite vagile and can or more species will eventually be found within be found on many remote, isolated islands, albeit the country's boundaries, which compares favor­ in reduced diversity. ably with the much better known faunas of Ame­ Taxonomy. Brooks & Barnard (1990) record about rica north of Mexico (81species: Penny etal., 1997), 1200 species of Chrysopidae placed in three ex­ southern Africa (77 species: Tjeder, 1966), Costa tant subfamilies (Nothochrysinae, Apochrysinae, Rica (72 species: unpublished data), Europe (56 and Chrysopinae) and four tribes of Chrysopinae species: Aspock etal.,1980),Australia (53 species: (Ankylopterygini, Belonopterygini, Chrysopini, New, 1980), Madagascar (39 species: Holzel & and Leucochrysini). Nothochrysinae are consid­ Ohm, 1990), and Japan (38 species: Tsukaguchi, ered the most primitive living subfamily (Adams, 1995). Mexico appears to be the center of diver­ 1967) and consists of nine genera and 20 species sity for Meleoma. However, there has not been ex­ distributed unevenly around the world in or near tensive collecting in much of the country and areas of Mediterranean climate. Apochrysinae (12 many more chrysopids are expected to be discov­ genera, 26 species) are very large, broad-winged ered within its borders. lacewings distributed in both Old and New World tropical forests. The Chrysopinae tribe Ankylop­ Key to the Genera of Mexican Chrysopidae terygini is only found in the Old World tropical (modified from Brooks & Barnard, 1990) forests. Belonopterygini are thick-bodied lacew­ 1. In forewing Psm continuous with inner row of NEUROPTERA (NEUROPTERIDA) /571

gradate crossveins; jugal lobe well developed - Hindwing with two gradate series of cross- ...... (subfamily Nothochrysinae) Pimachrysa veins 9 - Inforewing SPsm meets outer gradate crossveins; jugal lobe reduced or absent . 8. Forewingwithone gradate series of crossveins...... (subfamily Chrysopinae) 2 ...... Eremochrysa (Chrysopiella) - Forewingwith two gradate series of crossveins.... 2. Pterostigma with dark spot anteriorly on both ...... Eremochrysa (Eremochrysa) forewing and hindwing; antennae longer than forewing; outer gradate series meeting Psm at 9. Antennal scape parallel-sided; male head often low angle in forewing...(tribe Leucochrysini) 3 modified with frontal horns and elongated an­ - Pterostigma pale although more heavily scle­ tennal scape Meleoma rotized on both forewing and hindwing; anten­ - Antennal scape somewhat concave on medial nae shorter than forewing; outer gradate series surface; head of male unmodified .lO meeting Psm at almost a 90° angle .5 10.Male ninth sternite subapically constricted cre­ 3. Forewing marked with black or dark brown ating small apical node; red gena only facial spots; pterostigmalspotextends to radial vein... markings; no scape stripes Chrysoperla ...... Gonzaga - Male ninth sternite may have lateral lobes or be - Forewing usually unmarked, but if markings are apically upturned, but evenly tapered to apex; present these are restricted to dark shading ad­ frons may have antennal rings or transverse red jacent to crossveins; pterostigmal spot extends or dark bands; antennal scape mayor may not only to subcostal vein...... (genus Leucochrysa) 4 have stripes or spots ll

4.Cell im quadrangula...... Leucochrysa (Leucochrysa) 11. Male genital elements include gonapsis 12 - Cell im ovate or triangular...Leucochrysa (Nodita) - Male genital elements lack gonapsis .l3

5.Antennalflagellomeres broaderthanlong; prono­ 12. Gonapsis elongate, rod-like; forewing cross­

tum much broader than long; forewing cell C1 veins usually without dark margining of cross- veins longer than cell C2 subcostal crossvein located Ceraeochrysa ; halfway between origin of M2 and R+M 1 api­ - Gonapsis cross-shaped with lateral arms; fore­ cal palp segment swollen . wing crossveins heavily bordered with dark ...... (tribe Belonopterygini) 6 margining Pseudomallada - Antennal flagellomeres more than twice as long as wide; pronotum longer than broad (except 13. Pseudopenis present; tignum usually pre­ Chrysopodes collaris and Nineta); forewing cell sent 14 ; - Pseudopenis present; tignum absent 17 C2 as long or longer thanC1 subcostalcrossvein ; much closer to origin of M, than origin ofR+M1 apical palp segments cylindrical and apically 14. Partial or complete dark ring around base of tapered (tribe Chrysopini) 7 antennae Chrysopa (in part) - Noindicationof darkring at base of antennae...l5 6. Forewing longitudinal veins alternating dark and pale; rows of darkspotson thoracic nota..... 15. Pronotum bears no spots or stripes ...... Abachrysa . Chrysopa nigricornis - Forewing longitudinal veins pale; pronotum - Pronotum bears four red or orange spots...... l6 may have longitudinal red stripe, but no dark spots Nacarina 16. Pronotum bears four red spots; radial cross- veins darkened in middle Plesiochrysa 7. Hindwing with one gradate series of crossveins - Pronotum bears four orange spots; radial cross­ ...... (genus Eremochrysa) 8 veins darkened at ends ...Chrysopa quadripunctata 5721 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

17. Dark lateral scape stripe; hindwing generally of insects (i.e., Heteroptera, Diptera, Lepidoptera, with only 1-3 inner gradate veins....Yumachrysa Homoptera, Neuroptera, and Coleoptera), but pol­ - No lateral scape stripe; hindwing bears 5 or more len grains and plant tissue were also found. They inner gradate veins 18 observed two main tactics for prey capture: noc­ turnal flights with their legs stretched like traps 18. Black transverse bands above and below an­ (with further prey consumption in a sitting posi­ tennal bases Nineta tion), and diurnal waylaying for potential moving - Frons pale or with two red transverse bands..... victims. Courtship and mating in myrmeleontids ...... Chrysopodes is almost entirely unknown, but several species have been found to communicate via male phero­ Myrmeleontidae mones secreted by specialized exocrine glands (cf, The Myrmeleontidae (Fig.33.2),orantlions, are the Henry et al., 1992), and reproductive isolation by largest family of Neuroptera with about 2000 spe­ specific mixtures of compounds forming unique cies worldwide (New, 1991) and several hundred scents has been proposed (cf, Henry, 1997). Most species in the Neotropics. They include some of the species are probably univoltine, but many species most impressive examples of the order as both are known to take more than one year to complete adults and larvae. Adults resemble damselflies development, and there is a record of one species (Odonata: Zygoptera) because of their elongate requiring six years to complete developmentin cap­ bodies and generally feeble flight, but they have tivity (New, 1991). As previously mentioned this shorter bodies (typically), longer clubbed antennae family is cosmopolitan, however some broad bio­ and entirely different wing venation. The wings geographic patterns are apparent in the distribu­ may be clear, but in many species they are boldly tions of some higher taxa. For instance, the Pal­ patterned with cryptic spots and lineations. Wing­ parinae, a group with brightly colored or strongly span ranges from 35 to ca. 148 mm (New, 1991; patterned wings, are most diverse in the drier ar­ Henry et al., 1992).Antlion larvae are bizarre look­ eas of Africa, with a single genus, Dimares, found ing, typically having a ferocious aspect because of in the Neotropics. Species of Dimares are notable the prominent, sickle-shaped, mandibles. for their sexual dimorphism, with males having en­ Biology and geographical distribution. The biol­ tirely translucent wings, and females with exten­ ogy of the larvae of a few species has been studied sively patterned wings (Henry et al., 1992). thoroughly, but very little is known about the ma­ Taxonomy. The Myrmeleontidae belong in the jority of taxa. Most entomologists think of antlion monophyletic superfamily Myrmeleontoidea larvae as 'doodle-bugs' ('hormigas leon' in Span­ (Stange, 1994)or Myrmeleontiformia (Henry, 1997), ish) awaiting prey within their sand pits, but actu­ together with the Psychopsidae, Nemopteridae, ally only some species of the tribes Myrmeleontini Nymphidae, and Ascalaphidae. The higher classi­ and Dendroleontini build pits (Henry et al., 1992). fication of antlions has been somewhat unstable. Larvae of other subfamilies vary in habits, from Stange & Miller (1990),based primarily on studies actively pursuing prey on the ground, under the of larvae, recognized the subfamilies Stilbopterygi­ ground, or on trees. Free-living larvae differ from nae (Australia), Palparinae (worldwide, except the specialized digging habitus of pit builders (i.e., Australia and North America; 3 tribes), and Myrrne­ a ventrally curved neck region holding a flattened leontinae (worldwide, 5 tribes). Henry et al. (1992) head) and they might be similar to larvae of Asca­ recognized a slightly different scheme, Acantha­ laphidae. Most adults are cryptically colored. They clisinae (a tribe of Myrmeleontinae in the above typically rest during the day and become active at classification), Palparinae, Myrmeleontinae, and dusk or at night, although a few species are diur­ Stilbopteryginae. Stange (1994), in a cladistic analy­ nal. The adults of many species are attracted to sis of a large and varied group of taxa, recognized lights, often in great numbers in desert regions. 11 tribes and the subfamily Stilbopteryginae. Adults of nearly all species are predatory. In a study Keys for some Neotropical taxa are provided in Central Europe, Stelzl &Gepp (1990)found that in Henry et al. (1992; to genus), and the works of the fecesof four species contained mostly fragments Stange (1970a, b) and Miller & Stange (1989). NEUROPTERA (NEUROPTERIDA) I 573

Mexican Fauna. There are currently 97 valid spe­ ever, the two families are not distinct from each cies recorded in Mexico; however, the actual num­ other in all larval characters (Stange & Miller, ber of species in the country is higher and several 1990),and there is at least one antlion with reduced new species have been already detected and await claws (Navasoleon). description (Dr. Lionel Stange, pers. comm.). Be­ Biology and geographical distribution. Owlflies cause of incomplete sampling and unrecorded are cosmopolitan and can be found in temperate specimens in museums, the Mexican distribution and warmer climates, including arid environ­ data of most species are fragmentary. The currently ments. Adults of different species exhibit a wide known Mexican fauna slightly outnumbers the variety of activity modes, from distinctly diurnal, better studied fauna of America north of Mexico to crepuscular and nocturnal. Adults of many spe­ (93 valid species; Penny et al., 1997). cies have a distinctive resting position in which Following the classification scheme in Stange's the head is oriented downward and the abdomen catalog of world Myrmeleontidae (unpublished is held at a sharp angle from the resting substrate manuscript), and according to the incomplete cur­ (usually a narrow plant stem) so that it resembles rent record, Brachynemurini are the most species a twig or leaf petiole. Adult owlflies are typically rich group in Mexico (47 spp.), followed by Ne­ strong fliers and include the most supremely aerial moleontini (25 spp.), Myrmeleontini (12 spp., ge­ of all Neuropteran species. Adults of some spe­ nus Myrmeleon), Gnopholeontini (8 spp.), Acan­ cies reach flying heights of 10 m or more after com­ thaclisini (4 spp.), and Dendroleontini (1 sp.). plete darkness (Borror et al., 1989), which accen­ tuates difficulties for their capture by nonspecial­ Ascalaphidae ized collectors. The large size of the adult eyes is The family Ascalaphidae (Fig. 33.3), owlflies, is a probably related to the crepuscular or nocturnal unique and uncommonly collected group of Neu­ behavior of many species. Adult owlflies are ac­ ropterans. Adults of this family are large and ro­ tive aerial predators, many of which appear to use bust flyers with wingspans of 35 to 130mm (Henry their hairy legs to filter 'insect plankton' from the et al., 1992) and huge compound eyes (hence the air (Henry et al., 1992). common name). Many species superficially re­ Larval scalaphids are similar to antlions in be­ semble dragonflies in general body shape and ing strong predators with long sickle-shaped flight characteristics, although closer examination jaws. All ascalaphid larvae appear to be sit-and­ clearly reveals them as Neuropterans. Adult and wait, ambush style predators. Some species are larval owlflies are morphologically very similar known to camouflage themselves with particu­ to antlions (Myrmeleontidae), and these families late matter from their environment so that they are probably phylogenetic sister groups. The are indistinguishable from the substrate (e.g., Ulu­ adults of these groups can be distinguished as fol­ lodes). Larvae of other species (e.g., Ascaloptynx, lows. Owlflies typically have antennae about as Ascalobyas, and Haploglenius) are leaf litter dwell­ long as the body, while the antennae of antlions ers or live on leaves and branches of trees (Henry are usually only about as long as the head and et al., 1992). thorax combined. The eyes of ascalaphids are some Taxonomy. There are about 440 species described times subdivided near the middle by a shallow, in 90 genera. The family is subdivided in three horizontal, sulcus; the eyes of myrmeleontids are subfamilies: Albardiinae, Ascalaphinae, and always entire, never divided. Several details of the Haplogleniinae (New, 1989).The Albardiinae have venation and genitalia are also distinctive for these only one unusual species from tropical Brazil. The groups. Owlflies are often sexually dimorphic in Ascalaphinae are characterized by bisected com­ wing coloration and shape, and some times also pound eyes. Some species have a constricted waist in antennallength (Penny et al., 1997). The larvae and a yellow-and-black color pattern that makes of nearly all owlflies and antlions can be differen­ them closely resemble certain wasps. This group tiated by the presence of enlarged hind pretarsal occupies xeric to tropical and open to dense forest claws in antlions (related to antlion larval digging environments, with their larvae generally living behaviors), which are lacking in owlflies. How­ on the ground. The entire-eyed Haplogleniinae 574/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY tend to live in mesic, forested habitats (Henry et ASPOCK, H., U. ASPOCK & H. HOLZEL. 1980. DieNeurop­ al., 1992) and their larvae typically have arboreal teren Europas. 2 vols, Goecke and Evers, Krefeld,Germany. 495 and 355 p. habits. Most of the genera recorded from Mexico ASPOcK, U. 1986.The present state of knowledge of the fam­ can be identified using the keys in Henry et al. ily Berothidae (Neuropteroidea: Planipennia), pp. 87-101. (1992), except Ascaloptynx, which was not in­ InRecent Research in Neuropterology. Proceedings of the 2nd cluded. Other useful taxonomic references for the International Symposium on Neuropterology, J. Gepp, H. Mexican fauna are the monograph of the family Aspock and H. Holzel, eds., Graz, Austria. [Symposium held in Hamburg, 1984]. 176 p. by Weele (1909)and an unpublished PhD. disser­ ----.1992.Crucial points in the phylogeny of the Neu­ tation treating the Nearctic fauna by Shetlar (1977). roptera (Insecta), pp. 63-73.In Current Research in Neuropte­ Mexican fauna. Twenty one species are currently rology. Proceedings of the Fourth International Symposium reported from Mexico. This is almost three times on Neuropterology, M. Canard, H. Aspock, and M.W. Mansell, eds. Toulouse, France. the number of species known from north of Me­ ----& M. W. MANSELL. 1994.A revision of the fam­ xico (8 spp.; Penny et al., 1997), most of which (6 ily Rhachiberothidae Tjeder, 1959, stat. n. (Neuroptera). spp.) also occur in Mexico. Most species appear to Syst. Ent,. 19: 181-206. be Widespread in the country and extend south ----& H. L. NEMESCHKAL. 1998. A cladistic analy­ sis of the Berothidae (Neuroptera). in Panelius, S. P.(ed.). into Central America (e.g., Ululodes). Neuropterology 1997. Proceedings of the Sixth Interna­ tional Symposium on Neuropterology. Helsinki, Finland, 13-16July 1997. Acta Zoo1. Fenn., 209: 45-63. ACKNOWLEDGMENTS BANKS,N. 1897.NewNorth American neuropteroid insects. Trans. Amer. Ent. Soc., 24: 21-31. ----.1912. Notes on Nearctic Mantispidae. Proc. Ent. We are very grateful to the following friends and Soc. Wash., 14: 178-179. colleagues for providing information about Mexi­ BELKIN, J. N. 1954.Additional records of Oliarces clara in Ca­ can records of several groups: David E. Bowles lifornia and Arizona (Neuroptera, Ithonidae). Bull.South. (Texas Parks & Wildlife Department, Sisyridae), California Acad.Sci., 53: 65-68. BORROR,D.J.,C.A. TRIPLEHORN &N. F.JOHNSON. 1989. Kevin M. Hoffman (California Department of An introduction tothestudy ofinsects. 6th edition. Saunders Food and Agriculture, Mantispidae), Victor J. College Publishing, Philadelphia, USA, 875 p. Monserrat (Universidad Complutense de Madrid, BROOKS, S.J.1994.Ataxonomic review of the common green Hemerobiidae), and Lionel A. Stange (Florida State lacewing genus Chrysoperla (Neuroptera: Chrysopidae). Bull. Brit.Mus. Nat. Hist. (Ent.), 63(2): 137-210. Department of Agriculture, Myrmeleontidae). We ----& P.c. BARNARD. 1990. The green lacewings of also thank our home institutions for general sup­ the world: a generic review (Neuroptera: Chrysopidae). port, and in particular CAS and TAMU for help Bull. British. Mus. Nat. Hist. (EnU, 59(2): 117-286. with the elaboration of collection records data BROWN, H. P.1952.The life history of Climacia areolaris (Ha­ bases, on which to a great extent the species list in gen),a neuropterous 'parasite' of freshwater sponges. Amer. MidI. Nat., 47: 130-160. this chapter is based. Finally, thanks to J. Llorente BRUSHWEIN,J.R.1987. Bionomics of Lomamyia hamata (Neu­ and J. J. Morrone for their kind invitation to con­ roptera: Berothidae). Ann. Ent. Soc. Amer.,80: 671-679. tribute this chapter and especially for their encour­ CARPENTER, F.M. 1940.A revision of the Nearctic Hemero­ agement after we went beyond the deadline. biidae, Berothidae, Sisyridae, Polystoechotidae and Di­ laridae (Neuroptera). Proc. Amer. Acad.Arts and Sci., 74: 193-280. DEVRIES, P.J. 1986.TheButterflies ofCosta Rica andtheir Natural REFERENCES History. Papilionidae, Pieridae, Nyrnphalidae. Princeton Uni­ versity Press. Princeton, New Jersey. 327 p. ADAMS,P.A. 1950.Notes on Oliarces clara Banks (Neuroptera, ENDERLEIN, G. 1910. Klassifikation der Mantispiden nach dem material des Stettiner Zoologischen Museums. Stett. Ithonidae). Pan-Pac. Ent., 26: 137-138. Ent. Ztg., 71: 341-379. ----.1967.A review of the Mesochrysinae and Notho­ ERICHSON, W. F.1839. Beitrage zu einer Monographie von chrysinae (Neuroptera: Chrysopidae). Bull. Mus. Compo Mantispa, mit einleitenden Betrachtungen tiber die Ord­ Zoo1., 135: 215-238. nungen der Orthopteren und Neuropteren. Zeitschr. Ent. ----.1970. A review of the New World Dilaridae. Pos­ (Germar), 1: 147-173. tilla, 148: 1-30. FABRICIUS, J.C. 1775.Systema entomologiae, sistens insectorvm ----.1989. Anew genus of Berothidae from tropical classes, ordines, genera, species, adiectis synonyrnis, locis, des­ America, with two new species. Psyche, 96: 187-193. criptionibvs, observationibvs. Flensbvrgi et Lipsiae, 832 p. NEUROPTERA (NEUROPTERlDA) I 575

FAULKNER, D. K. 1990a. Current knowledge of the biology LAMBKIN, K. J. 1986. A revision of the Australian Mantispi­ of the moth-lacewing Oliarces clara Banks (Insecta: Neu­ dae (Insecta: Neuroptera) with a contribution to the clas­ roptera: Ithonidae), pp. 197-203In Advancesin Neuropterol­ sification of the family. I. General and Drepanicinae. Austr. ogy.Proceedings oftheThirdInternational SymposiumonNeu­ f. Zool., Suppl. Ser., 116: 1-142. ropterology, M. W. Mansell and H. Aspock, eds. South Af­ ----.1986b. A revision of the Australian Mantispidae rican Department of Agricultural Development: Pretoria. (Insecta: Neuroptera) with a contribution to the classifi­ [Symposium held in Berg en Dal, Kruger National Park, cation of the family II. Calomantispinae and Mantispinae. 1988.] 298 p. Austr. f. Zool., Suppl. Ser., 117: 1-113. ----.1990b.Phantom of the desert. Biology of the little­ LINSLEY,E. G. & J. W. MACSWAIN. 1955. Two new species known moth lacewing. Envir.West, San Diego 1990 (Win­ of Plega from Mexico (Neuroptera, Mantispidae). Pan-Pac. ter): 17-19. Ent., 31: 15-19. GALLARD, L. 1932.Notes on the feeding habits of the brown MACARTHUR, R. H. & E.O. WILSON. 1967.Thetheory ofisland moth-lacewing, Ithone fusca. Austr. Natur., 8: 168-170. biogeography. PrincetonUniversityPress.Princeton,New Jersey. GURNEY,A. B.1947.Notes on Dilaridae and Berothidae, with MACLEOD, E. G. 1964. A comparative morphological study special reference to the immature stages of the Nearctic of the head capsule and cervix of larval Neuroptera (In­ genera (Neuroptera). Psyche, 54: 145-169. secta). Ph.D. dissertation. Harvard University, Cambridge, HAGEN, H. A. 1861. Synopsis of the Neuroptera of North Massachusetts, USA. [iii] + 528 p. America, with a list of the South American species. Smith­ ----& P.A. ADAMS. 1967. A review of the taxonomy son. Misc. Coll., 4(1): xx + 1-347. and morphology of the Berothidae, with the description ----.1877. Symphrasis, eine neue Mantispiden-Gat­ of a new subfamily from Chile (Neuroptera). Psyche, 74: tung. Stett. Ent. Ztg., 38: 208-311. 237-265. HENRY, C. S. 1980. The importance of low-frequency, sub­ ----&P.E.SPIEGLER.1961.Notesonthelarvalhabi­ strate-borne sounds in lacewing communication (Neurop­ tat and developmental peculiarities of Nallachius america­ tera, Chrysopidae). Ann. Ent. Soc. Amer., 73: 617-621. nus(McLachlan)(Neuroptera: Dilaridae).Proc. Ent.Soc. Wash., ----.1997. Modem mating systems in archaic Holo­ 63: 281-286. metabola: sexuality in neuropterid insects, pp. 193-210 In MEINANDER, M. 1972. A revision of the family Conioptery­ J. C. Choe & B.J. Crespi (eds.). Theevolutionof mating sys­ gidae (Planipennia). Acta Zool. Fenn., 136: 1-357. tems in insectsandarachnids. Cambridge University Press, ----.1974a.Coniopterygidae from western North Am­ Cambridge, U.K. erica (Neuroptera). Ent. Scand.,5: 217-232. ----.,N. D. PENNY &P.A. ADAMS. 1992.The neurop­ ----.1974b. Coniopterygidae from South and Central teroid orders of Central America (Neuroptera and Me­ America (Neuroptera). Not. Ent., 54: 97-106. galoptera), pp. 432-458 In D. Quintero & A. Aiello (eds.). ----.1975.Coniopterygidae from North America (Neu­ Insects ofPanama andMesoamerica. Oxford University Press, roptera). Not. Ent., 55: 28-32. Oxford, 692 p. ----.1981. A review of the genus Coniopteryx (Neurop­ HOFFMAN, K. M. 1992.Systematics of the Mantispinae (Neu­ tera, Coniopterygidae). AnnalesEntomologici Fennici [=Suo­ roptera: Mantispidae) of North, Central, and South Am­ men hyonteistieteellinen aikakauskirja] 47: 97-110. erica. Ph.D. Dissertation. Clemson University. 501 p. ----.1983. Coniopterygidae from South and Central ----. & J. R. BRUSHWEIN. 1989. Species of spiders America II (Neuroptera). Neuroptera Int., 2: 179-198. (Araneae) associated with the immature stages of Mantispa ----.1990. The Coniopterygidae (Neuroptera, Plani­ pulchella (Neuroptera: Mantispidae). f. Arachnol., 17: 7-14. pennia). A check-list of the species of the world, descrip­ ----.& .1990. Spider (Araneae) taxa asso­ tions of new species and other new data. Acta Zool.Fenn., ciated with the immature stages of Mantispa interrupta (Neu­ 189: 1-95. roptera: Mantispidae). Ent. News, 101: 23-28. ----.1995.Coniopterygidae (Neuroptera, Planipennia) HOLZEL, H. & P. Ohm. 1990. Zoogeographical features of from Costa Rica and Nicaragua. Brenesia, 43/44: 61-70. Madagascan Chrysopidae (Insecta: Neuroptera), pp.167­ MILLER, R. B. & L. A. STANGE. 1989. Revision of the genus 181 In Current Research in Neuropterology. Proc. Fourth Int. Dimarella Banks (Neuroptera: Myrmeleontidae). Insecta Symp. Neuropterology, M. Canard, H. Aspock, and M. Mundi, 3: 11-40. Mansell, eds. Toulouse, France. MONSERRAT, V.J. 1984a. Contribuci6n al conocimiento de HUNGERFORD, H. B. 1931. Concerning the egg of Polystoe­ los Coniopterigidos de Mexico (Neuroptera, Planipennia, choiespunctatus Fabr. (Neuroptera). Bull. Brooklyn Ent. Coniopterygidae). EOS, 60: 211-222. Soc., 26: 22-23. ----.1984b. Sobre las especies americanas del genero JOHNSON,J.B.&K. S.HAGEN. 1981.Aneuropterous larva uses Notiobiella Banks, 1909,1. (Neuroptera, Planipennia, He­ an allomone to attack termites. Nature (London), 289:506-507. merobiidae), pp. 85-97 In J. Cepp, H. Aspock, and H. Hol­ KUKALOVA-PECK, J. 1991. Chapter 6. Fossil History and zel (eds.). Progress in World'sNeuropterology. Proceedings of the Evolution of Hexapod Structures. In CSIRO (ed.). The the First InternationalSymposium on Neuropterologv, Graz, Insects of Australia. Volume 1. Cornell University Press, Austria, 265 p. Ithaca, New York, 542 p. ----.1985.Lista de los tipos de Mecoptera y Neuropte­ KUWAYAMA, S. 1962. A revisional synopsis of the Neurop­ ra (Insecta) de la colecci6n L. Navas, depositados en el tera in Japan. Pac. lns.; 4: 325-412. Museo de Zoologfa de Barcelona. Misc. Zool., 9: 233-243. 576/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

----.1986. Longinos Navas, his neuropterological comments on the evolution of the hemerobiid forewing work and collection, pp. 173-176 In RecentResearch in Neu­ radial vein. Syst. Ent., 18: 363-370. ropterology. Proceedings of the 2nd International Symposium ----.1994. A new phylogenetically basal subfamily on Neuropterology, J. Gepp, H. Aspock, and H. Holzel, eds. ofbrownlacewingsfrom Chile (Neuroptera: Hemerobiidae). Craz, Austria. 176 p. Ent. Scand., 25: 295-302. ---. 1990a. Systematic studies on Hemerobiidae (In­ PARFIN, S. 1.& A. B.GURNEY. 1958.The spongilla-flies, with secta: Neuroptera), pp. 67-88 In M. W. Mansell and H. special reference to those of the western hemisphere (Sisy­ Aspock (eds.), Advances in Neuropterology, Proceedings of ridae, Neuroptera). Proc. U. S. Natl. Mus., 105: 421-529. theThirdInternational SymposiumonNeuropterology, Preto­ PARKER, F. D. & L. A. STANGE. 1965. Systematic and bio­ ria, R. South Africa, 298 p. logical notes on the tribe Platymantispini (Neuroptera: ---. 1990b. Asystematic checklist of the Hemerobi­ Mantispidae) and the description of a new species of Plega idae of the world (Insecta: Neuroptera), pp. 215-262 In M. from Mexico. Can. Ent., 97: 604-612. W. Mansell and H. Aspock (eds.), Advancesin Neuropterol­ PENNY, N. D. 1978. Lista de Megaloptera, Neuroptera e Ra­ ogy,Proceedings oftheThirdInternational SymposiumonNeu­ phidioptera do Mexico, America Central, ilhas Carafbas ropterology, Pretoria, R. South Africa, 298 P: e America do Sul. Acta Amaz., 7(4)(Suplemento): 1-61. ---.1996. Revisi6n del genero Hemerobius de Latino­ ----. 1983. Neuroptera of the Amazon Basin. Part 8. america (Neuroptera, Hemerobiidae). Fragm. Eni.,Roma. Berothidae. Acta Amaz., 13: 689-695. 27(2): 399-523. ----.1996. A remarkable new genus and species of ---.1997. Revision del generoMegalomus de Latino­ Ithonidae from Honduras (Neuroptera). J. Kansas EnL Soc. america (Neuroptera, Hemerobiidae Fragm. Eni., Roma, 69: 81-86. 29(1): 123-206. ----,P.A. ADAMS & L. A:STANGE. 1997.Species Cata­ ----.1998.Nuevosdatos sobre los hemer6bidos de Ame­ log of the Neuroptera, Megaloptera, and Raphidioptera rica (Neuroptera: Hemerobiidae). J. Neuropierol., 1: 109-153. of America north of Mexico. Proc. Calif.Acad. Sci., 50(3): ---.2000. Revision del Cenero Nusalala (Neuroptera, 39-114. Hemerobiidae). Fragrn. Eni., Roma, 32(1): 83-167. ---- & J. R. ARIAS. 1982. Insects of an Amazon Forest. ----. & J. BAYO. 1995. Nuevos datos sobre los coni­ Columbia University Press, New York, New York. 269 p. opterigidos de Mejico central (Neuroptera: Conioptery­ ---- & C. A. da COSTA. 1985. Mantispideos do Brasil gidae). Bol. Asoc. Espanola Ent., 19: 125-130. (Neuroptera: Mantispidae). Acta Amaz., 13: 601-687. ----.& Z. DERETSKY. 1999.New faunistical, taxonomic ---- & V. J. MONSERRAT. 1985. Neuroptera of the and systematic data on brown lacewings (Neuroptera: He­ Amazon Basin. Part 10. Hemerobiidae. Acta Amaz., 13 merobiidae). J. Neuropierol., 2: 45-66. (1983): 879-909. NAVAs, L. 1914a. Nevropteres nouveaux de I'Amerique du ----.& H. STURM. 1984. A flightless brown lacewing Nord. Heme serie. Ent. Zeitschr., Frankfurt am Main, 28: from Colombia (Neuroptera, Hemerobiidae). Spixiana, 18-20, 25-26. Miinchen,7: 19-22. ----.1914b. Mantispidos nuevos (Segunda [II] serie). POIRRIER, M. A. & Y.M. ARCENEAUX. 1972. Studies on Mem. Real Acad. Cienc. Artes Barcelona, (3)11: 83-103. southern Sisyridae (spongilla-flies) with a key to the third­ ---. 1928. Insectos neotr6picos. 3a serie. Rev. Chil. instar larvae and additional sponge-host records. Amer. Hist. Nat., 31: 316-328. MidI. Naiur., 88: 455-458. NEW, T. R. 1980. A revision of the Australian Chrysopidae PORION, T. 1994.Fulgoridae 1. Catalogue illustre de la faune (Insecta: Neuroptera). Austr.J. Zool., Suppl. Ser.,77: 1-143. americaine. Sci. Nat. Venette, France, 72 p. ----.1989. Planipennia (Lacewings). Handbuch der PUPEDIS, R. J. 1987. Foraging behavior and food of adult Zoologie, Vol. IV; Part30. Walter de Gruyter, Berlin, 123 P: Spongila-flies [sic] (Neuroptera: Sisyridae). Ann. Ent. Soc. ---.1991. Neuroptera, pp. 525-542 In CSIRO (ed.). Amer., 80: 758-760. The Insects of Australia, Vol. I. Cornell University Press, REDBORG, K. E. & E. G. MACLEOD. 1983.Climaciella brunnea Ithaca, New York, 542 p. (Neuroptera: Mantispidae): a mantispid that obligately OPLER, P. A. 1981. Polymorphic mimicry of polistine wasps boards spiders. J. Nat. Hist., 17: 63-73. by a neotropical Neuropteran. Biotropica, 13: 165-176. ----& . 1984. Maintenance feeding of first OSWALD, J. D. 1988. A revision of the genus Sympherobius instar mantispid larvae (Neuroptera, Mantispidae) on spi­ Banks (Neuroptera: Hemerobiidae) of America north of der (Arachnida, Aranea) hemolymph.J. Arachnol., 11:337-341. Mexico with a synonymicallist of the world species. J. ----& .1985. The developmental ecology New York Ent. Soc., 96: 390-451. of Mantispa uhleriBanks (Neuroptera: Mantispidae). Illi­ ----.1990. Revision of the Neotropical brown lacew­ nois BioI. Mon., 53: 1-130. ing genus Nomerobius (Neuroptera: Hemerobiidae). Ann. REHN, J.W.H. 1939. Studies in North AmericanMantispidae. Eni. Soc. Amer., 83: 18-29. Trans. Amer. Ent. Soc., 65: 237-264. ----.1993a.Revision and cladistic analysis of the world RIEK, E.F.1974. The Australian moth-lacewings (Neuroptera: genera of the family Hemerobiidae (Insecta: Neuroptera Ithonidae), J. Austr. Ent. Soc., 13: 37-54. J. New YorkEnt. Soc., 101(2): 143-299. SAY, T. 1824. Order Neuroptera, pp. 303-310 In Narrative ofan ----.1993b. A new genus and species of brown lacew­ expeditionto the sourceof St. Peter's River,LakeWinnepeek, ing from Venezuela (Neuroptera: Hemerobiidae), with Lakeof the Woods, &c. &c. performed in the year 1823, by NEUROPTERA (NEUROPTERIDA) /577

order of the Hon. l.C. Calhoun, Secretary of War, under the G. Rudebeck, eds. Volume 12. Swedish Natural Science command of Stephen H. Long, Major U.S.T.E., WH. Keating, Research Council, Stockholm. ed. Vol. 2. Carey and Lea, Philadelphia. ----.1968. The genus Mucroberotha Tjed. and its sys­ ----. 1825. Mantispa. Plate 25 and unpaginated let­ tematic position (Neuroptera). Ent. Tidskr., 89: 3-19. terpress [7 pp1in American entomology, or descriptions TSUKAGUCHI, S.1995. ChrysopidaeofJapan (Insecta, Neurop­ of the insects of North America. Vol. 2. Philadelphia. iera). Privately published, Osaka, Japan. 223 p. SHETLAR, D. 1977. The biosystematics of the Nearctic Asca­ WEELE, H. W., VAN DER. 1909. Ascalaphiden. Monogra­ laphidae (Insecta: Neuropteroidea, Planipennia), with notes phisch Bearbeitet. Collections zoologiques du Baron Ed­ on biology and morphology. Unpublished PhD. disser­ mond de Selys Longchamps. Cat. Syst. Descr.. Fasc. VIII, tation, Pennsylvania State University, 233 p. 326p. STANGE, L. A. 1970a. A generic revision and catalog of the WEISSMAIR, W. 1999. Prairnaginale Stadien, Biologie und western Hemisphere Glenurini with the description of a Ethologie der europaischen Sisyridae (Neuropterida: Neu­ new genus and species from Brazil (Neuroptera: Myr­ roptera). Stapfia, 60: 101-128. meleontidae). Los Angeles County Mus., Contrib. Sci., 186: WELCH, P.S.1914. The early stages of the life history of Polys­ 1-28. toechotes punctatus Fabr. Bull. Brooklyn Ent. Soc; 9: 1-6. ----. 1970b. Revision of the ant-lion tribe Brachyne­ WESTWOOD,J.O.1867. Descriptions of new species of Man­ murini of North America (Neuroptera: Myrmeleontidae). tispidae in the Oxford and British Museums. Trans. Eni, Univ. Calif. Publ. Eni., 55:vi + 1-192 p. Soc. London, (3)5: 501-508. ----.1994. Reclassification of the New World antlion WIESENBORN, W. D. 1998. High seasonal rainfall precedes genera formerly included in the tribe Brachynemurini Oliarces clara Banks (Neuroptera: Ithonidae) spring emer­ (Myrmeleontidae). Insecta Mundi, 8(1-2): 67-119. gence. Pan-Pac. Eni., 74: 217-222. ----& R. B. MILLER. 1990. Classification of the Myr­ WITHYCOMBE, C. L. 1925. Some aspects of the biology and meleontidae based on larvae (Insecta: Neuroptera), pp. morphology of the Neuroptera. With special reference to 151-169 In M. W. Mansell & H. Aspock (eds.). Advances in the immature stages and their possible phylogenetic sig­ Neuropterology. Proceedings of the Third International nificance. Trans. Ent. Soc. London 1924: 303-411. Symposium on Neuropterology. Ber en Dal, Kruger Na­ tional Park, R. S. A., 1988, Government Printer, Pretoria, South Africa, 298 p. APPENDIX 33.1. STELZL, M. & J. GEPP. 1990. Food analysis of imagines of central European Myrmeleontidae (Insecta: Neuroptera), pp. 205-210 In M. W. Mansell & H. Aspock (eds.). Advances Checklist of the Neuroptera of Mexico. in Neuropterology. Proceedings of the Third International Symposium on Neuropterology. Ber en Dal, Kruger Na­ The followingchecklistcontains the names ofallvalid, tional Park, R. S. A., 1988, Government Printer, Pretoria, South Africa, 298 p. extant, Neuropteran species that have been reported TAUBER, C. A. 1969. Taxonomy and biology of the lacewing from Mexico in the scientific literature or that are genus Meleoma(Neuroptera: Chrysopidae). Univ. Calif. Publ. known to the authors to occur in Mexico from mate­ Ent., 58: vi + 1-94 p. rial that they have examined or from other reliable ----& T. de LE6N. 2001. Systematics of green lacew­ sources. Several species that are known junior syn­ ings (Neuroptera: Chrysopidae): Larvae of Ceraeochrysa from Mexico. Ann. Eni. Soc. Amer., 94: 197-209. onyms, but which have not yet been formally desig­ ----&M.J. TAUBER. 1968. Lomamyia latipennis (Neu­ nated as such in the literature, have been omitted from roptera, Berothidae) life history and larval descriptions. the list to avoid the misleading impression that they Can. Ent., 100: 623-629. are valid species.Distributions are reported at the state TAUBER,M. J.,c. A. TAUBER, KM. DANNE & KS. HAGEN. level,where known. Extralimital distributions are not 2000. Commercialization of Predators: Recent Lessions from Gree Lacewings (Neuroptera: Chrysopidae: Chryso­ given, but the distributions of most species extend perla). Amer. Eni., 46(1): 26-38. either north or south (or both) of Mexico. TILLYARD,R.J. 1922.The life-history of the Australian moth­ The checklist is presented in alphabetical or­ lacewing, Ithone[usca, Newman (Order Neuroptera Plani­ der by family-group name, then by genus and pennia). Bull. Ent. Res., 13: 205-223. species. Abbreviations: n. comb.= newcombination; TJEDER, B. 1959. Neuroptera-Planipennia. The Lace-Wings of Southern Africa. 2. Family Berothidae, pp. 256-314 In n. rec.= new record for Mexico. South African Animal Life, B. Hanstrom, P. Brinck and G. Rudebec, eds. Vol. 6. Swedish Natural Science Research Ascalaphidae Council, Stockholm. Ascalaphinae ----.1966. Neuroptera-Planipennia. The Lace-Wings 1. Ameropterus mexicanus (Van der Weele, 1909) CHIS of Southern Africa. 5. Family Chrysopidae, pp. 228-534 2. A. trivialis (Gerstaecker, 1888) CHIS In South African Animal Life, B. Hanstrom, P. Brinck and 3. A. versicolor (Burmeister, 1838) VER 578/ JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

4. Cordulecerus mexicanus Van der Weele, 1909 CHIS TAB 42. C. quadripunctata Burmeister, 1838 CHili, CHIS 5. C. subiratus (Walker, 1853) Mexico (no further locality) 43. C. comanche (Banks, 1938) BCS, CHili, CHIS, COAH, DF, 6. Ululodes arizonensis Banks, 1907 BCS, CHIS, CHili, HGO, DGO, GRO, GTO, JAL, MICH, MOR, NAY, NL, OAX, PUE, JAL, NAY, OAX, SIN, SON, VER SIN, SON, TAMPS, VER, ZAC 7. U. bicolor (Banks, 1895) BCS, CHIS, JAL, MICH, NAY, NL, 44. C. exotera (Navas, 1914) BCS, CHIS, COAH, DF, JAL, OAX, SIN, SLP, SON, TAMP, VER MICH, MOR, NAY, NL, OAX, SLP, SON, VER 8. U. cajennensis (Fabricius, 1787) BCS, CHIS, GRO, SIN, TAB, 45. C. extema (Hagen, 1861) BCS, CHIH, CHIS, COL (includ­ TAMP, VER, YUC ing I. Revillagigedo), DF,DGO, GRO,JAL, MICH,MOR, NAY, 9.U. maclenyanus (Gooding, 1825)CHIH, CHIS, COL, GRO, HGO, OAX, PUE, SIN, SLp, TAMPS, YUC JAL,MICH, NL, OAX,QR, SIN, SLP,SON, TAMPS, VER,YUC 46. C.harrisii (Fitch, 1855) Mexico (no further locality) 10. U. mexicanus (Mclachlan, 1871) CHIS, GRO, JAL, MOR, 47. C. mexicana Brooks, 1994 SIN OAX, SON, TAMPS 48. C. plorabunda (Fitch, 1855) AGS, BCS, CHili, DF, HGO, 11. U. quadripunctatus (Burmeister, 1838) BCS JAL, MICH, NAY, NL, PUE, SIN, SLP, SON, VER, ZAC 12. U. subvertens (Walker, 1853) BCS, CHIS, OAX, SIN, SLP, 49. C. rufilabris (Burmeister, 1838) CHIH, COL, DF, GTO, JAL, SON, TAMPS, VER NAY, TAMP, VER 13. U. tuberculaius (Banks, 1901) QR 50. Chrysopodes (Neosuarius) collaris (Schneider, 1851) BCS, CHIS, COAH, COL (I. Revillagigedo), GRO, NAY,OAX, SIN, Haplogleniinae SON, TAMP, VER, YUC 14.Amoea immaculata (Olivier, 1790) CHIS, DGO, GRO, JAL, 51. Dichochrysa perfecta (Banks, 1895) [nocomb.] BCS, SIN NAY,SIN, VER 52. D. sierra (Banks, 1924) [nocomb.] BC 15.A. iniqua (Walker, 1853) CHIS 53. D. triangularis (Adams, 1978) [no comb.] HGO, JAL, 16.A. vacua (Gerstaecker, 1894) CHIS, YUC MICH,PUE 17.Ascalobyas albistigrna (Walker, 1853) CHIS, TAB 54. Eremochrysa (Chrysopiella) sabulosa (Banks, 1897) CHIH, 18.A. microcerus (Rambur, 1842)DGO, NAY, SLP,TAB,VER HGO, SLp, SON 19. Ascaloptynx appendiculata (Fabricius, 1793) CHIS, MOR, 55. E. (EJ altilis Banks, 1950 BC NAY,SIN, SON, TAMP 56. E. (E.)diguetiNavas, 1911 BC 20. Haploglenius luteus (Walker, 1853) NAY, VER 57. E. (E.) fraterna (Banks, 1897) BC 21.Neohaploglenius flavicomis (Mcl.achlan, 1871) CHIS, HGO, 58. E. (E.)hageni Banks, 1903 BC JAL, QR, SIN, VER, YUC 59.E.(EJ punctinerois (Mclachlan,1869)AGS,BCS,CHIH, COAH, DGO, HGO, JAL, NL, OAX, PUE, SLP,SON, ZAC Berothidae 60. E. (E.) tibialis Banks, 1950 BCS, DGO, ZAC Berothinae 61.Meleoma adamsi Tauber, 1969 MEX, MICH 22.Lomamyia squamosa Carpenter, 1940 BCS, CHIS, NAY,OAX, 62. M. antennensis Tauber, 1969 CHIS PUE, QRO, SLP, TAMPS 63. M. carapana Adams, 1969 MICH 23. Lomamyia sp. TAMPS 64. M. colhuaca Banks,1949 DF,HGO, MEX, MOR, QR, QRO 65. M. dolicharthra (Navas, 1914) CHIS, HGO, MICH Chrysopidae 66. M. emuncia (Fitch, 1855) CHIH, TLAX Chrysopinae: Beionopterygini 67.M.furcata (Banks, 1911) CHili 24. Abachrysa sp. JAL 68. M. hageni Banks, 1949 DF, MEX 25.Nacarina cordillera (Banks, 1910) [nocomb.] CHIS, DGO, PUE 69. M. innovata (Hagen, 1861) Mexico (no further locality) 70. M. macleodi Tauber, 1969 CHIS, DF, HGO, MEX, MICH, PUE Chrysopinae: Chrysopini 71. M. mexicana Banks, 1898 MEX 26.Ceraeochrysa cincta (Schneider, 1851) CHIS,COL, NAY,SIN 72. M. nahoa (Banks, 1949) DF, MEX 27. C. claveri (Navas, 1911) VER 73. M. pinalena (Banks, 1950) CHili, MICH 28. C. cubana (Hagen, 1861) SON, TAMP, VER 74. M. pipai Tauber,1969DGO, HGO, MEX,MICH, PUE, TLAX 29. C.effusa (Navas, 1911)Mexico (no further locality) 75. M. powelli Tauber, 1969 DGO 30. C. elegans Penny, 1998 TAMP 76. M. schwarzi (Banks, 1903) MEX 31. C. everes (Banks, 1920) OAX, YUC 77. M. tezcucana (Banks, 1949) DF, HGO, MOR 32. C.gradata (Navas, 1913) CHIS 78. Ninetananina (Banks, 1911) MICH 33. C. indicata (Navas, 1914) VER 79. Plesiochrysa brasiliensis (Schneider, 1851) CHIS,JAL,NAY, SON,VER 34.C.lateralis (Cuerin-Meneville, 1844) [nocomb.] CHIS, VER 80. P. elongata (Navas, 1913) CHIS, MOR, PUE, VER 35. C. placita (Banks, 1908) MICH 81.Yumachrysa apache (Banks, 1938)BCS,CHIH,JAL, PUE, SON 36. C. sanchezi (Navas, 1924) BCS, OAX, VER 82.Y.incerta (Banks, 1895)[no comb.] BC,MOR, OAX, TAMP,ZAC 37. C. smithi (Navas, 1914) BCS, COAH, VER 38. C. valida (Banks, 1895) BCS, COL, JAL, NAY, OAX, SLP, Chrysopinae: Leucochrysini TAB, TAMP, VER, YUC 83. Gonzaga torquatus Navas, 1913 CHIS, COL, VER 39. Chrysopa coloradensis Banks, 1895 BCS 84. Leucochrysa (L.) arizonica (Banks, 1906) JAL, MEX, MICH 40. C. nigricomis Burmeister, 1838 DGO 85. L. (L.) negata (Navas, 1913) DF 41. C. oculata Say, 1839 CHili, DF, DGO, JAL 86. L. (L.) serrula Adams, 1979 MICH, MOR, SLP NEUROPTERA (NEUROPTERIDA) I 579

87. L. (Nodita) americana Banks, 1897 BCS, OAX 130. C. californica Meinander, 1972 DF 88. L. (NJ antennalis (Navas, 1932)Mexico (no further locality) 131. Semidalis angusta (Banks, 1906), BCS, DGO, eTO, MEX, 89. L. (NJ antennata Banks, 1905 BCS, PUE, VER MICH, MOR, NAY (I. Marias), OAX, QRO, SIN, SON 90. L. (NJ askanes (Banks, 1946) OAX 132. S. boliviensis (Enderlein, 1905) CHIS, VER 91. L. (NJ callota Banks, 1915 ZAC 133.S. byersi Meinander, 1972DGO, eTO, HeO, MICH, MOR, 92. L. (NJ explorata (Hagen, 1861) Mexico ("Cordova") NAY, PUE, VER 93. L. (N.) mexicana Banks, 1900 VER 134. S. deserta Meinander, 1974 BC 94.L. (NJ navasi (Kimmins, 1940) Mexico (no further locality) 135.S.faulkneri Meinander, 1990 SIN 95. L. (NJ nigrinervis (Banks, 1939) BC 136. S.flinti Meinander, 1972 BC, BCS 96. L. (NJ pavida (Hagen, 1861) Mexico ("Cordova") 137. S.frommeri Meinander, 1974 SON 97. L. (NJ salleana (Navas, 1911)JAL 138. S. manausensis Meinander, 1980 VER 98. L. (NJ sulcata (Navas, 1921) JAL 139. S. mexicana Meinander, 1972 BCS, DGO, HeO, MICH, NL, SIN, SLP,SON Chrysopinae: tribe/genus Incertae Sedis 140. S. soleri Monserrat, 1985 VER 99."Chrusopa" aztecana Banks, 1903Mexico (no further locality) 100."C." brevihirta Banks, 1946Mexico (no further locality) Dilaridae 101. "e." fascialis Banks, 1905Mexico (no further locality) Nallachiinae 102."e." leptana Banks, 1914Mexico (no further locality) 141. Nallachius pulchellus (Banks, 1938) NAY 103.T." senior (Navas, 1928)Mexico (no further locality) Hemerobiidae Nothochrysinae Hemerobiinae 104. Pimachrysa albicostales Adams, 1967 BC 142. Hemerobiella sinuataKimmins, 1940 CHIS 143. Hemerobius alpestris Banks, 1908 CHIS, DF, DCO, MEX, Coniopterygidae VER, JAL, MICH, OAX, PUE, TAMP Aleuropteryginae: Aleuropterygini 144. H. bolivari Banks, 1910 CHIS 105. Aleuropteryx longipennis Meinander, 1974 BC 145. H. discretus Navas, 1917 AeS, BC, CHIS, COL, DF, DCO, 106. A. maculipennis Meinander, 1972 BCS eRO, eTO, HeO, JAL, MEX, MICH, MOR, NL, OAX, PUE, 107.A. punctata Meinander, 1974 BCS,SIN TAMPS, TLAX, VER 108. A. simillima Meinander, 1972 BC, SIN, SON 146. H. edui Monserrat, 1991 HeO, MICH, PUE, SLP 109. A. unicolor Meinander, 1972 BCS 147. H. elongatus Monserrat, 1990 MEX 148. H. gaitoi Monserrat, 1996 VER Aleuropteryginae: Fontenelleini 149. H. hernandezi Monserrat, 1996 CHIS, OAX, VER 110. Bidesmida morrisoni Johnson, 1977 BC 150. H. hirsuticornis Monserrat & Deretsky, 1999TAMPS 111.Neoconis inexpectata Meinander, 1972 JAL 151. H. humulinus Linnaeus, 1758 VER 112.N. marginata Meinander, 1972 BC, BCS,JAL, SON 152. H. jucundus Navas, 1928 CHIS, DF, eRO, HeO, JAL, 113. N. unam Monserrat, 1985 VER MEX, MICH, OAX, PUE, TAMPS, VER 153. H. kokaneeanus Currie, 1904 BC Coniopteryginae: Coniopterygini 154.H. martinezae Monserrat, 1996CHIS,eRO,MICH,OAX,VER 114. Coniopteryx (e.) callangana Enderlein, 1906 YUC 155. H. pacificus Banks, 1897 BC, DF, MEX, OAX, SON, VER 115.e.(e.) delta Johnson, 1981 VER 156. H. stigmaStephens, 1836 BCS, MEX 116. e. (C.) fitchi Banks, 1895 eTO, MICH, PUE 157. H. tibialis Navas, 1917 [nomen dubium] Mexico (no fur­ 117.e. (C.) mexicana Meinander, 1974 SIN ther locality) 118.e. (e.) minutaMeinander, 1972 BCS,SIN 158. H. tolimensis Banks, 1910 MEX, VER 119. C. (C.) palpalis Meinander, 1972 BCS, DCO, eTO, HeO, 159. H. withycombei (Kimmins, 1928) TAB, VER MICH, OAX, QRO, VER 160. Wesmaelius longipennis (Banks, 1920) CHIH, DCO, MOR 120.e. (e.) simplicior Meinander, 1972BCS,COL (SocorroIsland) 121. e. (C.) westwoodi (Fitch, 1855) QRO Megalominae 122. C. (Scotoconiopteryx) flinti Meinander, 1975 SLP 161.Megalomus fidelis (Banks, 1897) NL 123.e. (S.) isthmicola Meinander, 1972SLP 162. M. minorBanks, 1905 CHIS, COL, JAL, MOR, NAY,SLP, 124. e. (Xeroconiopteryx) diversicornis Meinander, 1972 BCS, TAB, TAMPS, VER DCO,NL, VER 163. M. moestus Banks, 1895 COL, JAL, MOR, SON, TAMPS, 125.e.(XJ meinanderi Johnson, 1981 SIN VER,ZAC 126. e.(X.) texana Meinander, 1972 BC 164.M. pictus Hagen, 1861Mexico (no further locality) 127. e.obscura Navas, 1934 MOR 128. Parasemidalis fuscipennis (Reuter, 1894) BC, DeO Microminae 165. Micromus fuscatus (Nakahara, 1965) CHIS, MICH, QRO Coniopteryginae: Conwentziini 166. M. posticus (Walker, 1853) CHIS, DF, eRO, MICH, MOR, 129. Conwentzia barretti (Banks, 1898) BC, CHIS, DF, DCO, NAY,NL, SLP,TAMPS, VER eTO, HeO, MEX, MICH, MOR, PUE, VER 167. M. subanticus (Walker, 1853) BCN, BCS, CHIH, COAH, 580 I JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

JAL, NL, SIN, SON 200. P. dactylota Rehn, 1939 [norec.] BCS, CHIH, COAH, JAL, 168. M. variolosus Hagen, 1886 BCN, CHIH, SON SIN'SON 169. Nusalala championiKimmins, 1936 VER 201. Pfratercula Rehn, 1939 [norec.] BCS, CHIH 170.N. kruegeri Nakahara, 1965 Mexico (no further locality) 202. P.fumosa Linsley & MacSwain, 1955 MICH 203. P melitomae Linsley & MacSwain, 1955 CHIS Notiobiellinae 204. P. signata (Hagen, 1877) BCS 171. Notiobiella mexicana Banks, 1913 JAL, SLP 205. P.variegata Navas, 1928 PUE, VER 172. N. spinosa Monserrat & Penny, 1983 Mexico (no fur­ 206. P. yucatanaeParker & Stange, 1965 CHIS, YUC ther locality) 207. Trichoscelia banksiEnderlein, 1910 CHIS 208. T. santareni (Navas, 1914) QROO, TAB Sympherobiinae 209. T. tobari (Navas, 1914) TAB 173. Sympherobius angusius (Banks, 1904) Be, COAH, DF, DCa, GTO, JAL, MEX, MICH, NL, OAX Mynneleontidae 174. S. arizonicus Banks, 1911 BCN, BCS, CHIH, OAX, PUE, Myrmeleontinae: Acanthaclisini SON,ZAC 210. Paranihaclisis congener (Hagen, 1861) BCS 175. S. axillaris Navas, 1928 Mexico (no further locality) 211.P hageni (Banks, 1899) BCS,CHIH, COAH, NAY,SLp,SON 176. S. barberi (Banks, 1903) BCN, BCS, CHIS, COAH, COL, 212.Vella fallax (Rambur, 1842) BCS,CHIS, COAH,HGO,JAL, HGO, MEX, OAX, PUE, SON, TAMPS MICH, NAY, OAX, PUE, SIN, SLP, SON, TAB 177. S. bifaseiatus Banks, 1911 BCS, DGO 213. V. mexicana Navas, 1932 Mexico 178. S. catifornicus Banks, 1911 BCN, BCS 179. S. distinetus Carpenter, 1940 GRO Myrmeleontinae: Brachynemurini 180. S. killingtoni Carpenter, 1940 BCN, CHIH, COAH, ZAC 214.Abatoleon dorsalis (Banks,1903)CHIS,NL,TAMPS,VER,YCC 181.S.perparous(McLachlan, 1869) BCS,CHIH, COAH,SON 215. Atrieholeon parkeri (Stange, 1970) COL, PUE 182.S.p~tus(Banks,1904)NL 216. A. tubereulatus (Banks, 1899) JAL, SIN, SON 183 S. pupillus Navas, 1915 [nomen dubium] Mexico 217. Brachynemurus abdominalis (Say, 1823) SIN 184 S. similis Carpenter, 1940 CHIS, MICH, MaR, NL, VER 218. B. californicus Banks, 1895 BCS, SIN, SON 185 S. subeostalis Monserrat, 1990 CHIS, JAL, VER, YUC 219. B. divisus (Navas, 1928) MICH, PUE 220. B. elongatus Banks, 1904 CHIH Ithonidae 221. B.ferox (Walker, 1853) BCS, SON 186. Narodona mexicana Navas, 1930 COL 222. B.fuscus (Banks, 1905) GRO, JAL, MICH, MOR, NAY, 187. Oliarces n. sp. BCS SIN,SON 223. B. hubbardii Currie, 1898 CHIH, DCa, MICH, SIN, 50:'\. Mantispidae VER,ZAC Calomantispinae 224. B. mexieanusBanks, 1895 CHIH, DCO, JAL, MICH, NAY, 188. Nolima praeliator Navas, 1914 GRO OAX, PUE, SIN 189. N. victor Navas, 1914 CHIS, GRO, OAX 225. B. pulehellus Banks, 1911 BCS, SON 226. B. sackeni Hagen, 1888 BCS, CHIH, COAH, DGO, HGO, Mantispinae JAL, MICH, NL, PUE, SLp, SON, TAMP 190. Climaciella brunnea (Say in Keating, 1824) BCS, CHIH, 227. B. uersuius (Walker, 1853) DF, GTO, JAL, MEX CHIS, COL, DCa, GRO, HGO, JAL, MICH, MaR, NL, OAX, 228. Chaetoleon pusillus (Currie, 1899) BCS, CHIH PUE, QR, QRO, SIN, SLP, TAMPS, VER 229. C. tripunctatus (Banks, 1922) SON 191.Enianoneura batesella (Westwood, 1867) In. ree.] CHIS,JAL 230. C. variabilis Banks, 1942 BCS 192.Mantispacompellens (Walker, 1860) CHIS, OAX, SLP, VER 231. Clathroneuria arapahoe Banks, 1938 SON 193. M. inierrupta (Say, 1825) [no rec.] BCS, CHIH, CHIS, 232. C. eoqui/letti (Currie, 1898) BCS, CHIH, SON, ZAC COAH, COL (1. Revillagigedo),GRO, HGO,JAL, MICH, MaR, 233. C. navajo Banks, 1938 SON NAY,NL, OAX, PUE, SIN, SLP, SON, TAMPS, VER, YUC 234. C. schwarzi (Currie, 1903) BCS, COAH, NL 194. M. minuta (Fabricius, 1775) In. rec.] CAMP, CHIH, CHIS, 235. Dejuna mimica (Stange, 1970)CAMp, CHIS, OAX, VER,YUC COL, GRO, HGO, JAL, MaR, NAY, NL, OAX, PUE, QR, SIN, 236. D. persimilis (Stange, 1970)CAMP, CHIS, OAX, VER,voc SLP, TAB, TAMPS, VER, YUC 237. D. setosa (Stange, 1970) CHIS, OAX, VER 195. M. pulchella (Banks, 1912) BCS, CHIH, CHIS, HGO, JAL, 238. D. straminea (Stange, 1970) SIN, SON MICH, NL, OAX, SIN, SLP, TAMPS, VER 239. Enrera mexicana Navas, 1915 [nomen dubium] TAMPS 196. M. sayi (Banks, 1897) CHIS, DCa, NL, QR, SIN, TAB, 240. Mexoleon mixteeus (Stange, 1970) PUE TAMPS, VER 241. M. papago (Currie, 1899) BCS, CHIH, NL, SON 197. M. scabrosa (Banks, 1912) NL 242. Seotoleon carrizonus(Hagen, 1888) BCS, CHIH, NL, SIN, 198. M. viridula (Erichson, 1839) In. rec.] COL, JAL, NAY, SON, TAMPS QROO, SLP, VER 243. S. digueti (Navas, 1913) BCS 244. S. dissimilis (Banks, 1903) BCS, CHIH, SIN, SON Symphrasinae 245. S. eiseni (Banks, 1908) BCS, SON 199. Plega banksiRehn, 1939 In. rec.] CHIH, SIN, SON 246. S. expansus (Navas, 1913) BCS, CHIH, SON NEUROPTERA (NEUROPTERIDA) / 581

247. S. fidelitas (Adams, 1957) BC, SON 280.M. uniformisNavas, 1920CHIS,JAL,NAY, OAX,SON, VER 248. S. intermedius (Currie, 1903) BC, SON 281. Nehornius dampji Navas, 1928 YUC 249. S.longipalpis (Hagen, 1888)BCS,CHIH, COAH, DCa, SON 250. S. minusculus (Banks, 1898) BCS, CHIH, COAH, SON, Myrmeleontinae: Nemoleontini TAMPS, VER 282. DimareIla (D.) menkei Stange, 1963 COL, SIN, SON 251. S. minuius (Adams, 1957) BC, SON 283. D. (D.) mixteca Miller in Miller & Stange, 1989 CHIS, 252. S. niger (Currie, 1898) BC, CHIH, SON OAX, VER 253. S. nigrescens (Stange, 1970) BC 284. D. (D.) nayaritaStange in Miller & Stange, 1989JAL, NAY 254. S. nigrilabris (Hagen, 1888) BC, CHIH, DCa, SON 285. D. (D.) psammophila Stange, 1963 CRO, NL, OAX, SLP, 255. S. pallidus (Banks, 1898) BC TAB, VER 256. S. peninsulanus (Banks, 1942) BCS, SIN, SON 286. D. (D.) totonecaMiller in Miller & Stange, 1989 VER 257. S. peregrinus (Hagen, 1861) BCS, CHIH, SIN, SON 287. Elachyleon punctipennis Esben-Petersen, 1927 CHIS 258. S. quadripunciaius (Currie, 1898) BCS, SON 288. Eremoleon femoralis (Banks, 1942) BCS, SIN, SON 259. S. singularis (Currie, 1903) BCS 289. E. genini (Navas, 1924)Mexico (no further locality) 260. S. yavapai (Currie, 1903) BC 290. E. gracilis Adams, 1957 BC 291. E. insipidus Adams, 1957 BC Myrmeleontinae: Dendroleontini 292. E. longior Banks,1938 YUC 261. Dendroleon speciosus Banks, 1905 BC 293. E. macer (Hagen, 1861) MaR 294. E. nigribasis Banks, 1920 BCS, DCa, SON Myrmeleontinae: Gnopholeontini 295. E. paIlens Banks, 1941 BC 262. Cnopholeon barberi (Currie, 1903) BC, SIN 296. Clenurus luniger Cerstaecker, 1894 BCS, CHIS, JAL, 263. C. delicaiulus (Currie, 1903) BC SIN, SON, VER 264. C. zapotecus Stange, 1970 MaR, OAX 297. C. proi Navas, 1930 CHIS, SON 265. Maracandula apicalis (Banks, 1901) DCa, JAL 298. C. snowii Banks, 1907 BC, JAL, NAY, SIN, SON 266. M. n. sp. OAX 299. Psammoleon albouarius (Banks, 1942) BCS, SON 267. M. pygmaea (Hagen, 1861) Mexico (no further locality) 300. P. arizonensis Banks, 1935 SON 268. Menkeleon bellulus (Banks, 1905) BC, CHIH 301. P. bistictus (Hagen, 1861) Mexico 269. Tyttholeon puerilis Adams, 1957 BC 302. P. connexus (Banks, 1920) BCS 303. P.guttipes Banks, 1906 TAMP Myrmeleontinae: Myrmeleontini 304. P. normalis Banks, 1942 BCS 270. M. arizonicus Banks, 1943 BCS, CHIS, COL (1. Revilla­ 305. P. paraIlelus Banks, 1935 Mexico gigedo), NAY (1. Marias), SIN, SON, VER 306. P. sinuaius Currie, 1903 BC, CHIH 271. M. californicus Banks, 1943 BC 272. M. exitialis Walker, 1853 BC, SIN Polystoechotidae 273. M. immaculaius DeCeer, 1773 ZAC(?) 307. Polystoechotes punciaia (Fabricius, 1793) CHIH, DCa, 274. M. laemargus Navas, 1914 SIN MICH, MaR, SIN 275. M. mexicanus Banks, 1903 OAX 276. M. n. sp. SON Sisyridae 277. M. rusiicus Hagen, 1861 BCS 308. Climacia areolaris (Hagen, 1861) SLP,TAMPS 278. M. iexanus Banks, 1900 TAMP 309. C. califomica Chandler, 1953 SON 279. M. timidus Cerstaecker, 1888 CHIS, COL, NAY, SLP, 310. C. chapini Parfin & Gurney, 1956 CHIH, CHIS, VER SON, VER 311. Sisyra apicalis Banks, 1908 QR