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Home , Chrysopini, Climaciella, Coniopteryx, Hemerobiinae, Nallachius, Neuropterida

(NEUROPTERIDA)

John D. Oswaldl, Atilano Contreras-Ramos2& Norman D. Penny3

RESUMEN.En este capitulo se presenta un panorama difficult to encounter. They probably attain their sobre la sistematica, biologia y distribution geogrAfi- greatest abundance (but not diversity) in desert ca de 10s Neuroptera (Planipennia) de Mexico, con communities and in a variety of temperate habi- una onentacion hacia la literatura taxonomica. 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 parbntesis): Myrme- dows are often covered with hundreds of adult leontidae (97), (81), -(44), lacewings attracted to the lights. (36), (22), Neuroptera have two distinctive characteristics (21), Sisyridae (4), Ithonidae (2), (2), Dila- that make them fascinating creatures. First, they ridae (1) y Polystoechotidae (1).La fauna total de Neu- are predators, especially as larvae, giving them the r~~teraactualmenteregistrada en el pais suma 311 es- distinction of helping protect us from a wide vari- pecies. Como en otros casos, el orden ha sido estudiado ety of agricultural and horticultural pests (Tauber so10 superficialmente en Mexico, por lo 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, , and Ascalaphidae can be quite colorful, and at a distance can be mis- Neuroptera, or lacewings, , and their al- taken with large butterflies. lies, is one of the smaller orders of , 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 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 cons~cuousgroups, such as butter- Department of , Texas A & M University, College Station, Texas 77845-2475, USA. e-mail: [email protected] (DeVries, 1986) and fulgorid bugs (Porion, 2Centrode Investigaciones Biol6gicas, Universidad Aut6no- 1994),however, the tropical parts of the Neotropi- ma del Estado de Hidalgo, Apdo. Postal 1-69, Plaza Jugrez, cal region appear to contain the richest taxonomic I Pachuca, Hidalgo 42001, MBXICO. assemblage in the world. Penny (1978) listed 1100 I e-mail: [email protected] species and subspecies of Neuroptera for America I Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, the United Or than 25% USA. e-mail: NPennyWalAcademy.org the known world fauna, including 171 taxa from 560 1 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

Mexico. Our taxonomic knowledge of the has 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 are restricted to specialists in the country. Also, we are not aware of Australia. Species of , , graduate students being trained in systematics of Rapismatidae and 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 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.).It is evident, therefore, that there is room for entomologists within the country to fe cus on the study of the different taxonomic and bie 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 Naviis 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., Yale University, Florida State graphs it is now possible to identify almost all Collection of ). One of us (ACR) has adult Mexican Neuropterans to (e.g., Henry started collecting efforts in Central Mexico, par- et a1 ., 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 4 family level, although Stange (1994) provides keys to tribes of larval Myrmeleontidae and gen- PHYLOGENY era of larval Brachynemurini. Tauber & De Le6n (2001) have published a key to larval Ceraeo- The evolution of Mexican Neuroptera must be .t chrysa for Mexico. The vast majority of larval viewed in a global context. Neuroptera have a rich I NEUROPTERA (NEUROPTERIDA)1561 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 inkhe lineage containing Psy- much more diverse past. *. chopsidae, vymphidae, Nemopteridae, Myrme- We are in great need of a better phylogeny for lebntidae, and Ascalaphidae. extant lineages of Neuroptera. ~;chof the basis Elements of subfamilial and tribal phylogeny for current relationships 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. Under Withycombe'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 instars (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, , biogeographical regions -the Nearctic region in Berothidae, Rhachiberothidae and Mantispidae. the north and higher elevations and the Neotro- Within this lineage Coniopterygidae have anrun- pica1 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,-aiound 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 IVevrorthidae. 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 Aspijck & Mansell, 1994; Aspijck & 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 . More problematical but untoothed, larval mandibles to grasp prey is the presence of the ithonid genera Narodona and 562 1 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NO IRMAN D. PENNY

Oliarces. Ithonidae has a bimodal distribution in TAXA IN MEXICO Australia and the region from southwestern U.S.A. to Honduras. The simplest explanation 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 rnm), 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 Mymzeleon larval pits nalic structures. Because of their small size and under houses and bridges, Ceraeochysa in or- waxy body covering, coniopterygids often super- chards, and in grasslands, pastures, ficially ressemble adult whiteflies. and backyards. When life histories of these insects 'AlthoughJuncommonly~ coll&cted by iion- are studied, almost invariably they 'are fohd to neuropterists; -coniopterygids can be extremely be beneficial to mankind. abundant lokally where favorable prey conditions As for many other groups of plants Ahd 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 adult coniopterygidshave 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 that it was de- made without dissection after a working knowled- 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- Adults are also generally predaceous, but may 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 prey, including mites, scale insects, help develop this foundation. insect eggs, coccids, aphids and phylloxerans. NEUROFTERA (NEUROFTERIDA)1 563

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 undesaibed 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 very likely be more than 50. 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 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. 100 spe- 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 northem 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 231 in 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 rarely collected because of their remote habitats and the rapid rate of description of new species in this unusual life histories. 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 Ithonefusca (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 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 on the fluids of plant root 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 southem 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, spongllaflies,is a small group gations, generally associated with a local elevated of Neuropterans that is closely associated with prominence, to which females 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 Oliarces 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 (Wiesenbom, 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 Ithonefusca. 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 Ithone and Oliarces. 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: Oliarces 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 Merendbn, Cusuco). At least two additional generally covered by the female with a fine silken NEUROITERA (NEUROITERIDA)I 565

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). Bio1ogy:and geographical distribution. Most of Taxonomy and geographic distribution. The fam- what we know about the biology of Dilaridae ily Sisyridae contains ca. 60 described species that comes from.the studies of are collectively distributed throughout the New (McLachlan) bytMacLeod & Spiegler (1961). Lar- and Old Worlds. The New World fauna consists vae were collectedander 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 insectalarvae 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 Mexican sisyrid 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 clzapini, C. calfomica, 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 southern U.S.A.and extends south into northeast- Africa. In the New World they are found from ern Mexico. The second, 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. calfornica, 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- I apicalis, is known from Cuba, southeastern U.S.A., chiinae. There are seventeen described species, Central America and northern South America, and has most from South America. Only four species are been collected on the Yucatan Peninsula @. 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 pulchellus, 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 nocturna, 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. pulchellus distinctive family of Neuropterans. They are rela- @a&) exist from 16 miles north of Mazatlh (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 [Navis] 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. 566 /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 and hyaline wings appearance of small preying mantids. The family with variable dark markings. The phylogenetic is cosmopolitan in tropical and temperate parts of affinitiesof 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, but one species, (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 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 I 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 extinct over large parts Larvae of 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 4 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 (southern Canada 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 I 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 '4 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 9 medium-sized group of Neuropterans (300 species; subfamilies (Lambkin, 1986).Two African genera, Aspock et al., 1980) immediately recognizable by Mucroberotha Tjeder and Rhachiberotha Tjeder, vari- 2 NEUROPTERA (NEUROPTERIDA)1567

Figs. 33.1-3. Habitus of adult Neuroptera: 1) Plega signata (Hagen), Mantispidae (redrawn from Henry et a1.,1992); 2) Chaetoleon pusillus (Currie), Myrmeleontidae (redrawn from Stange, 1994); 3) Ululodes mexicanus (McLachlan), Ascalaphidae (redrawn from Henry et a1 ., 1992). ously been have placed in the Berothidae CTjeder, unpublished Ph.D. thesis).This thesis contains much 1959)and Mantispidae (Tjeder, 1968),but currently 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 (Aspijck et a1 ., 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 & NOIRMAN 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 Taxonomy and 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, 1sp ., 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 , 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 sheet at 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 thisc initial phylogenetic hypothesis of intergeneric re- family are those of Tauber & Tauber (1968), for lationships within the family. Lomamyia latipennis, and Brushw>ein(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, Loma- are apparently obligate termitophiles in the gal- myia 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 Lomamyia species are known to occur in the termites. Prey are apparently immobilized by a desert regions the southwestern United States. It is poison injected with a quick thrust of the straight almost certain that future sampling will reveal at larval jaws. The report by Johnson & Hagen (1981) least some of these in northern Mexico. It is also that L. 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 oviposition behav- research due to their economic importance as po- NEUROPTERA (NEUROPTERIDAI 1 569 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 theractual number of valid Neotropical 199Clb).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). mrn (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 andlyses of intra- MAbeing 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 subfamily has been 1992 [fig.28.171). 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 a1 ., 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 (17 spp.) and Symphero- tats occupied by 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, howeverdt 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. food as well. They are usually medium-sized (wing length about Examples of cosmopolitan genera are Hemero- 15 rnm), pale green wing venation and body, with bius and Micromus, but many genera are small and golden eyes. The general green coloration and trans- geographically restricted (e.g., Biramus from Ven- parent wings makes them very difficult to see when ezuela [Oswald, 1993b1, Adelphohemerobius from resting under leaves. Adults of some species, such Chile [Oswald, 19941). In the Nearctic, Penny et al. as those in the genus , 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 1 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY biologcal 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 15 to more than 30 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 can be 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 Ch ysoperla, genera and approximately 180 described species, have short setae on the body and bear no debris, all but 15 of them in the genus Leucoch ysa. Mem- while other larvae, such as Ceraeoch ysa and Latco- bers of this tribe are easily recognized by their long ch ysa, have long thoracic and abdominal setae on antennae, dark pterostigmal spot, and low angle tuberacles and carry bits 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- may be one to several generations per year. 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. a 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 (81 species: Penny et a1 ., 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 ) and four tribes of Chrysopinae species: Aspock et al., 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 (Adarns, 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 gradate crossveins; jugal lobe well developed - Hindwing with two gradate series of cross- ...... (subfamily Nothochrysinae)Pimach ysa veins...... 9 - In forewing SPsm meets outer gradate crossveins; jugal lobe reduced or absent ...... 8. Forewing with one gradate series of crossveins...... (subfamily Chrysopinae) 2 ...... Ere a (Ch ysopiella) -Forewing with two gradate series of crossveins.... 2. Pterostigma with dark spot anteriorly on both ...... Eemoch a (Eremochysa) 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 homs 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 unrno'dified ...... 10 meeting Psm at almost a 90° angle ...... 5 10. Male ninth stemite subapically constricted cre- 3. Forewing marked with black or dark brown ating small apical node; red gena only facial spots; pterostigmal spot extends to radial ve in... markings; no scape stripes ...... Ch ysoperla ...... Gonzaga - Male ninth stemite 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- from may have antennal rings or transverse red jacent to crossveins; pterostigmal spot extends or dark bands; antennal s'cape may or may not only to subcostal ve in...... (g enus Leucoch ysa) 4 have stripes or spots...... I1

4. Cell im quadrangula...... Leucoch ysa (Leucochysa) 11. Male genital elements include gonapsis...... l2 - Cell im ovate or triangular ...Leucochysa (Nodita) - Male genital elements lack gonapsis ...... 13

5. Antennal flagellomeres broad& than long; prono- 12. Gonapsis elongate, rod-like; forewing cross- I tum much broader than long; forewing cell C, veins usually without dark margining of cross- longer than cell C,subcostal crossvein located veins...... Ceraeoch ysa halfway between origin of M2and R+M,; api- - Gonapsis cross-shaped with lateral arms; fore- cal palp segment swollen ...... wing crossveins heavily bordered with dark ...... tribe Belonopterygini) 6 margining ...... Pseudomaflada - Antennal flagellomeres more than twice as long as wide; pronotum longer than broad (except 13. Pseudopenis present; tignum usually pre- Chrysopodes collaris and ); forewing cell sent ...... 14 C, as long or longer than C,; subcostal crossvein - Pseudopenis present; tignurn absent ...... 17 much closer to origin of M2than origin of R+M,; apical palp segments cylindrical and apically 14. Partial or complete dark ring around base of tapered...... ( tribe Chrysopini) 7 antennae ...... Ch spa(in part) -No indication of dark ring at base of antennae... 15 6.Forewing longitudinal veins alternating dark and pale; rows of dark spots on thoracic nota ..... 15. Pronoturn bears no spots or stripes ...... Abach ysa ...... Ch ysopa nigricornis ! - Forewing longitudinal veins pale; pronotum - Pronotum bears four red or orange spots...... 16 may have longitudinal red stripe, but no dark spots...... Nacarina 16. Pronotum bears four red spots; radial cross- veins darkened in middle ...... Plesioch ysa 7. Hindwing with one gradate series of crossveins - Pronotum bears four orange spots; radial cross- 572 1 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY i

17. Dark lateral scape stripe; hindwing generally of insects (i.e., Heteroptera, Diptera, , with only 1-3 inner gradate veins....Yumach ysa 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 ...... Ch ysopodes is almost entirely unknown, but several species have been found to communicate via male phero- Myrmeleontidae mones secreted by specialized exocrine glands (cj., The Myrmeleontidae (Fig. 33.2), or antlions, 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 (cj., 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 development, and there is a record of one species (: Zygoptera) because of their elongate requiring six years to complete development in 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 a1 ., 1992). 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 a1 ., 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 larvae as 'doodle-bugs' ('hormigas le6n1 in Span- (Stange, 1994) or (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 a1 ., 1992). fication of antlions has been somewhat unstable. I 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 subfamiliesStilbopterygi- i ground, or on trees. Free-living larvae differ from nae (Australia), (worldwide, except I I the specialized digging habitus of pit builders (i.e., Australia and North America; 3 tribes), and Myrme- I a ventrally curved neck region holding a flattened leontinae (worldwide, 5 tribes). Henry et al. (1992) I 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 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, recogruzed , lights, often in great numbers in desert regions. 11 tribes and the subfamily Stilbopteryginae. Keys for some Neotropical taxa are provided i Adults of nearly all species are predatory. In a study in Central Europe, Stelzl & Gepp (1990) found that in Henry et al. (1992; to genus), and the works of the feces of four species contained mostly fragments Stange (1970a, b) and Miller & Stange (1989). NEUROFTERA (NEUROFTERIDA) I573

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 (~hvasoleon). 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. ~dultsof 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 130 mrn (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 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 Hiploglenius) 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 antenna1 length (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 thPir larvae generally living behaviors), which are lacking in owlflies. How- on the ground. The entire-eyed Haplogleniinae 574 1 JOHN D. OSWALD, ATILANO CONTRERAS-RAMOS & NORMAN D. PENNY

tend to live in mesic, forested habitats (Henry et ASPOCK, H., U. ASPWK & H. HOLZEL.1980. Die Neurop- al., 1992) and their larvae typically have arboreal term Europas. 2vols. Goecke and Evers, Krefeld, Germany. 495 and 355 p. habits. Most of the genera recorded from Mexico ASPWK, 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- In Recent 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, 19841.176 p. by Weele (1909) and an unpublished Ph.D. disser- 1992. Crucial points in the phylogeny of the Neu- tation treating the Nearctic fauna by Shetlar (1977). roptera (Insects), pp. 63-73. In Current Research in Neuropte- Mexican fauna. Twenty one species are currently rology. Proceedings of the Fourth InternationalSymposium 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. Arevision 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, 13-16 July 1997. Acta Zool. Fenn., 209: 45-63. ACKNOWLEDGMENTS BANKS, N. 1897. New North 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 ~e~artmentof An introduction to the study of insects. 6th edition. Saunders Food and Agriculture, ~antis~idaer~ktor J. College Publishing, Philadelphia, USA, 875 p. Monserrat (~niversidad~om~lutknse de Madrid, BROOKS, S. J. 1994. Ataxonomic review of the common green Hemerobiidae),and Lionel A. Stange (Florida State lacewing genus Chysoperla (Neuroptera: Chrysopidae). Department of Agriculture, Myrmeleontidae).We Bull. Brit. Mus. Nat. Hist. (Ent.), 63(2): 137-210. & 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. (Ent.), 59(2): 117-286. with the elaboration of collection records data BROWN, H. P. 1952. The life history of Climacia areolans (Ha- bases, on which to a great extent the species list in gen), a neuropterous 'parasite' of freshwater sponges.Amer. Midl. Nut., 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. The Butterflies of Costa Rica and their Natural REFERENCES Histoy. 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. Comp. Mantispa, mit einleitenden Betrachtungen iiber die Ord- Zool., 135: 215-238. nungen der Orthopteren und Neuropteren. Zeitschr. Ent. 1970. Areview of the New World Dilaridae. Pos- (Germar), 1: 147-173. tilla, 148: 1-30. FABRICKJS, J. C. 1775. System entomologiae, sistens insectorvm 1989. A new genus of Berothidae from tropical classes, ordines, genera, species, adiectis synonymis, locis, des- America, with two new species. Psyche, 96: 187-193. criptionihs, obseruationibvs. Flensbvrgi et Lipsiae, 832 p. FAULKNER, D. K. 1990a. Current knowledge of the biology LAMBKIN, K. J. 1986. A revision of the Australian Mantispi- of the moth-lacewing 01zarces clara Banks (Insecta: Neu- dae (Insecta: Neuroptera) with a contribution to the clas- roptera: Ithonidae), pp. 197-203In Advances in Neuropterol- sification of the family. I. General and Drepanicinae. Austr. ogy. Proceedings ofthe Thrrd International Symposium on Neu- I. ZOO^., 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 11. Calomantispinae and Mantispinae. 1988.1 298 p. Austr. I. 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. Envlr. West, San Diego 1990 (Wi- 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. 0.WILSON. 1967. The theory oflsland moth-lacewing, Ithone fusca. Austr. Natur., 8: 168-170. biogeography. Princeton Umversity Press. 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. sects). 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. Smlth- & P. A. ADAMS. 1967. A review of the taxonomy son. Mlsc. Coll., 4(1): xx + 1-347. and morphology of the Berothidae, with the description 1877. Symphmsls, 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. Notes on the larval habi- strate-bome sounds in lacewing communication (Neurop- tat and developmental peculiarities of Nallachius amerlca- tera, Chrysopidae). Ann. Ent. Soc. Amer., 73: 617-621. nus (McLachlan)(Newoptera:Dilaridae). Proc. Ent. Soc. Wash., . 1997. Modern matlng systems in archaic Holo- 63: 281-286. metabola: sexuality in neuropterid insects, pp. 193-210 In MEINANDER, M. 1972. Arevision of the family Conioptery- J. C. Choe & B. J. Crespi (eds.). The molution ofmating sys- gidae (Planipennia). Acta Zool. Fenn., 136: 1-357. tems in rnsects and arachnids. Cambridge University Press, 1974a. Coniopterygidae from westem North Am- Cambridge, U.K. erica (Neuroptera). Ent. Scand., 5: 217-232. ., N. D. PENNY & P. A. ADAMS. 1992. The neurop- 1974b. Conlopterygidae 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.Conlopterygidae from North America (Neu- Insects of Panama and Mesoamenca. Oxford University Press, roptera). Not Ent., 55: 28-32. Oxford, 692 p. 1981.Areview of the genus Comopteryx (Neurop- HOFFMAN, K. M. 1992. Systematics of the Mantispinae (Neu- tera, Coniopterygidae).Annales Entomolopa Fennlci [=Suo- roptera: Mantispidae) of North, Central, and South Am- men hyonteistieteellinen aikakauski ja] 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 I1 (Neuroptera). Neuroptera Int., 2: 179-198. (Araneae) associated with the immature stages of Mantispa . 1990. The Coniopterygidae (Neuroptera, Plani- pulchella (Neuroptera: Mantispidae). I. 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. Nms, 101: 23-28. .1995. Coniopterygidae (Neuroptera, Plampennia) 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). lnsecta Symp. Neuropterology, M. Canard, H. Aspock, and M. Mundi, 3: 11-40. Mansell, eds. Toulouse, France. MONSERRAT, V. J. 1984a. Contribuci6n a1 conocimiento de HUNGERFORD, H. 8.1931. Concerning the egg of Polystoe- 10s Coniopterigidos de Mexico (Neuroptera, Planipennia, chotes punctatus Fabr. (Neuroptera). Bull. Brooklyn Ent. Coniopterygidae). EOS, 60: 211-222. SOC.,26: 22-23. 1984b.Sobre las especies americanas del gknero JOHNSON, J B. & K. S. HAGEN. 1981. Aneuropterous lama uses Notlobzella Banks, 1909, I. (Neuroptera, Planlpennia, He- an allomone to attack termites.Nature (London),289: 506-507. meroblidae), pp. 85-97 In J. Gepp, H. Aspock, and H Hol- KUKALOVA-PECK, J. 1991. Chapter 6. Fossil History and zel (eds.). Progress in World's Neuropterology .Proceed~ngs of the Evolution of Hexapod Structures. In CSIRO (ed.). The the First International Symposzum on Neuropterology, Graz, Insects of Australia. Volume 1. Cornell University Press, Austria, 265 p. Ithaca, New York, 542 p. 1985. Lista de 10s tipos de y Neuropte- KUWAYAMA, S. 1962. A revisional synopsis of the Neurop- ra (Insecta) de la colecci6n L. Navks, depositados en el tera in Japan. Pac. Ins.,. 4: 325-412. Museo de Zoologia 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 Recent Research in Neu- radial vein. Syst. Ent., 18: 363-370. ropterology. Proceedings of the Znd Internatlonal Symposium 1994. A new phylogenetically subfamily on Neuropterology, J. Gepp, H. Aspock, and H. Holzel, eds. of brown lacewings from Chile (Neuroptera:Hemerobiidae). Graz, Aushia. 176 p. Ent. Scdnd., 25: 295-302. 1990a. Systematic studies on Hemerobiidae (In- PARFIN, S. I. & A. B. GURNEY. 1958. The spongilla-flies,wlth sects: 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. the Third International Symposium on Neuropterology, 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. A systematic 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.),Advances in Neuropterol- PENNY, N. D. 1978. Lista de , Neuroptera e Ra- ogy, Proceedings of the Third Internatlonal Symposium on Neu- phidioptera do Mexico, America Central, ilhas Caraibas ropterology, Pretoria, R. South Africa, 298 p. e America do Sul. Acta Amaz., 7(4)(Suplemento):1-61. 1996. Revision del genero Hemerobius de Latino- 1983. Neuroptera of the Amazon Basin. Part 8. america (Neuroptera, Hemerobiidae). Fragm. Ent., Roma, Berothidae. Acta Amaz., 13: 689-695. 27(2): 399-523. . 1996. A remarkable new genus and species of . 1997. Revision del genero Megalomus de ~atino- Ithonidae from Honduras (Neuroptera). J. Kansas Ent. Soc. america (Neuroptera, Hemerobiidae Fragm. Ent., Roma, 69: 81-86. 29(1): 123-206. ,P. A. ADAMS & L. A:STANGE. 1997.Species Cata- 1998. Nuevos datos sobre 10s hernerbbidos de Am& log of the Neuroptera, Megaloptera, and Raphidioptera rica (Neuroptera:Hemerobiidae). J. Neuropterol., 1: 109-153. of America north of Mexico. Proc. Callf. Acad. Sci., 50(3): ,2000. Revision del Genero Nusalala (Neuroptera, 39-114. Hemerobiidae). Fragm. Ent., Roma, 32(1):83-167. & J. R. ARIAS. 1982. Insects ofan Amazon Forest. . & J. BAYO. 1995. Nuevos datos sobre 10s 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. Espafiola Ent., 19: 125;130. (Neuroptera: Mantispidae). Acta Amaz., 13: 601-687. . & Z. DERETSKY. 1999. New fahistical, taxonomic & V. J. MONSERRAT. 1985. Neuroptera of the and systematic data on brown lacewings (Neuroptera: He- Amazon Basin..Part 10. yep~robiida-e.Acta Amaz., 13 , > merobiidae). J. Neuropterol., 2: 45-66. (1983)': 879-909. % . . . - .- . NAVAS, L. 1914a. Nevroptsres nouveaux de 1'Amerique du . &?:'sTUkM. 1984. A flightless l~;o&n lacewlng Nord. Iisme serie. Ent. ~eitschr.:Frankfurt am Main, 28: from ~olombia'(Newoptera, Hemefobiidae). Splxlana, 18-20, 25-26. Munchen, 7: 19-22. 1914b. Mantispidos nuevos (Segunda [11] serie). POIRRIER, M. A. & Y. M. ARCENEAUX. 1972. Studies on Mem. Real Acad. Cienc. Artes Barcelona, (3)ll:83-103. southern Sisyridae (spongilla-flies)with a key to the third- . 1928. Insectos neotropicos. 3a serie. Rev. Chil. instar larvae and additional sponge-host records. Amer Hist. Nut., 31: 316-328. Mldl. Natur., 88: 455-458. I NEW, T. R. 1980. A revision of the Australian Chrysopidae PORION, T. 1994. Fulgoridae 1. Catalogue illustre de la fame (Insecta: Neuroptera). Austr. J.Zool., Suppl. Ser., 77: 1-143. americaine. Sci. Nut. Venette, France, 72 p. . 1989. Planipennia (Lacewings). Handbuch der PUPEDIS, R. J. 1987. Foraging behavior and food of adult Zoolope, Vol. Iy Part 30. 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 ofAustralta, Vol. I. Cornell University Press, REDBORG, K. E. & E. G. MACLEOD. 1983. Cllmac1ella brunnea Ithaca, New York, 542 p. (Neuroptera: Mantispidae): a mantispid that obligately 1 OPLER, P. A. 1981. Polymorphic mimicry of polistine wasps boards spiders. J. Nut. Hlst., 17: 63-73. by a neohopical Neuropteran. Biotropica, 13: 165-176. & 1984.Maintenance feeding of first I 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 (Arachnids, Aranea) hemolymph. I. Arachnol., ll:337-341. Mexico with a synonymical list of the world species. J. & 1985. The developmental ecology New York Ent. Soc., 96: 390-451. of Mantispa uhleri Banks (Neuroptera: Mantispidae). Illz- 1990. Revision of the Neohopical brown lacew- nozs Blol. Mon., 53: 1-130. i ing genus Nomeroblus (Neuroptera: Hemerobiidae). Ann. REHN, J.W. H. 1939.Studies in North American Mantispidae. ! Ent. 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 York Ent. Soc., lOl(2): 143-299. SAY, T. 1824. Order Neuroptera, pp. 303-310 In Narrativeof an 1993b. Anew genus and species of brown lacew- expedition to the source of St. Peter's River, Lake Wlnnepeek, ing from Venezuela (Neuroptera: Hemerobiidae), with Lake ofthe Woods, bc. bc. performed in the year 1823, by I NEUROPTERA (NEUROYTERIDA)/ 577

order of the Hon. J.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., W.H. Keatzng, Research Council, Stockholm. ed. Vol. 2. Carey and Lea, Philadelphia. 1968. The genus Mucrobuotha Tjed. and its sys- . 1825. Mantispa. Plate 25 and unpaginated let- tematic position (Neuroptera). Ent. Tzdskr., 89: 3-19. terpress [7 pp] in American entomology, or descriptions TSUKAGUCHI, S. 1995. Chysopidae of Japan (Insecta, Neurop- of the insects of North America. Vol. 2. Philadelphia. tua). Privately published, Osaka, Japan. 223 p. SHETLAR, D. 1977. The biosystematiis 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 Ph.D. disser- mond de Selys Longchamps. Cat. Syst. Descr.. Fasc. VIII, tation, Pennsylvania State University, 233 p. 326 p. STANGE, L. A. 1970a. A generic revision and catalog of the WEISSMAIR, W. 1999. Pralmaginale Stadien, Biologie und western Hemisphere Glenurini with the description of a Ethologie der europaischen Sisyndae (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.0.1867. Descriptions of new species of Man- murlnl of North America (Neuroptera: Myrmeleontldae) tispidae in the Oxford and British Museums. Trans. Ent. Unzv. Callf. Publ. Ent., 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 Mundz, 8(1-2):67-119. gence. Pan-Pac. Ent., 74: 217-222. & R. B. MILLER. 1990. Classification of the Myr- WITHYCOMBE, C. L. 1925. Some aspects of the biology and meleontldae based on larvae (Insecta: Neuroptera), pp. morphology of the Neuroptera. With special reference to 151-169 In M. W. Mansell & H. Aspock (eds.).Advances zn 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- tlonal 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. zn Neuropterology. Proceedings of the Thlrd Internatjonal Symposium on Neuropterology. Ber en Dal, Kruger Na- The following checklist contains the names of all valid, tlonal 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. Pu bl. 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 LE~N.2001. ~istematicsof green lacew- sources. Several species that are known junior syn- ings (Neuroptera: Chrysopidae): Larvae of Ceraeochysa from Mexico. Ann. Ent. 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 hlstory 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, K.M. DANNE & K.S. HAGEN. level, where known. Extralimital distributionsare not 2000. Commercialization of Predators: Recent Lessions from Gree Lacewings (Neuroptera: Chrysopidae: Chyso- given, but the distributions of most species extend perla). Amer. Ent., 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, Ithonefusca, 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 Llfe, 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 CHIH, CHIS 5. C. subiratus (Walker, 1853) Mexico (no further locaiity) 43. C. comanche (Banks, 1938) BCS, CHIH, CHIS, COAH, DF, 6. Ululodes arizonensis Banks, 1907 BCS, CHIS, CHIH, 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 (NavBs, 1914) BCS, CHIS, COAH, DF, JAL, OAX, SIN, SLP, SON, TAMl', VER MICH, MOR, NAY, NL, OAX, SLP, SON, VER 8. U.cajennensis (Fabricius, 1787) BCS, CHIS, GRO, SIN, TAB, 45. C. externa (Hagen, 1861) BCS, CHIH, CHIS, COL (includ- TAMP, VER, YUC ing I. Revillagigedo), DF, DGO, GRO, JAL, MICH, MOR, NAY, 9. U.macleayanus (GuiIding, 1825) CHIH, CHIS, COL, GRO, HGO, OAX, PUE, SIN, SLP, TAMPS, WC 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, CHIH, 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. tuberculatus (Banks, 1901) QR 50. Chysopodes (Neosuarius) collaris (Schneider, 1851) BCS, CHIS, COAH, COL (I. Revillagigedo), GRO, NAY, OAX, SIN, Haplogleniinae SON, TAMP, VER, WC 14. Amoea immaculata (Olivier, 1790) CHIS, DGO, GRO, JAL, 51. Dichochysa perjecta (Banks, 1895) [n. comb.] BCS, SIN NAY, SIN, VER 52. D. sierra (Banks, 1924) [n. comb.] BC 15. A. iniqua (Walker, 1853) CHIS 53. D. triangularis (Adams, 1978) [n. comb.] HGO, JAL, 16. A. vacua (Gerstaecker, 1894) CHIS, WC MICH, PUE 17. Ascalobyas albistigma (Walker, 1853) CHIS, TAB 54. Eremochysa (Chrysopiella) sabulosa (Banks, 1897) CHIH, 18. A. rnicrocerus (Rambur, 1842) DGO, NAY, SLP, TAB, VER HGO, SLP, SON 19. Ascaloptynx appendiculata (Fabricius, 1793) CHIS, MOR, 55. E. (E.)altilis Banks, 1950 BC NAY, SIN, SON, TAMP 56. E. (E.) digueti NavBs, 1911 BC 20. Haploglenius luteus (Walker, 1853) NAY, VER 57. E. (E.)fraterna (Banks, 1897) BC 21. Neohaplogleniusf7avicornis(McLachlan, 1871) CHIS, HGO, 58. E. (E.) hageni Banks, 1903 BC JAL, QR, SIN, VER, WC 59. E. (E.)punctineruk (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 Adam, 1969 MICH 23. Lomamyia sp. TAMPS 64. M. colhuaca Banks, 1949 DF, HGO, MEX, MOR, QR, QRO 65. M. dolicharthra (NavBs, 1914) CHIS, HGO, MICH Chrysopidae 66. M. emuncta (Fitch, 1855) CHIH, TLAX Chrysopinae: BeIonopterygini 67. M.furcata (Banks, 1911) CHIH 24. Abachysa sp. JAL 68. M. hageni Banks, 1949 DF, MEX 25. Nacarina cordillera (Banks, 1910) [n. comb.] 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. Ceraeochysa cincta (Schneider, 1851) CHIS, COL, NAY, SIN 72. M. mhoa (Banks, 1949) DF, MEX 27. C. claveri (Navas, 1911) VER 73. M. pinalena (Banks, 1950) CHIH, MICH 28. C. cubana (Hagen, 1861) SON, TAME', VER 74. M. prpm' Tauber, 1%9 DGO, HGO, MFX, MICH, PUE,TLAX 29. C. eflw (Navhs, 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. meres (Banks, 1920) OAX, WC 77. M. tezcucana (Banks, 1949) DF, HGO, MOR 32. C.gradata (Navb, 1913) CHIS 78. Nineta nanina (Banks, 1911) MICH 33. C. indicata (Navb, 1914) VER 79. Plesioch ysu brasiliensis (Schneider, 1851) CHIS,JAL, NAY, SON, VER 34. C. lateralis (Gukrin-Mkneville, 1844) [n. comb.] CHIS, VER 80. P. elongata (NavL, 1913) CHIS, MOR, PUE, VER 35. C. placita (Banks, 1908) MICH 81. Yumachyw apache (Banks, 1938) BCS, CHIH, JAL, PUE,SON 36. C. sanchezi (NavBs, 1924) BCS, OAX, VER 82. Y.incerta (Banks,1895) [n.comb.] BC, MOR, OAX, TM,ZAC 37. C. smithi (Navb, 1914) BCS, COAH, VER 38. C. valida (Banks, 1895) BCS, COL, JAL, NAY, OAX, SLP, Chrysopinae: Leucochrysini TAB, TAMP, VER, WC 83. Gonzagn torquatus NavBs, 1913 CHIS, COL, VER 39. Chysopa coloradensis Banks, 1895 BCS 84. Leucochysa (L.) arizonica (Banks, 1906) JAL, MEX, MICH 40. C. nigricornis Burmeister, 1838 DGO 85. L. (L.) negata (NavBs, 1913) DF 1 41. C. oculata Say, 1839 CHIH, DF, DGO, JAL 86. L. (L.) serrula Adam, 1979 MICH, MOR, SLP NEUROPTERA (NEUROPTERIDA) 1 579

87. L. (Nodita)americana Banks, 1897 BCS, OAX 130. C. californica Meinander, 1972 DF 88. L. (N.) antennalis (NavBs, 1932) Mexico (no further locality) 131. Semidalis angusta (Banks, 1906), BCS, DGO, GTO, MEX, 89. L. (N.)antennata Banks, 1905 BCS, PUE, VER MICH, MOR, NAY (I. Marias), OAX, QRO, SIN, SON 90. L. (N.) askanes (Banks, 1946) OAX 132. S. boliviensis (Enderlein, 1905) CHIS, VER 91. L. (N.) callota Banks, 1915 ZAC 133. S. byersi Meinander, 1972 DGO, GTO, HGO, MICH, MOR, 92. L. (N.) explorata (Hagen, 1861) Mexico ("Cordova") NAY, PUE, VER 93. L. (N.) mexicana Banks, 1900 VER 134. S. deserta Meinander, 1974 BC 94. L. (N.)navasi (Kimmins, 1940) Mexico (no further locality) 135. S.faulkneri Meinander, 1990 SIN 95. L. (N.) nigrznervis (Banks, 1939) BC 136. S.flinti Meinander, 1972 BC, BCS 96. L. (N.)pavida (Hagen, 1861) Mexico ("Cordova") 137. S.fiommeri Meinander, 1974 SON 97. L. (N.)salleana (NavBs, 1911) JAL 138. S. manausensis Meinander, 1980 VER 98. L. (N.)sulcata (Navb, 1921) JAL 139. S. mexicana Meinander, 1972 BCS, DGO, HGO, MICH, NL, SIN, SLP, SON Chrysopinae: tribelgenus Incertae Sedis 140. S. soleri Monserrat, 1985 VER 99. "Chysopa" aztecana Banks, 1903Mexico (no further locality) 100. "C." brarihirta Banks, 1946 Mexico (no further locality) Dilaridae 101. "C." fascialis Banks, 1905 Mexico (no further locality) Nallachiinae 102. "C." leptana Banks, 1914 Mexico (no further locality) 141. Nallachius pulchellus (Banks, 1938) NAY 103. "C." senior (Navhs, 1928) Mexico (no further locality) Hemerobiidae Nothochrysinae Hemerobiinae 104. Pimachrysa albicostales Adarns, 1967 BC 142. Hemerobiella sinuata Kimmins, 1940 CHIS 143. Hemerobius alpestris Banks, 1908 CHIS, DF, DGO, MEX, Coniopterygidae VER, JAL, MICH, OAX, PUE, TAMP Aleuropteryginae: Aleuropterygini 144. H. bolivari Banks, 1910 CHIS 105. Aleuropteyx longipennis Meinander, 1974 BC 145. H. discretus NavBs, 1917 AGS, BC, CHIS, COL, DF, DGO, 106. A. maculipennis Meinander, 1972 BCS GRO, GTO, HGO, 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 HGO, MICH, PUE, SLP 109. A. unicolor Meinander, 1972 BCS 147. H. elongatus Monserrat, 1990 MEX 148. H. gaitoi Monserrat, 1996 VER

Aleuropteryginae: Fontenelleini i 149. H. hernandezi Monserrat, 1996 CHIS, OAX, VER 110. Bidesmida morrisoni Johnson, 1977 BC 150. H. hirsuticornis Monserrat & Deretsky, 1999 TAMPS 111. Neoconis inexpectata Meinander, 1972 JAL 151. H. humulinus Linnaeus, 1758 VER 112. N. marginata Meinander, 1972 BC, BCS, JAL, SON 152. H. jucundus NavBs, 1928 CHIS, DF, GRO, HGO, JAL, 113. N. unam Monserrat, 1985 VER MEX, MICH, OAX, PUE, TAMPS, VER 153. H. kokaneeanus Currie, 1904 BC Coniopteryginae: Coniopterygini 154. H. martineme Monserrat, 1996 CHIS, GRO, MICH, OAX, VER 114. Coniopteyx (C.) callangana Enderlein, 1906 YUC 155. H. pac$cus Banks, 1897 BC, DF, MEX, OAX, SON, VER 115. C. (C.) delta Johnson, 1981 VER 156. H. stigma Stephens, 1836 BCS, h4EX 116. C. (C.)fitchi Banks, 1895 GTO, MICH, PUE 157. H. tibialisr~avb,1917 [nomen dubium] Mexico (no fur- 117. C. (C.) mexicana Meinander, 1974 SIN ther locality) 118. C. (C.) minuta Meinander, 1972 BCS, SIN 158. H. tolimensis Banks, 1910 MEX, VER 119. C. (C.) palpalzs Meinander, 1972 BCS, DGO, GTO, HGO, 159. H. withycombei (Kimmins, 1928) TAB, VER MICH, OAX, QRO, VER 160. longipennis (Banks, 1920) CHIH, DGO, MOR 120. C. (C.)simplicior Meinander, 1972 BCS, COL (Socorro Island) 121. C. (C.) westwoodi (Fitch, 1855) QRO Megalominae 122. C. (Scotoconiopteryx)flintiMeinander, 1975 SLP 161. Megalomusfidelis (Banks, 1897) NL 123. C. (S.) isthmicola Meinander, 1972 SLP 162. M. minor Banks, 1905 CHIS, COL, JAL, MOR, NAY, SLP, 124. C. (Xeroconiopteryx) diversicornis Meinander, 1972 BCS, TAB, TAMPS, VER DGO, NL, VER 163. M. moestus Banks, 1895 COL, JAL, MOR, SON, TAMPS, 125. C. (X.) meinanden' Johnson, 1981 SIN VER, ZAC 126. C. (X.) texana Meinander, 1972 BC 164. M. pictus Hagen, 1861 Mexico (no further locality) 127. C. obscura NavBs, 1934 MOR 128. Parasemidalisfuscipennis (Reuter, 1894) BC, DGO Microminae 165. Micromusficscatus (Nakahara, 1965) CHIS, MICH, QRO Coniopteryginae: Conwentziini 166. M. posticus (Walker, 1853) CHIS, DF, GRO, MICH, MOR, 129. Conwentzia barretti (Banks, 1898) BC, CHIS, DF, DGO, NAY, NL, SLP, TAMPS, VER GTO, HGO, MEX, MICH, MOR, PUE, VER 167. M. subanticus (Walker, 1853) BCN, BCS, CHIH, COAH,