Neuroptera (Neuropterida)
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NEUROPTERA (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), Chrysopidae (81), Hemerobiidae-(44), lacewings attracted to the lights. Coniopterygidae (36), Mantispidae (22),Ascalaphidae Neuroptera have two distinctive characteristics (21), Sisyridae (4), Ithonidae (2), Berothidae (2), Dila- that make them fascinating creatures. First, they ridae (1) y Polystoechotidae (1).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, Nemopteridae, and Ascalaphidae can be quite colorful, and at a distance can be mis- Neuroptera, or lacewings, antlions, and their al- taken with large butterflies. lies, is one of the smaller orders of insects, with approximately 6000 described species, of which only about 4000 species are estimated to be valid SYSTEMATICS AND DIVERSITY taxa (Aspock et al., 1980). They are found in all temperate and tropical parts of the world, but Documenting biodiversity for insects is a difficult rarely constitute a major component of the insect task. Insects are often small and elusive, and pass fauna. Neuropterans probably reach their great- many months of the year in immature and resting est diversity in tropical forest communities, but stages. Even when abundant, they are often cryp- seldom are very abundant there, and often are very tically colored to avoid being eaten by visual predators, like birds and lizards. Thus, few com- parative statements have been made about the diversity of insect groups for any- region.- For some of the more cons~cuousgroups, such as butter- Department of Entomology, Texas A & M University, College Station, Texas 77845-2475, USA. e-mail: [email protected] flies (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 order 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 Nymphidae are restricted to specialists in the country. Also, we are not aware of Australia. Species of Nevrorthidae, Psychopsidae, graduate students being trained in systematics of Rapismatidae and Rhachiberothidae are found only Neuroptera within or out of the country that in the in the Old World. Representatives of Nemopteridae near future might engage in the study of the Mexi- and Osmylidae are found in northern South can fauna. There has been, however, considerable America, but have never been collected in Central work on the Mexican fauna, mostly by non-Mexi- America or Mexico. All eleven other families can can specialists describing species as part of revision- be found in some part of the country, including all ary work (e.g., Victor J. Monserrat from Spain on major diverse and speciose families of the order. Hemerobiidae, etc.).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 Arthropods). One of us (ACR) has adult Mexican Neuropterans to genus (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