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The neuropteroid orders of Central America (Neuroptera and Megaloptera)

CHARLES S. HENRY, NORMAN D. PENNY, AND PHILLIP A. ADAMS

INTRODUCTION strap-like, coriaceous front wings, not unlike the elytra of beetles. The order is among the most biologically and mor­ Imagine that you are a student in a course on insect identi­ phologically diverse in all Insecta. fication, and you are presented by your instructor with an /~ the Neuroptera is no success story, compared with other 'unknown' to classify, merely to order. It has extremely long, knobbed antennae, like a butterfly; perhaps it is a lepidopteran. Yet its wings are membranous, narrow, and strong, and it has ( \ !}l':'I /. legs that are spiny and obviously modified for seizing small I insect prey items during flight; these are characteristics of a , dragonfly, order Odonata. But yellowish-white markings on the rather fat abdomen, together with a narrow waist, give the .II/~·~~ specimen a wasp-like appearance: it could be in the Hymen­ , optera, or maybe a good wasp-mimicking fly (order Diptera). Examine the compound eyes: each is extremely large, as one expects of an odonate, but also divided by a deep sulcus into an upper and lower half, as in certain male mayflies-is the insect an aberrant ephemeropteran? Finally, you decide that your instructor has glued together the parts of many insects, to fool you. In fact, however, you have just been introduced to the peculiar, fascinating order of insects known as Neuroptera, and you are looking at a member of one of its most curious families, the Ascalaphidae or owlflies (Fig. 28.1). One learns to expect the unexpected in the Neuroptera. bisected "" Within the order, there are insects that trap their prey with eye funnel traps built in the sand (Myrmeleontidae), chase small insects across the desert floor on stilt-like legs (Nemopteridae), browse under the water on freshwater sponges (Sisyridae), live among ants and feast on the colony's brood (Chrysopidae: Italochrysa), and even kill their termite prey by blasting them with a lethal gas from the rear end (Berothidae). Some neur­ opterans (Ascalaphidae) in Europe have wide, brightly coloured wings, and for this reason were once placed in the butterfly genus Papilio; others possess extremely long, thread-like hind wings that trail out behind them in flight (Nemopteridae). The members of one widespread family faithfully duplicate nearly every major feature of the praying mantises, including their powerful raptorial front legs and elongated prothoraces (Fig. 28.5), yet as youngsters, these 'mantispids' (Mantispidae) para­ c.s.~ sitize spiders and social insects and look like maggots. And Fig. 28.1 Ululodes mexicana, nonnal resting position (Ascalaphidae) at least one aberrant dusky-wing (Coniopterygidae) possesses [Arizona, original]. THE NEUROPTEROID ORDERS 433 insect orders. At a mere SOOO-odd species, the order is min­ RsMA MP1+2 uscule compared with the Hemiptera (100 000 species), Diptera (85 000), Hymenoptera (c. 200 000), Lepidoptera (120 000), or Coleoptera (400000). Nor is it particularly well-defined: whereas each of the large orders above is homogeneous and united by clear synapomorphies (unique, specialized features shared among its members), neuropteran families are united principally by primitive (plesiomorphic) features, often shared with other ancient orders. 'Neuroptera' is also used differently by different authors. To some, it includes the Megaloptera base (dobsonflies and alderflies) and Raphidioptera (snakeflies), while to others, it is restricted to the so-called planipennian families, characterized by larvae that possess piercing, sucking mouthparts of a particular sort. Here, we choose to restrict Neuroptera, but we acknowledge the possible monophyly of Neuroptera, Megaloptera, and Raphidioptera by including all three orders in our discussion; to avoid confusion, we will refer to the three orders together as 'neuropteroids'. Oddly, the closest living relatives of the neuropteroids are probably Coleoptera, the most speciose of insect taxa. Most of the smaller, Fig. 28.2 Platyneuramus sarar (Hagen) (Corydalidae) (redrawn from older orders of holometabolous insects are also related evol­ Glorioso and Flint 1984). CuA, cubitus anterior; CuP, cubitus posterior; utionarily to species-rich groups: the Trichoptera to the Lepi­ MA, media anterior; MP1 + 2, fused anterior and posterior branches of doptera, the Mecoptera to the Diptera, and the Symphyta to the media posterior; Rs, radial sector; lA, first anal vein. aculeate Hymenoptera. In each of these cases, as in that of the neuropteroids, it seems that some evolutionary breakthrough, sopidae and Ascalaphidae, which have experienced more recent originating within a derivative taxon of the parental, primitive evolutionary success, attain highest diversity in the humid order, precipitated rapid speciation and adaptive radiation of tropics. those inheriting the valued feature. Eventually, the vigorous Central America and Panama possess a diverse neuropteroid child, favoured by its new technology, far outgrew its parent. fauna, perhaps because the region serves as a crossroads Relatively species-poor orders like the three comprising the between the distinctive adaptive radiations of North and South neuropteroid complex, then, are relict collections of diverse, America. Consequently, the larger, more cosmopolitan taxa are generally primitive taxa that happen to have escaped extinc­ disproportionately represented here: each of the major families tion; and because extinctions have removed many of the inter­ Mantispidae, Hemerobiidae, Chrysopidae, Ascalaphidae, Myr­ mediate, transitional forms that once existed in the complex, meleontidae, and Coniopterygidae is a major component of the neuropteroids present major problems to entomologists inter­ Central American entomofauna, and consists largely of genera ested in classification and phylogeny. with their centres of diversity in South America. Unfortunately, As relict, ancient groups, the neuropteroid orders and their some of the most ancient and bizarre families, mentioned in the families show very interesting, often disjunct geographical dis­ preceding paragraph, are absent from the New World tropics. tributions, which reflect the past excursions of the continents. Nevertheless, the neuropteroids ofthis region hold great interest A good number of families reach maximal diversity in bio­ for amateur and professional entomologists alike. geographically isolated regions like Australia or South Africa, and several taxa have typical Gondwanaland (southern hemi­ sphere) distributions. For example, Nymphidae is confined to GENERAL BIOLOGY AND the Australian region, while Psychopsidae (silky lacewings) shows high generic endemism in South Africa, Central Asia, EVOLUTION and Australia. In addition, several other families, including the two neuropteran sister-taxa Berothidae and Mantispidae, show Neuropteroids are endopterygote or holometabolous insects, striking phylogenetic ties between the South American and meaning that the life cycle is characterized by internally devel­ South African representatives (Aspock and Asp6ck 1988; oping wing pads and complete metamorphosis. Complete meta­ Tjeder 1959). Other families with peculiar, relict distributions morphosis, in which the adult insect differs radically in include Rapismatidae, which is scattered through parts of appearance and habits from its earlier immature, larval stages, central Asia, and Polystoechotidae (giant lacewings), restricted confers great evolutionary advantages, principally through to the New World from parts of North America to Chile. In the exploitation of different ecological niches by different Iife­ general, the oldest neuropteroid groups are more diverse in stages of the same organism. However, the process of metamor­ temperate, desert, or montane areas, while taxa like Chry- phosis is very difficult and risky, and the more generalized 434 28. HENR Y, PENNY, AND ADAMS

throughout larval life, because the hindgut and rectum are closed off from the midgut, and are eliminated as a 'meconial pellet' by the new adult while the wings and body sclerites are hardening. The larvae of the vast majority ofneuropteroids, and all of those in the New World tropics, are voracious predators of other invertebrates. The Ithonidae, whose larvae ingest flUids from decaying plant litter, is the only family for which a non­ predatory immature has been documented (New 1986a)~ Although the larvae of most neuropteroids are terrestrial, those of a few families are semi- or totally aquatic. All Mega­ loptera, for example, are underwater predators during their larval stages, breathing through specialized abdominal gills. Fig.l8.3 Glenurus luniger Gerst. (Myrmeleontidae) [Arizona, original] CuA, Sisyridae, too, have subaquatic larvae, which feed on fresh­ cubitus anterior; CuP, cubitus posterior; CuP + I A, fused CuP and first anal water sponges. A few Osmylidae and Neurorthidae are also veins; MP, media posterior. closely associated with fresh water, but these two families are not found in Central America. At the other extreme are neur­ opteroids like Myrmeleontidae, Ascalaphidae, and Nemop­ neuropteroids like Megaloptera and Raphidioptera seem not to teridae, whose larvae can survive in extremely arid areas. have perfected it; pupation and transformation consume much Most neuropteroids pass through three larval stadia prior to time and energy. Even the more specialized Neuroptera are pupation, although a larger number of larval instars char­ characterized by a primitive, biting pupa that must gnaw its acterizes the Ithonidae (Gallard 1932) and most members of way out of the cocoon and walk about before moulting. This the orders Megaloptera and Raphidioptera. Silk production, for type of pupa contrasts sharply with the sleek, passive pupae the cocoon, is confined to the order Neuroptera. Silk is spun of most other endopterygotes, like butterflies, social wasps, or from the anus and is manufactured within two (of eight) spe­ higher flies. cialized malpighian tubules, which empty into the rectum. The The families of the dominant neuropteroid order, Neuroptera, cocoon is usually spherical and double-walled, and may include fall quite naturally into two major phylogenetic groups, the debriS, sand, or parts of leaves in its construction. The typical Hemerobiiformia and the Myrmeleontiformia (MacLeod 1964). neuropteroid spends the majority of its time within the cocoon These distinct evolutionary lineages are defined principally by as a prepupa (dormant late third instar larva), moulting to the larval feeding specializations (MacLeod 1970), because adults pupal stage a few days or weeks before eclosing. Many tem­ within the order present a bewildering array of plesiomorphies, perate zone species overwinter within the cocoon. However, specializations, parallelisms, and convergences which obfuscate several well-known taxa endure the winter perfectly well as evolutionary trends. Hemerobiiform Neuroptera include the eggs (some Sisyridae; Pupedis 1980), larvae (some l'vlallada spp. primitive, more generalized feeders and consequently show of the Chrysopidae), or even adults (Chrysoper1a, of Chrysopidae; larval head and jaw morphology of a less extreme type than do Semeria 1977). Changes in photoperiod most commonly Myrmeleontiformia. The maxillary blade in larvae of the former control the life cycle in temperate zone neuropteroids (Tauber matches or exceeds the mandible in size and strength, the jaws and Tauber 1986), but in tropical areas like Central America, are toothless and only slightly curved, and the head capsule predictable changes in rainfall, humidity, or temperature are has only average sclerotization. Typically, only soft-bodied prey more important. are attacked. In contrast, myrmeleontiform larvae possess As a rule, neuropteroid adults are delicate insects with rela­ much more powerful, Sickle-shaped, often toothed jaws and tively large, attractive, intricately veined Wings. Flight is slow a heavily reinforced box-like head. Such features permit the and rather clumsy, except in specialized aerial predators like myrmeleontiforms to capture and subdue much larger arthro­ the Ascalaphidae. Most species fly willingly at any temperature, pods than can their hemerobiiform relatives. As typified by the and are thus dispersed over large distances by winds (Gressitt et pit-making antlions, myrmeleontiform larvae frequently 'sit (11961), e\'en to isolated oceanic islands (Zimmerman 1957). and wait' for their food, rather than choosing prey of suitable Adult feeding habits vary, but the vast majority ofneuropteroids size and strength, so their more powerfully built feeding struc­ are predators or scavengers of soft-bodied arthropods or, alter­ tures can be viewed as adaptations to unpredictably large prey. natively, are nectar and honeydew feeders. Some, including The larvae of all neuropteroids except Megaloptera and most members of the Megaloptera, consume little if any real Raphidioptera possess suctorial mouthparts, in which the elon­ food during their short adult life-spans. All possess generalized, gated maxilla fits into a groove on the under-surface of the mandibulate mouthparts, occasionally enlarged in males (e.g. mandible to form a channel. Poisons (Henry 1977) and diges­ Megaloptera) for use in intraspecific reproductive contests or tive enzymes can be injected into prey through this channel; courtship. Overall, adult external morphology is conservative later, the extra-orally digested tissues of the prey are imbibed and primitive, reflecting the ancient ancestry of the superorder. through the same passageway. Solid wastes are retained Wing venation is particularly archaic, making it difficult to ~;. 'y,­ T~

~, ····~··· ,i.····." . ;:l ~ • THE NEUROPTEROID ORDERS 435

axillary

Fig. 28.4 Neohaploglenius sp. (Ascalaphidae) [redrawn from Penny 1981]. CuA, cubitus anterior; CuP+ lA, fused cubitus posterior and first anal vein; MP, media posterior; MPl, anterior branch of MP; MP2, posterior branch of MP; MP2 + CuA, fused MP2 and CuA; Rs + MA, fused radial sector and media anterior; 2a, second anal vein.

distinguish neuropteroids from primitive Trichoptera, Plecop­ grapes or other susceptible crops, has even met with modest tera, and Mecoptera in the fossil record. They are as con­ commercial success (Hagen et al. 1970). servative internally as they are externally, displaying little Courtship and mating are not well studied within the neur­ fusion of the segmental components of the central nervous opteroids. It has been assumed, from the superficial and anec­ system, heart, or ventilatory system. However, as mentioned dotal observations of several early naturalists, that courtship is earlier, each family is characterized by a number of unique, simple and short in most families (Balduf 1939; MacLeod 1962, often bizarre specializations, making it quite easy for the know­ unpublished). Commonly seen elements are mouthpart contact ledgeable amateur to identify the families and discriminate between the partners and some sort of abdominal jerking or each from every other. drumming. We now know that courtship in some green lace-

Fig. 28.5 Plega signata (Hagen) (Man­ tispidae) [California, original].

In general, populations of neuropteroids are small and of low wings can be extremely complex, involVing the reciprocal density. Consequently, the impact of individual species on the exchange between the sexes of vibrational 'songs' produced environment is seldom noticeable, although impressively large by controlled abdominal movement (Henry 1986). As more swarms of polystoechotids once occurred in parts of boreal research focuses on courtship and mating, patterns of com­ North America (Welsh 1914). Because most neuropteroids are parable complexity will undoubtedly be found in other neur­ predatory, they are never important crop pests; in fact, many opteroid taxa. are classified as beneficial insects. Chrysopidae, in particular, Eggs ofneuropteroids are typically smooth-shelled and ovoid, have been recognized as important agents in the biological without much ornamentation, and conservatively pigmented. control of aphids, scale insects, and other damaging plantlice They are laid individually, but often closely packed. For some (New 1975). Mass culture of green lacewings, for release on reason, eggs on thin, silken stalks have evolved independently 436 28. HENRY, PENNY, AND ADAMS

tive' (unfertilized) eggs have defensive and trophic functions in New World Ascalaphidae (Henry 1972). Fertilization in most t neuropteroid species occurs through an obvious micropyle, and hatching is often facilitated by a line of weakness that encircles :1 one end of the egg at the conclusion of embryonic development. ~~I Hatchlings may be gregarious for a short while, but invariably j disperse before they start feeding.

IDENTIFICATION -j .•, As in many holometabolous insects, the evolution of neur­ £ opteroid immatures has not been closely coupled with that of ;! j- adults. It is as though each species is represented by two sep­ ~ arate organisms, each with its own independent evolutionary history. The aggressive, ravenous larva gives no clue to the appearance and habits of the fragile adult; the larva's job is to gather resources, while the adult's is to court, mate, and Fig. 28.6 Notiobiella mexicana Banks (Hemerobiidae) [redrawn from Mon­ disperse. To identify some neuropteroid taxa with confidence, serrat 1983]. branch, first forking of first radial sector; R, radius; first Rs, it is best to examine both larvae and adults, which in turn first radial sector; second Rs, second radial sector. requires one to have immatures positively associated with par­ ticular adults by rearing. Of course, this is seldom possible; i many species are known only from adults, and occasionally in a number of distantly related families: such a condition from adults of only one sex. In the following keys, we have used characterizes Chrysopidae, Berothidae, many Mantispidae, and both adult and larval characters to aid in the identification of ;~ several Nymphidae (Riek 1970). Presumably, the stalk serves problematical orders, families, or genera, but the emphasis to protect the egg from predators, although this widely accepted throughout is on adult morphology. ~ 'i hypothesis has not received rigorous testing. Specialized 'abor­

:'1J Key to the families of Central American Neuropteroidea

{. 1 Hindwings broader at base than forewings, and with an enlarged anal area which in ~ Corydalidae is folded fanwise at rest (Fig. 28.2); larvae aquatic with respiratory gills on , first eight abdominal segments Order Megaloptera 2

- Fore- and hindwings similar in shape, the hindwings without an enlarged anal area that is folded fanwise at rest; larvae aquatic or terrestrial with hollow suctorial mouthparts; larvae, when aquatic, small, living on freshwater sponges, with gills only at base of thoracic legs Order Neuroptera 3 ~" .1f ~ 2 Very large insects; bearing three ocelli; fourth tarsal segment cylindrical; larvae bearing paired hooked anal prolegs, but no terminal filament Family CorydaIidae

- Medium-sized insects with dark smokey wings; ocelli absent; fourth tarsal segment bilobed; larvae lacking anal prolegs, but bearing terminal filament Family SiaIidae I~

3 Size 3 mm or less; covered with a whitish powder Family Coniopterygidae

- Size 4 mm or greater; not covered with a whitish powder 4

4 Antennae clubbed or knobbed; insects with the abdomen long and slender, and resembling damselflies or dragonflies in appearance; larvae with robust, curved jaws 5 THE NEUROPTEROID ORDERS 437

- Antennae filiform, moniliform, or pectinate, not clubbed or knobbed; usually not particularly resembling damselflies or dragonflies in appearance; larvae with thin, straight or slightly curved jaws 6

5 An elongate cell behind point of fusion of Sc and R1 (Fig. 28.3); antennae apically swollen, but not knobbed, and no longer than head and thorax combined Family Myrmeleontidae - No elongate cell behind point of fusion ofSc and R1 (Figs 28.1, 28.21); antennae apically knobbed, and usually as long as entire body, but can be as short as head and thorax combined Family Ascalaphidae

6 Prothorax elongate; forelegs greatly enlarged, raptorial (Fig. 28.5); second and third instar larva sedentary and maggot-like Family Mantispidae - Prothorax not elongate; forelegs not greatly enlarged or raptorial; larvae not maggot-like 7

7 ForeWings with two or more branches of Rs arising from the apparently fused stems of R1 and Rs (Figs 28.6, 28.17) Family Hemerobiidae - Forewings with all branches of Rs arising from a single sector separate from R1 8

8 Antennae moniliform in both sexes; ovipositor not projecting; cross-veins few 9 - Antennae of male pectinate (Fig. 28.7); female with an exserted ovipositor as long as body Family Dilaridae

9 Humeral cross-vein recurved and branched; first apparent r-m cross-vein (base of MA) in hindwing longitudinal and sigmoid (Fig. 28.8); large species with a wing spread of 40­ 75 mm Family Polystoechotidae

Fig. 28.7 Nallachius limai Adams (Dilaridae) [from Adams 1970]. 438 28. HENRY, PENNY, AND ADAMS

- Humeral cross-vein not recurved; first apparent r-m cross-vein (base of MA) in hindwing variable; smaller species 10

10 Vertex convex; wing· venation relatively simple, radial sector of forewings with few cross­ veins (Fig. 28.9), Sc and Rl coalesced near tip of wing, costal cross-veins not forked, r-m cross-vein of hindwings in the axis of wing; size small, 6-8 mm in length; larvae aquatic, feeding on freshwater sponges Family Sisyridae - Vertex flattened; single radial sector in forewings with many cross-veins; Sc and Rl may or may not be coalesced near tip of wing, costal cross-veins mayor may not be forked; hindwings with r-m cross-vein oblique, transverse, or absent (Fig. 28.10); size larger 11 I 11 Wings rounded, not falcate; costal cross-veins not forked, Sc and Rl free at tip, Rs swinging away from Rl, cell Rl broad and containing many cross-veins (Fig. 28.10) Family Chrysopidae ~ - Apical portion of the hind margin of forewings straight or concave (falcate), leaving the :f apex more or less acute; wings and body hairy, especially the hind margin of wings; wings ~~ may bear scales along base of longitudinal veins; costal cross-veins forked, cell Rl narrow ~; :}. and almost devoid of cross-veins (Fig. 28.11) Family Berothidae ~f i Key to the genera of Central American Corydalidae ! 1 Lateral cephalic flange expanded into flattened plate behind eyes, especially prominent in males (Fig. 28.12); mandibles of males not enlarged and elongated Platyneuromus t-lit .~ - Lateral angle of head behind eyes forming flattened tooth, but not expanded into plate; males mayor may not have elongate mandibles 2 1 Bright lemon yellow colour with black margining of cross-veins in life, fading to pale straw t yellow after death; male mandibles never elongate Chloronia I - Reddish brown to smoky grey, often with forewings clouded with variable darkening and i small white points; male mandibles can be moderately to extremely elongate Corydalus

I}: ~ ~. '" ~ .-z{ ~:: '~~

Fig. 28.8 Polyswechotes punctatus (Fabricius) (Polystoechotidae) [original]. THE NEUROPTEROID ORDERS 439 Key to the genera of Central American Coniopterygidae (modified from Meinander 1972)

1 Two radio-medial cross-veins in middle of forewing; in hindwing Rs branching off from R very near base of wing (Fig. 28.13); plicaturae (Fig. 28.14) present on abdomen; antennae of larvae about as long as labial palpi; jaws projecting from beneath labrum Neoconis - One radio-medial cross-vein in middle of forewing; in hindwing Rs not branching off from R very near base of wing; no plicaturae present on abdomen; antennae of larvae about twice as long as labial palpi; jaws not projecting from beneath labrum 2

2 M of hindwing unforked Coniopteryx - M of hindwing forked Semidalis

Key to the genera of Central American Sisyridae (from Pamn and Gurney 1956)

1 A series of outer gradate cross-veins present in fore- and hindwings; wings predominantly pale with extensive dark markings Climacia - No series of outer gradate cross-veins present in either wing; wings entirely smoky brown Sisyra

Key to the genera of Central American Mantispidae (modified from Penny 1982)

1 Anterior tarsi each with only one claw and without arolia (Fig. 28.15) Subfamily Mantispinae 4 Anterior tarsi each with two claws and an arolium 2

2 Anterior first tarsal segment with a dentiform process (Fig. 28.16); adult female bearing ovipositor; male penisfilum coiled; subcosta entering costal margin in area of pterostigma; fore tarsal claws not forked Subfamily Symphrasinae 3 - Anterior first tarsal segment without dentiform process; adult female not bearing ovipositor; male penisfilum not coiled; subcosta of forewing entering costal margin far anterior to pterostigma; fore tarsal claws bifid Subfamily Calomantispinae Nolima

3 Sub-basal sprne of fore femur present Plega - Sub-basal spine of fore femur absent Trichoscelia

4 Cubitus of hindwing nearly straight and never coming close to first anal vein; vespid wasp mimic with dark coloration along costal margin of Wings Climaciella - Cubitus of hindwing bending sharply towards first anal vein, then bending sharply away again; wings transparent or perhaps with some coloration in subcostal area, but never }' covering all costal area of wings 5 440 28. HENRY, PENNY, AND ADAMS

Rs+MA

Fig. 28.9 Sisyra sp. (Sisyridae) [redrawn from Partin and Gurney 1956]. CuA, cubitus anterior; CuP, cubitus posterior; MA, media anterior; MP, media posterior; R, radius; Rs, radial sector; Rs + MA, fused Rs and MA; lA, first anal vein. MA Rs+MA

5 Crescent or semicircular marks on prozonal region of pronotum Entanoneura Prozonal region of pronotum with longitudinal stripes or all transverse marks, but without semicircular yellow markings Mantispa

Key to the genera of Central American Hemerobiidae

1 Forewing with two radial sectors 2 l ,.~; k~·filoi Forewing with three or more radial sectors 3 r ' ."., / '.rl / 1/ .• 1 , ) 2 First radial sector branching well before origin of second Rs Sympheromima [) First radial sector branching at level of origin of second Rs (Fig. 28.6) Notiobiella

3 Forewing with narrow costal area at base and no recurrent costal vein Nusalala Forewing with relatively wider costal area at base, and recurrent costal vein 4

4 A conspicuous cross-vein present in the forewing between MA and R5 (or R4 + 5) near the latter's origin Wesmaelius No such cross-vein present 5

5 Forewing with three radial sectors (Fig. 28.17); male ectoprocts caudally directed; male parameres absent Hemerobius Forewing with five or more radial sectors; male ectoprocts ventrally directed; male parameres present Megalomus THE NEUROPTEROID ORDERS 441

Fig. 28.10 Berchmansus adumbratus Navas (Chrysopidae) [Brazil, original]. CuP, cubitus posterior; imc, intramedian cell; MP1, anterior branch of media posterior; psc, pseudocubitus; psm, pseudomedia; psx, presectoral cross-vein.

Fig. 28.11 Lomamyia squamosa Carpenter (Berothidae) [Belize, female].

Key to the genera of Central American Chrysopidae

Basal antennal segments wider than long; rather stout, wide-bodied species; tignum absent, gonapsis (Fig. 28.18) may be present as two separate projections or a two-homed plate, female often with praegenitale (Fig. 28.19) Tribe Be1onopterygini Nacarina - Basal antennal segments longer than wide; rather thin, slender-bodied species; gonapsis, if present, not a two-horned plate; female lacks praegenitale 2

2 No dark spot at base of stigma; antennae usually shorter than 1.3 times forewing length, tignum or gonapsis (Fig. 28.18) may be present Tribe Chrysopini 7 - A dark spot at base of stigma, antennae distinctly longer than 1.3 times forewing length; no tignum or gonapsis (Fig. 28.18) 3

3 In forewing, basal subcostal cross-vein and intramedian cell absent; distinct blisterlike bullar spots on foreWing (Fig. 28.20) Tribe Apochrysini 4 In forewing, basal subcostal cross-vein and intramedian cell present; bullar spots absent Tribe Leucochrysini 12

4 Costal area of forewing with cells undivided (isolated divided cells may be disregarded) 5 Costal area of forewing composed of two or more rows of cells, or densely reticulated Lainius

5 Forewing with three to seven elevated bullae, often arranged in an arc from midway along Psm to apex of wing Loyola - Forewing with not more than two elevated bullae Domenechus 442 28. HENRY, PENNY, AND ADAMS

;>:: :1:.-'···. ....i ·1 iii :11 , ~~. ,:fl ~; ":t, lateral ":" cephaliC~ flange I

. '

:t:·.····.··.··..;.{•.

". "~._ ...•.. ~,., 1-.:.­ '<', ~:~C;~J!~:;j, Fig. 28.13 Neoconis dentata Meinander (Coniopterygidae) (modified from Fig. 28.12 Platyneuromus soror (Hagen), head of male (Corydalidae) Meinander 1972]. CuA, cubitus anterior; CuP, cubitus posterior; M, media; [redrawn from Glorioso and Flint 1984]. R, radius; Rs, radial sector; Sc, subcosta.

6 Tignum absent (Fig. 28.18) 9 - Tignum present 7

7 Pseudopenis present (Fig. 28.18) 8 - Pseudopenis absent Chrysoperla , 'e'f. 8 Males with enlarged or modified antennal bases and/or with facial horns; otherwise with stridulatory apparatus on abdominal base; gonapsis usually present Meleoma - Males with normal heads and antennae; no stridulatory mechanism or gonapsis Chrysopa (Plesiochrysa) 'f

9 Two hornlike structures on gonarcus or arcessus; gonapsis elongate Ceraeochrysa - Hornlike structures absent from gonarcus-arcessus; gonapsis absent or present, but not elongate 10

10 Pseudopenis present (Fig. 28.18) Meleoma ~ - Pseudopenis absent 11 ":1'

:#'.g,. 11 Mandibles blunt-tipped; wings usually with narrow costal area; gonapsis never present "~, Chrysopodes (Neosuarius)

Mandibles with fang-like tip; wings usually with wide costal area; gonapsis rarely present ;;;': Chrysopodes(Chrysopodes)

:i\t,'lI':;i 12 A cross-vein before radial sector; usually but five psm cross-veins beyond the intramedian ~;' cell (Fig. 28.10) Berchmansus No cross-vein before radial sector; usually six or more psm cross-veins beyond intramedian rell 13 ·:I·~ .." f' TZEn

THE NEUROPTEROID ORDERS 443

13 Intramedian cell triangular, MP2 joining MP1 before end of cell Nodita - Intramedian cell quadrangular, MP2 not joining MP1 14

14 Pterostigma with a large black spot extending back to radial sector; costal stigmal area usually twice as broad as subcostal stigmal area Gonzaga - Pterostigmal mark much smaller; costal stigmal area rarely twice as broad as subcostal stigma area Leucochrysa

Key to the genera of Central American Ascalaphidae (modified from Penny 1981)

1 Eyes entire, not divided by transverse sulcus Subfamily Haplogleniinae 2 - Eyes divided by transverse sulcus into two parts, superior and inferior, equal or unequal Subfamily Ascalaphinae 5

2 Forewing with prominent axillary angle at base of anal margin (wing cells distal to axillary angle shorter than within angle) (Fig. 28.4) Neohaploglenius - Forewing without prominent axillary angle at base of anal margin (Wing cells distal to axillary angle equal to or longer than along basal curve) 3

3 Antennae longer than distance to second Rs fork of forewing 4 - Antennae shorter than distance to second Rs fork of forewing Ascalobyas

4 Hindwing with 2A present, long; dark coloration along costal margin of forewing Haploglenius - Hindwing with 2A absent or extremely short; transparent clear coloration along costal margin of forewing Amoea

5 In hindwing, CuA nearly straight; antennae frequently as long as forewing (Fig. 28.21) Ameropterus - In hindwing, CuA clearly sinuous; antennae frequently much shorter than forewing 6

6 Hindwing long and narrow, approximately as wide at mid-length as near wing base (Fig. 28.1) Ululodes - Hindwing subtriangular, much wider at mid-length than near wing base Cordulecerus

Key to the genera of Central American Myrmeleontidae (modified from Stange 1970a,b)

1 Sensory pit of distal labial palpomere slitlike; hind leg with elongate sense hair near base of femur present or absent 2 - Sensory pit of the distal labial palpomere oval-shaped; femoral sense hair present on foreleg and midleg, but not on hindleg Subfamily Myrmeleontinae 3 444 28. HENR Y, PENNY, AND ADAMS

Iplicatura Fig.28.14 Aleuropteryx juniperi Ohm (Coniopterygidae). Abdomen showing plicaturae [modified from Meinander 1972]. Fig. 28.15 Mantispinae, fore tarsus (Mantispidae) [original].

2 Hindfemur with an elongate sense hair near base; male with a dorsal, eversible, and long­ haired sac at antero-lateral margin of intersegmental membrane between abdominal segments VI and VII; pretarsal claws strongly arched near base; thorax very hairy; no pigmentation on wing membrane of either sex; size very large, 57-60 mm forewing length; labial palpus not threadlike, not longer than head Subfamily Acanthaclisinae Vella - Hindfemur without an elongate sense hair; male without eversible sac between abdominal segments VI and VII; pretarsal claws nearly straight; thorax not extremely hairy; both fore and hindwings of female with strong pigmentation pattern; size moderate, 30-38 mm forewing length; labial palpus threadlike, twice as long as head Subfamily Palparinae Dimares J~ .' 3 Forewing vein 2A evenly curved from base to hind margin, widely separated from 3A at least at basal one-half (Fig. 28.22) Tribe Brachynemurini Brachynemurus Forewing vein 2A runs close to vein CuP + 1A for a short distance, then abruptly angles towards hind margin or veins 2A and 3A closely associated (Fig. 28.3) 4

4 Radial sector of hindwing arises beyond cubital fork, four or more presectoral cross-veins present; pilula axillaris (Fig. 28.24) present Tribe Myrmeleontini 9 Radial sector of hindwing arises well before cubital fork, one (rarely two or three) presectoral cross-veins present (Fig. 28.3); pilula axillaris absent (Fig. 28.24) Tribe Glenurini 5

5 Pretarsal claws capable of closing against distal tarsomere 6 Pretarsal claws not capable of closing against distal tarsomere 7

6 Posterior fork of forewing vein CuA and vein CuP + 1A parallel with each other and hind margin for a long distance; antennal fossa separated by more than greatest width of antennal pedicel from ocular rim; male ectoproct with elongate post-ventral lobe Dimarella Posterior fork of forewing vein CuA at an oblique angle to hind margin; antennal fossa separated by less than greatest width of antennal pedicel from ocular rim; male ectoproct without elongate post-ventral lobe Elachyleon

7 Forefemur swollen, at widest point much wider than interantennal distance, with abundant pubescence, especially on closing surface; femoral sense hair of foreleg elongate, over one- half length of femur; female ectoproct often with ventral projection Psammoleon - Forefemur slender and elongate, at widest point width about equal at most to interantennal distance, with sparser pubescence; femoral sense hair of foreleg less than one-third length of femur; female ectoproct not produced ventrally 8 THE NEUROPTEROID ORDERS 445

8 Apical one-fifth of hindwing and usually forewing with predominant dark brown suffusion in marked contrast to basal one-fifth (Fig. 28.3); posterior gonapophysis of female weakly produced as a swelling . Glenurus - Apical one-fifth of wings predominantly transparent, sometimes many scattered dark brown spots present; posterior gonapophysis of female well developed, digitiform Eremoleon

9 A series of transverse veins between cross-veins in costal area basal to pterostigma, forming a short longitudinal vein Hagenomyia - No veins connecting costal cross-veins basal to pterostigma (Figs 28.22, 28.23) Myrmeleon

genus from Chiapas, Mexico. Because of the broad distribution pattern of this species, intensive collecting is certain to yield many more specimens. All of the Neotropical sialid species have at one time or another been placed in the genus Protosialis. When Weele (1909) originally described this genus, he designated a Nearctic species as type, leading Ross (1937) formally to synonymize Sialis and Protosialis. Neotropical species (with the exception of Fig. 28.16 Symphrasinae, fore tarsus (Mantispidae) [original]. S. chilensis) generally have narrower wings and more orange on the head and thorax than do their Holarctic relatives. There are only two branches of R2 in the forewing of Neotropical species, while Holarctic species have three or more branches. DISCUSSION OF THE However, any attempt to redefine the generic status of these NEUROPTEROID FAMILES OF species should be incorporated into a broader redefinition of the other austral genera and Holarctic species groups. Until such CENTRAL AMERICA a study is undertaken, it is probably best to leave Central American specimens in the genus Sialis. Order Megaloptera Family COR YDALIDAE (dobsonflies) Dobsonflies or fishflies are large to very large megalopterans, Family SIALID AE (alderflies) up to 160 mm in Wingspan. Three ocelli and unmodified, cyl­ Sialids are medium-sized insects ranging from 10 to 15 mm in indrical fourth tarsomeres make it easy to distinguish the adults length. Adults are smoky to black in colour and possess poorly from Sialidae. Adults are further characterized by flattened sclerotized chewing mouthparts, filiform to moniliform anten­ lateral plates on the head; filiform, moniliform, or pectinate nae, and no ocelli. They may be distinguished from Cory­ antennae; and non-feeding adult habits. They are found near dalidae by their proportionately shorter pronotum, bilobed water and are strongly attracted to lights, although they can fourth tarsomeres, and largely diurnal rather than nocturnal neither walk nor fly very well. Many species show great sexual habits. The subaquatic larvae prefer well-oxygenated, rapidly dimorphism: in Corydalus, males have been seen to duel with flOWing water. Adults are awkward fliers, and courtship each other and to prod the female with their huge mandibles involves abdominal vibration in some species. The eggs are laid during courtship. Some, like alderflies, drum their abdomens in masses of 200 to 1000, during the few days of the female during their sexual displays. Adults live for only 1 or 2 weeks, adult's life. Fewer than 100 species of the family have been but they make a deep impression on those who encounter them, described, although the dominant genus, Sia[is, has an Holarc­ because of their huge size and intimidating appearance. tic distribution. Although adult dobsonflies do not feed, larvae are voracious Alderflies can be locally abundant along streams and lakes predators in ponds and streams, feeding upon bottom-dwelling in temperate climates, but in more tropical climates they are invertebrates, small fishes, and amphibians. They hatch from very difficult to find, although present throughout the New eggs that are deposited near water in one- to five-layered masses World. Only a single species has been described from this area: of 300 to 3000. Grown larvae are commonly used as fish bait Sialis mexicana Banks from Jalapa, Veracruz, Mexico. The Cali­ by fishermen, where they have acquired the names 'hellgram­ fornia Academy of Sciences has four more specimens of this mites' and 'toe-biters' for their aggressive behaviour. A pair of 446 28. HENRY, PENNY, AND ADAMS

11:' _o} Fig. 28.17 Hemerobius sp. (Hemerobiidae) [redrawn from 1A Richards and Davies 1977]. C, costa; CuA, cubitus anterior; CuA CuP MA CuP, cubitus posterior; MA, media anterior; MP, media , MP posterior; R, radius; Sc, subcosta; 1A, first anal vein. I .~. .~ ..~ t;:;

< i .~. t ,J hooked prolegs at the tip of the abdomen replaces the anal dalinae and Chauliodinae, with only the former group found filament of the sialid larva, permitting easy identification. In in Central America. Corydalinae seems the more specialized of the temperate zone, the life cycle is 1-5 years long. Life spans the two, based on venational and genitalic features. The genus of Central American corydalids are not known. Platyneuromus (Figs 28.2, 28.12) is endemic to this region, and Corydalidae is usually divided into two subfamilies, Cory­ its three species were revised by Glorioso and Flint (1984). The i• ~ genus Chloronia was revised by Penny and Flint (1982) and ';j four species were reported from Central America. The genus Corydalus is in need ofrevision. Penny (1977) lists seven species 1 and two subspecies from this region, several of which are ~ probably synonyms. S f t Order Neuroptera, Suborder Hemerobiiformia ! ap-ect ~ Family CONIOPTERYGIDAE (dusty-wings) Adult dusty-wings differ radically from other Neuroptera in their small size, much reduced wing venation (Fig. 28.13), distinctive hamuli-like wing coupling mechanism, and hypo­ dermal wax glands. Additionally, the ganglia of their central ~ nervous systems are extensively fused, and their malpighian tubules number six rather than eight and are histologically i unique. Because the dusty-wings are covered with a whitish 11 powder, they closely resemble whiteflies (Hemiptera, suborder i Homoptera: Aleyrodidae), upon which they feed. Larvae are f equally peculiar. Like the adults, they are covered with a f whitish, mealy wax, are serially fused internally, and feed I principally upon mites, red spiders, and plant lice. They pupate in a flattened, double-walled cocoon. Coniopterygidae share Fig.28.18 Chrysopidae, generalized male genitalia [from Adams and Penny i 1987]. acu, acumen; ap-ect, apodeme of ectoproct; ap-gs, apodeme of few synapomorphies with other neuropteroid taxa, suggesting gonarcus; ent, entoprocessus; gcr, gonocristae; gp, gonapsis; gs, gonarcus; early origin and differentiation of the family. gsac, gonosaccus; mi, microtholi; mu, mediuncus; ti, tignum; 9S, sternite The general biology of coniopterygids is poorly known. 9. Apparently, adults are quite long-lived and active. CourtshiP I ~ 5 I

THE NEUROPTEROID ORDERS 447

involves the seizure of the female's legs by the jaws and legs of 1931), but in nature the story might be quite different. Our the male. Fluid semen is then transferred to the female; this is present understanding suggests a close phylogenetic relation­ in contrast to all other neuropteroids, in which insemination ship between the Polystoechotidae and Ithonidae, as first pro­ occurs via a spermatophore (MacLeod 1962, unpublished). The posed by Withycombe (192 5). small, ovoid eggs, with their conical micropylar projections, This is the smallest family of Neuroptera, with only three are deposited singly or in small groups on leaves and conifer species-two in North America and one in Chile. Poly­ needles. There is some potential for the use of adults and larvae stoechotids are highly attracted to lights, and in regions where in the biological control of mites and piantlice in orchards (New lights occur have soon disappeared from the fauna. In North 1986a). America, they can no longer be found over much of their former Members of this family are seldom collected, but in fact may distribution. at times be among the most abundant Neuroptera in a given There are three specimens of Polystoechotes punctatus (Fig. locality. They can be taken by beating bushes and shrubs, 28.8) in the California Academy ofSciences collected at 1585 m where the adults flutter weakly away from the branches. Often elevation in Chiriqui PrOVince, Panama. The Museum of Com­ they feign death and can be picked from beating sheets. Dusty­ parative Zoology, Harvard University, has another specimen wings are also taken in flight-traps and light-traps, but if the from Lino, Panama, and the US National Museum holds a trap containers hold liquids the distinctive white powder dis­ specimen from Mount POlis, Costa Rica. These are the only appears and they look much like small four-winged flies. known records of this species from outside North America. About 250 species exist, throughout the world. The two principal subfamilies Aleuropteryginae and Coniopteryginae are very distinct. The former seems the more specialized of the Family SISYRIDAE (spongillaflies) two, although the dominant genera of the family, Coniopteryx These interesting neuropterans are small (6-8 mm) lacewings and Semidalis, are coniopterygines. The aberrant Bru­ that superficially resemble the hemerobiids (brown lacewings), cheiserinae constitutes a third subfamily, with three species but may be distinguished from the latter by their simple, open from arid regions of southern South America (Riek 1975). Wing venation and one-stemmed origin of the branches of the Meinander (1986) indicates the presence of only two species radial sector (Fig. 28.9). They occur throughout the world near of Aleuropteryginae: Neoconis dentata Meinander from Guat­ lakes and streams, but can be taken at lights a mile or more emala and Costa Rica, and Neoconis unam Monserrat from from water. The greenish or brownish larvae of spongillaflies southern Mexico (Fig. 28.13). Within the Coniopteryginae, are aquatic predators (parasites?) of freshwater sponges, imbi­ Meinander (1981, 1986) listed ten species of Coniopteryx from bing the liqUid contents of the 'prey' through extremely elong­ Central America: five species in the subgenus Scotoconiopteryx and five species in the westwoodi group of the subgenus Con­ iopteryx. The only species of Semidalis known from Central b.g. America is S. panamensis (Meinander 1974). C.g.

Family POLYSTOECHOTIDAE (giant lacewings) The giant lacewings are an exclusively New World family of peculiar, apparently ancient Neuroptera, which share many primitive (plesiomorphic) features with Megaloptera. In fact, the larva is the most structurally primitive in the Hemero­ biiformia, and could easily be mistaken for a terrestrial hell­ grammite (megalopteran). Head and jaw morphology of the polystoechotid larva is ithonid-like, but the larva lacks the fossorial scarabeiform specializations of the moth-Wing imma­ ture. Like the ithonids, larval giant lacewings may be detri­ tivores or even herbivores rather than predators; their natural habits and habitats are unknown. Adults, which measure 40­ 70 mm across the wings, also possess several specializations in ovd common with the Ithonidae. Little is known of their habits, except that they are predatory and sometimes associated with the banks of streams and rivers. An occasional mass emergence of Polystoechotes punctatus (FabriciUS) has been recorded in 5P pg 5g temperate North America (Fyles 1903). Eggs of polystoechotids Fig. 28.19 Chrysopidae, generalized female genitalia [original]. a.g., are like those of ithonids, with a buttonlike micropyle; they are accessory gland; b.d., bursal duct; b.g., bursal gland; cop. b., copulatory laid singly and scattered about by captive females (Hungerford bursa; ovd, oviduct; pg, praegenitale; sg, subgenitale; sp, spermatheca. 448 28. HENRY, PENNY, AND ADAMS

Family MANTISPIDAE (mantid-flies) These medium-sized insects are strikingly convergent on praying mantids (Dictyoptera), possessing elongated pro­ thoraces and procoxae, enlarged and spined profemora, and t~ 1 recurved protibiae (Fig. 28.5). As adults, like mantises, they ~ ~ capture and eat a variety of active insects with their raptorial i ~ front legs, but as larvae they are specialist predators or parasites. ,';j Adults can be active nocturnally, or diurnally; species that t ! forage during the day often congregate in large numbers on i it flowers, and even take some nectar (Opler 1981). A number of I New World taxa, like Climaciella brunnea, are extremely con­ f vincing mimics of social wasps, and are usually found in close i association with their specific models (Boyden 1984). A long ,~ ovipositor is present in the more primitive species. ~ '; There are four distinct subfamilies of Mantispidae, according to Lambkin (1986). Members of the morphologically homo­ geneous subfamily Mantispinae possess a single pretarsal claw on the prothoracic leg (Fig. 28.15) and are associated as larvae with spiders and their egg cases. The other three subfamilies, Fig. 28.20 Lainius constellatus Navas (Chrysopidae: Apochrysinae) [from which until recently were united in the Platymantispinae, Kimmins 1952]. exhibit more diversity of both form and habits, although all are characterized by two pretarsal claws on the prothoracic legs of the adult. The symphrasine subfamilies are often obligate associates of polybiine wasps, feeding as larvae on the brood of the social insect host. However, some have been found among § ate, flexible, hairlike jaws. The mature larva respires by means bee broods, within moth cocoons, or in association with scarab of seven pairs of segmental abdominal gills, and swims in a beetle pupae (New 1986a). In fact, it may turn out that the I vertical position by using a snapping motion of the body (Parfin larvae of genera such as Plega and Nalima are generalist pred­ and Gurney 1956). ators (MacLeod and Redborg 1982). Adults feed on organic detritus, pollen, soft-bodied insect Mantispid larvae are among the most highly modified in all prey, and honeydew (Duelli 1983). Activity is crepuscular and of the Neuroptera, being quite maggot-like. They are considered nocturnal, although adults have been seen to swarm on sunny by many to be true parasitoids, displaying heteromorphic days at some locales. Courtship in Climacia consists of the male development and often a high degree of host specificity. An extending and fanning one foreWing toward the female's head. active triungulin hatchling seeks out spider egg sacs or hymen­ The elongate eggs are deposited singly or in small masses in opteran nests; it then typically transforms into a nearly legless, depressions on objects overhanging the water, and are protected physogastric grub, which spends the rest of its life in one place. by a thin layer of silk spun from the tip of the mother's abdomen Some spider-associated mantispine taxa must jump aboard host (Pupedis 1980). females before they are able to recognize spider egg sacs as food This is one of the smaller families of Neuroptera with only (Redborg and MacLeod 1983). Blood-feeding on the spider by about 50 species known. Their phylogenetic affinities are the attached triungulin has been reported in Mantispa spp. unclear, although recent work on female reproductive anatomy (Redborg 1982). Possibly, symphrasine triungulins penetrate

oJ, has linked them closely to the Neurorthidae and more generally the colonies of social insects in a similar manner, by hitchhiking ,;:, to the Berothid/Mantispid/Dilarid complex (Pupedis, unpub­ on the foraging workers. lished). Mature larvae pupate in a silk cocoon spun within or near the There is no subfamilial or tribal classification for the spon­ consumed host. Upon emergence, adults engage in elaborate t , l gillaflies. Two of the three known genera occur in Central courtship activity that often involves visual and olfactory dis­ ~ America, and are easily separated by their basic wing color­ plays. Spermatophore production accompanies sperm transfer. ation: Sisyra has smoky, dark Wings, and Climacia displays pale The minute, pale eggs are often laid on very short stalks (Riek t •1.' wings with dark spots and stripes. Two species of Sisyra are 1970). ~ known from Central Africa, S. apicalis Banks and S. panama Mantispidae are closely related to Berothidae; in fact, taxa of ! Parfin and Gurney, both from Panama. There are also two intermediate morphology and habits exist in South Africa, species of Climacia in Central America: C. striata Partin and South America, and Australia. For example, Tjeder (1959) Gurney from Panama and C. tenebra Partin and Gurney from described a subfamily of Berothidae from southern Africa Honduras. (Rhachiberothinae) with raptorial forelegs like Mantispidae, li • ~ •

THE NEUROPTEROID ORDERS 449

but lacking the tarsal segmentation and other specializations (Gerstaecker). This species is rather large, with bright yellow of primitive Mantispidae. In a subsequent paper Tjeder (1968) meso- and metascutella and crescent-shaped yellow markings indicated that the males of Rhachiberothinae had four tarsal on the pronotum. The largest genus by far is Mantispa. Members segments, like Platymantispinae, while females had five tarsal of this genus vary widely in size, markings, and male genitalia, segments, like other Berothidae. Certainly, the relationship although they are outwardly similar. Penny (1983a) recog­ between these two families is in need of re-evaluation, and nized several species groups within this genus for the Brazilian there may be cause to unite both as a single family (Aspock species, and probably most North and Central American species and Aspock 1988). The nearest relatives of the Bero­ would fit into these same species groups. A similar cladistic re­ thidae/Mantispidae are probably the dilarids, sisyrids, and evaluation of all members of the genus is needed. There have neurorthids, with dilarids (Fig. 28.7) being the most similar in been ten species of Mantispa described from Central America, larval morphology and adult appearance. nine of them by Padre Longinos Navas. There is certainly Three of the four subfamilies of Mantispidae are represented much synonymy still present in this genus, and a revision and in Central America: Calomantispinae, Symphrasinae, and redescription of the Central American species is necessary. Mantispinae. Drepanicinae, with its greatest diversity in Aus­ tralia, is absent from North and Central America. Calomantispinae is the most easily recognized subfamily Family BEROTHIDAE (beaded lacewings) because of the very short subcostal vein and bifid fore tarsal Although the family Berothidae has a Wide, albeit discon­ claws, which give each foreleg the appearance of having four tinuous, world distribution, records have not been published claws. This subfamily has but a single Central American genus, from Central America. However, Penny (1983b) recently Nolima, with two species: N. infensus Navas from Costa Rica, described a species of the North American Lomamyia (Fig. and N. pugnax (Navas) from Guatemala. 28.11) from northern South America, and in the P. Adams Symphrasinae includes some of the smallest mantispids, and collection there is a series of L. squamosa Carpenter from Mount is the subfamily most closely related to the rhachiberothine Pine Ridge, Belize. An unidentified species of Lomamyia was Berothidae. The elongate projection of the first fore tarsal collected on Barra Colorado Island, Panama. There is also an segment, as well as the sub-apical position of the second tarsal undescribed genus ofBerothidae, with one species in Costa Rica segment and the presence of only four tarsal segments on the and another in Venezuela. foreleg (Fig. 28.16), make this a distinctive group. There are As mentioned above, beaded laceWings are close relatives of only two genera found in the region, separated by a sub-basal the Mantispidae, and are thus not well delimited by unique, femoral spine in Plega which is absent in Trichoscelia. The genus specialized features. They are named for their characteristic Trichoscelia is predominantly South American, but three species moniliform antennae. Adults are small- to medium-sized [T. banksi Enderlein, T. santaremi (Navas), and T. tobari (Navas)] insects, with seedlike scales and long hairs on the wings and are known from tropical southern Mexico. The genus Plega bodies of the females of some species. They are crepuscular and appears to be found farther north, from the Amazon Basin of relatively sedentary, resting on the undersides of leaves and South America to the south-western USA. Two species of Plega feeding on nectar, honeydew, pollen, or small insect prey have been described from tropical southern Mexico, P. variegata (MacLeod and Adams 1967; Tjeder 1968). Courtship is simple Navas and P. yucatanae Parker and Stange. A third species, P. and swift and involves Wing-fluttering by both sexes (MacLeod signata (Hagen), has been cited from Colombia, but its known and Adams 1967). The female deposits her elongate, ovoid eggs distribution is usually considered to be the south-western USA. on hollow, basally enlarged stalks, in clumps with fewer stalks Mantispinae is the most speciose subfamily in Central than eggs (Toschi 1964). America, with three genera and 12 species described. Climaciella Larvae closely resemble Dilaridae (below) in the construction includes a number ofvespid wasp mimics, with C. brunnea (Say) ofthe larval head capsule and mouthparts. Habits of the imma­ haVing several morphs mimicking different vespids species in tures are poorly known. Larvae of Australian Spermophorella different areas (Opler 1981). Only the one species, C. brunnea, are apparenty free-liVing on soft-bodied prey, but those of North has been collected in Central America. The genus Entanoneura American Lomamyia are associated with termites and ants and is also monotypic in Central America, represented by E. limbata transform at the first moult into inactive, mantispid-Iike grubs;

C Sc R Fig. 28.21 Ameropterus delicatulus (MacLachlan) (Ascalaphidae), hindwing; nDte CuA [Driginal]. C, costa; CuA, cubitus anterior; CuP, cubitus pDsteriDr; CuP+ lA, fused CuP and first anal vein; MA, media anteriDr; MP 1, anteriDr branch Df media posteriDr; MP2, posteriDr branch of media pDsteriDr; Rs, radial sector; Rs + MA, fused radial sector and media anterior; Sc, subcosta; 2A, secDnd anal vein; 3A, third anal vein. 450 28. HENR Y, PENNY, AND ADAMS

~-:===--O-O-CJ-C=--:::::-~C oratory, however, larval dilarids have been reared on a variety of disabled, soft-bodied insect prey (MacLeod and R­ Spiegler 1961). A Eurasian species of Dilar may tunnel in soil. ~~MA Adams (1970) divides pleasing lacewings into two sub­ ~~~MP families, based on profound differences in male claspers. Dilari­ ~~~V~CuA nae, including about 22 species in several genera, is mostly -v confined to the Old World. Nallachinae has but a single genus, 1A fold Nallachius, with about 13 species, all in the New World. Oddly, no dilarids are present in the Australian region. One species

~2A has been recorded from Central America: N. championi (Navas) ~ from Guatemala. With more intensive collecting, additional species should be discovered. Fig. 28.22 Brachynemurus sp. (Myrmeleontidae), base of forewing. Note vein 2A [modified from Stange 1980]. C, costa; CuA, cubitus anterior; CuP, Family HEMEROBIIDAE (brown lacewings) cubitus posterior; MA, media anterior; MP, media posterior; R, radius; Sc, subcosta; lA, first anal vein; 2A, second anal vein; 3A, third anal vein. The brown lacewings constitute a large family of small- to medium-sized insects which are predators, both as larvae and they then resume activity in the third stadium. Food capture adults. Although they are usually brownish (as the common by Lomamyia latipennis is the stuff of science fiction, and is name suggests), many species shine with iridescence. The adult unique among all Insecta. The first or third instar larva releases habits are poorly known, partly because the insects are cryptic an air-borne chemical from its anus, called an 'aggressive in colour and behaviour, and have a crepuscular or nocturnal allomone', which rapidly paralyses and eventually kills activity cycle. Most species are found on trees and large bushes, termites. Once immobilized, the prey is consumed at the bero­ but a few seem to be associated with more open, grassy areas. thid's leisure (Johnson and Hagen 1981). A number of unrelated taxa within the family have evolved Fewer than 60 species of beaded lacewings have been flightlessness, presumably in response to isolated habitats; this described, yet these few species form four distinct subfamilies trait is very rare in other neuropteroid insects. In general, containing nearly 25 genera. Australia has a particularly rich hemerobiids are far more abundant in temperate than tropical berothid fauna, with nearly 30 described species in six genera climates, and are extremely tolerant of low temperatures. (Aspock 1983). However, a few genera, such as Notiobiella, seem to thrive in the tropics, including Central America. Larvae are very similar in appearance to Chrysopidae, Family DlLARIDAE (pleasing lacewings) although more elongate than the latter, possessing relatively Pleasing lacewings are small, rather nondescript insects that shorter jaws and smaller heads, and lacking the setose tubercles are easily confused with brown lacewings (Hemerobiidae). typical of so many green laceWing aphis lions. They also possess However, dilarids possess a few features not found in hemero­ only three stemmata on each side of the head, instead of the biids, including ocelli and a long ovipositor in the female. six of chrysopids, and never carry trash (debris) on their bodies. Also, males of Dilaridae are immediately recognizable by their spectacular, coarsely pectinate or flabellate antennae, and both sexes display striking patterns of spotting and banding of the wings (Fig. 28.7). Unfortunately, when collected in containers of fluid in traps with other insects, segments of the male's LJOOOcJ[s: antennae break off very quickly, giving them the appearance R-~MA of small moths lacking wing scales. This is a tropical group, seldom encountered in temperate climates. They are considered rare, but can be collected with ~~::A some regularity in the tropics by placing a light-trap in the forest canopy during the dry season (Penny and Arias 1980). "~'I, Immature stages of only Nallachius americanus are known with 3A '\: certainty. Eggs of that species are elongate, not stalked, and are ~i.··· closely packed into spaces within bark or decaying wood. The \' larva is long in body but short-limbed, possessing enlarged ":: fossorial front legs and eyes that are reduced to a single stemma Fig. 28.23 Myrmeleon sp. (Myrrneleontidae), base of forewing. Note vein per side. Larvae and pupae have been recovered from weevil 2A. [modified from Stange 1980] C, costa; CuA, cubitus anterior; CuP, galleries in dead tulip trees (Gurney 1947), suggesting that cubitus posterior; MA, media anterior; MP, media posterior; R, radiUS; Sc, Nallachius pursues weevil larvae in their tunnels. In the lab­ subcosta; lA, first anal vein; 2A, second anal vein; 3A, third anal vein. i "'0>. If'' ~ THE NEUROPTEROID ORDERS 451

Feeding habits are much the same as in green lacewings: the (Fig. 28.17) can generally be used qUickly to separate Hemero­ larvae take aphids, whiteflies, scales, and other soft-bodied, bius from all other Central American hemerobiids. sessile prey from leaves, conifer needles, or young stems. Some The most diverse genus of Hemerobiidae in this region is species are probably prey specialists, which may account for Megalomus. These truly brown laceWings are easily identified the pronounced habitat specificity or preference observed in by the large number of radial sectors (4-6) and presence of a certain taxa (New 1984). They pupate within a loosely con­ recurrent costal cross-vein. Six species of this genus are known structed cocoon. Adults court and mate in a complex ritual from Central America: M. angustatus Navas, M. exterior Navas, involving abdominal vibration. The female can live for several M. insignis Kimmins, M. minor Banks, M. punctatus Kimmins, months, producing up to 2500 fertile eggs during that period and M. serrinus Navas. A revision of this genus is needed, and (Laffranque and Canard 1975; Miermont and Canard 1975; some synonymy and new species probably exist for the region. Neuenschwander 1975). Eggs are unstalked, ovoid, and laid in small batches. Because of their efficient foraging strategies, Family CHRYSOPIDAE (green lacewings; golden-eyes; preference for plantlice, and high fecundity, Hemerobiidae are serious contenders for important roles in the biocontrol of crop stinkflies) pests (New 1975; Neuenschwander and Hagen 1980). Green laceWings are the group of Neuroptera that most people There has never been a Widely accepted subfamilial or tribal think of first. With their pale green wings, golden eyes, and classification within this family, although three to five sub­ fluttering flight they are a common and memorable part of families are usually recognized. The 800 or so species and temperate faunas. On warm summer nights they can almost 80 or more genera of the family are badly in need of revision. always be seen at lights and windows. They are an important In Central America, six genera and 20 species are known. The part of agricultural ecosystems, where they are highly valued genus Sympheromima appears to be a closely related Neotropical agents of biological control. However, in the tropics, they are replacement for the antitropical Sympherobius. Only a single often much harder to find. Those which do manage to flutter species has been described within Sympheromima, S. marginata into lights and windows at night are frequently gobbled up by Kimmins, from 4000 to 5000 feet elevation in Guatemala. The geckos. Despite their paucity in most natural habitats in the type specimen lacks an abdomen, making comparisons difficult. tropics, they are still common in agricultural situations, where When this species is better known, Sympheromima may be they help protect crops from pests such as aphids, whiteflies, synonymized with Sympherobius. scale insects, and small caterpillars. Notiobiella (Fig. 28.6) includes two Central American species: Adult Chrysopidae are qUite conservative externally, dis­ ,'J. rubrostigma Navas, and N. spinosa Monserrat and Penny. playing very similar wing venational patterns and body mor­ Recent descriptions of all New World species appear in Mon­ phology in most species. The best diagnostic traits are found in serrat (1983) and Monserrat and Penny (1983). the male and female genitalia (Figs 28.18, 28.19). Colour Nusalala appears to be the Neotropical replacement for the patterns of the head and thorax can also be helpful, but much otherwise cosmopolitan genus Micromus. The two genera can of this coloration, being located in the subcuticular hypodermis, be separated by the three rows of gradate veins, more complex fades soon after death; field notes on colour patterns of live endoprocessus, shorter ectoproct projection, and elongate individuals are therefore valuable. They are medium-sized central shaft of the parameres in Nusalala (Penny and Monserrat insects, usually of various shades of green, although the 1983). Four species have been described from Central America: endemic Hawaiian genus Anomalochrysa can show brilliant N. championi Kimmins from Guatemala and Panama, N. krugeri pigmentation. In contrast to Hemerobiidae (Figs 28.6, 28.17), Nakahara from Veracruz, Mexico, N. marginata Navas from the Rs vein of chrysopids runs parallel to R1 for most of its Costa Rica, and N. uncata Kimmins from Panama. length, and a pseudomedia is present (Fig. 28.10). They possess Wesmaelius is generally considered to be a northern tem­ generalized mandibulate mouthparts, and many species are perate genus, with most species in North America being found predators of soft-bodied plantlice. Others, like the closely related in the northern part of the continent or at high elevations along genera Chrysopiella and Eremochrysa, seem to feed mainly on the highest mountain chains. One species, W. magna Kimmins, pollen Oohnson 1982). Another significant fraction of lace­ has been described from 2100 to 3150 m elevation in the Wings, notably many species of the worldwide genus Chryso­ QUiche Mountains and Totonicapam of Guatemala. perla, survive and reproduce solely upon honeydew and nectar Another genus normally associated with cooler temperate (Hagen et al. 1970). These sugar-feeding taxa acqUire the climates is Hemerobius. However, three species of this genus amino acids necessary for abundant egg production from sym­ described from Central America are: H. jucundus Navas from biotic Torulopsis yeasts which occupy specialized, heavily tra­ MeXico, H. tibialis Navas from Costa Rica and Mexico, and H. cheated tissues of the midgut region of the insect's intestinal tolimensis Banks from Guatemala. Kimmins (1928) separated tract (Hagen and Tassan 1966). one additional Wide-winged form, H. withycombei Kimmins, as Many adult chrysopids produce foul-smelling odours when a new genus, Anotiobiella, but without a more thorough study disturbed or during courtship, which is the basis for the appel­ of the diversity within the genus Hemerobius this decision seems lation 'stinkflies'. The odoriferous secretions are produced by premature. The three radial sectors and recurrent cross-vein the prothoracic glands, emptying near the base of the head. 452 28. HENRY, PENNY, AND ADAMS

One of the active constituents of these secretion is scatole, carrying species. In the temperate zone, the insect may over­ , which gives faeces and carrion their characteristic and familiar winter in this stage, or as an adult or even a larva. In many t perfume (Blum et al. 1973). It has been suggested that these members of Chrysoperla, adult winter diapause, mediated by ~ prothoracic gland secretions repel ants, which would be a trait photoperiod (Sheldon and MacLeod 1974), leads to a radical of great value to those lacewing species that feed upon aphids change in the insect's pigmentation, from solid green to reddish and scale insects: most plantlice are vigorously defended by or yellOWish brown. The green colour gradually returns as formicids. However, a positive correlation between predatory diapause terminates and reproduction begins or resumes. habits and stinkiness, which would support the hypothesis, has Courtship and mating behaviour of green lacewings is quite yet to be established. complex. We have already mentioned the presence of strong­ smelling compounds released during the courtship displays of presectoral crossveins some stinkflies; Chrysopa, Meleoma, and a number of other ~-rlr genera typify this condition. In Meleoma, the female feeds during courtship on the glandular secretions produced by specialized tubercles and cavities present on the heads of the males (Toschi 1965). However, the most remarkable reproductive behaViour is the elaborate 'acoustical' communication that has thus far been found in Chrysoperla and Chrysopiella (Henry 198 Sa, b, 1986). In those genera, both males and females produce vibrational, substrate-borne signals of great compleXity, by jerking their abdomens rapidly up and down. In a given species, Fig. 28.24 Myrmeleoll sp. (Myrmeleontidae), base of hindwing of male, the 'songs' of both sexes are usually identical, and the two showing piJula axillaris and presectoral cross-veins [modified from Stange individuals of a courting pair reciprocally exchange songs in a 1980]. precise duet. If a stable, prolonged duet cannot be established between the potential mates, copulation will not occur; the Chrysopid larvae are known as aphis lions, and feed songs of the two partners must match if courtship is to succeed. voraciously on a variety of soft-bodied arthropod prey. They The songs thus have a reproductive isolating effect between vary in shape from the elongate, thin larvae of Hypochrysa to closely related, potentially interfertile species, and are probably the very stout, robust Italochrysa from Europe. Their jaws are important in the process of speciation in certain subgroups of thinner and their heads more robust than is typical of hemero­ acoustical genera. ln fact, several valid, reproductively isolated biids. Trash-carrying behaviour is distributed widely in the species of Chrysoperla are morphologically indistinguishable in family; in fact, the habit is found in all Central American taxa, every respect, and can be told apart only by their calls (Henry excepting Chrysoperla. Trash carrying is always correlated with 1983). the presence of dorsolateral setigerous tubercles and hooked, Green lacewings lay stalked eggs, in various configurations. glandular setae. In many cases, the trash packet is constructed Most often, eggs are oviposited singly, but localized batches of from very specific materials-the husks of consumed prey, par­ eggs, or even many eggs sharing a single, thickened stalk, are ticles of a certain size, the fluff scraped from hirsute or pilose found in some species. Hawaiian Anomalochrysa is the only leaves, or even the lichens growing on rocks and tree branches. genus characterized by unstalked eggs. Individual females of Most chrysopid larvae forage actively in uncropped grasses, Chrysopidae are not quite so fecund as those of Hemerobiidae, shrubs, or trees, but a few species prefer the litter of the forest although recent work (Henry and Busher 1988) documents floor. Trash carriers are usually less mobile than naked types. lifetime production of well over 1000 fertile eggs by females of When threatened by predators, the larva manipulates the trash several North American taxa. More than any other neur­ packet ('casita') as a shield. opteroid family, the green laceWings have been successfully The most highly modified larval habits and morphology are employed in comprehensive programs of biocontrol, par­ found in the Old World genus Italochrysa, a member of the ticularly in the temperate zone (New 1975). unique tribe Belonopterygini. These grub-like, slow moving The present classification of Chrysopidae is based on Adams insects are inquilines or parasites of ants. Larvae of ltalochrysa (1967,1978) and Adams and Penny (1987). These authors italica, for example, have been found only within the nests of recognize two extant subfamilies, the primitive Nothochrysinae Crematogaster ants, where they feed on the early stages of their and the successful and dominant Chrysopinae. Separating the hosts (Principi 1946). Australian relatives of 1. italica, as well two taxa is the development in Chrysopinae of a special pres­ as New World belonopterygines such as Nacarina, have similar sure-sensitive tympanal organ, which occupies a swollen i associations with various Formicidae (New 1983; Weber region of the radial vein of the forewing. This unusual 'ear' can ~~ 1942). Extremely unusual larval morphology and coloration detect the ultrasonic pulses of echolocating bats, enabling the also characterizes the Hawaiian lacewing genus Anornalochrysa. insect to take evasive action in the presence of such aerial !: Pupation takes place in a compact, closely woven cocoon, predators (Miller 1984). .£ which may incorporate debris and broken setae in the trash- Nothochrysinae has a wide, discontinuous distribution, but

..j~'. • .. 7777; 7In 'P .. ,'.

THE NEUROPTEROID ORDERS 453 does not occur in Central America. In contrast, four tribes of and overall coloration, but males of most species have elab­ Chrysopinae are well represented in this region. The largest orately developed antennal bases and have horns between the and perhaps loveliest of the Central American green lacewings antennae on the frons. Often these horns are found in frontal are the Apochrysini (considered to have subfamily status by depressions or pits; as mentioned earlier, females lick secretions Brooks (1990) in his generic revision of the family). These from these horns and cavities during courtship. Males of a few dramatic insects, with their very broad wings (Fig. 28.20) and species and females of all species are much less distinctive. long antennae, appear to be confined to mature forests and are Male genitalia (Fig. 28.18) include a gonarcus with arcessus, seldom collected in large numbers. In fact, no more than half pseudopenis, entoprocessus, gonapsis, and gonocristae (Adams a dozen specimens are known of any of the Neotropical species. 1962). The three species known from tropical Central America The insects of this tribe measure 40-70 mm across the out­ are M. antennensis Tauber, from Chiapas, Mexico; M. dolich­ stretched wings. Many species have shiny dark pustules or dark arthra (Navas), from Chiapas and Guatemala (Tauber 1969); pigmentation in the middle ofthe wings. There are three genera and M. macleodi Tauber, from Chiapas and Honduras. and three species known from Central America: Domenechus Probably the best known genus of Chrysopini is Chrysoperla. mirifica (Gerstaecker), Lainius constellatus Navas, and Loyola This is the genus that has been studied most intensively for croesus (Gerstaecker). biological control in temperate agriculture, even though the The tribe Belonopterygini contains the stoutest members of adults are largely non-predatory and cannot contribute to the the Chrysopidae, both as adults and larvae. They have a broad destruction of plantlice. In tropical climates, Chrysoper1a does pronotum which is wider than long, and stocky antennae with not constitute such a large percentage of any local fauna, basal segments that are also wider than long. As mentioned but can be locally abundant in grasslands and agricultural earlier, species of Belonopterygini in both Europe and South situations. Morphologically, the genus is quite uniform, with America have been documented or implicated as ant nest species displaying stable and similar patterns of coloration. The associates. The only description of a larval belonopterygine in antennae are pale; there is a red patch on the gena just behind the New World is that of Weber (1942), of a blue larva of the mandibles; and a pale stripe courses down the middle of the Nacarina in a nest of Camponotus ants in Guiana, South America. thorax and abdomen. In New World species, the ninth sternum A single genus of Belonopterygini is known from Central of the male consistently bears a node at the apex, and male America, including the species Nacarina balboana (Banks) from genitalia always bear an elongate, apically curved arcessus. Panama, Costa Rica, and southern Mexico, and N. titan (Banks) Many valid species in the Holarctic Region are impossible to from Costa Rica. separate by morphological criteria, and the same may be true Our understanding of the Neotropical members of the import­ in the tropics. Two species of Chrysoper1a are known from ant tribe Chrysopini has been much advanced in recent years Central America, C. exotera (Navas) and C. externa (Hagen). through the descriptions of Ceraeochrysa (Adams 1982a) and They can be told apart by genitalic characters, described in Plesiochrysa (Adams 1982b), as well as by surveys of the Adams (1962). Amazon Basin species (Adams and Penny 1987). All four The fourth tribe of Chrysopinae, the Leucochrysini, is a genera of Chrysopini of the Amazon region are also found in complex taxon which reaches its greatest development and Central America. Two species of Chrysopa (Plesiochrysa) are species richness in central and northern South America (Banks found throughout Central America: C. brasiliensis Schneider 1945). Berchmansus can be separated easily from the rest of the and C. e10ngata Navas. The genus Ceraeochrysa reaches its tribe by the presectoral cross-vein of the forewing (Fig. 28.10). greatest development in this area, with at least 12 known Only a single species of this genus has been found in the Central species (Adams 1982a). The genus Chrysopodes can be further American region: Berchmansus cinctipes (Banks) from Panama. subdivided in this region into C. (Chrysopodes) and C. (Neo­ The other three genera encompass sibling species complexes suarius). Chrysopodes (Neosuarius) reaches its greatest diversity that, like many Chrysoperla, can be separated only with diffi­ and abundance in the Andean region, where many of the culty. In general, the species with the darkest wings have species are darkly pigmented. In the lower, warmer, moister been placed in Gonzaga; those with a quadrate intramedian cell habitats the species are green. The species C. (Neosuarius) co1­ have been directed to Leucochrysa; and species with a triangular laris (Schneider) appears to be distributed throughout Central intramedian cell have gone into Nodita. Unfortunately, the America. Three other species of Chrysopodes (Neosuarius) also shape of the intramedian cell is closely linked to overall wing have been collected from this region. The more typical Chry­ size, with larger specimens usually having the quadrangular sopodes (Chrysopodes) reaches its greatest diversity in the moist shape. Thus, specimens of the same species can sometimes be lowland forests of northern South America. Nonetheless, this keyed to different genera. The present classification is unsat­ subgenus of Chrysopodes has about six representatives from the isfactory and needs revision. Central American region, including C. (C.) nevermanni (Navas), Only a single species of Gonzaga is found in Central America: C. (C,) costalis (Schneider), and C. (C.) jubilosa (Navas). G. torquatus Navas from Guatemala and Panama. Nine or ten One primarily Nearctic genus of Chrysopini, Meleoma, has species of Leucochrysa and approximately 25 species of Nodita three species in northern Central America. This genus appears have been described from this region. The last two genera superficially similar to the other Chrysopini in wing venation especially are in need of revision. 454 28. HENRY, PENNY, AND ADAMS

facture (Henry 1972). Split-eyed ascalaphids, such as Ululodes Order Neuroptera, Suborder spp., are characterized by highly modified, oil-covered repagula Myrmeleontiformia that effectively prevent ants or other predators from approach_ ing the fertile eggs. This 'barrier' function also protects the i Family ASCALAPHIDAE (owlflies) newly hatched larvae, which remain on the egg twig for several Ascalaphidae is a truly unique family, and unfamiliar to most days after eclosion. Entire-eyed ascalaphids possess more t t people. Its members are large, handsome insects (35-130 mm normal-looking abortive eggs with no apparent capacity to 1; wingspan), but they are cryptic and crepuscular in their habits repel enemies. Here, the function of repagula is trophic: newly and inactive by the time most entomologists set up light-traps hatched larvae walk down to the abortive eggs and take their % at night; consequently, they are rarely encountered unless first meal. The larvae then regroup on the egg shells and actively :( specifically sought after, and their behaviour is poorly known. defend themselves en masse from enemy attack. Oddly, repagula i~ Many of the largest ascalaphids resemble antlions, but others do not seem to be present in Old World owlflies of either subfam­ :~ look very much like dragonflies, except for the extremely long, ~" ily (Henry 1978; New 1986b). 'If ,.' knobbed antennae incongruously attached to the head. The Ascalaphidae is a diverse family of perhaps 400 species in ~;. l;~ dragonfly-like taxa are swift, skilful fliers and aerial predators, 65 genera, distributed throughout the temperate and warmer ({ filtering small 'insect plankton' from the air with their hairy regions of the world. In the Old World, many species are day iill legs. Also in the manner of dragonflies, courtship and mating fliers, with broad, brightly pigmented, and patterned Wings, .j! ~. takes place on the wing (Henry 1977). A thick covering of hair like butterflies. The precise limits of the family are not well is present on the thorax, insulating the flight muscles and defined, as some of the entire-eyed taxa merge morphologically permitting them to operate more efficiently at a higher tem­ with the antlions (Myrmelontidae). The studies of Stange and perature. In fact, many ascalaphids are unable to fly at all Miller (1985) further assaulted the boundaries between the without a warm-up period of muscle shivering. Huge, owl-like two families: several features once thought to be the exclusive eyes are another characteristic feature of the adults. In members province of ascalaphid larvae have now been found in acan­ of one subfamily, the Ascalaphinae, each eye is split by a deep thaclisine Myrmeleontidae. sulcus into dorsal and ventral portions, with different spectral Taxonomic ambiguity notwithstanding, the family is nor­ sensitivities (Gogala 1967). Clearly, the large size of the eyes mally divided into three subfamilies, the Albardinae, Haplo­ facilitates visual prey-capture under low light conditions in all gleniinae (= Neuroptynginae or Ascaloptynginae of other ascalaphids, but it is not known why the eye is bisected in authors), and Ascalaphinae. Albardinae is an aberrant, mono­ j .~~ ascalaphine owlflies. typic taxon from Brazil, of uncertain phylogenetic position and Larvae of Ascalaphidae are just as peculiar, fascinating, and biological habits. Ascalaphinae is defined 'by the bisected com­ enigmatic as the adults. They are robust predators, not easily pound eye, but its members are additionally characterized by f-f separable from the antlions, characterized by powerful, sickle­ smaller size and sophisticated flight ability. Also, the constricted f shaped jaws and prominent occipital (posterior) lobes on the waist and yellow-and-black patterning of some ascalaphine head capsule. One or two rows of setose, finger-like extensions species make them strikingly wasp-like in overall appearance. occur along the lateral margins of the body, making the larva They occupy xeric, grassy, open habitats, and their larvae live quite flattened and stabilizing it in its encounters with large on the ground. In contrast, the entire-eyed Haplogleniinae are prey. Larval ascalaphids are formidable predators, capable of larger insects with a more fluttering and clumsy flight. They securing, paralysing, and consuming arthropods twice their generally live in mesic, forested habitats, where they cling to own size. They are not active foragers; instead, they 'sit and understorey vegetation. Their larvae tend to be more arboreal wait' for their prey, which they capture with lightning-fast than those of Ascalaphinae. Adults of both major subfamilies closure of their powerful, trap-like jaws. Some species, like spend much of the inactive daytime hours resting head down those of Ululodes, live on the ground, completely covering their on branches or vegetation with the wings folded and with the dorsal surfaces with sand grains, so that they are impossible to abdomen projecting upward at a sharp angle (Fig. 28.1). In see. Others, like Ascaloptynx, Ascalobyas, and Haploglenius, rely that position, they are extremely difficult to spot, because they upon their own pigmentation and specialized scale-like hairs look just like dead twigs. for crypsis, and may live either in leaf litter or on tree branches, Panama and Central America have a rich owlfly fauna, stems, or leaves high above the ground. although it is not so diverse as that of Australia and tropical The large, ovoid eggs of Ascalaphidae are deposited in several Asia. Four genera of Haplogleniinae are known from Central linear or spiralling rows near the tips of dead twigs or tapering America. Neohaploglenius is represented by two species, N. angu­ leaves, usually 1 m or less from the ground. A line of weakness latus (Gerstaecker) and N.j1avicornis (MacLachlan) [Fig. 28.4], around one pole permits hatching via a hinged cap (Henry which can be separated by the darker pterostigmal and apical 1977). New World taxa additionally deposit a number of infer­ regions of the foreWings of N. j1avicornis. Superficially, Neo­ tile, abortive eggs, usually in several rings around the twig just Iwploglenius is similar to Haploglenius and Ascalobyas, but is below the main egg mass. These abortive eggs, called repagula, separable from both of those genera by the narrower wings are produced in ovarioles that are dedicated to their manu­ with single anal vein and accentuated axillary area (Fig. 28.4), .tee· FJ ""'turm?U't

~ 1- rr­,~ :". THE NEUROPTEROID ORDERS 455

and from Ascalobyas by the longer antennae. The genus Asca­ ham's organ' at the base of the hindwing in others (Eltringham Iobyas is known from a single species, A. albistigma (Walker), 1926), or by eversible abdominal glands in some others (Tjeder distributed from Honduras to Panama. Similarly, just one 1954). However, details concerning the use of pheromones by species of Haploglenius has been reported from Central America: courting antlions are sparse, and copulation has actually been H. Iuteus (Walker), also distributed from Honduras to Panama. observed very rarely. Oviposition habits are also a mystery for The genus Amoea is known from two species in Central America, the most part, although deposition of eggs on sand or dirt seems A. Iatipennis (Navas) from Guatemala and A. vacuus (Ger­ to be common (Baba 1953; Stange and Miller 1985). staecker) from Honduras. At present these two species cannot Everyone is familiar with the pits formed in protected dusty be separated without further study of Navas' specimens of A. or sandy areas by doodle-bugs, the larvae of antlions. However, latipennis. very few people realize that only one subfamily of antlions, the The three Central American genera of the split-eyed Asca­ Myrmeleontinae, makes these pits; within that subfamily, pit­ laphinae have poorly defined generic limits and are all closely building is confined to the tribes Myrmeleontini and a few related. There is still a great deal of unresolved synonymy in Dendroleontini. Larvae of other groups live in a wide variety of the subfamily, and much work must be done before identi­ other situations, and are not adapted in any way for pit building. fications can be made with confidence. Part of the problem is Specializations of pit-builders include a narrow, dorsally dished that male genitalia are very similar in all New World asca­ head for throWing sand, stout digging setae at the tip of the laphine taxa, giving few clues to identification or phylogeny abdomen, and a peculiar and unique neck region that permits other than weak wing venational and colour characters. The powerful vertical motions of the head and jaws. They can move smallest Central American ascalaphines are in the genus Amer­ only backwards, and possess fusion of the metathoracic tibia opterus (Fig. 28.21), which includes A. consors (Gerstaecker), and tarsus to facilitate such locomotion. Pit construction A. mexicanus (Weele), and A. trivialis (Gerstaecker). The broad­ usually proceeds by backward spiralling movements of the winged, robust genus Cordulecerus is represented by four Central larva, accompanied by vigorous tossing of sand out of the American species, C. inquinatus Gerstaecker, C. mexicanus enlarging pit (Bongers and Koch 1981), although direct down­ Weele, C, praecellens (Gerstaecker), and C. subiratus (Walker). ward digging is an option in some taxa (Turner 1915). The There are four species of the genus UIulodes: U. mexicana (Mc­ insect finally takes up residence buried at the bottom of the pit Lachlan) [Fig. 28.1], U. macIeayanus (Guilding), U. opposita with only its jaws exposed. Arthropod prey which pass too close Banks, and U. tuberculata (Banks 1901). to the pit's rim fall down its steep, unstable slopes. The antlion encourages its fall by tossing sand at the struggling prey item, which is detected Dy means of tufts of setae on the thorax that Family MYRMELEONTIDAE (antlions) are very sensitive to substrate vibrations (Devetak 1985). The It probably comes as a surprise that Myrmeleontidae is the prey is then seized, poisoned, injected with digestive enzymes, largest family of Neuroptera, exceeding even the Chrysopidae and sucked dry. in species diversity. They have been well studied, but not always Antlion larvae that do not construct pits are quite different in a comprehensive manner; for example, the biology of the from those that do. They have a more generalized morphology, larvae of a handful of species has been analysed exhaustively, giving them an ascalaphid-like appearance; in fact, there are but almost nothing is known about the vast majority of taxa. no larval synapomorphies that clearly separate the two families. Although nearly everyone has heard of the pit-making 'doodle­ The head capsule is robust and square, with powerful, curved bug' larvae of antlions, adults of Myrmeleontidae are cryptic jaws, minute antennae, and seven stemmata on each ocular and unfamiliar even to many entomologists. Most antlion adults tubercle, while setose tubercles project from the margins of the superfically resemble damselflies (order Odonata, suborder Zy­ thorax and abdomen. Usually soil-surface or debris dwellers, goptera) in appearance and flying ability. They are medium­ these myrmeleontids may also be subterranean or arboreal. sized to large insects (35-120mm wingspans), with long, thin Most are cryptically pigmented; however, Brachynemurus abdomens; narrowly elongate, often beautifully patterned nebulosus (Olivier) in Florida is brilliantly pigmented with wings; and short antennae, thickened toward their tips or even orange, red, white, and black, apparently in mimicry of vicious mildly knobbed. Most are weak fliers and are active only at velvet ants (Brach 1978). Some species have been seen to dusk or night-time, but the more brightly coloured ones have pursue their prey actively, but the majority of non-pit-building diurnal habits. Adults of the majority of species are predatory; antlions must wait for their prey to come to them. All myr­ those with spurs on the legs are aerial predators, while spineless meleontid larvae can survive long periods of starvation, and forms feed on plantlice or even pollen (Stange 1970a). In the are consequently quite opportunistic and unpredictable in the manner of their close relatives, the Ascalaphidae (Fig. 28.1), timing in their life cycles. Depending upon the species, the they rest for long periods on vegetation, tWigs, or even rocks spherical cocoon is spun in sand, on rocks, in litter, on or under with the abdomen flexed dorsally (Stange 1970a). bark, or attached to the higher branches or leaves of shrubs Courtship and mating in myrmeleontids undoubtedly and trees. Usually, large quantities of material from the insect's involves communication by male pheromones, secreted by environment are incorporated in its walls. abdominal hairpencils in some taxa (New 1982), by 'Eltring­ The higher classification of the Myrmeleontidae is not well 456 28. HENRY, PENNY, AND ADAMS understood, and is much in need of attention. Various sub­ somewhat longer antennae than other genera. In the Amazon familial and tribal divisions have been proposed, but not con­ Region they were collected deep in very wet forests (Penny and sistently applied by antlion specialists. Here, we recognize four Arias 1980). There are four Central American species known, subfamilies, three of which are well represented in Central E. genini (Navas) from Veracruz, Mexico, and Costa Rica, B. America. longior Banks from Yucatan, Mexico, E. macer (Hagen) from '<" The subfamily Acanthaclisinae consists of large to very large Veracruz, Mexico, and E. triguttatus (Navas) from Honduras. insects, with robust, formidable, free-living larvae. The species Lastly, there are five species of Psammoleon known from Central Stiphroneura inclusus (Walker) from Vietnam has a hindwing America: P. banksi Esben-Petersen from Honduras, P. bistictus length of 75 mm, and the North American species Vella fallax (Hagen) from Yucatan, Mexico, P. debilis (Gerstaecker) from (Rambur) can measure as much as 115 mm across the out­ Panama, P. iniquus (Navas) from Honduras, and P. parallela stretched wings. This subfamily is represented in Central Banks from Honduras. America by the single genus Vella, with one Widespread species, One last subfamily, Stilbopteryginae, is found in Australia. V. fallax, along the Atlantic coastal areas from Mexico to Brazil. This subfamily was formerly included within the Ascalaphidae, The larva of this species is apparently a subsurface predator in largely because of the relatively long, knobbed antennae. coastal sand dunes, actively moving backward through the However, recent work on immature stilbopterygines indicates sand in search of prey. that they are specialized antlions. The subfamily Palparinae is a group of large insects with brightly coloured wings. They reach their greatest development in the drier areas of Africa, but a single genus, Dimares, is found in the Neotropics. One species of this genus, D. amoenus Navas, has been described from Central America, but with no specific LITERATURE CITED details as to distribution. Within this genus, males and females are highly dimorphic; males have completely transparent Adams, P. A. (1962). Taxonomy of Hawaiian Chrysopa (Neuroptera: wings, but females display an extensive pattern of dark pig­ Chrysopidae). Proc. Hawaiian Entomol. Soc., 1962: 221-3. mentation on both wings. Adams, P.A. (1967). A review of the Mesochrysinae and Notho­ Stange (l970a) has divided New World Myrmelontinae into chrysinae (Neuroptera: Chrysopidae). Bull. Mus. Compo Zool. Harv., four separate tribes, only three of which occur in Central 135: 215-38. Adams, P. A. (1970). A review of the New World Dilaridae. PostiIla, America. Myrmeleontini is represented here by two very closely 148: 1-30. related genera, Porrerus and Myrmeleon, which can be sep­ Adams, P.A. (1978). Zoogeography of New World Chrysopidae, a arated by the longitudinal vein in the costal area of the forewing progress report. Folia Entomol. 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