Systematic Entomology (1981) 6, 253--290

Systematics of the dobsonfly subfamily Corydalinae (Megaloptera :Corydalidae)

MICHAEL J. GLORIOSO* Department of Entomology, Ohio State University, Columbus, Ohio, U.S.A.

ABSTRACT. The genera of Corydalinae are redefined, and representative characters are figured for each genus. New character sources, such as mouthparts and internal female genitalia, are investigated, as well as traditional male genitalia and wings. Allohermes is synonymized with Protohermes, Doeringia with Platy- neuromus. Intergeneric relationships are hypothesized on the basis of a cladistic analysis. Acanthacorydalis and the New World genera form a monophyletic group, as do Protohermes and Neurhermes, and Neuromus and Neoneuromus. Chloroniella belongs in the Acanthacorydalis - New World lineage, but exact placement is uncertain. A phyletic sequence classification is proposed on the basis of the cladistic analysis.

Introduction American Covydulus cornutus are excellent angling bait, while dried larvae of Pvotoizevmes The Megaloptera, long considered among the grandis, referred to as 'magotaro mushi', were most primitive of holometabolous insects considered a remedy for infant emotional (Weele, 191 O), contains two families, Sialidae irritation in Japan (Kuwayama, 1962). and Corydalidae. The Corydalidae, easily Relationships within the Corydalinae are separated from the Sialidae by presence of poorly understood, and generic limits are ocelli, non-bilobed fourth tarsomeres, and poorly defined. In this study I redefine the large size, contains two subfamilies, Coryda- genera and postulate intergeneric relation- linae and Chauliodinae. The Corydalinae are ships based on shared derived character states, restricted to North and South America, South or synapotypies. In searching for sufficient Africa and Asia, while the Chauliodinae also characters to more confidently establish occur in Australia and New Zealand. relationships I also hope to generate enough Adult Corydalinae are among the largest characters and knowledge of character states and most bizarre appearing of living insects, to eventually resolve intrageneric relationships. males of Ac~ur~thucorydu1l.uand ('orydalu~ being noted for disproportionately large Tuxonomzc lz~story mandibles. The formidable appearance of Davis (I 903) divided the Sialidae into the Acanthacorydalis has led at least one collector Sialidinae and Corydalinae, and Weele (1909) to regard this insect as extremely dangerous divided the Corydalinae into two tribes, (McLachlan, 1899). Neuromini and Chauliodini. Some authors While not as spectacular, the larvae may be (Esben-Petersen, 1924; Barnard, 1931 ; of specialized importance. Those of the North Kimmins, 1948; Kuwayama, 1962) recog- * Editors' note: We record, with regret, the un- nized Neuromini, while Tillyard (19 18) and timely death of Ilr Michael J. Clorioso on 17 Octoher many subsequent authors called this group the 1980, and thank Professor C. A. Triplehorn and Dr 0. S. Flint for their help with proof reading. Corydalinae. Lestage (1927) noted the name should be based on reducing Correspondence: Professor Charles A. Tripleh~n, C'orydulus, Department of Entomology, Ohio State University, Neuromini to a junior synonym. 1735 Neil Avenue, Columbus, Ohio 43210, U.S.A. Some early authors (Tillyard, 1918;

0307-6970/81/0700-0253 $02.00 O 1981 Blackwell Scientific Publications 751 Lestage, 1927; Barnard, 193 1) considered the 2200 specimens of Corydalinae and Chaulio- Sialidae and Corydalidae distinctive enough to dinae borrowed from museum and private merit familial status, with Weele's tribes ele- collections. Individuals and institutions pro- vated to subfamilies, while others followed viding material for study were: P. C. Barnard, Weele's categorical rankings (Banks, 1940, British Museurn (Natural History), London, 1943; Kimmins, 1954). More recently (e.g. England (BMNH); D. H. Kavanaugh, Chandler, 1956; Flint, 1973; Evans, 1978) the California Academy of Sciences, San Sialidae and Corydalidae have been considered Francisco, California (CASC); G. Ekis, separate families. Carnegie Museum of Natural History, Weele (1910) was the first to revise the Pittsburgh, Pennsylvania (CMNH); J. E. H. Corydalinae, recognizing eight genera and Martin, Canadian National Collection of thirty-four species and menhoning possible Insects, Biosystematics Research Institute, relationships. Navas (1 9 15) proposed Ncur- Ottawa, Ontario (CNCC); L. L. Pechuman, hermes to replace Gray's Hermrs, homony- Cornell University, Ithaca, New York mous with a gastropod genus, and Esben- (CUNY); J. Kethley, Field Museum of Natural Petersen (1 924), Navhs (1 925) and Lestage History, Chicago, Illinois (FMNII); L. A. (1927), proposed additional genera. Lestage Stange, Florida State Collection of Arthro- (1927), in his catalogue of Indo-Chinese Cory- pods, Gainesville, Florida (FSCA); N. D. dalinae, included a list of world genera and Penny, Instituto Nacional de Pesquisas da postulated affinities based on wing venation. Amazonia, Manaus, Brazil (INPA); L. Since 1927 work on Corydalinae has been Dieckman, Institut fiir Pflanzenschutzfor- mainly species description, though Banks schung Kleinmachnow, Eberswalde, DDR (1940) included a key to Chinese genera, (IFPK); C. L. Hogue, Los Angeles County Kimmins (1948) reviewed the Protohc~rmesin Museum of Natural History, Los Angeles, the British Museum, and Penny (1977) listed California (LACM); M. M. Pearce, Museum the New World species. of Comparative Zoology, Ilarvard University, Cambridge, Massachusetts (MCZC); M. J. Glorioso, Columbus, Ohio (MJGC); J. With the exception of Corydalus c.orrzutu.s, Legrand, MusCum National dlHistoire the natural history of Corydalinae is poorly Naturelle, Paris, France (MNHN); C. A. known, though the larvae are probably all Triplehorn, Ohio State University, Entomo- lotic inhabitants. Larval lifespan varies from 1 logy, Columbus, Ohio (OSUC); N. W. Britt, to 5 years (Evans, 1978), this being correlated Columbus, Ohio (NWBR); P. A. Adams, with latitude in C.cornutus (Brown & Fullerton, California (PAAC); G. W. Byers, Fitzpatrick, 1978). Some species have been Snow Entomological Museum, University of observed occupying specific microhabitats Kansas, Lawrence, Kansas (SEMK); S. J. within a stream (Geijskes, personal communi- Merritt, Texas, A & M University, College cation), while label data indicate a correlation Station, Texas (TAMU); W. F. Barr, Univer- between species and stream type or altitude. sity of Idaho, Entomology, Moscow, Idaho Adults are crepuscular or nocturnal and (UIDC); H. P. Brown, University of readily collected at lights. Though some Oklahoma, Zoology, Norman, Oklahoma evidence exists indicating feeding on liquids (UOKL); 0. S. Flint, Jr, P. J. Spangler, United (Parfin, 1952), adult life is generally short, States National Museum of Natural History. ranging from 3 to 13 days for C.cornutus Entomology, Smithsonian Institution, (Davis, 1903; Parfin, 1952). Adult emergence Washington, D.C. (USNM); K. K. Gunther, of some species is seasonal in certain regions, Zoologisches Museum, Humboldt-Universitat, while little seasonality is observed in other Berlin, Germany (ZMHU). localities. Seasonality may be influenced by temperature or rainfall, but also occurs in Methods regions with no distinct seasons. Criteria for gerzera delimitution Material Mayr (1969) considers a genus to be a This study is based on examination of about monophyletic category containing a single Systemutlcs oj the dobsonfly subfam~lyCorydal~nae 255 species or group of species separated from However, the splitting of genera only on the other genera by a decided gap. I likewise basis of number of species should be conser- believe genera should be monophyletic, but vative to avoid proliferation of genera. in the sense of Hennig (1965, 1966), and, In this study several autapotypies involv- ideally, genera should be recognizable by dis- ing male genitalia define distinct lineages, continuities in character states. As the degree with no intergradation between lineages. As of difference between genera or other supra- these monophyletic units are also recognizable specific taxa is subjective, criteria for recog- on the basis of other characters, I propose nition of supraspecific categories should be that each be given generic status. explicitly stated to allow subsequent workers to judge the validity of such taxonomic deci- Phylogrnefzc and class~jccatorymethods sions (Ekis, 1977b). Before supraspecific classifications are My phylogenetic hypotheses and classifi- developed, worldwide knowledge of the cations are based on the results of a cladistic investigated taxon should be available (Erwin, analysis, the principles of which were outlined 1975). This requires investigation of inter- by Hennig (1 965, 1966). Phylogenetic classifi- specific, but not intraspecific variation cations, including phyletic sequencing, a (Herman, 1970), and an attempt should be means of deriving a classification from an made to examine all species of the larger asymmetrical branch of a cladogram without taxon before inferring generic limits. While superfluous supraspecific categories, have knowledge of all interspecific relationships been discussed by Wiley (1979b). Proponents is not essential, major monophyletic lineages of phyletic sequencing include Nelson (1972, based on synapotypies must be defined, as 1974), Cracraft (1974), Schuh (1976) and well as the relationships of each lineage to the Wheeler (1979a, b). others. After these relationships have been hypothesized discontinuities between lineages Dissect~ngmethods can be evaluated, and distant lineages (defined by autapotypies) can be given categorial The tip of the abdomen was removed, status, e.g. generic rank. cleared overnight in KOH or lactic acid, Stability and predictability of classifi- rinsed with water, and stored in a micro- cations should be considered when establish- vial with glycerin. Female internal structures ing generic limits. Where monophyletic were removed through a slit below the genital lineages coincide with preexisting genera, opening or were observed after inversion of classification should be conservative, i.e. the the abdominal tip. Mouthparts were relaxed preexisting classification should be retained. If with Barber's fluid (53 parts 95% ethanol, 49 the preexisting classification is incongruous parts water, 19 parts ethyl acetate, 7 parts with monophyletic lineages, paraphyletic and benzene). One maxilla and labial palp were polyphyletic genera should be converted into removed, cleared in lactic acid or KOH, rinsed monophyletic ones, as polyphyletic classifi- in water, and stored in microvials with cations are of limited predictive value, and glycerin. Examination of genitalia and mouth- paraphyletic taxa are less predictive than parts was in glycerin or lactic acid. monophyletic ones (Platnick, 1978). These discrepancies may be rectified by incorpora- General methods tion of excluded lineages in paraphyletic taxa or fragmentation into component mono- Size ranges were estimated by measuring phyletic units. the largest and smallest available individuals A consideration of some authors, e.g. of each genus, selected by eye. Length was Heyer (1974), is that genera should contain a measured from tip of clypeus to tip of convenient number of species. If I were to abdomen; wingspan was determined by consider this premise, then consolidation of summing width of the mesothorax and twice small genera would seem logical, with the the length of the forewing. Genitalic and corollary that splitting would be appropriate mouthpart dissections were made for at least where consolidation creates unwieldy genera. one male and female of each available species. Generic descriptions should be understood to include characters denoted in the subfamilial descriptions, and are based on specimens listed in Appendix A. As species concepts in most genera are poorly understood, necessi- tating revision before accurate names can be placed on many specimens, many specimens listed in the appendix are denoted as spp.; species I have confidently identified are listed separately. Lists of material only include localities and repositories, other data being summarized under natural history. Due to extreme numbers, Covydalus cornutus locali- ties are given by state only, Mexican Clutea records are listed as states, and ('.lzltru records from the United States are listed as counties. Geographic ranges of genera were determined from label data and reliable literature records. Drawings were made with a drawing tube on a Wild M5A stereoscopic microscope. Scale bars accompanying figures represent 1 mm unless otherwise indicated. 1:lG. 1. Corydalinae head, showing postocular ridge, spine, and plane.

Comparative morphology I4'IGS. 2 and 3. Gular region of Corydalinae: 2, Platyneuromus auritus; 3, Neurhermes maculipennis. Justifications for thorough morphological analyses in revisionary studies were given by Ekis (1977a). As knowledge of internal struc- and development of the lateral postocular ture of Corydalinae is limited to C'ouydu1u.r margins of the cranium. In Proto1zc~rrne.sand cornutus, this discussion is restricted to Neurhrvrnrs the head is robust, about as high integumental characteristics. Further, due to as wide, while in other genera it is noticeably lack of immatures, I treat only imaginal flattened. features. The postocular ridge (Fig. 1) extends Although fluid preserved material of most laterally behind the eyes to the occiput. Its Corydalinae is scarce, C.cornutus specimens dorsal extension curves anteriorly at the have always been readily available. Conse- posterior angle of the head, delimiting the quently both internal and external structures postocular plane. This plane is generally less have been described in considerable detail. rugosely sculptured than adjacent regions of Leidy (1848) discussed the digestive, repro- the head. In P1arj~neurornu.s(Fig. 61) the post- ductive and nervous systems, Hilton (1909) ocular plane occupies the postocular flange, treated the tracheal system, and Kelsey the explanate postocular cranial margin. The (1954, 1957) detailed the external morpho- postocular spine arises from the posterior logy and musculature of the head and thorax. angle of the head, forming a lateral acumina- The larval nervous system was treated by tion of the postocular ridge. This spine, as Krauss (1884), Hammar (1908) and Hilton well as the ridge itself, is barely noticeable in (191 1). Protoherrr~esand Neurhermes (Figs. 56 and 57), moderately developed in C'hlororlia and Head Neurornus (Figs. 58 and 62), and very con- spicuous in other Corydalinae. In some Typical features of the Corydallnae head Plutylleuromus the spine is fused with the are embodied in C'orydalus cortiutus as postocular flange. described by Kelsey (1954). Major variations The cranial disk, though rugose, is devoid involve the degree of flattening of the head of processes in most genera. However, the disk Systematzcs oj' the dobsonfly subjamily Corydalinae 257 of Acar2thacorydalis (Fig. 60) is provided with as long as the head plus prothorax, though in a pair of lateral spines. male Corydalus they may be longer than the The ocelli occur on nearly contiguous body. The antennifer is a ridged area at the prominences in most genera. The posterior postero-median margin of the transverse margins of the lateral ocelli are nearly in line anterior tentorial pit. The scape is cylindrical, with the posterior margins of the compound slightly longer than wide, and, together with eyes, occasionally being slightly anterior. The the shorter, thinner pedicel, parallels the median ocellus is generally transverse, its epistomal sulcus when extended laterally. posterior margin about at the anterior margin The pedicel and scape are rugosely sculp- of the lateral ocelli. In most Neurhermes and tured, while the antenna1 articles are covered some Protoherrnes the median ocellus is with small tubercles or short setae. Tubercles distinctly anterior to the lateral ocelli. are more common on proximal articles, distal The antennae consist of thirty-five to articles are moderately setose. All articlcs seventy-five articles, and are generally about are moderately setose in Protoherrnes and

FIGS. 4-1 1. Labrum and clypeal margin of Corydalinae: 4, Protohermes sp.; 5, Neurhermes maculi- pennis; 6, Acanthacorydalis orientalis; 7, Corydalus cornutus; 8, Neoneuromus sikkimensis; 9, Neu- romus testaceus; 10, Chloronia sp.; 11, Platyneuromus soror. 18 Neurhermes, with all setae very short. The Corydalus (Fig. 13). Membrane and two band- antenna is filiform to subserrate, anterior like subgaleal sclerites separate the galea from serrations being most apparent in Proto- the stipes, with the lower sclerite often being hermes and Neurhermes. divided by membrane. Microsculpture of the The clypeal margin may be entire, as in galeal base and stipal membrane consists of Protohermes and Neurhermes (Figs. 4 and S), diagonal series of acuminate scales. The galea or medially incised to varying degrees, ranging is densely covered with short and long setae, from the shallow notch in Chloroniu (Fig. many of the long setae in Neurhermes and 10) to the deep incision of Acanthacory- Protohevmes being flattened lanceolate. The dalis (Fig. 6). The clypeal margin of many apex, rounded in Protoiiermes and Neur- ('orj~dalusdisplays a pair of antero-lateral and hermes, acute in other genera, usually bears a median projections, with the incision between sensory peg with five to seven microsetae. The the median projections. In some C'orj~dalus sensory peg is poorly developed in Neurowrus the incision is obliterated by fusion of the and Neoneuromus, absent in Neurhermes. median projections (Fig. 7). The cylindrical five-segmented maxillary palp The triangular to broadly truncate or ovoid (four-segmented in Covydalus) arises from a labrum (Figs. 4-1 1) partly covers the man- distinct palpifer. The penultimate segment dibles, though in ('orydulus and Acantha- bears an apical crown of alternating longer corydalis it is slightly deflexed between the and shorter setae; basal segments usually have mandibles. It is glabrous dorsally, moderately similar crowns obscured by additional setae. setose ventrally, and bears two median pairs The terminal segment is conical (Fig. 12) or of long setae and several short, fine setae. In broadly rounded apically (Fig. 13), and has Neurornus and Neoneurornus (Figs. 8 and 9) one or two membranous areas densely covered the short setae are more abundant, being with microsetae. especially predominant in Nytwrornus. The labrum of Neuromus often has a deflexed anterior margin with a slight indentation. The mandibles are subfalciform, triangular in cross section, and vary in length from one- half to three-quarters the length of the head. In female Acanthacorydalis the mandible is as long as the head, while it is longer in male Acunthucoryytlulis and most male ('orydalus (Figs. 60 and 63). The inner margin of the mandible is entire basally, dentate distally, with one apical and three inner teeth. The dorsal and ventral surfaces are flattened, and exhibit weak longitudinal depressions. The cuticle is smooth ventrally near the condyle, rugose elsewhere. Mandibular articulation occurs near the anterior margin of the eye. The maxilla articulates at the lateral margin of the submentum, with the triangular cardo bearing several short setae on the outer margin. The subcylindrical stipes, one-and-a- half to twice as long as wide, bears several setae on its outer margin. These setae are normally inconspicuous, but are well deve- loped in Neurhermes and Protohermes (Fig. 12). The triangular lacinia is fused basally with the stipes, closely paralleling the stipes and galea distally. Lacinia vestiture consists of stout setae; three long apical setae are present in all but Neuromus, Acantlzacorydalis and Sy.~tematicso/'tllc dohsonjZy .~uh/LI~n~l.yCorydalinae 259

The labium, broadly lo~nedto the sub- SC mentum, consists of a nearly quadrate men- tum, a short prementum, a pair of three- or four-segmented palps, and a bilobate Iigula. - , 2 7 Sclerotization of the mentum is most ev~dent , . \ '\ basally; numerous short setae are present \ dorsally. The rectangular prementum IS ,A c.,> , ,/ M3*4 14 -- - weakly sclerotized, w~ththe lateral palpigers more heavily sclerotized. The membranous ligula is moderately setose; the labial palp is usually similar to the maxillary palp, although the terminal segment IS triangular in Tome ('orydulu c The submentum and gula are vested with -;\i4 microsetose shallow depressions. The posterior - i \_- - -- tentorial pits extend obliquely forward from - <- --- y - T-, the gular sutures, diverging anteriorly. They are linear in Protohemley and Neurhrrrrzc3~ (Fig. 3), arcuate in other genera (Fig. 2). --- -

The thorax is fairly homogeneous through- out the Corydalinae, and has been amply detailed by Kelsey (1954, 1957). The pro- sternum of some C'or.ydulus and Platy- I*'IGS. 14-1 7. Forewings of Corydalinae. Scale bar neuromus bears a pair of small antero-median represents 1 cm. 14, Chloronia mexicanu; IS, Cory- acuminations. dalus cornutus; 16, Platyneuromus soror; 17, The elliptical forewing is 3-4 times as long Acun thucorvdalis on'en tulis. as wide and a quarter to half again longer than the body; the hindwing is slightly wider and of sectorial veins varies within a species, and about four-fifths as long. Colour varies from some individuals may have additional veins nearly hyaline in L'hloroniu, Neuromus, many or vein bifurcations involving only one of a Protohermes, and some Neoneuromus to pair of wings. brownish or light grey with darker markings in The medial vein is nearly fused basally Corydalus, Platyneuromus, Acanthacorydalis with the radius, diverging approximately and Neoneuromus or to black with white one sixth of its length distally. and M3+4 maculations in Neurhermes and some Proto- separate slightly distal to the base of Rs. Vein hermes. may consist of two branches (C'hloronia, Wings are densely covered with greyish or Chloroniella, Platyneuromus and some Cory- brownish microtrichia and are fringed with dulus) (Figs. 14 and 16), two bifurcate longer macrotrichia. Fine black bristlelike branches (many Neurornus and some Neo- macrotrichia are present dorsally on radial neuromus) or four to nine branches (Proto- and medial veins and scattered in the anal hermc,s and many Neoneuromus). Vein M3+4 area. Several rows of short, stout macro- may be undivided (Chloroniella, Chloronia trichia occur on the costa. and C'ovydalus), bifurcate (Acanthucorydalis, The costal vein tapers to the apex of the Neuromus, Neoneuromus, Neurhermes and wing, while a fine vein continues around the Platyneuromus) or consist of up to four wing. The subcosta, closely paralleled by and branches (Protohermes). distally fused with R1, reaches the wing The cubitus diverges from the media margin slightly before the apex. The radial basally; Cu2 separates from Cul slightly past sector contains four to sixteen veins, the last the point of divergence of the media in the bifurcate in all Corydalinae except C'ory- forewing, basally in the hindwing. Cul has one dalus (Fig. 15) and Chloroniella. The number to six accessories, Cuz is unbranched. 260 Michael .I. Glorioso

Though venation in the anal area of fore veins between each sectorial branch. Chloronia and hindwing is similar, the anal area of the and Plut~~neurorriushave three crossveins hindwing in much larger, forming a small anal between the radial sector and MI, Nc,urornu.s fan. Vein 1A is normally two-branched; it is and Neurher~r~c~shave four, and other genera three-branched in Neurhrrme.~ and Proto- have three to eight. The basal r-m crossvein in hcrrnes (Figs. 18 and 19). Vein 2A is bifur- the hindwing is long and oblique, connecting cate, the upper branch approaching 1A and Rs with the base of M. occasionally appearing to be a branch of 1A There are two medial crossveins in Platy- due to a short crossvein. Vein 3A is often nc,uromus and Clzlororziu, three in Neur- sinuate and, due to a short crossvein, hcrrnc~s,Neurorrzus and many Neoneuromus, occasionally appears two-branched, with the while Corydalus, Acanthacorydalis and upper branch being the lower branch of 2A. Protohermes have as many as eight. There are Extra longitudinal veins may also be present. usually one to three crossveins between Costal crossveins are extremely variable in branches of M, rarely up to seven. Three number; there may be anywhere between crossveins are often present between M4 and eighteen and sixty veins, with numbers vary- Cul, with a fourth connecting M and Cu, ing between wings of an individual. Crossveins though Neurhermes and Protohermes have may be contiguous basally, especially in two; A canthacorydulis and, rarely, Corydulus Corydalus and Protohevmes, or in a reticulate have as many as eight crossveins. pattern (restricted to male Acunthucorydulis) There is one cubital crossvein, except for (Fig. 17). Acanthacorydalis, which may have two or Three crossveins connect R1 and Rs in three. The cubital accessories are linked by a Corydulus, Chloro~?iu,Chloroniellu and Platy- single crossvein in Neuvomus and Neonru- neurornus; other Corydalinae have at least rornus (Figs. 20 and 21), two to five in four crossveins. There are one to four cross- Acuntliac~orydulis. Often one crossvein occurs between Cu and lA, though extras are occasionally present. Anal crossveins are as described in the discussion of the anal area, though .A curl thacorydulis often has extra anal area crossveins.

Abdomen Non-genital segments in Corydalinae are similar to those of Chauliodinae, as described by Maki (1936) for Neochaullodes fovmo- .sunus. However, Plutyneuromu~, Chloron~a and some C'orydalus females have a sternal pouch on the sixth abdominal sternum (Fig. 43). This structure is membranous, semi- circular, and opens in the intersegmental membrane. It has previously been reported only for Chloronia (Weele, 1910; Flint, 1970), and its function remains unknown.

Male genitalia Male genitalia consist of elements from the ninth and tenth segments, though much controversy exists over the exact homologies of these structures. The terminology used

FIGS. 18-2 1. Forewings of Corydalinae: I 8, Neur- here is adapted from Acker (1 960). hermes maculipennis; 19, Protohermes davidi; 20, The ninth tergum is slightly more heavily Neuromus intimus; 2 1, Neoneuromus sp. sclerotized than preceding segments, and has Systematics of'the dobsonfly suhfurnily Corydalinae 261

of the ninth tergum, and may be unguiform or clavate. The clavate stylus (Acanthacory- dalis, Platyneurowrus, Chloronia, Corydalus) (Figs. 22, 25, 27 and 29) is densely setose, directed postero-medially, and terminates in a small chitinous claw, the claw being absent in many Corydalus. Unguiform styli are directed medially and may be glabrous or sparsely setose with a glabrous tip (Figs. 3 1, 33, 35 and 37). The gonostylus of Neur- hermes (Fig. 37) is directed postero-medially in the basal half, dorsally in the distal half. The apodeme of the ninth gonostylus usually parallels the lateral margin of the ninth tergum, though it is directed medially in Protohermes and N~urhermes(Figs. 35 and 37). The ninth sternite or genital valve is generally quadrate, though it is attenuate in Neoneuromus (Fig. 33), broadly and deeply incised medially in Protohermes (Fig. 35). Neurhermes displays a median projection (Fig. 37), sometimes with a median incision; Acanthacorydalis has postero-lateral lobes (Fig. 22); Uzloronia bears long setae on feebly to well developed lateral lobes (Fig. 27). The setae of Acanthaccrrydalis are short and stout while those of Cl~loroniaare fine. The tenth tergites (superior appendages of Weele, 1910; surgonopods of Crampton, 1918; dorsal arms of basimere of Snodgrass, 1957; anoprocesses of Tjeder, 1970) are FIGS. 22 and 23. Acanthacovydalis orientalis, male generally tubular or clavate, slightly sigmoid, genitalia: 22, ventral view; 23, dorsal view. and about the length of the ninth tergum. The tenth tergites of Chloronia are 2-3 times as an internal inflection basally which may be long as the ninth (Figs. 26 and 27), while arched (Figs. 30, 32, 34 and 36) or inverted those of some Neurorrzits are flattened (Figs. V-shaped (Figs. 23, 24, 26 and 28). In 30 and 31). Neurhermes exhibits a biramous Neurhermes the tergum is much wider than tergite with the dorsal arm shorter than the long, with a shallow postero-median incision ventral (Figs. 36 and 37); some Protohermes (Fig. 36), while it is nearly quadrate in other have a short ventral and a long dorsal arm, genera. though most are uniramous (Figs. 34 and 35). The ninth gonocoxites are fused with the The tenth sternite is represented by the ninth tergite; in Neoneuromus the dorsal fused gonocoxites (penis of Weele and incisions of the tergum (Fig. 32) may be Crampton, parameres of Tjeder, aedeagus of vestiges of the ancestral line of fusion between Chandler and others) and the gonostyli tergum and gonocoxite (Acker, 1960). In all (mammiliform processes of Weele, penis other Corydalinae fusion is complete. hooks of Crampton). The gonocoxites The ninth gonostyli (inferior appendages of normally form a bandlike sclerite, though a Weele, 1910; subgonopods of Crampton, postero-median projection is present in 1918; lower lobes of the cerci of Chandler, Neoneuromus (Fig. 39). The styli are sparsely 1956; ventral arms of the basimere of setose and usually digitiform. In most genera Snodgrass, 1957; catoprocesses of Tjeder, they are paired antero-medially; they are 1970) articulate at the postero-ventral margins located lateral to the median projection in Neoneuromus, while they are broadly fused sparsely to moderately setose. The area in many Neuromus (Fig. 31). In C'orytlalu.~ between the sternite and ovipositor is and C'hloronia the gonostyli are shorter and generally membranous, though two weak, papilliform (Figs. 40 and 4 1 ). setose sclerites are present in Neurhrrtne.~ A pair of genital papilli (Figs. 22, 37 and (Fig. 44). 38) (utriculi of Tjeder) may be present lateral The ninth tergum is moderately sclero- to the genital opening, and possibly represent tized and usually membranous dorsally. tenth sternal apophyses (Acker, 1960), or The tenth tergum is divided dorsally by the may be of ninth sternal origin. They may be anus, and is bilobed, nearly divided by the present in all members of a genus (Proto- round, setiferous cercus. The ovipositor of hermes, Ncurhermes, Acanthacorydulis), some Corydulus has been described by Mickoleit members of a genus (Platyneuromus, (1973), and is similar to that of other Neurornus), or unknown in any members Corydalinae, major variations being develop- (Corydulus, C'hloroniu, Nconeuromus). ment of the lateral sclerite and articulation of The cerci are located between the tenth the ninth gonostylus. tergites and ninth gonostyli, and are small, The lateral sclerite (Fig. 43), strengthening hemispherical, and setiferous. A sclerite over the ninth gonocoxite, is usually well sclero- the anus of Neoneuromus (Fig. 32) may repre- tized, though it is only weakly to moderately sent the eleventh tergum (Acker, 1960). sclerotized in Chloronia. The gonostylus is articulated with the posterior tip of the gono- coxite in Neurhermes and Yrotohermes (Fig. 44), while it is fused in other genera. The eighth sternite is quadrate or with The genital opening is in the membrane convex margins, moderately sclerotized, and at the base of the ovipositor, and opens into a membranous saclike (Fig. 45) or muscular tubular (Figs. 46 and 47) bursa copulatrix. The spermathecal duct is short and biramous T-shaped in genera with a saclike bursa, while in genera with a tubular bursa (Neurornus and Neor~euromus)it is long, twisting, and con- tinuous with the bursa. The paired, ovoid spermathecae are located at the end of the spermathecal duct, on either arm of the 'T-shaped duct, though only one sperma- theca is present in Neuromus. The common oviduct enters the saclike bursa near the opening of the spermathecal duct, while it enters in the anterior third of the bursa- spermathecal duct complex of Neurornus and Neoneuromus. A pair of glands located on the ovipositor immediately behind the genital opening may be homologous to the accessory glands of Sialidae as described by Heberdey (1 93 1). These glands are absent in Corydalus, Chloronia and Platyneuromus. while they are longer than the bursa-spermathecal duct complex in Protohermes and Neurhermes (Figs. 48 and 49) and one fourth to one half as long in Neuromus, Neoneuromus and Acanthacorydalis (Figs. 50-52). They are nearly linear in Neuromus and Neoneuromus, FIGS. 24 and 25. Platyneuromus soror, male geni- sigmoid in Protohermes, Neurhermes and talia: 24, dorsal view; 25, ventral view. Acanthacorydalis. Systemut~csof the dohsonjly subfum~lyCorydul~nue 263

The colleterial gland, opening near the life history, and ecological sources. However, centre of the ovipositor, is membranous lack of living specimens and life history data and nearly as long as the abdomen. This gland often precludes such thorough investigations, secretes a chalky substance used in coating the necessitating reliance on morphological eggs at oviposition. characters. Moreover, lack of suitably pre- served material may prevent thorough exami- nation of internal structures, restricting Cladistic analysis character investigation to integumental characteristics. As confidence in evolutionary analyses Such is the case in this study - little is increases with number and diversity of known about life histories of most Coryda- character sources investigated, it would be linae and lack of fluid-preserved material ideal to investigate all possible sources, includ- prevents comparisons of delicate internal ing internal and external morphological, structures. I have partially alleviated this physiological and biochemical, karyological, problem by thorough investigations of

FIGS. 26 and 27. C'hlovonia sp., male genitalia: 26, dorsal view; 27, ventral view. FIGS. 28 and 29. Corydalus comutus, male genitalia: 28, dorsal view; 29 ventral view IAHLE: 1. Character states.

Character Plesiotypic Apotypic

. ------1. Gular setation Conspicuously setose Microsetose 2. l'ostocular ridge Ahsent Present (Fig. I) 3. Postocular plane Absent Present (Fig. I) 4. Head shape Robust Ilattened 5. l'ostocular flange Absent Present (Fig. 6 1) 6. Postocular spine Feebly developed (Figs. Moderately to well developed 56 and 57) (I'igs. 58-63) 7. Posterior tentorial pits Linear (Fig. 3) Arcuate (I'ig. 2) 8. Cranial disk Without spines With spines (Fig. 60) 9. Antennae Feehly subserrate (I'igs. 1;iliform 56 and 57) 10. Clypeal margin I.:ntire (b'igs. 4 and 5) Incised medially (E'igs. 6-1 1) 1 1. Male mandibles Not enlarged Enlarged (I'igs. 60 and 63) 12. k'emale mandihles Three-fourths head length or Length of head shorter 13. Labrum shape Triangular (Figs. 4-7) Ovoid (E'igs. 8 and 9) 13'. Broadly truncate (l'igs. LO and 1 1) 14. Labral position Over mandibles Between mandibles 15. Labral anterior margin Sparsely setose I'imbriate (Fig. 9) 16. Maxilla shape Short and broad (Fig. 12) Relatively elongate (Fig. 13) 17. Lacinia apex With three elongate setae Without elongate setae (Fig. 12) (Fig. 13) Stipal setae ('onspicuous (Fig. 12) Inconspicuous (Fig. 13) Galeal setae blattened, lanceolate Hristlelike Galeal sensory peg Well developed (Figs. 12 and 13) Poorly developed Absent Maxillary palp Five-segmented (l'ig. 12) I'our-segmented (Fig. 13) Maxillary palp apex Conical (I'ig. 12) Broadly rounded (Fig. 13) Maxillary palp setae Long Short Maxillary palp sensory areas One at apex Two at apex Labial palp Four-segmented Three-segmented Labial palp sensory areas One at apex Two at apex Last branch of radial sector Bifurcate (Figs. 14, 15, 17-21) Not bifurcate (I'ig. 16) M,+2branches Two (1;igs. 14- 17) 1;our or more (Figs. 18-2 1) M,+, branches Two (Figs. 16-2 1) One (Figs. 14 and 15) 1A branches Two (Pigs. 14-17, 20 and 21) Three (Figs. 18 and 19) K ,-Ks crossveins Three (Figs. 14-16) Four or more (I'igs. 17-21) Medial crossveins Two (Ia'igs. 14, 16 and 18) Three or more (Figs. 15, 17 and 19-2 1) 33. M-Cu crossveins Three (Fig. 18) Four (I'igs. 14-17, 20 and 21) 33'. Six or more (Fig. 17) 34. C'ubital accessory crossveins Absent (1,igs. 14-16, 18-19) Present (Figs. 17, 20 and 21) 35. Costal crossveins Vertical or oblique Reticulate (k'ig. 17) 36. Male ninth sternum Hind margin not as in Fig. 35 Hind margin as in 1:ig. 35 37. Male ninth sternum More or less quadrate Attenuate (1;ig. 33) 38. Male ninth sternum Without median projection With median projection (Fig. 37) 39. Male ninth sternum Without postero-lateral lohes With postero-lateral lobes (Fig. 22) 40. Male ninth sternum Without setiferous lateral With setiferous lateral protuberances protuberances 41. Male ninth sternum With normal setae With short, stout setae 41'. With fine setae 42. Male ninth sternum Not sclerotized dorsally Sclerotized dorsally (Fig. 37) 43. Male ninth sternum Without internal ridges With internal ridges (Pigs. 31 and 33) 44. Male ninth sternum Not locking with tenth sternite Locking with tenth sternite 45. Male ninth tergum More or less quadrate Short, with broad median incision (Fig. 36) 46. Male ninth tergum With dorso-lateral incisions (Fig. 32) Without dorso-lateral incisions 47. Ninth tergal internal Without median fossa With median fossa (Figs. 30 inflection and 32) Systematics oj the dobsonjly subfamily Corydalinae 265

TABLE 1 (Continued)

Character I'le\lotyp~c Apotypic ------48. Ninth tergal internal Arched (I~igs.30, 32, 34 and 36) Inverted V-shaped (Figs. 23, inflection 24, 26 and 28) 49. Membrane between ninth Thin Thickened. bilobate and tenth sternites 49'. Thickened, regularly convoluted 50. Genital papillae I'resent (bigs. 22, 37 and 38) A hsen t 5 1. Tenth gonocoxites Without median projection With median projection (Fig. 39) 52. Tenth gonocoxites Antero-lateral corner without Antero-lateral corner with acute acute projection (Figs. 36, 39 projection (Figs. 40 and 41) and 42) 5 3. Tenth gonostylus Digitiform (Figs. 38, 39 and 42) Short, broad (I:ig. 31) 53'. Papilliform (Figs. 40 and 41) 54. Ninth gonostylus Unguiform (Figs. 31, 33, 35 ('lavate (Figs. 22, 2 5, 27 and 37) and 29) 55. Ninth gonostylus Relatively short Long, bent (Fig. 37) 56. Ninth gonostylus Sparsely setose Ilensely setose 57. Ninth gonostylus apodeme Parallels ninth tergum (Figs. 22, Ilirected rnedially (Figs. 35 25, 27, 29, 31 and 33) and 37) 58. Tenth tergites Short Long, thin (Figs. 26 and 27) 59. Tenth tergites Not as in I'igs. 36 and 37 Biramous as in Figs. 36 and 37 60. Eleventh tergum Present (Fig. 32) Absent 61. Lateral sclerite of ovipositor Well scleroti~ed Weakly sclerotized 62. (;onostylus Articulated with gonocoxite Fused with gonocoxite (I.ig. 44) (Fig. 43) 63. Sternal pouch Absent I'resent (Fig. 43) 64. Sclerites between eighth Absent (k'ig. 43) Present (Fig. 44) sternum and gonopore 65. Bursa copulatrix Not saclike (Figs. 46 and 47) Saclike (Fig. 45) 66. Number of spermathecae 'l'wo (Figs. 45 and 46) One (1,ig. 47) 67. Spermathecal duct Continuous with bursa (Figs. Separate from bursa, T-shaped 46 and 47) (Fig. 45) 68. Accessory glands Present Absent 69. Accessory glands Short Long 70. Accessory glands Sigrnoid (Figs. 48-50) Linear (E igs. 5 1 and 52)

available integumental sources, including polarities were determined by the outgroup previously overlooked or under-utilized method, whereby joint possession of a sources. character state by the ingroup (taxon under Sources investigated include mouthparts investigation) and the outgroup (sister group and internal female genitalia, which have or next highest taxon) is considered indicative never been incorporated into Corydalinae of inheritance from a primitive common evolutionary hypotheses, and male genitalia ancestor rather than independent derivation. and wing venation. While male genitalia and Polarity determinations for character states general appearance have been used by many not present in the outgroup were accom- authors, including Weele (19 1O), wing vena- plished by using other characters to establish tion has been used sparingly by most authors monophyletic units within the Corydalinae to except Lestage (1927). serve as ingroups and outgroups. For any of these characters to be useful we Assuming monophyly of the Corydalidae, must know the variation within each taxon I looked outgroup to the Chauliodinae, which and the polarity, or relative apotypy or must either be, or contain, the sister group of plesiotypy of each character state. Variation Corydalinae. The Sialidae, the probable sister was determined by examination of as many group of Corydalidae, were also examined. species of each genus as available, although Taxa examined include Chuuliodes pectini- lack of Chloroniella specimens necessitated cornis, C.rastricornis, Nrochauliodes simplex, reliance on literature references. Character Neochauliodes spp., Neohermes concolor, Neoilc~rmt~sspp., Nigronia fusciutu.r, N.srrri- cornis, Protochauliodrs cir~eruscc,rl.rand Siu1i.s spp. Wing venation and genitalia for additional Chauliodinae genera were analysed by reference to works by Esben-Petersen (1 924) and Barnard (193 1) (Plat~~chuuliodes,TacJnio- chauliodes), Kimmins (1938, 1954) (Anachau- liodes, Archichauliodes, C'tenoc~hauliodes), Munroe (1953) and Chandler (1956) (Uysmi- cohermes), and Ponomarenko (1967) (Creto- chaulus). Characters used in the cladogram and discussion are listed in Table 1, with references to illustrations in parentheses. Numbers in parentheses in the discussion and cladogram refer to apotypic character states.

Character phylogenj~ The presence of a postocular ridge (2) and postocular plane (3) is regarded as autapo- typic for the Corydalinae as neither structure is present in the Chauliodinae or Sialidae. The microsetose gular region (1) of Corydalinae is considered autapotypic; the gular region of Sialidae and Chauliodinae is conspicuously setose. Dorso-ventral compression of the head (4) within Corydalinae is apotypic as indicated by the robust condition in the primitive Neur- hermes and Protohemmer, the Chauliodinae and Sialidae. Expansion of the lateral margin of the cranium into a flange (5), unique to Plat~~neuromus,is considered autapotypic, as are the spines on the cranial disk of Acantha- corydalls (8). Development of the postocular spine varies from feeble expression in Neur- hermes and Protohermes to well developed (6) in other genera. Linear posterior tentorial pits, subserrate antennae, and an entire clypeal margin are primitively found in Neurhermes and Protohermes as well as in the FIGS. 30 and 31. Neuromus intimus, male genitalia: Chauliodinae; apotypic arcuate tentorial pits 30, dorsal view; 31, ventral view. (7), filiform antennae (9) and incised clypeal margin (10) indicate the other genera form a monophyletic group. primitive species, has normal length man- Enlarged male mandibles (1 1) and a dibles. A triangular labrurn is considered deflexed labrum (14) are convergent apo- plesiotypic as indicated by the identical typies in Acanthacorydalis and most Cory- condition in Chauliodinae. In Neuromus dalus, while female mandibles the length of and Neoneuromus the labrum is ovoid (1 3), the head (12) are restricted to Acantha- with a fimbriate anterior margin (15) auta- corydalls. Enlarged male mandibles in Cory- potypic for Neuromus. A broadly truncate dalus are not considered apotypic for the labrum (1 3') may have been independently genus as C.cephalotes, probably the most derived in Platyneuromus and Chloronia, or Systematics of'the dobsonfly subjamily Corydalinae 267 may have evolved in the common ancestor to The maxillary and labial palps have five the New World lineage, and has undergone and four segments respectively in most reversal of this character state in Corydalus. Corydalinae; the four-segmented maxillary The maxilla of Neurhrrmes and Proto- (2 1) and three-segmented labial (25) palps of hermes, as in the Chauliodinae and Sialidae, is Corydalus are autapotypic. Terminal palp relatively short and broad with conspicuous segments of most Corydalidae are conical, stipal setae and flattened galeal setae. The and the broadly rounded apex in the New more elongate maxilla (16) with incon- World genera indicates monophyly. Reduc- spicuous stipal setae (1 8) and bristlelike galeal tion of the setae on the last two palp segments setae (19) indicates the common ancestry of has occurred in all Corydalinae except Neur- the rest of the Corydalinae. The lacinia bears hermes; this may have occurred twice, once three elongate apical setae in most Cory- in Protohevmes and again in the ancestor dalidae; independent loss of these setae (17) to the other genera, or may have occurred has occurred in A canthacorydalis, Nc>uvomus several times. One sensory area is primitively and Corydalus. The well-developed sensory present at the tips of the palps as indicated peg at the tip of the galea has been reduced in by the homologous state in Chauliodinae; two the Neuromus-Neoneuromus lineage (20) areas (24 and 26) have evolved in the Proto- and lost in Neurl~errnes(20'). hermes-Neurhermes lineage and the New

FIGS. 32 and 33. Neoneuromus sp., male genitalia: 32, dorsal view; 33, ventral view. FIGS. 34 and 35. Protohermes costalis, male genitalis: 34, dorsal view; 35, ventral view. World lineage, with possible secondary loss of one maxillary palp sensory area in C'hloro~ria. Most Corydalinae wings exhibit a bifurcate last branch of the radial sector, while that of ('orydalus is unbranched (27). As presun~ably primitive Chauliodinae (Dysrnicokerrnc.~, Chandler, 1956; C'retoc,huulus, Ponomarenko, 1976) display the bifurcate condition, and C'orydalus is a highly derived genus, the non- bifurcate condition is considered apotypic. In most Corydalinae, Sialidae and pre- sumed primitive Chauliodinae M3+4 consists of two branches, while these have fused (29) in the ancestor of Corydalus and ('hloronia. The opposite trend is apparent in MI+* - again the primitive state is two branched, but the derived state, present in the Protohrvrrres Neurhermes and Neurornus-Neoneuromus lineages, consists of four or more branches (28). In Protohermes and Neurhermes 1A is three-branched (30), while the primitive two- branched state is present in all other Coryda- linae, most Chauliodinae, and Sialidae. As indicated by comparison with the Chauliodinae and Sialidae, crossvein addition has been a general trend. Three crossveins between R1 and Rs and two medial crossveins are primitive states, while four or more cross- veins between R1 and Rs (31) and three or more medial crossveins have evolved in the Protohermes-Neurhermes and Neuromus- Neoneuromus lineages and in Acunthucoyy- FIGS. 36 and 37. Neurhermes maculipennis, male dalis. Alternatively, the apparent plesiotypies genitalia: 36, dorsal view; 37, ventral view. in the New World genera may be secondarily derived. Medial crossvein number has also increased in C'orydalus. Three crossveins are Chloron~a (4 1'). Internal longitudinal ridges plesiotypically present between M and Cu in (43) are shared by Neuromus and Neo- Protohevmes and Neurhermes, while the other neuvomus, while these ridges are deep and genera have four (33), except for Acuntha- platelike in Neuromus (44), forming a coupl- corydalis, with six or more (33'). Crossveins ing mechanism with the tenth sternite. between cubital accessories have evolved in The male ninth tergum of both Chaulio- the Neuromus-Neoneuromus lineage and in dinae and Corydalinae is nearly quadrate, Acanthacorydalis, while a reticulate costal except for the autapotypically shortened crossvein pattern (35) is autapotypic for tergum of Neurhermes (45). If the dorso- Acanthucorydu1i.s. lateral slit of Neoneuromus represents A more or less quadrate male ninth remnants of the division between tergum and sternum is plesiotypic, as indicated by this ninth gonocoxite (Acker, 1960), then com- state in Chauliodinae and Sialidae. The broad, plete fusion (46) has occurred in all other deep medial incision of Pro tohermes (36), genera. attenuate shape of Nc~oneuromus (37), In the internal inflection at the anterior postero-median projection and sclerotized margin of the ninth tergum is primitively a dorsal surface of Neurhermes (38 and 42) are simple arch in Protohermes and Neurhermes, autapotypies, as are the short, stout setae of similar to that of Chauliodinae. The deeper Acanthucorydalis (41) and the fine setae of arch with a median fossa (47) is shared by Systematics of the dobsonfly subfamily Corydulinae 269

Neuromus and Neoneuromus, while an and elongate, digitiform styli of other Cory- inverted V-shaped inflection (48) indicates dalinae lineages. The short, broad gonostyli of the monophyly of Acanthacorydalis and the Neuromus are autapotypic. New World genera. As the ninth gonostylus of the Acantha- The membrane immediately behind the corydalis-New World lineage is clavate ninth sternum, primitively thin, is thickened (54) while those of the more primitive Proto- (49) in the New World genera. Two distinct hermes-Neurhermes and Neuromus-Neo- lobes are formed in Platyneuromus and neuromus lineages are unguate, the clavate Chloronia, and regular convolutions are state is regarded as apotypic. Correlated with present in Corydulus (49'). As more characters this, but also present in advanced Neuromus indicate a sister group relationship between species, is dense setation (56). The extremely Chloronia and Corydulus than between elongate, bent stylus (55) of Neurhermrs is Chloronia and Plat.ynruromus, the regularly autapotypic, while the apodemes of the ninth convoluted state is regarded as more derived. gonostylus are directed mesad (57) in Proto- As the genital papillae are present through- hrrmes and Nrurherm~s, functioning as out the primitive Protohermes-Neurhermes support for the membranous genital area. lineage and in Acunthacorydalis, the most The tenth tergites of Corydalinae are rela- plesiotypic of the Acanthacorydalis-New tively short and cylindrical; tergites at least World lineage, these are regarded as primitive. 5 times as long as wide (58) and biramous Presence of these papillae in the most primi- tergites (59) are autapotypic for C'hloronia tive Neurornus and Platyneuromus species is and Neurhermes respectively. The structure further evidence of their plesiotypy; they have interpreted as the eleventh tergite (Acker, been lost (50) in Neoneuromus, Chloronia 1960) is present only in Neoneurornus, with and most Nruromus and Platyneuromus. loss of this structure (60) occurring at least The tenth gonocoxites are generally with- three times (in Neuromus and the Proto- out median projections; presence of a broad hermes-Neurhermes and Acanthacorydalis- median projection (5 1) in Neoneuromus is New World lineages). inferred as autapotypic. Pointed antero- The weakly developed lateral sclerite of the lateral projections (52) and papilliform tenth ovipositor of Chloronia (61) is considered gonostyli (53') have evolved in the relatively autapotypic as this sclerite in other Coryda- derived Corydalus-Chloronia sister pair, as lidae and Sialidae is well developed. The indicated by the small anterior projections gonostylus is freely articulated with the tip

1:IGS. 38-42. Male tenth sternites: 38, Pvotohermes costalis, showing genital papillae; 39, Neoneu- romus sp.; 40, Chloronia sp.; 41, Corydalus cornutus; 42, Platynezrromus souor. 270 Michael J. (;lovioso

lateral rclerite

common oviduct

I'IGS. 43 and 44. Female abdomens: 43, Chloronia sp.; 44, Neurhermes maculipennis. FIGS. 45-47. Bursa and spermathecae: 45, Corydalus cornutus; 46, Neoneuromus sp.; 47, Neuromus in tim us. FIGS. 48-52. Female accessory glands: 48, Neurhermes maculipennis; 49, Protohermes sp.; 50, Acan- thacorydalis fruhstorferi; 5 1, Neuromus intimus; 52, Neoneuromus sp, Systematzcs of the dobsonfly subfamily Corydalinae 27 1 of the gonocoxite in Sialidae and many monophyletic, and only individual species Chauliodinae, while fusion of these structures within each taxon can give rise to new taxa, (62) indicates at least two lineages in Coryda- ancestor-descendant relations among supra- linae, with that lineage excluding Protohermes specific taxa are impossible (Wiley, 1979a). and Nrurhernles being monophyletic. Therefore I consider the genus level clado- Autapotypic setiferous sclerites between gram of Fig. 53 a logical hypothesis of the gonopore and eighth sternum (64) are Corydalinae phylogeny, based on seventy unique to Neurhermes, while the sternal characters investigated in eight non-monotypic pouch (63) is restricted to the New World genera. Although no ancestors are represented genera. Monophyly of the New World genera on the cladogram, an early dichotomy into is further indicated by lack of accessory two ancestral species is suggested, one of these glands (68), while glands much longer than leading to the Protohermes--Neurhermes those of any other Megalopteran (69) are lineage, the other eventually giving rise to all shared by Prorohrrmes and Neurhermes. As the other genera. A dichotomy of this species these glands are signloid in Acanthucoryilalis led to the Neuromus-Neoneuromus and the and the Protohermes-Neurhermes lineage, Acanthacorydalis-New World lineages, with the linear state (70) in the Neuromus--Nee- subsequent speciation events leading to the neuromus lineage is considered apotypic. ancestors of each genus. A separate saclike bursa copulatrix (65) has The probable position of C'hloroniella evolved independently in the Protohermes- (not shown in Fig. 53 or discussed previously) Neurhermes and Acanthacorydalis- New can best be resolved after examination of World lineages, as indicated by the plesiotypic actual specimens; characters deciphered from state of the Chauliodinae, Sialidae and the previous descriptions indicate inclusion in the Neuromus Neoneuromus lineage. A biramous Acanthacorydalis- New World lineage, but T-shaped spermathecal duct non-continuous conflicting character states indicate relation- with the bursa (67) is correlated with the sac- ship with either Acanthucorydalis, the most like bursa, while a single spermatheca (66) is plesiotypic member of this lineage, or Cory- autapotypic for Neuromus. dalus, the most apotypic. No discussion is given of size and colour A well-developed postocular spine (6) and characters as polarities remain uncertain. The four m-cu crossveins (33) are shared with all large size of Acai~thacorydalis is possibly genera except Protohevines and Neurhermes, apotypic, but, due to lack of knowledge of the male ninth tergal internal inflection (48) ecology and its influence on size, this remains indicates placement in the Acanthacorydalis uncertain. Luteous coloration occurs in some -New World lineage, an unbranched M3+4 members of all major lineages, and may be (29) is shared with Corydalus and Chloronia, due to plesiotypies or adaptive convergences. and a non-bifurcate last radial sector branch Likewise, dark coloration in the Protohermes (27) indicates Corydalus is the sister genus. Neurhermes lineage may be synapotypic, However, postero-lateral lobes of the male with the lighter colour of many Protohermes ninth sternite (40) indicate affinity with autapotypic for those species, or dark colora- Acan thacorydalis. tion may be independently derived in Neur- The actual relationship may be somewhere herrnes and several Protoherrnes species. between Acanthacorydalis and Platyneuromus, in which case Chloroniella should share at least some of the apotypies defining the monophyly of the New World lineage, such as Although species level cladograms do not a thickened membrane behind the male ninth rule out ancestor-descendant relations sternite (49), presence of a sternal pouch (63), among extant taxa, and therefore are not or absence of accessory glands (68) in females. necessarily representations of the true phylo- Presence of all these apotypies plus those geny, but rather, representations of several indicating the sister group relationship of possible phylogenetic trees (Platnick, 1977), Corydalus and Chloronia would indicate supraspecific cladograms can be interpreted common ancestry with Corydalus, especially as representing phylogeny. As each supra- if any other apparent autapotypies of Cory- specific taxon on the cladogram is presumed dalus are shared. P 1-0 N 1-11 Nrir Ncu ~\ca I'la Ch I Lor

FIG. 53. Cladogram showing relationships of Corydalinae genera. Pro = Protohermes, Nrh = Neur- hevmes, Nnr = Neoneuromus, Neu = Neuromus, Aca = Acanthacorydalis, Pla = Platyneuromus, Chl = Chloroniu, Cor = Corydalus. For key to characters, see Table 1.

Biogeographic considerations Neuromus~-Neoneuromus and Protohermes -Neurhermes lineages are Asian, while the As readily seen in Figs. 54 and 55, the Acanthacorydalis-New World lineage has distribution of Corydalinae is mainly tropical one genus in Asia, one in South Africa, and Asian and American, although C'hloroniella three in the New World. is restricted to South Africa, Covydalus Many Neoneuromus occupy Asian cornutus ranges to southern Canada, and montane regions, while its sister genus, Neuro- Protohermes grandis is present in Japan. The rnus, is apparently lowland and occurs on Syctc,matrcs of thc dob.conj7y suhfamlly ('orydallnae 273

=CO~ II~I~~IPI~BAca BNeu m~~~ FIG. 54. Distribution of Corydalinae. Abbreviations as in Fig. 53.

BChl CII mNrh mPro

FIG. 55. Distribution of Corydalinae. C11= Chloroniella; other abbreviations as in Fig. 53. some Malaysian and Indonesian islands. romus is restricted to Central America. Platy- Nrurherme.c and its sister group, Protoh~rmrs, tzeuromus probably diverged from an ancestral may have both montane and lowland species, South American stock after dispersal into, and occur on islands and the mainland. Acun- and subsequent vicariance in Central America. thac,orydallr has a similar distribution to Because of the diversity and presence of Neorzeurornus the most primitive species of Corydalus and The New World lineage is most diverse in P1ut)~nruromu.r in the Amazonian region, I tropical South America, although tJlatyneu- hypothesize this as the ancestral range of the 19 Lomrnon dnce\tor ot these genera, with subse- necessitat~ngsynonomy. As I have heen unable quent range extension into Central Amer~cd to satisfactorily establish the position of after d~vergenceof ('l~lorotz~uand C'orydulur ('hlorotrlrllu I list it at the end of the classifi- ('i~loronla inexrcu/zu, the northernmost cation under 'placetnent uncertain.' C'hlo/onzu species, appears to be h~ghly der~ved,ar does ('orydulut cornutmr and also C luteu, the northernmost C'ovydulus rpecles CORYDALINAE

Corydalinae Davis, 1903: 443 (type-genus Tuxonomic conclusion.\ Corydulur Latreille, 1802: 290). Tillyard, Before conversion of the cladogram into a 1918: 820. Lestage, 1927: 75, 93. Banks, phyletic sequence classification a suprageneric 1940: 179 (as Corydalini). category must be established for the three Neurorninae Weele, 1909: 250; 1910: 8 (as major lineages. Although this category is Neuromini) (ty pe-genus Neuronzus Rambur, between genus and subfamily, making tribal 1842: 441). Barnard, 193 1: 170. Kuway ama, status appear logical, I believe such a desig- 1962: 327. nation adds little to predictability of the classification. Tribal status should imply signi- Diagnosis. Quadrate head with postocular ficant difference between lineages. However, ridge, spine, and plane; microsetose gular the difference between the three major line- region; males with well developed ninth gono- ages of Corydalinae is no greater than that styli. between genera. Therefore I am assigning each Description. Size: length 15-70 mm ; wing- of these lineages the informal status of genus span 45--170 mm. Coloration: often luteous assemblage, this category being merely for or stramineous with piceous markings, sorne- taxonomic convenience and without nomen- times fuscous to piceous; wings nearly hyaline clatural significance. with darker markings, sometimes smoky or The classification I propose for Coryda- black with lighter maculations. Head: linae (Table 2) reflects the phylogeny of the quadrate, robust to flattened, slightly rugose subfamily with minimum nomenclatural with smooth dendriform patterns; gular region change. Two of the eleven genera recognized microsetose; posterior tentorial pits oblique, by Lestage, lloeringiu and Allohermes, do not linear or arcuate; postocular ridge and plane appear in this classification due to new present, postocular spine feebly to well deve- synonomies. As Dorrirlgiu appears to be a loped; ocelli centered between posterior rather derived member of the Plut.)~neuromus margins of eyes, hind margin of median lineage, exclusion from Pl~rtl~t7~~~4ro~rlu.rwould ocellus rarely anterior to lateral ocelli; require recognition of Pluryne~trornusas para- antenna near anterior of eye, with thirty to phyletic. Likewise Allohermes is an apparently eighty-five articles, usually filiform, about derived ineniber of the Protohcrrrre.r lineage, length of head plus prothorax; clypeal margin entire or incised medially; labrum triangular, ovoid, or broadly truncate, usually partly

TABLE 2. Classification of Corydalinae. covering mandibles; mandibles usually half to three-quarters length of head and with three Protohermes assemblage prominent inner teeth, rarely greatly enlarged Pro tohrrmes and without distinct teeth. Thorax: prothorax Neurhermes rectangulate or trapezoidal, slightly narrower Nelrromus assemblage than head, about length of meso- plus meta- Neurornus thorax; meso- and metathorax quadrate, ~\reonetrromus about equal size, slightly wider than hind margin of pronotum; legs about length of Corydalus assemblage Acan thacorydulis head plus prothorax, femur about length of Plutyneuromus tibia, tibal apex with two ventral spines, tarsi Chloroniu with five articles, first and fifth about equal Corydalus in length, articles two to four combined Chloronella - placement uncertain about length of first, tarsal claws feebly Systematics of the dobsonfZy subfamily Corydalinae 275 dilated basally. Wings: elongate elliptical, fore- 2 M3+4 bifurcate (1;ig. 16); postocular region wing half again length of body, hindwing four- explanate (Fig. 61); Central America Plutyneuromus fifths length of forewing, with small anal fan; - M3+4a single branch (Figs. 14 and 15); postocular Rs four- to sixteen-branched, two- to region not explanate ...... 3 nine-branched, M3+4 one- to four-branched, 3 Last branch of radial sector hranched (Pig. 14); Cul with one to six accessories, Cu2 one South and Central America ...... Chloroniu branch, 1A two or three branches, 2A two - Last branch of radial sector unbranched (Fig. branches, 3A one or more branches; three, 15) ...... 4 or four to fourteen R1-Rs crossveins, three to 4 Male ninth gonostylus clavate (Fig. 29); males eight Rs-M crossveins, basal one in hindwing usually with enlarged mandibles; North to South America ...... Corydalus long, oblique; two to eight medial crossveins - Male ninth gonostylus clawlike; South Africa with one to seven between branches of M; Chloroniella three to eight nl-cu's, one (rarely up to three) 5 Occipital disk with two prominent spines (Fig. cu's with zero, one, or two to five crossveins 60); males with enlarged mandihles; large, dark between accessories; costal crossveins eighteen species; Asia ...... Acunthacorydulis to sixty, vertical or oblique, rarely in reticu- - Occipital disk without spines; males never with late pattern. Males: ninth sternite and tergite enlarged mandibles ...... 6 usually quadrate; ninth gonostyli well deve- 6 Posterior tentorial pits arcuate (Fig. 2); post- loped, clavate or unguate, apodeme usually ocular tooth well developed; 1A two branched (F'igs. 20 and 21) ...... 7 paralleling margin of ninth tergum; tenth - Posterior tentorial pits linear (Fig. 3); postocular gonocoxites fused medially; tenth gonostyli tooth poorly developed; 1A three branched sparsely setose, usually digitiform. Females: (I?igs. 18 and 19) ...... 8 eighth sternum usually quadrate; ovipositor 7 Male ninth sternite attenuate (Fig. 33); female composed of ninth gonocoxites, reinforced with two spermathecae (Fig. 46); clypeal margin by usually well-sclerotized lateral sclerite; with few setae (Fig. 8); mostly large, darkspecies; Asia...... Neonruromus gonostylus fused or articulated with gono- - Male ninth sternite quadrate (Is'ig. 31); female coxite; genital opening near anterior of ovi- with one sperrnatheca (1;ig. 47); clypeal margin positor; sper~nathecaeovoid, usually paired, fimbriate (Fig. 9); moderate sized, pale species; rarely single; spermathecal duct usually about Asia...... Neurornus length of bursa; accessory glands i~nrnediately 8 Male tenth tergites always biramous, ventral arm behind genital opening when present; slightly longer than dorsal (Figs. 36 and 37); colleterial gland large, opening at midlength head and body black, pronoturn often orange, wings hlack with light maculations; galea without of ovipositor. sensory peg; Asia ...... Nrurhermrs Distribution: Temperate and tropical North - Male tenth tergites simple (I'igs. 34 and 35) or and South America, South Africa, temperate with dorsal arm much longer than ventral; body and tropical Asia (India to China and usually pale, wings hyaline to smoky, sometimes Indonesia, Japan). with lighter markings; galea with sensory peg; Asia...... Protvhrrmes Natural history: Larvae are aquatic and long-lived, while adults are often collected at lights, many being found in montane regions. Protohermes Weele (Figs. 4, 12, 19, 34, 35, Remarks: Due to great intraspecific varia- 49, 55 and 56) tion within this subfamily, some genera contain many nominal species, yet the total 1'rotohcrrnc.s Weele, 1907: 243 (type-species number of species in Corydalinae may be less Protohermes anlicus Weele, 1907: 243, not than fifty. Relatively little material of Asian llrvrnes anticus Walker, 1 853: 207 ; desig- genera is available, making generic revisions nation by Weele, 1910); 1909: 251, 1910: 35, difficult. Banks, 1908: 28; 1940: 179. Lestage, 1927: 99. Kimmins, 1948: 52. Key to genera of Corydalinae Allohermes Lestage, 1927: 100 (type species Protohc~rn~cstluvitli Weele, 1909: 254; original designation). Syn.n. 1 'l'hree crossveins between K, and Rs (1,'igs. 14-16)...... 2 Diagnosis. - lbour or more crossveins between K, and Rs L~nearposterior tentorial pit; (bigs. 17 21)...... S galeal sensory peg well developed; usually FIG. 56. Prototzevmes sp., head and prothorax. lighter than Ncurhermes; male ninth sternite short and broad; stipes with conspicuous broadly incised; male tenth tergite uniramous, setae; lacinia with three elongate apical setae; or with short ventral and long dorsal arm. galeal sensory peg well developed; maxillary Description. Size: length 20- 60 mm; wing- palp four-segmented, labial palp four-seg- span 60130 mm. Coloration: stramineous to mented; terminal palp segments conical, with fulvous, two piceous prothoracic vittae, often two sensory areas. Wings (Fig. 19): radial sector with head maculations, rarely entire body eight to eleven branches, last branch bifurcate; dark; wings stramineous, veins often darker, MI+, four to nine branched; M3+4 two to four wings occasionally smoky with lighter macu- branches; Cu, with two or three accessories; lations. Head (Fig. 56): robust; posterior 1A three branches; six to fourteen crossveins tentorial pit linear; postocular spine feebly between R1 and Rs, four or five (rarely three) developed; antennae feebly subserrate, about between Rs and M; two to five medial cross- length of head plus prothorax, of thirty-five veins, with one to three between branches to fifty articles; clypeal margin entire, labrum (rarely up to eight, with up to seven between triangular (Fig. 4); maxilla (Fig. 12) relatively branches); three to seven m-cu's, one cubital Systematzcs of the dohsonfly subfamily Corydallnae 277 crossvein, twenty-four to fifty-eight costal Diagnosis. Linear posterior tentorial pit; crossveins. Males: ninth sternite (Fig. 35) galeal sensory peg absent: dark colour, usually broadly and deeply incised medially; genital with orange prothorax; male tenth tergite papillae present; internal inflection of ninth biramous, lower arm slightly longer than tergum arched; ninth gonostyli unguate, upper. nearly glabrous, apodemes directed mesad, Description. Size: length 15-30 mm; wing- occasionally uniting; tenth tergites short, span 45-85mm. Coloration: black, pro- tubular with setal brush directed medially thorax usually orange, rarely black or with (Figs. 34 and 35), or flattened triangular, black maculations; wings black with white occasionally with long dorsal and short ventral maculations. Head (Fig. 57): robust; posterior arm; tenth gonostyli sparsely setose, long, tentorial pit linear (Fig. 3); postocular spine digitiform. Females: without sternal pouch; feebly developed; antennae feebly subserrate, gonostylus setose, articulated with gonocoxite; about length of head plus prothorax, of forty bursa saclike, separate from spermathecal duct; to fifty-five articles; clypeal margin entire, accessory glands sigmoid, longer than bursa labrum triangular (Fig. 5); maxilla relatively plus spermathecal duct (Fig. 49). short and broad; stipes with conspicuous Distribution: Northeastern India to setae; lacinia with three elongate apical setae; Indonesia and China, Japan (Fig. 55). galeal sensory peg absent; maxillary palp five- Natural history: Label data indicate that segmented, labial palp four-segmented; adult emergence occurs all the year round in terminal palp segments conical, with two sen- some localities, with some species seasonal sory areas; palp setae relatively long. Wings or present in montane regions. (Fig. 18): radial sector seven or eight Remarks: Although there are many branches, last bifurcate; four branches; nominal species, many names may require M3+4 two (rarely three) branches; Cul with synonymizing, reducing the number of valid two accessories; 1A three branches; five to species to around a dozen. eight crossveins between R1 and Rs, four Lestage's Allohermes is placed in the Proto- between Rs and M; three medial crossveins, hevmes lineage by the apotypic incised male with one between branches; one cubital ninth sternite. Lestage based his genus on the crossvein, twenty-five to forty-six costal complex venation of P.davidi, though other crossveins. Males: ninth sternite with promi- species, such as P.grandis, have similar vena- nent (Fig. 37) or short, notched median pro- tion. As there are no major genitalic jection, upper surface sclerotized; genital differences, and davidi may be a relatively papillae present; internal inflection of ninth derived species of the Protohevmes lineage, tergum arched, ninth tergite short, broadly separation of Allohevmes from Protohermes incised posteriorly (Fig. 36); ninth gonostyli may result in a paraphyletic group. There- elongate, setose, distal half narrower, directed fore I place Allohermes in synonymy with dorsally, apodemes directed mesad, occasion- Protohermes. ally uniting; tenth tergites biramous, dorsal arm slightly shorter than ventral (Figs. 36 and 37); tenth gonostyli sparsely setose, long, digitiform. Females: without sternal pouch; Neurhermes Navas (Figs. 3, 5, 18, 36, 37,44, gonostylus setose, articulated with gono- 48, 55 and 57) coxite (Fig. 44); setiferous sclerites present between eighth sternite and gonopore; bursa Neurhermes Navds, 1915: 39 1 (type-species saclike, separate from spermathecal duct; Iiermes macullpennzs Gray, 1832: 33 1; accessory glands sigmoid, longer than bursa original designation). Lestage, 1927: 96. plus spermathecal duct (Fig. 48). Hermes Gray, 1832: 33 1 (type-species Distribution: Northeastern India to Indo- Iferinrs maculipennls Gray, 1832: 331; by nesia (Fig. 55). monotypy). Nom. praeocc. (not Hermes Natural history: Label data indicate that Montfort, 1810: 398, in Molluscs). Walker, specimens were collected from late March to 1853: 20. Weele, 1906: 208; 1909: 251; May at altitudes of 2000-3500 ft. Weele 1910: 40. Banks, 1908: 29; 1940: 179. (19 10) mentions specimens collected between Okamoto, 1910: 258. October and February. FIG. 57. Neurhermes maculipennis, head and prothorax.

Remarks: Navris's Nrurherme~is a replace- margin firnbriate; male ninth sternite quad- ment name for Gray's Ifrrmrs, homonymous rate; female with one spermatheca. with a gastropod genus. Of the six nominal Description. Size: length 30-35 mm; species of Neuvhevrnes, four are based on wingspan 75- 100 mm. Coloration: luteous; colour pattern of females, and may be varia- antennae, tips of mandibles, four spots on tions of the other two. pronotum piceous; wings nearly hyaline, with piceous forewing veins. Head (Fig. 58); moderately flattened; posterior tentorial pit Neuromus Rambur (Figs. 9, 20, 30, 31, 47, arcuate; postocular spine moderately deve- 5 1, 54 and 58) loped; antenna filiform, about length of head plus prothorax, of forty-five to sixty-five Neuvomus Rambur, 1842: 44 1 (type-species articles; clypeal margin incised medially, Neuvomus testaceus Rambur, 1842: 442; labrum ovoid, anterior margin deflexed and designation by Weele, 1907: 235). Brauer, fimbriate (Fig. 9); maxilla relatively long and 1868: 400. McLachlan, 1869: 36. Davis, narrow; stipes with indistinct setae; lacinia 1903: 465. Weele, 1906: 208; 1907: 235; without elongate apical setae; galeal sensory 1910: 27. Banks, 1908: 29; 1940: 179. peg moderately developed ; maxillary palp Lestage, 1927: 96. five-segmented, labial palp four-segmented; terminal palp segments conical, with one Diagnosis. Smaller and paler than most sensory area. Wings (Fig. 20): radial sector Neoneuvomus; no spines on cranial disk; labral eight to ten branches, last bifurcate; Sv~temurrc.coj tkc dohtotrjly ruh/utnllj~Corj~dulznue 279

FIG. 58. Nrurornus restuceus, head and prothorax.

two or three branches; M3+4 two branches; 30 and 31); tenth gonostyli setose, broadly Cu, with two or three accessories; 1A two fused to gonocoxite, often fused medially. branches; four or five crossveins between Females: without sternal pouch; gonostylus R1 and Rs, four between Rs and M and fused with gonocoxite; bursa tubular, fused between M and Cu; three medial crossveins, with spermathecal duct; spermathecal duct with one or two between branches; one short (Fig. 47) or extremely long, coiled; cubital crossvein, with one crossvein between one spermatheca; accessory glands linear, accessories, twenty-eight to thirty-five costal half length of bursa (Fig. 5 1 ). crossveins. Males: ninth sternite (Fig. 3 1 ) Distribution: Northeastern India to nearly quadrate, slightly incised medially, Indonesia (Fig. 54). narrower than eighth sternite, with two Natural history: Label data indicate that longitudinal, flangelike dorsal ridges; genital specimens were collected in September at papillae present or absent; internal inflec- 450 ft. tion of ninth tergum arched, with median Remarks: Neuvomus contains three fossa; ninth gonostyli unguate, sparsely to nominal and at least one undescribed species. moderately setose; tenth tergites cylindrical, Two of the nominal species are probably feebly sigmoid, or flattened triangular (Figs. synonymous. PIC:. 59. Nroneuromus sp., head and prothorax.

Neoneuromus Weele (Figs. 8, 21, 32, 33, 39, and narrow; stipes with inconspicuous setae; 46, 52, 54 and 59) lacinia with three elongate apical setae; galeal sensory peg moderately developed; maxillary Neoneuromus Weele, 1909: 252 (type-species palp five-segmented, labial palp four- Neuromus fenestrulis McLachlan, 1869: 42; segmented; terminal palp segments conical, designation by Weele, 1910); 1910: 24. with one sensory area. Wings (Fig. 2 1): radial Lestage, 1927: 96. sector ten to thir&en (usually eleven) branches, last bifurcate; three to eight Diagnosis. Usually larger and darker than branches; M3+4 two branches; Cul with two Neuromus; no spines on cranial disk; labral to four accessories; 1A two branches; four to margin sparsely setose; male ninth sternite seven crossveins between R1 and Rs, four attenuate; female with two spermathecae. (rarely five or six) between Ks and M; three Description. Size: length 35-50 mm; wing- (rarely four) medial crossveins, with one to span 80--140mm. Coloration: ferruginous to three between branches; four m-cu's; one fuscous, rarely luteous; often with fuscous or cubital crossvein, with one between acces- piceous vittae on head and prothorax; wings sories, thirty to fifty costal crossveins. Males: fulvous to fuscous, usually with darker ninth sternite (Fig. 33) attenuate, extending tessellations. Head (Fig. 59): moderately to tip of tenth tergites, slightly incised medi- flattened; posterior tentorial pit arcuate; post- ally, with two longitudinal ridges internally; ocular spine well developed; antenna filiform, internal inflection of ninth tergum arched, about length of head plus prothorax, of forty- with median fossa; posterior margin of tergum five to sixty-five articles; clypeal margin incised dorsolaterally incised; ninth gonostylus medially, labrum ovoid, anterior margin unguate, sparsely setose; tenth tergites clavate, sparsely setose (Fig. 8); maxilla relatively long feebly sigmoid (Figs. 32 and 33); tenth gono- Systernutics of the dohsonJ1y suhfumlly Cor.ydalinue 281 coxites with postero-median projection (Fig. 39); tenth gonostyli lateral to projection, sparsely setose, digitiform; eleventh tergite present over anus. Females: without sternal pouch; gonostylus fused to gonocoxite; bursa tubular, continuous with spermathecal duct (Fig. 46); accessory glands linear, half length of bursa plus spermathecal duct (Fig. 52). Distribution: Northeastern India to Malay Peninsula and China (Fig. 54). Natural history: Label data indicate that most specimens were collected in June or July, one at 4400 ft. Banks (1940) indicates collec- tion in July and August at altitudes ranging from 2000 to 8000 ft. Remarks: Only eight nominal species have been described, and the number of actual species may only be two or three.

Acanthacorydalis Weele (Figs. 6, 17, 22, 23, 50, 54 and 60)

Acanthacorydalis Weele, 1907: 228 (type- species Corydalus asiatica Wood-Mason, 1884: 1 10; present designation); 1909: 250; 1910: 22. Banks, 1908: 29; 1940: 179. Lestage, 1927: 95.

Diagnosis. Large size; spines on cranial disk; male mandibles enlarged. Description. Size: length 60-80 mm; wing- span 140- 175 mm. Coloration: piceous, often with testaceous to ferruginous markings on head, thorax, and abdomen; wings smoky, with darker veins and tesselations. Head (Fig. 60): moderately flattened; posterior tentorial FIG. 60. Acan thacorydalis orien talis, head and pro- pit arcuate; postocular spine well developed; thorax. one pair of spines on cranial disk; antenna filiform, about length of head plus prothorax, of seventy to eighty-five articles; clypeal margin accessories; 1A two branches; usually six deeply incised medially, labrum triangular crossveins between R1 and Rs, four to eight (Fig. 6), deflexed between mandibles; male between Rs and M and between MI+,and mandibles up to length of body; female man- M3+4; three to five crossveins between dibles length of head; maxilla relatively long branches of media; six to eight m-cu's; one to and narrow; stipes with indistinct setae; three cubital crossveins, with two to five lacinia without elongate apical setae; galeal between accessories; twenty-five to forty- sensory peg well developed; maxillary palp five costal crossveins, median ones forming five-segmented, labial palp four-segmented; reticulate pattern in males. Males: ninth ster- terminal palp segments conical, with one nite ovoid, with lateral lobes (Fig. 22), setae sensory area. Wings (Fig. 17): radial sector short, stout ; genital papillae present; internal eight to eleven branches, last bifurcate; MI+:! inflection of ninth tergite inverted V-shaped; and M3+, bifurcate (M3+4 of hindwing rarely ninth gonostyli clavate, densely setose, with with three branches); Cul with two to four terminal chitinous claw; tenth tergites clavate or tubular, weakly sigmoid (Figs. 32 and 33); Iloc~ringiu Navis, 1925: 207 (1 y pe-species tenth gonostyli sparsely setose, long, digiti- Iloeringia christcl Navis, 1925: 209; by form. Females: without sternal pouch; gono- monotypy). Lestage, 1927: 94. Syn.n. stylus fused with gonocoxite; bursa saclike, separate from spermathecal duct; accessory Diagnosis. Presence of postocular flange, gland sigmoid, half length of bursa plus Central American distribution. spermathecal duct (Fig. 50). Description. Size: length 20-50 nim; wing- Distribution: Northeastern India to span 50- 1 10mm. Coloration: luteous to Vietnam and China (Fig. 54). fulvous; pronotum with fuscous to piceous Natural history: Label data indicates that vittae, vittae often on head; wings fulvous specimens were collected from late March to with darker tessellations. Head (Fig. 61): May at 1800-3500 ft. broad, flattened; posterior tentorial pit Remarks: Weele (1907: 230) included in arcuate (Fig. 2); postocular flange at least Acunthacorydalis the species Corydalus asia- width of eye; postocular spine usually promi- ticu Wood-Mason and C.uricntalis McLachlan. nent, occasionally fused with postocular I designate C.asiaticu, the first described flange; antenna filiform, about length of head species, as the type of the genus. Only three plus prothorax, of forty to fifty-five articles; other nominal species have been described, clypeal margin feebly incised medially, labrum and there may be only two or three valid broadly truncate, rounded at corners (Fig. species. 11); maxilla relatively long and narrow; stipes with indistinct setae; lacinia with three apical elongate apical setae; galeal Platyneurornus Weele (Figs. 2, 1 1, 16, 24, 25, sensory peg well developed; maxillary palp 42,54 and 61) five-segmented, labial palp four-segmented; Platyneuromus Weele, 1909; 252 (type-species terminal palp segments broadly rounded, with Corydalus soror Hagen, 1861 : 193; by mono- two sensory areas. Wings (Fig. 16): radial typy); 1910: 23. Lestage, 1927: 94. sector eight to ten (usually ten) branches,

FIG. 61. Platyneuromus soror, head and prothorax. Systematics ojtkr dohsonj7y suhjurnily Corydalinae 283 last bifurcate; and M3+4 each two branches; Cul with four or five accessories; 1A two branches; three crossveins between R1 and Rs and between Rs and M; two medial crossveins, four m-cu's; one cubital crossvein, eighteen to thirty-five costal crossveins; first R1 cell 1-1s times length of third. Males: ninth sternite quadrate (Fig. 25), membrane behind sternite thickened, bilobate; genital papillae present or absent; internal inflection of ninth tergum inverted V-shaped; ninth gonostyli clavate or nearly unguiform, with terminal chitinous claw; tenth tergites clavate, sometimes with crenulate inner margin (Figs. 24 and 25); tenth gonocoxites broad (Fig. 42); gonostyli long, digitiform, sparsely setose. Females: with sternal pouch; gono- stylus fused with gonocoxite; bursa saclike, separate from spermathecal duct; accessory glands absent. Distribution: Central America (Fig. 54). Natural history: Species in Panama are highly seasonal (Wolda, personal communi- cation) and label data from Mexican specimens indicate adult emergence from May to July. The larvae may inhabit slow-moving rivers (Adams, personal communication). Remarks: Navas's Doeringia ckristel appears to be a highly derived member of the Platyneuromus lineage, having male genitalia similar to those of P.soror, and an extremely PIG. 62. Clzloronia sp., head and prothorax developed postocular flange. To avoid recog- nition of Platyneurornus as paraphyletic I place Doeringia in synonomy with Platy- Description. Size: length 20-40 mm; wing- nc.uromus. Besides these two species, Platy- span 50-90 mm. Coloration: luteous, with neuromus contains one additional nominal four piceous markings on pronotum, occasion- species and one variety, and at least one ally with other markings on head and meso- undescribed species. thorax; wings nearly hyaline, luteous or stramineous, often with darkened veins and maculations. Head (Fig. 62): moderately flattened; posterior tentorial pit arcuate; Chloronia Banks (Figs. 10, 14, 26, 27,40,43, postocular spine moderately developed; 55 and 62) antenna filiform, of thirty-five to sixty articles; clypeal margin feebly incised Ckloronia Banks, 1908: 30 (type-species medially, labrum broadly truncate or feebly Hermes corriplens Walker, 1860: 180, desig- pointed anteriorly (Fig. 10); maxilla relatively nation by Weele, 1909: 252); 1943: 60. long and narrow; stipes with indistinct setae; Weele 1909: 252; 1910: 30. Lestage, 1927: lacinia with three elongate apical setae; galeal 94. sensory peg well developed; maxillary palp five-segmented, labial palp four-segmented; Diagnosis. Usually smaller and paler than terminal palp segments broadly rounded, Corydalus; third R1 cell longer than first; labial palp with two sensory areas, maxillary last branch Rs forked; South and Central with one. Wings (Fig. 14): radial sector seven American distribution. to nine (usually eight) branches, last bifurcate; two branches, M3+4one; Cul with one to four (usually three) accessories; IA two branches; three crossveins between R1and Rs and between Rs and M; two medial cross- veins; four m-cu's; one (rarely two) Cu cross- veins, twenty-two to thirty-six costal cross- veins; third R1 cell at least as long as first. Males: ninth sternite more or less quadrate (Fig. 27), with long, fine setae, often on lateral projections, membrane behind sternite thickened, bilobate; internal inflection of ninth tergum inverted V-shaped; ninth gono- styli clavate, with terminal chitinous claw; tenth tergites elongate tubular, 3-8 times longer than wide (Figs. 26 and 27); tenth gonostyli sparsely setose, papilliform (Fig. 40). Females: with sternal pouch (Fig. 43); lateral sclerite of ovipositor weakly sclero- tized; gonostylus fused with gonocoxite; bursa saclike, separate from spermathecal duct; accessory glands absent. Distribution: Central and South America (Fig. 55). Natural history: Label data and field work (Wolda, personal communication) indicate a high degree of seasonality in at least some tropical areas. Mexican specimens were collected from May to August, Panamanian specimens from May to June, and South American specimens from December to January. Remarks: Banks (1 908) originally pro- posed Cl~loroniaas a subgenus for the New World species then placed in Neurnmus. Eleven nominal species have been described.

Corydalus Latreille (Figs. 7, 13, 15, 28, 29, 45, 54 and 63)

Corydalus Latreille, 1802: 290 (type-species Hemerohzus cornutus Linnaeus, 1758: 55 1; by monotypy). Weele, 1909: 251; 1910: 9. Lestage, 1927: 94. Comstock, 1940: 287. Banks, 1943: 60. Chandler, 1956: 232. Gurney & Parfin, 1959: 976. Evans, 1978: 138. Corydal~s Latreille, 1804: 44. 'Lapsus calami. ' Palisot, 182 1: P1. 1, Fig. 1. Olivier, 1825: 59. Burmeister, 1839: 950. Rambur, 1842: 534. Hagen, 1861: 192. McLachlan, 1869: 36. Davis, 1903: 470. Banks, 1908: 29. Needham, 1918: 935. Townsend, 1935: 27. Imms, 1957: 492. FIG. 63. Corydalus cornutus, habitus. Diagnosis. Three- and four-segmented labial luted riffles of small streams. Adults of some and maxillary palps respectively; last branch species may be collected nearly all the year of radial sector not bifurcate; first R1 cell round in tropical areas, while others emerge much longer than third; males usually with in spring and summer. enlarged mandibles; New World distribution. Remarks: Much intraspecific variation is Description. Size: length 20--60 rnm; evident in C'o~ydalus;consequently species wingspan 70- 150 mm. Coloration: fuscous limits are poorly defined. Of the thirty-five to fulvous, occasionally with lighter head and nominal species and subspecies, only about pronotum markings; wings usually fuscous a dozen appear valid (Glorioso, in prep.). with darker markings, often with white dots in forewing cells. Head: moderately flattened; posterior tentorial pit arcuate; postocular Chloroniella Esben-Petersen spine prominent; antenna filiform and length of head plus prothorax in female, usually ('hloronlc~lla Esben-Petersen, 1924: 15 1 (type- stouter, filiform to subserrate or denticulate species C'hloror~~~~IluperingucJvl Esben- in male, may be as long as entire body; clypeal Petersen, 1924: 152; by rnonotypy). Lestage, margin feebly to moderately incised medially, 1927: 93. Barnard, 1931: 171. labrum triangular (Fig. 7), deflexed between mandibles; male mandibles usually without Diagnosis. Last branch of radial sector not conspicuous teeth, may be length of body bifurcate; ninth gonostylus unguate; South (Fig. 63); maxilla relatively long and narrow African distribution. (Fig. 13); stipes with indistinct setae; galeal Description. Size: forewing 33 mm. Colora- sensory peg well developed; maxillary tion: luteous to fulvous, with piceous dorso- palp four-segmented, labial palp three- lateral head and pronotal vittae; wings hyaline segmented; terminal palp segments broadly with fuscous veins. Head: moderately rounded, with two sensory areas. Wings flattened, postocular spine present; male (Fig. 15): radial sector six to fourteen antenna serrate, three-quarters length of branches, last not bifurcate; two to five forewing; labrum triangular. Wings: radial (usually three) branches, M3+4 one; Cul with sector five branches, last not bifurcate; three to six accessory veins; IA two branches; two branches, M3+4 one; Cu, with one acces- three crossveins between R and Rs, three to sory; 1A two branches; three crossveins six between Rs and M; three to seven medial between R1 and Rs, four between Rs and crossveins, one to three between M1 and MZ; M; two medial crossveins, four rn-cu's; one four to eight rn-cu's; one (rarely two) cubital cubital crossvein, twenty-two to twenty-six crossveins, thirty to sixty costal crossveins; costal crossveins. Males: ninth sternite first R1 cell 1%--2% times length of third. moderately incised medially, with lateral Males: ninth sternite quadrate (Fig. 29), rarely lobes; internal inflection of ninth tergum with median projection, membrane behind inverted V-shaped; ninth gonostyli sternite thickened, regularly convoluted; unguate; tenth tergites obtusely conical, internal inflection of ninth tergum inverted convex dorsally, slightly concave ventrally. V-shaped; ninth gonostyli clavate, rarely with Distribution: South Africa (Stellenbosch; terminal chitinous claw; tenth tergites Wellington) (Fig. 55). cylindrical or clavate, often uncinate or Natural history: Adults and one pupa dilated basally (Figs. 26 and 27); tenth gono- were collected in mountains in November, one styli sparsely setose, papilliform (Fig. 41). specimen at 3000 ft. Larvae were collected in Female: sternal pouch present or absent, March. rarely feebly developed; gonostylus fused with Remarks: I have been unable to obtain gonocoxite; bursa saclike, separate from specimens of the only known species due to spermathecal duct (Fig. 45); accessory glands its scarcity in collections, and base this absent. description on those of Esben-Petersen and Distribution: North, Central and South Barnard. America (Fig. 54). Barnard states that the ninth gonostyli Natural history: Larvae of some species, arise from the tenth tergites, but without such as C.cornutus, are common in unpol- examination of a specimen I cannot ascertain whether this is true or is an artifact of preser- Brauer, F. (1 868) Verdeichniss der his jet/; bekannten vation. If Barnard examined dried material Neuropteren im Sinne Linne's. Vtrrhandlun~c~n without clearing the genitalia, the ninth gono- der Zoologisch-Botanischen Gesellschaf't in Wien, 18, 359-416. styli may appear distorted and fused with the Brown, A.V. & l:itzpatrick, L.C. (1978) Life history tenth tergites. and population energetics of the dobsontly, Corydalus cornutus. Ecology, 59, 1091-1 108. tlurmeister, H. (1839) Ifandhuch der Bntomologie, Vol. 2. Riemer, Berlin. Chandler, H.P. (1956) Megaloptera. Aquatic Insects of California (ed. by R. L. Usinger), pp. 229- Acknowledgments 233. University of California Press, Berkeley. Comstock, J.ti. (1 940) An Introduction to Bntomo- I thank Dr Ginter Ekis, Steven W. Nichols, logy, 9th edn. Comstock I'ubl., Ithaca, New Dr Charles A. Triplehorn, Larry E. Watrous York. Cracraft, J. (1974) Phylogenetic models and classifi- and Quentin D. Wheeler for useful discussion cation. Systematic Zoology, 23, 7 1-90. and criticism pertinent to this manuscript. I Crampton, G.C. (1918) The genitalia and terminal also thank Dr Henk Wolda for generously abdominal structures of male Neuroptera and donating Panamanian material and Dr Paul Mecoptera with notes on the Psocidae, Iliptera Spangler for arranging the availability of and Trichoptera. Psyche, 25, 47-58. Davis, K.C. (1903) Sialididae of North America. Ecuadorian Covydalus collected by the Bulletin of the New York State Museum, 68, Ecuador-Peace Corps-Smithsonian Insti- 442-487. tution Aquatic Insect Survey. To Dr Oliver Ekis, G. (1977a) Classification, phylogeny, and S. Flint, Jr, I am especially grateful for the zoogeography of the genus Perilypus (Cole- optera: Cleridae). Smithsonian Contributions hospitality shown during a visit to the U.S. to Zoology, 227. National Museum, and for valuable comments Ekis, G. (1 977b) Classification and evolution of the regarding my studies. Dr Phillip A. Adams Central American heetle genus Colyphus (Cleri- offered his broad insights on Megaloptera and dae). 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'I'ONKIN (North 114-122. Viet Narn). (MNIIN); Montes Mauson (ZMHU). Ramhur, P. (1842) 6Iistoirc. Naturelle das Insectes A.orientalis (McLachlan). 6 specimens, including NPvropteres. Paris. type of A.ko1bei Weele (CHINA. SZECHWAN: Mt. Schuh, 11.1'. (1 976) I'retarsal structure in the Miridae Omei Shan - ZMHIJ). CHINA. HUI'FLH: Ichang (Hemiptera), with a cladistic arialysis of relation- (MCZC). SZECHWAN: Kuanshien (MCZC, USNM). ships within the family. American Museum Novi- Kouy Tcheou (MNHN). INDIA. ASSAM: United tates, 2601, 1-39. Jair~tiaKhasi Hills. Mynso (CNCC). Snodgrass, K.E. (1957) A revised interpretation of the external reproductive organs of male insects. Chloronia Banks. 6 species. Smithsonian Miscellaneo~tsCollections, 135 (6), C.mexicanu Stitz. 19 specimens. MEXICO. 1-60. CHIAI'AS: Chiapa de Corm (CNCC); 20-25 miles Tillyard, R.J. (1 9 18) Australian Megaloptera or N Huixtla (CNCC); I'alenque (CNCC, MCZC): 32 alderflies, descriptions of new genera and species. miles W San Cristdbal (CNCC). SAN LUIS PO'TOSI: Proceedings of the 1,innean Society of New El Salto (SF,MK). VERACRUZ: Cordoba (CNCC). South Wales. 43, 8 19-825. Tjeder, B. (1970) Neuroptera. Taxonomists Glossary C.pa1lidus (Ihvis). type in USNM. of Genitalia in Insects, 2nd edn (ed. by S. L. Chloronia spp. 39 specimens. BOLIVIA. Kio Longa Tuxen), pp. 89--99. Copenhagen. (MCZC). BRAZIL. SAO PAULO: Salisopolis Townsend, L.H. (1935) Key to larvae of certain (('ASC). Annapolis (MCZC). COLOMULA. META: families and genera of Nearctic Neuroptera. Villavicencio (MCZC). COSTA KlCA. CAKTAGO: Proceedings of the Entomological Society of (MCZC). GUATEMALA. SOLOLA: Olas de Moka Washington. 37, 26-30. (MCZC). PANAMA. CHIKIQUI: Hugaba (MCZC); Walker, 1'. (1853) Catalogue of the Specimens of Fortuna (MJGC); Lino (MCZC). PERU. HUANACO: Neuropterous Insects in the Collection qf the 'l'ingo Maria (SE:MK); Manson Valley (CASC). British Museum. London. VENISZUE1.A. Guarquka (MC'ZC); Northern Kange Walker, F. (1 860) Characters of undescrihed Neur- (MCZC). optera in the collection of W. W. Saunders. Truns- Corydalus Latreille. 13 species actions of the Entomological Society (~fI.ondon, 5, 176--199. C.affinis Burmeister. 40 specimens. ARGENTINA. Weele, H.W. van der (1906) hersicht der Sialiden BUENOS AIKES: (MCZC). MISSIONES: Arroyo des Indo-Malayischen Archipels. Notes fiom the Coati, 15 km E San Josk (USNM); Arroyo I.iso, Leyden Museum. 26, 207-222. 8 km W Gal Guemes (USNM); Arroyo Piray Guazu, Weele, H.W. van der (1907) Notizen iiber Sialiden N San Pedro (IJSNM); Arroyo Piray Mini, W ?os and Beschreibung einiger neuen Arten. Notes Hermanas (USNM); Panambi (PAAC). TUCUMAN: from the Leyden Museum, 28, 227--264. Ciudad Universitaria (USNM); Tafi de Valle (USNM). Weele, H.W. van der (1909) New genera and species BRAZIL. (MCZC, USNM). MINAS GEKAIS; Vicosa of Megaloptera Latr. Notes from the Leyden (CASC, USNM). PAKANA: (BMNH). RIO I)E Museum. 30, 249-264. JANIIIRO: Itatiaia, N Park (PAAY). SANTA Weele, H.W. van der (1910) Megaloptera. Collections ('A'I'AKINA: Nova Uremen (CUNY). SAO PAULO: Zoologiques du Baron Edm. de Selys Long- Salesopolis (CNCC); Sao Jose dos Campos (CUNY). champs, fasc. 5. COLOMBIA: Medellin (CASC). PAKAGUAY Wheeler, Q.D. (1979a) Slime mold beetles of the (MCZC). genus Anisotoma (Leiodidae): classification and evolution. Systematic Entomology, 4, 25 1 - C.armatus Hagen. Around 400 specimens. 309. AKGENTINA. JUJUY: (CUNY, LACM). Wheeler, Q.D. (1979b) Revision and cladistics of the TUCUMAN: (LACM); Quebrada de Lules (USNM). Middle American genus Creagrophorus Matthews Salto Pocitos (I:S(:A). BOLIVIA. COCHABAMBA: (Coleoptera, Leiodidae). Quaestiones Entomo- (PAAC); Chappare (PAAC, E'SCA). LA PAZ: logicae, 15, 447-479. Chulumani (USNM): Coroico Exp. Sta., N Yungas Wiley, E.O. (1979a) Cladograms and phylogenetic Prov. (USNM). COLOMBIA. (LACM). ANTIOQUIA: trees. Systematic Zoology, 28, 88-92. Jerico (USNM). META: Villavicencio (MCZC). Wiley, E.O. (1979b) An annotated Linnaean Hier- VALLE DE CUACA: (CASC); Altaflor (BMNH); archy, with comments on natural taxa and Anchicaya (CNCC); Cali (MCZC); Costa Rica competing systems. Systematic Zoology, 28, (CASC); La Esperansa (CASC). Buena Vista 308-337. (USNM); Medellin (USNM); Monterredondo (PAAC); Wood-Mason, J. (1884) Description of an Asian Praetera (MCZC); San Augustin (MCZC); San Jose species of the neuropterous genus Corydalis. (MCZC); San Vicente (MCZC); St. Antonio (MCZC). Proceedings of the Zoological Society of London, COSTA KICA. (MCZC, USNM). CAKTAGO: 1884, 1 10. (MCZC, USNM); Piedra Negra (USNM); Turrialba (LACM, USNM); Volcan de Turrialba (CMNH). Accepted 1 July 1980 GUANANCASTE: (SEMK). LIMON: Guapiles (SEMK); Los Diamantes (LACM). PUNTARENAS: (CUNY, MJGC, NWBR, UIDC, USNM); California Monte Verde (LACM, USNM); 4 miles E Palmar (CASC, USNM); Connecticut (USNM); District of (SEMK); Rincon de Osa (LACM); San Vito de Java Columbia (USNM); Florida (CNCC, CUNY, FSCA, (I.'SCA). IrCUA1)OK. (BMNH). EL ORO: 9 miles MCZC, USNM); Georgia (CNCC, USNM); Illinois S Santa Rosa (CASC'). GUAYAS: Guayanquil (CNCC, I'MNH, USNM); Indiana (CNCC, MCZC, (USNM). MORONA-SANTIAGO: (BMNH). NAPO: USNM): Iowa (UIIIC, IISNM); Kansas (CUNY, Puyo (CUNY, USNM); Tena (USNM). PASTAZA: MCZC); Kentucky (CNCC, OSUC, USNM); (CUNY, USNM). PICHINCHA: Quito (MCZC); Louisiana (CASC); Maine (USNM); Massachusetts Santo Domingo de Los Colorados (CASC); Tandapi (MCZC, USNM); Michigan (CUNY, FMNH); (SEMK). ZAMORA-CHINCHIPE: Rancho Allegro, Mississippi (I'SCA, USNM); Missouri (CASC, CNCC, 15 miles below Zamora (CASC). Balzapamba FMNH, NWBK, OSUC, USNM); New Hampshire (USNM); Banos (MCZC). Nanegalito (UIDC); Playas (CNCC); New Jersey (USNM); New Mexico (MCZC); de Montalva (MCZC). GUATEMALA. CHIMAL- New York (CUNY, MCZC, USNM); North Carolina TENANGO: Yepocapa (USNM). SACATEPEQUEZ: (SEMK, USNM); Ohio (CUNY, MJGC, NWBK, Finca Moca (USNM); 1;inca Parax6 (USNM). El USNM); Oklahoma (CUNY, FMNH, MCZC, UOKL, Cacao (USNM); Trece Aguas (USNM); San Marcos USNM); Pennsylvania (MCZC, USNM); Rhode Island (CASC). BRITISH HONDURAS. Middlesex, Stann Cr. (MCZC); Tennessee (FMNH); Texas (CASC, CUNY, Distr. (LACM). MEXICO. CHIAPAS: (PAAC). I'MNH, MCZC, TAMU, USNM); Utah (MCZC, VERACRUZ: Cordoba (LACM); IIos Amates USNM; Virginia (CIJNY, MCZC, NWBK, USNM). (PAAC). PANAMA. CHIRIQUI: Bugaba (MCZC, C.ecuadorianus Banks. 5 specimens, including holo- USNM); Fortuna (MJGC); Miramar (MJGC); k'otre- type (ECUADOI<: Banos - MCZC). COLOMBIA: rillos (ITSCA); Volcan de Chiriqui (BMNH, CASC). Anchicaya (CNCC); St Antonio (MCZC). PERU. (MNHN, SEMK). AMAZONAS: 5 km N Pomacocha (CASC). CAJAMAKCA: Ja6n Pucara C.flavicomis Stitz. 32, specimens, including type (CASC). HUANUCO: 10 miles N Huanta (CASC); (PANAMA: CHIRIQUI - ZMHU). COLOMBIA. Tingo Maria (CASC, USNM); Yurac, 67 miles E ANTIOQUIA: Jerico (USNM). Buena Vista (USNM). Tingo Maria (CASC). LIMA: Verrugas (USNM). COSTA KICA. LIMON: Hacienda Tapezco, 29 km PASCO: (LACM): Chontilla, 22 km SE Iscozazin W Tortuguero (LACM). PUNTAKENAS: 1.8 miles W (LACM); Pan de Azucar (LACM). Chanchamayo Rincon Oso Penninsula (LACM). ECUADOR. LOS (USNM, CASC); Chauchorango (MCZC); Chosica RIOS: Quevedo (USNM); 29 km W St I)omingo, (SEMK); Paltaybamba (USNM); San Miguel (USNM), Manabi (USNM); Quevedo (MCZC, USNM). Sinchono (USNM). VENEZUELA. (BMNH, MCZC). C.lutea Hagen. Around 500 specimens. COSTA ARAGUA: Kancho Grande (CUNY, USNM). RICA. GUANACASTE: 5 km N Canas (TAMU). near MERIDA: (BMNH, MCZC, MNHN, USNM). Paranos C:ua'piles (CUNY); 7 miles NW Liberia (USNM); Rio (MCZC). MIKANIIA: Caracas (MCZC). El Valle Corobici, Las Carias (USNM). EL SALVAIIOR. (USNM). Lomas Verde (TAMU); San Nicolas Lempa (USNM). C.batesi McLachlan. 15 specimens, including holo- GUATEMALA. CHIMALTENANGO: Yepocapa type (BRAZIL. AMAZONAS: hga - BMNH). (USNM). GUATEMALA: E SE Guatemala City BRAZIL. AMAZONAS: Manaus (INPA), Reserva (USNM). SAN MAKCOS: Puente Talisman (CUNY). Campinas (INPA), Reserva Diike (USNM). BOLIVA. SUCHITEP~QUEZ: Cuyotenango (USNM). ALTA SANTA CKUZ: (MCZC). COCHABAMBA: (CUNY). VERA PAZ: Trece Aguas (USNM). BRITISH ECUAI3OK. NAPO: Lago Agrio (USNM). GUYANA. HONDURAS. BELIZE: Belize (USNM). STANN (BMNH). ESSEQUIBO: 6 miles S Winniperu, CREEK: Middlesex (PAAC). HONDURAS. El Picrewana Island (USNM). Zamorano (OSUC); Siguatepeque Escuela Sup. Cienc. For. (PAAC). MEXICO: Chiapas. (CASC, C.bolivari Banks. 2 9 specimens, including types TAMU, USNM);.Colima (I&IS,rYM); Durango (OSUC); (VENEZUELA. Appun - MCZC). COLOMBIA. Guerrero (F'scA);' E5idatgb.i~~~~);Morelos (FSCA, (PAAC). VENEZUELA. ARAGUA: Kancho SEMK); Nayarit (SEMK); Nuevo Leon (CUNY, Grande (CUNY, PAAC, USNM). MERIDA: La FSCA, TAMU, USNM); Oaxaca (TAMU); Puebla Mucuy (CUNY). Appun (MCZC); Los Canales, (USNM); Queritaro (TAMU); San Luis Potosi (VSCA, Naiguata (MCZC); Priramo la Negra (MCZC); San SEMK, TAMU, USNM); Sinaloa (CNCC); Tamaulipas belipe Yara (CUNY). (CUNY); Veracruz (CASC, CNCC, PAAC, SEMK, C.cephalotes Kambur. 12 specimens. BRAZIL. TAMU, USNM); Zacatecas (OSUC). NICARAGUA. (MCZC). Chapada (BMNH, MCZC); Espirito Matagalpa (USNM). PANAMA. CANAL ZONE: Santo (BMNH); Sahara-Bello-Horizonte, Rio das Barro Colorado Island (CASC, CNCC, CUNY, INPA, Velhao (BMNH). MCZC, MJGC, SE,MK, USNM). CHIKIQUI: Miramar (MJGC)., COLON: Santa Rita Kidge (INPA). C.cornutus (Linnaeus). Around 900 specimens. PANAMA: Pearl Island (USNM). UNITFD STATES. CANADA: Ontario (CNCC); Quebec (CNCC). TEXAS: Bexar Co. (MCZC); Gillespie Co. (TAMU); MEXICO: Baja California Sur (CASC); Colima Hays Co. (MCZC); Hidalgo Co. (USNM); Kimble Co. (USNM); Distrito Federal (IJSNM); Durango (OSUC); Kleburg Co. (CUNY); Sabine Co. (USNM); (OSUC:, USNM); Guerrero (USNM); Jalisco Uvalde Co. (USNM); Webb Co. (TAMU). (('UNY, MCZC'); Morelos (CUNY, SEMK, USNM); Puehla (SEMK); Sinaloa (CNCC); Sonora (CUNY). C.nubilus ,Efichwn. 2 specimens, including holotype UNIT1:I) STATES: Arizona (CASC', CNCC. CUNY, (BRITISI-I GUYANA - ZMHU). 14RENCH T'MNH, MCZC, SEMK, lJll)C, USNM); Arkansas GUYANA: Saut-Maripa Oyapock (MNHN). 20 C.primitivus Weele. 12 specimens. AKGE;N'TINA. N.testaceus Karnbur. 5 specimens. INDONESIA. CATAMAKCA: Rio Portrero, near Andalgala JAVA: Saeka Poemi (MNHN). SUMATRA: (MCZC): (USNM). JUJUY: (CUNY). MJSSIONES: Campo l,ort Dekoch (MCZC); Redjang-Lebong (ZMHU). Viera (PAAC). TUC'UMAN: Quebrada de Lules Neuromus sp. 1 specimen. MALAYSIA. SABAH: (USNM). BOLIVIA. Rio Longa (USNM). BRAZIL. Mt Kinabalu (USNM). Chapada (MCZC). PARAGUAY. 2 km S Cerro Cora (USNM). PERU. PASCO: Chontilla, 22 km SI*: Platyneuromus Weele. 4 or 5 species. Iscozazia (USNM). P.auritus Kimmins. 10 specimens. GUATEMALA. C.tridentatus Stitz. 18 specimens, including type Cayuga (USNM). HON1)URAS. (BMNH); La Lima (BRAZIL - ZMHU). BRAZIL. (MCZC). (USNM). MEXICO. CHIAPAS: Chiapa (CASC); ARGENTINA. MISSIONES: Arroyo I'iray Mini, W Ocozoautla (CASC); Tuxtla Gutierrez (MCZC), Dos Hermanos (USNM); Piray Guazu (USNM). PAAC). Covydalus sp. 16 specimens. BOLIVIA. Cavinas Beni P.chuiste1 (Navis). 9 specimens, including type (USNM): Guayarin (MCZC). BRAZIL. AMAZONAS: (COSTA KICA, Turialba - IFPK). MEXICO, VERA- Manaus (INPA); Para Urua, 65 km SW Itaituha CKUZ (USNM); CHIAPAS: Tuxtla Guitierrez (INPA); Porto Velho, Kio Madieros (MCZC); Reserva (USNM). PANAMA. CHIRIQUI: Fortuna (MJCC); Campines (INPA). COLOMBIA. Buena Vista Santa Clara (INPA). No data. (BMNH, CASC). (USNhl); Puerto Berrio (MCZC). ECUAljOR. Huigra (MCZC). I'KENCH GUYANA. Saut-Maripa-Oyapock P.sorou (Hagen). 40 specimens, including types (MNHN); Sikini-Oyapock (MNHN). GUYANA. (Mb:XICO, Koppe - MCZC). COSTA RICA. Heredia, ITinca la Selva (USNM). HONDUIIAS. Bartica (BMNH): Rockstone, Essequibo River (BMNH). MEXICO. CHIAPAS: 6 miles S Puebla (USNM). PARAGUAY. (MCZC). PERU. Huallaga, Nueva (CASC); Tuxtla Gutierrez (MCZC). Aguatia (PAAC). Ithaca, New York (CUNY) - mis- MORELOS: Cuernevaca (MCZC). PUEBLA: Villa labelled. Juarez (MCZC). SAN I,UIS IJO'rOSI: F.1 Salto Neoneuromus Weele. 2 or three species. (CASC, MCZC, SEMK, USNM); El Naranjo (USNM); Nsikkin~ensisWeele. 2 specimens, including type I'alitla (USNM). VEKACKUZ: Ciudad Mendoza (INDIA, SlKKlM - ZMHU). INDIA. ASSAM: ((:AS(:); Fortin de Ins Flores (NWIjK); 5 miles N Khasia Hills (MCZC). Huatusco ('TAMU); Misantla (I3MNH); Teacalco Neoneuromus spp. 27 specimens, including types of (1JSNM); Tuxtla (USNM). Jalapa (BMNH); N.ferzestralis marluchlani (Weele) (TONKIN, Montes Rinconada (BMNH); Temescaltqpec (C'ASC); Xilitla Mauson - MC'ZC) and N.latratus tonkinerrsis (USNM). PANAMA. ('HIKIQUI: Fortuna (MJGC); (Weele) (CHINA. SZI-CHWAN: Mt. Omei Shan - Santa Clara (INI'A); Volcan de Chiriqui (BMNH). MCZC). CHINA, HUPEH: Suisap, Lichuan Distr. Platyneuromus spp. 10 specin~ens. GUATEMALA. (CASC). KIAN(;SI: Hong San (MC'ZC). I'anzos (USNM); Quirigua (USNM); Trece KWANGTtJN(;: Tsin Leongsham (CASC). Aguas (USNM). H0NL)URAS. I.omhardia (USNM). Mohkanshan (CASC); Yim Na San, 17 Ewantung MEXICO. CHIAl'AS: Palenque (C'NCC). VEKA- (MCZC); Yuman L:ou, Tau-Nin Kai (MNHN). INDIA. CRUZ (USNM). ASSAM: Kameng (Rhareli River), Dirdmq Ilzong (CNCC), Nakha (C'NC'C'), Salar (CNCC'). MANII'UK: Puotohermes Weele. 8 or 9 species. Chatrick (CNCC); Khamassom (CNCC). SIKKIM: P.costalis (Walker). 3 specimens. CHINA. HUPEH: Monqang (CNCC'); Simqhik (CNC'C'); Yugang Suisap, Lichuan Distr. (C'ASC). Kwaysi, Pui Liaw (CNCC). (MC'ZC). INDIA. ASSAM: Khasia Hills (MCZC). Neurherrnes Navis. 2 or 3 species. P.davidi Weele. Type (NOIITH VIET NAM. Mou N.maculipennis (Gray). 9 specimens. INIIONESIA. I'in - MNHN). SUMATRA: Medan (MCZC). JAVA: (MCZC); Mont Gedeh (MNHN). MALAYSIA. KEllAI1: Kedah Peak P.,qrandis (Thunberg). 9 specimens. JAPAN. (MCZC). SABAH: Mt Kinabalu (MNHN). kIOKKAIDO: (MCZC): Sapporo (MCZC). HONSHU: SELANGOK: Ulu Langat (MCZC). No data (MCZC). Kofu (CASC); Nikko (('AS('); Nippon Moyen, env. Tokyo (MNHN). Kyushu: Ashiya (CAS('). N.cosratostriatu (Weele). 2 specimens. INIIIA. ASSAM: United Jaintia Khasi Hills, Mawpran Prorohermrs spp. 43 specimens. CHINA. HUPKH: (CNCC). Mowshun (CNCC). Suisapa, Lichuan Ijistr. (CASC). SZECHWAN: (MCZC); Chengtu (CASC, MCZC); Chiang Nsien Neurhermes spp. 6 specimens. INDIA. MANII'UK: (MCZC); Mt Omei Shan (MCZC). Ta Hao, Eldipa Kambiron (C'NCC'); Lagairong (CNCC); Luaglang (MCZC). INDIA. ASSAM: Kameng (Bahareli River), Kunov (CNCC'). I ONKIN (North Viet Nam). Montes Uilo La (CNCC), Domkja (CNCC), Nakhu (CNCC), Mauson (MCZC). Nyukmadong (CNCC), Rupa (CNCC), Salar (CNCC), Neurotntrs Kamhur. 3 species. Shergaon (CNCC); United Jaintia Khasi Hills, Mawja N.intimzls McLachlan. 11 specimens. INDIA. (CNCC). MANIPUK: Chimasao (CNCC:); Khangoi ASSAM: Lushai (Hills), Sonai (CNCC). Nb:PAL. near (CNCC); Mapum (CNCC): WEST BENGAL: Lingsoka Birganj Lothar (CNCC). SOUTH VIE7 NAM. Pleiku (CNCC). INDONESIA. BORNEO: (MCZC). TAIWAN Prov., Nui Hong Kong (UIDC). (FORMOSA). Hori (MCZC); Rokki (MCZC). Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 2625

Reference Citation: Glorioso, M. J. 1981 [1981.??.??]. Systematics of the dobsonfly subfamily Corydalinae (Megaloptera: Corydalidae). Systematic Entomology 6:253-290.

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File: File produced for the Bibliography of the Neuropterida (BotN) component of the Global Lacewing Digital Library (GLDL) Project, 2007.