Victor Johnson.
Review of the Coniopterygidae (Neuroptera) of North America, with a Revision of the Genus Aleuropteryx.
Psyche 87:259-298, 1980.

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REVIEW OF THE CONIOPTERYGIDAE (NEUROPTERA) OF NORTH AMERICA WITH A REVISION OF THE
GENUS ALEUROPTERYXl
The smallest members of the Neuroptera belong to the family Coniopterygidae, which has several characteristics unusual within the order. The wings have reduced venation with few cross-veins and without the terminal branching of longitudinal veins found in other families. Coniopterygids produce a characteristic waxy secre- tion from wax glands on the head, thorax and abdomen which gives them their common name "dusty-wings"; no other neuropterans produce wax. Another unusual feature is the presence of a scler- otized penis in several genera and species. The only other neurop- terans with a sclerotized penis are some species of Mantispidae (Tjeder 1956).
The first major U.S. work on this group was a Nearctic revision by Banks (1906) who reported seven U.S. species and provided a key to genera based on wing venation. Because wing venation is not always bilaterally symmetrical, a single specimen may key out to different genera depending upon the wing used. The family re- mained in a state of confusion until Meinander (1972) published his world revision, in which he described 22 new species from North America. He described an additional nine species in 1974 and four more in 1975. Henry (1974) recorded the introduction and establish- ment of Aleuropteryx juniperi Ohm from Europe into the eastern part of the United States. Johnson (1976) described the genus 'This paper (No. 80-7-205) is published with the approval of the Director of the Kentucky Agricultural Experiment Station and is based on research performed in partial fulfillment of the requirements for the Ph.D. degree in Entomology. former research associate in the Department of Entomology, University of Kentucky, Lexington, Kentucky 40546. Present address: USDA-APHIS-PPQ, Box 182729, Erianger, Kentucky 41018.
To simplify information in this publication, trade names of some products are used. No endorsement is intended, nor is criticism implied of similar products not named.
Manuscript received by the editor February 2, 1981.



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260 Psyche [vol. 87
Bidesmida with one species (B. morrisoni) bringing the total to 40 species in 8 genera. This present study describes 5 additional new species. There is little doubt that there are many more undescribed species in North America.
Coniopterygids have been considered rare by most entomologists, mainly because of their small size, inconspicuous appearance and superficial resemblance to Psocoptera and Aleyrodidae. In addition, Withycombe (1922) stated that only the males are attracted to incandescent lights. I have found males and females only rarely attracted to incandescent lights, but both sexes are readily attracted to blacklights and the use of blacklight traps is the most efficient method of collecting adults.
Rather than being rare, many coniopterygids are quite common. Several species can be easily collected in numbers in the southern part of the United States where they appear to be widely and uniformly distributed. I have noted large numbers of Semidalis vicina (Hagen) congregating on oak and maple shrubbery in Kentucky, Tennessee and Arkansas and have seen this species and several others in large numbers in Texas feeding on an unidentified aphid on hackberry. Withycombe (1923) considered several coniop- terygids to be "exceedingly common, at least in England. . . ". He found Conwentzia sp. to be one of the "commonest insects in London parks" on Quercus and Platanus with Semidalis sp. and Coniopteryx sp. frequently plentiful.
The life histories of very few of the Coniopterygidae are known. Withycombe (1923) made a major contribution with studies of several species in England, Muma (1967) studied S. vicina in Florida and Henry (1976) reported on A. juniperi in Pennsylvania. Other studies have been made but these frequently include ambiguous species identification.
Both larvae and adults of coniopterygids are predaceous and several have been studied as biological control agents. Muma (1967) evaluated S. vicina for control of mites on citrus. He found this species to have several characteristics which made it suitable for biological control of citrus pests: wide distribution, stable mean population and wide food tolerance range, including rust mites, spider mites, whiteflies and scale insects. Individuals were found to consume 29-83 citrus red mites (eggs, larvae, and pupae) per day of development. Quayle (1 9 12) reported Conwentzia hageni Banks (Conwentzia california Meinander) to be one of the commonest



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19801 Johnson - Review of Coniopterygidae 26 1 enemies of spider mites in Southern California citrus, consuming 6-15 mites per day of development. Collyer (1951) reported C. pineticola Enderlein to consume 15-40 mites per day. Arrow (19 17) found Conwentzia psociformis Curtis to feed on Phylloxera sp. on oak, attacking all stages with "great voracity". Withycombe (1924) found that the main check on populations of Phylloxera punctata Light in England was C. psociformis. Henry (1976) stated that A. juniperi preys on two species of scale insects: juniper scale, Caru- laspis juniperi (Bouche) and the minute cypress scale, Carulaspis minima (Targioni-Tozetti). Ward (1970) also reported this species feeding on C. juniperi in Great Britain. In addition to feeding on live prey, adults have been reported to feed on other substances such as honeydew (Withycombe 1923 and Arrow 1917), scale secretions and honey water (Henry 1976). Several parasites are reported for coniopterygids. Withycombe (1923) reported two species of Hymenoptera to parasitize cocoons of C. psociformis in Great Britain; the female of Lygocera sp. tears a small hole in the cocoons for insertion of the parasite's eggs. The parasite larvae feed on the coniopterygid pupa, then pupate and emerge from the host cocoon the following summer. Ceraphron sp. have been found to emerge from coniopterygid cocoons during July and August. Muma (1967) reported three parasites of the pupae of C. vicina: Dendrocerus conwentziae Gah., Camptoptera sp. and Cosmoccidea morilli How.
In view of their predaceous habits and potential for biological control of mites and small insects, there is a need for biological information on this group. Life histories are known for very few species and few species have correlation between males and females and between larvae and adults. This present study was undertaken to advance the taxonomic status of this very interesting group so that life history investigations might be more easily completed. All specimens were examined either in glycerol or ethyl alcohol. Specimens on pins or points were treated with household cleaner, Formula 409@, for 2-24 hours, washed in water and then placed in glycerol. The Formula 409 softened the specimens and many times returned them to a state indistinguishable from specimens which have been stored in alcohol.




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262 Psyche [vo~. 87
For examination of genitalia, either the terminal abdominal segments or the entire abdomen was removed and treated with 10% KOH at room temperature for 2-10 hr. In some cases, specimens were placed in a cleaning sonicator for a few seconds, but this was too severe a treatment for general use. Specimens were then washed with water and placed in glycerol.
A dissecting microscope at 95X magnification was used for determination and for illustrations. A Zeiss Photomicroscope II@ was used for more detailed examination at magnifications up to 400X.
Illustrations were prepared freehand with the use of a counting grid mounted in one ocular of the dissecting microscope. Measure- ments are accurate to .04 mm. Illustrations are accurate to .02 mm. After examination and illustration, all specimens which had been dissected for examination of genitalia and all specimens which were fragile and/ or broken were placed in glycerol in genitalia vials for permanent storage. All other specimens were placed in vials of 80% ethyl alcohol with a few drops of glycerine. Specimens used in this study were from the collection of the author (AUTH) or borrowed from one of the following collections: Arizona State University (ARIZ), Illinois Natural History Survey (INHS) or Utah State University (UTAH). Types were deposited in one or more of these collections or in the National Museum of Natural History (USNM). Meinander (1972) types are deposited in the California Academy of Sciences (CAS), University of California at Berkeley (UCA) or the National Museum of Natural History. Due to the recent world revision by Meinander (1972), some material which is readily available in that revision was omitted from this study. Synonomies were limited to original descriptions and to publications after 1972. In the interest of standardization, I have followed Meinander's format regarding descriptions and illustra- tions and have used the same terminology where possible. I have noted my differences in interpretation from Meinander. I have prepared illustrations of previously undescribed species and other species where necessary.
The descriptions include only those characters which have been found to vary within the genus. Keys are provided to the species.



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19801
Johnson - Review of Coniopterygidae
Species are discussed in alphabetical order. Under the heading, new material examined, new country and state records are indicated by an asterisk. CONIOPTER YGIDAE Burmeister, 1839
Coniopterygidae Burmeister, l838:77 1. Type: Coniopteryx Curtis, 1834.
Eggs: Light colored, ovoid with micropylar end pointed and chorion reticulated. Incubation period varies with temperature. Withycombe (1924) reported 6-2 1 days for incubation. Larvae: Spindle-shaped, generally smooth, except for rows of small setae on each abdominal segment. Head small, rounded, prognathous; slightly retracted 'into thorax. Eyes formed of 4-5 facets. Antennae 2 segmented. Labrum projecting, covering the mandibles completely in Coniopteryginae and partially in Aleurop- teryginae. Mandibles enlarged basally, sharply pointed, slightly grooved ventrally. Maxillae pointed, with external barbs. No maxillary palpi present. Labrium reduced. Thorax large, soft, sharply marked off from abdomen. Three segments subequal. Legs well developed, long; tarsi one segmented with two simple claws and a well developed empodium. Tibia and tarsi not freely articulating.
Abdomen 10-segmented, tapering, with 10th segment adhesive. Notes: Meinander (1972) reported the labial palpi to be 2 segmented but Withycombe (1923) reported a pair of three seg- mented labial palpi.
The larval stages require about 2-4 wk for development. Withy- combe (1923), working with several species, and Henry (1976) working with A. juniperi, reported three larval instars, while Muma (1967) working with S. vicina and Quayle (1912) working with C. hageni reported 4 larval instars. Meinander (1 972) considers the family to have 3 larval instars. I have found 3 species in 2 genera of the Coniopteryginae to have 4 larval instars. Prior to pupation, the larvae build a flat, ovoid cocoon composed of an inner and an outer layer of white silk. The silk is produced by modified malpighian tubules (Meinander 1972) connected to an eversible anal papilla (Withycombe 1924). The cocoon is similar to that produced by some spiders and the emerging adult coniopterygids have been mistakenly thought to be parasites of spiders. Withycombe (1923) stated that



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264 Psyche [vo~. 87
there are 2 types of development in Conwentzia in England. During the early and mid-summer, pupation takes place normally and within about 2 weeks the adults emerge; in late summer, the larvae do not pupate immediately, but overwinter as prepupae in the cocoon and pupate early in the following spring. Henry (1976) reported A. juniper! in Pennsylvania overwintering as 2nd instar larvae and to be active during warm periods during the winter. Pupa: Quadrate, short with the head bent forward at an angle to thorax.
Notes: Just prior to emergence, a hole is cut in the cocoon through which the adult subsequently emerges. The pupal skin is either left entirely or partially within the cocoon or the pupa emerges and then sheds the pupal skin.
Adults: Hypognathous, strongly sclerotized head capsule. Ocelli absent, compound eyes well developed and large. Antennal sclerite and antennal condyle small. Antennal sockets often with unscler- otized area of frons between them. Male antennal segments modified and broader than in females. Mandibles small; maxillae well developed with 5-segmented palpi; labium well developed with 3- segmented palpi.
Prothorax short and lightly sclerotized; meso- and metathorax well developed, strongly sclerotized, with mesothorax being larger. Meso- and metathorax of some species with distinct lateral shoulder spots darker than surrounding areas.
Legs slender with 5-segmented tarsi, 4th segment flattened. Males with clasping setae on anterior femur.
Most with 2 pairs of well-developed wings. Exceptions are some species of Helicoconis and Coniopteryx and most species of Con- wentzia which have reduced metathoracic wings. Wings held roof- like over body.
Venation of fore wing: Costa (C) much reduced and visible only at base. Subcosta (Sc) runs parallel with margin and is distally furcate; posterior branch resembles a cross-vein (Withycombe 1922). Radius (R) in most species basally apparently crossing base of medius (M), but no actual crossing occurs. Radial sector (Rs) branches off in middle of wing. Media forked except in Conio- compsa. Basal cross-vein may be present along with radio-medial cross-vein (Aleuropteryginae have 2 radio-medial cross-veins). Cubitus (Cu) forks near base into 2 branches. One medio-cubital cross-vein present, striking M on stem or posterior branch. Two



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19801 Johnson - Review of Coniopterygidae 265 anal veins. Costal area with 2 or fewer cross-veins. Two basal cross- veins present between Cu2 and AI.
Venation of hind wing: Costa and subcosta similar to fore wing. Rs forks off near base in Aleropteryginae, more distally in Coniop- teryginae; generally furcate. Radial cross-vein between RI and Rs, strikes either stem of Rs or its anterior branch. M forked except in Coniopteryx and some Coniocompsa. A radio-medial cross-vein present, striking anterior branch of M except in Coniopteryx, where M is unforked. Cu forked near base. Two anal veins present. Two or fewer cross-veins in basal part of costal area. Membrane of wings mostly clear or unicolorous, but some with markings. Marginal fringes usually present and short. Frequently a group of hairs along base of posterior margin of hind wing and along base of costa of hind wing.
Abdomen weakly sclerotized except for genitalia. Segments 1-8 of Aleuropteryginae and segments 1-7 of Coniopteryginae with spiracles. Lateral and dorsal wax glands form transverse bands. Aleuropteryginae with paired organs called plicaturae on some segments; function unknown.
Notes: Newly emerged adults are pale and do not have the characteristic waxy power on the body and wings. Within a short time, the wax is secreted in long spirals from the wax glands, primarily located on the abdomen, head and thorax. The hind femora are used to apply the wax to the wings. The adults are normally active and run rapidly over plant surfaces. When disturbed, they either feign death or fly. Flight is fluttery and usually ends with a landing on the underside of vegetation.
Meinander (1972) treated the genitalia extensively in his revision; therefore a comprehensive study is not included here. Discussions of genitalia are included under each genus and species heading. With few exceptions, Meinander followed the interpretation and termi- nology of Tjeder (1970). I have followed the terminology of Meinander, although occasionally differing from his interpretation. Mating in Coniopterygidae is apparently accomplished by either of 2 methods: Parasemidalis sp. (Withycombe 1922) and C. pineti- cola (Collyer 1951) mate in a staggered parallel position with the male bending the tip of the abdomen dorsally to meet the female terminalia; A. juniperi mate with the male and female in a tail-to-tail position (Henry 1976). I have found 4 species of the subfamily



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266 Psyche [vo~. 87
Coniopteryginae (2 in Semidalis and 2 in Conwentzia) to mate in the staggered parallel position similar to that reported by Collyer. Eggs are laid singly on vegetation, frequently along leaf margins (Withycombe 1923) or on the underside of leaves near the midrib (Quayle 19 12). Muma found S. vicina to produce 2-5 eggs daily for 16-25 days for a maximum of 266 eggs per female. Adult longevity varies with environmental conditions. Muma (1967) found 5'. vicina adults to live about one month. The Coniopterygidae are divided into 2 subfamilies: Key to subfamilies
One radio-media cross-vein in middle of fore wing. ........... Galea one-segmented. No plicaturae on abdomen ........................................ Coniopteryginae Apparently two radio-medial cross-veins in middle of fore wing (either actually two radio-medial cross veins or one and Mi+z and R~+s briefly coalesce with the part of R4+5 between the point of fusion and the stem of Rs appearing as a second cross-vein). Galea ............. three-segmented. Plicaturae present on abdomen ........................................ Aleuropteryginae Subfamily Aleuropteryginae Enderlein, 1905 Aleuropteryginae Enderlein, 1905:225. Type: Aleuropteryx Low. Head capsule dorso-ventrally elongated in lateral view, genae long. Galea with basigalea and terminal knob. Fore wing with 2 radio-medial cross-veins and possible basal cross-vein. M has 2 setae in middle of wing, the bases of which are enlarged (except Bidesmida and possibly 2 other genera not in the area of this study). M frequently narrowed between setae. In hind wing, M and Cul run close to each other for more than half the length of Cul; frequently without any membrane showing between them. Rs branches off from R near base of wing. Radio-medial cross-vein strikes stem of Rs basally of radial cross-vein and strikes M (when M is forked) on the anterior branch. Plicaturae present on sterna 3-6. Wax glands of abdomen in a narrow transverse band on each of first eight terga (except one genus not in area of present study) and surrounding plicaturae. Eighth segment normally with stigmata. Ninth segment of males strongly sclerotized.




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19801 Johnson - Review of Coniopterygidae 267 Larvae with antennae and labial palpi of about equal length. Jaws long, slender, projecting from beneath labrum for more than half their length.
Notes: There are 4 known genera of this subfamily within the area of this study.
Key to North American genera of Aleuropteryginae 1. Radial cross-vein in hind wing strikes Rs on branch R2+3 ...
...................................................... 2 Radial cross-vein in hind wing strikes Rs on stem or on fork Aleuropteryx
........................................... 2. Seventh sternum of male with plicaturae.. ............... .3 Seventh sternum of male without plicaturae ..... Helicoconis 3. Bases of medial setae distinctly thickened .......... Neoconis Bases of medial setae not distinctly thickened. Medial setae not prominent ..................... Bidesmida Genus Aleuropteryx Low, 1885
Aleuropteryx Low, 1885:79. Type, by monotypy, Aleuropteryx Lowii Klapalek, 1894.
Antennae distinctly separated. Eyes small to medium. Head cap- sule and palpi usually dark.
Antennae 19-27 segmented. Scape and pedicel elongate; male pedicel with distinct ventral spine. Flagellar segments varying in length, usually as long as or longer than broad; hairs not arranged in distinct whorls.
Palpi unicolorous and slender, last segments of both labial and maxillary palpi only slightly broader than preceding segments. Basal segments of labial palpi distinctly longer than broad. Fore wings (Fig. 1) short, subtriangular, usually slightly more than twice as long as broad. Usually 2 basal cross-veins in costal area. Sc forks distally of radial cross-vein, which usually strikes R2+3, but sometimes strikes fork or stem of Rs. R4+5 coalesces with M1+2 in most specimens, but may be connected by a cross-vein which superficially appears to be a branch of M. M not thickened at bases of setae. Cross-veins rs-m and m-cul strike M close to each other in middle of wing. Cu2 sinuous, parallel with hind margin in all North American species. Sc, RI, distal end of R2+3 and basal parts of Cul, A1 and A2 thicker than other veins and bearing hairs. Longitudinal



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Psyche [vo~. 87
Figure 1. Wings of Aleuropteryx werneri
veins usually dark and cross-veins light, except cui-cu2 and posterior part of cross-vein m-cui, which are dark. Marginal fringes short. Hind wings usually slightly more than twice as long as broad. Usually 2 basal cross-veins in costal area. Sc forking distally of radial cross-vein, which strikes stem of Rs. Sc and distal parts of RI and R2+3 thicker than other veins and having hairs. Marginal fringe short.
Abdomen weakly sclerotized with limits of terga and sterna somewhat indistinct. Distinct plicaturae on segments 3-6 in both sexes and occasionally discernible on segment 2. Single narrow dorsal band of wax glands on segments 2-8 and surrounding plicaturae.
Male genitalia strongly sclerotized and basically internal. Ninth tergurn obliterate. Ninth sternum modified, dorsally fused in all



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19801 Johnson - Review of Coniopterygidae 269 North American species to form a ring through which the penis protrudes. Ventrally and anteriorly, 9th sternum has lateral apoph- yses (except in 2 species not within the area of this study). Ventrally and posteriorly, 9th sternum has an unimpaired structure called the process of the 9th sternum. Originating anteriorly of the process are paired structures called the appendages of the 9th sternum. They are usually backwards and dorsally directed, bear hairs at their apical ends and are either fused distally or are connected to a transverse plate. Appendages of 9th sternum are frequently membranous and difficult to delimit. Penis prominent with a pair of dorsolateral apodemes and a somewhat conical anterior one. Ectoprocts un- sclerotized in all North American species, but sclerotized or other- wise modified in some other species.
Female genitalia with ectoprocts often indistinct. Gonapophyses laterales forming a sclerotized ventral plate. Bursa copulatrix sclerotized and differing interspecifically. Notes: The male genitalia of Aleuropteryx differ from those of other genera and it is difficult to find homologous structures in other genera. The naming of structures in this case is for conveni- ence and an attempt is made to standardize terminology. This has not always been the case in the past (Meinander 1972). The shape of the bursa copulatrix and associated structures allow the females to be determined to species. The larval stages of only one North American species (A. juniperi) is known.
There are 14 species of this genus within the area of this study, 5 of which are herein described as new.
Key to species - males
Wings with distinct brownish spots .............. punctata ..........................
Wings without distinct spots .2
A pair of upward directed processes anterior to the process of the 9th sternum ....................................... 3 ............................
No such processes present .7
Processes anterior to process of 9th sternum longitudinally .........................
flattened, plate-like and short .4
Processes anterior to process of 9th sternum spine-like and ................................................. long 5



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270 Psyche [vo~. 87
Process of 9th sternum ending in a central, caudally directed, .................................
pointed process cupressi
Process of 9th sternum not as above.. ............ werneri Process of 9th sternum with 4 caudally directed processes, 2 ........................ central and 2 lateral.. dragoonica Process of 9th sternum ending in 3 processes, 1 central, 2 lateral .................................................... 6 Caudal projections of process of 9th sternum subequal in ......................................
length. arceuthobii
Central caudal projection of process of 9th sternum much larger ..............................
than lateral ones knowltoni
Sclerotized structure present just ventral to dorsal roof of 9th