The Green Lacewings of the Genus Chrysopa in Maryland ( Neuroptera: Chrysopidae)

Home , Chrysopa, Chrysopa nigricornis, Chrysopa oculata, Chrysopa quadripunctata, Chrysopidae

Ralph A. Bram and William E. Bickley Department of Entomology

INTRODUCTION

Tlw green lacewings which are members of the genus Chrysopa are extremely lwndicia1 insects. The larvae are commonly called aphislions and are well known as predators of aphids and other injurious insects. They play an important part in the regulation of populations of pests under natural conditions, and in California they have been cultured in mass and released for the control of mealy- bugs ( Finney, 1948 and 1950) . The positive identification of members of the genus is desirable for the use of biological-control workers and entomologists in general. Descriptions of most of the Nearctic species of Chrysopidae have relied heavily on body pigmentation and to a lesser extent on wing shape, venational patterns and coloration. Specimens fade when preserved in alcohol or on pins, and natural variation in color patterns occurs in many species ( Smith 1922, Bickley 1952). It is partly for these reasons that some of the most common and relatively abundant representatives of the family are not easily recognized. The chrysopid fauna of North America was treated comprehensively by Banks ( 1903). Smith ( 1922) contributed valuable information about the biology of the green lacewings and about the morphology and taxonomy of the larvae. He also pro- vided k<'ys and other help for the identification of species from Kansas ( 1925, 1934) and Canada ( 1932). Froeschner ( 194 7) similarly dealt with Missouri species. Bickley and MacLeod ( 1956) presented a review of the family as known to occur in the N earctic region north of Mexico. This paper included keys to the eight genera and to certain species groups as then understood. Subsequently Adams ( 1956) described the genus Pimachrysa. The task of analysing taxonomic characters and evaluating generic and specific relationships is so complex that progress is very slow. Some of the diffi- culties were discussed by Bickley and MacLeod ( 1956). The work here reported represents an attempt to clarify the status of a limited number of species. We recognize the desirability of a more thorough treatment, but we believe that many years will elapse before completion of a revision. In the meantime a key and othPr means for identifying Maryland species of Chrysopa should be helpful to any- one dealing with the fauna of Maryland and other eastern states. The species ?f Chrysopa included are undoubtedly the dominant representatives of the genus in North America east of the Rocky Mountains and north of Florida and south- ern Texas. More extensive collect'ing will make possible studies which will change distribution patterns and narrow the field of usefulness of the key. (_ .. The clarification of the status of the 10 species has been accomplished by Utilizing structures of the rnale genitalia as taxonomic characters. The term, external genitalia, is commonly reserved for the sclerotized structures of the seg- rne~t which contains the genital aperture. Terminalia is the generalized term which refers to all sclerotized structures of the 8th, 9th, 10th, and 11th abdominal segments. PREPARATION OF TERMINALIA FOR STUDY To prepare male t~rmin~.Iia f?r study, the la~t four or five ab~ominal segments were removed with nncrosc1ssors. The portion removed was nnmersed in 15-percent KOH solution and heated in a water bath for 15 minutes. It was then gently flushed with a 1 ml. hypodermic syringe equipped with a 27-gauge needle, which was inserted through the open end of the abdomen. The specimen was then rinsed in distilled water. StaininK~ suggested by Dr. P. Adams ( per~onal communication), was accomplished by 'immersing the entire abdomen for 2 to :3 minutes in a drop of 5-percent Chlorazol Black E aqueous solution. The specimen was again rinsed in distilled water and pJaced in one drop of glycerine on a slide. Specimens were permanently stored in microvials containing glycerin<\

MORPHOLOGY AND TERMINOLOGY OF THE MALE TERMINALIA Banks ( 1903) pointed out a few differences in the gross structure of the external genitalia. Smith ( 1932) recommended the study of the male genitalia and suggested staining with Biebricht'~ scarlet. Tjeder ( 1936) described the male genitalia of several Chinese chrysopids. Killington ( 1936 and 1937) figur~d the male genitalia of many British species, and Principi ( 1949) made detailed studies of the genitalia of several Palearctic chrysopids. The male genitalia of four Australian chrysopids were described by Kimmins ( 1952). Tjeder ( 1954 and 1956) attempted to stabilize the terminology of gen'ital structures in the order Neuroptera. Howe~er, not all subsequent workers have accepted thP "neutral terms" which he proposed. Although Hwang and Bickley ( 1961 ) followed the terminology of Tjeder in describing the structures of Chryso/Ja oculata Say, Zimmerman (1957) followed, to a degree, the terminology of Kill-- ington in describing Hawaiian chrysopids. Adams ( 1959) applied some na nws differently when describing Micronesian forms. A morphological interpretation of the male terminalia of all neuropteran families was undertaken by Acker ( 1960) . He found that more than 125 terms have been applied, and in the case of the chrysopids his usage of terms varies to some extent from that of other authors. Perhaps the basis for disagreement jn the interpretation of genitalic organs lies in the origin of the genitalia. Gustafson ( 1950) presented a historical survey of this problem and pointed out that the male genitalia in insects are usually regarded as belonging entirely to the ninth segment. In contrast_, \Vheeler ( 189~1), Else ( 1934), and Roonwal ( 1937) suggested that the genitalia arise from the lateral and sternal regions of the tenth segment. But Snodgrass ( 1935) reported that the male phallic organs ari~e from the conjunctiva] membrane behind the ninth sternum. Acker ( 1960) concluded that the genitalia of neuroptnans arise from the tenth sternite and coxopodite. Since there is such a diversity in taxonomic terms available for the description of male tenninalia, there seems to be no purpose in adding new terminology to an already confused situation. In referring to the male termina.lia, the n.omenclatorial system followed by Tjcder ( 1954 and 1956) and modified by Hwang and Bickley ( 196 I ) is used. The structures of the terminalia which are employed as taxonomic characters a re: the gonocristae, the transverse arch, the gonarcus, the entoprocessus, the pseudopenis, and the processus. The gonocristae ( fig. 26-31) 1 are a pair of retort-shaped, sclerotized pb te~

1The illustrations are grouped into five Plates, pages 14:-18. Text, refert>tH'f's art> to 1tlafr sc•d 1011.., bearing arabic numerals, which run consecutivel;v through the fh'e illustrations.

2 which are located on the dorsal surface of the hypovalva (fig. 12). These plates carry scale-like, overlapping teeth and in some species are connected by a transverse band of teeth. The gonocristae have been found in the genus Chrysopa only in species having oculata-like and lineaticornis-like terminalia. The transverse arch ( fig. 5) is composed of a pair of arched, anteriorly projecting arms which fuse posteriorly, producing a small backward-projecting tubercle. The arch terminates anteriorly with small hooks projecting laterally, which are attached just inside the callus cerci ( fig. 3) . The posterior tubercle may be seen to project from the abdomen just below the anus. The transverse arch has been found only in those species of Chrysopa having carnea-like term inalia. The gonarcus ( fig. 5) is situated internally between the anal segment and the ninth sternite. It is an arched structure with the paired, concave wings projecting anteriorly and fused posteriorly. In species having carnea-like terminalia, the wings of the gonarcus fuse posteriorly in a "V" -shaped juncture ( fig. 6) . In oculat a-like terminalia the wings of the gonarcus are parallel and are fused posteriorly by a thin, chitinous transverse bridge ( fig. 14) . Small, posteriorly directed processes may arise from the ventral surface of the transverse bridge. The point of fusion may also serve as a point of attachment for the pseudopenis in carnea-like tcnninalia ( fig. 6) . Directly beneath, and attached to the gonarcus is the entoprocessus ( fig. 5). In species with oculata-like terminalia, the entoprocessus is composed of paired, posteriorly directed processes which are fused to the ventral edge of the wings of the gonarcus (fig. 12). In species having carnea-like terminalia, the entoprocessus is represented by small triangular sclerites which are fused directly to the ventral edge of the wings of the gonarcus ( fig. 5) . The pseudopenis is a curved, tubelike, unpaired organ which Tjeder ( 1954 and 1956) considers to be derived from the fused parameres. In species with carnea-Iike terminalia, the pseudopenis is joined to the gonarcus at the point of fusion of the wings ( fig. 5) . It appears to articulate with the gonarcus. In oculata- like terminalia, the pseudopenis is not articulated with the gonarcus and appears as a large pointed structure (fig. 13) . In species possessing lineaticornis-like terminalia, a heavily sclerotized, unpaired process articulates at the hypovalva ( fig. 23) . This structure is internally directed anteriorly and terminates before reaching the eighth segment. A homo- logous process in the corydalids was called the columna by Crampton ( 1918 a and 1918 b). Tjeder ( 1954 and 1956) named a similar prolongation in a different corydalid species the processus. Tjeder's terminology is adopted here. The callus cerci are considered by most authors to be vestiges of the eleventh coxopodites or cerci. In the Neuroptera the cerci have been reduced to a pair of small pads located on the tenth tergite and containing sensory setae set in rosette sockets called trichobothria ( fig. 3). The callus cerci have been found present in every species of Chrysopa examined, but are considered to have little diagnostic value. The hypandrium internum was considered by Acker ( 1960) to be derived from the tenth sternite. It is situated on the dorsal lip of the gonopore and is a very small, keel-shaped sclerite, the poi1;t aim~d posteriorly and the conve~ curva- ture pointing ventrad. This structure 1s considered to have no taxonom1c value and will not be used as a diagnostic character. Below the gonarcus is a pair of hair-bearing membranous cavities which were termed by Killington ( 1936) "membranous sacs." These structures were found to be present in all species of Chrysopa examined, but their taxonomic significance has not been fully evaluated.

3 It is the hope of the authors that the key to species groups based on male terrninalia and the descriptions of the male terminalia will be used to supplement and confirm identifications made by other methods.

KEY TO THE SPECIES OF ~.~YSOPA KNOWN TO OCCUR IN MARYLAND 1. Antennae except the basal segment black, or with at least the basal fourth blackish ______2 Antennae except second segment entirely pale or with basal fourth pale; apical third may be brownish but not dark-brown ------_ ~1 2. Basal segment of antennae with a dark-red or black line on the lateral surface ______linl'al icorn is Basal segment of antennae unmarked ------nigricornis 3. Antennae with a black or brown ring on the second segment ______4 Antennae unmarked ------5 4. Interantennal spot forming an X ------"'·------chi Interantennal spot extending laterally toward eyes, not reaching vertex and not forming an X ------oculata 5. All veins entirely pale, or at most with only an occasional dark cross vein ____ 6 Gradates and some other veins marked with black or brown ______8 6. A definite narrow black or dark-red band from eye to mouth over the genae; varying amounts of red suffusion adjacent to black band; hind wing bluntly rounded at apex ( fig. 1) ------earn ea Genae lacking a definite black or dark-red band from eye to mouth ------7 7. Red suffusion on genae; hind wings angulate at apex ( fig. 2) ______lzarrisii Without red suffusion on genae; with or without black spots on genae and/or clypeus; large species ------nigri£:ornis 8. Pronotum with 2 or 3 orange spots on each side; orange spots usually on thorax and abdomen and often on head near eye ______quadriJJUntlala Pronotum, thorax and abdomen without orange spots ------9 9. A red stripe on genae from eye to mouth ------rufilabris Genae lacking red stripes ------10 10. A black band across the frons beneath antennae ------incomplcta Frons without any markings; clypeus usually with a pair of b.lac_k . spots ------__ ---- __ -- -- __ -- -- __ --- 1Z. zgrZC() r ll lS

KEY TO GROUPS OF MARYLAND SPECIES OF CHRYSOPA BASED ON MALE TERMINALIA 1. Transverse arch present; gonocristae absent ------carnea, harrisii and rufilah,·i_,- 2. Transverse arch absent; gonocristae present ------~1 3. Pseudopenis joined to gonarcus ______/ineaticorni., Pseudopenis separated from gonarcus ------______o cu lat a, inc om p let a, chi, quadri Jm n c I at a, and n l.[!J i 01 rn i \.

Chrysopa carnea Stephens, 1836 Until recently, this species has been known from the Nrarctic rt:'gion ~,s Chrysopa plorabunda Fitch. Tjeder ( 1960) synonymized plorabunda with earn ca on the basis of genital structures. According to Tjeder ( 1960) this species is now the most common and widely distributed one of the world fauna of Chrysopidac. It occurs throughout Europe, North Africa, Asia Minor, Palestine, Persia, Meso- potamia, Afghanistan, India, Siberia, China, Japan, Formosa_, tlw Azores, and

4 throughout North America. It is characterized by having pale-green wing venation, a whitish median, dorsal stripe, and a straight brown line on the gena. Killington ( 19 3 7) figured the male terminalia of this species. The terminalia of this species lack gonocristae ( fig. 3) . In a freshly collected specimen, the pseudopenis and the tubercle of the transverse arch may be seen projecting from the terminal end of the abdomen (fig. 3). The gonarcus is a wing-shaped, sclerotized structure, the wings being fused posteriorly ( fig. 5 and 6). At the point of fusion, the pseudopenis is attached. The entoprocessus ( fig. 5) is a triangular sclerite which is fused to the ventral edge of the wings of the gonarcus. The pseudopenis is distinctive in this species. The organ proceeds down from its base and is incurved at the terminal end. The dors:al surface of the pseudopenis contains longitudinal striations distally ( fig. 4). Variation in the pseudopenis was slight, and every specimen examined contained the dorsal, longitudinal striations. Differences in the general shape of the pseudopenis were minor. These con- clusions are based on the study of about 75 specimens from Maryland, Alask~, Arkansas, California, Idaho, Illinois, Kansas, Michigan, New Jersey, Texas, Utah, Virginia, and Washington. Rearing experiments were conducted to observe possible structural variation in terminalia. Three gravid females were collected in the College Park, Maryland, area during late August 1959. The progeny of these females were reared, and 46 male and 35 female adults survived as the first generation. Three successful "crosses" were made among individuals of the first generation. These three "crosses" resulted in 44 individuals in the second generation. Six "crosses" between members of the second generation were made, including various combi- nations and one "back cross." These "crosses" resulted in 57 progeny in the third generation. All specimens reared were pinned and the male terminalia studied. No variation in the structure of the male terminalia was noted in any of the specimens examined.

Chrysopa rufilabris Burmeister, 1839 This species was described from "Mittel Amerika and Mexiko" and has been recorded from many localities in eastern North America. Bickley and Mac- Leod (1956) reported that the collection of the California Academy of Science contains specimens from California, Arizona, Texas., Nevada, Oregon, and Mon- tana. The species is characterized by: green wing venation with the gradate~ and the ends of some cross ve:ins black, a pale yellow median dorsal stripe, and a straight red line on the gena. Terminalia in rufilabris are carnea-like (fig. 7 and 8). The gonarcus in this species differs from carnea and harrisii in general shape. The entoprocessus is represented by triangular sclerites arising from the ventral edge of the gonarcus. Each half of the transverse arch differs from that of carnea by being shorter, and from that of harrisii by being less abruptly curved. The pseudopenis is distinct from that of carnea in lacking the dorsal, longitudinal striations and being only slightly curved distally rather than incurved. The pseudopenis differs from harrisii in being only obtusely angulate basally. ~etae may be present on the dorsal surface of the pseudopenis. The setae are set in small tubercles, and their number varies not only among specimens but also on each side of the pseudopenis. Specimens have been seen with from O to 11 setae on the pseudopenis. · An attempt was made to discover the extent of variation in the setae on the pseudopenis. A sample of 149 male specimens collected in Maryland was studied

5 and compared with a sample of 100 male specimens collected in New Orleans, Louisiana. In the sample from Louisiana 4 percent lacked setae and 82 percent had from two to seven setae. In the sample from Maryland 75 percent lacked setae, and 15 percent had from two to seven setae. A minor difference in the colorational patterns in the two populations was noted. Specimens from the Louisiana population, in general, had a slightly broad- er red genal line than did specimens from the Maryland population. The setae of the membranous sac were, in general, sligh'tly longer in individuals of the Louisiana population. Specimens with setae on the pseudopenis have also been seen from Kansas, Texas, and Florida. Eleven of 12 males, offspring of a single Maryland female, lacked setae on the pseudopenis. The twelfth reared male had one seta. A series of 24 male rufilabris from 9 different localities in California was studied. Examination of the pseudopenis revealed that all spedimens possessed dorsal longitudinal striations and lacked setae. Chrysopa medialis Banks, 1903-NEW SYNONYMY. Banks described this species from a few specimens collected at High Island, Maryland, in late Sep- tember. It was characterized by a red line on the gena; a dorsal median red stripe from the pronotum to ~he tip of the abdomen; green wing venation with the gradates and the tips of almost all cross-veins marked with black. Since the original description, the name has appeared only in Banks' ( 1907) catalogue and in the 1956 synopsis of Bickley and MacLeod. Bickley and :Mac- Leod ( 1956) suggested that this species is very near rufilabris and quadrijnmctata and that the validity of the species is questionable. The type is a female. A male specimen which exactly fits the description of medialis was collected in Colll'gc Park, Maryland on January 29, 1954. Examination of the male termina1ia revealed that the specimen was actually rufilabris. Inasmuch as all available evi- dence indicates that medialis was no more than a winter form of rufilahris, Chrysopa medialis Banks is here considered to be a synonym of C. rufilabris Burmeister. Chrysopa interrupta Schneider, 1851-NE1W SYNONYMY. This species has been extremely difficult to distinguish from rufilabris. Banks ( 1903) reported that interrupta is very close to rufilabris and does not appear to be common any- where. Bickley and MacLeod ( 1956) also suggested that this species is \·cry close to rufilabris. The species was described from Pennsylvania, and has been recorded from seven eastern states and from Arizona, California, Kansas, and Illinois. Chrysopa interrupta is characterized by being pale-straw yellow througfo>t1L The gradate series and most of the other cross-veinlets of the forewing 3re brmvn. A reddish streak extends from each eye to the mouth on the gena, and there is no pale median dorsal stripe. These color characters are hopelessly inadeq ua tc for the separation of this species from rufilabris. By studying Schneider's ( 1851 ) col- ored lithograph one may obtain a concept of interrupta. Banks and Roger ( '.. Smith had a concept of this species as being like rufilabris, but pale. \Ve hau~ examined three male specimens identified as interrupta by Banks and six male specimens considered by Smith to be interrupta. The genitalia of these nine specimens are identical with those of rufilabris. The type of interrupta was deposited in the Berlin Museum by Schneider. We have corresponded with several European workers over an extended period and have been unable to locate the type. Although the type has not been examined, Chrysopa interrupta Schneider is here adjudged to be a synonym of C. rufilabris Burmeister.

6 Chrysopa harrisii Fitch, 1856 Chrysopa harrisii was described from New York and is widely distributed throughout the Nearctic region. Thfa species is often difficult to separate from C. carnea~ particularly in preserved material. The species is characterized by having pale wing venation, an acute wing tip, a pale yellowish median, dorsal stripe, and red suffusion on the gena. The terminalia of this species are carnea-l'ike. No gonocristae are present, and the pseudopenis is joined to the point of fusion of the wings of the gonarcus. The gonarcus differs from carnea in being more slender dorso-ventrally; the entoprocessus is triangular but not as prominent; and the transverse arch is more widely extended laterally than in carnea (fig. 9 and 10). The pseudopenis is quite distinct. It is only slightly curved distally and lacks the dorsal longitudinal striations present in carnea ( fig. 11). Basally, the pseudopenis is acutely angulate near the point of articulation, whereas the pseudopenis in carnea is rounded basally. No variation in male terminalia was noted in approximately '20 specimens examined from Maryland, Michigan, California, Virginia, and Mex'ico.

Chrysopa oculata Say, 1839 This species was described from the "U. S." and is the most common lacewing throughout many areas of the Nearctic region. It is an extremely variable species, and there are 16 names which may be applied to identify individuals which differ primarily in color markings on the head and wings. Bickley ( 1952) concluded that the varietal names have little value. Chrysopa oculata is characterized by having a hlack crescent under each eye; a broad blackish band under the antenna! sockets; and the sockets above are margined by a narrow blackish line. Between the antennae a reddish spot ascends and the vertex has two submedian black dots. Wing venation is green with many of the cross-veinlets black. Hwang and Bickley ( 1961) described the reproductive system of this species. The terminalia of oculata are quite distinct from those species having carnea- like terminalia ( fig. 12, 13 and 14) . Gonocristae are present on the dorsal surface of the hypovalva. The pseudopenis is separated from the gonarcus. A transverse arch is not present, and the entoprocessus is extended into two posteriorly directed processes. The gonocristae are retort-shaped structures with the pointed, elongated teeth overlapping ( fig. 26) . The gonarcus is a paired, arched structure, with the concave wings directed anteriorly ( fig. 13) . The wings are parallel and are fused posteriorly by a thin transverse bridge. Processes project from the posterior edge of the gonarcal bridge ( fig. 14). The processes of the entoprocessus are fused anteriorly beneath the gonarcus and are directed posteriorly with the pointed ends slightly downcurved. The processes of the entoprocessus are fused to the ventral edge of the wings of the gonarcus ( fig. 13 and 14) . The pseudopenis is sparsely spiculate basally ( not illustrated) and the tube-like, unpaired organ gradually flattens toward the pointed end.

Chrysopa incompleta Banks, 1911 Chrysopa incompleta was described from North Carolina, and specimens have been examined by the authors from Maryland, Massachusetts, New Jersey, Virginia, and Georgia. This species is characterized by the absence of the dark band around the second segment of the antennae and by the presence of a black band across the

7 face just beneath the antennae, a black spot on each gena, and one on each side of the clypeus; two black stripes on the vertex; anterior outer edge of pronotum black; wings with longitudinal veins green, gradates and cross veins black. This species is very similar in appearance to oculata but differs in the clypeal marks and the less marked antenna! sockets. The male terminalia of incompleta are oculata-Iike ( fig. 15 and 16). The gonocristae are similar to oculata but differ in general conformation ( fig. 27). The posterior edge of the gonarcal bridge has no posteriorly directed processes as does oculata~ but the entire gonarcal bridge is greatly extended posteriorly and the posterior border is serrate. A pair of small, posteriorly projecting processes are present on the anterior ventral edge of the gonarcal bridge as in quadripunctata. The entoprocessus is extended into posteriorly directed processes with the distal end downcurved. The pseudopenis is sparsely spiculate basally ( not illustrated) and terminates in a pointed, incurved tip. Eight male specimens of Chrysopa incompleta were examined including ttw type. No variation in the terminalia was observed.

Chrysopa chi Fitch, 1856 This species was described from New York and has been recorded from the Northeastern United States as far south as Tennessee. It has also been reported from seven Canadian provinces, Minnesota, California, and we have examined specimens from Montana, Michigan, and Alaska. Chrysopa chi is characterized by having a black ring on the second antenna] segment; a conspicuous, Y-shaped, black mark between the antennae and a pair of black spots below the bases of the antennae ( the inter-antenna! mark and the subantennal spots may, or may not, be connected) ; black spots also present on the clypeus and upper border of eye; pronotum and mesonotum with symmetric a] black spots dorsally; wing with longitudinal veins green, gradates and cross veins black. The male terminalia of chi are oculata-like ( fig. 17 and 18). The gonocristae differ somewhat in general shape and placement of individual teeth ( fig. 28). The posterior edge of the gonarcal bridge is not projected into a pair of posteriorly directed processes as in oculata, but exhibits a pair of small, posteriorly projecting processes on the anterior ventral edge as in quadripunctata. The processes of the entoprocessus meet anteriorly and are directed posteriorly terminat- ing in a pointed, slightly downcurved end. The pseudopenis is quite distinctive. It is exceptionally long and broad and terminates in a sharp point. A thin, chitin- ized shield-like structure flanges out at the apical tip of the pseudopenis and the basal portion is sparsely spiculate ( not illustrated). Nineteen male specimens of Chrysopa chi were examined fr01;1 Maryland. New York, Michigan, Montana, Canada, Massachusetts, and Alaska. All terrninalia examined appeared consistent.

Chrysopa quadripunctata Burmeister, 1839 Chrysopa quadripunctata was described from North America and has been recorded from seven eastern states, six central states, and from VancounT Island. Three specimens from Kelseyville, California, collected in 1933 are in the colll'c- tion of the Univers1ity of California, Berkeley. All three specimens are females. and they exactly fit the description of this species. Another female quadripunctala was collected in Stockton, California on July 17, 1960 by Mr. G. Taussig, and is in the collection of the University of Maryland. The known range of this specie~ is thus extended into central California southward from British Columbia.

8 This species is characterized by having a reddish line on the g-ena; four or six bright orange spots on the thorax; and the abdomen is marked with orange patches dorsally. vVing venation is greenish yellow with the gradates and the ends of some cross-veins black. The male terminalia of quadripunctata are oculata-like (fig. 19 and 20). The gonocristae are not separated from each other as in oculata, but are con- m~cted by a broad band of scale-like teeth ( fig. 29). The posterior edge of the gonarcal bridge is not projected into a pair of posteriorly directed processes as in oculata. On the anterior ventral edge of the gonarcal bridge two small processes project posterio-laterally (fig. 20). The processes of the entoprocessus meet anteriorly and are directed posteriorly with the pointed end downcmved. The pseudopenis is separated from the gonarcus and is slightly curved and pointed distally. No spicules are present basally on the pseudopenis. Five male quadripunctata collected at light traps in Maryland during the summer of 1959, and three specimens collected in Virginia during the summer of 1949 were studied. The male terminalia of all specimens examined appeared consistent.

Chrysopa nigricornis Burmeister, 1839 This species was described from North America and is widely distributed in the Nearctic region. It is a large species characterized by having the basal fourth of the antenna! flagellum dark and by having a black spot on each side at the base of the clypeus. After studying more than 200 specimens from various parts of the Nearctic Region we have concluded that the characterization of C. nigricornis must be drastically changed. The most reliable characters are found in the male terminalia and are described below. As currently understood Chrysopa nigricornis may or may not have the basal fourth of the antenna.I flagellum black. The black spots on the clypeus were present in all but one of the specimens examined. In this specimen a minute brown spot can be recognized on one side of the clypeus. It is so faint that it would not be observed without painstakingly ma- nipulating the specimen under strong light with high magnification. In several other specimens these spots were greatly reduced and scarcely detectable. It is estimated that 90 percent of a representative population of the species will have dark spots on the genae in addition to those on the clypeus. Usually the venation is green, with the exception of the gradates and costal cro5s-veinlets. However, additional dark cross veins sometimes occur. In a few cases the gradates are not dark, and one has the impression that aH veins are green; but by a careful search three or four dark cross veins are discovered. Specimens with pale antennae and without clypeal marks may be confused with C. harrisii if the gradates are pale or with C. rufilabris if the gradates are dark. Male genitalia of specimens from widely scattered areas in the Nearctic Region are consistent and are indicative of the species. The male terminalia of Chrysopa nigricornis are oculata-like (fig. 21 and 22). Gonocristae are present, forming a continuous band across the hypovalva. Laterally the teeth are small and do not overlap, but centrally there is a group of about 24 large teeth which overlap (fig. 30). Each tooth is broad and short with a pointed tip. On the anterior ventral edge of the gonarcal bridge two small processes project posteriorly ( fig. 21 and 22), however, these processes are longer than those in quadripunctata. The entoprocessus is extended into posteriorly directed processes with the distal end downcurved. The entoprocessus is situated considerably below the gonarcus and is fused to it by a sclerotized connection. The pseudopenis is similar to oculata but is not spiculate basally.

9 Chrysopa majuscula Banks, 1898-NEW SYNONYMY. This species was described from California and has been recorded from British Columbia, Canada , ' Minnesota, and New Mexico. The species w~s characterized by having a black spot on each side near the base of the clypeus. Wing venation is green with black gradates. The costal and radial cross-veinlets are black medially and green term- inally. Smith ( 1932) reported that this was a very common species and resembled nigricornis except for the pale antennae. The type of this species ( called ery- throcephala by Banks in 1898 and changed to majuscula in 1906) is a female. A male specimen in the Museum of Comparative Zoology which Banks undoubtedly considered to be majuscula and five male specimens from Maryland, Cali- fornia, and Utah which fit the description of majuscula have genitalia identical with those of nigricornis. C. majuscula is therefor adjudged to be a synonym.

Chrysopa lineaficornis Fitch, 1856 Chrysopa lineaticornis was described from New York, and has been recorded from Maryland, Michigan, New England, North Carolina, Quebec, Tennessee, and Virginia. It is characterized by a black line on the antenna! scape, and the basal fourth of the antenna] flagellum is black. The prothorax sometimes has a red stripe on the lateral edge. Wing venation is green, but the gradates, costals, and often the radial cross-veinlets are black; all pterostigmas are distinct. The male tenninalia of this species are quite distinct from both the c:arnca- like and the oculata-Iike types. Gonocristae are present. The teeth, which form a continuous transverse band across the hypovalva, are very small and distinctly separated from one another (fig. 31). The processus terminates in the area of the callus ce:rci ( fig. 23). A transverse arch is absent. The gonarcus is a wing- shaped, sderotized structure with posteriorly fused wings ( fig. 24 and 25) . The entoprocessus is reduced to posteriorly directed processes which are slightly downcurved termiinally. The pseudopenis is extremely specialized. It articulates from a ventral extension of the gonarcus. Another articulation occurs about midway on the pseudopenis. On the basal portion of the pseudopenis are two posteriorly directed processes which slightly overlap the median articulation. Distally, the pseudopenis narrows somewhat and terminates in a pointed projection. Laterally at the distal end paired cheliform processes are present. This species is superficially similar to representatives of C. nigricornis \vhich have the basal fourth of the antennal flagellum dark, but the dark lines on the antennal scape have been diagnostic. Also the clypeal spots which are usually present in nigricornis are absent in lineaticornis. A study of a small number of specimens from Massachusetts, Maryland and Georgia has shown that the lines on the antenna] scape are not always prominent. \Vhen the lines are very faint the specimen may be distinguished from certain specimens of nigricornis only by size and by geni talic characters. Chrysopa columbiana Banks, 1903-NEW SYNONYMY. This species was described from Washington, D. C. and was considered by Banks to be related to n.igricornis but dist'inct in having the clypeus unmarked. The type is a male, and the genitalia are identical with those of C. lineaticornis. The lines on the antennal scape are extremely faint. It may be assumed that they were so indistinct that d1ey were overlooked by Banks. Because the male genitalia of columbiana are identical with those of lineaticornis the former is here treated as a synonym.

10 SUMMARY This is a report of work undertaken to clarify the taxonomic status of the species of green lacewings ( aphislions) known to occur in Maryland and neigh- boring states. Studies of male terminalia were made following clearing in potassi- um hydroxide and staining with aqueous Chlorozol Black E. The species were evaluated and consideration was given to genitalic characters, as well as previously used characters such as color of antennae, color markings on the head and thorax, color of wing veins and venational patterns. The structures of the male terminalia which vary among species by their presence or absence or by their conformation are: the gonocristae, the transverse arch, the gonarcus, the entoprocessus, the pseudopenis, and the processus. A key to the ten species is presented. A key to groups of species based on male genitalia is also given with the hope that it can be used by nonspecialists to supplement and confirm identifications made by other methods. Chrysopa carnea 1== jJlorabunda auct.], C. harri')ii) and C. rufilabris fall into one group; C. lineaticornis represents another group; and the third group consists of C. oculata, C. inconz plcta) C. chi) C. quadripunctata) and C. nigricornis. C hryso pa media/is Banks, 1903 and C. interru pta Schneider, 1851, were with C. rufilabris Burmeister, 1839. C. majuscula Banks, 1898 was synonyrnized with C. nigricornis Burmeister, 1839. As currently understood, C. niRricornis may or may not have the basal fourth of the flagellum black, and in some specimens the clypeal spots are scarcely detectable. C. columbiana Banks~ 1903 was synonymized with C. lineaticornis Fitch, 1856.

LITERATURE CITED Acker, T. S. 1960. The comparative morphology of the male terminalia of Neuroptcra (Insecta). Microentomology 24: 25-84. Adams, P. A. 1956. A new genus and new species of Chrysopidae from the western United States, with remarks on the wing venation of the family (Neuroptera). Psyche 63: 67-74. 1959. Neuroptera: Myrrneleontidae and Chrysopidae. Insects of Micronesia 8(2): 13-33. Honolulu: Bernice P. Bishoo Museum. Banks, N. 1903. A revision of the Nearctic Chrysopidae. Trans. Amer. Ent. Soc. 24 : 13 7-16 2. 1907. Catalogue of the neuropteroid insects of the United States. American . Entomological Society, Philadelphia. pp. 1-53. Bickley, \IV. E. 1952. Inheritance of· some varietal characters in Chrysopa oculata Say. Psyche 59: 41-46. Bickley, \V. E. and E. G. MacLeod. 1956. A synopsis of the Nearctic Chryso- pidae with a key to the genera ( N europtera) . Proc. Ent. Soc. Wash. 58 : 177-202. Crampton, G. C. 1918 a. A phylogenetic study of the terminal abdominal structures and genitalia of male Apterygota, Ephemerida, Odonata, Plecop- tera, Neuroptera, Orthoptera, and their allies. Bull. Brooklyn Ent. Soc. 13: 49-68. 1918 b. The genitalia of Neuroptera and Mecoptera with notes on Psocidae, Diptera, and Trichoptera. Psyche 25: 47-59. Else, F. L. 1934. The developmental anatomy of male genitalia in Melanoplus difjerentialis. ]our. Morph. 55: 577-609.

11 Finney, G. L. 1948. Culturing ChrysojJa californica and obtaining eggs for field distribution. Jour. Econ. Ent. 41: 719-721. 1950. Mass culturing Chrysopa californica to obtain eggs for field distribution. Jour. Econ. Ent. 43: 97-100. Froeschner, R. C. 1947. Notes and key to the Neuroptera of Missouri. Ann. Ent. Soc. Amer. 40: 123-136. Gustafson, J. F. 1950. The origin and evolution of the genitalia of the Insecta. Microentomology 15: 35-67. Hwang, J. C. and W. E. Bickley. 1961. The reproductive system of ChrysojJa oculata Say (Neuroptera: Chrysopidae). Ann. Ent. Soc. Amer. 54: 422-429. Killington, F. J. 1936. A monograph of the British Neuroptera. Vol. 1. Lon- don: Ray Society, 269 pp. 1937. A monograph of the British Neuroptera. Vol. 2. London: Ray So6e- ty, 306 pp. Kimmins, D. E. 1952. Some new Australian Chrysopidae. Ann. Mag. Nat. History 12 : 49 : 69-81. Principi, M .. M. 1949. Contributi allo studio dei Neurotteri Italiana VIII. Morfologia, anatomia e funzionamento degli apparati genitali nel gen. C hry- sopa Leach. Boll. Inst. Univ. Bologna 17: 314-362. Roonwal, M. L. 1937. Studies on the embryology of the African migratory lo, cust, Locusta migratoria migratorioides (R. and F.) II. Organogeny. Philosophical Trans. Royal Soc. Lond., B. 227: 175-244. Schneider, W. G. 1851. Symbolae ad monographiam generis Chrysopac: Leach. Vrotislaviae, 178 pp., 60 pls. Smith, R. C. 192'2. The biology of the Chrysopidae. N. Y. (Cornell) Agr. Expt. Sta. Mem. 58: 1'285-1'377. 1925. The Neuroptera and Mecoptera of Kansas. Bull. Brooklyn Ent. Soc. 20: 165-171. 1932. The Chrysopidae (Neuroptera) of Canada. Ann. Ent. Soc. Amer. 25: 579-601. 1934. Notes on the Neuroptera and Mecoptera of Kansas with keys for the identification of species. Jour. Kansas Ent. Soc. 7: 120-145. Snodgrass, R. E. 1935. Principles of insect morphology. New York: McGraw- Hill. pp. 1-66 7. Tjeder, B. 1936. Schwedisch-Chinesische wessensc haftliche nach den nordwest- lichen provinzen Chinas. 62. Neuroptera. Arkir Zoology 29A: 1-36. 1954. Genital structures and tern:inology in the order Neuroptera. Entorn. Medd. 27: 23-40. 1956. Neuroptera. In Tuxen, S. L., Taxonomist's glossary of genitalia :n insects. Copenhagen: Munksgaara, pp. 76-83. 1960. Neuroptera from Newfoundland, Miquelon, and Labrador. Opusc. Ent. 25: 146-149. Wheeler, W. M. 1893. A contribution to insect embryology. Jour. Morph. 8: 1-160.

Zimmermah, E. C. 1957. Ephemeroptera-Neuroptera-Trichoptera. Insects 0 : Hawaii 6: 1-209. Honolulu: Univ. of Hawaii Press.

12 FIGURES

EXPLANATION OF ABBREVIATIONS

Bgrc, gonarcal bridge hyv, hypovalva cc, callus cerci pr, processus epr, entoprocessus psp, pseudopenis gcr, gonocristae TA, transverse arch grc, gonarcus

13 2

cc \ ______TA

--·-·· psp ------hyv 3

·-·-----·-·-·-----·-·-·-·-·grc ------·-· I ~------TA------I ..... ___ epr ·-·-·-psp ·-·---·-·---- 5

-·-·-·-·-·-·-·-·-·-·-·-·-·-·-grc------

------TA----·-·-·-·-·

7 -·-·-psp ·-·-·-·-·-·-·------8

PLATE I Fig-. 1. Left meta.thoracic wing of C. carnea. Fig. 2. Left matathoracic wing of C. harrisii. Fig. 3. Lateral view of the terminal portion of the male abdomen of C. carnea. Fig. 4. Dorsal view of pseudopenis of C. carnea. Fig. 5. Lateral view of genital armature of C. carnea. Fig. 6. Dorsal view of same. Fig. 7. Lateral view of genital armature of C. rufi,labris. Fig. 8. Dorsal view of same.

14 /-- .. epr ~-/ ,, ------TA ------n -·-·-·-·-·psp -·-·-·- ·-·-·-· ·-·------l--~-- 9 10

-----hyv ----gcr

11 12

psp --·-·-·-·-·-·-·-·-·-•4

------·-----· grc------

.------epr------.

psp \ \ \ 13 \ 14 \ Bgrc------~

PLATE II

Fig. 9. Lateral view of genital armature of C. harrisii. Fig. 10. Dorsal view of same. Fig. 11. Dorsal view of pseudopenis of C. harrisii. Fig. 12. L:: teral view of the terminal portion of the male abdomen of C. oculata. Fig. 13. Lateral view of genital armature of C. oculata. Fig. 14. Dorsal view of same.

15 -·-·-·-·----grc

epr ----

------. ._ ...... ---·---·---- psp

Bgrc ---·--- 16 psp ·-·-·-·-

r~--- ... ' \ ----- I' I\ ·-·-·-grc --·-· / // \ / ,~ \ ______,' ,/ /

-----epr I I I I ·, 17 I' psp ____ :~ Bgrc----···· I

psp ---. ___ _ ...... _ ·------·-·-grc ·-·-·---.

-----·-·------·-·-----·-- psp

epr ------¼ ' 1 \ 19 \ Bgrc ---·i 20

PLATE III

Fig. 15. Lateral view of genital armature of C. in com pleta. Fig. 16. Dorsal view of same. Fig. 17. Lateral view of genital armature of C. chi. Fig. 18. Dorsal view of same. Fig. 19. Lateral view of genital armature of C. quadr£punctata.

16 ·------...... __ ___ ·------grc------

----r•, ': I '· ! \/ \/ : : ------. '·'epr ... ______I\\ \\- z;'1 , I ------. psp ------~ '\ 21 Bgrc----~, 22

--pr

·----hyv 23

-·-·····-·------grc------·

24 25

PLATE IV

Fig. 21. Lateral view of genital armature of C. nivicornis. Fig. 22. Dorsal view of same. Fig. 23. Lateral view of the terminal portion of the male abdomen of C. lineaticornis. Fig. 24. Lateral view of genital armature of C. lineaticornis. Fig. 25. Dorsal view of same.

17 PLATE V

Fig. 26. Left gonocrista of C. oculata. Fig. 27. Left gonocrista of C. incompleta. Fig. 28. Left gonocrista of C. chi. Fig. 29. Left gonocrista and broad connective band of teeth of C. quadripunctof(I. Fig. 30. Gonocris.tae of C. niRricornis. Fig. 31. Gonocristae of C. lineaticornis.

18 Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 1653

Reference Citation: Bram, R. A.; Bickley, W. E. 1963 [1963.??.??]. The green lacewings of the genus Chrysopa in Maryland (Neuroptera: Chrysopidae). University of Maryland Agricultural Experiment Station Bulletin A-124:ii + 1-18. 5 paginated plates [#I-V] with 31 figures [#1-31]. [BotN ref#1653]

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File: File produced for the Bibliography of the Neuropterida (BotN) component of the Lacewing Digital Library (LDL) Project, 2018.