Taxonomy of United States Leucochrysa (Neuroptera: Chrysopidae).
Psyche 84(1):92-102, 1977.
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TAXONOMY OF UNITED STATES LEUCOCHRYSA
BY PHILLIP A. ADAMS
Department of Biological Sciences
California State University
Fullerton, California 92634
This paper is one of a projected series dealing with the taxonomy and identification of United States Chrysopidae. Both of our spe- cies of Leucochrysa prove to range well into the tropics, where the genus is diverse and abundant.
Bickley and MacLeod 1956 discuss at length the history and pos- sible validity of Allochrysa. Banks 1903 separated Allochrysa from Leucochrysa on the basis of its having a quadrangular rather than a triangular intramedian cell, despite the fact that L. varia, the type species of the genus, also has a quadrangular cell. This oversight was rectified by Navas 1917, who correctly synonymized Allochrysa with Leucochrysa, and erected the genus Nodita for the species with a triangular intramedian cell, previously placed in Leucochrysa by Banks. Banks followed Navas' usage for tropical species, but inexplicably continued referring all the U. S. species to Allochrysa. Bickley and MacLeod took a non-conimittal stand, justifying re- tention of Allochrysa by suggesting that characters might someday be found which show that these genera are distinct. My examina- tion of many neotropical species has not turned up any such char- acters. Banks himself referred the Antillean population of L. in- sularis to Leucochrysa, and the United States population of the very same species to Allochrysa. There appears to be no basis whatever for the continued recognition of Allochrysa. It may be superfluous to point out that since L. insularis is the type species of both Protochrysopa Kolbe 1888 (by monotypy) and Allochrysa Banks 1903 (L. virginica, by original designation), these genera are synonymous.
In the tropics, the genus Nodita merges with Leucochrysa. Thus far, the only characters useful for its separation are the venational ones discussed by Banks 1945, which have proved variable and unreliable in many cases, as Banks himself was aware. An example "Manuscript received by the editor June 5, 1977 92
19771 Adams - Leucochrysa 9 3
is Leucochrysa risi Navas, in which many specimens have an intra- median cell resembling that of Nodita; this species could be assigned to either genus equally well. The case of L. negata is comparable (see below). Species of Nodita and Leucochrysa from the United States, however, can be separated with fair reliability by the form of the intramedian cell.
Occasional aberrant speciniens of other genera with a quadran- gular intramedian cell may mistakenly be referred to Leucochrysa if other characters are not considered. This is exemplified by Al- lochrysa parvula Banks 1903: 143, the unique type of which is Chrysopa lineaticornis Fitch: No data, (Runnyniede, Fla., accord- ing to description) male, MCZ 11405, "C. columbiana, det. E. G. MacLeod" (new synonymy).
The genitalia of both sexes of Leucochrysa are fully illustrated here for the first time. In the niale, the eighth sternite is more or less distinctly demarked from the ninth; sternites except ninth usu- ally with niicrotholi. Tignum, gonapsis and gonocristae are absent. The mediuncus (=arcessus) usually bears a small curved median tooth or hook flanked by notches (Fig. 10, 1 l), no entoprocesses (gonocoxites), In the female, subgenitale small, or entire area pos- teriorly to seventh sternite broadly sclerotized (insularis, arizonica, singularis), ventral pit sonietinies far anteriorly (insularis, arizon- ica) or on a separate sclerite (internata). Spermatheca pillbox-shaped as in Chrysopa, or more frequently elongate and bent (internata, dolichocera), sperniathecal duct short (insularis, arizonica) to ex- tremely elongate (internata, angrandi), bursal duct sometimes elab- orated (magnifica), two bursal glands.
The female genitalia, although tedious to prepare for exan~ina- tion, are surprisingly diverse, offering excellent taxonomic char- acters. It is imperative that preparations of critical speciniens retain the copulatory bursa with its glands, ducts, and connection to the sperniatheca intact. Removal of the sperniatheca destroys the as- sociated structures, and should be avoided if at all possible. Leucochrysa colombia Banks
Allochrysa colombia Banks 1910: 150. A specimen from "Sta. Margarita, W. Colombia, July, 2700 m" MCZ No. 11999 (not dissected) is designated lectotype.. A "cotype" female from Canon del Monte Tolima, Colombia, 1700 rn, in the BMNH is designated a paratype. Leucochrysa colombia, Banks, 1944: 32.
94 Psyche [March
Leucochrysa claveria Navas 1927. Banks 1944: 135 (synonymy). Although the type could not be located in the Navas collection in 1974, the description is sufficiently con~plete to indicate Banks' action is probably correct. Leucochrysa californica Navas 1928: 235, new synonymy. Holotype: "California", Riksrnuseum, Stockholm, female. Allochrysa californica, Banks 1938: 122. Allochrysa virginica (incorrectly), Bickley and MacLeod, 1956: 184. The identity of L. calijiornica has long been a puzzle, as Navas' description does not fit any known species from the United States. The specimen proved to be heavily plastered with moth scales, accounting for his inconiplete description of the head markings. The printed label "California" dates froni the late nineteenth cen- tury (Per Inge Persson, pers. comm.), and perhaps replaced an original handwritten label, now lost, such as "Col" or perhaps even "Cali". As this species is otherwise known only froni Colombia, we may presume that the type originated there also. Description. Head (Fig. 4) pale, labruni, clypeus and frons suf- fused with wine red; red genal stripe present; on vertex a dark blackish red V-mark bordering the antenna1 fossae, jointed pos- teriorly by a pair of slender transverse red marks; membrane of fossae pink or red suffused. Scapes dorsally suffused with blackish red, bearing medial black stripe; pedicel black ringed, short an- terior blackish red, flagellar stripe. Pronoturn pale yellow green, anterior corners pink suffused, occasionally with two small brown spots. Abdominal tergites 6 and 7 heavily black-marked. In the forewing, the inner gradate series extends far distally, intersecting all but one to three "intern~ediates" (branches of Rs extending from Rs to pseudoniedia, and appearing almost as crossveins). Costal area broad, tallest cell 4.3 tinies as tall as wide, niarginal fork op- posite outer psm crossvein, 5 tinies as long as wide. Dark spot surrounding the outer psni crossvein, and 6 or 7 crossveins near the center of the outer gradate series are dark, narrowly brown bordered. Forewing 23.5 nim long. In female, subgenitale (Fig. 1) narrow, bearing a sniall pit on the inner anterior surface. Copu- latory bursa (Fig. 2) sniall, bursal duct delicate, moderately long, two bursal gland ducts; sperniatheca pillbox-shaped, duct short (Fig. 3).
This species is easily recognized by the inner gradate series ex- tending far basally, broad wings with long costal and niarginal cells, and head markings.
Adams - Leucochrysa
Leucochrysa arizonica (Banks)
Figures 5-9, 17- 1 8
Allochrysa arizonica Banks 1906: 98. Holotype: Palmerlee, Ark, July, male, M.C.Z. No. 11403 (not dissected). Banks 1938: 122; Bickley and MacLeod 1956: 184. Head, palpi, antennae pale except red marks on gena, vertex, pronotum, niesoprescutum, as in Figure 9. Wings pale, forewing with short red-black marks on middle of several costals, and on anals, pseudocubitals, niarginal forks behind pseudocubitus, pseu- domedials, basal radials and branches of Rs. First two medial crossveins wholly red, gradates brown. Membrane suffused with brown at basal inner gradate, and at distal pseudoniedial crossvein; no brown spot at base of stigma. Height of tallest c-cells 3.0 times width, 19-20 radials, 1 1-12 inner gradates, series follows psni basad, proximal inner gradate nearly perpendicular to and ending on psm; 10-1 1 outer gradates, marginal fork nearest last pseudomedial cross- vein 5.6 times as long as wide; 10 apparent pseudomedial cross- veins, distal crossvein oriented as an extra outer gradate. Hind wing veins pale, dark spot at base of stigma. Male: sternites 2-8 with niicrotholi (Fig. 5); sternite 9 bears a pair of denticulate forcipate processes; a field of small niicrotrichia posteriorly. Ectoprocts notched, ventral lobe hairless; callus cerci black posteriorly. Gonarcus (Fig. 7-8) heavily sclerotized, 3 thin plates form low hood above bluntly hooklike niediuncus; gonosetae sparse, small.
Female: Subgenitale (Fig. 17, 18) as broad as seventh sternite, conspicuously exposed even in dried material, shiny red-brown, laterally bearing downturned angular process; apical pit deep, bor- dered by two thin nearly vertical ridges. Dorsally to the subgenitale, membrane surrounding oviducal opening and forming floor of cop- ulatory bursa expanded and tanned. Spermatheca pillbox shaped. Copulatory bursa niuch as in L. insularis. Specimens examined: Arizona: Santa Rita Mts., Madera Can., 12- 13-VI- 1968, female, Menke and Flint, USNM. Mexico: Mi- choacan, Jct Hwy 4 and Huetanio Rd., 15 mi. E. Morelia, 8-VII- 1947, male, T. H. Hubbell (Univ. of Michigan); Jalisco, Ajijic, 16-18-VII-1966, Flint and Ortiz, 6 specimens (USNM and PAA). This is a niuch more robust insect than L. insularis, easily rec- ognized by the color markings of the head, and unusual external
L. colombia (holotype of L. californica). Fig. 1, subgenitale, ventral view.
Fig. 2, copulatory bursa and spermatheca, left lateral view. Fig. 3, sper- matheca, ventral view.
Fig. 4, head and pronotum. Figures 5-9, L. arizonica (Michoacan. Mexico). Fig. 5, Male abdomen, lateral. Fig. 6, apex of ninth sternite, ventral. Fig. 7, gonarcus and mediuncus, lateral. Fig. 8, same, dorsal. Fig. 9, head and thoracic markings. Abbreviations used in Figs. 1-9: bd - bursal duct, bg - bursal gland, gs - gonarcus, h - hood, mu - mediuncus, spd - spermathecal duct.
19771 Adams - Leucochrysa 97
genitalia in both sexes. Not previously reported from Mexico, it now appears to be a tropical species, Arizona representing the prob- able northern range boundary. L. negata (Navas) 19 13: 3 16 is very similar, but the gonarcus hood is much more developed, extending almost to cover the mediuncus viewed dorsally; apical process of ninth sternite separated by a distinct v-cleft and with larger teeth, pronoturn and head more lightly marked. I suspect that when more material is available, it will prove to be a geographic or develop- mental variant of L. arizonica. This species was described from "Guatemala: Amula, Guerrero, 6000 ft, Aug., H. H. Smith." The only Guerrero listed for Guatemala is at 15'29'N, 88'35'W, at an altitude of less than 2000 ft. It seems probable that L. negata is from Guerrero, Mexico, which is much closer to localities for L. arizonica.
The holotype of L. singularis Navas 19 13: 3 16 was collected si- multaneously with that of L. negata; both are identical but for the structure of the intramedian cell, the specimen with a quadrangular cell being named Allochrysa [now Leucochrysa] negata, and the other, with a triangular cell, Leucochrysa [now Nodita] singuZaris (New synonymy; L. negata has precedence). Navas' overlooking the identity of these two species, while describing them one after another, seems quite in character.
Leucochrysa insularis (Walker)
Figures 10- 16
Chrysopa insularis Walker 1853: 269. Holotype: "Janiaica/ insularis", male, British Museum (Natural History) (examined). Protochrysopa insularis Kolbe 1888: 74. Chr.ysopa virginica Fitch 1856: 91. New synonyn1y. Holotype (not seen, probably lost): Cartersville, Va. Con~parison of a male from Virginia with the holotype of L. insularis revealed no significant differences. Nothochrysa virginica, Banks 1895: 3 15. Allochrysa virginica, Banks 1903: 143, Bickley and MacLeod 1956: 184. Nozhochrysa phantasms MacGillivray 1894: 170. Six cotypes are present in the MCZ, a male from "West Chop Mass., Aug. 8, 1893, MCZ Type 10479" is designated lectotype. Banks 1895: 3 15. Leucochrysa cerverai Navas 1923: 325. New synonynly. Type not found. A long series from the type locality, Santiago de las Vegas, Habana, Cuba, F. Z. Cer- Vera, Navhs det., is in the MCZ and there is no doubt as to the identity of this species. The synonymy with L. insularis was suggested by Alayo 1968: 57. Lxucochrysa joannisi Navas 1925: 13, "Santiago de las Vegas, Cuba, July 17, Aug. 20, 1924, F. Cervera". No type specimen designated, and no type found in the Navas collection, 1974. Alayo 1968: 57 (synonymy). There seems to have been no fornial description published of this species; it is the color variant of
98 Psyche [March
insularis with brown instead of pale lateral thoracic markings. Allochrysa virginica ocala Banks 1938: 122. Type: "Lloyd Sink, Jefferson Co., Fla., G. Fairchild coll., Aug. 9, 1935", female, MCZ No. 23184 (exan~ined). Maculation of this species shows geographic as well as individual variation. Specimens froni Florida, Alabama, and the Antilles have a red or brown V niark on vertex bordering antenna1 fossae and genae are red marked (L. insularis ocala). In Georgia and Alabania the V niark, if present, is faint. North of Georgia, the vertex marks are absent and genae are brown niarked or pale. The niesonotuni bsars two brown spots on the prescutal-scutal suture on lightly niarked speciniens; on heavily niarked exaniples ("L. joannisi") nearly the entire pteronotuni is brown or black. The holotype of L. insularis has niost of the transverse veins in the forewing dark marked. In Florida specimens, the inner gradates are paler, costals pale; dark transverse veins include outer gradates (except apical 3 or 41, ends of proxinial r's, 1-3 m, the cubitals, and ends of the anals. The gradates are often bordered by a dark streak. North of Flor- ida, transverse veins are paler.
Male genitalia. Sterna except ninth with n~icrotholi; apex of ninth sternuni slightly notched with a sniall field of lanceolate gono- cristae each side (Fig. 10). Mediuncus with median sclerotized band and hook, confluent with sen~iniembranous lateral lobes? gonosac- cus without setae (Fig. 11). Laterally to gonarcus a pair of delicate digitiforni nienibranous sacs (Fig. lo), ventrally a pair of lightly sclerotized lobes connected by a nienibranous flap (shown with- drawn in Fig. 10). A nienibranous lobe between gonopore and ninth sternite.
Feniale genitalia. Su bgenitale (Fig. I 6) broadly sclerotized, ex- panded anteriorly as pit-bearing lobe adjacent to seventh sternite. Bursa1 gland ducts very long, unbranched, bursa1 duct inconspicu- ous.
The forni of the niediuncus is typical for a Leucochr~vsa, but the ventral sclerotized lobes are highly distinctive. Distribution. Coastal states froni Massachusetts to Florida, Puerto Rice? Cuba and Janiaica, also West Virginia, Tennessee (Bickley and MacLeod 1956), Alabania, Mississippi, Arkansas, and Missouri. Range extensions: Ala.: Wilson Dani, F. Q. 9-VIII-1941, J. N. Belkin, LAM. Miss.: Clinton, Hinds Co., 20-V-1960, Bryant Mather, USNM. Ark.: Devil's Den St. Pk., Wash. Co. 12-VI- 1966, R. W. Hodges, USNM. Mo.: Colunibia, Malaise trap, 7
Adams - Leucochrysa
Figures 10-1 6.
I!,. insularis. Fig. 10, male abdomen, lateral, sclerotized lobes withdrawn. Fig. 11, gonarcus, n~ediuncus and sclerotized lobes, dorsal view. Fig. 13, hypandriun~ internun~. Fig. 14, spern~atheca, dorsolateral. Fig. 15, female ab- domen, showing broadly sclerotized subgenitale, and internal structures. Fig. 16, subgenitale, ventral view. Figures 17-18, L. arizonica. Fig. 17, apex of seventh sternite and subgenitale, ventral. Fig. 18, same, lateral. Abbreviations used in Figs. 10- 16: bc - copulatory bursa, bg - bursa1 glands, nls - n~en~branous sac, sg - subgenitale, s1 - sclerotized lobes.
100 Psyche [March
a.n~.-7 p.nl., 22-IX-1967, F. D. Parker, USNM; 5 nii. S. Joplin, 25-VI-1968, E. L. Todd, USNM.
Gumilla longicornis (Walker)
Osmylus longieornis Walker 1853: 235. Holotype: "Georgia," "Type, Osmylus longi- cornis Walker, det, D. E. Kin~n~ins", "Gumilla longicornis Walk., Long. Naviis det," BMNH.
Meleoma longieornis, Hagen 186 I : 2 10. Lmcochrysa longicornis, Banks 1907: 26.
Gumilla ? longicornis, Navas 19 12: 189, Kruger 19 13: 22 I. Allochrysa longicornis, Bickley and MacLeod 1956: 184. Not A. longieornis (Gray), Banks 1920: 339, as cited by Bickley and MacLeod 1956: 184. This insect is not a chrysopid, but an osn~ylid in which the ocelli are lost, and the antennae are exceedingly elongate. Hagen's re- ferral of G. longicornis to the chrysopid genus Meleoma seems to have been based solely upon the nature of the antennae. Navhs' illustration of the wing characters is reasonably accurate. Addi- tional features are: pronotuni articulating above n~esothoracic spi- racle, claws simple, single aroliar pad, two nygnlata between RS- MA and MP in forewing. In hindwing, basal piece of MA ("sinu- ate crossvein") absent, steni of MP runs close to R; MP2 fused briefly with and appearing to be a continuation of GuA; CUP un- branched, clearly separate from 1A for its entire length. Wing n~en~brane n~icrotrichiated, marginal vein entire except for a few marginal dots near wing apex.
The only other species of this genus, Gumilla aspersus Navas 19 12: 189, known from a single male specimen from Brazil (Vienna Museun~, not seen) is probably the same; Navas' figure of the fore- wing does not differ in any in~portant respect from that of G. longi- cornis. Two additional specinlens from Brazil, Langsdorf, are in the Berlin Museun~. There thus seems little question that the type of G. longicornis is also from Brazil, not Georgia. The following have graciously made material available for study: peter Barnard - British Museun~, Natural History (BMNH), 0. S. Flint - U. S. National Museuni (USNM), K. K. Gunther - Mu- seum fur Naturkunde, Berlin, Charles Hogue - Los Angeles County
19771 Adams - Leucochrysa 10 1
Museun~ of Natural History (LAM), T. H. Hubbell - Univ. of Michigan, John Lawrence - Museunl of Conlparative Zoology, Harvard (MCZ), Per Inge Persson - Riksniuseun~, Stockholnl. ALAYO, P.
1968. Los Neuropteros de Cuba. Poeyana Ser. B(2): 1-127. BANKS, N.
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New Chrysopidae and species new to the United States. Canad. Enton~. 70: 118-122.
Neuroptera of Northern South An~erica. Part 3. Chrysopidae. Boletin de Enton~ologia Venezolana 3: 1-34.
A review of the Chrysopidae (Nothochrysidae) of Central America. Psyche 52: 139-1 74.
BICKLEY, W. AND E. MACLEOD.
1956. A synopsis of the nearctic Chrysopidae, with a key to the genera (Neu- roptera). Proc. Enton~. SOC. Washington 58: 177-202. FITCH, A.
1856. First and second reports on the noxious, beneficial, and other insects of the State of New York. Albany, 336 pp. GRAY, G.
In Griffith: Animal kingdom 15: 33 1, pl. 72, Fig. 3 (not seen). HAGEN, H.
1861. Synopsis of the Neuroptera of North America. Sn~ithsonian Misc. Coll. 347 pp.
Die geographische Verbreitung der Neuroptera und Pseudoneuroptera der Antillen. Arch. f. Naturgesch. 54 Bd. I, Hft. 2: 153-178, pl. 13. KRUGER, L.
I9 13. Osn~ylidae. IIIa. Nachtrag zur Literatur und Katalog. Stettiner ento- n~ologische Zeitung 74: 2 18-224.
Insectos neuropteros nuevos o poco conocidos. Men~orias de la Real Academia de Ciencias y Artes de Barcelona Ser. 111, 10: 135-202. 1913.
Les chrysopides (Ins. Nkvr.) du Muske de Londres. Annales de la So- cietk scientifique de Bruxelles 37: 292-330.
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Insecta nova, series XII. Men?. Pontif. Accad. Nuovi Lincei Ser. 11, 10: 1-10.
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MACGILLIVRAY, A. D.
New species of Nothochrysa. Canad. Enton?. 26: 169- 17 1. WALKER, F.
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