1869 THE LIBRARY 1. ‘J t I ' I ' X i ' * ■ - iiw t ■W*^ 4' I I I 6^7 ) - Vol. LXXXVII MARCH 1979 of the New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General CX2a The New York Entomological Society Incorporating The Brooklyn Entomological Society Incorporated May 21, 1968 The New York Entomological Society Organized June 29, 1892 — Incorporated February 25, 1893 Reincorporated February 17, 1943 The Brooklyn Entomological Society Founded in 1872 — Incorporated in 1885 Reincorporated February 10, 1936 The meetings of the Society are held on the third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St. & Central Park W., New York, N. Y. 10024. Annual dues for Active Members; including subscription to the Journal, $15.00. Members of the Society will please remit their annual dues, payable before January 15th, to the Treasurer. Officers for the Year 1979 President, Dr. Louis D. Trombetta Isaac Albert Research Institute, Brooklyn, N.Y. 11203 ‘ Vice-president , Dr. Betty Faber I American Museum of Natural History, New York 10024 Secretary, Dr. Joseph Cerreta Columbia University, New York 10032 ^ Assistant Secretary, Dr. Gerard Iwantsch -j Fordham University, New York 10458 j Treasurer , Dr. Ivan Huber Fairleigh Dickinson University, Madison, New Jersey 07940 Assistant Treasurer, Mr. Kenneth Sherman Rutgers University, New Brunswick, New Jersey 08903 Trustees Class of 1979 Dr. Peter Moller Dr. Howard Topoff Class of 1980 Mr. Louis Sorkin Mr. Charles Calmbacher Publication Business Manager Mrs. Irene Matejko Fordham University, New York 10458 Mailed April 26, 1979 The Journal of the New York Entomological Society (ISSN (X)2S-7199) is published quarterly for the Society by Allen Press Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New Brunswick, New Jersey and at addi- tional mailing office. Known office of publication; Waksman Institute of Microbiology, New Brunswick. New Jersey 08903. Journal of the N.Y. Entomological Society, total No. copies printed 750. Paid circulation 490. mail subscription 470. free distribution by mail 23, total distribution 493. left-over 257 copies each quarter. Journal of the New York Entomological Society VOLUME LXXXVII MARCH 1979 NO . 1 EDITORIAL BOARD Editor Dr. Karl Maramorosch Waksman Institute of Microbiology Rutgers University New Brunswick, New Jersey 08903 Associate Editors Dr. Lois J. Keller, RSM Dr. Herbert T. Streu Publication Committee Dr. Randall T. Schuh American Museum of Natural History Dr. Daniel Sullivan Fordham University Dr. Felix J. Bocchino The College of Mt. Saint Vincent CONTENTS The taxonomic position of three North-West Indian species commonly referred to the genus Schrank (Lepidoptera: Pyralidae) H. S. Rose and H. R. Pajni 2-8 Effect of low host density on oviposition by larval parasitoids of the alfalfa weevil Robert V. Dowell 9-14 Mite predators in Eastern New York commerical apple orchards R. W. Weires and G. L. Smith 15-20 Strategies of gall formation in Pemphigus aphids Daniel P. Faith 21-37 European katydid Meconema thalassinum (De Geer) recorded from new location on Long Island, New York (Orthoptera: Tettigonidae) Burke Smith, Jr. 38^1 A new species of Megaris and the status of the Megarididae McAtee & Malloch and Canopidae Amyot & Serville (Hemiptera: Pentatomoidea) F. J. D. McDonald 42-54 Autographa californica nuclear polyhedrosis virus (NPV) in a vertebrate cell line: localization by electron microscopy Arthur H. McIntosh, Karl Maramorosch and Russell Riscoe 55-58 Two new species of Laccobius from eastern North America (Coleoptera: Hydro- philidae) Stanley E. Malcolm 59-65 The evolution of eyespots in tropical butterflies in response to feeding on rotting fruit: an hypothesis Allen M. Young 66-77 Nest of the wasp Clypearia weyrauchi (Hymenoptera, Vespidae) Robert L. Jeanne 78-84 Effectiveness of native parasites against Agromyza frontella (Rondani) (Diptera: Agromyzidae), an introduced pest of alfalfa R. M. Hendrickson, Jr., and S. E. Barth 85-90 Possible phylogenetic significance of complex hairs in bees and ants U. N. Lanham 91-94 Book reviews 95-96 NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 2-8 THE TAXONOMIC POSITION OF THREE NORTH-WEST INDIAN SPECIES COMMONLY REFERRED TO THE GENUS PYRAUSTA SCHRANK (LEPIDOPTERA: PYRALIDAE) H. S. Rose and H. R. Pajni Abstract. — Three species of subfamily Pyraustinae referable to the un- revised genus Pyraiista Schrank were collected from North-West India be- tween November 1972 and November 1974. A critical examination of these species leads to the erection of a new genus Rattana with type species P. euryphaea Meyrick and the description of a new species, Coclebotys mu- tuuri. The characterization of the new genus and the description of the new species are recorded. Introduction The authors collected ninety two species of subfamily Pyraustinae from North-West India between November 1972 to November 1974. Out of these, three species, as followed through Hampsonian key (1896, 1898), are referable to the old Pyrausta Schrank. This genus, however, has been re- cently found to contain very heterogeneous material and has been divided into two sections, with and without a frenulum hook, restricting genus Py- rausta to the section with a frenulum hook and with P. cingulata (Linnaeus) as its type species. Munroe (1950, 1958, 1958a) and Munroe and Mutuura (1968c, 1969a, 1969c, 1970a, 1971) have revised the other section and erected several new genera based on other species. One of three species in our possession, machoeralis Walker is referable to the genus Pyrausta, as restricted by Munroe (1950). The second species Abbreviations I A, First anal vein; 2A, Second anal vein; 3A, third anal vein; ANT.APO, Anterior apophyses; CO. Costa; CRN, Cornuti; CRP.BU, Corpus bursae; Cu,. First Cubital vein; Cuj, Second cubital vein; DU.BU, Ductus bursae; JX, Juxta; M,, First median vein; Mj, Second median vein; M3, Third median vein; PO.APO, Posterior apophyses; R,. First radial vein; Rj, Second radial vein; R3, Third radial vein; R4. Fourth radial vein; R5, Fifth radial vein; Rs. Radial sector; SA, Saccus; Sc, Subcosta; Sc + R,. Stalk of Sc and R,; SIG, Signum; SL, Sacculus; SSCA, Subscaphium; TG. Tegmen; TRAH, Half transtilla; TU.A, Tuba analis; UN, Uncus; VIN, Vinculum; VLV, Valva. Figs. 1-7. Rattana euryphaea, 1. photograph of adult; 2. forewing; 3. hindwing; 4-6. different parts of male genitalia; 7. female genitalia. 1 mm VOLUME LXXXVII, NUMBER 1 3 1 4 mm 4 NEW YORK ENTOMOLOGICAL SOCIETY is clearly congeneric with coclesalis Walker the type species of the mono- typic genus Coclebotys (Munroe and Mutuura, 1969c) but is distinctly dif- ferent from the former and also from all the species described under Py- rausta Schrank, and hence a new species. The third species Pyrausta euryphaea Meyrick is also referable to the section lacking the frenulum hook but it does not go under any of the genera erected or revalidated by Munroe (1950, 1958, 1958a) and Munroe and Mutuura (1968, 1968a, 1968b, 1968c, 1969, 1969a, 1969b, 1969c, 1970, 1970a, 1971, 1971a). Accordingly a new genus Rattana with Pyrausta euryphaea Meyrick (new name for Botys signatalis Walker, preoccupied) as its type species is proposed. The diag- nosis of the genus Rattana and a complete description of Coclebotys mu- tuuri, n. sp., are here presented. The terminology for different parts of gen- italia has been adopted from Klots (1970). Rattana new genus (Figs. 1-7) Type species: Pyrausta euryphaea Meyrick Exot. Micr., 4, p. 318 (1932). Labial palpus porrect and straight; third joint covered with scales from second segment. Maxillary palpus strongly dilated with scales. Frons round- ed. Antenna simple, shorter than fore wing. Fore wing with vein R, arising from very well before anterior angle of cell; R2 approximated to R3+4; R5 curved and approximated to R3+4; Mj, M3 and Cu, from posterior angle of cell. Hind wing with outer margin not excised below apex; discal cell one- third the length of cell; Rs anastomosing with Sc + Rj beyond cell for some distance; Mg and M3 closely approximated at base for some distance; Cu, from lower angle of cell; frenulum hook absent. Male genitalia. — Uncus very long, strongly curved, with basal half narrow and distal half dilated and spoon-shaped, dorso-distal and ventro-distal sur- faces fringed with setae, extreme tip naked; gnathos absent; tuba analis much shorter than uncus; scaphium not developed; subscaphium strongly sclerotized, supporting tuba analis throughout its length; tegumen arched, roughly as broad as long and strongly sclerotized; vinculum produced an- teriorly into a well sclerotized saccus; saccus short. Valva long, the costal margin greatly arched and the saccular margin straight, rounded distally; costa broadly inflated, supported by thin sclerotized lines; sacculus con- spicuous; harpe missing. Transtilla with each half sharply triangular, being Figs. 8-14. Coclebotys mutuuri n. sp.. 8. photograph of adult; 9. forewing; 10. hindwing; 11-13. different parts of male genitalia; 14. female genitalia. VOLUME LXXXVII, NUMBER 1 5 6 NEW YORK ENTOMOLOGICAL SOCIETY broad at costa and pointed at apex, weakly sclerotized; juxta more or less rectangular. Aedeagus long, broad at anterior end and slightly narrow pos- teriorly; vesica armed with two or three spinose patches. Female genitalia. — Corpus bursae exceptionally reduced, more or less ir- regular in outline and completely membranous; signum absent; ductus bur- sae quite long, with a membranous dilation near anterior end and posterior portion sclerotized; anterior apophyses long, each with a triangular thick- ening near base, base sharply pointed; posterior apophyses short. Ovipos- itor lobes narrow, setose with long and short setae. Coclebotys mutuuri n. sp. (Figs. 8-14) Holotype. — 9, Uttar Pradesh: Dehra Dun, 21st September, 1973 (H. S. Rose), from fluorescent tube. Allotype. — S , same data. Holotype pinned and allotype slides (i.e. wings and gentalia) in Zoological Museum, Entomological Section, Department of Zoology, Panjab University, Chandigarh (India). Head. — Vertex covered with light ochreous scales and with a whitish tinge; frons oblique and somewhat flattened, smoothly scaled with ochreous scales, scales along inner margins of eyes white. Antenna filiform, shorter than fore wing; scape covered with light brown scales; flagellum with weak annulations on upper surface, the lower almost naked, slightly compressed in male. Eye large, with a row of ochreous scales behind. Ocellus well developed. Labial palpus porrect, exceeding head by less than length of latter; first segment and a small basal portion of second segment on ventral surface covered with white scales, the remainder being fulvous scaled; base of third segment hidden by scales of second segment. Maxillary palpus prominent, strongly dilated with brownish scales distally. Proboscis large, covered with white scales at base. Posterior end of head adorned with long and erect ochreous brown scales, the latter surrounding the white ones. Thorax. — Ochreous brown dorsally; white ventrally. Forewing. — Costal margin curved near apex; apex rounded and narrowly acute; termen oblique and somewhat curved; tornus obtuse; anal margin curved near base. Ground color ochreous brown, with costal and outer area slightly darker; a wavy antemedial line from costa to inner margin; a discocellular speck present; a postmedial line from anterior to posterior margin, strongly excurved between M, and Cu,; marginal fringe ochreous brown. Discal cell slightly less than half the length of wing. Rj from before anterior angle of cell; R2 from upper angle of cell, approximated to R3+4; R3 VOLUME LXXXVII, NUMBER 1 7 and R4 stalked; R5 not approximated to R3+4; M, from somewhat behind upper angle. M2, M3 and Cu, from lower angle of cell, fairly spaced around the angle; CU2 from cell at two-third the length of cell; 3A making loop with 2 A at base. Hindwing. — Anterior margin straight; apex, termen and tornus rounded. Ground color ochreous brown; outer-marginal area with fuscous suffusion; a postmedial line from Rs to vein Cu, excurved between M2 and CU2; mar- ginal fringe ochreous brown. Discal cell less than half the length of wing; discocellular straight and oblique; cell closed. M2, M3 and Cu, from posterior angle of cell; M2 and M3 closely approximated at base; CU2 from cell at two- third the length of cell; three anals present. Legs. — Clothed with white scales; tibia of prothoracic leg covered with fuscous scales; outer anterior and posterior spurs of male hind tibia minute; outer spur of mid tibia of male and all outer spurs of female about two-third the length of inner spurs. Abdomen. — Ochreous brown dorsally; pure white ventrally. Male genitalia. — Uncus relatively reduced, somewhat triangular and round- ed at extreme end, heavily setose with anteriorly directed setae; gnathos present; tuba analis longer than uncus; scaphium not developed; subsca- phium thin strap-like; tegumen parallel sided posteriorly and well sclero- tized; vinculum produced anteriorly into a short saccus. Valva long and of moderate width, costal and saccular margins almost parallel, tip unsym- metrically rounded; costa weakly inflated; sacculus differentiated and car- rying a rounded setose lobe, the latter partly underlying the basal part of harpe; harpe represented by a setose lobe, bearing four dorsally directed scale-like strong setae. Transtilla relatively reduced, with each half trian- gular; juxta moderately stout, with its walls well sclerotized. Aedeagus long and slender, walls well sclerotized, with a short conical projection from a strap-like thickening of aedeagal wall; vesica with three well defined cornuti, along with strap-like thickenings at posterior end. Female genitalia. — Corpus bursae bag-like and with an irregular boundary; signum with its margin serrate, lateral angles somewhat produced and me- dial angles rounded; ductus bursae fairly long, strongly sclerotized at the proximal portion; anterior apophyses long and thickened near middle; pos- terior apophyses short and stout; ovipositor with relatively narrow lobes, each bearing macro and micro setae. Alar expanse. — Male: 28 mm. Female: 22 mm. to 27 mm. Paratypes. — 5 9 9, same data as type, collected between 1.9.1973 to 21.9.1973. (Zoological Museum, Entomological Section, Department of Zo- ology, PanJab University, Chandigarh, India.) 8 NEW YORK ENTOMOLOGICAL SOCIETY Acknowledgment The authors are much indebted to Mr. P. E. S. Whalley and Mr. M. Shaffer of the British Museum (Natural History) London, for the identifi- cation of Pyrausta euryphaea and for supplying relevant literature. Literature Cited Hampson, G. F. 1896. Fauna of British India Moths, 4: 1-594. Taylor & Francis Ltd., London. . 1898. A revision of Moths of subfamily Pyraustinae and family Pyralidae. Part 2. Proc. Zool. Soc. London, 1898:590-761. Klots, A. B. 1970. Lepidoptera in “Taxonomist’s Glossary of Genitalia in Insects” (ed. S. L. Tuxen), 2nd ed. Munksgaard, Copenhagen, pp. 115-130. Munroe, E. 1950. The generic position of some North American species commonly referred to Pyrausta Schrank (Lep. Pyralidae). Canad. Ent. 82:217-231, 28 figs. . 1958. Far Eastern Pyralidae (Lepidoptera). Canad. Ent. 90:249-254, 9 figs. . 1958a. A revision of genus Epicorsia Hiibner (Lep. Pyralidae). Canad. Ent. 90:293- 310, 10 figs. and A. Mutuura. 1968. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. I. Canad. Ent. 100:847-861, 33 figs. . 1968a. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. II. Canad. Ent. 100:861-868, 16 figs. . 1968b. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. III. Canad. Ent. 100:974-986, 35 figs. . 1968c. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. IV. Canad. Ent. 100:986-1001, 25 figs. . 1969. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. V. Canad. Ent. 101:299-305, 9 figs. . 1969a. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. VI. Canad. Ent. 101:897-906, 18 figs. . 1969b. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. VII. Canad. Ent. 101:1069-1077, 22 figs. . 1969c. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. VIII. Canad. Ent. 101:1239-1248, 16 figs. . 1970. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. IX. Canad. Ent. 102:294-304. . 1970a. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. X. Canad. Ent. 102:1489-1507. . 1971. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. XL Canad. Ent. 103:173-181. . 1971a. Contributions to a study of the Pyraustinae (Lep. Pyralidae) of Temperate East Asia. XII. Canad. Ent. 103:503-506. Department of Zoology, Panjab University, Chandigarh- 1600 14 (India). Received for publication April 6, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 9-14 EFFECT OF FOW HOST DENSITY ON OVIPOSITION BY FARVAL PARASITOIDS OF THE ALFALFA WEEVIL* I 1 [ Robert V. DowelP’^ i i Abstract. — I examined the effect of different host densities at low host: : parasitoid ratios (1: 1-6:1) on parasitism by four larval parasitoids of the I alfalfa weevil in the laboratory. The mortality inflicted was independent of I changes in host density. This is due to the short handling times (<5 sec) of I the Bathyplectes spp. and to the ability of Tetrastichus incertus (Ratzeburg) I to parasitize only 4-5 hosts/day. I Introduction I The alfalfa weevil, Hypera postica (Gyllenhal) is one of the most impor- tant pests of alfalfa in the United States. Since 1957 it has been the object 1 of a biological control effort by the U.S.D.A. and cooperating states. A I total of seven parasitoid species utilizing various developmental stages of I the weevil have been established and credited with reductions in alfalfa ' weevil numbers in the Northeast (Dysart and Day 1976). The current emphasis on pest management models for the alfalfa weevil ' system has required a greater understanding of the facts influencing host- ! parasitoid interactions and how these interactions affect the ability of the I parasitoids to stabilize host numbers (Latheef et al. 1977; Yeargan and Lath- i eef 1977). Previous studies with Bathyplectes anurus (Thompson)^ and Bathyplectes curculionis (Thompson)^ in open-choice experiments showed no relationship between the various host densities exposed (2, 4, 8, 16 and 32) and the mortality inflicted by the parasitoids (Latheef et al. 1977; Year- gan and Latheef 1976). Thus both parasitoids appear to satisfy the definition of a density-independent mortality factor (van den Bosch and Messenger 1973) and as such are incapable of stabilizing host numbers. Yet B. curcu- lionis is credited with substantial biological control of the alfalfa weevil in several sections of the United States (van den Bosch 1971; Michelbacher 1940). Here I report the results of my investigations into the relationship between ' Coleoptera: Curculionidae. ^ Submitted to the Graduate School of The Ohio State University in partial fulfillment of the requirements for the Degree of Doctor of Philosophy. Former address: Department of Ento- mology, The Ohio State University, 1735 Neil Avenue, Columbus, Ohio 43210. ’ Research partly supported by funds from the Department of Entomology, The Ohio State University. Fla. Agricultural Experiment Station Journal Series No. 989. ^ Hymenoptera: Ichneumonidae. 10 NEW YORK ENTOMOLOGICAL SOCIETY host density and parasitism by the 4 larval parasitoids of the alfalfa weevil; B. anurus, B. curculionis, Bothyplectes stenostigma (Thompson) and Te- trastichus incertus Ratzeburg.'^ In particular, I dealt with very low host: parasitoid ratios (1; 1-6:1) to determine how the parasitoids responded to changes in host density at such levels. Methods and Materials I reared host larvae from eggs of field collected adults (Dowell 1977a). Twenty-four hours prior to their being exposed to parasitoids, I placed 1- 6 host larvae on three 7-cm-long alfalfa stems in a cotton plugged 5-cm glass vial to allow feeding and the accumulation of feces. Host larvae and their feces stimulate searching behavior in both the Bathyplectes spp. (Dowell 1977, McKinney and Pass 1977) and T. incertus (Dowell and Horn 1977). The 2nd instar hosts used for the Bathyplectes spp. and the 3rd instar hosts used for 7. incertus are within the preferred host range of each parasitoid (Dowell 1977; Dysart and Day 1976). The source and care of the various parasitoids are described elsewhere (Dowell 1977; Dowell and Horn 1976). Prior to use, each female was held for 24 hours without hosts and then 24 hours with hosts to allow for ovary maturation (all species), to prevent an excessive accumulation of eggs in the lateral oviducts of the Bathyplectes spp. and to allow 7. incertus to feed on host exudates from ovipositional wounds (Dowell 1978). Each mated female was then exposed to different number of hosts daily for 2-5 successive days. Care was taken to avoid having the total number of hosts exposed exceed the expected fecundity of the parasitoid species (Dowell 1977). Host larvae were exposed to individual parasitoids in 10 x 10x9 cm plastic containers (Dowell 1977) for 24 hours in a Percival environmental chamber held at 21°C and 12 hour photophase. After use, all parasitoids were dissected to be certain that their ovaries were functional (Dowell 1976), and only those females with functional ovaries were included in the data analysis. A min- imum of four replicates/density/species were run. After exposure the host larvae were reared until pupation to determine whether they were parasit- ized. The data for each host density were averaged by parasitoid species and mortality was expressed as k-values (k-logio initial # — log,o survivors) (Varley and Gradwell 1965). The effect of host density on mortality (k-values) was examined by use of regression analysis. A fungal disease similar to that described by Dysart and Coles (1971) killed most of the larvae exposed to B. stenostigma. Results and Discussion Table 1 shows the results of the regression analysis of the k-values on host density for each parasitoid. There was no significant {P < 0.05) affect Hymenoptera: Eulophidae. VOLUME LXXXVII, NUMBER 1 II Table 1. Results of regression analyses of k-values (y) of the larval parasitoids on alfalfa weevil density (x). Parasitoid species Regress. coeffs. Intercept Standard error of estimate r t-value H„; b = 0 B. anurus -0.01 0.39 0.24 -0.05 -0.17* B. curculionis 0.04 0.14 0.18 0.31 0.05* T. incertus -0.01 0.44 0.10 -0.25 0.42* B. stenostigma 0.15 0.15 0.11 0.98 5.01t * Not significantly different from b = 0 at P = 0.05 as determined with a t-test. t Significantly different from b = 0 at P = 0.05 as determined with a t-test. However there were only 4 data points for this analysis; one replicate at each of 4 densities. j exerted by changes in host density on the mortality inflicted by B. anurus, B. curculionis or T. incertus. Only the mortality inflicted by B. stenostigmo I significantly {P > 0.05) changed with changes in host density. However, the previously mentioned disease limited this data analysis to the 1 replicate that survived at each of 4 densities. Previously I (Dowell 1977; 1978) have shown that the biology and reproductive tactics of B. stenostigma differ little from its congenors. Based upon these previous studies, I feel that B. stenostigma probably responds to changes in host density in a manner sim- ilar to its congenors and that the small number of replicates analyzed were insufficient to show this. The density-independent response of B. anurus and B. curculionis to changes in host density at these low host:parasitoid ratios agree with pre- vious studies using greater ratios (5:1-100:1) (Barnety et al. 1977; Latheef et al. 1977; Yeargan and Latheef 1977). That T. incertus also responded in a density-independent manner is surprising since its reproductive biology differs greatly from that of the Bathyplectes spp. (Dowell 1978; Dowell and Horn 1977). Recent studies have shown that the field mortality inflicted by B. anurus and B. curculionis in Ohio (Lewis 1977) and by B. curculionis and T. incertus in Ontario (Harcourt et al. 1977) is independent of changes in host density. I believe that the density-independent response of the Bathyplectes spp. and T. incertus to changes in host density have different origins but that both can be explained by the effect of several biological traits on the disc equation (1) of Holling (1959). N a aT,Np l + aThN„ (1) Where: Na = Number of hosts parasitized a = area of discovery 12 NEW YORK ENTOMOLOGICAL SOCIETY T( = exposure time Th = Handling time/host = Number of hosts available. The handling time for the Bathyplectes spp. is <5 sec/larva (Dowell 1977) making it approximately equal to zero when compared to the exposure time of 12 hr. When this happens; equation 1 can be reduced to the following: Na = aT,N„ (2) The numbers of hosts parasitized now becomes a linear function of the number of hosts available as ‘aTt’ is a constant in this equation. According to both this study and others cited herein, the slope of this linear function does not significantly differ from zero. The handling time for T. incertus is >23 min/larva (Horn 1970) and is large enough to keep equation from reducing. When the average number of hosts at each density is fitted to the disc equation by the method described by Holling (1959) and Messenger (1968) the result is equation 3. 0.68 No “ 1 + 0.07 No (3) According to equation 3, T. incertus should show a density-dependent response to changes in host numbers until all the available eggs have been laid. I believe that T. incertus is capable of parasitizing an average of only 4-5 hosts/day. This low figure is limited in two manners. The first is that egg production in T. incertus is dependent upon a continual supply of pro- teins gathered by feeding on ovipositional wounds on the host (Dowell 1978). This combined with the low number of ovarioles/ovary (n = 12) (Dowell 1978) limit the number of eggs formed/day. In addition T. incertus lays an average of 5-6 eggs/host (Streams and Fuester 1967) further restricting the number of host larvae it can parasitize on a daily basis. The final result is that a density-dependent process appears density-independent due to the limited number of hosts the parasitoid can parasitize on a daily basis. The density-independent mortality inflicted by the larval parasitoids of the alfalfa weevil, whether based upon the rapid handling time/larvae or the limited number of larvae that can be parasitized/day, make them incapable of stabilizing host numbers. While their presence does increase host mor- tality, they alone are not responsible for long-term decreases in host num- bers. Acknowledgments I thank my committee, Drs. Gordon R. Stairs, David G. Nielsen and Rodger Mitchell for their help and guidance in my work, the Entomology VOLUME LXXXVIl, NUMBER 1 13 Department of The Ohio State University for its financial support, Drs. K. V. Yeargan, M. A. Latheef and B. C. Pass for graciously allowing me access to their data and unpublished manuscripts, and John K. Fessel, Richard J. Dysart and K. V. Yeargan for supplying me with parasitoids at various times. I especially thank my advisor and friend David J. Horn for his help, understanding and loan of his cabin throughout my graduate career. Literature Cited Dowell, R. V. 1976. Non-functional ovaries in Bathyplectes spp. (Hymenoptera: Ichneumon- idae), larval parasitoids of the alfalfa weevil (Coleoptera: Curculionidae). J. N. Y. Ento- mol. Soc. 94:384-5. . 1977. Biology and intrageneric relationships of Bathyplec tes stenostigma. a parasite of the alfalfa weevil. Ann. Entomol. Soc. Amer. 70:845-8. . 1978. Ovary structure and reproductive biologies of larval parasitoids of the alfalfa weevil. Can. Entomol. 110:507-12. and D. J. Horn. 1977. Adaptive strategies of larval parasitoids of the alfalfa weevil. Ibid. 109:641-8. Dysart, R. J. and L. W. Coles. 1971. Bathyplectes stenostigma, a parasite of the alfalfa weevil in Europe. Ann. Entomol. Soc. Amer. 64:1361-67. and W. H. Day. 1976. Release and recovery of introduced parasites of the alfalfa weevil in Eastern North America. USDA Prod. Res. Rpt. No. 167, 61 p. Harcourt, D. G., J. C. Guppy and M. R. Binns. 1977. The analysis of intrageneration change in Eastern Ontario populations of the alfalfa weevil, Hypera postica (Coleoptera: Curculionidae). Can. Entomol. 109:1521-34. Holling, C. S. 1959. Some characteristics of simple types of predation and parasitism. Ibid. 91:385-98. Horn, D. J. 1970. Oviposition behavior of Tetrastichus incertus, a parasite of the alfalfa weevil. J. Econ. Entomol. 63:303-4. Latheef, M. A., K. V. Yeargan, and B. Pass. 1977. Effect of density on host-parasite inter- actions between Hypera postica (Coleoptera: Curculionidae) and Bathyplectes anunts (Hymenoptera: Ichneumonidae) Can. Ent. 109:1057-62. Lewis, D. R. 1977. Analysis of the life table of the alfalfa weevil in Ohio. Ph.D. Dissertation, The Ohio State University. Messenger, P. S. 1968. Bioclimatic studies of the aphid parasite Praon exsoletum 1. Effects of temperature on the functional response of females to varying host densities. Can. Ent. 100:728-71. McKinney, T. R. and B. C. Pass. 1977. Olfactometer studies of host seeking in Bathyplectes curcuHonis Thoms. (Hymenoptera:lchneumonidae). J. Kansas Entomol. Soc. 50:108- 112. Michelbacher, A. E. 1940. Effect of Bathyplectes curcuHonis on the alfalfa weevil population in lowland middle California. Hilgardia 3:81-99. Streams, F. A. and R. W. Fuester. 1967. Biology and distribution of Tetrastichus incertus, a parasite of the alfalfa weevil. J. Econ. Entomol. 60:1576-79. van den Bosch, R. 1971. Biological control of insects. Ann. Rev. Ecol. and Systematics 2:45- 66. and P. S. Messenger. 1973. Biological Control. In-text Educational Publishers. 180 p. Varley, G. C. and Gradwell, G. R. 1965. Interpreting winter month population changes. Proc. XII Int. Cong. Ent. (London) pp. 377-78. 14 NEW YORK ENTOMOLOGICAL SOCIETY Yeargan, K. V. and M. A. Latheef. 1976. Host-parasitoid density relationships between Hy- pera postica (Coleoptera:Curculionidae) and Bathyplectes curculionis (Hy- menoptera:Ichneumonidae). J. Kansas Entomol. Soc. 49: 551-6. University of Florida, Agricultural Research Center, 3205 S.W. 70th Ave., Ft. Lauderdale, FL 33314. Received for publication May 18, 1978. I I NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 15-20 MITE PREDATORS IN EASTERN NEW YORK COMMERCIAL APPLE ORCHARDS' I R. W. Weires and G. L. Smith j Abstract. — A survey for predators of apple leaf-feeding mite species was j made during mid-summer of 1975, 1976, and 1977. Apple leaves were col- ! lected from 36 commercial orchards throughout the 10 major fruit-growing : counties of Eastern New York. The phytoseiid mite predators Amblyseius I fallacis (Garman), Typhlodromus pomi (Parrott), and Typhlodromus sp. I were found in 26, 2, and 1 of the orchards, respectively. The stigmaeid I Zetzellia mail (Ewing) was found in eight of the orchards, while the coc- I cinellid beetle Stethorus punctum LeConte was found in one. Results of 1 this survey suggest that Eastern New York apple growers could utilize cer- tain of these mite predators in integrated mite control programs similar to ; those developed in other areas. I Introduction i Recent emphasis on more efficient utilization of chemical pesticides in deciduous orhcards has encouraged the development of integrated mite con- trol programs employing mite natural enemies in addition to chemical and cultural control methods (Glass, 1975). The development of widespread re- sistance to organophosphate insecticides among predatory mites of the fam- ily Phytoseiidae (Croft and Brown, 1975) has provided much of the impetus for such programs. Dean (unpublished manuscript) listed four species of plant-feeding mites on apple trees in the Hudson Valley including the European red mite, Pan- onychus ulmi (Koch), twospotted spider mite, Tetranychus urticae Koch, clover mite, Bryobia praetiosa Koch, and apple rust mite, Aculus schlecht- endali (Nalepa). The aforementioned species plus the McDaniel spider mite, Tetranychus mcdanieli McGregor, have been reported (Brann, 1976) in the Champlain Valley. Dean (unpublished manuscript) listed the ladybird beetle, Stethorus punc- tum LeConte, as the most common mite predator in the Hudson Valley and reported that two bugs Hyaloides vitripennis (Say) and Orius insidiosus Say, known to feed on the European red mite had been largely eliminated from most orchards by DDT sprays. Dean also found predaceous phyto- ' Approved by the Director of the New York State Agric. Exp. Sta. for publication as Journal Paper No. 3129, January 27, 1978. 16 NEW YORK ENTOMOLOGICAL SOCIETY seiids but considered their influence slight. Specimens of the predaceous stigmaeid, Zetzellia mali (Ewing), were collected during 1969 from the Hud- son Valley Laboratory orchards by Dr. Dean and identified by Dr. E. W. Baker. Phytoseiid mite predators were found where reduced rates of miticides were used in large plot mite control studies conducted during 1974 in the Champlain Valley. Subsequent trials in both the Hudson and Champlain Valleys indicated that the phytoseiids were providing biological control of the European red mite (Weires, unpublished data). A limited survey for mite predators was conducted in Ulster and Clinton counties during 1975, while the survey was expanded during 1976 and 1977 to include these as well as eight other counties (Eig. 1) throughout Eastern New York. Materials and Methods I Leaf samples were collected from six orchards in Clinton and Ulster coun- I ties in 1975, from 22 orchards throughout Ulster, Clinton, Saratoga, Orange, I Rockland, Putnam, and Greene counties in 1976, and from twelve orchards H in Columbia, Dutchess, Rennselaer, Rockland and Orange counties in 1977. I| Orchards were randomly selected with the help of Cooperative Extension H fruit agents. Sample size varied, but at most locations four to eight blocks ; which included at least two varieties (one of them McIntosh) and ranged in size from 5-15 acres were sampled by collecting 10 leaves/tree from five , trees in each block. More intensive sampling was conducted where greater ‘ precision was required to determine predator-prey ratios. The number of ( samples/orchard, leaves/sample, and leaves/tree, as well as the sampling < date and location are presented in Table 1. • Mites were brushed from the leaves onto a glass plate coated with glue using a mite brushing machine (Henderson & McBurnie, 1943). All mite species and stages were counted and growers were sent a report of the counts. Predatory mites were carefully removed from the plates, placed in 95% EtOH, and mounted in PVA modified Heinz medium. Dr. Chand Watve (Geneva, NY) helped collect, mount, and identify the phytoseiids during 1975, while in 1976 and 1977 the phytoseiids were mount- ed and identified by Mr. Smith and confirmed by Dr. Watve. Collections of the stigmaeidae were compared with original specimens determined by Dr. Baker or sent to Dr. Watve. The coccinellids were determined by Dr. John Leeper (Geneva, NY). Results and Discussion Three species of phytoseeid predators were found. Amblysieus fallacis (Garman) was the predominant species. Typhlodromus pomi (Parrott) was j found in one orchard during 1975 and 1976 (Table 1). Another unidentified , Typhlodromus species was also found in one orchard in 1975. . VOLUME LXXXVII, NUMBER 1 17 Fig. I. Location of commercial apple orchards surveyed for mite predators in Eastern New York, 1975-1977. Amblysieus fallacis was found in 5 of the 6, 14 of the 22, and 11 of the 12 orchards surveyed in 1975, 1976, and 1977, respectively. Zetzellia nuili was found in three of the orchards surveyed each season. Stethorus punctum was found in the same orchard in both 1976 and 1977. 18 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Mite predators in Eastern New York Commercial Apple Orchards — 1975-1977. Date sam- pled No. sam- ples No. No. leaves/ leaves/ sample tree Predator species Orchard City or village County A.fal- lacis z. malt 5. punc- tum 1975 7/29 A New Paltz Ulster 48 25 25 X' X O 7/29 B Clintondale Ulster 4 50 25 X o o 7/29 C High Falls Ulster 4 50 25 X o o 7/29 D Highland Ulster 39 25 25 X X 0 7/29 E Keesville Clinton 48 25 25 X 0 0 7/29 F Peru Clinton 4 25 25 o X o 1976 7/22 G Brewster Putnam 4 50 25 X X 0 7/26 H Pomona Rockland 12 50 25 0 0 0 8/3 I Slate Hill Orange 5 50 10 0 0 X 8/3 J Warwick Orange 5 50 10 x^ X 0 8/18 K Walden Orange 6 50 10 X o 0 8/3 L Walden Orange 8 50 10 X o o 8/2 M Plattekill Ulster 4 50 10 0 o o 8/2 N Plattekill Ulster 4 50 10 X 0 0 8/2 O Marlboro Ulster 4 50 10 X o o 7/20 P Milton Ulster 40 25 25 X' o 0 8/2 Q New Paltz Ulster 5 50 25 X 0 0 6/24 D Highland Ulster 36 25 25 X X 0 8/2 R Ulster Park Ulster 4 50 10 o o o mi S Athens Greene 4 50 10 0 0 0 mi T Malta Saratoga 4 50 25 X 0 0 mi U Clifton Park Saratoga 4 50 25 X 0 0 mi V Rexford Saratoga 4 50 25 o 0 o 7/13 E Keesville Clinton 52 25 25 X o 0 8/13 W Peru Clinton 4 50 10 X o 0 8/13 X Chazy Clinton 4 50 10 0 0 0 8/13 Y Chazy Clinton 4 50 10 0 0 0 8/13 Z Chazy Clinton 2 50 10 X o o 1977 8/1 AA Claverack Columbia 6 50 10 0 o o 8/2 BB Red Hook Dutchess 6 50 10 X o 0 8/2 CC Germantown Columbia 8 50 10 X 0 o 8/2 DD Greenport Columbia 8 50 10 X o 0 8/3 EE Red Hook Dutchess 4 50 10 X o o 8/4 I Slate Hill Orange 8 50 10 X 0 X 8/4 FF New Hampton Orange 4 50 10 X 0 o 8/4 H Pomona Rockland 8 50 10 X X 0 8/5 GG Valatie Co Columbia 8 50 10 X X 0 8/5 HH Kinderhook Columbia 6 50 10 X 0 o 8/5 II Castleton Rensselaer 6 50 10 X o 0 8/5 JJ West Ghent Columbia 4 50 10 X X o X = present. O = absent. ' Several Typhlodromus pomi. ^ Several Typhlodromus sp?. VOLUME LXXXVII, NUMBER 1 19 Amblysieus fallacis was found where a variety of orchard spray programs were used, whereas T. pomi and Z. mali were only found in young, seldom- sprayed orchards or in older orchards being brought back into production or receiving a very minimal spray program. Stethorus punctum was found in an orchard which received regular sprays, but in which European red mite populations exceeded five mites/leaf at the time of sampling. Knisley and Swift (1972) found eight species of phytoseiid mite predators in New Jersey apple orchards. Amblysieus fallacis was the predominant species in orchards receiving pesticide sprays while Typhlodromus longi- pilus Nesbitt was found less frequently. Typhlodromus pomi was the most common phytoseiid in abandoned orchards. The stigmaeids Z. mali and Agistemus fleschneri (Summers) were found in both abandoned and com- mercial orchards (Knisley and Swift, 1972). Growers throughout Eastern New York should be able to utilize A. fal- lacis in integrated mite control programs similar to those developed for other areas (Swift, 1968; Holdsworth, 1974; Croft, 1975). This is in contrast to the situation in Western New York where both A. fallacis and Typhlodromus pyri Scheuten are found with T. pyri predominant (Watve and Lienk, 1976 and pers. communication). The absence of T. pyri in our collections is considered a benefit (1) because of the threat it poses for A. fallacis through interspecific competition; (2) because of its low level of tolerance to the most commonly used organophosphate insecticides (Watve and Lienk, 1976); and (3) because there is some question as to its predatory effective- ness (Croft, 1976). Utilizing Z. mali in an integrated program does not appear promising. Zetzellia mali is apparently susceptible to present pesticide programs. Re- cent work also suggests that because of spatial heterogeneity coupled with lack of feeding on female tetranychids, Z. mali may not be able to control phytophagous mites below economic damage levels (Santos, 1976). In ad- dition Croft and McGroarty (1977) have observed that A. fallacis seemed unable to express its normal reproductive and predation potential in or- chards where A. fleschneri and Z. mali were present at similar densities to A. fallacis. Stethorus punctum was found in only one orchard during our collections. Croft and McGroarty (1977) reported that S. punctum occurred very spo- radically in Michigan apple orchards but that its occurrence was unrelated to pesticide use patterns. Growers in the southwestern portion of the Hud- son Valley should be able to utilize 5. punctum in an integrated mite control program but would probably have to adapt the Pennsylvania integrated mite control practices (Tetrault et al., 1977) designed to protect Stethorus. 20 NEW YORK ENTOMOLOGICAL SOCIETY Literature Cited Brann, J. L., Jr. 1976. Fruit insect control recommendations for New York State. 39 pp. In 1976 New York State Insecticide, Fungicide, and Herbicide Recommendations. Cornell Univ. Extension Ser. 343 pp. Croft, B. A. 1975. Integrated control of apple mites. Mich. State Univ. Ext. Bull. E-825, 12 p. . 1976. Establishing insecticide-resistant phytoseiid mite predators in deciduous tree fruit orchards. Entomophaga 21:383-399. , and A. W. A. Brown. 1975. Responses of arthropod natural enemies to insecticides. Annu. Rev. Entomol. 20:285-335. , and D. L. McGroarty. 1977. The role of Amblyseius fallacis (Acarina: Phytoseiidae) in Michigan apple orchards. Mich. State Univ. Agric. Exp. Sta., Farm Sci. Res. Rept. 333. 22 p. Glass, E. H. 1975. Recent developments in deciduous orchard pest management in the United States. EPPO Bull. 5:101-111. Henderson, D. F., and H. Y. McBurnie. 1943. Sampling technique for determining populations of citrus and red mite and its predators. U.S. Dep. Agr. Circ. 671. 11 p. Holdsworth, R. P. 1974. Integrated control of apple pest insects in Ohio. In 1974 commercial fruit spray recommendations for Ohio. Bull. Ohio State Univ. Extension Ser. 506:20- 26. Knisley, C. G. and F. C. Swift. 1972. Qualitative study of mite fauna associated with apple foliage in New jersey. J. Econ. Entomol. 65:445^48. Santos, M. 1976. Evaluation of Zetzellia mali as a predator of Panonychus ulmi and Aculus schlechtendali. Environ. Entomol. 5:187-191. Swift, F. C. 1968. Population densities of the European red mite and the predaceous mite Typhlodromus (A.) fallacis on apple foliage following treatment with various insecti- cides. J. Econ. Entomol. 61:1489-1491. Tetrault, R. C., D. Asquith and W. M. Bode. 1977. Apple integrated pest management pro- gram. p. 44-52. In 1977 Tree Fruit Production Guide. Penn. State Univ. Extension Ser. 77 pp. Watve, C. M. and S. E. Lienk. 1976. Resistance to carbaryl and six organophosphorus in- secticides of Amblyseius fallacis and Typhlodromus pyri from New York apple or- chards. Environ. Entomol. 5:368-370. Department of Entomology, New York State Agricultural Experiment Station, Highland, New York 12528. Received for publication Eebruary 10, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 21-37 STRATEGIES OE GALL EORMATION IN PEMPHIGUS APHIDS'- ^ I Daniel P. Eaith i I Introduction I This is a study of the various aspects of the strategies of gall formation I by two species of aphids which form galls on the leaves and petioles of the I Eastern Cottonwood, Populus deltoides. Pemphigus galls are initiated when ! a young stem mother or fundatrix (the name for the first parthenogenetic ! generation after the sexual generation) begins feeding on the newly emerging leaves of Populus. On Long Island, the two common gall-forming species I on cottonwood are Pemphigus populicaulis Eitch and P. populitransversus I Riley. These induce “oblique” and “transverse” galls, respectively. The I oblique gall is formed by a swelling and twisting of the leaf at the point i where the blade meets the petiole. The transverse gall, by contrast, is formed entirely on the leaf petiole. In each case, feeding by the fundatrix induces changes in the growth of the plant tissue, resulting in a chamber enclosing her. The parthenogenetic offspring of the mature fundatrix usually remain within the gall until after a final molt that, in all cases, results in winged individuals. These winged forms (alate fundatrigeniae, called “alates” for short in this paper) migrate to the roots of a secondary host plant (members of the Cruciferae or Compositae). After one or more par- thenogenetic generations on the secondary host, winged forms (“sexupar- ae”) are produced that return to the primary host (Populus) and give birth parthenogenetically to sexual forms. Each fertilized female deposits, in cracks in the bark of the tree, a single egg which in turn develops into a fundatrix, completing the life cycle (Harper 1959). The exact timing of these life history stages for P. populicaulis and P. populitransversus will be dis- cussed later in this paper. Initiation of Pemphigus galls has been observed only on young, newly unfolding leaves. P. deltoides produces leaves continuously throughout the summer; early leaves are preformed in the winter bud and morphologically distinct late leaves are formed from leaf primordia after the expansion of the early leaves (Kozlowski 1971). As a result, feeding sites for young fun- datrices are continuously available from spring until late summer. However, once the fundatrix initiates a gall, it does not appear to change its feeding site. For purposes of this paper, I shall define “gall position” as one plus the number of leaves on a shoot preceding (older than) the leaf bearing the gall. ' This research was supported by Grant No. BMS720221 1 A04 from the National Science Foundation to Robert R. Sokal. ^ This paper is publication no. 155 for the Graduate Program in Ecology and Evolution at the State University of New York at Stony Brook, Stony Brook, New York 11794. 22 NEW YORK ENTOMOLOGICAL SOCIETY For example, if the gall position is “6,” then the gall is located on the sixth leaf of the shoot, counting from the base of the shoot. Obviously, the proximal causes of a given gall position are the time the fundatrix nymph emerges and initiates the gall, the time of bud break of the shoot, and the subsequent rate of leaf expansion of the shoot. The ultimate causes of gall position, that is, the selective pressures affecting the timing of gall initiation, are discussed in this paper. Whitham (1974) first considered the adaptive significance of the position of Pemphigus galls on cottonwoods. He showed that galls sharing the same leaf were generally smaller than single galls, and argued that the aphids were resource (sap) limited. He found a significant correlation of gall size with leaf size for oblique and transverse galls, and also showed that galls of both species are found on larger-than-average leaves. He suggested that the obliques may achieve this through the timing of gall formation and the trans- verse through the “choice” of longer shoots bearing larger leaves. My preliminary observations on one tree showed that of 129 oblique galls, 97 (75.2%, with 95% confidence levels 66.5-82.8%) were found on the larg- est leaf of their shoot. However, of 154 transverse galls, only 38 were found on the largest leaf (24.7%, limits 19.1-33.0%). The correlation of leaf length with gall diameter was 0.87 (n = 129) for oblique galls and 0.39 (/? = 154) for transverse galls. This suggested that, to the extent that fitness is related to gall size, leaf length might influence the success of transverse galls less than that of oblique galls. In this paper a dimorphism in gall position of P. populitrcmsversus is demonstrated. Then, the possible factors affecting relative success of these galls are determined with the goal of discovering whether there is differential fitness at different gall positions. Finally, for both P. populitransversus and P. populicaulis, position is analyzed as a possible response to such selection pressures. Materials and Methods Observations and collections were made from cottonwood trees in the vicinity of Stony Brook, Long Island, starting in early May, 1975. Mea- surements of the cottonwoods were made in the field and galls were re- moved, tagged, and placed in jars of alcohol. Populus measurements in- cluded tree height and for shoot samples, the number of leaves on the shoot, gall position, position of the largest leaf, petiole lengths, and leaf lengths. In the laboratory, each gall was measured and its aphid population censused under a dissecting microscope. The numbers of winged forms and individ- uals in three arbitrarily chosen size classes of nymphs were recorded for all galls. Galls that showed signs of predation or other disturbances were not used for this study. VOLUME LXXXVII, NUMBER 1 23 Relative reproductive success of different fundatrices was estimated by censuses of galls in which some winged forms had developed. This seems to be a reasonable estimate of the reproductive potential of a gall for the season. In a study of six species of Pemphigus in Alberta, including P. popidicaulis and P. populitransversus, Harper (1959) found a correlation of 0.99 between the average number of aphids emerging from a gall for a given species over the summer and the average population counts in galls in a separate sample of that species. I will also assume that, within a species, differences in population counts among galls reflect differences in the num- ber of migrants that could be expected from the galls over the summer. Dimorphism in the Timing of Gall Initiation in P. populitransversus Gall initiation by Pemphigus populitransversus was found to occur at two clearly separate times during the summer of 1975. Gall initiation is first detected as a slight swelling and bending of the petiole at the point where a fundatrix nymph has begun feeding. In this early stage, the young fundatrix is not enclosed and appears as a small black spot in the pocket formed by the swelling and bending of the petiole. Transverse galls first began forming, along with oblique galls, around May 15 on cottonwoods in the vicinity of Stony Brook. At this time, the early leaves of the tree were expanding and provided suitable sites for gall initiation. The trees were checked carefully on successive days, and after two or three days, there was no further gall initiation. Then, on June 15, more transverse galls began forming on the newly formed late leaves. This dimorphism in the timing of gall initiation resulted in a set of galls on the early leaves and a set of galls on the late leaves of the shoots. The early transverse galls had an average position of 6.2 ± .39 (n = 48), over several trees, while the average position of the late galls was 16.2 ± .35 (n = 85). A dimorphism in gall shape, life history, and aphid morphology has been described for this species by Senner and Sokal (1974). The morph they refer to as ''elongate" produces galls that are more elongate in shape than those of the other morph called "globular." The question immediately arose whether the presence of early and late transverse galls was related to the globular-elongate dimorphism. An analysis of the early and late transverse galls confirms the suspected relationship (Table 1). The late galls are larger than the early galls, and more spherical as shown by the ratios of gall dimensions. The values in Table 1 resemble those of the sample from West Point, Georgia, shown in Table 3 of Senner and Sokal (1974). The early and late galls were significantly different for each character. Corroborating tests were run following methods developed by Senner and Sokal (1974). The gall dimensions and their ratios split a mixed sample of 71 galls into early and 24 NEW YORK ENTOMOLOGICAL SOCIETY Table I. Means of gall characters for 39 early and 31 late transverse galls in Stony Brook, New York, 1975. Gall character Early transverse Late transverse Fs Length 10.9 14.7 38.9 Width 7.5 14.3 19.4 Depth 8.0 13.4 12.4 Length/width 1.46 1.03 77.2 Length/depth 1.38 1.11 20.5 Note: Measurements are in millimeters. is the sample F-statistic testing differences between the means. All values of are significant at F < 0.001. late classes fairly well. Only two galls would be misclassified by time of gall formation when the sample was split according to gall depth and only 4 galls would be so misclassified on the basis of length/width ratio. It is highly probable that the early and late galls correspond to the elongate and globular morphs, respectively. Population Sizes in Globular and Elongate Galls Senner and Sokal (1974) found that globular galls had larger within-gall population sizes than elongate galls. Table 2 summarizes the data on pop- ulation size for several trees on Long Island. In a pooled sample from sev- eral trees, the population size of globular galls was higher than that of elon- gate galls; the average population size of globular galls was 377.6 ± 19.2 (// = 60) compared with 290.9 ± 13.6 (n = 52) for elongate galls, for trees on which both types of galls were present. These values were significantly different (Fg = 10.4 > F oo.^(i,ui))- For individual trees, the results were somewhat variable (Table 2). The difference in population size between elongate and globular morphs was significant for both tree 4 and tree 8 (Fg 9.8 > F.oo.mi.46)^ Fs = 7.6 > F.o2.5(i.2o)- For tree 3, the difference in popula- tion sizes of the two morphs was not significant (Fg = . 128 < F (,00.24)) • Thus, average population size is somewhat variable among trees for each morph, but in general, stem mothers of globular galls produce more offspring than those of elongate galls. Table 2. Mean population size (± standard error) of globular and elongate galls for different trees in Stony Brook, New York, 1975. Tree 3 Tree 4 Tree 8 Globular 376 ± 46.3 (14) 409 ± 29.0 (29) 454 ± 37.3 (12) Elongate 357 ± 26.4 (II) 237 ± 20.1 (18) 294 ± 19.1 (10) Note: Sample sizes (number of galls) in parentheses. VOLUME LXXXVII, NUMBER 1 25 Table 3. Analysis of covariance of population sizes of globular galls on characteristics I of the galls, and correlation matrix for these variables. Source of variation Sum of squares df Mean squares F. Corrected total 1638165.15 73 Adjusted treatment 212100.79 2 106150.40 13.53*** Regression 1073545.41 6 178924.24 22 8^*** Error 509518.67 65 7838.75 Regression homogeneity Error for regression 111766.55 12 9313.88 1. 24ns homogeneity 397752.13 53 7504.76 Partial regression coefficients Adjusted treatment means GL GW GLP LAG GPL GLL Tree number 23.0 30.6 -6.5 4.0 -20.1 -.7 2 3 4 314.5 478.7 396.3 Correlation matrix Population GL GW GLP LAG GPL GLL size GL .618 .008 -.217 -.498 .000 .612 GW .000 .116 -.327 .200 .684 GLP .008 .000 J26 -.097 LAG .000 .299 .131 GPL .000 -.284 GLL .009 Note: Treatments are differences among three trees. Gall characteristics used as covariates are: GL — gall length, GW — gall width, GP — gall position, LAG — number of leaves after gall position, GPL — number of galls per leaf, GLL — length of leaf bearing the gall. Significance values are symbolized by *** — P .001, ** — .001 P ^ .01, * — .01 P « .05, ns — P > .05. All correlation coefficients significant at P ^ .01 are in italics. ' What factors account for the variation in population size among galls i within each morph? Can factors relating to the difference in gall position of the two morphs account for the greater population sizes of the globular galls? Also, is there evidence of competition between galls? Such evidence would suggest some limiting factor on population size. I shall address these questions in turn. The population size of globular galls was used as the dependent variable in an analysis of covariance (Table 3). Gall length, gall width, gall position, the number of leaves after (distal to) the gall position, the number of galls per leaf, and the length of the gall-bearing leaf were used as covariates and the 3 trees sampled served as treatments. The regression was significant and homogeneous over trees. However, the adjusted treatment means were sig- 26 NEW YORK ENTOMOLOGICAL SOCIETY nificantly different = 13.5 > F 00K2.6.5)); therefore, the regression does not completely account for the variation in population size among trees. The partial regression coefficients for gall length, gall width and gall posi- tion were significant = 19.8 > F^ = 23.9 > F „„ki.65), and Fs = 3.86 > F o.5(i,6.5), respectively). As noted earlier, population size in- creased with gall dimensions. However, it decreased with position. Age of the gall may be responsible for this effect, since galls with lower positions will be slightly older and hence more populated than those at higher positions. There seems to be some competition among galls on the same leaf, as suggested by the significant negative correlation of number of galls per leaf with gall length and gall width, and the negative correlation of number of galls per leaf with population size (significant at F < 0.025). The effect of the number of galls on a given leaf or shoot on population size in the galls seems to be realized through the effect of number of galls on gall dimensions. For tree 2 alone, competition was tested between galls on the same shoot. The number of galls per shoot was seen to increase with the number of leaves on the shoot (r = .60, n = 31). The regression of population size on the number of galls per shoot, holding the number of leaves constant, was also significantly negative (Fj = 8.4 > F.ook2,6.'5))- There would appear to be competition for resources among globular galls on the same shoot. Gall leaf length was found to increase with gall position; that is, leaves were larger toward the tip of the shoot. However, the length of the leaf did not seem to be important to the success of the gall. Gall leaf length had no significant correlation with population size, or with gall dimensions. Thus, larger populations are found in larger galls and in earlier positions, but this is not related to leaf size. A similar analysis of covariance was performed for three trees with elon- gate galls (Table 4). The regression is significant and, in this case, it does account for the difference in the average population sizes of the three trees, since the adjusted treatment means were not significantly different (F. = 2.98 < F,.o,2,39.). Again, gall dimensions correlate highly with population size in the gall, but only the partial regression coefficient on gall width was significant (Fj = 4.33 > F 05(1,39)). Gall leaf length, as in the case of globular galls, increases with gall leaf position and does not show a significant correlation with pop- ulation size or gall dimensions. However, in contrast to the globular case, the gall position correlated positively with population size. To investigate further any effect due to variation in leaf length with position, population size was regressed on gall position, holding gall leaf length constant. This regression was significant (Fj = 5.37 > F.o5(i,40)). Therefore, in elongate galls, there must be some factor other than leaf size relating to gall position that affects population size. VOLUME LXXXVII, NUMBER 1 27 Table 4. Analysis of covariance of population sizes of elongate galls on character- istics of the galls, and correlation matrix for these variables. Source of variation Sum of squares df Mean squares Corrected total 427501.48 47 Adjusted treatment 31698.23 2 15849.14 2.98ns Regression 121292.59 6 20215.43 3.80** Error 207382.82 39 5317.51 Regression homogeneity Error for regression 55584.99 12 4632.08 .82ns homogeneity 151797.83 27 5622.14 Partial regression coefficients Adjusted treatment means GL GW GLP LAG GPS GLL Tree number 10.4 33.3 4.5 -1.9 11.1 -.60 3 4 8 329.2 285.5 228.1 Correlation matrix Population GL GW GLP LAG GPS GLL size GL .445 .253 .110 -.198 .156 .430 GW .009 .144 -.181 .063 .462 GLP -.298 .150 .528 .462 LAG .496 .455 -.170 GPS .494 .423 GLL .276 Note: See Table 3. For elongate galls, competition was examined in terms of galls on the same shoot. Surprisingly, the number of galls per shoot correlated positively with population size within a single tree. The number of galls per shoot correlated with shoot length (r = .496, n = 48). Longer shoots produce nu- merous and more populated galls for unknown reasons. To find the causes of the differences in the average population sizes of globular and elongate galls, these morphs were used as treatments in an analysis of covariance (Table 5) with gall length, gall width, gall leaf length, and the number of leaves after the gall as covariates. The two treatments were significantly different for each of these variables. The treatment means of populations sizes were 378 ± 19.2 for globular and 291 ± 13.6 for elongate galls. As expected, the partial regression coefficients for gall length and gall width were significant {F, = 8.7 > F.oo.5d.,28), Fs = 12.0 > F.ooui.izs))- These gall dimensions account for the differences in population size of the two 28 NEW YORK ENTOMOLOGICAL SOCIETY Table 5. Analysis of covariance of population sizes between globular and elongate galls on characteristics of the galls, and correlation matrix for these variables. Source of variation Sum of squares df Mean squares Fs Corrected total 2599665.20 133 Adjusted treatment 243.97 1 243.97 .026ns Regression 1096649.72 4 274162.43 28.86*** Error 1215984.13 128 9499.87 Regression homogeneity Error for regression 63056.38 4 1574.10 1.70ns homogeneity 1 152927.75 124 9297.80 Partial regression coefficients Adjusted treatment means GL GW LAG GLL Globular Elongate 15.5 26.9 .24 2.3 352.25 356.24 Correlation matrix Population GL GW LAG GLL Size GL .580 -.020 .140 .612 GW -.164 .313 .676 LAG .003 -.064 GLL .240 Note: Treatments are differences between morphs. Gall characteristics used as covariates are: GL — gall length, GW — gall width, GLL — length of leaf bearing the gall, and LAG — number of leaves after the gall position. *** — P « .001, ** — .001 P « .01, * — .01 P « .05, ns — P > .05. Correlation coefficients significant at P « .01 are in italics. morphs, since there was no significant difference between the adjusted treat- ment means {F^ = .026 < F.o!5(i,i28))- Within tree number 8, the correlation of gall leaf length and population size, for both morphs, was .560 {n — 20). However, overall, the correlation of leaf length with population size (.240) and gall dimensions (.140, .313) was low, and within each morph, gall leaf length did not account for the variation in population size. Therefore, it seemed useful to try another vari- able that might explain gall size and the relationship of gall position to population size in the elongate morph. Petiole length and width were tried as variables that might affect gall dimensions. Since transverse galls grow on petioles, the size of the latter might be more closely related to gall dimensions than leaf length. For a new pooled collection of 9 globular and 10 elongate galls from a single tree, petiole width explained 78% of the variance in gall width (F^ = 63.3 > F. 001(1. i8))i and petiole length explained 41% of the variance in gall length (Fg = 12.3 > F 00,5(1.18))- Petiole length and width, then, are better indicators of gall dimensions than gall leaf length for transverse galls. 29 I ! VOLUME LXXXVII, NUMBER 1 14 16 18 20 22 24 26 28 NUMBER OF LEAVES PER SHOOT Fig. 1. Position of globular gall (crosses and dashed line) and position of the longest petiole (x’s and solid line) on the total number of leaves on the shoot. The lines represent significant least squares regression lines. The ordinate is position and the abscissa is total number of leaves. Samples based on tree 2, Stony Brook, New York, 1975, Note that for most shoot sizes the position of globular galls is earlier than that of maximal petiole size. I In this new sample, gall length of the two morphs was not significantly different (Fs = .5 < F 0.5n.i8)), but width of the globular morph was signifi- cantly greater than that of the elongate morph (Fs = 14.7 > F.oo.sn.is))- Pet- iole width for leaves with globular galls was 4.5 ± .36 mm and for leaves ! with elongate galls was 3.1 ± .22 mm. My field observations suggest that I this difference in width is not due to the presence of the galls on the petioles; I measurements of many late and early leaf petioles without galls showed that petiole width of late leaves was rarely as low as 3.0 mm, which was the usual width of early petioles. This difference in petiole width between early and late leaves may indirectly account for the general difference in popu- lation size of the two morphs through its effect on gall width. An analysis I of covariance was carried out with gall width as the dependent variable, I globular and elongate as treatments, and petiole width as a covariate. The 1 regression, controlling for petiole width, was significant (Fs = 52.4 > 1 ^.001(2. 17))- Thus, although petiole width does explain 78% of the variance in I gall width for elongate and globular galls taken together, other (possibly I genetic) determinants of the differences in gall width of the morphs remain. In summary, the greater population sizes of globular galls are partly at- tributable to the greater petiole widths of the late leaves on which these 30 NEW YORK ENTOMOLOGICAL SOCIETY "time" Fig. 2. Regression of petiole length on gall position, holding the total number of leaves constant (P < .05). Numbers of graph indicate number of times a given point occurs. The ordinate is petiole length (cm) and the abscissa is "time” (deviation of gall position from regression on total number of leaves per shoot). Sample is made up of globular galls taken from tree 2, Stony Brook, New York, 1975. galls are found, relative to the petiole widths of the early leaves, which are the sites of the elongate galls. Position of Globular Galls among the Late Leaves Position of globular galls, as expected, increases with the total number of leaves on the shoot (Figure 1). Any given time of gall initiation will result in a later position on faster than on slower growing shoots, because the faster shoots will have produced more leaves in the same amount of time. Although population size of a gall did not correlate with gall position, it may correlate with time of gall initiation, which would suggest selective pressure for earlier or later gall initiation. A regression of population size on gall position, controlling for the number of leaves per shoot relates population VOLUME LXXXVIl, NUMBER 1 31 Fig. 3. Globular gall position ( + ’s and dashed line) and longest petiole position (x’s and solid line) on the total number of leaves on the shoot. The solid line represents a significant regression line. Ordinate: position; Abscissa: total number of leaves. Samples are from tree 8, Stony Brook, New York, 1975. size to time; however, this regression was not significant, so population size is not dependent on time in the data from tree 2. Since petiole length has been shown to correlate with gall length, I also investigated the possible relationship of position of globular galls with the variation in petiole size of late leaves. Does the position of globular galls maximize petiole size among late leaves? For tree 2, petiole length was regressed on leaf position, holding the total number of leaves constant (Fig- ure 2). Petiole length significantly increased with time, suggesting that later gall initiation may imply an increased population size through the effect of petiole length on gall length. The relationship of the actual positions of globular galls to petiole size is shown in Figure 1. For most shoot sizes, the globular galls appear earlier than the leaf with the longest petiole. The results were similar for another tree (8), as shown in Figure 3. There is no evidence, then, that the globular galls were positioned as an adaption to maximal petiole length among the late leaves. Positions of Elongate and Oblique Galls among the Early Leaves Eor elongate galls alone, petiole length explains 62% of the variation in gall length (Fg = 17.9 > F ooki.id)- Gall leaf length, on the other hand, does 32 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 4. Pemphigus galls, (a) A "typical” gall off. populicaulis (“oblique”), (b) A "typical” gall of P. popiditransversus ("transverse”). Drawings by Lorenz Rhomberg. not have a significant correlation with gall length (r = .156, n = 48). Since gall length was found to correlate highly with population size (r — .50, n = 25), for tree 4, it might be expected that there would be selection for timing of gall initiation such that the resultant position of the gall provided a larger petiole, on the average. Such selection would be possible if petiole length can be predicted by timing of gall initiation. By contrast, oblique gall di- ameter correlates more highly with leaf length (r = .874, n = 25) than with petiole length (r = .209, n = 25); and gall position might similarly reflect a prediction of leaf size through timing. The reasons for the differences in correlations are clear, since elongate galls are formed on leaf petioles while oblique galls are formed by a twisting together of the base of the leaf blade (Figure 4). The relative and absolute positions of oblique and elongate galls are not constant from tree to tree (Table 6). Trees 1 and 4 each show elongate galls significantly earlier in position than oblique galls. Tree 3 showed the same trend but the difference was not significant. Table 6. Mean (± standard errors) of oblique and elongate galls of different trees sampled in Stony Brook. New York, 1975. Oblique N Elongate N Fs Tree number 1 7.17 ± .44 15 5.59 ± .38 22 12 3 8.80 ± .20 109 8.30 ± .20 98 3.21ns 4 7.45 ± .37 21 5.21 ± .55 19 1 2 0*** 5 6.70 ± .38 25 — — — VOLUME LXXXVII, NUMBER 1 33 Fig. 5. Leaf length as a proportion of maximum length (solid line) and petiole length as a I proportion of maximum length (dashed line) on leaf position. Ordinate: proportion of maximum I length; Abscissa: leaf position. Data are for tree 1, Stony Brook, New York, 1975. For a more detailed examination of the shoots on tree 1 we turn to Figure : 5. Only shoots without galls having 9, 10, or 1 1 early leaves were considered I to limit the variability of patterns of leaf or petiole size due to variation in ! the number of early leaves of the shoots. The abscissa gives the leaf posi- i tion. The ordinate is the ratio of the length for a given position on the shoot I to the maximum length for the shoot. The solid line shows the average j proportion of maximum leaf length (length divided by the length of the i largest leaf on the shoot) and the dashed line shows the average proportion of maximum petiole length for each position on these shoots. The average petiole length at position 6, the first peak, was 7.92, and at position 8, the second peak, was 7.08. Because of the small sample size {n = 6) for each group, these differences are only suggestive {P = .08). Leaf length follows the reverse pattern with the average at position 8 (12.1), greater than that of position 6 (10.7) {F^ - 4.25 > F.osd.io))- Again this is only suggestive, but it appears that the position with the greatest average petiole length is before the position with the greatest average leaf length for this tree. Another expression of this difference is that for shoots without galls on tree 1, the average position of the longest leaf, 7.6, was significantly greater than the average position of the longest petiole, 6.1 (F^ = 30.8 > F’.ooiu.ao))- Is this difference reflected in the positions of oblique and elongate galls? The frequency distributions of elongate and oblique gall positions for tree 34 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 6. Frequency polygons of elongate (solid line) and oblique (dashed line) gall positions for tree 1. Ordinate: frequency; Abscissa: leaf position. 1 are shown in Figure 6. The solid line shows the frequency distribution of the position of elongate galls, and the dashed line that of oblique galls. The mean gall positions of the two types differ for these shoots (Fj = 12.3 > ^.00.5(1 .47))- It is apparent from a comparison of Figures 5 and 6 that both elongate and oblique galls are, on the average, in a position that should result in greater fitness for each. Also, although the figures do not show it, the last 1 or 2 leaves among the early leaves usually have smaller blades and petioles. The oblique and elongate galls, then, are usually on one of the leaf positions that have the greatest average leaf and petiole sizes, respec- tively. Similar data are shown for tree 3 in Figures 7 and 8. Here, the positions of oblique and elongate galls are not significantly different (Figure 8). The mean positions are marked as crosses on the lines. The average petiole lengths at the peaks at positions 6 and 8 are also not significantly different (P = .70). Elongate and oblique galls again are near the optimal positions, but this time with more overlap between the two species. It may be that the timing of gall initiation of one or both species on tree 3 is slightly different from that of tree 1, but the observed difference in positions is better ex- plained by a greater rate of early leaf expansion in tree 3, creating a higher mean position of each species and at the same time, less separation in position between the two. VOLUME LXXXVII, NUMBER 1 35 g i— cr o Q. O cc CL 0.90 0.70^ / / J I I l \ I L 3 4 5 6 7 8 9 LEAF POSITION Fig. 7. Average proportion of maximum leaf length (solid line) and petiole length (dashed line) on leaf position. Ordinate: proportion of maximum length; Abscissa: leaf length. Data are for tree 3, Stony Brook, New York, 1975. Fig. 8. Frequency polygons of elongate (solid line) and oblique (dashed line) gall positions. Ordinate: frequency; Abscissa: gall positions. Data from tree 3, Stony Brook, New York, 1975. 36 NEW YORK ENTOMOLOGICAL SOCIETY It appears that the average earlier timing of elongate galls relative to oblique galls may be a strategy to obtain longer petioles, thereby increasing population size. Similarly, oblique gall position seems to correspond to the largest leaf position among the early leaves. If trees had growth patterns similar to that of tree 1, later elongate gall initiation would be selected against. The earlier initiation of elongate galls relative to oblique galls may be a result of the growth pattern in which petiole length, on the average, reaches its peak before leaf length among the early leaves. Summary The aphid. Pemphigus populitransversus , producing galls on the petioles of cottonwoods, Populus deltoides, was shown to comprise two forms dif- fering in time of gall initiation and resulting in galls on early leaves and late leaves of the shoots. This dimorphism in position corresponds to the elon- gate-globular dimorphism previously described for this species. The factors determining population size in the galls of these two morphs were explored. In each case, gall length and width correlated strongly with population size. Length of the leaf bearing the gall showed no correlation with population size within each morph. However, leaf length did help ac- count for differences in population size between the two morphs. The greater population sizes of globular galls are related to their larger size relative to elongate galls. Globular galls were shown to be significantly greater in length, width, and depth measurements. Petiole size was shown to account for much of the variation in gall size in P. populitransversus, both among and within morphs. The larger petioles of the late leaves help account for the larger size of the globular galls; hence, the greater population sizes of globular galls is due, in part, to the strategy of late gall formation, through the indirect effect of petiole size on population size. Within the late leaves, globular galls tended to be formed on leaves before those with maximal petioles. Among the early leaves, elongate galls did seem to be in an average position that matched the position of the longest petiole. The galls of P. populicaulis were in a position such that leaf size was nearly maximized. The dimensions of elongate galls were dependent more on petiole size and those of oblique galls more on leaf size. This agrees with their sites of gall formation. The tendency among early leaves for petiole size to reach a maximum before leaf size may explain the slightly earlier initiation of elongate compared to oblique galls. VOLUME LXXXVII, NUMBER 1 37 Literature Cited Harper, A. M. 1959. The gall aphids of Poplar in Alberta II. The Canadian Entomologist 61:680-685. Kozlowski, T. T. 1971. Growth and Development of Trees. Academic Press, New York. 394 pp. Senner, J. W. and R. R. Sokal. 1974. Analysis of dimorphism in natural populations with consideration of methodological and epistemological problems. Systematic Zoology 23:363-386. Whitham, T. 1974. Strategies of Pemphigus aphids in using resources. Unpublished Master’s thesis. Ohio State University. Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, New York 11794 Received for publication March 31. 1978 NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 38^1 EUROPEAN KATYDID MECONEMA THALASSINUM (DE GEER) RECORDED EROM NEW LOCATION ON LONG ISLAND, NEW YORK (ORTHOPTERA: TETTIGONIDAE) Burke Smith, Jr. Abstract. — Establishment of the European katydid Meconema thalassi- num (De Geer) is confirmed on Long Island, New York. Thirteen specimens taken at Garden City in July 1977, add a third geographic location on Long Island for the species, previously found in much smaller numbers at King’s Park (1968) and Little Neck (1957 and 1959). No other North American records are known. Distinguishing morphological characteristics of the species are described and illustrated, and habits (including sound production and food preferences) are discussed. Thirteen specimens of the Europeak katydid Meconema thalassinum (De Geer) were collected by the author at Garden City, Long Island, New York, in July 1977. Known as the Oak Bush Cricket, or Eichenschrecke, this species is widely distributed in Europe (Beier, 1966), but is a rarity in North America, where it has been found previously at only two locations, both on Long Island. It was first recorded at Little Neck, Long Island, where two males and two females were taken in July and August 1957; two other specimens were taken from the same locality in 1959. The specimens from Little Neck were all collected by Mr. John K. Torres, editor of Audubon Magazine, and were reported by A. B. Gurney ( 1960, 1960a). The next Long Island record was at King’s Park, Suffolk County in 1968, where a single male was taken by R. M. Emberson, reported by D. E. Johnstone (1970). The present find establishes a third location for the species on Long Island, with a larger number of specimens taken than heretofore. There are no other North American records known for Meconema thalassinum. When first found on Long Island twenty years ago, Gurney (1960) con- sidered it to be an established adventive, probably introduced from eggs on imported nursery stock. The scarcity of records concerning it since that time (only one specimen being recorded during the seventeen-year interim) would indicate that the species has existed in small localized populations, possibly subject to cyclical variations, or that it may even have died out and been reintroduced. It will be interesting to see what additional records of the species may occur in the future. The specimens from Garden City were collected on the porch and grounds of a town-house during the period of July 7 through July 19, 1977. Most were captured in late evening walking around a light on an open porch VOLUME LXXXVII, NUMBER 1 39 ceiling. Several specimens were taken when they were at rest on the porch ceiling during the day. In all, ten males and three females were collected; Dr. Irving J. Cantrall, University of Michigan Museum of Zoology, identi- ! fied the species as Meconema thalassinum (De Geer). Specimens have been i deposited in the University of Michigan Museum of Zoology, the Field j Museum of Natural History, and the Academy of Natural Sciences, Phila- j delphia. I Meconema thalassinum is a rather small katydid, of the subfamily Me- ! conematinae, having a body length of 11-15 mm, with tegmina extending I about mm beyond the apex of the abdomen (Fig. 1), and is thus not I likely to be confused with the larger and more robust North American ka- I tydids of the subfamilies Phaneropterinae and Pseudophyllinae. Gurney j (1960) has pointed out that the species does superficially resemble some of the native meadow grasshoppers of the subfamily Conocephalinae (genera ; Orchelimum and Conocephalus) which however have the tympana of the I fore-tibia covered, except for a slit-like opening, whereas the tympana of Meconema thalassinum are exposed (Fig. 5). Also, in Meconema thalas- sinum, the tegmina of males and females closely resemble each other and are without any special sound-producing organs. Meconema thalassinum may also be distinguished by special structural characteristics of the genital areas as shown in Figs. 3, 4, 6, and 7. The male cerci are about 4 mm long, I strongly curved, without the lateral projections found in native North Amer- i ican katydids, and with the apices scarcely specialized; the male sub-genital j plate is short, apically truncate and bears two small moveable lateral ap- j pendages; the ovipositor is about 9 mm long, slightly curved and pointed, and without serrations. The habits of this species in Europe, as described below, have been well described in handbooks by various authors, including Chopard (1951), Harz (1957) and Ragge (1965). Gurney (1960) and Johnstone (1970) have also summarized and commented on observations made by previous investi- gators. Meconema thalassinum is part of the Palaearctic arboreal fauna in Eu- rope, occurring on oaks and other deciduous trees, as well as on pines. It may be found intermittently in parks, gardens and tree-rows in urban areas, especially where there are large, old trees. The location in Garden City, Long Island, provides such a habitat with its large well-established maple trees. The species is nocturnal, individual insects remaining quiet during the day and becoming active at twilight. Perhaps the scarcity of records on Long Island may be due in part to these habits. Although some authors (Chopard, 1951; Ragge, 1965) have stressed its carnivorous habits, the species appears to be essentially omnivorous (Johnstone, 1970), feeding on oak and other deciduous leaves as well as on a variety of insects. 40 NEW YORK ENTOMOLOGICAL SOCIETY Figs. 1-7. Meconema thalassinum (De Geer). 1. Male, lateral view. 2. Head, dorsal view, showing shape of fastigium. 3. Dorsal view of supra-anal plate and cerci of male. 4. Ventral view of sub-genital plate and cerci of male. 5. Front leg. showing open tympanum. 6. Lateral view of ovipositor. 7. Ventral view of ovipositor, showing sub-genital plate of female. I VOLUME LXXXVII, NUMBER 1 41 The males lack the stridulatory apparatus which is found almost univer- sally in other Tettigoniidae, but nonetheless make a “purring” or “drum- ming” sound. There have been conflicting reports on how this sound is , produced. Some investigators (Currie, 1953; Harz, 1955) have indicated that it is caused by the rapid beating of a body part (probably a hind leg) on the substratum. However, Cappe de Baillon (1921) suggested that in connection with the rapid body movements associated with sound production, the teg- I mina, which are held above the body, may be rubbed together to produce the actual sound. There are microscopic teeth on the dorsal distal portion of the tegmina which could be used in sound production, but there have been no confirming observations that this is actually the case. Further ob- servations are obviously needed in this respect. In Europe eggs hatch in May, with adults being found from July through October. Females come down from the tree-tops to lay eggs in the cracks of rough-barked trees, or in lichens covering the tree trunks, from the be- ginning of September until the last of October. Whether the species has modified any of its behavior patterns or life span on Long Island, remains to be determined. Literature Cited Beier, M. 1966. Orthopterorum Catalogus, Pars 9: Tettigoniidae; Subfam. Meconematinae, Mecopodinae, Phyllophorinae. Junk, Gravenhage, Netherlands, pp. 248-342. Cappe de Baillon, P. 1921. Note surle Mechanisme de la Stridulation C\\tz Meconema varium Fabr. (Orthoptera — Phasgonuridae). Ann. Soc. Ent. France 90:69-80. Chopard, L. 1951. Faune de France. Orthopteroides. P. Lechevalier, Paris, 56:359 pp., 531 Figs. Currie, P. W. E. 1953. The "Drumming” of Meconema thalassinum Fabr. Entom. Record 65:93-94. Gurney, A. B. 1960. Meconema thalassinum, a European Katydid New to the United States. Proc. Ent. Soc. Wash. 62:95-96. . 1960a. Meconema Taken in the United States in 1957. (Orthoptera: Tettigoniidae). Proc. Ent. Soc. Wash. 62:279. Harz, K. 1955. Das Trommeln der Eichenschrecke {Meconema thalassinum De Geer). Nachrichtenbl. Bayer. Ent. Miinchen 4:91-93. . 1957. Die Geradfliigler Mitteleuropas. G. Fischer, Jena, 494 pp., 255 Abb. Johnstone, D. E. 1970. Notes on the Palaearctic Grasshopper, Meconema thalassinum (De Geer). (Orthoptera: Tettigoniidae; Meconematinae) Established in Long Island, N.Y. Ent. News 81:62-65, 2 Figs. Ragge, D. R. 1965. Grasshoppers, Crickets and Cockroaches of the British Isles. F. Warne, London, 299 pp., 22 PI., 130 Figs. Associate, Division of Insects, Field Museum of Natural History, Chi- cago, Illinois 60605. Received for publication April 12, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 42-54 A NEW SPECIES OF MEGARIS AND THE STATUS OF THE MEGARIDIDAE MCATEE & MALLOCH AND CANOPIDAE AMYOT & SERVILLE (HEMIPTERA: PENTATOMOIDEA) F. J. D. McDonald Abstract. — A new species of Megaris is described. Descriptions are given of the male genitalia of Megaris stalii, M. constricta, Canopus caesus, C. orbicularis and C. impressus. The spermatheca and external female genitalia are described for Megaris laevicollis, M. atratula, Canopus burmeisteri, C. impressus, C . fabricii, C. caesus and C. orbicularis. From a consideration of all available morphological evidence the family status of Megarididae and Canopidae is confirmed. Neither family is closely related to the Plataspidae. Very little work has been done on the interesting and unusual insects of the genera Canopis Fabricius and Megaris Stal since McAtee and Malloch’s revision in 1928. Barber (1939) added Megaris puertoricensis and Kormilev (1956) described Megaris vianai. Another species is added in this paper. The general characteristics of these two genera have been very clearly set out in McAtee and Malloch’s (1928) paper. I have attempted in this paper to examine these genera in more detail and come to some conclusion as to their correct status within the Pentatomoidea. The two genera were put in separate subfamilies within the family Pentatomidae by McAtee and Mal- loch. However the Pentatomidae of these authors is equivalent to the Pen- tatomoidea in most current classifications. Descriptions of the pygophore and external female genitalia are given to facilitate species recognition. Megaris rotunda n. sp. (Figs. 1-9) Typical small oval megaridid (Fig. 1) with the scutellum completely cov- ering the dorsal surface of the abdomen. Ventral surface flat (Fig. 2), dorsal surface steeply convex. Head tucked into the prothorax and protruding but little beyond it. Shiny rich reddish brown all over. Head. — Head broader than long, eyes prominent, globular. Jugae (Fig. 3) short, converging, not completely meeting at apex of tylus. Antennae four segmented; all segments provided with very long fine setae; first segment short, second longest, third and fourth subequal. Bucculae very short, indistinct. First rostral segment extending beyond bucculae by almost a half its length; rostrum not exceeding hind coxae. i VOLUME LXXXVII, NUMBER 1 43 0 07mm Figs. 1-9. Megaris rotunda n. sp. 1. Dorsal view. 2. Ventral view. 3. Apex of head, dorsal view. 4. Stink gland aperture. 5. 5th abdominal sternite. 6. Pygophore. 7. Right clasper. 8. Aedeagus. lateral view. 9. Aedeagus, dorsal view. Dorsal margin (D.m.), endophallic duct (E.d.), ventral margin (V.m.). 44 NEW YORK ENTOMOLOGICAL SOCIETY Thorax. — Pronotum steeply declivous anteriorly, lateral angles with a rounded protuberance. Mesoscutellum covering abdomen entirely excepting corium where exposed by incision in anterolateral margins of mesoscutel- lum. Hemelytron (Fig. I) heavily sclerotized along anterior half of margin, this portion terminating in large triangular red callus; remainder membra- neous, smoky brown. Membrane with a number of faint cross veins. Pos- terior wing membraneous with a number of sclerotized veins. Hemelytron folded under scutellum; fold occurs at junction of callus and membrane. Propleura raised anteriorly into collar, deeply grooved by sternum. Meso- and metapleura flat and plate-like. Metathoracic stink gland (Fig. 4) orifice minute, represented by an oval raised area; no evaporative area developed. Coxae small, flattened, lying close to sterna; trochanters about same size as coxae, fused to the femora. Femora elongate, swollen medianly. Tibiae shorter than femora, uniform in diameter, provided with numerous fine short setae. Tarsi two segmented; first segment half the length of the second, latter bearing a pair of claws and pulvilli; both segments with a number of short fine setae. Abdomen. — Sterna bowed cephalad medianly; sutures delimiting each seg- ment terminating before reaching lateral margin. Spiracles near lateral mar- gins of segments 2-7; trichobothria paired (Fig. 5), placed one behind other, very slightly laterad of spiracles. Male genitalia. — (Figs. 6-9.) Pygophore opening facing caudad (Fig. 6), dorsal and ventral margin flattened into a broad rim surrounding the open- ing. Proctiger small, box-like. Claspers (Fig. 7) minute, L-shaped, apically acute. Aedeagus (Figs. 8, 9.) Basal plates large in relation to theca, latter squat, cylindrical, lightly sclerotized. Conjunctival appendages not apparent, mem- braneous sheath surrounding apex of endophallic duct; latter short and tu- bular. Female genitalia — not seen. Diagnostic measurements: Length 2. 18 mm Breadth 1 .87 mm Width between eyes 0.37 mm Rostrum 1.15 mm Antennal segment I 0.22 mm 11 0.50 mm III 0.44 mm IV 0.43 mm Type. — Holotype, male, labeled: Brasilien, Nova Teutonia, 27°1TB 52° I VOLUME LXXXVII, NUMBER 1 45 0-1 mm 0 08mm 0-lmm 14. O-Imm Figs. 10-12. Megaris stall. 10. Pygophore. 11. Right clasper. 12. Aedeagus, dorsal view. Figs. 13-15. Megans constricta. 13. Pygophore. 14. Right clasper. 15. Aedeagus, lateral view. , 23'L, 300.500 m. 11 . 1.1973. Fritz Plauman. (Deposited United States Nation- I al Museum.) 1 Description of the Genitalia of Some Species of Megans and Canopus Male genitalia. Megaris stalii McAtee and Malloch (Figs. 10-12). Pygophore (Fig. 10). Opening dorsal, surrounded by wide flattened area. Ventral border slightly sinuous with a vertical outer face forming caudal face of pygophore. Dorsal border bearing spine-like protuberances on each side above claspers; central border broadly arched, partially enclosing base of box-like proctiger. Clasp- ers (Fig. 1 1), minute, conical, flattened, lying in rounded recess at each side beneath the ventral border. Aedeagus (Fig. 12). Theca small, tubular, lightly sclerotized. One pair of strap-like conjunctival appendages. Ejaculatory duct straight, tubular, re- tracted wholly within theca when at rest, moderately sclerotized. Megans constricta McAtee and Malloch (Figs. 13-15). Pygophore (Fig. 13). Opening facing dorsad, surrounded by wide flange, mainly concealed beneath scutellum; ventral face vertical and heavily sclerotized. Dorsal mar- gin deeply emarginate centrally, bearing small projections on each side. 46 NEW YORK ENTOMOLOGICAL SOCIETY Ventral margin straight. Proctiger tubular, lightly sclerotized, bearing a num- ber of stout setae on dorsal surface. Claspers (Fig. 14) very small with a stout stem, apically tapering to a slender curved process. Aedeagus (Fig. 15). Theca small cylindrical. One pair of membraneous conjunctival appendages (not expanded). Endophallic duct small tubular and slightly sinuous, moderately sclerotized. Canopus caesus (Germar) (Figs. 16-19). Pygophore (Fig. 16). Opening facing dorsad, ventral face of pygophore vertical. Ventral border straight; internally lying below the border is a thin arched shelf attached mesally and projecting caudad so that apex of arch is in line with ventral border. Dorsal border omega-shaped, merging lateroventrally with ventral border. Claspers (Fig. 17), lying on each side of ventral shelf, L-shaped; apical arm rod-like, bearing near apex on outer margin a long slender curved process. Medianly clasper expanded and flattened, bearing a number of long setae; basal stem narrow flattened. Aedeagus (Figs. 18, 19). Theca small, squat, heavily sclerotized except for a band around apical margin and basally. One pair of strap-like mem- braneous conjunctival appendages, basally with a small lobe. Median penial lobes fused, membraneous except for outer margin which is heavily scler- otized, basally surrounding ejaculatory duct; latter thick, cylindrical, heavi- j ly sclerotized, bearing on dorsal surface a small elongate apically acute process. Canopus orbicularis Horvath (Figs. 20-22). Pygophore (Fig. 20). Opening facing dorsad; ventral border straight; internally ventral wall of pygophore bearing an oblong plate attached midway at its base; plate projects freely upwards, bowed. Dorsal border omega shaped, surrounding base of proc- tiger centrally; latter box-like. Claspers (Fig. 21), L-shaped; apical arm flat- tened, blade-like, bearing near apex on outer margin a long thin curved process; mid portion of clasper expanded and somewhat flattened, bearing a number of long fine setae; stem of clasper cylindrical. Aedeagus (Fig. 22). Theca squat, heavily sclerotized except for apical rim. One pair of membraneous strap-like conjunctival appendages, apically bifid with two short rounded lobes. Median penial lobes fused into trough- like structure beneath vesica; ventral surface well sclerotized, dorsal surface Figs. 16-19. Canopus caesus. 16. Pygophore. 17. Right clasper. 18. Aedeagus, lateral view. 19. Aedeagus, ventral view. Figs. 20-22. Canopus orbicularis. 20. Pygophore. 21. Right clasper, lateral view. 22. Ae- deagus. lateral view. Figs. 23-26. Canopus irnpressus. 23. Pygophore. 24. Right clasper, lateral view. 25. Ae- deagus, lateral view. 26. Aedeagus, ventral view. Conjunctival appendage (C.a.), sclerotized process (S.p.). VOLUME LXXXVII, NUMBER 1 47 0 15mm 0 5 mm 48 NEW YORK ENTOMOLOGICAL SOCIETY membraneous. Endophallic duct large, tubular, heavily sclerotized, bearing on ventral surface a heavily sclerotized process, apically triangular, tapering basally. Canopus impressus (Fabricius) (Figs. 23-26). Pygophore (Fig. 23). Ven- tral border straight, medianly somewhat thicker than in C. orbicularis, in- ternally bearing a stalked oval plate-like projection somewhat similar to one found in C. orbicularis. Dorsal border with deep median V-shaped incision, laterally merging with ventral border; proctiger oblong, flattened, enclosed partially in median incision in dorsal border. Claspers (Fig. 24), shallow hook-shaped; anterior arm cylindrical apically, broadly rounded, bearing small curved process near apex on outer margin; clasper flattened medianly, bearing a number of long fine setae; stem cylindrical, tapering basally. Aedeagus (Figs. 25, 26). Theca squat, cylindrical, bearing oblong shield- like extension on ventral margin; lateral and ventral apical margins unscler- otized. One pair of flattened blade-like conjunctival appendages sclerotized except for apices, latter bifid, produced into two short rounded arms. Me- dian penial lobes fused into sclerotized trough-like structure lying below endophallic duct, latter large, tubular, sclerotized, bearing elongate process on ventral margin. Notes. — These three species are very similar showing a distinct gradation in characters. The process on the clasper is smallest in C. impressus and longest in C. caesus and C. orbicularis. The plate lying beneath the ventral i margin varies in shape in the three species being almost free in C. impressus and more elongate in C. caesus. The aedeagus also is very similar in the three species. Canopus burmeisteri McAtee and Malloch (Figs. 27-30). Pygophore (Fig. 27). Opening surrounded by wide flat flange, dorsal border smoothly arched, ventral border straight. Proctiger small, tubular. Claspers (Fig. 28) elongate rod-like, apically tapering into a long thin curved process, a number of long setae found mid-laterally. Aedeagus (Figs. 29, 30). Theca small, tubular, bearing large ring-like col- lar when expanded out, normally recessed within theca. One pair of elongate tubular conjunctival appendages, feebly sclerotized basally, apically mem- braneous. Median penial lobes fused into sclerotized trough-like structure lying beneath the endophallic duct; latter stout, oblong, heavily sclerotized. Canopus germari McAtee and Malloch (Figs. 31-35). Pygophore (Fig. 31). Opening surrounded by wide flat flange, ventral margin straight, dorsal margin arched with two sharp V-shaped projections on either side of base of proctiger. Latter small, tubular. Claspers (Fig. 32) broad centrally, bear- ing a number of long setae, apically tapering and drawn into a fine hook; basally clasper with a tubular stem. Aedeagus (Figs. 33-35). Theca small, squat, cylindrical, bearing mem- VOLUME LXXXVII, NUMBER 1 49 Figs. 27-30. Canopus burmeisteri. 27. Pygophore. 28. Right clasper, lateral view. 29. Ae- deagus, lateral view. 30. Aedeagus, ventral view. Figs. 31-35. Canopus gennari. 31. Pygophore. 32. Right clasper, lateral inner view. 33. Aedeagus, lateral view. 34. Aedeagus, ventro-apical view. 35. Median penial lobes. ' braneous collar apically, invaginated within theca when at rest. One pair of ' flat oblong conjunctival appendages (probably tubular when fully expanded), ; apically membraneous, sclerotized towards base. Median penial lobes fused I into a pyriform body (viewed ventrally), lying beneath the apex of the en- 50 NEW YORK ENTOMOLOGICAL SOCIETY dophallic duct; margins of lobes curved inwards forming a trough-like struc- ture on the dorsal surface. Endophallic duct tubular, slightly sinuous, mod- erately sclerotized, apex diffuse, the whole structure lying in the trough formed by median penial lobes. Canopus fahricii McAtee and Malloch (Figs. 36-38). Pygophore similar to C. burmeisteri, ventral margin slightly thickened. Claspers (Fig. 36), very similar to C. burmeisteri, with a large number of setae and somewhat stouter centrally. Aedeagus (Figs. 37, 38). Theca squat, cylindrical, bearing apical sheath normally retracted within theca. One pair of conjunctival appendages, each bifid, one short arm and one longer tapering arm, longer arm membraneous slightly sclerotized basally. Median penial lobes fused into a boat-like struc- ture lying below the endophallic duct; latter stout, sinuous. Notes. This species differs from C. burmeisteri in the following regards. Conjunctival appendages bifid, single in burmeisteri, median penial lobe smaller and differing in shape, burmeisteri more trough-like; ejaculatory duct slender, thicker and stouter in burmeisteri. Female genitalia. Megaris laevicollis Stal (Figs. 39. 40). Spermatheca (Fig. 39) simple glob- ular structure surrounded by a larger membraneous sac. No sclerites present around spermathecal opening. External genitalia (Fig. 40). Paratergites 8 fused centrally by narrow me- dian band. Paratergites 9 lying one on either side of oval tenth segment, each paratergite concave centrally and fitting around tenth segment. First gonocoxae large, plate-like, mostly hidden under abdominal sterna; apical margins of each gonocoxa turned upwards forming all that is visible exter- nally, which consists of a small oblong plate tapering to a fine point laterad. Megaris atratula StM (Fig. 41). Spermatheca a simple bulb with a small elongate appendix apically. No sclerites surrounding spermathecal opening. External genitalia (Fig. 40) similar to M. laevicollis. Canopus burmeisteri McAtee and Malloch (Fig. 42). Spermatheca (Fig. 41) consisting of well developed pump with proximal and distal flanges con- nected to sclerotized wheel-like spermathecal dilation. Duct from pump en- ters one side of spermathecal dilation centrally and a duct exits from other side centrally connecting spermatheca to vulva. Spermathecal dilation pro- vided with a number of internal circular canals. Entrance of spermatheca into vulva without surrounding sclerites. Large paired sac-like accessory glands found ventrally, one on each side of spermathecal entrance. Vulva provided with paired interlocking sclerotized rami. External genitalia described by McAtee and Malloch. Figs. 36-38. Canopus fabricii. 36. Right clasper, inner view. 37. Aedeagus, lateral view. 38. Aedeagus, ventral view. Figs. 39-40. Megaris laevicoUis. 39. Spermatheca. 40. Female genitalia. Fig. 41. Megaris atratula. Spermatheca. Fig. 42. Canopus burmeisteri. Spermatheca. Fig. 43. Canopus impressus. Spermatheca. Fig. 44. Canopus orbicularis. Spermatheca. Figs. 45-46. Canopus fabricii. 45. Spermatheca and accessory glands. 46. Spermathecal bulb. Accessory glands (A.g.), spermathecal dilation (D.), first gonocoxite (1 Gx), pump (P.). paratergite 8 (Pt. 8), paratergite 9 (Pt. 9). 52 NEW YORK ENTOMOLOGICAL SOCIETY Canopus impressus Fabricius (Figs. 43, 44). Similar to C. bunneisteri, interlocking rami and accessory glands present. Canopus fahricii McAtee and Malloch (Figs. 45, 46). Similar to C. bur- meisteri. Accessory glands (Fig. 45) large, covered with minute spines. Spermathecal dilation with internal striations. Interlocking rami present. Canopus orbicularis Horvath. Similar to C. bunneisteri. Duct between spermathecal opening and dilation longer and more coiled than in C. bur- meisteri. Interlocking rami and accessory glands present. Discussion It is clear from an examination of the male and female genitalia and of other characters cited by McAtee and Malloch (1928) that the genera Me- garis and Canopus each warrant family status. The Canopidae have a well developed series of parallel veins in the fore- wing whereas the Megarididae have no veins or only one major vein in the membrane. The male genitalia of the three megaridids examined are very simple, consisting of an unsclerotized conjunctiva surrounding a tube-like endophallic duct. The female spermatheca in two species examined is also simple and sac-like without a complicated pumping mechanism. The canopids examined all show much more highly evolved male and female genitalia. The male genitalia have well defined conjunctival appendages to- gether with conjunctival processes associated with the endophallic duct. The females have a complex spermatheca with a well developed circular dilation or reservoir together with a well developed pumping mechanism. The vulva has paired interlocking rami on each side resembling those found in some species of Scutelleridae, and paired accessory glands were also found resembling some species of Cydnidae (McDonald, 1966). Preliminary examination of the male and female genitalia of some species of Australian Plataspidae indicates that neither the canopids nor megaridids are closely related to this family (McAtee and Malloch, 1928). In the struc- ture of the wing venation, however, the Canopidae do show some affinity with the Plataspidae in possessing parallel venation in the membrane. The spermatheca in the plataspids (McDonald, 1970) has a well developed pump but no dilation or reservoir in the duct. The males have a well developed aedeagus (Figs. 47-53) with conjunctival appendages, but these do not re- semble the type of aedeagus found in either the Megarididae or Canopidae. The abdominal sutures in the nymphs (McAtee and Malloch, 1928) also distinguish these two families from the Plataspidae. From the information so far obtained it would appear that the Megaridi- dae, Canopidae and Plataspidae, while superficially resembling one another in possessing a well developed scutellum, are quite clearly separate families VOLUME LXXXVII, NUMBER 1 53 Figs. 47-49. Brachyplatys flavipes. 47. Pygophore. 48. Aedeagus, lateral view. 49. Aedea- gus, ventral view. Figs. 50-51. Coptosoma falloui. 50. Vesica, lateral view. 51. Vesica, dorsal view. Figs. 52-53. Coptosoma hemispherica. 52. Vesica, lateral view. 53. Vesica, dorsal view. 54 NEW YORK ENTOMOLOGICAL SOCIETY and are not closely related. The Megarididae are very primitive and are probably an early offshoot from the Pentatomoid line of evolution. The Canopidae are more highly evolved and show some affinity to the Scutel- leridae. The Plataspidae have some affinity with the Pentatomidae but are not closely allied to them. Acknowledgments I should like to thank the following: Dr. R. C. Froeschner, United States National Museum; Dr. Randall T. Shuh, American Museum of Natural His- tory; Dr. Per Inge Persson, Swedish Museum of Natural History; Mr. W. R. Dolling, British Museum (Natural History); Dr. Paul H. Arnaud Jr., California Academy of Sciences; and Dr. Tamas Vasarhelyi, Hungarian Natural History Museum for the loan of type and other material used in this study. I should also like to thank Professor L. H. Rolston for his help and en- couragement on this project which was carried out in the Department of Entomology, Louisiana State University, Baton Rouge. I am also indebted to him for reading and criticising the manuscript. Literature Cited Barber, H. G. 1939. Scientific survey of Puerto Rico and the Virgin Islands, Vol. XIV (3). Insects of Puerto Rico and the Virgin Islands — Hemiptera — Heteroptera (excepting the Miridae and Corixidae). N.Y. Acad. Sci., N. Y.:263-441 . Kormilev, N. A. 1956. Notas sobre Pentatomoidea neotropicales iv, v. (Hempitera). Acta scientifica de los Institutos de Investigacion de San Miguel 3:1-12. McAtee, W. L. and J. R. Malloch. 1928. Synopsis of Pentatomid bugs of the Subfamilies Megaridinae and Canopinae. Proc. U.S. Nat. Mus. 72:1-21, 2 pis. McDonald, F. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera: Het- eroptera). Quaest. Ent. 2:7-150. . 1970. The morphology of Lestonia haustorifera China (Heteroptera: Lestoniidae). J. Nat. Hist. 4:413^17. Department of Plant Pathology and Agricultural Entomology, University of Sydney, Sydney, 2006, Australia. Received for publication September 15, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 55-58 ; AUTOGRAPHA CALIFORNICA NUCLEAR POLYHEDROSIS i VIRUS (NPV) IN A VERTEBRATE CELL LINE; I LOCALIZATION BY ELECTRON MICROSCOPY j Arthur H. McIntosh, Karl Maramorosch and Russell Riscoe I Abstract. — The nuclear polyhedrosis virus (NPV) of Autographa califor- nica has been localized by electron microscopy in the cytoplasm of poikilo- thermic vertebrate cells (VSW). C-type particles carried by this cell line were also visualized together with baculovirus particles in a vacuole-like I organelle. Inoculation of VSW cells with A. californica NPV had a retarding ! effect on cell growth. Nuclear polyhedrosis viruses (NPVs) are the agents of choice for the I control of agricultural and forests pests (WHO 1973). Their selection is based on their specificity for insect hosts and their apparent innocuous na- i ture for higher animals (Heimpel 1966; Heimpel and Buchanan, 1967; Ig- noffo and Heimpel 1965). However, it is important that both in vivo and in vitro systems become available for the evaluation of such agents prior to widespread use in the field. There have been no reports concerning the successful replication of NPVs in vertebrate cells other than that of Himeno et al. 1967. Amongst the theories purported to explain this is that virus does not penetrate vertebrate cells. In the present report we tested this premise. A vertebrate viper cell line (VSW) known to replicate an insect Chilo irri- descent virus (McIntosh and Kimura, 1974) was inoculated with A. califor- nica AC NPV (Vail et al. 1970) at a multiplicity of infection of 1.5 (TCID50/ cell). Cultures were incubated at 28°C for 5 days, then prepared for electron microscopy as described previously (McIntosh and Kimura 1974). The results of the electron microscopy study are present in Figs. 1 and 2. NPV particles (arrows) can be readily seen intracellularly amongst the C-type particles which are carried by this line (Fig. 1). In Fig. 2 a NPV particle with envelope partially removed can be seen extracellularly with many C-type virions. No increase in virus titer could be observed over the 5-day incubation period and this confirms our previous finding (McIntosh and Shamy 1975). However, it was observed that many more C-type par- ticles were present in inoculated cultures than in unchallenged cultures. Since C-type particles do not produce a cytopathic effect, it is not possible to titrate them until such a system is made available. Particle counting using the electron microscope would be an alternative means of quantitating them. An interesting result was the marked retardation in cellular growth fol- lowing challenge with AC NPV. At the end of 1 week incubation cell counts 56 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 1. Intracellular localization of A. californka NPV particles (arrows) in a vertebrate VSW cell. Note the presence of C-type particles. x60,000. were twice as high in control cultures as in inoculated cultures. When sub- cultures were made by making a 1:2 split the subculture from the inoculated VSW did not grow out and the pH became very alkaline. Control VSW cells challenged with extracts of the insect cell line TN-368 (Hink 1970) in which AC NPV is propagated, grew normally. The present findings show that AC NPV is incorporated intracellularly into the vertebrate VSW cell line. The means by which such incorporation occurs are unknown. It is also apparent that intracellular localization of virus particles has a retarding effect on cell growth. It thus appears that failure of AC NPV to replicate in the VSW cell line is not due to lack of uptake by the cells. Acknowledgments This investigation was supported in part by a National Science Founda- tion grant BMS 74-13608, a Rockefeller Foundation grant 75074 GA AGR 7621, a PHS grant AI- 14509, and a Charles and Johanna Busch award. We thank Mrs. R. Shamy for preparation of the electron microscopy plates. VOLUME LXXXVIl, NUMBER 1 57 Fig. 2. Extracellular abundance of C-type particles following inoculation of VSW with A. californica NPV. A virus particle (arrow) can be seen with partially removed envelope. X 45 ,000. Literature Cited Himeno, M., F. Sakai, K. Onodera, H. Nakai, T. Fukada and Y. Kawade. 1967. For- mation of nuclear polyhedral bodies and nuclear polyhedrosis virus of silkworm in mam- malian cells infected with viral DNA. Virology, 33:507-512. Hink, W. F. 1970. Established insect cell line from the cabbage looper, Trichoplusia ni. Nature (London) 226:466-467. Heimpel, A. M. 1966. Exposure of white mice and guinea pigs to the nuclear polyhedrosis virus of the cabbage looper, Trichoplusia ni. J. Invertebr. Pathol. 8:98-102. Heimpel, A. M. and C. K. Buchanan. 1967. Human feeding tests using the nuclear polyhe- drosis virus. J. Invertebr. Pathol. 9:55-57. Ignoffo, C. M. and A. M. Heimpel. 1965. The nuclear polyhedrosis virus of Heliolhis zea and Heliothis virescens. V. Toxicity-pathogenicity of virus to white mice and guinea pigs. J. Invertebr. Pathol. 7:329-340. McIntosh, A. H. and M. Kimura. 1974. Replication of the insect chilo irridescent virus (CIV) in a poikilothermic vertebrate cell line. Intervirology 4:257-267. McIntosh, A. H. and R. Shamy. 1975. Effects of the nuclear polyhedrosis virus (NPV) of Autographa californica on a vertebrate viper cell line. Ann. N. Y. Acad. Sci. 266:327- 331. 58 NEW YORK ENTOMOLOGICAL SOCIETY Vail, P. V., D. L. Jay and D. K. Hunter. 1970. Cross infectivity of a nuclear polyhedrosis virus isolated from the alfalfa looper, Autographa californica. Proc. 4th Internat. Colloq. Insect. Pathol, pp. 297-304. College Park, Maryland, U.S. World Health Organization Technical Report Series. 1973. The use of viruses for the control of insect pests and disease vectors. No. 531. pp. 5-48. Waksman Institute of Microbiology, Rutgers University, P. O. Box 759, Piscataway, New Jersey 08854. Received for publication July 15, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(I), 1979, pp. 59-65 TWO NEW SPECIES OE LACCOBIUS EROM EASTERN NORTH AMERICA (COLEOPTERA: HYDROPHILIDAE) Stanley E. Malcolm Abstract. — Laccohius reflexipenis and L. spangleri are described from eastern North America. Holotypes and paratypes are designated. Male gen- italia and pronotal maculation are figured. Distribution maps are included. In “The Water Beetles of Maine” (Malcolm, 1971) I mentioned two species of Laccobius to be described by two other taxonomists, Brian S. I Cheary and Ronald B. Willson. Because my paper included significant char- acters and appeared while those of Cheary and Willson were in press, I inadvertently became descriptor of the two species. In order to clarify this unfortunate situation, I include full descriptions of the two species and des- I ignate types. The publications of Cheary and Willson have not yet appeared. I The following standard abbreviations for collections are used: BMNH j — British Museum (Natural History); CAS — California Academy of Sci- I ences; CNC — Canadian National Collection; DCM — David C. Miller; j EMNH — Field Museum of Natural History; SEM — Stanley E. Malcolm; I UCR — University of California at Riverside; UM — University of Maine; i USNM — U.S. National Museum of Natural History, j Laccobius reflexipenis n. sp., Malcolm Laccobius reflexipenis "Cheary.” Malcolm 1971. Univ. Maine Agri. Exp. Sta. Tech. Bull. 48:40^2. Holotype. Male; Maine, E. Corinth, Penobscot Co. July 17, 1969, Stan Malcolm; USNM type number 71356. Form oval; length 2.8 mm; width 1.7 mm; head with frequent medium sized punctures overlying a strongly microreticulate surface, color metallic bronze-green except a triangular pale area anterior to the eye; pronotum punctured and reticulate as head, disc metallic bronze-green, margins pale, (for pronotal spot pattern see Fig. la); scutellum metallic bronze-green; elytra pale, medium sized dark brown punctures arranged in imperfect series overlying moderately microreticulate surface, some puncture marks co- alesced along suture at midpoint to form a dark spot, punctures at apices and margins only slightly darkened; ventral surface dark brown, covered with dense hydrofuge pubescence; coxae dark brown, trochanters and fern- 60 NEW YORK ENTOMOLOGICAL SOCIETY A B Fig. I. Pronotal spot patterns: a. Laccohiiis reflexipenis Malcolm, b. Laccobiiis spangleri Malcolm. ora brown except lighter at distal ends, tibiae and tarsi pale, forefemora with dense hydrofuge pubescence on proximal third; palpi and antennae pale except antennal club brown; aedeagus with median lobe filiform, bowed in lateral view; parameres with recurved extensions in ventral view, tips of parameres downcurved, rounded (Fig. 2a). Variation. Length 2.4 to 3.4 mm, males average 2.6 mm, females 3.0 mm; metallic colored areas often bronze-red, sometimes brown; pale background coloration often darker, obscuring pronotal spot pattern and changing (often enhancing) elytral maculation. The great variation between individuals in coloration and size necessitates the use of male genitalia to distinguish L. reflexipenis from sympatric species. Specimens examined. 323; Distribution is plotted on map I. Paratypes bearing the cited locality data have been deposited in the following collec- tions: 1 <3, 1 9, Indiana, U.S.A., Starke Co., W. S. Blatchley Coll. (BMNH); 2 c3. Mo. Ripley Co., Buffalo Creek at Route C, 5.5 mi N. of Briar, 5-VIII-1967, collector H. B. Leech (CAS); 1 d, Boiestown, N. B., VII-1 1-1928, W. J. Brown, 1 d, Knowlton, Que., VI-26-1928, G. H. Fisk (CNC); 1 d, MAINE, Corinth, Penobscot Co., 8-13-1969, Stan Malcolm (DCM); 1 d, IOWA, Scott Co., Davenport, V-10-1964, leg. S. B. Peck (FMNH); 1 d, 1 9, E. of Beddington, ME, VIII-1-1970, Rt. 9, Stan Malcolm (SEM); 1 d, Indiana, U.S.A., Starke Co., W. S. Blatchley Coll. (UCR); 2 d, MO., 7 mi W. of Weldon Spgs., VI-25-1954, Paul Spangler, 1 d, 8 mi W. Steelville, MO. Meramec River, X-10-1953, M. C. Grabau (USNM). 62 NEW YORK ENTOMOLOGICAL SOCIETY Map 1. Distribution of Laccobius reflexipenis Malcolm. Laccobius spangleri n. sp., Malcolm Laccobius spangleri “Willson.” Malcolm 1971. Univ. Maine Agri. Exp. Sta. Tech. Bull. 48:40^2. Holotype. Male; Dixmont Ctr., Maine VII-20-1970, S. Malcolm; USNM type number 73365. Form oval; length 2.5 mm; width 1.6 mm; head with frequent medium sized punctures overlying a strongly microreticulate surface, color metallic bronze-green except a triangular pale area anterior to the eyes; pronotum punctured and reticulate as head, disc metallic bronze-green, margins pale, (for pronotal spot pattern see Fig. lb); scutellum metallic bronze-green; elytra pale, medium sized metallic bronze-green punctures arranged in im- VOLUME LXXXVII, NUMBER 1 63 I Map 2. Distribution of Laccobius spangleri Malcolm. perfect series alternating with less regular rows of punctures overlying light- ly microreticulate surface, some puncture marks coalesced along suture at midpoint to form a dark spot, other vague dark spots similarly formed dis- tributed in a pattern typical of many Laccobius species, punctures at apices and margins with little darker marking; ventral surface and coxae dark brown, covered with dense hydrofuge pubescence; legs brown proximally, gradually lighter from mid-femora distally, forefemora with dense hydrofuge pubescence in basal third; palpi and antennae pale except antennal club brown; aedeagus with median lobe filiform, bowed in lateral view; para- meres with overlapping semi-membranous extensions in ventral view, tips of parameres downcurved, rounded, concave below (Fig. 2b). 64 NEW YORK ENTOMOLOGICAL SOCIETY Variation. Length 2.3 to 3.2 mm, males average 2.6 mm, females 2.9 mm; metallic colored areas often bronze-red, sometimes brown; background col- oration may be lighter or considerably darker than holotype, obscuring pro- notal spot pattern and changing elytral maculation. As with the preceding species, the great variation between individuals in coloration and size ne- cessitates the use of male genitalia to distinguish L. spangleri from sym- patric species. Specimens examined. 454; Distribution is plotted on map 2. Paratypes bearing the cited locality data have been deposited in the following collec- tions: I (5, 1 9, Indiana, U.S.A., Starke Co., W. S. Blatchley Coll. (BMNH); 1 c?, 1 9, 7-VII-1946, Cornwall, CONN., Spring Brook, Cham- berlain Collector (CAS); 1 d, 1 9, Ottawa, Ont., 15-V-1930, W. J. Brown (CNC); 1 d, 1 9, Dixmont Ctr., ME., VII-20-1970, S. Malcolm (DCM); 1 d, 1 9 , MAINE, Oxbow, Aroostook Co., July 7, ’69, Stan Malcolm (SEM); 1 d, Mohawk, MICH., Gratiot River, Keweenaw C., 7 July 1964, R. B. Willson, 1 d, Allendale, MICH., Ottawa Co., 17 July 1963, R. B. Willson (UCR); 2 d, MAINE, Harmony, Somerset Co., Aug. 3, 1969, Stan Malcolm (UM); I d, 1 9, 4 mi E. Levering, Emmet Co., Mich. 43b, VIII-5-1952, Paul J. Spangler (USNM). For both species the male genitalia are diagnostic. Males can be recog- nized by their generally smaller size and dilated second foretarsal segment. As with many species of Laccobius, females can only certainly be deter- mined by association with males. The pronotal spot pattern is variable, but may be of use in separating these species from other Laccobius. In Maine I have collected these species together and with L. minutoides. D’Orchymont ( 1942) included a figure of the male genitalia in his description of L. minutoides . Acknowledgments I thank the following individuals for the loan of specimens and for valuable advice: Mr. M. E. Bacchus (BMNH), Dr. David H. Kavanaugh (CAS), Dr. A. Smetana (CNC), Dr. David C. Miller (DCM), Mr. Eric H. Smith (FMNH), Dr. Saul Frommer (UCR), Dr. John Dimond (UM), Dr. Paul J. Spangler (USNM), Dr. Brian S. Cheary, and Mr. Ronald B. Willson. 1 am most grateful to Drs. James A. Slater, David C. Miller, and Paul J. Spangler for critical review of this manuscript. My sincere thanks also to the illus- trators. Ms. Mary Jane Spring and Ms. Molly Hubbard. Literature Cited D'Orchymont. A. 1942. Revision des Laccohius Americains (Coleoptera Hydrophilinae Hy- drobiini). Bull. Mus. R. d'Hist. Nat. Belg. 18:13-18. VOLUME LXXXVll, NUMBER 1 65 Malcolm, S. E. 1971. The Water Beetles of Maine: Including the Families Gyrinidae, Hali- plidae, Dytiscidae, Noteridae, and Hydrophilidae. Univ. Maine Agri. Exp. Sta. Tech. Bull. 48. 49 pp. Biological Sciences Group, Box U-43, The University of Connecticut, Storrs, CT 06268. Received for publication July 21, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 66-77 THE EVOLUTION OF EYESPOTS IN TROPICAL BUTTERFLIES IN RESPONSE TO FEEDING ON ROTTING FRUIT: AN HYPOTHESIS Allen M. Young Abstract. — A substantial portion of the butterfly species in tropical forests of Central and South America feed primarily as adults, on rotting fruits and to a lesser degree, on moldy (fermenting) sap issuing from wounds in trees. In general, exploitation of fruit frequently occurs on the ground, where volatile odoriferous substances, released in decay, attract butterflies in the three nymphalid subfamilies Satyrinae, Brassolinae, and Morphinae; simi- larly, sap and hanging injured or rotting fruit are exploited arboreally by several genera of the Nymphalinae. However, a few nymphalines also feed on fallen fruit and experimentally placed fruit baits on the ground. This paper explores in a preliminary way the possible adaptive relations between (1) feeding on the ground versus feeding arboreally, (2) dispersal agents (vertebrates) as opportunistic predators of feeding butterflies, (3) functional role of eyespot markings (eyespots) on the undersides of wings, and (4) impairment of escape behavior of butterflies from intoxication acquired by feeding on rotting fruits. The general thesis is advanced that butterflies be- come intoxicated in nature from feeding on rotting fruits, and that eyespots increase the margin for successful escape when normal behavior has been impaired. Since fallen fruits ferment quickly, the intoxication of butterflies is greatest on the forest floor and eyespots are most prevalent in the bras- solines, satyrines, and morphos, the three groups that thrive here. Very often, feeding takes place in patches of forest floor directly exposed to sunlight, and at times of the day when such exposure is maximized. Under conditions of direct sunlight, eyespot markings are very noticeable. A high selective value is placed on eyespots as an additional line of evasive escape | behavior since animals attracted to fallen fruit (for dispersal purposes and feeding on insects) can be predators on butterflies. Eyespots are known to ■ lure pecking and biting away from the body of a butterfly or moth. Alter- natively, it is also known that eyespots frighten away animals. Either way, they function to reduce the likelihood of attack on an insect by a vertebrate predator. It is easy to conceptualize how the three groups of butterflies , developed similar eyespots as they are closely linked phylogenetically. For | the arboreal-feeding nymphalines, eyespots are expected to be less func- tional as indicated by their conspicuous absence in most species. Such mark- i ings are also generally absent from most flower-visiting butterflies. The de- composition of fallen fruits and the yeasting of sap flows are processes that | play a major role in maintaining the butterfly community of tropical forests. | VOLUME LXXXVII, NUMBER 1 67 As microbes render fruit less savory for vertebrates that act as dispersal agents, they open a feeding niche for butterflies. Introduction Each year, the lowland tropical rain forests of Central and South America contain collections of fallen fruits from various tree species. Most of these, such as Coumarouna oleifera (Leguminosae) are sweet-smelling rotting, fleshy fruits which attract a variety of butterflies, predominately in the nym- phalid subfamilies Satyrinae, Brassolinae, and Morphinae (e.g.. Young 1972a). Two major features of these interactions between fruits and butter- flies are: (a) although the fallen fruit collection can be large under a parent tree, only certain fruits are returned to repeatedly by butterflies, and (b) an individual of a butterfly species participating in the interaction often returns to the site many times over several days or weeks. The observed extension of the proboscis on, or into, the fruit indicates that the butterfly is ingesting fluids seeping from the fruit. After a fruit matures, it generally falls to the ground where it sits for some period that may be terminated by dispersal away from the area by a ver- tebrate (Smythe 1970); if such dispersal does not occur, the ripe fruit spoils and begins to rot as a result of microbial action on the fruit wall. Spoilage is usually accompanied by the emission of a sweet or tangy odor from the fruit. At this stage, the fruit becomes a feeding patch for many insects, including butterflies. The selective advantages for microbe species to attack fallen fruits prior to the onset of dispersal by vertebrates have been discussed (Janzen 1977). Along with the microbes causing the decay, the insects associated with these fruits, including Lepidoptera, are also conceptualized as being in competi- tion with vertebrates (fruit and seed dispersal agents) for the fruit as a food resource (Janzen 1977). Although the feeding on rotting fruits and fermented fruit products by Lepidoptera and other insects has been documented ex- tensively (e.g., Dethier 1947; Frost 1928; Norris 1936; Barcant 1970; Gilbert 1972; Muyshondt 1973a, b; Young 1972a; Gomez 1977 and many others), little attention has been given to the likely ecological consequences of such feeding behavior. The purpose of this note is to call attention to some pos- sible features of these interactions that relate to a predicted loss in fitness resulting from such feeding, and the role of cryptic coloration and behavior of day-flying Rhopalocera in offsetting these effects. These comments are made as an argument for further documentation of these interactions, es- pecially in the tropics. 68 NEW YORK ENTOMOLOGICAL SOCIETY Unusual Butterfly Foods Adults of many temperate and tropical butterflies imbibe juices from ripe fleshy fruit, and rotting fleshy fruits (e.g., Comstock 1895; Young 1972a ). In the American tropics, many forest-dwelling satyrines, brassolines, and morphos, exhibit this behavior (Brown 1972). In addition, many nympha- lines visit fresh arboreal sap flows (e.g., Muyshondt 1973a, b). In general, Nymphalinae such as Anaea, Prepona, Adelpha, Historis, and Marpesia exhibit this behavior. Among the brassolines, genera such as Calico, Bras- solis, and Opsiphanes have been observed to visit rotting fallen fleshy fruits (e.g., Harrison 1963; Young 1972a; Young and Muyshondt 1975). In addition to feeding on rotting fleshy fruits, both under natural conditions and when baited (Young 1972a; Young and Muyshondt 1973; Young 1973; Young and Thomason 1974; Young 1975), Murpho has also been seen imbibing fluids from moldy growths on sap flows of fallen trees (Young and Muyshondt 1973 and Fig. 1). Furthermore, Morpho feeds on yeasts associated with disposed dairy products in garbage heaps at the edge of forest at the Mon- teverde farms in Costa Rica (anonymous). An outstanding morphological feature of the Satyrinae, Brassolinae, and Morphinae, generally absent in the Nymphalinae and of significance in feed- ing behavior, is the presence of distinctive eyespots on the undersides of the wings (Fig. 1). In addition to being generally absent from the Nym- phalinae, including genera that feed arboreally on sap flows, these markings are absent from most flower-visiting butterflies. Owing to the positioning of the wings in a vertical closed position while feeding (Fig. 1), the eyespot markings (eyespots) are exposed maximally to the visual field of an ap- proaching vertebrate. All four subfamilies of fruit-and-sap-feeding butter- flies are cryptically colored in terms of the background coloration of the undersides of the wings. Spatial Temporal Variation of the Butterfly Community Owing to the very great diversity and local species abundance of nym- phalines, brassolines, and morphos in the wet forests of tropical America, it seems that the exploitation of ripe or rotting fleshy fruits and fermenting sap flows is a significant factor in determining and maintaining this subset of the butterfly community in a forest habitat. The Morphinae and Brasso- linae are strictly tropical groups, while most Satyrinae are also strictly trop- ical (Ehrlich and Raven 1965). Furthermore, the Nymphalinae are relatively far more abundant in the tropics. In southern Brazil, nymphalines, morphos, satyrines, and brassolines reach very high levels of diversity and many members of these groups exhibit little or no seasonal variation in abundance at a locality throughout the year (Brown 1972). Thus the subset of the adult VOLUME LXXXVIl, NUMBER 1 69 Fig. 1. Above: live individual of Morpho peleides, wild-caught in northeastern Costa Rica. Note vertebrate-type eyespots on the undersides of both sets of wings. Below: Morpho peleides feeding on soupy fungal growth on sap issuing from a fallen tree in northeastern Costa Rica. Time of day is late afternoon; eyespots fully exposed while feeding. 70 NEW YORK ENTOMOLOGICAL SOCIETY butterfly community associated with ripe and rotting fleshy fruits and moldy sap flows is generally active throughout much of the year, and this is also the case for the northeastern tropical wet lowland forest region of Costa Rica (pers. obs.), and presumably along much of the Atlantic watershed region of southern Central America. Owing to the generally high spatial patchiness of parent individuals of many tree species in tropical forests (e.g.. Black et al. 1950), and the sea- sonality of fruit production in many of these species (Frankie et al. 1974), rotting fallen fruits for butterflies are expected to be a variable resource from place to place, and from time to time at a given place. Thus the but- terflies are expected to exploit a succession of fruit species in space and time as availability of each fruit species changes; during periods of peak fruit production of a preferred species, and at places where the fruit is abundant, it is predicted that populations of butterfly species exploiting this food supply will be residential. Young (1972a) found that individuals of a given species of satyrine, bras- soline, or morpho tend to return to the same fruit baits over several days. Observations such as these and others for Morpho (e.g.. Young 1973; Young and Thomason 1974) suggest indirectly that the food niche of these butter- flies is probably narrow at a given time of the year. The observed tendency for individuals to return repeatedly to the same area of fallen fruit or bait each day (e.g.. Young 1972a; Young and Thomason 1974) suggests that these butterflies become familiar with certain portions of the habitat where food resources are found; there is less of a tendency to wander through large areas and rather more of a tendency to become localized near food supplies. What the relationships, if any, of such distributional patterns have to locations of larval food plants has not been determined. In general, though, these observations and those of the preceding paragraph suggest that the subset of the tropical forest butterfly community using rotting fruits and moldy sap flows is a fairly predictable collection of species in space and time. Such conditions may increase predation by home-ranging small ver- tebrates on adults. Cryptic Properties of Adult Tropical Butterflies Cryptic wing coloration in the Lepidoptera evolved as a means of allowing adults to match the surrounding environment (e.g., Wickler 1968). Most of the Neotropical fruit-eating groups such as the morphos, brassolines, and satyrines possess distinct eyespots on the undersides of the wings. These eyespots closely resemble vertebrate eyes, and several of them often occur on the wings (Fig. 1). In direct light, the eyespots are very noticeable to the human observer, while in shade they tend to blend in with the cryptic back- ground hues of the wings, and of the forest floor. It is believed that such markings stimulate pecking by birds away from the body (Blest 1957; Wick- VOLUME LXXXVII, NUMBER 1 71 ler 1968). Such markings presumably lower the risk of an individual butterfly being eaten while resting or feeding with the eyespots exposed to the visual field of predators. These markings, in combination with generally subdued colors and generally motionless posture while feeding or resting, are effec- tive deterrants for vertebrate predators. It is also known that eyespots fright- en away some animals, rather than inviting attack (e.g.. Mailman 1977). It is not known which of these types of interactions occur in tropical butterflies feeding on the forest floor, but the frequent occurrence of individuals with uncanny symmetrical sections missing from the hindwings in Morpho and Caligo suggests that these insects are attacked by small vertebrates (rep- tiles) while feeding or resting. With a few exceptions, such as Prepona, most Neotropical nymphalines lack eyespots. Yet the general coloration of the undersides of the wings is subdued and cryptic; feeding proceeds generally arboreally from moldy sap flows (e.g., Muyshondt 1973a, b). It appears, therefore, that the occurrence of small eyespots is most frequent among Neotropical butterfly groups that feed on or near the ground in forest habitats, these being the Morphinae, Brassolinae, and Satyrinae. In general, there is a burst of butterfly feeding activity just before sunset on clear, sunny days in Costa Rican lowland tropical rain forests (pers. obs.). While sections of the forest floor receive varying amounts of direct sunlight during the day and butterflies feeding on rotting fruit sometimes sit in patches of sunlight, Mailman ( 1977) mentions that Daniel Janzen has noted that direct sunlight is almost horizontal near sunset in tropical forests. Thus near-sunset feeding in butterflies may also occur in patches of direct sun- light. These conditions enhance the visual perception of eyespots by verte- brate predators while the butterflies are feeding. Some recent data from temperate zone forests indicate that vertebrates that disperse the fallen fruits of some tree species are most active in areas where direct sunlight is more abundant (Thompson and Willson 1978). Dam- age of fallen fruit by various invertebrates was cited as the major selection pressure favoring rapid dispersal of fruit in that study. Thus, a similar pat- tern may exist in tropical forests, where light gaps and other types of ex- posed areas are common. This means that small vertebrates which disperse fruit and are opportunistic predators of butterflies, could be active in sunny places where feeding butterflies maximize exposure of eyespots while feed- ing. Intoxication and Modification of Behavior Fallen ripe fruits are attacked by a wide variety of microbes, these or- ganisms producing toxins, antibiotics and other mechanisms of substrate degradation that result in fruit spoilage or rotting (many references in Janzen 1977). This rotting or fermentation releases volatile substances that attract 72 NEW YORK ENTOMOLOGICAL SOCIETY insects, including many Lepidoptera; fermenting fruit juices become avail- able as food for butterflies as the result of seeping through breaks in the fruit wall, or from general seepage through a decaying fruit wall. Rotting fruits and moldy (yeasty) sap flows are common sources for the isolation of yeasts, which are specialists on eating ripe fruit (Janzen 1977), this inter- action forming the basis for feeding by butterflies. While the spoilage of the fruit makes it less available to vertebrates, it becomes a major resource for butterflies. It is not known exactly what kinds of nutrients butterflies obtain from rotting fruits or moldy sap flows, but alcohol is imbibed since some species exhibit intoxicated behavior after feeding (e.g., Gomez 1977). There are far more observations of occasional intoxication of birds and mammals from feeding on rotting fallen fruits, rendering them easier to capture (Janzen 1977). Owing to the high humidity of the lowland tropical rain forest and the relatively small body size (and nervous system) of insects, it is likely that intoxication in butterflies is more widespread than for larger animals exploiting these food sources. Adults of Morpho peleides with abdomens prodigiously distended and swollen, as the result of extended periods of feeding at baits of rotting bananas, are unable to fly away when disturbed (pers. obs.). In some instances where feeding is less intense at baits, adults fly away but with great difficulty, making them very easy to capture with a net (pers. obs.). My experience with bait studies of Morpho and Caligo in Costa Rica from 1968 through 1976 is that the longer adults are left un- disturbed feeding at these fruits (usually bananas) the greater the level of apparent intoxication as reflected in the impairment of typical escape be- havior. Butterflies, when recently arrived at a bait and disturbed, do not exhibit such modified behavior, presumably because there has been insuf- ficient feeding for intoxication to occur. When intoxicated butterflies such as Morpho and Caligo are picked up, they excrete large quantities of fluid; when left undisturbed for a period, most fluids are excreted and eventually these individuals “sober up” and fly away. For morphos and brassolines the sequence of events in typical escape behavior elicited when they are deliberately disturbed, shortly after arriving at a bait, is a brief period of rapid, jerky wing movements followed by walking and flight. When disturbed following considerable feeding, the be- havioral sequence is to remain motionless and then an attempt to fly away. In some instances, a heavily intoxicated Morpho peleides remains motion- less even after an aerial net is placed over it. As with other animals, symp- toms of intoxication in butterflies include the tendency to sit quietly, ap- parently the result of depression of the nervous system. A consequence is that recovery time is extended and it is this interruption of the typical escape behavior that renders intoxicated butterflies easy prey for vertebrates. Intoxicated butterflies are frequently discovered at fruit baits deliberately VOLUME LXXXVII, NUMBER 1 73 placed in small piles on the forest floor (e.g.. Young 1972a; 1973; Young and Thomason 1974) whereas butterflies observed at natural food supplies are seldom intoxicated (e.g., Young 1975). Therefore, the frequency of severe intoxication under natural conditions is questionable. Introduced baits, sev- eral centimeters across and placed close-by in an area represents a highly available food supply as most natural supplies come in smaller patches and the patches are very dispersed. Thus although the fallen fruits of Coumci- rouna trees are abundant seasonally and attract Morpho amathonte, Mor- pho peleides, and many satyrines, the sweet-smelling pulpy cover on each fruit occurs as a thin coat and this coat generally dries up in a few days. Other natural food supplies include small, yeasty fresh sap flows at the base of large forest trees (Young 1975) attracting Morpho and several Satyrinae. The single exception to these observations would be the availability of large quantities of rotting banana (Afi/5a(Musaceae)) in plantations, and areas where there are extensive plantings of other fruit trees such as mango (Mangifera ( Anacardiaceae)) and guava (Psidium (Myrtaceae)). In these habitats, large quantities of rotting fruit are available to butterflies for sev- eral months each year, and intoxicated individuals of Morpho and Ccdigo have been seen (pers. obs.). Patterns of Butterflies at Fruits As the result of capture-mark-resight field studies, it has been found that by far the majority of individuals appearing at both experimental and natural rotting fruits and moldy sap flows for Morpho, Caligo and several satyrines (e.g., Antirrhea, Piriella, Caerois, Taygetis) are males (Young 1972a; Young 1973; 1975; Young and Muyshondt 1973; Young and Thomason 1974; pers. obs.). Females tend to show up more irregularly than males (Young 1972a). The scarcity of females at natural food supplies in forest habitats is less understood. But Young and Thomason (1974) found that females of Morpho peleides were far less abundant than males at fallen fruits of Giiazuma idmifolia Lam. (Sterculiaceae). A similar pattern of abundance between the sexes was seen on the sap flows of Samanea saman Merrill (Leguminosae) for the same population at a different time (Young 1975). The sex ratio of laboratory-reared Morpho peleides is unity (Young and Muyshondt 1973). It seems, therefore, that for at least Morpho peleides, there is a definite behavioral difference in feeding activities between males and females. The observed general tendency for males to predominate at bait for several species of satyrines and brassolines in addition to morphos, suggests that such a behavioral difference is widespread among these butterflies, although more data are needed to confirm this prediction. In addition to the above considerations in relation to feeding, eyespots undoubtedly function to enhance survival from predators while butterflies 74 NEW YORK ENTOMOLOGICAL SOCIETY are resting. As with feeding on the ground, morphos, satyrines, and bras- solines generally spend the night hanging from leaves near the ground (pers. obs.). During these periods the butterflies are exposed to nocturnal verte- brate predators such as lizards and small mammals. Discussion Although many insects including some nymphalines of temperate zones exploit rotting fruits and moldy sap flows (e.g., Simon and Enders 1978), such feeding behavior is more prevalent in the American tropics, where members of four major subfamilies of nymphalids, the Nymphalinae, Mor- phinae, Brassolinae, and Satyrinae (following Ehrlich 1958), contain genera and species whose adults feed almost exclusively upon these food sources. While it is known that some temperate-zone families of Lepidoptera are attracted to volatile fermentation products (e.g., Utrio and Eriksson 1977), less is known about the attraction of tropical butterflies to rotting fruits and moldy sap flows. Bait studies indicate that butterflies become intoxicated from extended periods of feeding on an abundant fruit supply and fruit products (i.e., large resource patch). Although intoxication is less likely for butterflies exploiting natural food sources which tend to be smaller and dispensed in space and time, it is expected that some individuals of a species would experience intoxication, thus increasing the likelihood of being de- tected by predators. In addition to insects being attracted to these food sources, vertebrate predators may also be attracted to rotting fruit and moldy sap flows. Such conditions favor the evolution of morphological cryp- sis and associated behavior patterns that provide effective passive protec- tion of intoxicated individuals. Although the functional role of eyespots on the undersides of lepidopteran wings has been examined (e.g., Stradling 1976), I wish to advance a new hypothesis to account for the adaptive role of these markings. The small eyespots so familiar on many different species of satyrines, brassolines, and morphines, and the noticeable lack thereof in most nymphalines (including fruit and sap-feeding forms) suggests that they function to provide additional protection for butterflies feeding from the forest floor; butterflies generally feeding arboreally, including most nymphalines that feed on moldy sap flows, lack eyespots, suggesting that such markings are less adaptive to butterflies in these feeding niches. Owing to the increased likelihood that rotting fruits have already fallen from the parent tree by the time they are degraded to suitable food for butterflies, most incidents of intoxication prob- ably occur on the forest floor. Here, the eyespots (Eig. 1) provide additional protection to an individual butterfly that perhaps is partially intoxicated to a point of impairment of escape behavior. If such markings act as effective decoys for pecking by birds and flash attacks by other vertebrate predators, they provide additional time for an escape. VOLUME LXXXVIl, NUMBER 1 75 Owing to the close evolutionary history of the Satyrinae, Brassolinae, and Morphinae (Miller 1968), eyespots probably evolved in one of these groups and were then carried over to the others as they evolved. It is less clear as to how these markings, so strikingly similar among most members of these three groups, could have also evolved in the Nymphalinae, a very large group exhibiting manifold patterns of resource exploitation, both as larvae and adults. Most members of the Morphinae as well as all members of the Satyrinae and Brassolinae are monocot-feeders as larvae (Ehrlich and Raven 1965; Miller 1968). Virtually all Nymphalinae are dicot-feeders (Ehrlich and Raven 1965). Although it is difficult to determine whether or not the Bras- solinae and Satyrinae were derived phylogenetically from the Morphinae, or the reverse (e.g.. Young 1972b; Ehrlich and Raven 1965; Durden and Rose 1978), the three groups are closely linked in evolutionary history. The collections of fallen, rotting fruits available each year in the tropical wet forests are a major food resource for many butterflies, and the abun- dance and local diversity of these insects may be determined in part by these resources. The various sweet-smelling to fermenting odors given off by these fruits attract butterflies and other insects, and the activities of these animals in turn hasten the decay processes, making the fruit less attractive to larger animals (Janzen 1977). In terms of the insects, the fruits are a predictable resource in time and space and such predictability may be a major selective factor in the establishing of breeding populations of individ- ual species in a region of tropical forest. The repeated exploitation of a fruit ' crop in an area by the subset of the butterfly community that feeds on fruits reduces the likelihood that these fruits will be carried away from the area by larger animals acting as dispersal agents. As discussed primarily for I microbes associated with rotting fruit (Janzen 1977), the butterflies and other I insects are entering a competitive interaction with larger animals for this I food supply. Such interactions are less likely at arboreal sap flows, although i both resources are probably limiting factors for butterflies. Thus the likeli- j hood that opportunistic predators on large insects (reptiles, birds, and small ‘ mammals), will be active near or at fruits on the forest floor is greater than ; for arboreal sapkflows; such conditions select for behavior and morpholog- ' ical adaptations that increase the survival of the butterflies feeding on the fruits. Eyespot markings have high adaptive value under these conditions, and females are less abundant in such places since males are the more expend- I able. If mortality of feeding butterflies is high, selection favors more cryptic I behavior in females. Although the high levels of intoxication of butterflies at fruit baits and fruit products (e.g., Gomez 1977) are probably not fre- ! quently attained in nature, save for the exploitation of fallen fruit in banana ' plantations, there exists a spectrum of intoxication levels affecting escape behavior. Breakage of the wall of a fruit by rotting, falling, or handling by 76 NEW YORK ENTOMOLOGICAL SOCIETY animals allows fruit-foraging butterflies to use the fruit as a food source, but at the same time it exposes a portion of the population to intoxication which in turn lowers fitness. This interaction is mediated by the microbe com- munity associated with the fruit. As discussed primarily for the microbes (Janzen 1977), the joint activities of dispersal agents (vertebrates) affects fitness in the populations of the plant (tree) species involved. The ideas developed in this paper on the adaptive significance of butter- flies feeding on rotting fruits and fermenting sap flows in the tropics may also be applicable to the temperate zone fauna. For nymphalids of the tem- perate zone that feed on sap flows and only occasionally on rotting fruit, there is a lack of eyespot markings. Most of these feed arboreally and the density of vertebrate predators is very likely lower than in comparable trop- ical habitats. Selection pressures favoring eyespots are expected to be most intense in the tropics, and to be most prevalent among butterflies that feed on rotting fruits on the forest floor. Given the great diversity of butterflies in the tropics, the occurrence of eyespots in three large groups, such ad- aptations may have high selective value hitherto underemphasized. Literature Cited Barcant, M. 1970. Butterflies of Trinidad and Tobago. London: Collins, 314 p. Black, G. A., T. Dobzhansky, and C. Paven. 1950. Some attempts to estimate species diversity and population density of trees in Amazonian forests. Bot. Gaz. 1 1 1:413^25. Blest. A. D. 1957. The function of eyespot patterns in the Lepidoptera. Behaviour 11:209- 256. Brown. K. S.. Jr. 1972. Maximizing daily butterfly counts. J. Lepid. Soc. 26:183-196. Comstock. J. H. 1895. A Manual for the Study of Insects. Ithaca, New York: Comstock Publishing Co.. 701 pp. Dethier, V. G. 1947. Chemical Insect Attractants and Repellants. Philadelphia: Toronto, 289 P- Durden, C. J. and H. Rose. 1978. Butterflies from the Middle Eocene: The earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, Texas Mem. Mus., No. 29: 1-25. Ehrlich, P. R. 1958. The comparative morphology, phylogeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull. 39:305-370. Ehrlich, P. R. and P. H. Raven. 1965. Butterflies and plants: a study in coevolution. Evolu- tion 18:586-608. Frankie, G. W., H. G. Baker, and P. A. Opler. 1974. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. J. Ecol. 62:881-919. Frost. S. W. 1928. Continued studies on baits for oriental fruit moth. j. Econ. Entomol. 21:339-348. Gomez, L. D. 1977. The behavior of an inebriated Opsiphanes cassiae (Brassolidae). J. Lepid. Soc. 31:203-204. Mailman, J. P. 1977. Optical Signals. Animal Communication and Light. Indiana: Indiana University Press. 362 p. Harrison. J. O. 1963. On the biology of three banana pests in Costa Rica. Ann. Entomol. Soc. Amer. 56:87-94. Janzen. D. H, 1977. Why fruits rot, seeds mold, and meat spoils. Amer. Natur. 111:691-713. VOLUME LXXXVIl, NUMBER 1 77 Miller. L. D. 1968. The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Mem. Amer. Entomol. Soc., No. 24, 174 p. Muyshondt, A. 1973a. Notes on the life cycle and natural history of butterflies of El Salvador. IIA. Eiphile adrasta adrasta (Nymphalidae-Catonephelinae). J. New York Entomol. Soc. 81:214-223. . 1973b. Notes on the life cycle and natural history of butterflies of El Salvador. IVA. Pseudonica flavilla canthara (Nymphalidae-Catonephelinae). J. New York Entomol. Soc. 81:234-242. Norris, M. J. 1936. The feeding-habits of the adult Lepidoptera Heteroneura. Trans. Roy. Entomol. Soc. 85:61-90. Simon, D. and F. Enders. 1978. Insects visiting sap-exudate of groundsel tree (Baciharis neglecia). Southwest. Natur. 23:303-313. Smythe, N. 1970. Relationship between fruiting seasons and seed dispersal methods in a neotropical forest. Amer. Natur. 104:25-35. Stradling, D. J. 1976. The nature of the mimetic patterns of the brassolid genera, Caligo and Eryphanis. Ecol. Entomol. 1:135-138. Thompson, J. N. and M. F. Willson. 1978. Disturbance and the dispersal of fleshy fruits. Science 200: 1161-1163. Utrio, P. and K. Eriksson. 1977. Volatile fermentation products as attractants for Macrolep- idoptera. Ann. Zool. Fennici 14:98-104. Wickler, W. 1968. Mimicry in plants and animals. London: Weidenfeld & Nicholson (English translation by R. D. Martin), 253 p. Young, A. M. 1972a. Community ecology of some tropical rain forest butterflies. Amer. Midi. Natur. 87:146-157. . 1973. The comparative ethology and ecology of several species of Morpho butterflies in Costa Rica. Studies Neotrop. Fauna 8:17-50. . 1975. Feeding behavior of Morpho butterflies in a seasonal tropical environment. Rev. Biol. Trop. 23:101-123. Young. A. M. and A. Muyshondt. 1972. Ecological and geographical expansion in tropical butterflies of the genus Morpho in evolutionary time. Rev. Biol. Trop. 20:231-263. . 1973. The biology of Morpho peleides in Central America. Carib. J. Sci. 13:1-49. . 1975. Studies on the natural history of Central American butterflies in the family cluster Satyridae-Brassolidae-Morphidae (Lepidoptera: Nymphaloidea). III. Opsiphanes tamarindi and Opsiphanes cassina in Costa Rica and El Salvador. Studies Neotrop. Fauna 10:19-56. Young, A. M. and J. H. Thomason. 1974. Demography of a confined population of the but- terfly Morpho peleides during a tropical dry season. Studies Neotrop. Fauna 9:1-34. Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233. Received for publication July 31, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 78-84 NEST OF THE WASP CLYPEARIA WEYRAUCHI (HYMENOPTERA, VESPIDAE) Robert L. Jeanne Abstract. — The nest of the Neotropical social wasp Clypearia weyrauchi Richards is described for the first time. It consists of a single comb of sessile cells built directly on a tree trunk and covered by a domed envelope with an entrance below the center. The envelope does not contact the comb and is not thickened. It is unusual in having its outer surface coated with a layer of transparent film, probably a glandular secretion. This film serves to strengthen the crumbly, granular carton, which consists primarily of stone cells. The same film coats and strengthens the cell walls. Despite the fact that one of the two colonies examined appeared to be completing its cycle, its nest had not been enlarged beyond its initial size, a rare phenomenon among social wasps. There are two reasons why the study of social wasps’ nests is important. First, since the nest serves as a boundary between the external and internal environments of the colony, it must be considered in any study of the be- havioral ecology of a species. Second, since nest construction behavior evolves, and since many of the details of nest architecture are species spe- cific, nests provide valuable behavioral input into phylogenetic studies of the social Vespidae. Though the range of diversity of major architectural types is known (Jeanne, 1975), for studies of the behavioral ecology and phylogeny of the wasps we need to know such details as the type of cell construction, manner of thickening the envelope, material used, texture and strength of the carton, and the extent of the use of oral secretion in the nest. This kind of knowledge exists for the nests of only a small fraction of the approximately 700 species of social wasps. The Neotropical social wasp genus Clypearia comprises seven species (Richards, 1978); the nests of two, C. apicipennis and C. angiistior, are known (Ducke, 1910; Araujo, 1951), though Ducke’ s description of the nest of C. apicipennis was limited to one sentence. The purpose of the present paper is to describe the nest of a third species, C. weyrauchi Richards. All three species construct a single comb of cells directly on the surface of a tree trunk or limb, and cover it with a single-layered envelope, but the nest of C. weyrauchi differs in several interesting details from those of its congeners. The following description is based on two nests collected near Santarem, Para, Brazil (2°32'S, 54°20'W). Both were located in the same grove of trees VOLUME LXXXVII, NUMBER 1 79 Standing in a brushy pasture at the edge of the vorzea (annually inundated floodplain of the Amazon River). The larger of the two nests (no. 1112) was collected at night on July 26, 1975. It was situated at a height of about 7 m on the underside of a large horizontal branch. Though well sheltered from rain, the nest was not ob- scured by foliage. The nest was 45 cm long by 18 cm wide and contained approximately 1000 cells. This colony was evidently in the last stages of decline, for the nest contained only 12 adults and the brood cells were empty except for a few scattered pupae. The second nest, illustrated in Fig. 1, was collected from a nearby tree of the same species before dawn on August 14, 1977. It was also approximately 7 m high and very exposed, but the colony had chosen a nearly vertical section of tree trunk. The nest was 25.5 cm long by 9 cm wide and contained 287 cells. The adult population comprised 74 females, and the cells contained brood in all stages of development. Envelope The envelopes were constructed entirely by the edge construction tech- nique, the primary form of construction of all vespid nests so far described (Jeanne, 1973). At no point were the envelopes thickened by the addition of pulp to their surfaces. The thickness of the carton varied between 0.75 and 0.85 mm. The absence of surface thickening means that the pattern of construction of the envelope remains unobscured. This pattern is due to the lines of growth representing individual loads of carton pulp added by the wasps, and is visible in Fig. 1. The pattern for each nest indicates that the envelope was built at the outset to its final size, and was not enlarged from a smaller original envelope. In the smaller nest (Fig. 1) the lines of growth indicate that the envelope was raised uniformly around the entire comb to a height of 1 .0-1.5 cm, but that then carton was added to the two ends, and especially the upper end, faster than at the sides. The entrance, which results from the incomplete closure of the envelope, is consequently positioned below the middle of the nest. In the smaller nest the entrance was 1 cm in diameter. There was no thickening of the carton around the rim of the entrance, nor was there any flaring outward of the rim to form a tube or spout. The larger nest had two entrances. The pattern of growth lines around one of these was very irregular, suggesting that this part of the envelope had been damaged and repaired, leaving the second entrance. The envelope was constructed of a non-fibrous, finely granular material. Under the dissection microscope the individual particles could readily be broken up into finer granules 0.01-0.02 mm in diameter. These appear to be stone cells, most commonly found in the cortex of young stems, bark, and in fruit and seed coats (R. C. Koeppen, personal communication). Micro- 80 NEW YORK ENTOMOLOGICAL SOCIETY scopic examination also revealed minute amounts of an almost clear, plastic- like film intermingled with the particles and serving to bind them loosely together. This substance is probably a salivary secretion that is mixed with the particles as they are masticated before being applied to the nest. This matrix is very weak and the resulting carton very crumbly. The envelope is given added integrity by a thin surface coating of what is presumably more of the same salivary secretion. This material is evidently added after the envelope is completed, for it forms a discrete film on the outer surface of the carton, even extending for several mm onto the adjacent substrate. It is this thin pliable film that gives the envelope what little strength it has. Flexing the envelope readily cracked the carton, and with a minimum of abrasion of the uncoated inner surface the carton crumbled away, leaving only the outer film and a few scattered particles of carton adhering directly to it. At the time of collection of each nest there were several such “win- dows," some several cm across, that appeared to have originated by such loss of the underlying carton (Fig. 1). It is not known whether these windows were intentionally produced, or whether they simply resulted from wear and tear. The intact nests in situ were rather inconspicuous. This was partly be- cause the sloping sides of the envelope helped eliminate shadows, and partly because the nest closely matched its substrate in color. The close color resemblance of carton and bark, and the fact that the carton is composed largely of stone cells, suggest that the source of carton material might have been the bark of the tree on which each colony nested. The translucent coating of secretion over the envelope further subdued any contrast between the nest and the tree trunk. Comb and Brood The comb of each nest was centered within the space enclosed by the envelope. A space of 1.3— 4.5 cm separated the edge of the comb from the line of attachment of the envelope to the substrate. Thus the envelope is not initiated by the elongation of the walls of peripheral cells, but is built up from the substrate completely independently of the comb (Fig. 2). The combs of both nests appeared to have been built in a single effort; there were no irregularities in outline, hexagonal pattern, cell height, or brood distribution to indicate that a smaller original comb had been en- larged. All cells in both nests were sessile; that is, they were built directly on the surface of the tree trunk, which formed the bottoms of the cells (Jeanne, 1973). Cells were 5. 2-5. 4 mm in diameter and reached a depth of 20 mm. Those of the nest on the vertical trunk were angled downward 15° with respect to the trunk surface (Fig. 2). VOLUME LXXXVII, NUMBER I 81 Fig. I. Intact nest of Clypearia weyniuchi (no. 2035). The nest is 25.5 cm long. The stria- tions indicate the pattern of construction of the envelope, and converge on the entrance. The dark blotches above the envelope are 'windows’ formed by the removal of carton, leaving only the translucent film coating the outer surface of the envelope. 82 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 2. The nest shown in Fig. 1 with its envelope removed. The line of attachment of the envelope is visible on the bark surface. The comb is centered within the surface enclosed by the envelope, and does not contact the envelope. VOLUME LXXXVII, NUMBER 1 83 The cells were constructed of the same crumbly carton as the envelope, and were similarly coated with a film of secretion. Thus the walls of the cells were layers of carton sandwiched between two layers of secretion, the latter providing the main strength for the cells. The coating lined the bottoms and walls of all cells, even shallow peripheral cells that had not produced adults, though in some cases the upper few mm near the rim of these cells lacked it, suggesting that these had recently been heightened by adding carton, but had not yet been coated. Eggs were attached to the wall of the cell just above the bottom, and always in the uppermost angle of the cell, so that they hung downward nearly parallel to the flat cell bottoms. This position indicates that the queens faced upward on the comb while ovipositing. Larvae about to pupate spin a silken cocoon completely lining the cell, including the bottom. This lining was distinguishable from the film of secre- tion lining the cells by virtue of its fibrous texture, greater strength, whiter color, and by its presence only in cells that contained meconia. The silk lining was relatively tough, and with care the entire cocoon, with the me- conium at the bottom, could be removed intact from a cell. ^ The caps of the cocoons were set 2-6 mm below the rims of the cells, ! were slightly domed, and were translucent at the edges, grading to opaque white in the center. After the cocoon is completed the adults add scattered streaks and blotches of carton to the cap. , After the adult ecloses from a cell, the only treatment the cell receives is I to have the cap trimmed away. The walls of the cell are not lowered, nor j are the meconia removed, before the cell receives another egg. 1 No droplets of stored nectar were found in any of the cells of either nest. ! Discussion I Though much detail is lacking from Araujo’s description of the nest of C. ' angustior (Araujo, 1951), and Ducke’s (1910) description of the nest of C. apicipennis is even less useful, a few points of comparison with that of C. I weyrauchi are possible and seem worth making. The most striking differences pertain to the envelope. While in C. wey- rauchi there is a wide space separating comb and envelope, Araujo (1951) makes it clear that C. angustior starts the envelope by elevating the outer I walls of the peripheral cells of the comb, so that the forms of the cells are ' visible on the sides of the covered nest. The two species also differ with respect to the nest entrance, that of C. angustior protruding somewhat, while that of C. weyrauchi is flat. Though Araujo (1951) states that the envelope of his nest of C. angustior contains numerous 'windows’ of transparent secretion, he is not clear on their size or nature. He states that they are "perfectly reminiscent” of those 84 NEW YORK ENTOMOLOGICAL SOCIETY in the nest of Metapolybia cingulata. In that species the 'windows’ are tiny and are incorporated as the envelope is being built. There is no film of secretion added later to the surface of the envelope. In C. weyrauchi, on the other hand, the windows are much larger and are made by the removal of carton from the inside of the film after the envelope is complete. It is not possible to be certain from Araujo’s description whether the 'windows’ and envelope of C. angustior are more like Metapolybia cingulata or C. wey- rauchi, and he does not mention the existence of a surface film of secretion. Finally, Ducke (1910) states that the nest of C. apicipennis seen by him had been enlarged, as indicated by irregularities in the outline of the enve- lope. On the other hand, neither nest of C. weyrauchi showed any evidence of having been enlarged, even though the larger one had evidently housed its colony for nearly a full normal developmental cycle. If C. weyrauchi, indeed, does not enlarge its nests once they are built, it is one of few species of New World polybiines not to do so. Acknowledgments I thank Roger Swain for assistance in the field. R. C. Koeppen kindly examined the carton material. I am grateful to the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico for granting me permission to work in Brazil. Research supported by the College of Agricultural and Life Sciences, University of Wisconsin, Madison, and by National Science Foundation grants GB 33619 and BNS 77-04081. Literature Cited Araujo, R. L. 1951. Contribui^ao para o conhecimento de Clypearia angustior Ducke. 1906 (Hym. Vespidae). Arq. Mus. nac. Rio de Janeiro 42(l):49-56. Ducke, A. 1910. Revision des guepes sociales polygames d’Amerique. Ann. Mus. nac. Hun- garici 8:449-544. Jeanne. R. L. 1973. Aspects of the biology of Stelopolybia areata (Say) (Hymenoptera: Ves- pidae). Biotropica 5(3): 183-198. . 1975. The adaptiveness of social wasp nest architecture. Quart. Rev. Biol. 50:267- 287. Richards, O. W. 1978. The social wasps of the Americas excluding the Vespinae. British Museum (Natural History). London vi -i- 581 pp. Department of Entomology, University of Wisconsin, Madison, WI 53706. Received for publication August 8, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 85-90 EFFECTIVENESS OF NATIVE PARASITES AGAINST AGROMYZA FRONTELLA (RONDANI) (DIPTERA: AGROMYZIDAE), I AN INTRODUCED PEST OF ALFALFA R. M. Hendrickson, Jr. and S. E. Barth Abstract. — Fourteen species of native parasites of the alfalfa blotch leaf- miner, Agromyza frontella (Rondani), an introduced European pest, were I recovered in the northeastern USA. Two additional species were recovered I by cooperators in Canada. This complex of native parasites is poorly syn- I chronized with the first 2 generations of the host, which may therefore reach I economic injury levels; however, it gives satisfactory control of the next 3 I generations. Season-long parasitism averaged 36%, but only 2% of the host puparia were parasitized. European host puparia collected in 1976-77 and shipped to the USDA quarantine facility at Newark produced 7 parasite species with a combined parasitism of 25.5%. All 7 species have been in- troduced in the USA. The alfalfa blotch leafminer (ABL), Agromyza frontella (Rondani), is a European species first reported in North America in Massachusetts in 1968 (Miller and Jensen, 1970). It has since spread throughout the northeastern states (Hendrickson and Barth, 1978a) and into the adjoining Canadian prov- inces (Bereza, 1977). In North America, ABL is frequently an economic I pest, though populations in Europe are usually held at low levels, with rare, i localized outbreaks (Bollow, 1955). We therefore surveyed the native North American parasites of ABL to 1 determine their effectiveness and their phenology in relation to the host. I Such information was useful in selecting parasite species from a large Eu- i ropean complex, studied by personnel at the USDA European Parasite Lab- oratory, Paris, France, for possible introduction into the USA. The order, family, and identifier of species mentioned in this paper are presented in Table 1. Sampling techniques. — Samples of 50 mined alfalfa leaflets, that is, leaf- lets visibly mined by any of the 3 agromyzid species present on alfalfa in the survey area (Agromyza frontella, Liriomyza trifoliearum, and Liriomyza trifolii) were collected randomly each week without regard for host species, instar, or condition of host from early May through November 1975 from each of 7 fields ( 1 at Newark, Del., 3 near Oxford, Pa., and 3 near Rancocas, N.J.). Collections were not made if the alfalfa was less than 12.7 cm (5 in.) high or bad weather interfered. The same procedure was followed in 1976 and 1977 except that in these years an additional field was surveyed in the 86 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Order, family, and identifier of species mentioned in this paper. Species HYMENOPTERA: BRACONIDAE Dacnusa dry as (Nixon)“ Dapsilarthra halteata (Thomson)'' Opius dureseaui Fischer® HYMENOPTERA: EULOPHIDAE Achrysocharella Formosa (Westwood)' Chrysocharis clarkae Yoshimoto'' Chrysocharis giraulti Yoshimoto® Chrysocharis punctifacies Delucchi" Closterocerus cinctipennis Ashmead’’ Closterocerus tricinctus (Ashmead)*" Closterocerus utahensis CrawTord'" Diaulinopsis callichroma Crawford* Diglyphus begini (Ashmead)*" Diglyphus intermedius (Girault)*" Diglyphus pulchripes (Crawford)*" Diglyphus websteri (Crawford)' Pnigalio minio (Walker)" Zagrammosoma multilineatum (Ashmead)*" HYMENOPTERA: PTEROMALIDAE Cyrtogaster sp." Halticoptera circulus (Walker)" Halticoptera laevigata Thomson" Miscogaster hortensis Walker" Miscogaster maculata Walker" DIPTERA: AGROMYZIDAE Agromyza front ella (Rondani)" Liriomyza trifoliearum Spencer" Liriomyza trifolii (Burgess)" Identified by: ® P. M. Marsh, *" G. Gordh, " G. C. Steyskal, and " E. E. Grissell, Systematic Entomology Laboratory, AR/SEA, USDA, do U.S. National Museum, Washington, DC 20560. '* M. Fischer, Naturhistorisches Museum, 2. Zoologische Abteilung, Vienna, Austria. " C. M. Yoshimoto, Biosystematics Research Institute, Agriculture Canada, Ottawa, On- tario KIA 0C6. ' R. M. Hendrickson, Jr. Newark, Del., area. Also, collections were made weekly, from mid- April through July, and thereafter at 2-week intervals. However, if the abundance of leaflets containing mining larvae or Liriomyza trifoliearum puparia (the other 2 species pupate in the soil rather than in the leaflet) was low, the sample consisted of the number that could be collected in each field in 20 minutes. In the collections over the 3 years, ABL larvae were more abun- VOLUME LXXXVIl, NUMBER 1 87 Fig. 1 . Ratio of unparasitized to parasitized alfalfa blotch leafminer larvae related to number of occupied mines per stem for 1977. Larvae were collected on alfalfa at 2 fields in Newark, Del., 3 fields near Rancocas, N.J., and 3 fields near Oxford, Pa. dant than larvae or puparia of Liriomyza spp. by ca, 9:1. In addition, we removed 20 stems from each field to determine the number of occupied and ; unoccupied mines per stem in 1977. Some host characteristics. — There were 5 generations per year of ABL j in the study area. Alfalfa is commonly cut 3 times each season in our area, the last time early in September, and a generation of ABL preceded each I cutting. Then the alfalfa continues to grow until the end of November, so there is a period of nearly 3 months during which a 4th and possibly a 5th generation of the ABL can be produced. That these generations do occur was indicated by the presence of some active larvae and adult flies in the field up to the end of November. Pupating larvae of the 5th generation then emerged as adults in late April or early May of the following year. The major peak in population occurred at the time of the 2nd generation when 70+ mines per stem were sometimes encountered (Fig. 1). Native parasite complex. — A total of 13,551 leaflets containing ABL lar- vae was placed in 5.08 cm (2 in.) diameter, tightly-sealed plastic petri dishes (one leaf per dish) with moistened filter paper on the bottom to maintain humidity. (So only a single host larva could be confined in each petri dish, any supernumerary insects were removed from the leaflet.) The petri dishes 88 NEW YORK ENTOMOLOGICAL SOCIETY Table 2. Parasite species and numbers reared from 13,351 alfalfa blotch leafminers col- lected in Delaware, southern New Jersey, and southeastern Pennsylvania in 1975-77. Recovered from Species Larval stage Pupal stage Total no. Diglyphus intermedins X 1323 Diglyphus pulchripes X 344 Pnigalia minio X 152 Diaulinopsis c alliehroma X 123 Closterocerus tricinctus X 78 Chrxsocharis clarkae X X 65 Closterocerus cinctipennis X 58 Achrysocharella formosa X 43 Diglyphus websteri X 29 Chrysocharis giraulti X X 15 Zagrammosoma multilineatum X 5 Closterocerus utahensis X 2 Hahicoptera laevigata X 2 Halticoptera circulus Unidentifiable® Total X 1 69 2309 ^ Adults were damaged or lost, or larvae were in diapause. were then maintained at 20°C ±1.1° (68°F ± 2°) with 14/24 h photophase. After 6 weeks, emerged parasite species were removed and identified. Initial recoveries were identified by authorities (see Table 1); subsequent recov- eries were determined by comparison with identified specimens. Species recovered and numbers are listed in Table 2. Two species of parasites re- covered from ABL from eastern Ottawa, Ontario, but not recovered by us in the USA, were the larval parasite Diglyphus begini and the larval-pupal parasite Cyrtogaster sp. (J. C. Guppy, pers. com.). Diglyphus begini was also recovered from ABL at St. Hyacinthe, Quebec (M. Guibord, pers. com.). There was no emergence of either parasites or ABL from 52% of the mined leaflets. This was probably the result of many factors: early instars killed by probing (without oviposition) by the most abundant parasite, Di- glyphus intermedins (Hendrickson and Barth, 1978b); overcrowding in the leaflet; host diapause; disease; predators; weather and other physical fac- tors; insecticides; and possible deficiencies in our emergence technique. At the peak of host larval populations in the 1st generation, an average of 11% of the larvae were parasitized (total live forms). The average was 16% in the 2nd generation. Parasites appeared 2-3 weeks too late in the spring to be effective against the 1st generation of ABL, and too few were present to control the 2nd host generation, when 70+ mines per stem were VOLUME LXXXVII, NUMBER 1 89 I * Table 3. Parasite species and numbers reared from 33,573 alfalfa blotch leafminer puparia I collected in Europe in 1976-1977 by personnel from the USDA European Parasite Laboratory, Paris, France. Parasites emerged in the quarantine facility at the USDA Beneficial Insects ' Research Laboratory, Newark, Delaware. Species No. % of total parasites Chrysocharis punctifacies 1656 42 Dacnusa dryas 1614 41 Miscogaster maculata, M. hortensis 571 15 Opiiis dureseaui 47 1 Halticoptera circidus 26 <1 Dapsilarthni hcdteata 6 <1 Total 3920 sometimes encountered. During the remaining 3 generations, the average parasitism was 55%, and the host populations remained well below the eco- nomic threshold. (A tentative economic threshold of 21% of the leaflets I mined by a single larva has been established by R. A. Byers, pers. com.). I Combined season-long parasitism was 36% (based on total emerged live ' forms; numbers of live forms varied for each host generation). Of the 14 native species recovered from ABL, only 4 species emerged 1 from ABL puparia (Table 2). These species emerged from 1.9% of the 4284 ; collected hosts which formed puparia. Origin of the native parasite complex. — A likely explanation for the ABL parasites is adaptation from native agromyzids found on alfalfa. Therefore, petri dishes were set up as before, for 1566 leaflets containing larvae or puparia of Liriomyza trifoliearum or larvae of Liriomyza trifolii (the latter species pupates in the soil rather than the leaflet). All but 2 of the 14 parasite species recovered from ABL were also recovered from Liriomyza spp. The species not recovered were Zagrammosoma multilineatum and Clostero- cerus utahensis, which together accounted for only 0.3% of the parasitism of ABL. Possibly with more extensive surveys, these 2 species would also be recovered from Liriomyza spp. We found that Liriomyza spp. on alfalfa are under excellent biological control. Season-long parasitism averaged ca. 63%. The complex of native parasites of ABL is therefore probably phenologically associated with that of Liriomyza spp. European survey. — The USDA European Parasite Laboratory, Paris, France, provided us with 33,573 ABL puparia collected from France, Den- mark, West Germany, Austria, Switzerland, and Liechtenstein in 1976-77. Parasites or ABL adults emerged from ca. Vi the puparia. Seven parasite species (Table 3) emerged from these collections, all of which have been 90 NEW YORK ENTOMOLOGICAL SOCIETY released in the USA with additional numbers shipped to us as adults. Par- asitism of the pupal stage averaged 25.5%. We therefore believe the introduction of parasite species capable of in- creasing parasitism of the pupal stage by 20-25% would reduce ABL to the extremely low population levels found in Europe. The most desirable species for establishment in the USA are Chrysocharis punctifacies and Dacnusa chyas, which together accounted for 83% of the parasites emerging from European puparia. These species have been established in Delaware (Hendrickson. 1978). Literature Cited Bereza. K. 1977. Alfalfa blotch leafminer. Ontario (Canada) Factsheet No. 77-067. 2 pp. Bollow, H. 1955. Ein Massenauftreten der Minierfiiege Agromyza frontanella ROND, an Luzerne. Pflanzenschutz 7:141-143. Hendrickson, R. M. Jr. 1978. Establishment of Dacnusa dryas (Nixon) (Hymenoptera: Bra- conidae) and Chrysocharis punctifacies Delucchi (Hymenoptera: Eulophidae), parasites of Agromyza frontella (Rondani) (Diptera: Agromyzidae) in Delaware. (Abstract) J. N.Y. Entomol. Soc. 86:295. Hendrickson, R. M. Jr. and S. E. Barth. 1978a. Biology of the alfalfa blotch leafminer. Ann. Entomol. Soc. Am. 71:295-298. . 1978b. Notes on the biology of Diglyphus intermedins (Hymenoptera: Eulophidae), a parasite of the alfalfa blotch leafminer, Agromyza frontella (Diptera: Agromyzidae). Proc. Entomol. Soc. Wash. 80:210-215. Miller, D. E. and G. L. Jensen. 1970. Agromyzid alfalfa leafminers and their parasites in Massachusetts. J. Econ. Entomol. 63:1337-1338. Beneficial Insects Research Laboratory, USDA-SEA-AR, Newark, Del- aware 19713. Received for publication 20 October 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 91-94 POSSIBLE PHYLOGENETIC SIGNIFICANCE OF COMPLEX HAIRS IN BEES AND ANTS U. N. Lanham Abstract. — If both larval and adult stages are taken into account, then ants and bees are the only major groups of aculeate Hymenoptera in which complex hairs are abundant. The possible phylogenetic significance of this is discussed. ' The first serious attempt at constructing a phylogeny of the higher acu- leate Hymenoptera was that of Borner (1919). In it he grouped the bees, ' scolioid wasps, and ants in a category (the Haplocnemata) which is con- trasted with the vespid, pompilid, and sphecid wasps (Diplocnemata). This scheme has been discussed and rejected by Bischoff (p. 546) and Malyshev (p. 255). The modern conception is that the Apoidea are little modified descendants of the Sphecidae, so closely related that both can be included I in a single superfamily (Bohart & Menke, p. 31). Borner, however, has made a considerable contribution to the array of characters available to discriminate higher categories, among them the strigil of the hind leg which, I while it establishes his major categories, also provides a useful means of distinguishing apoids and sphecids (Lanham 1960, and Bohart & Menke 1976:27.) The main difficulty with the concept that bees are essentially sphecid I wasps that have converted from a predatory to a pollen-gathering mode of : existence is that intermediate steps have not been found either in morphol- ogy or behavior, nor do behavioral patterns of our presently living forms provide material for demonstrating transition stages. If, on the other hand one takes, as a heuristic device, the theory that bees are closely related to ants, then a multitude of possibilities are opened up. Also, one is led into new ways of looking into problems of morphology or behavior. It is not the purpose of this paper to outline a route or routes by which a bee-ant tran- sition could be made, but rather to look at a morphological character whose distribution among the major higher groups of Hymenoptera has been ne- glected, namely the compound hairs which are generally thought to be char- acteristic only of the Apoidea (Saunders 1880:202, and most modern text- books). All adult bees that I have examined carefully possess at least a few com- pound (branched, or denticulate) hairs. Bees have in addition other types of complex hairs, such as the spirally twisted hairs in the pollen-transporting brush of the megachilids, or the spatulate hairs on the abdominal terga of some anthophorids, for example Svastra. 92 NEW YORK ENTOMOLOGICAL SOCIETY Rayment (1935:251) notes that compound hairs are present in mutillids, and Brothers { 1975:495) assigns the additional “scolioid” families Eotillidae, Typhoctidae and Anthoboscidae to the list of aculeates with such hairs. Brothers, on the basis of his sample of the ants, assigned to the formicids the character state “simple hairs.” It was news to me to find that it has long been known to myrmecologists that adult ants possess complex hair types, including compound hairs. Creighton’s key (1950) to North American ants mentions Acanthomyops ptumipilosis and a species of Triglyphothrix as having branched hairs. In a key to the genera of ants of the world Wheeler (1922) mentions a variety of hair types in adult ants: clavate, spatulate, hooked, and denticulate (in mellitology, the last would be called obscurely compound). Ants with these hairs appear to represent scattered species or genera. If one takes a larger view of the ants, taking into account the larvae, then the ants are not far behind the bees in having bizarre hair structure. About 90% of the genera of ants have quite hairy larvae (nearly all other aculeates have larvae with at best a few bristles). G. C. and J. Wheeler (1976) classify the hairs in nearly 30 categories, using such terms as denticulate, uncinate, anchor-shaped, branched, bifid, or clawlike. In this array, 20 types are com- pound, in the terminology of bee taxonomy. Data are not presented in such a way as to make a good estimate, but perhaps half the genera of ants, including at least some species of the primitive Australian Mynnecki, have larvae with compound hairs. The Wheelers regard these elaborate larval hairs as adaptations for life in a communal chamber, with the hairs holding them above the substrate, providing insulation and preventing desiccation by producing a dead air space, keeping the brood together by clumping, or helping the larvae cling to vertical surfaces. Probably the only other aculeate Hymenoptera with communal brood chambers (larvae not in individual cells) are a few genera of bees. The most interesting of these on theoretical grounds are the allo- dapine bees, which nest in hollow twigs, have hairy larvae, and in at least , one species, have compound (denticulate) hairs (Michener 1977). I am aware of no speculations concerning the function of complex hair types in adult ants. If one assumes that the presence of compound (or in broader terms, highly modified) hairs in both bees and ants indicates a close phylogenetic rela- tionship, he is of course faced with the difficulty that in ants it is mainly the larvae that have such hairs, while in bees they are found almost exclusively in the adults. One also has to make a judgment concerning the numerous, and so far as known at present, haphazard occurrence of such hairs in adult ants. Probably the key question is this: are compound hairs easy to originate I VOLUME LXXXVII, NUMBER 1 93 ' de novo in the evolutionary process, or does the establishment of this char- . acter require special conditions that are rarely met with. ; A significant fact bearing on the question of whether complex hairs are j easy to come by is that, so far as I can determine, the hairs are invaribly I simple among the pompilid, vespid, sphecid, scoliid, and tiphiid wasps, j which total approximately 15,000 species. Among the thousands of modes of life to be found in these wasps, one would expect that some species would encounter situations in which there would be adaptive value for com- ' pound hairs. Those of bees are of use in trapping pollen grains, although on many parts of the body they may also contribute to the density and resilience ! of the furry coat, affecting such properties as insulation or resistance to abrasion. Some of the vespid wasps gather pollen for their young, but none of these have compound hairs. It will be assumed here, for purposes of discussion, that compound hairs are difficult to achieve in the evolutionary process, requiring the assembly ' of a large number of genes into a stable and long-lasting complex. Such complexes then come under the control of a single suppressor or switch gene. Variability of this gene is subjected to selection pressure, establishing alleles which determine whether or not the complex expresses itself, or modulating its expression with respect to body region or stage in life cycle. I Thus, the genetic aspects of our problem come down to this, that the gene : complex for a given character, such as the morphology of a single-celled ' hair, may be multigenic, and may require a long period of time as well as long-lasting and unusually favorable environmental conditions for the as- i sembly of this gene complex. But the addition of a suppressor (switch or j control) gene can obscure this situation, making the character act in many t ways as if it were based on a single gene. Without direct genetic experi- mentation, one can only make a judgment on the basis of the distribution of the character within the taxonomic framework. T. D. A. Cockerell early in his career published a few notes on bee phylogeny, but never pursued the matter in print. That he was long interested in basic aspects of the problem is indicated in a paper by himself and Louise Ireland (1933): "... there is increasing proof that the genes for many characters lie latent for long periods, so that similar structures appear in different branches of the same family or order. It may of course be debated whether these reap- pearances are due to entirely new developments or have a common origin somewhere else in the ancestral germ plasm, or whether they are due to a sort of orthogenesis. It is not necessary to assume that every case may be developed in the same way, but among bees, at any rate, the evidence for germinal continuity and latency seems rather convincing.” Applying the concept of latency to the present problem, one could spec- ulate that the sphecids and other groups without compound hairs have an 94 NEW YORK ENTOMOLOGICAL SOCIETY 1 evolutionary background without gene complexes for such structures, and 1 that favorable circumstances for their establishment never arose. The ants < and bees both have a background that did include such complexes for pro- ducing varied hair types. If one assumes that the modulating capabilities of the suppressor gene make it possible to transfer the expression from larval to adult stage, or adult to larval, there is considerable room for speculative maneuver. Adult ants would be widely infiltrated with suppressed genetic complexes for varied hair types, which surface only here and there, while expression is characteristic of most larvae. In bees the situation is the other way around, with apparently all adults expressing the character, even if only in very rudimentary form. The allodapine bees, which have hairy lar- vae, are probably still too poorly known to make any judgement as to the extent to which complex hair types are present. Literature Cited Bischoff, H. 1927. Biologic der Hymenopteren. Berlin, Julius Springer. 598 pp. Bohart. R. M. & A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. Berkeley, University of California Press. 695 pp. Borner. C. 1919. Stammegeschichte der Hautfieugler. Biol. Zentralblatt 29:5-186. Brothers. D. J. 1975. Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae. U. Kansas Sci. Bull. 50:383-648. Cockerell. T. D. A. & L. Ireland. 1933. The relationships of Scrapter. a genus of African bees. Proc. Nat. Acad. Sci. 19:972-978. Creighton. W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585. Lanham, U. N. 1960. A neglected diagnostic character of the Apoidea. Ent. News 71:85-86. Malyshev. S. I. 1968. Genesis of the Hymenoptera and the phases of their evolution. London, Methuen. 319 pp. Michener, C. D. 1977. Discordant evolution and the classification of allodapine bees. System- atic Zoology 26:32-56. Rayment. Tarlton. 1935. A cluster of bees. Endeavour Press, Sydney. 752 pp. I Wheeler, G. C. & J. Wheeler. 1976. Ant larvae: review and synthesis. Memoir Ent. Soc. i Washington, no. 7. 108 pp. Wheeler, W. M. 1922. Keys to genera and subgenera of ants. Bull. Amer. Mus. Nat. Hist. t 45:631-710. University of Colorado Museum, Boulder, Colorado 80309. Received for publication 23 October 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(l), 1979, pp. 95-96 BOOK REVIEW George Ordish. The Year of the Ant. 139 p. 1978. Charles Scribner’s Sons. New York. $9.95. This is a delightful description of the ants and their social system, ex- plaining their role in the ecosystem and their ability to survive for millions of years. The explanation of the system, that was able to overcome potential disasters, such as fire, floods, DDT and dependance on slavery and drug addiction, is masterfully presented and of interest not only to the expert myrmecologist, but also to the layman. The author writes with an appealing sense of humor that pervades this volume and provides a real treat. Ants, as well as other social insects, have fascinated man since biblical time. Ants probably originated in North America wherefrom they spread to South America and to Eurasia over the land bridges that once connected the con- tinents. The ant colony is often described as a successful Marxist system, in which each ant gives according to its ability and receives according to its need. However, ants in a colony are not merely part of an automatic ma- chine. There is some free will, as ants enjoy some leisure, such as occasional sunning. Individual ants are capable of learning, and they make choices. The very limited scale of such activities can easily be overridden by the need of the colony and control is exercised by chemical secretions. The author describes this with scientific accuracy by presenting the life cycle of an individual wood ant of a colony. The story begins in March and the star performer is from a monogynous nest, situated in the woods in a clearing in the Adirondack Mountains. Nearby is another, older nest that is poly- gynous. The behavior of ants in both nests, their contact with aphids, care of eggs, dangerous mites and even more dangerous staphylinid beetle in- vasions that result in drug addiction, the emergence of males and mating, robber ants, loss of the queen and transformation of the worker into a queen, and finally the fight with another queen and the decapitation of the “hero- ine” are chronologically presented, from March till October of “the year of the ant.” The chapters are aptly illustrated by Clarke Hutton. There is a bibliography of books and original articles and a subject index. This book can be read profitably not only as an introduction to social insects, but also by the specialist. The book can be recommended to stu- dents, teachers, and research scientists, as well as for general academic libraries, public libraries and high schools. The good quality of the print and paper in a book costing less than $10.00 might also be mentioned. In short, the author succeeded in providing a book not only for entomologists but 1 96 NEW YORK ENTOMOLOGICAL SOCIETY also for a more general audience, that merits reading by all interested in biology. Several copies should be made available in the libraries of high schools and academic institutions. Karl Maramorosch Waksman Institute of Microbiology Rutgers University P. O. Box 759 Piscataway, New Jersey 08854 BOOK REVIEW Tahanini of Thailand above the Isthmus of Kra (Diptera: Tahanidae). John J. S. Burton. 165 pp. 1978. Entomological Reprint Specialists, Los An- geles, California. $15.00. Eemale Tabanini are well known blood suckers, whereas the males mostly feed upon honeydew and on juices of flowers. These flies play an important role in spreading agents of certain infectious diseases such as Surra to live- stock— especially virulent in equine species. Tabanini also play a role in the spread of tularemia to man. The biological diversity and economic impor- tance of gadflies has been the subject of many scientific papers. Although the oriental tabanid fauna is unusually rich, it remained the least known up to now. Therefore this excellent study is very important and it is useful not only insofar as the tabanid fauna of Thailand and southeast Asia is con- , cerned, but also for the whole oriental zoogeographic region. In twelve chapters the author presents detailed information about the economic importance of gadflies, collections made in Indochina, superspe- cific taxa of Tabanini with reference to Thailand, the species concept, field procedures, descriptions of species, and a key to females. The interesting ' and detailed study of the tribe Tabanini comprises 81 species, of which 31 are described as new and 24 nominal taxa are newly synonymized. For many species lectotypes are given and some new generic combinations are proposed. The study is illustrated with 80 excellent figures. The last chapter deals with other species involved or implicated in the transmission of disease agents in the Indochina area. D. Simova-Tosic Faculty of Agriculture Department of Entomology Zemun, Yugoslavia Journal of the New York Entomological Society VOLUME LXXXVIl JUNE 1979 NO. 2 CONTENTS A brief review of the genus Cicindela of Argentina (Coleoptera: Cicindelidae) William D. Sumlin, III 98-117 Reproductive behavior of Euresta bella and E. festiva (Diptera: Tephritidae), poten- tial agents for the biological control of adventive North American ragweeds (Ambrosia spp.) in Eurasia S. W. T. Batra 118-125 Acoustic attraction of herons by crickets Paul D. Bell 126-127 Community ecology and Pieris — Crucifer coevolution F. S. Chew 128-134 Interactions between blow flies (Calliphoridae) and Entomophthora bullata (Phy- comycetes: Entomophthorales) John Paul Kramer 135-140 Post-ingestive utilization of plant biomass and nitrogen by Lepidoptera: legume feed- ing by the southern armyworm J. Mark Scriber 141-153 Observations on aggregation and overwintering in the coccinellid beetle Coleo- megilla maculata (DeGeer) Allen H. Benton and Andrew J. Crump 154-159 On the identity of two species of Rhyparochrominae from Argentina (Hemiptera: Lygaeidae) James A. Slater 160-166 Introduced parasites of Agromyza frontella (Rondani) in the USA R. M. Hendrickson, Jr. and S. E. Barth 167-174 Photoperiod and temperature influences on egg number in Brachymeria inter- media (Hymenoptera: Chalcididae), a pupal parasitoid of Lymantria dispar (Lepidoptera: Lymantriidae) P. Barbosa and E. A. Frongillo, Jr. 175-180 Book Reviews 181-184 NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK2), 1979, pp. 98-117 A BRIEF REVIEW OF THE GENUS CICINDELA OF ARGENTINA (COLEOPTERA: CICINDELIDAE) William D. Sumlin, III Abstract. — The genus Cicindela L. (sensu W. Horn) of Argentina is brief- ly reviewed. Thirty-four taxa are discussed and known ranges are given. A key to the species is presented. Four new species and one new subspecies are described and figured. Three of the new species (C. halophila, C. sic- calacicola and C. hirsutifrons) share the same type locality: 24 mi. S. Re- creo, Cordoba Province. The fourth new species (C. stamatovi) is described from Tucuman. C. drakei latifascia n. ssp. is described from Termas de Rio Hondo, Santiago del Estero Province. Introduction The genus Cicindela of Argentina has not been adequately reviewed. It has been presented in the form of checklists by Horn (1905), Bruch (1910), , Horn (1915, 1926), Blackwelder (1946) and Rivalier (1954). j The latter author incorporated a study of male genitalia into his work and split the genus into several “genera.” Following Rivalier’s work, Vidal Sar- miento (1966) gave a detailed account of the male genitalia for many of the species. Although that work was detailed, with respect to the males of the species reviewed, it failed to incorporate a key or deal with females. The present work treats 34 taxa with five being described as new. I have endeavored to treat all specific and subspecific taxa contained in Horn’s (1926) definition of the genus Cicindela of Argentina with the exception of C. cribrata Brulle. I feel that Rivalier’s (1969) placement of cribrata into Pentacomia Bates is correct. I do not, however, subscribe to his splitting j up the New World Cicindela in 1954. Rivalier’s “genera” are herein treated i as subgenera. Many of the species treated in the present work are quite rare in collec- tions and as a result some were only represented by one or two specimens. . In some cases Argentine specimens were not available for study and spec- imens from other countries had to be utilized. Two species (C. argentinica Mandl and C. obsoletesignata W. Horn) were unavailable for study which necessitated culling key data from their original descriptions. All characters used in the key below are easily seen and interpreted. To facilitate usage, with respect to the terminology of the elytral maculation, users are directed to Figure 1 which deals with lunule nomenclature. VOLUME LXXXVII, NUMBER 2 99 Key to Species of Cicindela of Argentina - Elytra without distinct subsutural rows of large metallic foveae 2 3 4 5 6 7 8 9 10. 11. 12. 13. 14. 15. 16. 17. Elytra immaculate Elytra with some degree of maculation Dorsal color dark reddish brown Dorsal color black Labrum 3 toothed Labrum 4 or 5 toothed Genae glabrous Genae setose Maculation very wide, covering much of elytral surface Maculation thin or consisting of small spots Basal half of elytra sparsely covered with white setae Basal half of elytra without white setae Elytral apex immaculate Elytral apex with maculation Dorsal color black or dark brown Dorsal color green tinged cupreous red Elytra with distinct subsutural rows of large metallic foveae 2 4 rufoaenea 3 obsoletesignata morio 5 13 6 7 eugeni apiata 8 9 obscurella aureola 10 11 Size smaller (8 mm in length) Size larger (12 mm in length) Apex of elytra rounded (see Eig. 2a) Apex of elytra angular (see Eig. 2b) Humeral lunule broken — consisting of two small spots Humeral lunule complete and unbroken chlorosticta staudingeria argentata^ 12 misella^ stamatovi n. sp. 14 Pigment (ground color) contacting lateral edges of elytra Pigment not contacting lateral edges of elytra (i.e., maculated from humeral area to apical suture) 24 Apex of elytra without microserrations 15 Apex of elytra with microserrations 16 Labrum with a longitudinal black band running through its medi- an gormazi Labrum not as above sinuosa Erons with less than 6 setae confluentesignata Erons with more than 6 setae 17 Labrum with a longitudinal dark band running through its median chiliensis Labrum not as above 18 ' C. argentinica (Mandl) will also key out in this couplet. See under Discussion section. ^ Pentacotnia cribrata (Brulle) will also key out in this couplet. It can be separated from C. misella Chaudoir by its glabrous undersides. Also, see Rivalier (1969) for further discussion. 100 NEW YORK ENTOMOLOGICAL SOCIETY H F I E Fig. I. Left elytron of a hypothetical Cicindela species illustrating terminology used in key: A) basal dot, B) humeral lunule, C) marginal line, D) apical lunule, E) microserrations, F) sutural spine, G) middle band, H) subsutural row of foveae. 18. Labrum with more than 9 subapical setae mixtula - Labrum with 9 or less subapical setae 19 19. Elytral ground color black melaleuca - Elytral ground color brown or reddish brown 20 20. Maculation not contacting medial lateral edge of elytra ritsemai - Maculation contacting medial lateral edge of elytra 21 VOLUME LXXXVIl, NUMBER 2 101 21. I 23. i 24. t i - I 25. I “ 26. i 27. ’ 28. I 29. Marginal line not converging on humeral lunule drakei drakei Marginal line converging on humeral lunule 22 Size larger (12 mm in length) sinuosa Size smaller (9 mm in length) 23 Head and pronotum dark cupreous red drakei pseudochiloleuca Head and pronotum dark green drakei latifascia n. ssp. Frons completely covered from view with white decumbent setae hirsutifrons n. sp. Frons glabrous or with setae but not covered from view 25 Elytra dull, reflecting little light 26 Elytra shiny, reflective 28 Vertex glabrous confluentesignata Vertex setose 27 Antennal scape glabrous (except for subapical sensory setae) patagonica Antennal scape with several white setae (if setae are rubbed off, minute dimples are present) ramosa Clypeus glabrous halophila n. sp. Clypeus setose 29 Antennal scape glabrous (except for subapical sensory setae) siccalacicola n. sp. Antennal scape with several white setae nivea In the ensuing section only the new taxa are fully discussed. The remain- ing species and subspecies are briefly discussed. The known ranges and collecting data (where available) are presented alphabetically by province. Many of the study specimens are from older collections and carry labels simply stating “Argentina” or “Buenos Aires” or some other province with no other data. For this reason, there are no collecting data provided for several taxa. The phylogeny of Rivalier (1954), with slight modification, is followed. Subgenus Cicindelidia Rivalier, 1954 Cicindela rufoaenea W. Horn , Cicindela unicolor W. Horn 1892, Deutsche Ent. Zeitschr. p. 86 (preoccu- pied). Cicindela rufoaenea W. Horn 1915, in Wytsman, Gen. Ins. Cic. p. 402. This species is the only representative of the above subgenus known from Argentina. It can be distinguished immediately by its immaculate, red-brown 1 color and testaceous abdomen. Type locality. — “Argentinisches Hochgebirge.” Range. — Jujuy: 5 km. S. San Pedro, 5-V-64; Salta; Peru; Colombia. 2a 2b Fig. 2a. Left elytron of C. argentata showing a rounded apex. 2b. Left elytron of C. misella showing an angular apex. Subgenus Brasiella Rivalier, 1954 Cicindela argentata argentata Fabricius Cicindela argentata Fabricius 1801, Syst. Eleuth. I, p. 242. C. argentata, once thought to be a single species, has proven to be a “species swarm.” Many of the species within this swarm can only be sep- arated by characters contained within the aedeagus. Argentine samples of C. argentata run from well-marked light-brown in- dividuals to dark-brown or black individuals with broken lunules, seemingly in a random fashion. However, all the samples have several characters in common: truncate labrum, rounded elytral apices and the characteristic ar- gentata inner sac of the aedeagus. Type locality. — “America Meridionali.” Range. — Buenos Aires: San Fernando, III-1956; Corral del Cruz, 11-1964; Chaco: Cordoba: V. Dolores, 11-1959; Formosa: P. Irigoyen, XII-1950; Jujuy: Misiones: Salta: Gen. Ballivian; C. Olleros, 1-1958; San Luis: Santa Fe: Dpto. Garay, III-1950; Gran Guardia, 11-1953; Santiago del Estero: Cap. Aeropuerto, 11-1961; Tucuman: Tucuman, 1-51; Brazil; Venezuela; Colom- bia; Paraguay. VOLUME LXXXVII, NUMBER 2 103 Cicindela argentata semicircumscripta Mandl Cicindela argentata semicircumscripta Mandl 1958, Ent. Nachr. Osterr. Schweiz. Ent., 10(2), p. 23. This subspecies can be separated from the nominate form by its greenish elytra and long marginal line. Type locality. — “Santiago del Estero, El Pinto.” ; Range. — Santiago del Estero: El Pinto, XI-1962; Rio Saladillo, 29-XII- I 1975; Suncho Corral, 28-XII-1975; Paraguay. Cicindela argentinica (Mandl) Brasiliella (sic) argentinica Mandl 1963, Ent. Arb. Mus. G. Frey, 14, p. 583. No specimens of this species were available for study. In reading Mandl’s description and studying his accompanying figures, one quickly learns that 1 it is quite difficult to separate C. argentinica from C. argentata. According ' to the description, argentinica has an almost cylindrical pronotum and an apical lunule that extends a good distance up the suture. Type locality. — “Argentine, Rio Salado, Icano.” Range. — At present, known only from the type locality. Cicindela obscurella obscurella Klug Cicindela obscurella Klug 1829, Preis-Verz. Ins. Mus. Berlin, p. 3. This species can be identified by its lack of an apical lunule and its round- 1 ed, slightly produced labrum. Some specimens possess a spot representing the humeral lunule. Type locality. — “Sud-Brasilien. ’ ’ Range. — Buenos Aires: Corral del Cruz, 11-1964; Cordoba: Tucuman: Tucuman, 7-II-1908; Brazil; Uruguay. Cicindela aureola aureola Klug Cicindela aureola Klug 1834, Jahrb. Ins. I, p. 35. C. aureola can be separated immediately from the rest of the argentata complex by its green tinged cupreous red color. Apparently it is sexually dimorphic, with respect to the shape of the labrum; males possess short labra while the female’s is produced. Type locality. — “Sudlichen Brasilien.” Range. — Jujuy: Salta; Tucuman; Paraguay; Brazil. 104 NEW YORK ENTOMOLOGICAL SOCIETY Cicindela misella misella Chaudoir Cicindela misella Chaudoir 1854, Bull. Soc. Imp. Nat. Moscou, 27, p. 121. For years this species was thought to be confined to Central America and northern South America. Vidal Sarmiento (1966) proved its existence in Argentina. It can be separated from argentata by its angular elytral apices and characteristic inner sac. Type locality. — “Colombie.” Range. — Jujuy; Salta; Venezuela; Colombia north to Guatemala. Cicindela stamatovi n. sp. (Fig. 3a, b) Head. — Labrum white with a narrow, dark anterior border, equipped with a single row of subapical setae (8 setae), slightly produced, rounded, single- toothed; first four antennal segments metallic cupreous with green reflec- tions; scape with a single subapical seta; second, third and fourth segments with a few erect white setae; clypeus, frons and genae glabrous, finely ru- gose and striate; vertex glabrous (except for supraorbital sensory setae). Thorax. — Pronotum with sparse decumbent setae on lateral margins and disc, finely rugose, narrowed posteriorly; pro-, meso- and medial metaster- num glabrous; remainder of thorax with sparse erect setae. Abdomen. — Lateral edges of venter with sparse covering of decumbent setae, glabrous medially. Elytra. — Both sexes, nearly parallel-sided although slightly wider from ( basal third to apical fourth, then more or less oblique from apical sixth to ; apex; both sexes with small sutural spines, microserrate apical margins and i shallow green punctae; elytral surface granulate; maculation complete, hu- meral lunule long and slender, descending nearly to apical half, remainder of lunules of the argentata type. Color. — Head, dorsal and lateral portions of thorax metallic cupreous with green reflections; ventral thorax and abdomen metallic green to dull, dark green; elytra cupreous with metallic green punctae. Size. — Male, 6.4 mm length, 2.2 mm width; female, 6.5 mm length, 2.4 mm width. Type locality. — Tucuman, Argentina. Holotype. — Male. Tucuman, 11-50, Argt. Allotype. — Female. Same data as holotype. Holotype and allotype to the I American Museum of Natural History, New York City, New York. \ Etymology. — I take pleasure in naming this species after Dr. John Sta- ' i matov, who kindly submitted the specimens for study. i Diagnosis. — C. stamatovi appears to be closest to C. misella in the Ar- I gentine fauna. It can be separated from misella by its produced labrum and VOLUME LXXXVIl, NUMBER 2 105 complete humeral lunules. C. stamatovi is apparently the only species in the ""argentata complex” with a complete humeral lunule. Cicindela chlorosticta Kollar (Figs. 3, 4) Cicindela chlorosticta Kollar 1836, Ann. Wiener Mus. Naturg., 1 (2), p. 332. No Argentine representatives of this species were available for study. Key characters were drawn from specimens collected in Brazil. Type locality. — “Brasiliae Provincia Ypanema.” Range. — Jujuy\ Brazil. Cicindela staudingeria W. Horn I Cicindela staudingeri W. Horn 1892, Deut. Ent. Zeitschr., p. 368 (preoc- cupied). Cicindela chlorosticta staudingeria W. Horn 1915, in Wytsman, Gen. Ins. Cic., p. 404. No Argentine representatives of this species were available for study. Key characters were drawn from Brazilian specimens. It differs from chlo- rosticta by its larger size and coloration. Most specimens are a brilliant cupreous color. Type locality. — “Sao Paulo.” Range. — Argentina; Brazil. Cicindela obsoletesignata W. Horn Cicindela obsoletesignata W. Horn 1895, Deutsche Ent. Zeitschr., p. 91. No specimens of this species were available for study. It is apparently quite rare in collections. Horn (1895) compares it with C. hemichrysea Chevrolat but the coloration is similar to C. morio Klug. Type locality. — “St. Catharina.” Range. — Santa Fe = Chaco Santafecino; Brazil. Subgenus Cylindera Westwood, 1831 Cicindela confluentesignata W. Horn Cicindela confluens W. Horn 1893, Deutsche Ent. Zeitschr., p. 197 (preoc- cupied). Cicindela confluentesignata W. Horn 1915, in Wytsman, Gen. Ins. Cic., p. 407. 106 NEW YORK ENTOMOLOGICAL SOCIETY Eig. 3a. Dorsal aspect of holotype male of C. stamatovi n. sp. 3b. Dorsal aspect of labrum of C. stamatovi n. sp. Eig. 4a. Dorsal aspect of male C. halophila n. sp. 4b. Dorsal aspect of labrum of C. halophila n. sp. All scale lines indicate 1 mm. One Argentine representative of this species was available for study. Key characters were drawn from that exemplar and from Paraguayan specimens. It is a distinctive species, bronze-black in color, with many elytral foveae. Type locality. — “Minas Geraes.” VOLUME LXXXVII, NUMBER 2 107 Range. — Entre Rios: Pronunciamiento, XII-1961; Formosa, Santiago del Estero; Brazil; Paraguay. Cicindela eugeni Castelnau Cicindela eugeni Castelnau 1835, Etudes Ent., pt. 1, fasc. 1, p. 36. A distinctive species not treated by Rivalier (1954). Known by its con- fluent maculation and setose elytra. ' Type locality. — “Cordova.” Range. — Cordoba, Santiago del Estero: 50 km. NE. Santiago del Estero, ' 16-XII-1971. I Cicindela halopliila n. sp. (Eig. 4a, b) i Head. — Labrum white with a single row of subapical setae (4-5 setae), I slightly produced, single toothed; antennal scape metallic green with cu- I preous reflections, equipped with a single subapical seta; third antennal segment with 6-8 erect white setae; clypeus glabrous, metallic green; genae ' with sparse covering of white decumbent setae; frons and vertex glabrous I (except for two erect supraorbital setae, per side, near front of eye and one ! over eye), finely wrinkled. Thorax. — Pronotum with sparse covering of white decumbent setae (ex- treme medial area glabrous), finely rugose; proepisternum, prosternum, proepimeron, procoxa, mesepimeron, mesocoxa, metaepisternum, anterior edge of metasternum and lateral edge of metacoxa with covering of white i decumbent setae; mesepisternum with decumbent setae along ventral edge, mesosternum glabrous. Abdomen. — Lateral edges of venter with a covering of white decumbent setae, glabrous medially; posterior two, visible, abdominal segments red- testaceous. Elytra. — Male and female, nearly parallel-sided although slightly wider from basal third to apical third, then gradually rounded from apical fifth to apex; males without sutural spines; females with a small, slightly retracted spine; both sexes with microserrate apical margins and shallow punctae; maculation consists of a broad marginal band running from the humeral area along the lateral edge to the apex and thence to the suture at the apical fifth; humeral lunule enters disc obliquely; middle band begins descent into disc at apical half and descends nearly to apical fifth. Color. — Anterior portion of head metallic green with cupreous reflections; remainder of head cupreous metallic (except for genae, which are metallic green); pronotum metallic red cupreous with metallic green sulci; lateral portions of thorax metallic cupreous; undersides all metallic cupreous with 108 NEW YORK ENTOMOLOGICAL SOCIETY green tinges; abdomen dull green metallic with last two segments red-tes- taceous; elytra bright red-brown with green punctae throughout pigmented portion. Size. — Male, 7.5 to 7.9 mm in length, 2.5 to 2.8 mm in width; female, 8.1 to 8.5 mm in length, 2.8 to 3.0 mm in width. Type locality. — Argentina, Cordoba Province, 24 mi. S. Recreo. Holotype. — Male. 24 mi. S. of Recreo, Cordoba, R. A., II-9-51, Salt Flat, Ross and Michelbacher collectors. Allotype. — Female. Same data as holotype. Paratypes. — 4 males, 3 females (one female damaged), same data as ho- lotype; 2 males. La Colina B. A., Buenos Aires, Arg., 12-9-1938, Carl J. Drake; 1 female, Estrella, Chaco, Prg., XII-43, N:l. Holotype, allotype and three paratypes to the California Academy of Sciences, San Francisco, Cal- ifornia; three paratypes to the National Museum of Natural History, Wash- ington, D.C.; one paratype to the American Museum of Natural History, New York City, New York. The remaining paratypes are in the author’s collection. Etymology. — Name derived from the Greek for salt-lover, alluding to the new species’ occurrence on a salt playa. Diagnosis. — C. halophila n. sp. appears to be closest to C. eugeni. It can be separated from that species by slightly different markings, color, elytral lustre, and absence of setae on the anterior basal half of the elytra, and the presence of setae on the genae. Cicindela morio morio Klug Cicindela morio Klug 1834, Jahrb. Ins. I, p. 16. No Argentine representatives were available for study. Key data were obtained from Brazilian specimens. Type locality. — “Brasilien.” Range. — N. Argentina; Brazil. Cicindela sinuosa Brulle Cicindela sinuosa Brulle 1837, Voyage Orbigny, Ins. Col., p. 8. A single female specimen referrable to this species, labeled “Misiones,” was used for key data. Brulle compared sinuosa with C. trisignata Dejean and his description seems to maintain that comparison. The illustration by Horn ( 1938, Table 84, Fig. 1 1), however, bears little resemblance to Brulle’s description or the specimen currently at hand. The study specimen (badly damaged) resembles a broadly marked C. mixtula W. Horn. Type locality. — “Corrientes.” Range. — Corrientes', Entre Rios; Misiones. VOLUME LXXXVII, NUMBER 2 109 Cicindela mixtula mixtula W. Horn Cicindela mixta W. Horn 1892, Deutsche Ent. Zeitschr., p. 215. (preoccu- pied). Cicindela mixtula W. Horn 1915, in Wytsman, Gen. Ins. Cic., p. 409. Apparently a fairly common species in Argentina — at least it is frequently collected. Type locality. — “Tarija (Bolivia).” Range. — Catamarca: S. Maria, 19-1-1945; Formosa: Desmonte, XII- 1950; Jitjuy: Quemado, III-1926; Mendoza', Salta: La Termes, 17-X-1975; San Juan: Retamito, 17-III-1907; San Luis: Kahuel Mopa, 1-1964; Tucu- man: Cadillal, 4-XII-1975; Tucuman, 20-XI-1913, 2-XI-1916, 30-XI-1919; Vipos, IV- 1960; Bolivia. Cicindela drakei drakei W. Horn Cicindela drakei W. Horn 1892, Deutsche Ent. Zeitschr., p. 85. This species is easily confused with C. melaleuca reedi W. Horn but, may be distinguished by its brown color, instead of the black of reedi. Type locality. — “Mendoza.” Range. — Mendoza', Salta; Tucuman. Cicindela drakei pseudochiloleuca W. Horn Cicindela drakei pseudochiloleuca W. Horn 1908, Ent. Wochenblatt, 25, p. 209. Subspecies pseudochiloleuca is differentiated by its smaller size and the shape and extent of the lunules; it has a marginal line that is connected to the humeral lunule, something not found in the nominate form. Also, pseu- dochiloleuca possess a rather diffuse, indistinct descending portion of the middle band. Type locality. — “Tukuman (Argentien).” Range. — Tucuman. Cicindela drakei latifascia n. ssp. (Fig. 5) Similar in most respects to subspecies pseudochiloleuca but differing in that the new subspecies possesses a green head and pronotum (as opposed to the cupreous color of the former) and the lunules are much wider and more distinct. Type locality. — Terma-Rio Hondo, Santiago del Estero Province, Argen- tina. 110 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 5. Dorsal aspect of holotype male of C. drakei latifascia n. ssp. Scale line indicates 1 mm. Holotype. — Male. Argentina, Sgo. Estero, Termas de Rio Hondo, 30-XII- 1975, R. M. Bohart. Holotype to the University of California, Davis collec- tion. Etymology. — Name derived from the Latin lata (wide) and fascia (mark). Diagnosis. — The new subspecies can be easily separated from the nom- inate subspecies and pseudochiloleuca by its green head and pronotum and the very wide markings. After studying long series of pseudochiloleuca (to which latifascia appears closest) from Tucuman, I found no tendency to- ward wide lunules or green coloration of the head or pronotum. Cicindela melaleuca melaleuca Dejean Cicindela melaleuca Dejean 1831, Spec. Col. V, p. 238. VOLUME LXXXVII, NUMBER 2 111 C. melaleuca is a distinctive black species with a humeral lunule that normally comes in contact with the anterior-most part of the middle band. Type locality. — “Buenos-Ayres.” Range. — Buenos Aires: San Fernando, XII- 1962; Guamini, 11-1964; La Colina, 9-XII-1938; Mendoza: Patagonia. Cicindela melaleuca reedi W. Horn Cicindela reedi W. Horn 1895, Deutsche Ent. Zeitschr., p. 88. This subspecies has been placed under C. drakei by several workers including Horn (1926). Horn (1938), however, placed it with C. melaleuca, where I feel it justly belongs. The differences in genitalia and external morphologies between C. drakei and C. melaleuca are slight. When large samples are examined from their respective ranges it may be found that the two species are actually one — a brown form with its melanic counterpart. Type locality. — “Patagonia.” Range. — Patagonia. Cicindela siccalacicola n. sp. (Fig. 6a, b) Head. — Labrum white with a narrow dark anterior border, equipped with a single row of subapical setae (6-9 setae), slightly produced, rounded, single toothed; first four antennal segments metallic cupreous with green reflections; scape with a single subapical seta; second, third and fourth segments with several erect setae; clypeus and genae with sparse covering of partially erect white setae; frons and vertex with sparse covering of par- tially erect to erect white setae. Thorax. — Pronotum sparsely covered with partially erect white setae, finely rugose; pro- and mesosternum glabrous, remainder of thorax with sparse, erect setae. Abdomen. — Lateral edges of venter with a covering of decumbent setae, glabrous medially. Elytra. — Male, nearly parallel-sided although slightly wider from basal fourth to apical third, then gradually rounded from apical fourth to apex; female, markedly widened from basal third to apical third, then gradually rounded to apex; both sexes equipped with small sutural spines, microser- rate apical margins and shallow, green punctae; maculation complete, mar- ; ginal line broad, connected with humeral lunule and outer edge of apical I lunule; humeral lunule a knob pointing laterally; middle band elbowed at I center of elytral disc, descending to edge of apical third; apical lunule large, comma-shaped. 112 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 6a. Dorsal aspect of male C. siccalacicola n. sp. 6b. Dorsal aspect of labrum of C. siccalacicola n. sp. Fig. 7a. Dorsal aspect of holotype male of C. hirsutifrons n. sp. 7b. Dorsal aspect of labrum of C. hirsutifrons n. sp. All scale lines indicate 1 mm. Color. — Head predominately metallic cupreous with green reflections; pronotum cupreous with green reflections, medial sulcus metallic green; lateral portions of thorax metallic cupreous; ventral portions of thorax blue- green metallic with cupreous reflections; abdomen dark brown with green VOLUME LXXXVIl, NUMBER 2 113 reflections, cupreous laterally; elytra dark red-brown covered with metallic green punctae. Size. — Male, 10.5 mm length, 3.5 mm width; female, 11.5 mm length, 4.5 mm width. Type locality. — 24 mi. S of Recreo, Cordoba Province, Republic of Argentina. Holotype. — Male. 24 mi. S Recreo, Cordoba, R. A., 11-9-51, Salt Flat, Ross and Michelbacher collectors. Allotype. — Female. Same data as holotype. Paratypes. — 16 males, 12 females same data as holotype. Holotype, allotype and 17 paratypes to the California Academy of Sci- ences, San Francisco, California; one paratype each to the National Mu- seum of Natural History, Washington, D.C., and the American Museum of Natural History, New York City, New York. The remaining paratypes are in the author’s collection. Etymology. — The name of the new species is derived from the latin sicca (dry), lacuna (lake), and incola (dweller). Diagnosis. — This species is apparently not closely related to any other known from Argentina. With its unusual maculation pattern and near-glossy elytra it is quite distinct. Discussion. — The range of this species apparently extends into Buenos Aires Province as I have before me a female (not in the type series) collected i at La Colina. This specimen is larger (13.5 mm in length) than the females of the Cordoba series and is dark green in color. The status of this population is unknown. Cicindela ritsemai W. Horn Cicindela ritsemae W. Horn 1895, Notes Leyd. Mus., 17, p. 15. Cicindela ritsemai W. Horn 1915, in Wytsman, Gen. Ins. Cic., p. 409. A distinctive little species, apparently rare in collections. It slightly re- sembles C. drakei pseudochiloleuca, but can be separated from that taxon by the marginal line that fails to contact the elytral lateral margins. Type locality. — “Argentiniae (Provincia Catamarca).” Range. — Catamarca', Cordoba', Santiago del Estero: Termas de Rio Hon- do, XII-30-1975; Suncho Corral, XII-28-1975; Tucuman: Cadillal, XII-4- 1975. Cicindela gormazi Reed Cicindela gormazi Reed 1871, Ent. Mo. Mag., 8, p. 76. No Argentine specimens were available for study. Key data were drawn from Chilean specimens. 114 NEW YORK ENTOMOLOGICAL SOCIETY Type locality. — “Chile merid." Range. — Chubuf, Patagonia', Chile. Cicindela chiliensis Audouin & Brulle Cicindela chiliensis Audoin & Brulle 1839, Arch. Mus. Hist. Nat., 1, p. 133. No Argentine representatives of this species were available for study. Key data were extracted from Chilean specimens. Type locality. — “Chili.” Range. — Chubuf, Patagonia', Chile. Cicindela patagonica patagonica Brulle Cicindela patagonica Brulle 1837, Voyage Orbigny, Ins. Col., p. 7. This species is readily confused with C. ramosa Brulle. They are easily separated by the lack or presence of setae on the scape of the antenna. Nominate patagonica is a brown tiger beetle with fairly thin markings. Type locality. — “Rio Negro (Patagonie).” Range. — Buenos Aires', Patagonia. Cicindela patagonica cherubim Chevrolat Cicindela cherubim Chevrolat 1858, Ann. Soc. Ent. France, ser. 3, 6, p. 315. Similar to the nominate form except that the markings are much wider, the ground color is blue and the elytra are expanded laterally. Type locality. — “Montivedeo.” Range. — Buenos Aires', Uruguay. Cicindela patagonica bergiana W. Horn Cicindela patagonica var. bergiana W. Horn 1895, Anal. Mus. Nac. Buenos Aires, 4, p. 174. Identical with the preceding form except that the ground color is brown. Type locality. — “Montevideo.” Range. — Buenos Aires', Uruguay. Cicindela apiata apiata Dejean Cicindela apiata Dejean 1825, Spec. Col., 1, p. 86. Apparently the most common member of the genus is Argentina — at least it is collected more often than other species. VOLUME LXXXVII, NUMBER 2 115 Type locality. — “meridionale du Bresil.” Range. — Buenos Aires: Buenos Aires, 11-1936, XII-1940; La Colina, 29- XI-1938; R. Sauce Grande, 23-11-1968; San Fernando, 1-1962, Tandileofu, 23-11-1968; Veronica, 1-1922; Cordoba: 22 km. S. Alta Gracia, 18-X1-1975; Capilla del Monte; Cosquin, 1-9-111-1920; Entre Rios: Concordia; 4 km. N. Va. San Jose, 15-X1-1973; Mendoza', San Luis', Santiago del Estero: Ter- mas de Rio Hondo, 30-X11-1975; Tucuman', Brazil, Paraguay; Uruguay. Cicindela ramosa Brulle Cicindela ramosa Brulle 1837, Voyage Orbigny, Ins. Col., p. 7. A distinctive species with very broad markings. It is usually green in color. Type locality. — “baie de San-Blas.” Range. — Buenos Aires: Buenos Aires, Xll-1938; Guamini, 11-1964; Lag. Las Tunas, 4-1X-1961; Catamarca: Andaleala, Xl-1945; Patagonia', Uru- guay. Cicindela nivea nivea Kirby Cicindela nivea Kirby 1818, Trans. Linn. Soc. London, 12, p. 376. No Argentine representatives of this subspecies were available for study. Key data were drawn from Brazilian specimens. It is a large Argentine Cicindela (13 mm in length) with markings so broad they completely cover the elytra with white in most examples. Type locality. — “Brasilia.” Range. — Buenos Aires', Misiones', Entre Rios; Brazil; Uruguay. Cicindela nivea orbignyi Guerin-Meneville Cicindela intricata Brulle 1837, Voyage Orbigny, Ins. Col., p. 7 (preoccu- pied). Cicindela orbignyi Guerin-Meneville 1839, Rev. Zool., p. 296. Differentiated from the nominate form by the increased amount of pig- mentation on the elytra, i Type locality. — “la Patagonie.” ‘ Range. — Buenos Aires: Necochea, 11-1927, Xll-1963; Patagonia; Uru- guay. I Cicindela hirsutifrons n. sp. , (Fig. 7a, b) I Head. — Labrum testaceous with a narrow border of dark brown on an- I terior edge (labral color in a degreased specimen would probably be white). 116 NEW YORK ENTOMOLOGICAL SOCIETY equipped with a single row of subapical setae (4 setae), produced, rounded, single toothed; first four antennal segments green-metallic with cupreous reflections, scape with a single subapical seta, segments 3 and 4 nearly covered with broad, white decumbent setae; segment 2 glabrous; clypeus and genae with dense covering of wide, white decumbent setae; frons and anterior edge of vertex completely covered with wide, decumbent setae; remainder of vertex glabrous (except for supraorbital sensory setae); base of mandibles with a sprinkling of decumbent setae. Thorax. — Pronotum nearly covered with white decumbent setae, disc sparsely setose, finely rugose; proepisternum, proepimeron, procoxa, mes- epimeron, socoxa, metaepisternum, anterior edge of metasternum, ventral edge of mesepisternum and lateral edge of metacoxa with dense covering of white decumbent setae; mesosternum glabrous. Abdomen. — Lateral edges of venter with dense covering of white decum- bent setae, sparsely setose medially; posterior two visible, abdominal seg- ments red-testaceous. Elytra. — Male, nearly parallel-sided, slightly expanded from basal fourth to apical fourth, then gradually rounded to apex; female, unknown; male without sutural spines or microserrate apical margins, shallow punctae cov- er surface; maculation consists of a band of white starting at the humerus and running to the suture at the apical fourth gradually widening as it pro- gresses so that over 60% of the elytra are white; middle band, humeral and apical lunules are only hinted at, very confluent type of maculation. Color. — What can be discerned (due to extreme pilosity) of the anterior portion of the head is metallic green with cupreous reflections; genae me- tallic cupreous; vertex metallic cupreous with green reflections; pronotum metallic cupreous with medial sulcus metallic green; lateral and ventral por- tions of thorax metallic cupreous with green tinge; abdomen dark brown with metallic red reflections with last two segments red-testaceous; elytra bright red metallic. Size. — Male, 8.6 mm in length, 2.5 mm in width; female, unknown. Type locality. — 24 mi. S. Recreo, Cordoba Province, Republic of Argen- tina. Holotype. — Male (damaged). 24 mi. S. Recreo, Cordoba, R. A., 9-II-1951; Salt Flat, Ross and Michelbacher collectors. Holotype to the California Academy of Sciences, San Francisco, California. Etymology. — The new species name is a combination of the latin hirsuta (hairy) and frons (forehead). Diagnosis. — C. hirsutifrons appears to be closer to C. nivea than to any other species in Argentina. It can be separated readily from nivea by its smaller size, coloration, extent of pilosity, shape and type of maculation. Remarks. — Although the holotype is damaged (it appears to have been VOLUME LXXXVII, NUMBER 2 117 Struck with the edge of a collecting net); it is nonetheless complete, except for a portion of one antenna. Acknowledgments I would like to thank the following people for the use of material utilized in this study: Dr. George Byers, The University of Kansas, Lawrence, Kansas; Dr. Terry Erwin, National Museum of Natural History, Washing- ton, D.C.; Dr. Lee Herman, American Museum of Natural History, New York City, New York; Dr. David Kavanaugh, California Academy of Sci- ences, San Francisco, California; Mr. Robert Murray, Texas A&M Uni- versity, College Station, Texas; Dr. L. L. Pechuman, Cornell University, Ithaca, New York; Mr. Robert Schuster, University of California, Davis, California; Dr. John Stamatov, Armonk, New York; Dr. Rupert Wenzel, Field Museum of Natural History, Chicago, Illinois. I would also like to extend thanks to Mr. Bert Kohlmann for his assistance in German translation and Mrs. Marian Seibert for typing the manuscript. Literature Cited Bruch, C. 1910. Catalogo sistematico de los coleopteros de la Republica Argentina. Rev. Mus. La Plata 4: 143-166. Blackwelder, R. E. 1944. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part I. Bull. U.S. Natl. Mus., No. 185:1- 188. Horn, W. 1895. Zwolf neue Cicindeliden-Species. Deutsche Ent. Zeitschr. 1895:81-93. . 1905. Systematischer Index der Cicindeliden, 56 pp. Berlin. (Published in connection with Deutsche Ent. Zeitschr., 1905, Heft 2.) . 1915. Coleoptera. Adephaga. Family Carabidae, subfamily Cicindelinae. Genera in- sectorum dirges par P. Wytsman. Louis Desmet-Verteneuil, Brussels. Ease. 82C:209- 486, 8 pis. . 1926. Carabidae: Cicindelinae. In Junk, W., Coleopterorum catalogus. Berlin. 1:1- 345. . 1938. 2(KX) Zeichnungen von Cicindelinae. Ent. Beih., Berlin-Dahlem. 5:1-71, 90 pis. Rivalier, E. 1954. Demembrement du genre Cicindela Linne. II. Faune americaine. Rev. franc. d’Ent. 21:249-268. . 1%9. Demembrement du genre Odontochila (Col. Cicindelidae) et revision des prin- cipales especes. Ann. Soc. Ent. France. (N. S.). 5 (1): 195-237. Vidal Sarmiento, J. 1966. Las especies Argentinas de los generos Cicindelidia Riv., Brasiella Riv. y Cylindera Westwood. Rev. Mus. La Plata. (N. S.). 9 (68):25-46. Riverside County Agricultural Commissioner’s Office, Riverside, Califor- nia 92501. Received for publication November 1, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 118-125 REPRODUCTIVE BEHAVIOR OF EUARESTA BELLA AND E. FESTIVA (DIPTERA: TEPHRITIDAE), POTENTIAL AGENTS FOR THE BIOLOGICAL CONTROL OF ADVENTIVE NORTH AMERICAN RAGWEEDS {AMBROSIA SPP.) IN EURASIA S. W. T. Batra Abstract. — Larvae of Euaresta bella (Loew) and E. festiva (Loew) de- stroy seeds of Ambrosia artemisiifolia L. and A. trifida L., respectively; E. bella has been colonized in the USSR for ragweed control. The univoltine adults feed on honeydew and ragweed sap. In Maryland, E. bella adults reach peak abundance in mid-August, E. festiva a month later. Courtship behavior by both sexes is complex, involving several stereotyped wing movements, proboscis and foot contact, head butting, and male territoriality or lekking. Oviposition occurs in the young female flowers. Introduction The native North American ragweeds. Ambrosia artemisiifolia L., A. trifida L. and A. psilostachya DC., have become accidentally established in Eurasia, where they cause increasing agricultural and medical problems (Goeden et al., 1974). Ambrosia artemisiifolia and A. trifida are abundant in the northeastern United States, producing about 90% of the hayfever- causing late summer pollen to which about 4% of the population is sensitive (Dickerson and Sweet, 1971). These rapidly growing annuals are among the most important weeds in row crops, pastures, ornamentals and small fruits in the United States (Danielson et al., 1965). A. artemisiifolia and A. trifida have become established and troublesome in the southwestern USSR (Shu- tova, 1970), which is climatically similar to the eastern United States. The study of the bionomics of insects that attack ragweed in the northeastern United States was initiated by the author in 1976, and several species were shipped to the USSR. These included 425 adult Euaresta bella (Loew), which were colonized in 1977 at Sochi (O. V. Kovalev, personal comm.). Euresta bella is a small univoltine tephritid that occurs throughout the United States on A. artemisiifolia, its sole host (Foote, 1965; Wasbauer, 1972). In Ohio adults are found from mid-June to late September; repro- duction begins in August; the larvae develop and overwinter in the seeds, destroying 1-15%; and pupation occurs in spring (B. A. Foote, 1965; in litt. 1977). Euaresta festiva (Loew) is a larger univoltine species that attacks only A. trifida in the eastern and midwestem states. In Ohio adults appear VOLUME LXXXVII, NUMBER 2 119 in July; larvae destroy 2-25% of the seeds, where they overwinter; and pupation occurs in spring (B. A. Foote, 1965; in litt., 1977). The seed-feeding habit of larvae of these two tephritids contrasts with that of the larvae of Euarestoides acutangulus (Thomson), which destroy the staminate flowers of Ambrosia spp. (Piper, 1976). Field and cage observations of Euaresta bella and E. festiva were made between 1976 and 1978 at the USDA Agricultural Research Center, Belts- ville, Maryland. Caged adult flies were kept in a rearing room at 16L:8D hr. photoperiod, at about 25°C, or were kept in a greenhouse at ambient day- length at about 27-30°C. They were provided with vigorous potted host plants, water droplets, and food (4:1 sucrose /autolyzed yeast extract in water). Individual flies lived as long as two weeks. E. bella adults began to appear in the field at Beltsville in mid-July, the females at that time being noticeably slender, with translucent abdomens; at this time the host plants were not yet blooming. The adult population of this insect reached a maximum in mid-August, when the male and female flowers of A. artemisiifolia began to bloom, and abdomens of female flies at this time were swollen and opaque. The fly population subsequently de- clined rapidly as most ragweed plants set seed, although male flowers con- tinued to produce abundant pollen. By the end of August, E. bella adults were scarce. However, E. festiva reached its peak abundance in mid-Sep- tember on the later-blooming A. trifida. Both E. bella and E. festiva were relatively most active and phototactic i during the late afternoon. They flew poorly, E. bella hopping only a few centimeters from leaf to leaf; E. festiva flying erratically for one or two meters to adjacent plants. Dispersive ability of individual flies is apparently slight, as in Urophora jaceana (Hering), which did not cross a two meter wide cart track to host plants on the other side (Varley, 1946). Adult E. bella and E. festiva often licked ragweed leaves. Additionally, E. bella licked dew, honeydew of aphids and leafhoppers, the foam of cer- copid larvae on ragweed stems, sap oozing from damaged leaves, and the sucrose-yeast- water mixture. A droplet of food sometimes was repeatedly regurgitated and reingested by both sexes, behavior resembling nectar-whip- ping by bees, which reduces the water content. Honeydew and dew are often eaten by other tephritids (Christenson and Foote, 1960; Bateman, 1972). Courtship and Territoriality The courtship behavior of E. bella and E. festiva is variable and complex (Table 1), involving stereotyped movement of the conspicuously patterned wings, visual orientation, wing and abdomen vibration, proboscis and foot contact, head butting, and lekking or male territoriality. As in Valentibulla 120 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Major courtship behavior patterns of E. bella and E.festiva. C, common, R, rare, — , not seen. £. bella E. festiva s 9 6 9 (1) Visual orientation C c c c (2) Alternate wing waving with C c c c vibration (3) Both wings extended R — — — horizontally (4) Both wings extended with proboscis extended C c Front legs also extended (5) Both wings extended with C C R — head butting (6) Tapping with front feet — R R — (7) Rapid flicks of both wings simultaneously c C With proboscis extension (8) Territoriality (lekking) c — C — (9) Eollowing female, abdomen R — C — curved (10) Following female, wings — — C — flattened against abdomen (Wangberg, 1978), both sexes play active roles in courtship and the sequence of courtship displays is frequently interrupted and resumed at various stages. Visual orientation by both sexes toward conspecific flies of both sexes j (Table 1: item 1), and to other small insects was common. Orienting flies { turned to face these insects as they moved about, and grooming was oc- casionally begun by flies that had been watching conspecifics or other in- sects that were grooming themselves. Orientation is common in many te- phritids (Tauber and Toschi, 1965; Piper, 1976; Berube, 1978; Wangberg, 1978) and in Drosophila (Spieth, 1974). During orientation, random walking, and (in females) while mating, both ' species of Euaresta often slowly waved alternate wings. A slight vibration of the extended, vertical wing was sometimes seen, and side-stepping may occur (Table 1: 2). Occasionally an E. bella male or female faced another : fly of either sex while waving alternate wings (Fig. 1); the other fly respond- ed by synchronous ipsilateral wing waving. Similar behavior occurs in Eu- arestoides acutangulus (Piper, 1976), Tephritis dilacerata Loew (Berube, VOLUME LXXXVIl, NUMBER 2 121 Eigs. 1-4. Euaresta bella on Ambrosia artemisiifolia. 1. Eemale waving alternate wings while approaching a resting male. 2. Male performing the spread-winged display to an intruding male, in defense of its territority. 3 and 4. Females ovipositing through the floral involucre. 1978) and T. stigmatica (Coquillett) (Tauber and Toschi, 1965); it may be a general species recognition signal in E. bella. A behavior pattern peculiar to both sexes (but usually seen in males) of E. bella is a simultaneous horizontal spreading of both wings (Table 1: 3- 5). The vertically tilted wings are spread out for several seconds while the fly turns toward another fly of either sex or another small insect. A con- specific fly may respond by similarly spreading its wings while facing the instigator (Fig. 2); the abdomen may vibrate. After approaching, one of the 122 NEW YORK ENTOMOLOGICAL SOCIETY 6 Figs. 5-8. Euaresta festiva and E. bella, as drawn from color slides. 5. Proboscis contact or ‘kissing’ by male and female E. festiva. 6. Copulation of E. festiva. 7. Female E. festiva ovipositing, as male watches in precopulatory attitude. 8. Copulation of E. bella. VOLUME LXXXVII, NUMBER 2 123 flies may butt the other with its head (Table 1: 5), or both flies may make contact with each other’s proboscis (Table 1: 4); a female may extend her front feet in addition to her proboscis. Unlike Drosophila (Spieth, 1974), and E.festiva, this ‘kissing’ in E. bella is not only used for courtship, but appears to be related to territoriality, because one of the flies (usually not the initiator) abruptly leaves the area after kissing, butting, or wing spread- ing displays. Proboscis extension is usually initiated by males, but occa- sionally females approach other females in this way (20 6: 3 $ -initiated). Butting and chasing other E. bella (including a mating pair) or other insects (gnats, small Hemiptera, beetles, ants), was performed by both sexes (17 6: 6 9). Nonreceptive females thus deterred male advances. Male E.fes- tiva, before following females, extended their probosces to make contact with those of females (Fig. 5) in conjunction with abdomen vibration and wing flicking, as part of the courtship. Rapid, repeated, simultaneous flicking (10/sec.) of both wings is charac- teristic only of males in both E. bella and E.festiva (Table 1: 7). Male E. festiva flicked their wings while orienting toward both sexes, while mating, 1 and occasionally before the spread-winged display. i Lekking or male territoriality by male E.festiva and E. bella was common I (Table 1: 8). This behavior pattern also occurs in male Rhagoletis (Chris- I tenson and Foote, 1960), U.jaceana (Varley, 1946), and Valentibulla spp. i (Wangberg, 1978). Territories of male E. bella, on one or two leaves of the host, were about 7-10 cm sq., and appeared to be occupied by individual : males for only a few hours (males were not marked for recognition); the I same areas were usually not used the next day. The larger territories of i male E.festiva are about 1 m sq.; males returned to their territories after i flying away when they were disturbed by the observer. Male E. bella drove ] other insect species and other males from their territories by butting them 1 or running at them with their probosces and wings extended. Wandering I females were displayed to when they entered or approached male territories. The aggressiveness of males toward each other and toward nonreceptive females may be advantageous to the species, by aiding dispersal and sub- sequent oviposition in a larger number of plants. Female-female aggression, as in Dacus dorsalis Hendel (Christenson and Foote, 1960) also helps to disperse the population and avoid multiple ovipositions in the small Ambro- sia fruits. Male E.festiva, more often than male E. bella, were seen closely following females; their abdomens were ventrally curved (Table 1: 9, 10). These males kept their wings folded closely, one above the other, above their abdomens while following or observing females (Fig. 7); they waited 2-i cm away while females completed probing or ovipositing in flowers before they moved closer and attempted to copulate from the rear. Although mating in 124 NEW YORK ENTOMOLOGICAL SOCIETY E.festiva is associated with oviposition, as in T. dilacerata (Berube, 1978), ovipositing females of E. bella were not noticeably attractive to males. Copulation and Oviposition Mating in E.festiva occurred between 14:20 and 18:00 EOT; two copu- lations lasted 57 and 58 minutes (Fig. 6). Couples walked or flew and both sexes occasionally flicked their wings. In E. bella, mating occurred between 8:00 and 17:00 EDT, with most copulations about 16:00, when the flies generally were most active. Fifteen copulations lasted 20-60 minutes each (Fig. 8). Females walked, fed, performed the spread-winged display, and butted at nearby flies while mating; males occasionally flicked their wings. Fecund females of E. bella and E. festiva wandered over the leaves and stems of their hosts, but female flowers in the upper half of the plants were most attractive to ovipositing flies. Before ovipositing or probing the flowers with the ovipositor, flies walked about over the flowers, and occasionally contacted them with their probosces. E. festiva characteristically stood above the female flowers while probing or ovipositing (Fig. 7). The ovipos- itor was inserted for 10 to 60 seconds; the shorter times being apparently probes, since eggs were not subsequently found, although the sides of the flowers’ involucres and ovaries had been slit (brown slit marks, 0.3 to 0.7 mm long). In E. bella, such probing by the ovipositor slits numerous floral ovaries, causing necrosis and sterility of about 30% of the flowers that were probed, even when eggs were not laid. As described by Foote (1965), eggs of E. bella are partially inserted into young flowers at the edges of flower clusters or deposited on their surfaces; larvae then enter and totally destroy the developing seeds; eggs of E. festiva are placed inside the involucre against the seed coat (B. A. Foote, in litt.). During oviposition, E. bella typically penetrated flowers through the involucre from the side (Figs. 3 and 4), often with a rocking motion as the ovipositor was inserted. Probing and oviposition was seen between 13:30 and 17:05 EDT and each lasted 15 to 50 sec. Ovipositing females often probed several flowers in succession; one female probed six flowers in 15 minutes. Females usually groomed them- selves after probing each flower. Acknowledgments I thank Drs. B. A. Foote of Kent State University and R. H. Foote of the Systematic Entomology Laboratory, IIBIII, SEA, USDA, for references and information regarding these tephritids. Ms. I. Sunzenauer and Mr. D. Vincent assisted with observations, collection, and rearing. VOLUME LXXXVII, NUMBER 2 125 Literature Cited Bateman, M. A. 1972. The ecology of fruit flies. Annu. Rev. Entomol. 17:493-518. Berube, D. E. 1978. Larval descriptions and biology of Tephritis dilacerata (Dip.; Tephriti- dae), a candidate for the biological control of Sonchus arvensis in Canada. Entomophaga 23:69-82. Christenson, L. D. and R. H. Foote. 1960. Biology of fruit flies. Annu. Rev. Entomol. 5:171- 192. Danielson, L. L., W. B. Ennis, Jr., D. L. Klingman, W. G. Shaw, F. L. Timmons, J. E. Jernigan, J. R. Paulling and P. E. Strickler. 1965. A survey of extent and cost of weed control and specific weed problems. USDA Crops Res. Bull. ARS 34-23-1. 78 pp. Dickerson, C. T., Jr. and R. D. Sweet. 1971. Common ragweed ecotypes. Weed Sci. 19: 64-66. Foote, B. A. 1965. Biology and immature stages of eastern ragweed flies (Tephritidae). Proc. North Cent. Branch Entomol. Soc. Am. 20:105-106. I Goeden, R. D., O. V. Kovalev and D. W. Ricker. 1974. Arthropods exported from California to the USSR for ragweed control. Weed Sci. 22:156-168. Piper, G. L. 1976. Bionomics of Euarestoides acutangulus (Diptera: Tephritidae). Ann. Ento- ! mol. Soc. Am. 69:381-386. I Shutova, N. N., ed. 1970. A handbook of pests, diseases and weeds of quarantine significance. 1 Kolos Publ., Moscow: 240 pp. Spieth, H. T. 1974. Courtship behavior in Drosophila. Annu. Rev. Entomol. 19:385—405. I Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior of Tep/zr/r/i 5//gma//'ca i (Coquillett). Pan Pac. Entomol. 41:73-79. ! Varley, G. C. 1946. The natural control of population balance in the knapweed gall-fly (Uro- phora jaceana). J. Anim. Ecol. 15:140-187. I Wangberg, J. K. 1978. Biology of gall-formers of the genus Va/enr/7jz///a (Diptera: Tephritidae) I on rabbitbrush in Idaho. J. Kansas Entomol. Soc. 51:472-483. Wasbauer, M. S. 1972. An annotated host catalog of the fruit flies of America north of Mexico. I (Diptera; Tephritidae). Occas. Papers Bur. Entomol. Calif. Dept. Agric. 19:1-172. j Beneficial Insect Introduction Laboratory, Insect Identification and Ben- i eficial Insect Introduction Institute, Agricultural Research, SEA, USDA, I Beltsville, MD 20705. Received for publication April 6, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 126-127 ACOUSTIC ATTRACTION OF HERONS BY CRICKETS Paul D. Bell Abstract. — Male crickets of the species Anurogryllus celerinictus attract mates after sunset by giving a loud sustained calling song. A predator, the heron, Florida coerulea uses this sound to locate its prey. F. coerulea was also attracted to a tape recorded song of A. celerinictus. It was first suggested by Lutz (1924) that predators such as frogs, lizards and birds likely hear and respond to insect sounds. Walker (1964) experi- mentally demonstrated that domestic cats can acoustically locate singing orthopteran prey. Cade (1975) and Soper et al. (1976) reported on dipteran parasites orienting to the sound of their insect prey. Bird predators should be capable of hearing songs of certain Grylloidae (Marler, 1955). Yet, no one has shown that birds locate insect prey by using the prey’s sound(s). In May, 1978, at Ocho Rios, Jamaica I observed an adult little blue heron, {Florida coerulea) to orient to a tape-recorded calling song of the fast-calling, short-tailed cricket, Anurogryllus celerinictus (Walker). The heron(s) was observed to stalk, catch and eat crickets, at night between the hours of 20:00 and 22:00 EOT. This time period coincides with the crickets’ peak acoustical activity (Bell, In Press). Many of the villagers were familiar with these, “dark, cricket-eating sea gulls.’’ During daylight hours the same species of heron fished shallow, slow-moving freshwater streams. Two Philips tweeters were positioned 5 M apart, magnets down on a flat lawn. Over one speaker, using a Uher 4000 Report L, the recorded calling song of a male A. celerinictus was played for 20 min. periods. The second speaker was silent. On one occasion an adult F. coerulea was approximately 30 M from the speakers when a recording was begun. Immediately the heron turned toward the speakers. It approached the live speaker, stalking slowly and turning its head from side to side. The heron came within 1 m of the live speaker, and was never closer than 5 m from the silent speaker. After seeing myself and the recorder the heron turned away from both speakers, walked a few steps (1 m), and caught and ate a cricket. Male A. celerinictus call to attract females for the purposes of mating. Being nocturnal singers, and associated with burrows, these crickets escape most daytime predators. However, herons appear to locate these crickets in the dark by orienting to the cricket’s song. VOLUME LXXXVII, NUMBER 2 127 Acknowledgments I thank Maureen E. Bell for assistance in the field. This study was funded in part by the Zoology Department and Erindale College, University of Toronto, by Dr. Glenn K. Morris through the National Research Coun- cil grant #4946, and an Ontario Graduate Scholarship to PB. Literature Cited Bell, P. D. 1979. The thermoregulatory burrow of the fast-calling, short-tailed cricket, Anurogryllus celerinictus (Walker) (Orthoptera: Gryllidae). Ann. Entomol. Soc. Am. (In press). Cade, W. 1975. Acoustically orienting parasitoids: fly phonotaxis to cricket song. Science 190:1312-1313. Lutz, E. E. 1924. Insect sounds. Bull. Amer. Mus. Nat. Hist. 50:333-372. Marler, P. 1955. Characteristics of some animal calls. Nature 176(4470): 6-8. Soper, R. E., G. E. Shewell and D. Tyrrell. 1976. Colcondamyia auditrix nov. sp. (Diptera: Sarcophagidae), a parasite which is attracted by the mating song of its host, Okanagana rimosa (Homoptera: Cicadidae). Can. Ent. 108:61-68. Walker, T. J. 1964. Experimental demonstration of a cat locating orthopteran prey by the prey’s calling song. Ela. Entomol. 47:163-165. Department of Zoology, Erindale College, University of Toronto, Missis- sauga, Ontario L5L 1C6, Canada. Received for publication October 18, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK2), 1979, pp. 128-134 COMMUNITY ECOLOGY AND P/E/?/5— CRUCIFER COEVOLUTION‘-2 F. S. Chew The association of certain subfamilies of Lepidoptera with certain families of flowering plants whose members are exploited as larval food is well j known (e.g. Ehrlich and Raven, 1965; Fraenkel, 1959; Whittaker and Feeny, 1971). Among the Pierinae, for example, larvae of the genus Pieris are j confined to the Cruciferae, Capparidaceae, and a small number of other plant families whose members contain glucosinolates (mustard oil gluco- sides; Ettlinger and Kjaer, 1968). A number of glucosinolates, tested under laboratory conditions, appear to mediate specificity in these plant-insect associations by eliciting feeding and oviposition responses from various Pieris (e.g. David and Gardiner, 1966; Hovanitz and Chang, 1963) as well as from other crucivorous species (e.g. Thorsteinson, 1953; Tanton, 1965; Hicks, 1974). In addition, allylisothiocyanate, the volatile hydrolysis prod- uct resulting from enzymatic degradation of allylglucosinolate (sinigrin) when crucifer tissues are damaged, acts as an attractant to Pieris rapae and ' some chrysomelid beetles (Blau et al., 1978; Feeny et al., 1970; Matsumoto, j 1970). Specific electrophysiological responses of Pieris brassicae to indi- vidual glucosinolates (Schoonhoven, 1967) and laboratory behavioral tests : of insect ability to discriminate among glucosinolates (e.g. Hovanitz and : Chang, 1963; Hicks, 1974) suggest that Pieris and other crucifer specialists • may distinguish among various Cruciferae (and other glucosinolate-bearing taxa) on the basis of this chemical class of more than 70 compounds (Et- tlinger and Kjaer, 1968; Rodman, 1978). In this paper I present evidence j that Pieris preferences among several co-occurring crucifer species in a . community are correlated with the presence of specific glucosinolates in | these species. These results suggest that by affecting the intensity of Pieris !j herbivory on their allelochemically-similar neighbors, individual crucifer ( species may influence each other’s evolution. | How do Pieris choose foodplants among allelochemically-similar taxa in 1 a community? Chew (1974, 1975, 1977) characterized the behavioral and | growth responses of a population of Pieris napi macdunnoughii to Crucif- i ‘ Based on paper contributed to symposium “Components of Host-Plant Utilization in Lep- S idoptera” (P. Barbosa, organizer) presented at Eastern Branch Meetings, Entomological So- f ^ ciety of America, 27 September 1978. S t ^ Research supported by grants from the American Philosophical Society and the Marshall I Fund (Tufts University). VOLUME LXXXVII, NUMBER 2 129 erae in its montane habitat (Table 1). P. n. macdunnoughii responses to the native crucifer taxa in this community comprise three categories: a) a pre- ferred foodplant, Descurainia richardsonii (nomenclature follows Weber, 1976) which supports significantly faster larval development to pupation than other native foodplants; b) a group of foodplants which support similar larval growth rates and among which adults and larvae show no consistent preferences, viz., Arabis drummondii, Cardamine cordifolia, Draba aurea, and Thlaspi montanunv, and c) a crucifer which is unconditionally rejected by both ovipositing adults and feeding larvae — Erysimum asperum. The behavioral responses of P. n. macdunnoughii adults and larvae to these crucifers are thus consistent with each other: larvae are able to complete larval development on all crucifer species chosen by ovipositing females. In contrast, behavioral and growth responses of P. n. macdunnoughii to the naturalized weed Thlaspi arvense are inconsistent with each other: females accept T. arvense as an oviposition substrate and larvae feed on the plant, but die before completing development. Similar incongruity between female oviposition behavior and larval growth responses has been observed in a variety of Lepidoptera exposed to previously unencountered relatives of their foodplants (e.g. Straatman, 1962; Sevastopulo, 1964; Bowden, 1971). This inconsistency also characterizes the behavioral and growth responses of Pieris occidentalis, another native Colorado species, toward T. arvense. P. occidentalis places significantly fewer eggs on T. arvense than on native crucifers (Chew, 1977); however, unlike P. n. macdunnoughii avoidance of Erysimum asperum, P. occidentalis avoidance of T. arvense is statistical rather than absolute, with the result that substantial numbers of Pieris eggs are placed on this lethal plant. Since T. arvense is naturalized to this region, perhaps insufficient time has elapsed for the resolution of this incongruity. Botanical surveys of this region (see Chew, 1977) suggest that T. arvense has been abundant in this community for less than 100 years. Given the present pattern of behavior and mortality on T. arvense, P. n. macdun- noughii and P. occidentalis should eventually reject T. an’ense plants as oviposition sites; alternatively, because females currently oviposit on T. arvense and larvae feed on the plant before dying (Chew, 1975), selection may favor any change which permits larvae to develop successfully on this plant. Examination of the leaf glucosinolate profiles of these crucifer species shows that this community contains three distinct glucosinolate arrays (Ta- ble 2). The group of four species (in which P. n. macdunnoughii exhibits no consistent preference) is characterized by isopropyl glucosinolate and several biosynthetically related compounds. The unconditionally rejected plant. Erysimum asperum, contains methylsulfinylalkyl and methylsulfon- ylalkyl compounds, which are distinct from other major components of oth- er glucosinolate profiles. Since some plants in this genus and the related 130 NEW YORK ENTOMOLOGICAL SOCIETY Table 1 . Growth and behavioral responses of Pieris napi macdunnoughii to crucifer species growing in a montane Colorado community. These species include all locally abundant taxa which contain glucosinolates. Data are summarized from Chew (1974, 1975, 1977). Crucifer species Oviposition Feeding Growth to pupation (a) Descurainia richardsonii Prefer Prefer Easiest (b) Arabis drummondii Yes Yes Yes Cardamine cordifolia Yes Yes Yes Draba aurea Yes Yes Yes Thlaspi montanum Yes Yes Yes (c) Erysimum asperum No No No Thlaspi arvense' Yes Yes No ' Naturalized from Eurasia. genus Cheiranthus contain cardenolides (Hegnauer, 1964) Erysimum as- perum may be chemically distinctive in this community on that basis as well; however, this species has not been examined for cardenolide content. The third array (with allylglucosinolate as a major component) includes both the preferred foodplant Descurainia richardsonii and the naturalized lethal crucifer Thlaspi arvense. Unlike the first two glucosinolate arrays, which m correspond directly to crucifers in two behavioral-growth response cate- in gories, this third array comprises two crucifers whose effects on these Pieris ali are widely divergent. The correlation of glucosinolate profiles with Pieris larval and adult be- fli havior suggests that P. n. macdunnoughii distinguish among these crucifers co at least partly on the basis of their glucosinolate profiles. In the case of mi Erysimum asperum, lack of information on the possible cardenolide content gli of this species precludes determination of what allelochemic class(es) me- po diates rejection of this plant by Pieris (Nielsen, 1978). While the possible m physiological effects of individual glucosinolates on adapted crucivores such Tti as Pieris have only begun to be explored (e.g. Marsh and Rothschild, 1974; pa Aplin et al., 1975; Blau et al., 1978), Pieris behavior towards T. arvense cai suggests that allylglucosinolate is perceived as a signal of suitable larval sic food. The allyl compound, and other major components of these glucosi- gla nolate profiles, are probably not intrinsic indicators of foodplant suitability tio for larval Pieris. Rather, Pieris behavior toward specific individual gluco- gei sinolates shared by several taxa probably depends on the evolutionary as- 19] sociation of those glucosinolates with food resources of a particular quality. . or For these Colorado Pieris, it is likely that the consistent association of | liyi allylglucosinolate with the foodplant Descurainia richardsonii has been dis- i bui VOLUME LXXXVII, NUMBER 2 131 Table 2. Results of qualitative analysis of crucifer leaves for glucosinolates. Paper and gas- chromatographic methods are outlined by Rodman (1974). Data are summarized from unpub- lished data of Rodman and Chew. Crucifer species Major glucosinolates (aglycone moiety) Minor glucosino- lates (number only) I. Arabis drummondii' Isopropyl-, 2-hydroxyisoproyl-, 2-butyl- 5 Cardamine cordifolia Isoproyl-, 2-hydroxyisopropyl-, 2-butyl-, l-ethyl-2-hydroxyethyl- 1 Draba aurea Isopropyl- 2 Thlaspi montanum Isopropyl-, 2-butyl-, (p)-rhamnopyranosyloxy benzyl- 1 II. Erysimum asperum 3-methylsulfinylpropyl-, 4-methylsulfonylbutyl 0 III. Descurainia richardsonii Allyl-, 3-butenyl- 5 Thlaspi arvense Allyl- 2 ' Roman numerals refer to crucifer arrays described in the text. rupted by the relatively recent introduction of Thlaspi arvense to this com- munity; ovipositing adults may be confusing the lethal T. arvense with its allelochemically-similar associate D. richardsonii. These data suggest that variation among members of a single chemical class may be significant in modifying insect behavior toward plants which contain this chemical class (cf. Dethier, 1978). For glucosinolates, variation may occur in two ways. First, biosynthesis of the aglycone moiety of the glucosinolate may vary, giving rise, for example to the hydroxylated com- pounds found in Arabis drummondii and Cardamine cordifolia\ variation in the number of methylene groups occurs in Descurainia richardsonii. These changes probably involve minor modifications of existing biosynthetic pathways (e.g. Chew and Rodman, 1979 and references therein). In this case it is probably significant that electrophysiological work on Pieris bras- sicae shows that the receptors of this crucivore are more sensitive to the glucosinolates tested than to their corresponding isothiocyanates (Schoon- hoven, 1967). Alternatively, after biosynthesis, a single glucosinolate may generate different aglycone products upon enzymatic hydrolysis (Benn, 1977). Allylglucosinolate, for example, may give rise to allylisothiocyanate or its isomer allylthiocyanate; allylthiocyanate has been found as the major hydrolysis product of T. arvense by others (e.g. Gmelin and Virtanen, 1960) but preliminary tests of T. arvense from this montane community reveal 132 NEW YORK ENTOMOLOGICAL SOCIETY allylisothiocyanate as the major product. Feeny and colleagues (Feeny et al., 1970; Feeny, 1977) have shown that allylthiocyanate is much less at- tractive to crucivorous chrysomelid beetles under field conditions than the corresponding isothiocyanate. While the variation generated within each crucifer species will be largely constrained by existing biosynthetic pathways (cf. Atsatt and O’Dowd, 1976), the adaptive value of a particular glucosinolate variant in a particular crucifer species depends on its relation to glucosinolates already present in the community (Jones, 1968; Dolinger et al., 1973; Janzen, 1973; Atsatt and O’Dowd, 1976; Cates and Rhoades, 1977; Feeny, 1977, Moore, 1978b). Pier- is response to a novel compound in the glucosinolate profile of a particular crucifer will thus depend a) on whether it mimics the glucosinolate profile already produced by some other community associate (cf. Macior, 1970); b) on Pieris growth responses to species containing that glucosinolate; and c) on whether Pieris distinguish the plant containing the new glucosinolate from chemically-similar taxa in the community. To the extent that glucosi- nolates mediate the responses of Pieris towards their foodplants and to the extent that Pieris activity imposes differential mortality on different chem- ical morphs within crucifer populations (Jones, 1971; cf. Moore, 1978a; Morrow and LaMarche, 1978), it seems likely that these crucifer species evolve in relation to each other as well as in relation to the crucivorous Pieris in this community. Literature Cited Aplin, R. T., R. d’Arcy Ward and M. Rothschild. 1975. Examination of the large and small white butterflies {Pieris spp.) for the presence of mustard oils and mustard oil glucosides. J. Ent. (A) 50:73-78. Atsatt, P. R. and D. J. O’Dowd. 1976. Plant defense guilds. Science 193:24-29. Benn, M. 1977. Glucosinolates. Pure Appl. Chem. 49:197-210. Blau, P. A., P. Feeny, L. Contardo and D. S. Robson. 1978. Allylglucosinolate and herbiv- orous caterpillars: a contrast in toxicity and tolerance. Science 200:1296-1298. Bowden, S. R. 1971. American white butterflies (Pieridae) and English foodplants. J. Lepid. Soc. 25:6-12. Cates, R. G. and D. F. Rhoades. 1977. Patterns in the production of antiherbivore chemical defenses in plant communities. Biochem. Syst. Ecol. 5:185-193. Chew, F. S. 1974. Strategies of foodplant exploitation in a complex of oligophagous butterflies (Lepidoptera). Ph.D. dissertation. Yale University, New Haven. . 1975. Coevolution of pierid butterflies and their cruciferous foodplants. I. The relative quality of available resources. Oecologia 20:117-127. . 1977. Coevolution of pierid butterflies and their cruciferous foodplants. II. The dis- tribution of eggs on potential foodplants. Evolution 31:568-579. and J. E. Rodman. 1979. Plant resources for chemical defense, pp. 271-307 In Her- bivores: their interaction with secondary plant metabolites (ed. G. A. Rosenthal and D. H. Janzen). Academic Press, New York. VOLUME LXXXVII, NUMBER 2 133 David, W. A. L. and B. O. Gardiner. 1966. Mustard oil glucosides as feeding stimulants for Pieris brassicae in a semi-synthetic diet. Entomol. Exp. Appl. 9:247-255. Dethier, V. G. 1978. Other tastes, other worlds. Science 201:224-228. Dolinger, P. M., P. R. Ehrlich, W. L. Fitch and D. E. Breedlove. 1973. Alkaloid and predation patterns in Colorado lupine populations. Oecologia 13:191-204. Ehrlich, P. R. and P. H. Raven. 1965. Butterflies and plants: a study in coevolution. Evolution 18:586-608. Ettlinger, M., and A. Kjaer. 1968. Sulfur compounds in plants. Rec. Adv. Phytochem. 1:60- 144. Feeny, P. 1977. Defensive ecology of the Cruciferae. Ann. Mo. Bot. Gard. 64:221-234. , K. L. Paauwe and N. J. Demong. 1970. Flea beetles and mustard oils: hostplant specificity of Phyllotreta cruciferae and P. striolata adults (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Amer. 63:832-841. Fraenkel, G. 1959. The raison d’etre of secondary plant substances. Science 129:1466-1470. Gmelin, V. R. and A. 1. Virtanen. 1960. The enzymic formation of thiocyanate (SCN ) from a precursor! s) in Brassica species. Acta Chem. Scand. 14:507-510. Hegnauer, R. 1964. Chemotaxonomie der Pflanzen. Vol. 3: Dicotyledoneae: Acanthaceae- Cyrillaceae. Birkhauser, Basel. Hicks, K. L. 1974. Mustard oil glucosides: feeding stimulants for adult cabbage flea beetles, Phyllotreta cruciferae (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Amer. 67:261- 264. Hovanitz, W. and V. C. S. Chang. 1963. Comparison of the selective effect of two mustard oils and their glucosides to Pieris larvae. J. Res. Lepid. 2:281-288. Janzen, D. H. 1973. Community structure of secondary compounds in plants. Pure Appl. Chem. 34:529-538. Jones, D. A. 1968. On the polymorphism of cyanogenesis in Lotus corniculatus L. II. The interaction with Trifolium repens L. Heredity 23:453—455. . 1971. Chemical defense mechanisms and genetic polymorphism. Science 173:945. Macior, L. W. 1970. The pollination ecology of Pedicularis in Colorado. Amer. J. Bot. 57:716- 728. Marsh, N. and M. Rothschild. 1974. Aposematic and cryptic Lepidoptera tested on the mouse. J. Zool. 174:89-122. Matsumoto, Y. 1970. Volatile organic sulfur compounds as insect attractants with special reference to host selection, pp. 133-160 /« Control of insect behavior by natural prod- ucts (ed. D. L. Wood, R. M. Silverstein, N. Nakajima). Academic Press, New York. Moore, L. R. 1978a. Seed predation in the legume Crotolaria. I. Intensity and variability of seed predation in native and introduced populations of C. pallida Ait. Oecologia 34: 185- 202. . 1978b. Seed predation in the legume Crotolaria. II. Correlates of interplant variability in predation intensity. Oecologia 34:203-223. Morrow, P. A. and V. C. LaMarche, Jr. 1978. Tree ring evidence for chronic insect suppres- sion of productivity in subalpine Eucalyptus. Science 201:1244-1246. Nielsen, J. K. 1978. Host plant discrimination within the Cruciferae: feeding responses of four leaf beetles (Coleoptera: Chrysomelidae) to glucosinolates, cucurbitacins and car- denolides. Entomol. Exp. Appl. 24:41-54. Rodman, J. E. 1974. Systematics and evolution of the genus Cakile (Cruciferae). Contrib. Gray Herb. Harvard Univ. 205:3-146. . 1978. Glucosinolates; methods of analysis and some chemosystematic problems. Phy- tochem. Bull. 11:6-31. Schoonhoven, L. M. 1967. Chemoreception of mustard oil glucosides in larvae of Pieris brassicae. Proc. Roy. Acad. Sci., Amsterdam. 70C:556-568. 134 NEW YORK ENTOMOLOGICAL SOCIETY Sevastopulo, D. G. 1964. Lepidoptera ovipositing on plants toxic to their larvae. J. Lepid. Soc. 18:104. Straatman, R. 1962. Notes on certain Lepidoptera ovipositing on plants which are toxic to their larvae. J. Lepid. Soc. 16:99-103. Tanton, M. T. 1965. Agar and chemostimulant concentrations and their effect on intake of synthetic food by larvae of the mustard beetle, Phaedon cochleariae Eab. Entomol. Exp. Appl. 8:74-82. Thorsteinson, A. J. 1953. The chemotactic responses that determine host specificity in an oligophagous insect (Plutella maculipennis (Curt.) Lepidoptera). Can. J. Zool. 31:52- 72. Weber, W. 1976. Rocky Mountain Elora. Colorado Assoc. Univ. Press, Boulder. Whittaker, R. H. and P. P. Eeeny. 1971. Allelochemics: chemical interactions between species. Science 171:757-70. Department of Biology, Tufts University, Medford, Massachusetts 02155. Received for publication December 14, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 135-140 INTERACTIONS BETWEEN BLOW FLIES (CALLIPHORIDAE) AND ENTOMOPHTHORA BULLATA (PHYCOMYCETES: ENTOMOPHTHORALES) John Paul Kramer Abstract. — Studies of interactions between blow flies and Entomophthora bullata showed that: (1) the time between exposure to conidia and death of the fly ranged from 5 to 12 days with about 50% dying by day 6; (2) the fungus produced only conidia in 79% of the cadavers, resting spores in 18%, and a mixture of the two in 3% of them; (3) in flies Vi to 3 days old at exposure only conidia were formed in >80% of the cadavers, while in flies 4 to 5 days old at exposure only resting spores were formed in about 50% of the cadavers; (4) the death rates of flies were not influenced by their ages at exposure; and (5) the time required for the disease to kill flies was not related to the types of spores formed on or in their cadavers. Host species included Phormia regina, Phaenicia sericata, and Protophormia terraeno- vae. Introduction On August 10, 1973, I found two fresh cadavers of black blow flies, Phor- mia regina, that had apparently succumbed to a mycosis in a cage of mis- cellaneous muscoids taken alive two days earlier from a trap set in a wooded area near Ithaca, New York. I tied both cadavers to the wet wick of a water bottle which rested on a layer of moist sand within a glass battery jar. The jar, covered with clear polyethylene wrap to maintain a relative humidity close to saturation, was held overnight in a dark and cold room (14-16°C). By the following morning a reddish tan mycelial mat had grown over one cadaver; only the dorsum of its thorax remained uncovered. A thick layer of conidia discharged from this cadaver covered the aluminum foil collar of the water bottle and portions of the wick as well. The body surface of the second cadaver exhibited no growths and this specimen was removed from the wick for further study. A sample of the conidial shower produced by the first cadaver was also removed from the wick for microscopical exam- ination. The body cavity of the second cadaver was filled with spherical and hya- line bullate resting spores that ranged from 38 to 58 pm in diameter. The discharged conidia from the first cadaver were hyaline and elliptical with an evenly rounded apex and a papillate base. These conidia ranged in size from 25-37 X 11-17 ^m. In conformation and size these resting spores and con- 136 NEW YORK ENTOMOLOGICAL SOCIETY idia compare very favorably with those described by MacLeod et al. (1973) for Entomophthora bullata. I therefore concluded that the fungus found in these P. regina was the same species. The present study was done: (1) to develop baseline data on the in vivo culture of E. bullata and its effects on blow flies, and (2) to determine if an association exists between the age of flies at exposure to conidia and the time of their death due to E. bullata infections and the types of spores produced in their cadavers. Materials and Methods The P. regina cadaver that produced the conidial shower, noted in the first section of this paper, served as the source of inoculum for the infectivity tests. Thirty young, laboratory-reared P. regina adults were added to the battery jar containing the water bottle with this cadaver tied to its wick. As noted previously, the collar of the water bottle and portions of the wick were covered with a thick layer of conidia. The jar containing this contam- inated bottle and the healthy flies was returned to the dark and cold room. On the following morning the jar was placed on a laboratory bench at room temperatures (21-27°C). Here the flies received some artificial light from overhead fluorescent lamps during each day for the duration of the experi- ment. A small dish of dry sugar-milk was added to the jar as a source of nourishment for the flies. Several small holes were punched in the polyeth- ylene cover for ventilation. The flies were observed at daily intervals and dead ones were removed from the jar and examined for signs of E. bullata infections. The data pertaining to this specific test are presented in Table 1 (see Test 1). The same procedures were used in all of the subsequent infec- tivity tests. All of the P. regina, Phaenicia sericata, and Protophormia terraenovae tested came from clean laboratory cultures. The results given in Tables 2 through 4 are pooled data from 49 separate tests. The number of flies used in any single test was variable and depended upon the number of flies available at the time. While records on the ages of flies were maintained, records on the sex ratios within the test groups were not. Flies that died without E. bullata infections and flies that lived for more than 12 days following exposure to E. bullata conidia have been omitted from the data presented in Tables 2 through 4. No flies living for more than 12 days died with E. bullata infections. Results and Interpretations The data in Table 1 show that 27 serial passages of E. bullata were achieved from August 1973 to March 1974. The failure to effect transmission for additional passages was linked to a bacterial septicemia found in most VOLUME LXXXVIl, NUMBER 2 137 Table 1. Fly-to-fly passages of Entomophthora bullata in adult blow flies during an eight- month period under laboratory conditions. Test number Date test started Flies tested No. cadavers as inoculum sources* Percentage dying with E. bullata** Species Age (days) 1 13 Aug. P. regina 111 1 79 (23/30) 2 20 Aug. P. regina 4 2 40 (6/15) 3 28 Aug. P. regina 1/2 2 22 (4/18) 4 5 Sep. P. regina 5 1 1 1 (3/27) 5 13 Sep. P. regina 1/2 1 44 (10/23) 6 20 Sep. P. regina 1/2 1 31 (6/18) 7 26 Sep. P. sericata 1/2 1 53 (8/15) 8 4 Oct. P. regina 1/2 2 100(17/17) 9 10 Oct. P. terraenovae 1/2 2 100 (10/10) 10 17 Oct. P. sericata 5 1 86 (12/14) 11 26 Oct. P. regina 1/2 1 59 (17/29) 12 5 Nov. P. sericata 1/2 1 74 (17/23) 13 12 Nov. P. regina 1/2 3 92 (12/13) 14 21 Nov. P. sericata 1/2 1 79 (11/14) 15 28 Nov. P. regina 1/2 3 82 (28/34) 16 5 Dec. P. sericata 1/2 1 58 (11/19) 17 13 Dec. P. regina 1/2 3 83 (15/18) 18 26 Dec. P. sericata 1/2 1 100 (10/10) 19 2 Jan. P. regina 1 1 67 (8/12) 20 14 Jan. P. regina 1 1 29 (2/7) 21 25 Jan. P. regina 1 1 44 (4/9) 22 4 Feb. P. sericata 1 1 54 (15/28) 23 12 Feb. P. regina 1 5 40 (15/28) 24 21 Feb. P. regina 3 3 67 (4/6) 25 6 Mar. P. sericata 1 1 37 (7/19) 26 19 Mar. P. sericata 1 1 6 (1/18) 27 27 Mar. P. regina 1 1 50 (5/10) 28 4 Apr. P. sericata 2 3 0 (0/22) * Source for Test 1 was a field-collected P. regina; sources for subsequent tests were conidia-producing cadavers from the previous trial. ** By post-exposure day 12. of the E. bullat a -miecicd flies that died in Tests 23 through 27 (see Table 1) and this severely depressed the development of the fungus. Hence the numbers of infectious units to which the flies were exposed declined during these last five tests. The data in Table 1 show that the percentage of flies dying with E. bullata infections ranged from 6 to 100%, yet there is no obvious relationship between the numbers of cadavers used as inoculum sources and the percentages of flies that acquired the infection. In nearly all cases the cadavers were attached to the substrate by rhizoids. All three species of blow flies were susceptible to infection. 138 NEW YORK ENTOMOLOGICAL SOCIETY Table 2. Age of blow flies at time of exposure to Entomophthora bullata conidia and time of death due to infection. Age in days (No.) Post-exposure days and percentage dying during each period 1-4 5 6 7 8 9 10 u 12 to 1 P. regina (454) 0 14 45 15 14 6 4 0.5 1.5 P. sericata (225) 0 19 40 28 7 4 1.5 0.5 0 Both spp. (679) 0 16 43 19 11 5 4 0.5 1.5 2 P. regina 0 0 0 0 0 0 0 0 0 0 P. sericata (16) 0 0 0 56 0 13 25 0 6 Both spp. (16) 0 0 0 56 0 13 25 0 6 3 P. regina (46) 0 0 23 43 24 4 2 2 2 P. sericata (14) 0 57 43 0 0 0 0 0 0 Both spp. (60) 0 13 27 33 18 3 2 2 2 4 P. regina (18) 0 6 11 33 28 0 11 11 0 P. sericata (34) 0 12 34 29 17 3 6 0 0 Both spp. (52) 0 10 25 31 21 2 8 3 0 5 P. regina (18) 0 0 83 17 0 0 0 0 0 P. sericata (14) 0 0 86 0 0 0 7 7 0 Both spp. (32) 0 0 84 8 0 0 4 4 0 All ages in both species (839) 0 14 42 21 12 5 4 1 1 Flies died of E. bullata infections as early as day 5 and as late as day 12 following exposure to conidia with about 50% dying by day 6 and 75% by day 7 (see Table 2). The data presented also suggest that susceptibility to infection among flies Vi to 5 days old is about the same in all of these age classes, and in both species tested. The data on age classes presented in this table, however, do not suggest that age of the fly at exposure alters the time required for the disease to kill the fly, i.e. older flies die at about the same rate as younger ones. The data in Table 3 show that the fungus produced conidia in 79% of the cadavers, resting spores in 18%, and a mixture of conidia and resting spores in 3% of them. In this last group very light conidial showers were produced from scanty mycelial mats that grew on the surfaces of abdomens that har- bored some resting spores internally. The data in Table 3 suggest that age of the fly at time of exposure does influence the types of spores produced by the fungus in the cadavers; in more than 80% of the flies V2 to 3 days old at exposure only conidia were formed, while in about 50% of the flies 4 to 5 days old only resting spores were formed. This observation supports the suggestion of Wilding and Lauckner (1974) that resting spore formation oc- curs more frequently in older wheat bulb flies, Leptophylemyia coarctata, infected with E. muscae. VOLUME LXXXVIl, NUMBER 2 139 Table 3. Age of blow flies at time of exposure to Entomophthora bullata conidia and types of spores produced in their cadavers. Spore types with percentages of each Age Resting Conidia + in days (No.) Conidia spores resting spores 14 to 1 P. regina (454) 81 15 4 P. sericata (225) 88 9 3 Both spp. (679) 84 13 3 2 P. regina 0 0 0 0 P. sericata (16) 94 6 0 Both spp. (16) 94 6 0 3 P. regina (46) 86 7 7 P. sericata (14) 29 71 0 Both spp. (60) 73 22 5 4 P. regina (18) 22 78 0 P. sericata (34) 68 32 0 Both spp. (52) 48 52 0 5 P. regina (18) 56 44 0 P. sericata (14) 21 79 0 Both spp. (32) 41 59 0 All ages in both species (839) 79 18 3 Table 4. Time of death of Entomophth produced in their cadavers. ora bullata-infected blow flies and types of spores Post-exposure days and percentages dying during each period Spore types (No.) 1-4 5 6 7 8 9 10 11 12 Conidia P. regina (423) 0 13 37 22 15 6 4 1 2 P. sericata (242) 0 16 44 30 6 1.5 1.5 0.5 0.5 Both spp. (665) 0 14 40 25 12 5 3 0.5 0.5 Resting spores P. regina (94) 0 2 63 6 16 9 2 0 2 P. sericata (56) 0 23 41 16 9 0 7 4 0 Both spp. (150) 0 10 55 10 13 6 5 0.5 0.5 Conidia + resting spores P. regina (19) 0 0 42 58 0 0 0 0 0 P. sericata (5) 0 0 100 0 0 0 0 0 0 Both spp. (24) 0 0 54 46 0 0 0 0 0 All spore types in both species (839) 0 14 42 21 12 5 4 1 1 140 NEW YORK ENTOMOLOGICAL SOCIETY Conidia or resting spores may be formed in flies dying from days 5 through 12 (see Table 4). This phenomenon was observed in both P. regina and P. sericata. The conidia-resting spores mixture, found only in flies Vz to 3 days old at exposure, occurred in specimens dying on days 6 and 7 only. The data in Table 4 do not suggest that the time required for the disease to kill the fly is related to the type of spores formed on or in their cadavers. Discussion Prior to the discovery of E. bullata in P. regina reported in the present study, this fungus has been found parasitizing several other species of blow flies and flesh flies in nature. Povah (1935) found it in blue bottle flies and MacLeod (1956) extended its host range to include Sarcophaga aldrichi and a Calliphora species. MacLeod et al. (1973) observed an outbreak of E. bullata infections in a field population of 5. aldrichi, and suggested that the life cycle of the fungus involved alternating generations of conidia and rest- ing spores. The results of the present study, however, clearly indicate that conidia or resting spores, and in some cases a mixture of the two, may be produced on or in the dead bodies of flies that acquired the infection by exposure to conidia only. The host range of E. bullata in nature probably includes many species of blow flies that share the same habitat, e.g. P. regina, P. sericata, and T*. terraenovae. Literature Cited MacLeod, D. M. 1956. Notes on the genus Empusa Cohn. Can. J. Bot. 34:16-26. MacLeod, D. M., D. Tyrrell, R. S. Soper and A. J. DeLyzer. 1973. Entomophthora bullata as a pathogen of Sarcophaga aldrichi. J. Invert. Pathol. 22:75-79. Povah, A. H. W. 1935. The fungi of Isle Royale, Lake Superior. Pap. Mich. Acad. Sci. Arts Lett. 20:113-156. Wilding, N. and F. B. Lauckner. 1974. Entomophthora infecting wheat bulb fly at Rothamsted, Hertfordshire, 1967-71. Ann. appl. Biol. 76:161-170. Department of Entomology, Cornell University, Ithaca, New York 14853. Received for publication November 20, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 141-153 POST-INGESTIVE UTILIZATION OF PLANT BIOMASS AND NITROGEN BY LEPIDOPTERA: LEGUME FEEDING BY THE SOUTHERN ARMYWORM J. Mark Scriber Abstract. — Polyphagous southern armyworms, Spodoptera eridania, are extremely efficient at assimilating ingested biomass and nitrogen in foliage of forage legumes. Variable efficiencies of processing the digested food (E.C.D.) were observed, however, and these differences reflect metabolic costs which may be related to biochemical factors mediating varietal differ- ences in host plant resistance to other kinds of insects. Armyworms are apparently able to compensate for low conversion efficiencies with in- creased consumption rates such that their relative growth rates (mg. gained/ mg. tissue/day) are virtually independent of digestive efficiency. In fact, larval growth rates of southern armyworms on these 15 legume varieties were among the greatest observed in a comparison with 22 other species of Lepidoptera in 140 similar feeding experiments. It appears that the combi- nation of high leaf water content (80-91%) and high nitrogen content (4.2- 5.5% dry) of legume foliage are key factors permitting such growth. Alle- lochemics such as coumarin or the various saponins known to be present in these legumes were ineffective “barriers” to larval feeding and growth of armyworms in no-choice situations. Introduction The relative post-ingestive roles of allelochemics and plant nutritional quality (e.g. leaf water content and leaf nitrogen content) in determining larval growth efficiencies have been analyzed for 22 species of Lepidoptera larvae feeding upon a variety of normal foodplant species (Scriber, 1978a). Foodplants used in these experiments ranged from trees (with leaf-water and nitrogen contents of 50% and 1%, respectively) to various groups of forb species with leaf-water typically in the range of 80% to 90% and nitro- gen content ranging from 1.5% to 6.0%. In addition to a range of nitrogen concentrations provided by mature forb leaves, the Apiaceae (= Umbelli- ferae) (Kingsbury, 1964; Hegnauer, 1971) and the Cruciferae (Kjaer, 1974; Slansky and Feeny, 1977; and Chew, 1975) were chosen because of the unique array of allelochemics encountered in the various species. In the study to be reported here, a variety of species of the Leguminoseae were selected because of their high nitrogen as well as high water content, and also for the assortment of insect resistant genotypes (cultivars) which were 142 NEW YORK ENTOMOLOGICAL SOCIETY Leaf Water Content (%) Fig. 1 . The relationship of mean leaf nitrogen content and mean leaf water content of plants used for feeding experiments with various Lepidoptera. In this study the 15 legume varieties are indicated (“L”) for comparison with 21 Cruciferae varieties (“C”), various Umbelliferae (starred), and additional vine, shrub, and tree species (solid circles) from experiments reported by Scriber (1978a). The six additional legumes are described by Scriber (1978b). available for analyzing the relative growth suppressive effects upon the po- lyphagous southern armyworm, Spodoptera eridania (Cram.), The possible importance of leaf water content in determining the larval growth of certain tree-feeding Lepidoptera has been described by Scriber (1977, 1979a). A comprehensive study of various swallowtail butterfly (Pa- pilionidae) and silkmoth (Bombycoidea) species has revealed a striking cor- relation between larval growth and growth form of the foodplants (Scriber and Feeny, 1979). Leaf water content appears to provide a general indica- tion of the larval performance to be expected from a variety of species of Lepidoptera of different sizes and different degrees of feeding specialization (Scriber 1978a). Just as the variance from the regression on leaf water of nitrogen content of mature leaves increased above 75% leaf water content (Fig. 1), so does the variance in larval performance (Scriber, 1978a; Scriber and Feeny, 1979). Larval performance (the consumption, assimilation, and VOLUME LXXXVII, NUMBER 2 143 conversion of plant biomass and nitrogen for tissue growth) of forb feeders is generally much better than tree leaf feeders, and larval growth rate (mg. gain/mg. dry tissue/day) of herb feeders is generally 2x-6x that of tree feeders. Although the patterns of biomass and nitrogen accumulation rates be- tween tree- and forb-feeding larvae are rather striking (Scriber and Feeny, 1979), we do not know the relative contributions of low nitrogen, high fiber, high tannin or other factors which could be involved in suppressing larval growth rates upon mature tree leaves. Furthermore, seasonal variations in nutritional quality of tree leaves are certainly of prime concern to lepidop- teran herbivores (see Feeny, 1970, 1976; Rhoades and Cates, 1976). Sea- sonal variation in nutritional quality of forbs is also a very important factor determining larval growth rates and also may be more important than qual- itative plant allelochemics in determining utilization efficiencies of the Um- belliferae by ‘adapted’ herbivores (Finke, 1977). Of course, nutritional qual- ity will be much less important when qualitative (behaviorally deterrent) allelochemics are encountered by “unadapted” herbivores (see Erickson and Feeny, 1974; Feeny, 1975; Blau et al., 1978). Any limiting effects of leaf water would presumably be less obvious for larvae feeding on plant tissues with 75-90% water content (e.g., forbs). It may thus be more likely that leaf nitrogen or allelochemic content would play more significant roles in determining the post-ingestive utilization of forb tissues for larval growth. Slansky and Feeny (1977) analyzed utilization of various Cruciferae species by larvae of an ‘adapted specialist,’ Pieris rapae L. They suggest that nitrogen is indeed in limiting supply (81-90% leaf water content) and that larvae increase their feeding rates on plants which are low in nitrogen content in order to maximize the nitrogen accu- mulation rate (N.A.R. = Nitrogen Consumption Rate x Nitrogen Utiliza- tion Efficiency). The consumption rate of nitrogen was apparently increased up to a point beyond which declining nitrogen utilization efficiency (N.U.E.) offset the gain. It was suggested that consumption rate on any particular crucifer species would be adjusted to the lowest value at which maximal N.A.R. could occur. Although at least 19 different glucosinolates were re- ported from leaves of these foodplants, there was no detectable correlation between larval growth rate and these patterns of allelochemics. Perhaps the importance of nitrogen in this study of Cruciferae-adapted larvae may have obscured any subtle allelochemic effects. The independent roles and synergistic effects of leaf-water content, leaf nitrogen and allelochemics on larval growth are very difficult to assess. The use of allelochemics in artificial diet studies (Reese and Beck 1976a, b, c) or plants which are polymorphic for allelochemics (Scriber, 1978b) allow some control of variables. Regulation of diet water content in artificial diets (Reese, 1977; Reese and Beck, 1978) or in natural foodplant leaves (Scriber, 144 NEW YORK ENTOMOLOGICAL SOCIETY 1977) permits additional assessment of the relative importance of water in relation to other aspects of the diet. Fertilization to increase plant nitrogen content generally results in increased leaf water content as well (see Slansky and Feeny, 1977), and may not permit one to distinguish between effects to be attributed to these two variables. It was the intent of the present study to attempt an assessment of the effects on larval growth of various plant characteristics of the Leguminosae family. In this study, the leaf- water content was relatively high (80-91%) and the plant nitrogen content was also generally high (4. 2-5. 5%). It was thus hoped that any differences in larval performance on 15 forage cultivars would not primarily be reflections of leaf-water and nitrogen limitations, but instead might reflect the more subtle post-ingestive effects of various bio- chemical factors implicated in host-plant resistance. Methods and Materials Fifteen Legume species selected and bred for agronomic qualities and insect resistance were fed to larvae of the southern armyworm (S. eridania). Growth performances on alfalfa varieties (Apollo, Arc, Culver, Kanza, MSA-CW3-AN3, Ranger, Riley, Team, Vernal, and Weevlcheck) were com- pared and contrasted with bird’s foot trefoil, Lotus corniculatus; red clovers (Arlington and Lakeland); white clover; and yellow blossom sweet clover in these bioassays. Glasshouse-grown seedlings about 12 inches in height were used. Consumption rates, assimilation efficiencies, conversion efficiencies and relative accumulation rates of biomass and nitrogen were determined for larvae via gravimetric methods (see Waldbauer, 1968; and Scriber 1977, 1978b for methods). These values were compared to those for other Lepi- doptera as reported in Scriber (1978a), and in the figures presented here, the regression lines are calculated based on the means for the previous 140 experiments and do not include the 15 legume experiments. Environmental conditions were vitually identical for all experiments re- ported (16:8 hr. photo: scotophase with a corresponding temperature regime of 23:19 C°). Included in these comparisons are experiments of Slansky and Feeny (1977) and Finke (1977) as well as those of Scriber (1975, 1978a, 1978b, 1979b) and Scriber and Feeny (1979). Results and Discussion As larval performance is surveyed over a range of foodplants from trees (with leaf-water contents of 50-75%) to forbs (with leaf-water contents of 75-95%), it becomes obvious that herb leaves are generally consumed more rapidly and converted to larval biomass more efficiently than tree leaves (Scriber 1978a). Legumes analyzed in this study support these general pat- VOLUME LXXXVII, NUMBER 2 145 LEAF WATER % i Eig. 2. Approximate digestibilities (Eig. 2a) and nitrogen utilization efficiencies (2b) of 22 species of lepidoptera in their penultimate instar as a function of leaf water content of their I foodplants (n = 155). Trees, shrubs and vines are indicated by a solid circle, forbs are indicated by stars and triangles, and legumes by open circles (cf. Scriber, 1978a). I I 146 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 3. Approximate digestibilities (Fig. 3a) and nitrogen utilization efficiencies (Fig. 3b) of final instar Lepidoptera as a function of leaf water of their foodplants (n = 170 experiments). The legume experiments are indicated by open circles and the Cruciferae by stars (cf. Scriber, 1978a). Solid symbols represent tree species. terns, but more striking was the efficiency with which larvae were able to assimilate legume biomass and nitrogen in relation to penultimate instar lepidoptera in 140 other feeding experiments. The biomass assimilation ef- ficiency or approximate digestibility (A.D.) of legumes by penultimate instar d larvae ranges from 76 to 94% (Fig. 2a), and nitrogen utilization efficiencies ti (N.U.E.’s) from 89 to 96% (Fig. 2b). This pattern was also observed for VOLUME LXXXVll, NUMBER 2 147 Fig. 4. The relative growth rates of penultimate instar Lepidoptera larvae as a function of the average leaf water content of the foodplant reared upon. The regression line was calculated previous to the inclusion of the 15 legume experiments (open circles). legume-fed larvae in the final instar. The A.D.’s ranged from 64-92% and N. U.K.’s from 76-95% (Figs. 3a, 3b). When performance data of final instar Pieris rapae on various Cruciferae (Slansky and Feeny, 1977) were compared to those of Umbellifer- and Le- gume-feeding lepidoptera, it appears that the Cruciferae may uniformly re- strict the assimilation of biomass (A.D.) and nitrogen (N.U.K.) by P. rapae larvae (Fig. 3a and 3b respectively). While low efficiency of assimilating nitrogen and biomass could be related to nutritional qualities of the Crucif- 148 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 5. The nitrogen accumulation rate (N.A.R.) of penultimate instar larvae as a function of leaf water content. Symbols represent the same plant types as in figure 2 and figure 4. erae, perhaps glucosinolates should also be considered. These allelochemics may thus have suppressive effects even upon Cruciferae-adapted specialists. While growth of P. rapae is not affected even by artificially high concen- trations of allylglucosinolate (Blau et al., 1978), it appears that the larvae are able to compensate for low efficiency of assimilating crucifer leaf tissues by increased consumption rates (Slansky and Feeny, 1977; cf. Scriber, 1978a). Umbellifer-feeding specialists (Papilio polyxenes and P. zelicaon) studied by Scriber and Feeny (1978) and Finke (1977) exhibited intermediate effi- ciencies in relation to P. rapae on Cruciferae and 5. eridania on Legumi- nosae (Fig. 3a and 3b). Specialized forb feeders such as these species of VOLUME LXXXVIl, NUMBER 2 149 I I I I I 1 2 3 4 5 6 LEAF NITROGEN CONTENT % Fig. 6. Relative growth rate of penultimate instar larvae of 22 species of Lepidoptera as a function of leaf nitrogen content of their foodplants. Tree, shrub and vine plants are indicated by solid circles, forbs by stars, and legumes by open circles (See also Scriber, 1978a). Umbellifer-feeding Lepidoptera grew at rates 2x-3x those of the 18 species of tree, shrub and vine feeders (Fig. 4). Larval growth rates (R.G.R., Fig. 4) and nitrogen accumulation rates (N.A.R., Fig. 5) for the polyphagous S. eridania were also 2x-3x greater than tree feeders as a group, suggesting that plant quality rather than degree of feeding specialization may be the more important influence. Compared to the 22 species of Lepidoptera (Scriber, 1978a), larval growth rate is very fast for these army worms on legumes. High nitrogen content (Fig. 6) as well as high water content (Fig. 4) is certainly a key contributing factor. While efficiencies of assimilating biomass and nitrogen are high for larvae on all of these forage legumes (Fig. 2 and 3), the efficiency of con- version of digested (assimilated) biomass (E.C.D.) was extremely variable. The low efficiencies of biomass conversion on Team (18.9%), Arc (24.1%) and Culver (29.8%) alfalfa varieties reflect greater metabolic costs for larvae than observed for other varieties (e.g. larvae fed Vernal and Apollo alfalfa had E.C.D.’s = 76.6% and 68.3% respectively, with 71.3% on yellow blos- som sweet clover). 150 NEW YORK ENTOMOLOGICAL SOCIETY Relative Consumption Rate (mg. mass/mg./da/) Fig. 7. The gross efficiency of growth (E.C.I. = A.D. x E.C.D.) as a function of the rel- ative consumption rate (R.C.R.) of plant biomass. Each symbol represents the experimental mean values obtained for penultimate instar S. eridania larvae on a particular legume variety, (r = .937; n = 15 for linear regression Y = 74.2 - 26. 6X). Why are certain genotypes more costly to process (in terms of assimilated energy and biomass) than others? Team, Arc and Culver alfalfa have been bred specifically for insect resistance to alfalfa weevil, aphids and meadow spittlebugs (Wilson and Davis, 1960; Barnes et al., 1970 and 1977; Sorensen et al., 1972), while Vernal and Apollo have been bred primarily for agro- nomic qualities and yield potential in Wisconsin (Rohweder et al., 1978). Concentrations of saponins, coumarin, cyanogenic glycosides and nitrates were not determined in this study. However, allelochemic variations be- tween varieties can influence resistance to many insects (Horber, 1972; 1974; Manglitz et al., 1976; Pederson et al., 1976), and might be responsible for some of the variation metabolic costs experienced by armyworm larvae in the present study. Induction of the mixed function oxidase detoxication activity in 5. eridania can be very rapid (Brattsten et al., 1977); however, it is premature to suggest that possession or operation of these enzyme systems is directly related to differential metabolic costs as measured here. Nonetheless, polyphagous southern armyworm, S. eridania, is apparently able to compensate for the low efficiencies of conversion (i.e., high metabolic costs) on certain legume varieties by increased rates of food consumption (Fig. 7). This allows rapid growth in spite of the need to process a variety of potential allelochemics in legume tissues. This ‘power and efficiency’ VOLUME LXXXVII, NUMBER 2 151 trade-off (Odum and Pinkerton, 1955; Fig. 7) has not been observed for tree- feeding Lepidoptera (Scriber, 1978a). It has been suggested that in these cases low leaf water content may be a fundamental efficiency limiting factor and that increased consumption rates would perhaps only aggravate the problem (Scriber, 1977). Conclusions While some insects are effectively repelled, deterred or suppressed by different biochemicals in the 15 legumes used in this study, the southern armyworm is able to feed and grow successfully. Significant differences in metabolic costs of processing and converting plant biomass were reflected by E.C.D.’s (efficiency of conversion of digested food), which ranged from less than 20% to more than 75% on certain varieties. Larvae were, however, able to “compensate” by increasing their rate of consumption where effi- ciencies were low. It is suggested that this compensatory mechanism of maintaining larval growth rates near their maximum would be successful primarily where leaf water is not a limiting factor. Acknowledgments Thanks are extended to D. K. Barnes for supplying seed of certain alfalfa genotypes used in this study, and to Mark Finke, who assisted during the critical periods of the feeding experiments. The armyworm culture was orig- inally obtained from Dr. C. F. Wilkinson at Cornell University. This re- search was supported by the College of Agricultural and Life Sciences and the Graduate School of the University of Wisconsin at Madison, and in part by Hatch Project 5134. I would also like to thank M. D. Finke, E. Grabstein and F. Slansky, Jr. for their comments. Literature Cited Barnes, D. K., C. H. Hanson, R. H. Ratcliffe, T. H. Busbice, J. A. Schillinger, G. R. Buss, W. V. Campbell, R. W. Hemken and C. C. Blickenstaff. 1970. The development and performance of Team Alfalfa: A multiple pest resistant alfalfa with moderate resistance to the alfalfa weevil. U.S.D.A. A.R.S. Publication 34-115. 41 pp. , E. T., Bingham, R. P. Murphy, O. J. Hunt, D. F. Beard, W. Y. Skrdla and L. R. Teuber. 1977. Alfalfa germplasm in the United States: Genetic vulnerability, use, im- provement, and maintenance. A.R.S. (U.S.D.A.) Technical Bull. 1561. 21 pp. Blau, P. A., P. Feeny, L. Contardo and D. S. Robson. 1978. Allylglucosinolate and herbiv- orous caterpillars: A contrast in toxicity and tolerance. Science 200:1296-1298. Brattsten, L. B., C. F. Wilkinson and T. Eisner. 1977. Herbivore-plant interactions: Mixed function oxidases and secondary plant substances. Science 196:1349-1352. Chew, F. 1975. Coevolution of pierid butterflies and their cruciferous foodplants. I. The quality of available resources. Oecologia 20:1 17-127. 152 NEW YORK ENTOMOLOGICAL SOCIETY Erickson, J. M. and P. P. Eeeny. 1974. Sinigrin: A chemical barrier to the black swallowtail butterfly. Ecology 55:103-111. Eeeny, P. P. 1970. Seasonal changes in oak-leaf tannins and nutrients as a cause of spring feeding by winter moth caterpillars. Ecology 51:565-581. . 1975. Biochemical coevolution between plants and their insect herbivores pp. 3-19 in Coevolution in Animals and Plants (L. E. Gilbert and P. H. Raven, eds.). Univ. Texas Press, Austin. . 1976. Plant apparency and chemical defense. In: Biochemical interactions between plants and insects (J. Wallace and R. Mansell, eds.). Recent Adv. Phytochem. 10:1-49. Finke, M. D. 1977. Factors controlling the seasonal food-plant utilization by larvae of the specialized herbivore, Papilio polyxenes (Lepidoptera). M.S. Thesis. Wright State Univ. Dayton, Ohio. 83 pp. Hegnauer, R. 1971. Chemical patterns and relationships of Umbelliferae. p. 267-278. In: V. H. Heywood (ed.). The Biology and Chemistry of the Umbelliferae. Academic Press, N.Y. 438 pp. Horber, E. 1972. Alfalfa saponins significant in resistance to insects, p. 611-627. In: Insect and Mite Nutrition, J. G. Rodriguez (ed.). North Holland, Amsterdam. . 1974. Techniques, accomplishments and potential of insect resistance in forage le- gumes. p. 312-343. In Proc. Summer Institute of Biological Control of Plant Insects and Diseases (F. G. Maxwell and F. A. Harris, eds.). Univ. Miss. Press, Jackson. Kingsbury, J. M. 1964. Poisonous plants of the United States and Canada. Prentice Hall, Englewood, N.J. 626 pp. Kjaer, A. 1974. Glucosinolates in the Cruciferae pp. 207-220 in J. G. Vaughan, A. J. MacLeod and G. M. G. Jones (eds.). The Biology and Chemistry of the Cruciferae. Academic Press, London. 355 pp. Manglitz, G. L, H. J. Gorz, F. A. Haskins, W. R. Akeson and G. L. Beland. 1976. Interactions between insects and chemical components of sweetclover. J. Environ. Qual. 5:347-352. Odum, H. T. and R. C. Pinkerton. 1955. Time’s speed regulator: the optimum efficiency for maximum power output in physical and biological systems. Amer. Sci. 43:331-343. Pederson, M. W. et al. 1976. Effects of low and high saponin selection in alfalfa on agronomic and pest resistance traits and the interrelationship of these traits. Crop Sci. 16: 193-198. Reese, J. C. 1977. The effects of plant biochemicals on insect growth and nutritional physi- ology. Amer. Chem. Soc. Series 62. pp. 129-152. and S. D. Beck. 1976a. Effects of allelochemics on the black cutworm, Agrotis ipsilon: Effects of p = Benzoquinone, hydroquinone, and duroquinone on larval growth, de- velopment, and utilization food. Annals Entomol. Soc. Amer. 69:59-67. . 1976b. Effects of allelochemics on the black cutworm, Agotis ipsilon: Effects of Catecol, L-dopa, dopamine, and chlorogenic acid on larval growth, development, and utilization of food. Annals Entomol. Soc. Amer. 69:68-72. . 1976c. Effects of allelochemics on the black cutworm, Agotis ipsilon: Effects of resorcinol, phloroglucinal and gallic acid on larval growth, development, and utilization of food. Annals Entomol. Soc. Amer. 69:999-1003. . 1978. Interrelationships of nutritional indices and dietary moisture in the black cut- worm (Agrotis ipsilon) digestive efficiency. J. Insect Physiol. 24:473-479. Rhoades, D. F. and R. G. Cates. 1976. Toward a general theory of plant and antiherbivore chemistry. In Biochemical Interactions Between Plants and Insects (J. Wallace and R. Mansell, eds.) Recent Adv. Phytochem. 10:168-213. Rohweder, D. A. et al. 1978. Forage Crop Varieties and Seeding Mixtures for 1978. Univ. Wise. Extension Publ. A1525. Scriber, J. M. 1975. Comparative nutritional ecology of herbivorous insects: Generalized and VOLUME LXXXVII, NUMBER 2 153 specialized feeding strategies in Papilionidae and Saturniidae (Lepidoptera) Ph.D. The- sis. Cornell Univ., Ithaca, N.Y. 283 pp. . 1977. Limiting effects of low leaf-water content on the nitrogen utilization, energy budget, and larval growth of Hyalophora cecropia (Lepidoptera: Saturniidae). Oecologia (Berl.) 28:269-287. . 1978a. The effects of larval feeding specialization and plant growth form upon the consumption and utilization of plant biomass and nitrogen: an ecological consideration. Proc. 4th Internat. Insect/Host Plant Symp. (London, England June 4-9) Entomol. expt. appi 24:694-710. . 1978b. Cyanogenic glycosides in Lotus corniculatus: Their effect upon growth, energy budget, and nitrogen utilization of the southern armyworm, Spodoptera eridania. Oeco- logia. 34: 143-155. . 1979a. Effects of leaf-water supplementation upon post-ingestive nutritional indices of forb-, shrub, and tree-feeding Lepidoptera. Ent. Expt. app. 25:(in press). . 1979b. The effects of sequentially switching food plants upon biomass and nitrogen utilization by polyphagous and oligophagous Papilio larvae. Ent. Expt. appl. 25: 203-215. and P. P. Eeeny. 1979. Growth of herbivorous caterpillars in relation to feeding spe- cialization and to the growth form of their foodplants. Ecology (in press). Slansky, E. Jr. and P. P. Eeeny. 1977. Maximization of the rate of nitrogen accumulation by larvae of the cabbage butterfly on wild and cultivated foodplants. Ecol. Monogr. 47:209- 228. Sorensen, E. L., M. C. Wilson and G. R. Manglitz. 1972. Breeding for insect resistance, p. 371-390 In C. H. Hanson (ed.) Alfalfa Science and Technology. Amer. Soc. Agron. Madison, WI. Waldbauer, G. P. 1968. The consumption and utilization of food by insects. Adv. Insect Physiol. 5:229-288. Wilson, M. C. and R. L. Davis. 1960. Culver alfalfa, a new Indiana variety developed with insect resistance. Proc. N. Cent. Branch Entomol. Soc. Amer. 15:30. Department of Entomology, University of Wisconsin, Madison, Wiscon- sin 53706 Received for publication Dec. 14, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK2), 1979, pp. 154-159 OBSERVATIONS ON AGGREGATION AND OVERWINTERING IN THE COCCINELLID BEETLE COLEOMEGILLA MACULAE A (DeGeer) Allen H. Benton and Andrew J. Crump Abstract. — In the overwintering site studied, Coleomegilla maculata be- gan aggregating in late September to mid-October. The aggregation proce- I dure appeared to be governed by photoperiod rather than temperature. I The major aggregation site studied included three major physical com- ponents: large prominent willow trees {Salix nigra)', a small pond; and a south-facing slope. Migration to this site appeared to occur via a low level flight of positive anemotactic and hygrotactic nature, with the possibility of a hypsotactic component. No chemotaxis was demonstrated, although the possibility was not ruled out. Once at the aggregation site, beetles remained for some time on the leaves of wild raspberry {Rubus occidentalis) and sensitive fern {Onoclea sensi- bilis), especially the dead dry leaves of the latter. It is likely that this resting » period on dry leaves assists in the loss of excess water. Mortality of the beetles in the aggregation was 3% to 9% in the winter of ) 1973-74. Temperatures within the aggregation were generally higher than i ambient temperatures, and the aggregation had a modifying effect on sudden r temperature changes which might otherwise exceed the insect’s ability to supercool. Introduction The massing of Coccinellid beetles for overwintering has been observed i in many parts of the world. Aggregations are known to occur only in beetle >\ species with abundant individuals which are closely associated with ephem- > eral prey or other food supplies which undergo seasonal fluctuation. Such i is the case in Coleomegilla maculata (DeGeer) which, despite being mark- i edly phytophagous in comparison to closely related coccinellids, aggregates » to overwinter. Once the aggregation has been formed, the beetles exhibit a long dormancy period, often existing in below freezing temperatures. Over- i wintering C. maculata enter a state of ateleodiapause (after Mansingh 1971), 1 as the beetles, upon disturbance, are capable of immediate but somewhat i sluggish movement. In this state the overwintering adult generation survives for periods up to eight times the normal span of the summer generations. Present address: Department of Zoology and Applied Entomology, Imperial College of Sci- ence and Technology, London, England. VOLUME LXXXVII, NUMBER 2 155 The behavioral processes involved in migration toward an aggregation site, site selection and the formation of the aggregation vary widely in dif- ferent species. According to Hagen (1962), C. maculata and Hippodamia convergens are the only two coccinellid species which do not exhibit simple hypsotactic aggregation. In C. maculata, a number of physical factors in- teract to guide the beetle to its aggregation site, so-called climatotactic ag- gregation. These factors and their interaction are, in the case of C. macu- lata, poorly understood. This paper reports observations of aggregating C. maculata and attempts to illuminate some of the processes used in aggre- gation site selection and migration. Location An aggregation site of C. maculata was discovered in 1971 in the town of Pomfret, Chautauqua County, New York. This site has been used each year for at least five years by aggregating beetles and was studied intensively from May 1973 to October 1974, and monitored periodically until May 1976. Unless specifically stated, observations recorded herein refer to this aggre- gation. The aggregation was located near a small farm pond encircled by a clump of willow trees {Salix nigra), on the south-facing slope of a high bank which had resulted from digging the pond. The bank was generally covered with leaf litter and undergrowth. The area was about 100 meters north of exten- sive cornfields, and separated from them by a major rail line. Three other aggregations, two in Erie County, New York and one in Cayuga County, New York, were discovered during the course of the study and were monitored periodically. The Erie County sites were also at the base of large willow trees. The Cayuga County site was at the base of a large poplar {Pop ulus delt aides) in the front yard of a farm home and ap- proximately 100 meters west of a large cornfield, and 50 meters north of another cornfield. Field Observations In 1973, the first influx of beetles at the aggregation site occurred on October 1, and in 1974 on September 28. These dates were either identical or approximate to arrival dates at the three other sites. Since the 1974 date followed several days of unseasonably hot weather (20°C), we felt that pho- toperiod rather than temperature was the signal for aggregation. In 1975, however, beetles arrived at the Cayuga County site on October 15, and date of arrival at the main site could not be determined. A few beetles were found on December 1 , at a site about ten meters from the major aggregation sites of previous years, so that the exact time of aggregation is unknown. We believe that photoperiod is an important factor in the instigation of 156 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Temperatures (°E) at the three sites of overwintering beetles in the aggregation in Town of Pomfret, Chautauqua County, New York, in the winter of 1973-74. Month Site 1 Site 2 Site 3* Low Mean Low Mean Low Mean December 27 42 24 37 24 39 January 21 33 22 35 15 31 February 14 39 12 41 8 31 March 20 44 14 42 13 38 * No beetles were present at this site after December. processes leading to migration and aggregation, although it may not be the prime causal factor. After the arrival of the first beetles, the aggregation grew rapidly and appeared to be complete within seven days of the initial arrivals. During the initial influx, clusters of beetles were found in depressions in the ground or at the base of saplings around the pond. These sites proved to be transi- tional, with the beetles eventually joining the main aggregation. The beetles formed three distinct clusters within the aggregation area. The largest of these clusters (site 1) contained the most beetles and was located around the base of the largest willow tree. Site 2 was established in a large depres- sion near the entrance to a woodchuck hole, approximately 5 meters from site 1. The third site (3), containing the smallest number of beetles, was situated on the bank equidistant from sites 1 and 2. During early December the beetles at site 3 migrated to join those at site 1. The final two clusters at sites 1 and 2 covered little more than 0.5 square meters each, both being approximately 3 meters from the pond and elevated 1 meter above the water level. The location of the biggest cluster of beetles (site 1) near the base of the bank and at the base of the largest tree afforded maximum protection from the cold prevailing northerly winds moving in off nearby Lake Erie. At no time during the study were beetles found on the north-facing slope of the bank. Having migrated to the aggregation site, beetles alighted on the upper sides of the abundant undergrowth species, including wild raspberry (Rubus occidentalis). Soon after arrival the beetles congregated on the undersides of the leaves and particularly under or in dead, dry, rolled up leaves of the sensitive fern {Qnoclea sensiblis). Beetles on the undersides of leaves were generally immobile, while those on the upper surfaces (presumably newer arrivals) were more active. This pattern of arrival and subsequent associa- tion with dead vegetation was apparent throughout the aggregation forma- tion. The reason for this is unclear but may be connected with the need to lose excess water, the freezing of which is thought to be the cause of some mortality among overwintering beetles (Hodson, 1937). VOLUME LXXXVII, NUMBER 2 157 While in this situation, the beetles show a definite thermokinesis. Warmer periods induced greater activity but the beetles remained at the same levels on the vegetation. The thermokinetic response is, however, clearly com- plexed with other tactic responses as drops in temperature served to accel- erate or instigate movement to the ground. Once on the ground, the beetles huddled under the top layers of loose soil and dead and decaying vegetation, where they remained for the duration of the winter. Mortality Maximum-minimum thermometers were placed among the three original aggregating groups of beetles at sites 1, 2, and 3. The thermometer at site 3 was left in place after the migration of the beetles to site 1. Sudden drops in temperature, thereby not giving the beetles sufficient time to supercool, are known to be the major cause of death in overwintering beetles (Hodson, 1937). During the winter of 1973-74, the temperatures never dropped below the level to which C. maculata can supercool successfully. Temperatures taken during the winter months (Table 1) show two main points. First, the “low” temperatures at sites 1 and 2 were higher than those at the beetle- less site 3. Second, and more important, the “low” temperatures at site 2 were usually lower than those at site 1. This indicates the importance of the greater number of beetles at site 1 helping to maintain a higher temperature within the aggregation. Actual mortality was determined by taking groups of beetles from the aggregation, placing them in wire mesh containers which afforded little extra protection, and replacing them in their original aggregation. At the end of the winter, mortality figures of 3% for site 1 and 9% for site 2 were obtained. In addition to the greater number of beetles at site 1, most of the beetles at i this site were shielded from northwesterly winds by the presence of the willow tree whereas those at site 2 were not. This, together with the lower I number of beetles, may account for the higher level of mortality observed ' at the more exposed site 2. Discussion The mechanisms and responses involved in triggering aggregation and in the migration to the aggregation site are difficult to elucidate due to their . complex interactions. The possibility of a chemical cue being utilized in the selection of the aggregation site cannot be dismissed. Weiss (1913) felt that odor was important as an attractant to the aggregation site. However, during 1 olfactory tests carried out on C. maculata no chemotaxis was found, the only consistent positive response being towards moisture-laden air. Landis (1936) showed that there is a significant difference in the reassociation of the Malpighian tubules in climatotactic as opposed to hypsotactically ag- 158 NEW YORK ENTOMOLOGICAL SOCIETY gregating species. Hodson (1937) also indicated that C. maculata showed a marked preference for specific moisture situations, suggesting that these were significant in reducing the effect of cooling by the latent heat properties of water. C. maculata is known to be very susceptible to desiccation and the ability of the beetle to detect and move towards moisture-laden air has been shown to affect its distribution in its summer habitat (Crump and Ben- ton, in prep.). Adult C. maculata are larger and heavier than other closely related Coccinellids, including H. convergens which also undergoes cli- matotactic aggregation. The larger size, in conjunction with the predilection for moist areas and its profoundly pollenaceous diet, causes the low level flights and low distribution of C. maculata in vegetation (Ewert and Chiang, 1966). Short, low-level migratory flights would explain why many aggrega- tions are found at or near the edges of open fields. Migration to the aggre- gation site is determined by positive hygrotactic and anemotactic responses complexed with a hypsotactic component causing aggregation at the bases of prominent objects in an area of optimum moisture level. Aggregation of C. maculata serves as a method of enhancing winter sur- vival, there being two aspects to this enhancement. First, the large number of beetles present emphasizes the aposematic coloration of the beetle. Second, and more significant, large numbers of beetles provide a degree of heat conservation. These factors combined with the biochemical mecha- nisms involved in the ateleodiapause of C. maculata ensure a greater sur- vival rate over the arduous winter months. Acknowledgments This research was supported by National Science Foundation grant B036435. For assistance with field work, we are indebted to James Moore and Richard Northrup. Literature Cited Balduf, W. V. 1935. The bionomics of entomophagous Coleoptera. J. S. Swift Co., Inc., St. Louis. Crump, A. J. and A. H. Benton. In prep. Observations on spring and summer behavior in Coleomegilla maculata (DeGeer). Ewert, M. A. and H. C. Chiang. 1966. Dispersal of three Coccinellids in a corn field. Canad. Entomol., 98:999-1003. Hagen, K. S. 1962. Biology and ecology of predaceous Coccinellidae. Annual Rev. Entomol., 7:289-326. Hodson, A. C. 1937. Some aspects of the role of water in insect hibernation. Ecol. Mono- graphs, 7:271-315. Landis, B. J. 1936. Alimentary canal and Malpighian tubules of Ceratomegilla fuscilabris (Muls.). Ann. Entomol. Soc. Amer., 29:15-28. VOLUME LXXXVII, NUMBER 2 159 Mansingh, A. 1971. Physiological classification of insect dormancies. Canad. Entomol., 103:983-1009. Weiss, H. B. 1913. Some tropic reactions of Megilla maculata (DeGeer). Canad. Entomol., 45:85-87. Department of Biology, State University College, Fredonia, New York 14063. Received for publication December 14, 1978. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 160-166 ON THE IDENTITY OF TWO SPECIES OF RHYPAROCHROMINAE FROM ARGENTINA (HEMIPTERA: LYGAEIDAE)* James A. Slater Abstract. — Two species described by Berg from Argentina in 1883 and 1884 and placed by him in the otherwise Palearctic genus Tropistethus have been restudied. Both species, T. dubius and T. australis, are considered to be distinct species in the genus Cryphula. A lectotype is selected for Cry- phula dubia (Berg); redescriptions from type specimens are given for both species; a discussion is given of their relationships to other species of Cry- phula. Syngenicus Berg 1883, listed as a provisional new generic name for T. dubius, is placed as a junior synonym of Cryphula. The lectotype of Cryphula dubia is figured. In 1883 Berg described a new species of lygaeid from Argentina under the name Tropistethus dubius. Berg obviously had considerable doubt as to the correct generic position of this species as he stated that it might well rep- resent a new genus and, indeed, went so far as to reserve the name Syn- genicus for it in the event that it should prove to represent an undescribed genus. In the following year Berg (1884) described another new species from Argentina as Tropistethus australis. To my knowledge neither of these species has been subsequently discussed and the names only appear in fau- nal lists and catalogues (see Slater 1964). Tropistethus Fieber is a Palearctic genus (nine species) in the tribe An- tillocorini. I have recently been interested in the systematics and phylogeny of the Antillocorini of the Neotropics and have had an opportunity to ex- amine the Berg types. As might have been anticipated neither species is congeneric with the Palearctic species of Tropistethus. Both species belong in the genus Cryphula Stal, a member of the rhyparochromine tribe Le- thaeini. Both dubius and australis are, however, distinct from any described species of Cryphula. Syngenicus Berg 1883 (type species Tropistethus dubius) thus becomes a junior synonym of Cryphula Stal (1874) (type species Cryphula parallel- ogramma StM) NEW SYNONYMY. Cryphula dubia (Berg) new combination 1883. Tropistethus dubius Berg pp. 265-266. Berg described dubius from two males from Chacabuco (“Provincia Bon- aerensis”) collected by Felix Lynch. I have examined both specimens which ' This work was supported by a grant from the National Science Eoundation. VOLUME LXXXVII, NUMBER 2 161 have identical labels saying (1) “Typus” (2) “Chacab. F. Lynch.” Berg did not select a holotype and thus it is necessary to designate a lectotype. This is particularly important in the present instance as the two specimens prob- ably are not conspecific. Berg apparently thought he was dealing with brachypterous specimens. One syntype, indeed, is coleopteroid with only a narrow membrane vestige present. The second specimen, however, is a macropter with most of the membrane broken off, as is the distal portion of the right corium. There seems little doubt that Berg drew his description from the coleopter as he mentions the dark corial macula and testaceous legs which this specimen has but which is lacking on the mutilated macropter. Further a third label apparently in Berg’s handwriting saying "Tropistethus dubius Berg” is on the pin with the coleopter. Accordingly this specimen is selected as lecto- type and an appropriate label attached. Redescription of Male Lectotype of Cryphula dubia (Berg) (Figure 1) Head, pronotum, scutellum, a large ovoid macula in center of corium, and antennal segments three and four dark reddish brown. Tylus yellowish. A large macula at each humeral pronotal angle which narrows mesad, ex- treme apex of scutellum, entire hemelytra except as indicated above, first and second antennal segments, labium and legs bright yellow. Forefemora infuscated with brownish on proximal three-fourths. Pleural and ventral surfaces completely shining or subshining, reddish brown. Head and prono- tum strongly polished and shining. Dorsal surface, especially hemelytra, clothed with elongate, erect and conspicuous yellow hairs. Head broad, eyes almost in contact with anterolateral pronotal angles; tylus broad distally reaching distal two-thirds of first antennal segment; head length .52\ width .70, interocular space .50. Pronotum subquadrate, dis- tance across anterior portion nearly as broad as that across humeri, ante- rior lobe conspicuously convex, posterior lobe narrow, lateral margins slightly sinuate; posterior margin shallowly concave, pronotum length .64, width .94. Scutellum length .58, width .58. Clavus and corium completely fused, commissure meeting evenly down midline, membrane vestige a nar- row strip along apical margin of hemelytron and not extending posterior to caudolateral corner; lateral margins of hemelytron explanate and moder- ately arcuate, hemelytron length 1.40. Metathoracic scent gland auricle rather elongate, subtruncate, irregularly curving posteriorly. Evaporative area with distal (=dorso-lateral) margin evenly truncate, lacking shining in- trusion anteriorly. Forefemora strongly incrassate, armed below near distal end with three short sharp spines followed proximally by two or three ’ All measurements are in millimeters. 162 NEW YORK ENTOMOLOGICAL SOCIETY Eig. 1. Cryphula dubia (Berg) Lectotype. Dorsal view. VOLUME LXXXVII, NUMBER 2 163 very elongate conspicuous spines. All tibiae with coarse spines present over surface. Antenna stout, typical of genus, antennal segment lengths I .28, II .48, III .46, IV .58. Labium probably attaining hind coxae, length labial segments I .40, II .32, III .36, IV .30. The lectotype of Cryphula dubia has numerous elongate, erect hairs on dorsal surface. It will run to Cryphula abortiva Barber at couplet four in Scudder’s (1962) key to the species of Cryphula. It is, however, not closely related to abortiva, differing in having pale testaceous hemelytra with a dark median macula, pale yellow humeral maculae, pale first and second antennal segments and a highly polished pronotum. C. dubia is rather sim- ilar in habitus to the recently described Cryphula bennetti Baranowski and Slater by virtue of the pale first and second antennal segments and legs, dark corial macula, pale humeral spots and dark scutellum. C. bennetti, however, lacks elongate hairs on the dorsal surface, has the rows of he- melytral punctures outlined as a series of dark stripes, has much larger humeral maculae, more evenly arcuate lateral pronotal margins and a white posterior metapleural lobe. The other syntype of dubius is in poor condition. In addition to the broken membrane the antennae, fore tibiae and tarsi are all missing, and the spec- imen is greasy. It appears to be a species of Cryphula although in habitus it also resembles species of Valtissius Barber. The poor condition of the specimen makes it inadvisable to attempt to place it further taxonomically. Cryphula australis (Berg) new combination 1884. Tropisthetus [i/c] australis Berg, pp. 187-188. Berg described australis from a single female from “Tandil” collected by “Dr. Holmberg.’’ The holotype is a conventional looking Cryphula and runs in Scudder’s (1962) key to Cryphula apicata (Distant) at couplet 6. Through the kindness of Mr. W. R. Dolling of the British Museum I have been able to examine Distant’s lectotype of apicata, a male which was described from “S. Geronimo, Guatemala.’’ The types of australis and apicata are not conspecific. In australis the lateral pronotal margins are distinctly sinuate whereas in apicata they are evenly although narrowly arcuate. The mem- brane of apicata is dark brown with contrastingly pale veins, that of aus- tralis is white with a very large chocolate brown macula occupying most of the center of the surface. The posterior lobe of the metapleuron of aus- tralis is white and strongly contrasting with the anterior lobe, whereas in apicata the posterior metapleural lobe is reddish brown and concolorous with the anterior lobe. In apicata the metathoracic scent gland auricle is more elongate than it is in australis and slightly enlarged and subtruncate on the distal end, whereas it is subacute in australis. The adjacent evapo- rative area of apicata is invaded near its anterodorsad edge by a mesally 164 NEW YORK ENTOMOLOGICAL SOCIETY projecting "finger” of shining surface whereas in australis the dorsolateral margin of the evaporative area is straight throughout. C. apicata is ex- tremely closely related to C. nitens Barber with which it agrees in all of the above characteristics. In fact, the specific status of apicata and nitens needs further study. Barber (1955) differentiated nitens from apicata by the former having the antennae “provided with several long semi-erect setae,” by hav- ing “less conspicuous veins of the corium” and by differences in the color of the pronotum. I have examined specimens of nitens from Utah, California and Texas and cannot find differences in antennal hairs or degree of corial color differentiation between these specimens and the lectotype of apicata. C. apicata does have the pale coloration of the posterior pronotal lobe more prominently and extensively developed. I think the status of nitens ques- tionable and in need of careful analysis. Redescription of Female Holotype of Cryphula australis Head, pronotum (except anterior margin, humeral angles and irregular extensions over posterior lobe on either side of midline), scutellum (except apex) and a very large ovoid distally irregular macula in center of hemelytral membrane reddish to chocolate brown. Remainder of dorsal surface white to light testaceous, hemelytral punctures darker brown forming irregular dark stripes. Nearly uniformly dark reddish brown below, shining, with dorsal portion of posterior propleural lobe and all of hind lobe of metapleu- ron a strongly contrasting white. Legs light yellow with anterior femora yellowish brown on proximal three-fourths. Dorsal surface nearly glabrous lacking elongate upstanding hairs. Head slightly declivent, conspicuously convex between eyes, latter in contact with anterolateral angles of pronotum, head length .58, width .76, interocular space .50. Pronotum with lateral margins narrowly but conspic- uously explanate and markedly sinuate, posterior margin shallowly concave before scutellum, pronotum length .68, width 1.24. Scutellum length .74, width .76. Claval commissure length .36. Lateral margins of corium explan- ate and evenly arcuate. Distance apex clavus-apex corium .54; distance apex corium-apex membrane .58. Metathoracic scent gland auricle finger- like tapering distally, evaporative area with dorsolateral margins evenly ' truncate, complete. Forefemora strongly incrassate, armed below near distal ends with two very sharp spines followed proximally by three conspicuous elongate hairs. Antennae missing. Labium extending well between and prob- ably slightly beyond posterior margin of mesocoxae, perhaps reaching | metacoxae, length labial segments I .38, II .40, III .36, IV .30; total length 3.36. I VOLUME LXXXVII, NUMBER 2 165 Redescription of Lectotype of Cryphula apicata Distant Head, major part of pronotum and scutellum dark chocolate brown. An- terior marginal area of pronotum, elongate elliptical macula in area of humeri and three conspicuous spots along posterior margin of pronotum white to very pale testaceous. Hemelytra variegated, veins conspicuously white or j pale yellow, intervening areas reddish brown mottled with testaceous and with a conspicuous small pale elliptical macula present near apical margin I between radius and medius. Membrane brown with veins a strikingly con- trasted translucent white. Antennae yellowish, distal third of segment three I and all of segment four darker. Legs pale reddish brown becoming testa- 1 ceous distally. Ventral and pleural surfaces including posterior lobe of I metapleuron bright red brown but with pale yellow to white coloration pres- ent along dorsal portion of posterior lobe of propleuron. Dorsal surface I nearly glabrous lacking conspicuous upstanding hairs. Head slightly declivent, moderately convex between eyes, tylus broad reaching at least to distal fourth of first antennal segment, head length .50, I width .72, interocular space .46. Pronotum subquadrate, lateral margins ; narrowly explanate evenly curving or gently arcuate from humeri to antero- lateral angles, pronotum length .76, width 1.22. Scutellum length .76, width .72. Claval commissure length .38. Corium very slightly arcuate, nearly straight for greater portion of length. Distance apex clavus-apex corium .70, distance apex corium-apex membrane .44. Metathoracic scent gland auricle elongate slightly broadened and subtruncate at distal end, evaporative area with deep finger-like shining intrusion into dorsolateral margin near anterior end of metapleuron. Forefemoral armature and labial length obscured, latter apparently reaching at least well between mesocoxae. Antennal segment lengths I. 24, II .48, III .38, IV .46. Total length of body 3.32. Acknowledgments I am indebted to Dr. Luis De Santis (Universidad Nacional de la Plata, Argentina) and to Mr. W. R. Dolling (British Museum of Natural History) for the loan of type material from their respective institutions. My appre- ciation is extended to Mr. Steven Thurston (formerly University of Con- necticut) for preparation of the dorsal view illustration of Cryphula dubia and to Mrs. Betty Slater for aid in the preparation of the manuscript. Literature Cited Barber, H. G. 1955. The genus Cryphula Stil, with the description of two new species (Het- eroptera: Lygaeidae). Journ. New York Ent. Soc. 63:135-137. Baranowski, R. M. and J. A. Slater. In press. Notes on the biology of two species of 166 NEW YORK ENTOMOLOGICAL SOCIETY Cryphula (Hemiptera, Lygaeidae) in Trinidad with the description of a new species. Florida Entomologist. Berg, C. 1883. Addenda et emendanda ad Hemiptera Argentina. An. Soc. Cient. Argent. 15:241-269. . 1884. Addenda et emendanda ad Hemiptera Argentina. Bonariae; Pauli E. Coni pp. 1-213. Scudder, G. G. E. 1%2. The world Rhyparochrominae (Hemiptera: Lygaeidae) I. New syn- onymy and generic changes. Canad. Ent. 94:764-773. Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. University of Connecticut, 2 volumes, 1668 pp. St^l, C. 1874. Enumeratio Hemipterorum pt. 4, K. Svenska Vetensk. Akad. Hand). 12:1:1- 186. Systematic and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06268. Received for publication January 26, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 167-174 INTRODUCED PARASITES OF AGROMYZA FRONTELLA (RONDANI)> IN THE USA R. M. Hendrickson, Jr. and S. E. Barth Abstract. — From 1974 to 1978, we released 14 species of European par- asites (ca. 82,000 insects from 3 agromyzid host species on alfalfa) as part of a biological control effort against the alfalfa blotch leafminer, Agromyza frontella (Rondani), in the USA. A list of release locations, dates, and par- asite numbers is presented. Techniques for rearing 4 of the species are briefly described. Two species, Dacnusa dryas (Nixon) (Hymenoptera: Bra- conidae) and Chrysocharis punctifacies Delucchi (Hymenoptera: Eulophi- dae), have been established in Delaware. Sufficiently large field populations of these species were present in 1978 so that insects were collected by sweeping and moved to other release locations. The alfalfa blotch leafminer (ABL), Agromyza frontella (Rondani), a pest of European origin first reported in the USA in Massachusetts in 1968 (Miller and Jensen, 1970), is now found throughout the northeastern states and adjacent Canadian provinces. The USDA began the introduction of parasites for biological control of this pest in 1974. The purpose of this paper is to document parasite releases and recoveries. Sources of parasites released. — Foreign parasite material was provided by the USDA European Parasite Laboratory, Sevres, France. The material was received at our laboratory quarantine facility as adults or as immatures in host puparia. Several parasite species were reared successfully in the laboratory, and releases were made from these cultures. By 1978, 2 species of parasites, Dacnusa dryas (Nixon) and Chrysocharis punctifacies Deluc- chi, became so abundant in local fields that they were collected by sweeping and moved to new release locations. The order, family, and determiner of all parasite and host species mentioned in this paper are listed in Table 1. Releases of 14 species of European parasites (ca. 82,000 insects from 3 agromyzid host species on alfalfa) against ABL were made from 1974 to 1978. The species, numbers, and locations are found in Table 2. Techniques for rearing parasite species. — We attempted to rear nearly all of the 14 parasite species but were successful with only 4 of them. (Since the use of ABL as a host means that considerable space must be provided for potted alfalfa and relatively few ABL per plant are produced, we used alternate hosts whenever possible.) Diptera: Agromyzidae. 168 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Order, family, and determiner of species mentioned in this paper. ll Species ■ 1 HYMENOPTERA: BRACONIDAE | Chorebus poss. misellus (Marshall)® 5 Dacnusa dryas (Nixon)® I Dacnusa maculipes Thomson® Dacnusa sp.® Dapsilarthra balteata (Thomson)' Opius dureseaui Fischer® Opius maculipes (Thomson)® Phanomeris braconius (Haliday)® HYMENOPTERA: EULOPHIDAE Chrysocharis naenia (Walker)' Chrysocharis punctifacies Delucchi' Diglyphus intermedins (Girault)'’ Diglyphus isaea (Walker)'’ HYMENOPTERA: PTEROMALIDAE Halticoptera circulus (Walker)' Miscogaster hortensis Walker’’ Miscogaster maculata Walker’’ DIPTERA: AGROMYZIDAE Agromyza frontella (Rondani)’^ Agromyza nana Meigen' Agromyza parvicornis Loew’' Liriomyza congesta (Becker)' Liriomyza trifoliearum Spencer* ® P. M. Marsh, ’’ G. Gordh, ' G. C. Steyskal, and ’’ E. E. Grissell, Systematic Entomology Laboratory, USDA-SEA-AR, do U.S. National Museum, Washington, DC 20560. ' M. Fischer, Naturhistorisches Museum, Zoologische Abteilung, Vienna, Austria. I ' C. M. Yoshimoto, Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6. y [i Chrysocharis punctifacies, a solitary endoparasite of larvae, emerging j from puparia, was successfully reared on ABL by placing adult parasites in I cages containing alfalfa plants that had mining larvae in the leaflets. After the parasites oviposited, the plants were laid on their sides so mature par- > J asitized hosts dropped from the leaflets into trays of moist vermiculite from . which they eventually emerged. This is a slight modification of the tipped ' i plant method for rearing ABL described by Hendrickson and Barth (1977). 1 The parasite also oviposited readily on Liriomyza trifoliearum, reared on i bean plants, but this leafminer invariably died after transforming to the pupal i stage, and no parasites developed. Dacnusa dryas, another solitary endoparasite of larvae, emerging from 1 ^ puparia, was successfully reared on ABL by using the same techniques VOLUME LXXXVII, NUMBER 2 169 Table 2. Releases of European parasites against the alfalfa blotch leafminer in the USA. Original host species are indicated in footnotes. State County Locality Release year (19—) No. Chorebus poss. misellus^’’^ Delaware New Castle Newark 77 132 Chrysocharis naenia^-'^ Delaware New Castle Newark 77 28 New Jersey Warren Blairstown 78 5 Total 33 Chrysocharis punctifacies^ Delaware New Castle Newark 77 1,277 78 217 New Jersey Burlington Rancocas 78 100 Warren Blairstown 78 35 New York Cortland Cortland 78 255 Dutchess Standfordville 78 16 Herkimer Mohawk 78 163 Tompkins Dry den 78 338 Ohio Ashtabula Padanaram 78 29 Pennsylvania Chester Kemblesville 78 10 New London 78 10 Oxford 77 66 78 791 Total 3,307 Dacnusa dryas^ Delaware New Castle Newark 77 620 78 596 New Jersey Burlington Rancocas 77 12 78 156 Warren Blairstown 78 126 New York Cortland Cortland 78 1,839 Dutchess Stanfordville 78 30 Herkimer Mohawk 78 254 Ohio Ashtabula Padanaram 78 21 Pennsylvania Chester Kemblesville 78 50 New London 78 50 Oxford 77 55 78 1,398 Total 5,207 Dacnusa maculipes'’-’^ Delaware New Castle Newark 77 352 Pennsylvania Chester Oxford 77 13 Total 365 170 NEW YORK ENTOMOLOGICAL SOCIETY Table 2. Continued. State County Locality Release year (19-) No. Dacnusa sp. a,b,c Delaware New Castle Newark 76 10 77 653 New Jersey Burlington Rancocas 77 97 Pennsylvania Chester Oxford 77 139 Total 899 Dapsilarthra balteata^-’^ Delaware New Castle Newark 77 50 78 3 Pennsylvania Chester Oxford 78 7 Total 60 Diglyphus Delaware New Castle Newark 75 2,000 76 16,295 Maryland Cecil Fair Hill 75 600 76 2,000 Massachusetts Hampshire Hadley 75 600 New Hampshire Grafton Concord 75 600 New Jersey Burlington Rancocas 75 1,850 76 13,800 Warren Blairstown 75 1,150 76 2,500 New York Orange Windsor 75 400 Pennsylvania Chester Kemblesville 75 50 New London 75 1,400 76 4,000 Oxford 75 950 76 17,950 Schuylkill Port Clinton 75 400 Total 66,545 Halticoptera circulus^ Delaware New Castle Newark 78 5 Miscogaster hortensis^ and M. maculata^ Delaware New Castle Newark 75 34 76 5 77 355 78 270 Ohio Ashtabula Padanaram 78 3 Pennsylvania Chester Oxford 77 70 78 110 New Jersey Burlington Rancocas 78 8 Warren Blairstown 78 30 VOLUME LXXXVII, NUMBER 2 171 Table 2. Continued. Release year State County Locality (19 — ) No. New York Cortland Cortland 78 34 Herkimer Mohawk 78 3 Tompkins Dryden 78 71 Total 993 Opius dureseaui^-'' Delaware New Castle Newark 75 22 77 127 78 20 New Jersey Warren Blairstown 75 16 New York Cortland Cortland 78 51 Herkimer Mohawk 78 52 Ohio Ashtabula Padanaram 78 8 Pennsylvania Chester Oxford 78 13 Total 309 Opius maculipes^ Delaware New Castle Newark 77 10 Pennsylvania Chester Oxford 78 45 Total 55 Phanomeris braconius^’'' Delaware New Castle Newark 75 20 76 320 77 1,494 78 741 New Jersey Burlington Rancocas 74 107 77 148 78 240 Warren Blairstown 74 8 77 16 78 190 Hope 77 40 Sussex Vernon 74 31 Pennsylvania Chester Oxford 76 20 77 352 78 357 Landenberg 77 32 Total 4,116 ! ^ Agromyza frontella 1 Agromyza nana. ' Liriomyza congesta " Host relationships are uncertain due to a recent determination that 2 species of Miscogaster were involved, rather than an initial determination of 1 species. 172 NEW YORK ENTOMOLOGICAL SOCIETY described for C. punctifacies. When the parasite was exposed to L. trifo- liearum on bean plants, only a few males were produced. However, both sexes emerged from L. trifoliearum collected at the laboratory field in 1978. Diglyphus isaea, a larval ectoparasite of ABL, was initially reared on ABL infesting potted alfalfa plants. Once the parasites had oviposited on these larvae, the alfalfa stems were cut, and placed in emergence cages, and allowed to dry out. However, production of these parasites was limited by the relatively small numbers of host larvae available on potted alfalfa plants. Later, an easier and more productive method of rearing on L. trifoliearum on bean plants was devised (Hendrickson, 1975). In either case, the parasite had to be provided with 3rd-(mature) instar host larvae since it killed Isl- and 2nd-instar larvae by probing, usually without ovipositing. Phanomeris braconius, a larval ectoparasite, was successfully reared on ABL by the methods described for D. isaea. Thus L. trifoliearum on bean plants was an acceptable host, but the technique was inefficient because the emerging parasites were small and too many host larvae developed into adult flies. Best results were obtained by rearing P. braconius on Agromyza parvicornis, a native pest of corn. After the parasites had oviposited on the host larvae, the com culms (ca. 30-cm tall) were cut, placed in an emergence cage, and dried. Honey was provided as food for emerging parasites. The parasites obtained were slightly larger than those that developed on ABL. Parasite establishments. — The following species were released at the lab- oratory alfalfa field in 1977 and recovered by rearing from ABL or by sweep- ing in 1978. The sampling and emergence techniques employed were de- scribed by Hendrickson and Barth (1979). Dacnusa dryas was released at a rate of 750/ha. The parasite was first recovered from samples collected June 26, 1978 (2nd-cutting alfalfa) and was collected from subsequent samples through November. Dacnusa dryas was the most abundant parasite species during the period June 26-30, when it parasitized 14% of the ABL. On the basis of number of stems per m^, number of mines per stem, and percentage of mines producing the parasite, we calculate that this peak emergence resulted in ca. 175,000 host larvae/ha , later producing the parasite species. The rapid rate of increase in only a year is probably a result of a relatively high rate of parasite fecundity and the occurrence of 5 generations of ABL per year in the Delaware area. ■ Parasite adults were collected by sweeping (maximum recovery rate was 35/100 sweeps) and moved to other release locations. At first we aspirated |j the D. dryas adults from the sleeve cages where we placed the field material j from the 100 sweeps. This procedure was time consuming, and separation 1 of D. dryas from other insects was difficult. It was more efficient to put the j unseparated field material from 1,000 sweeps (collected 200 at a time) into f a large, styrofoam container cooled by a freeze pack wrapped in paper | VOLUME LXXXVII, NUMBER 2 173 towels. Then the material was taken to a release location. Mortality of material handled in this manner appeared to be slight. Chrysocharis punctifacies was released at a rate of 190 /ha in 1977. The parasite was first recovered on July 17, 1978 (3rd-cutting alfalfa) and was reared from samples collected through September. It was the most abundant parasite species during Sept. 18-22 when it parasitized 34% of the ABL. Adults were collected by sweeping (maximum recovery rate was 5/100 sweeps) and moved to release locations. We had the same difficulty sepa- rating adult C. punctifacies from field-collected material as we had in sep- arating D. dry as, but in addition, several common native Chrysocharis species cannot be distinguished from C. punctifacies except under the mi- croscope. Again this species was more efficiently disseminated locally by mass collections. However, we unexpectedly recovered a few C. puncti- facies from field-collected L. trifoliearum. Possibly size of host explains our inability to rear the parasite in the laboratory (field-collected insects are larger than laboratory-reared specimens). Parasites of uncertain establishment status. — A total of 86 Miscogaster \ hortensis and M. maculata, solitary endoparasites of larvae, emerging from I puparia, was released in 1975-77 at the laboratory alfalfa field (release rate i was 85/ha). (The European host relationships for Miscogaster spp. in Table j 2 are uncertain because the initial determination was that only a single j species was involved; later examination of a more extensive series of spec- I imens indicated 2 species were involved.) No Miscogaster spp. emerged from samples collected in 1978, but several adult M. hortensis were col- lected by sweeping. Although the species has survived at least one winter, I we believe recovery was too low to warrant a claim of establishment. The I species may eventually become sufficiently abundant for unquestioned es- I tablishment and it may produce significant results. i Miscogaster spp. comprised 15% of the parasites emerging from Euro- j pean ABL puparia processed at the laboratory quarantine facility in 1976- 77 and the genus was third in abundance after C. punctifacies and D. dryas (Hendrickson and Barth, 1979). Diglyphus isaea, the most abundant parasite of ABL in Europe (J. J. Drea, Jr., pers. com.), was reared in the laboratory and released in large numbers in 1975-76. There were few recoveries in those years and none subsequently. Since D. isaea is very similar morphologically to the most abundant North American parasite attacking ABL, D. intermedins, we ex- amined the hypothesis that these 2 species had hybridized. Crosses of D. intermedins 9 with D. isaea 6 produced fertile female progeny, but the reciprocal cross produced only males, which indicated that fertilization had not taken place. It therefore seems probable that some D. isaea genetic material has been added to the D. intermedins gene pool in the USA. How- 174 NEW YORK ENTOMOLOGICAL SOCIETY ever, D. isaea females appear to be reproductively isolated from D. inter- medius, so their disappearance may have resulted from competitive dis- placement with D. intermedins, or from mating (without fertilization) with D. intermedins males, which were far more abundant in the field than D. isaea males. Literature Cited Hendrickson, R. M. Jr. 1975. Mass rearing of Diglyphus isaea (Walker) (Hymenoptera: Eu- lophidae) on Liriomyza trifoliearum Spencer (Diptera; Agromyzidae). (Abstract). J. N.Y. Entomol. Soc. 83:243-244. and S. E. Barth. 1977. Techniques for rearing the alfalfa blotch leafminer. J. N.Y. Entomol. Soc. 85:153-157. . 1979. Effectiveness of native parasites against Agromyza frontella (Rondani) (Diptera: Agromyzidae), an introduced pest of alfalfa. J. N.Y. Entomol. Soc. 87:85-90. Miller, D. E. and G. L. Jensen. 1970. Agromyzid alfalfa leaf miners and their parasites in Massachusetts. J. Econ. Entomol. 63:1337-1338. Beneficial Insects Research Laboratory, USDA-SEA-AR, Newark, Del- aware 19713. Received for publication April 12, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 175-180 PHOTOPERIOD AND TEMPERATURE INFLUENCES ON EGG NUMBER IN BRACHYMERIA INTERMEDIA (HYMENOPTERA: CHALCIDIDAE), A PUPAL PARASITOID OF LYMANTRIA DISPAR (LEPIDOPTERA: LYMANTRIIDAE)* P. Barbosa and E. A. Frongillo, Jr.^ Abstract. — Oosorption in Brachymeria intermedia (Nees) is photoperi- odically induced by a short photophase. Low temperatures enhance the effects of photoperiod. Oocyte resorption proceeded more quickly at lower temperatures. Preconditioning of immature stages within its host or of young adults has an effect on rate of oocyte resorption. I Insects exhibit a variety of reproductive mechanisms including viviparity, ovoviviparity, oviparity, etc. In many species, particularly in parasitoid species, the timing of oviposition is critical. Appropriate timing of oviposition and conservation of nutrients may be enhanced by oosorption. Oosorption is a process characterized by cessation of vitellogenesis and degeneration of the vitellogenic ooctye within the ovary. It has been associated with ' various life history phenomena as an energy conserving reproductive strat- egy. That is, resorption is an adaptive mechanism which occurs when ovi- position would waste energy. Resorption of oocytes in diapausing females I has been described in a number of species including an ichneumonid para- sitoid. It is also found in insects avoiding adverse conditions, characterized j by low temperature or short day photoperiods (Bell and Bohm 1975). The reproductive system of Brachymeria intermedia (Nees), a European pupal parasitoid of the gypsy moth, is typically hymenopteran (D’Rozario 1 1942), containing two ovaries each of which is comprised of three polytro- phic ovarioles (Dowden 1935, Engleman 1970). A mature ovariole will usu- ally hold one chorionic and one or two other vitellogenic oocytes. Thus, B. intermedia is a synovigenic species with monootene ovarioles where ovu- lation is externally induced (Flanders 1950). Flanders (1950) suggested that hymenopteran parasitoids that have synovigenic females are more likely to ‘ Paper No. 2312 Massachusetts Agricultural Experiment Station, University of Massachu- setts at Amherst, MA. U.S.A. This research supported (in part) from Experiment Station Project No. 437 and U.S.D.A. sponsored program entitled, “The Expanded Gypsy Moth Re- search and Development Program" (CSRS Special Grant No. 516-15-57). ^ Present address — Dept, of Entomology, University of Maryland, College Park, Maryland 20742 and E. A. Erongillo: Division of Nutritional Sciences, Cornell University, Ithaca, New York 14853, respectively. 176 NEW YORK ENTOMOLOGICAL SOCIETY Fig. lA. The influence of photoperiod on the average number of vitellogenic eggs in B. intermedia.* be effective biological control agents, in part, because ovigenesis and oo- sorption are not totally host dependent. This study details the role of pho- toperiod and temperature on oosorption in B. intermedia. Materials and Methods B. intermedia were reared on wax moth pupae {Galleria mellonella), a standard laboratory host. Upon emergence, adults were maintained in 30 x 30 cm cages at 16 hr L:8 hr D, 50% R.H. and 21.1°C for two weeks before being used in experiments. In the laboratory, the achievement of reproduc- tive maturity in females is variable but in general, females are mature after about 7 days. For each set of experimental conditions in each experiment groups of 80 females and 25 males were placed in a cage and provided with honey, distilled water and paper strips (resting sites). Data on the initial state of the female reproductive system (week 0) was obtained by removing 10 females for dissection. On each of 6 successive weeks 10 females were dissected to determine the state of ovarian devel- opment. Each dissection involved the removal of the intact ovaries. Both the total number of opaque (vitellogenic) eggs and the number of terminal * Each data point represents the average egg number of the population sample dissected for any given week. VOLUME LXXXVII, NUMBER 2 177 Ul < s Ul ee Ul a. lA O O III u Z o « o z u 6 z z < Ul s WEEK Fig. IB. The influence of photoperiod on the average number of chorionic eggs in B. intermedia.* oocytes which were larger than 0.33 mm in length were recorded. The final stages of oocyte resorption were characterized by the disappearance of lipid yolk spheres. In order to characterize more fully the state of oocyte devel- opment, ovaries dissected in the 10 hr and 16 hr photophase experiments (see below) were stained with trypan blue for 20 minutes at 23°C (Telfer and Anderson 1968). This dye is an accurate index of the change from normal vitellogensis to oosorption, based on alteration of the oocyte membrane. Week numbers refer to the number of weeks in which the wasps were kept under experimental conditions. Experiments were conducted in envi- ronmental control chambers which provided 50% R.H. and photoperiods and temperatures appropriate to each set of experiments. To demonstrate the influence of photoperiod on oosorption, adults were exposed to 21.1°C and the following photophases: 10, 11, 12, 16 hr. The influence of exposure to short photoperiod prior to adult reproductive ma- turity was also investigated. Wasps were reared in their pupal host and kept after adult parasitoid emergence for 2 weeks in a 12 hr photophase and 21.rC and compared to those kept in a 16 hr photophase and 21.1°C for the same period. Finally, three cohorts of adults were kept each at an 11 hr * Each data point represents the average egg number of the population sample dissected for any given week. 178 NEW YORK ENTOMOLOGICAL SOCIETY Eig. 2. The influence of temperature on the average number of vitellogenic eggs in B. intermedia.* photophase and at 15.5°C, 21.1°C or 26.6°C to investigate temperature ef- fects on oosorption. Analyses of variance and Duncan’s new multiple range tests were per- formed to determine significant differences in the data. Results and Discussions Oosorption in B. intermedia is affected by photoperiod. (Figs. lA, B). That oosorption ensued in adults placed in a short photophase (10, 11, 12 hr) compared to those in long photophase (16 hr) is demonstrated by the decreasing egg numbers through the six weeks of the experiments {P < 0.01). Adults kept in a long photophase showed a less dramatic decline in oocyte number compared to short photophase (10, 11 and 12 yr). The over- all trend in total number of oocytes (Fig. lA) is similar to that of number of chorionic eggs (Fig. IB). This concurrence is found in all subsequent experiments and thus, no other data on chorionic eggs are presented. From * Each data point represents the average egg number of the population sample dissected for any given week. VOLUME LXXXVII, NUMBER 2 179 Eig. 3. The influence of photoperiod on the average number of vitellogenic eggs in im- mature and young adult B. intermedia (<2 weeks old.)* weeks 0 to 6 the relationship between treatment means, for any given week, I varies. This is due, in part, to the fact that weekly dissections necessitate ! the use of a new cohort of adults each week. j Adult female parasitoids have been reported to partially absorb eggs in I the absence of hosts (Flanders 1950). Although greater resorption has been observed in older females, no quantitative study of the effect of age on resorption is yet available (Bell and Bohm 1975). Thus, although age (6 : weeks) and absence of hosts may enhance oosorption, short photoperiod I does have a direct effect on oosorption since adults at long photoperiod I were of comparable age and they experience a similar lack of hosts in our j experiments. Low temperatures enhanced the effect of short photophase (Fig. 2). Oo- cyte resorption proceeded more quickly at lower temperatures as indicated by comparison of mean values at each temperature {P < 0.01). This is sig- nificant, particularly, since physiological processes generally advance more I rapidly at higher temperatures. The two series of experiments discussed above demonstrate the sensitiv- ; ity of 2-week and older females to photoperiod and temperature. Exposure * Each data point represents the average egg number of the population sample dissected for any given week. 180 NEW YORK ENTOMOLOGICAL SOCIETY of immature stages of B. intermedia (within its host) and of young adults (<2 weeks old) to short daylength had marked consequences on their ovar- ian development (Fig. 3). Adults preconditioned at a 12 hr photophase before being placed in a 10 or 16 hr photophase generally had fewer oocytes with yolk at week 0 than those preconditioned at a 16 hr photophase (P < 0.01). Those adults placed in a short photophase (10 hr) had totally resorbed ovaries laden with fatty tissues by week 5. Those placed in long daylength (16 hr) maintained a higher mean no. of eggs per female compared to adults in a short photophase. Females are the overwintering stage of B. intermedia. Short-day photo- periods and low temperatures characterize pre-overwintering conditions, when few, if any, hosts are available to B. intermedia. Oosorption of oocytes under these conditions (short photophase and low temperature) would represent a logical adaptive mechanism to conserve nutrients critical to the overwintering (diapausing) female. Indeed in 1942, Flanders stated that in the parasitic Hymenoptera the occurrence of oosorption is an ad- aptation for maintaining the reproductive capacity when environmental con- ditions are unfavorable for oviposition. Acknowledgments We thank Dr. Marguerite Bohm for her assistance in conducting this study, William Metterhouse, New Jersey Department of Agriculture, Tren- ton and the USDA, APHIS laboratory at Otis, MA, USA forB. intermedia and gypsy moth pupae. Literature Cited Bell, W. A. and M. K. Bohm. 1975. Oosorption in insects. Biol. Rev. 50:373-396. Dowden, P. B. 1935. Brachymeria intermedia (Nees), a primary parasite and B. compsilurae (Cwfd.) a secondary parasite of the gypsy moth. J. Agric. Res. 50:495-523. D’Rozario, A. M. 1942. On the development and homologies of the genitalia and their ducts in Hymenoptera. Trans. Roy. Ent. Soc. 92:363^15. j Engelmann, E. 1970. The Physiology of Insect Reproduction. Pergamon Press, Oxford. , Flanders, S. E. 1942. Oosorption and ovulation in relation to oviposition in the parasitic j Hymenoptera. Ann. Ent. Soc. Am. 35:251-266. . 1950. Regulation of ovulation and egg disposal in the parasitic Hymenoptera. Can. i Ent. 82:134-140. Telfer, W. J. and L. M. Anderson. 1968. Functional transformations accompanying the ini- ^ tiation of a terminal growth phase in the cercropia moth oocyte. Devel. Biol. 17:512- 535. Department of Entomology, University of Massachusetts, Amherst, Mas- sachusetts 01003. Received for publication March 8, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(2), 1979, pp. 181-184 BOOK REVIEW C. L. Mandahar. Introduction to Plant Viruses. 333 p. 1978. S. Chand & Co., Ltd., New Delhi. 20 rupees ($3.00). This book contains 13 chapters on subjects relating to plant viruses. A brief, interesting introduction presents the discovery of viruses, their nature, and hypothesis of their origin. A chapter on isolation and purification gives a description of methods and contains useful information on salient features of virus concentration and purity. An entire chapter is devoted to modern tools and techniques, such as the electron microscope. X-ray diffraction, fluorescent antibody technique, animal and plant tissue culture, as well as assays with indicator plants. Virus structure and chemistry are well pre- sented and illustrated. Current knowledge on multicomponent, incomplete, and satelite viruses is brought up to date. Other chapters deal with disease symptoms, translocation and distribution, infection and replication, and variability. Entomologists will be most interested in the chapter dealing with transmission. Here the author discusses in detail the various groups of vec- tors, such as aphids, leafhoppers, whiteflies, mealybugs, beetles, and others, and the interactions between plant viruses and vectors. Harmful and ben- eficial effects on arthropod transmitters, transovarial transmission, and vec- tor specificity are all expertly described. A separate chapter is devoted to virus nomenclature and classification. The recently discovered spiroplasma and mycoplasma-like agents of plant diseases are also included, since they were earlier considered as plant viruses. The last chapter concerns various methods used to control plant virus diseases. A subject index and author index are provided. In addition to text illustrations, there are 28 pages of figures, containing electron micrographs of viruses, quite well reproduced on glossy paper. This is a very useful book, in which each section is up-to-date. The chap- ters are relatively free of errors, and provide a concise but thorough source of information. This hard-cover book can be recommended as a text for graduate students in agricultural colleges and universities. It is authoritative ' and at its price a real bargain! It might also find its way, as a source of information on plant viruses, to public libraries and it certainly belongs to I private libraries of plant virologists and plant pathology teachers. I Karl Maramorosch Waksman Institute of Microbiology Rutgers University P. O. Box 759 Piscataway, New Jersey 08854 182 NEW YORK ENTOMOLOGICAL SOCIETY BOOK REVIEW Biochemistry of Insects. Morris Rockstein, ed. Academic Press. 649 p. 1978. $29.50. This volume consists of 14 chapters, covering all aspects of insect bio- chemistry, the protein synthesis, insect pigments, cuticle, chemical genetics and the functional role of carbohydrates. The authors of this treatise are among the leading authorities in the diverse areas of insect biochemistry. G. Michael Chippendale gives an in-depth analysis of the function of car- bohydrates in insect life processes. Roger G. H. Downer discusses the fun- damental role of lipids, their digestion, transport, biosynthesis and endo- crine regulation of lipid metabolism. Moises Agosin covers the functional role of proteins, and P. S. Chen protein synthesis in relation to cellular activation and deactivation. The insect cuticle, sclerotization and melani- zation are the subject of a chapter by A. Glenn Richards. The role of insect biochromes is presented by A. E. Needham. Lynn M. Riddiford and her husband James W. Truman combined forces to give a lucid and stimulating presentation of the present status of insect hormones and growth regulators. Nevin Weaver covers the intraspecific chemical control of behavior, while Wendell L. Roelofs deals with pheromones. The biochemical defenses are the subject of Muray S. Blum’s exciting presentation. R. D. O’Brien gives an up-to-date analysis of the biochemistry of toxic action of insecticides. The detoxication mechanisms are described by W. C. Dauterman and Ernest Hodgson. The last chapter, by Francisco J. Ayala, is on chemical genetics and evolution. Each chapter includes a general reference list of pertinent books and reviews for advanced students and research scientists. There is an extensive subject index. The volume is unique in providing an authoritative and com- prehensive coverage of a critical interdisciplinary area. The quality of the contributions is uniformly high and exceptionally stimulating. The contrib- utors covered thoroughly all aspects of insect biochemistry and thus pro- vided a volume that will be indispensable for researchers, students, and teachers and will have lasting value. This outstanding book will be of interest to workers in entomology, agriculture, toxicology, pesticides, bio-organic chemistry, pollution research, ecology, and biochemistry. Karl Maramorosch, Waksman Institute of Microbiology , Rutgers Uni- versity. BOOK REVIEW The Life that Lives on Man. By Michael Andrews. Toplinger Publishing I Co., New York. 183 p. 1976. Paperback $4.95; cloth $9.95. This is a delightful book, and its selection by Library Journal as one of i the 100 best science books of 1977 certainly seems justified. The “life” VOLUME LXXXVII, NUMBER 2 183 includes viruses, bacteria, yeasts, as welf as mites, ticks, fleas, lice and bed bugs that live on our skin. The author has a good sense of humor as well as the ability to present the subject matter in a very readable manner. Il- lustrations include numerous scanning electron micrographs and excellent photographs of, among others, delousing, antityphus squads, and drawings of various insects. The photograph of the Apollo 12 on the moon illustrates the revealing finding by Surveyor III astronauts and Houston scientists that the camera, which had spent 2 years on the moon’s surface, was still con- taminated with bacteria from earth. Although this book is written for the layman, it will be of considerable interest to professional entomologists as well as to teachers and students in entomology departments. The book be- longs in personal and institutional libraries, as well as in public libraries. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni- versity. BOOK REVIEW Introduction to Insect Biology and Diversity. By Howell V. Daly, John T. Doyen, and Paul R. Ehrlich. McGraw-Hill Book Co. 564 p. 1978. $19.50. This handsome volume is a novel and modern approach to an introductory text in entomology. It is intended for students who have completed a basic course in biology. The lectures are devoted to insect structure and function, ecology, behavior, and applied aspects. There are keys for insect identifi- cation and the combined volume provides a foundation for professional entomological training. The first part of the book deals with insects as or- ganisms and the excellent illustrations help the beginner to digest the com- plex nature and diversity of insects. The second part on population biology follows an earlier book by Erhlich et al. (1974), giving a description of the evolutionary mechanism underlying the great variability of insects. The third part discusses insects in relation to the environment: insects of soil and water, relation of insects to plants, vertebrates, microbes and helminths, as well as biotic and physiological factors of the environment. The fourth part deals with insect diversity. A glossary, a list of references, and a taxonomic and subject index complete the volume. Throughout the book, discussions of biological phenomena are cross-referenced to more detailed treatments in various parts of the volume. Although the book has been written by three authors, it gives a uniform impression of a single-authored text. The pho- tographs, credited to a large number of renowned entomologists, are all excellent, and it can be expected that many teachers of introductory ento- mology will adapt this text for their courses. Occasionally the book departs from the traditional boundaries of entomology, but the final result is most 184 NEW YORK ENTOMOLOGICAL SOCIETY satisfactory. There has been a need for a modern, stimulating entomology text and the authors are to be congratulated for preparing this impressive, modern, up-to-date introductory volume. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers Uni- versity. Journal of the New York Entomological Society VOLUME LXXXVll SEPTEMBER 1979 NO. 3 EDITORIAL BOARD Editor Dr. Karl Maramorosch Waksman Institute of Microbiology Rutgers University New Brunswick, New Jersey 08903 Associate Editors Dr. Lois J. Keller, RSM Dr. Herbert T. Streu Publication Committee Dr. Randall T. Schuh American Museum of Natural History Dr. Daniel Sullivan Fordham University Dr. Felix J. Bocchino The College of Mt. Saint Vincent CONTENTS Dr. Charles Paul Alexander Henry M. Knizeski, Jr. 186-188 I Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, sub- I families of Pentatomidae and tribes of Pentatominae (Hemiptera) L. H. Rolston and F. J. D. McDonald 189-207 ' Positional variation and modifications relating to the protergum in Hymenop- tera Malkiat S. Saini, Surjit S. Dhillon and Tarlok Singh 208-215 Insects associated with weeds in the northeastern United States. II. Cinque- foils, Potentilla norvegica and P. recta (Rosaceae) S. W. T. Batra 216-222 Insects associated with weeds in the northeastern United States. III. Chickweed, Stellaria media, and stitchwort, S. graminea (Caryophyllaceae) S. W. T. Batra 223-235 The life histories of the autodice and sterodice species-groups of Tatochila (Lepidop- I tera; Pieridae) Arthur M. Shapiro 236-255 A reconsideration of the genus Bakeriella (Hymenoptera: Bethylidae) Howard E. Evans 256-266 Book Reviews 267-268 NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(3), 1979, pp. 186-188 DR. CHARLES PAUL ALEXANDER Charles Paul Alexander was born in Gloversville, New York, September 25, 1889, the son of Emil and Jane Parker Alexander. He entered Cornell University in 1909 to study entomology and received the Bachelor of Sci- ence degree in 1913, and the Doctor of Philosophy degree in 1918. While a student, he served as assistant, and later as instructor, in general biology and natural history. He was curator of the Snow Entomological Collection of the University of Kansas from 1917 until 1919. At the Illinois State Nat- ural History Survey, Urbana, he served as curator of the insect collections from 1919 until 1922. In 1922, he went to the then Massachusetts Agricul- tural College as assistant professor of entomology. He assumed charge of entomology in 1930 and was named chairman of the Department of Ento- mology and Zoology in 1938. He was Acting Dean of the School of Science from 1945 until 1946, when he was appointed Dean, a position he held until 1952. He retired from active teaching in 1959. Charles P. Alexander is a Eellow of the American Association for the Advancement of Science and a member of many leading entomological so- cieties in the United States and abroad. He is an Honorary Member of the New York Entomological Society. In 1942 and 1943, he served as president of the Entomological Society of America. In 1917, he married Mabel M. Miller of Brookview, New York. She has been his constant companion and helper, collecting insects, doing all the driving on field trips, and coauthoring papers. Charles P. Alexander is probably best known as an authority on one of the largest groups of insects, the crane flies (Diptera: Tipulidae). His interest in the group started in 1906, when he visited Dr. E. P. Pelt, the state ento- mologist of New York, for help in identifying some specimens of crane flies. The very conspicuous species were easily identified, but there was no one available to identify the rest. Encouraged by Drs. James G. Needham and Charles W. Johnson, who assured him that the family was ripe for study, Alexander has been studying the crane flies to this day. He has collected crane flies regularly and widely. The lure of opportunities for crane fly collecting has always been strong, and many field trips were taken, some as far as Alaska. A typical western trip covered more than 10,000 miles, lasted 80 days, and resulted in the collection of about 5,000 crane flies and more than 25,000 other insects for the University of Massachusetts collec- tions. He has assembled what is probably the most complete collection of any major group of insects. It includes almost 13,000 species of crane flies, many represented by type material, acquired from museums, collectors, expeditions, and his own field trips. Of particular importance is a reference collection of more than 50,000 microscope slides showing the various struc- VOLUME LXXXVII, NUMBER 3 187 Dr. Charles Paul Alexander, 1950. tures needed in taxonomic study of the group. Alexander has described over 10,000 species of tipulids and published more than 1,000 papers on them. After his retirement from active teaching, he transferred his crane fly collection to his home in Amherst, Massachusetts, where he had a special annex built to contain the collection. In “Cranefiy Haven,” as this annex is known, Alexander has continued his entomological research free from academic responsibilities. With his comprehensive knowledge and incom- parable collection of tipulids, he has accomplished a tremendous amount of taxonomic work in a relatively short time. But as he realizes perhaps more than anyone else, knowledge of crane flies is still incomplete. Despite his work in taxonomy, the majority of species are represented only by adults. 188 NEW YORK ENTOMOLOGICAL SOCIETY Dr. Alexander in “Cranefly Haven,” Amherst, Massachusetts. Top left: With reprints of his publications. Top right: At work. Bottom left and right: Charles and Mabel Alexander, 1979. and little is known about the ecology, behavior, and biology of all stages of most of the species. However, few other groups have such a strong taxo- nomic foundation upon which to build. In recognition of this great entomologist’s intensive efforts and impressive accomplishments over the past 68 years, this issue of the Journal of the New York Entomological Society is dedicated to Dr. Charles Paul Alex- ander on his 90th birthday. Dr. Henry M. Knizeski, Jr., Department of Natural Sciences, Mercy Col- lege, Dobbs Ferry, New York 10522. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIIO), 1979, pp. 189-207 KEYS AND DIAGNOSES FOR THE FAMILIES OF WESTERN HEMISPHERE PENTATOMOIDEA, SUBFAMILIES OF PENTATOMIDAE AND TRIBES OF PENTATOMINAE (HEMIPTERA) L. H. Rolston and F. J. D. McDonald ^ Abstract. — The families of Pentatomoidea of the Western Hemisphere (Acanthosomatidae, Canopidae, Corimelaenidae, Cydnidae, Cyrtocoridae, ; Dinidoridae, Megarididae, Pentatomidae, Phloeidae, Scutelleridae, Tessar- atomidae), the subfamilies of Pentatomidae (Asopinae, Discocephalinae, I Edessinae, Pentatominae, Podopinae), and the tribes of Pentatominae (Hal- , yini, Mecideini, Pentatomini, Sciocorini) are keyed and diagnosed. Edes- 1 sinae is raised from tribal status and Pantochlora Stal included in this I subfamily. All Western Hemisphere genera formerly included in Halyini except Brochymena Amyot and Serville are removed from this tribe. Car- I acid Stal, Marghita Ruckes and Janeirona Distant are transferred to Pen- tatomini. Janeirona is a senior synonym of Zimmerana Ruckes. I Identification and classification of many pentatomoids of the Western 1 Hemisphere have been hampered by the lack of a recent conspectus of the taxon. This contribution to such a conspectus may resolve some difficulties. Our opinions regarding the classification of Western Hemisphere penta- I tomoids are based on study of the external morphology and genitalia. The ' genital structure of suprageneric taxa has been characterized when sampling I for dissection was sufficiently broad that generalizations could be drawn i with confidence, and when such generalizations seem useful in defining a j taxon. We consider 11 taxa of the pentatomoids represented in the Western Hemisphere to merit family rank: Acanthosomatidae, Canopidae, Corime- laenidae, Cydnidae, Cyrtocoridae, Dinidoridae, Megarididae, Pentatomidae, Phloeidae, Scutelleridae, and Tessaratomidae. This classification is tradi- tional, although the canopids, corimelaenids, and megaridids are sometimes regarded as subfamilies of the cydnids. These 4 taxa were treated as sub- families of the Pentatomidae by McAtee and Mallock (1928, 1933) in the most recent revision of the megaridids, canopids, and corimelaenids. How- ever, the Pentatomidae in the classification of the above authors is equiv- alent to the Pentatomoidea as currently understood, and their subfamilies should be valued in context. Froeschner (1960) also excluded the canopids, corimelaenids, and megaridids from the cydnids in his revision of the Cyn- idae of the Western Hemisphere. Kormilev (1955) argued eloquently in favor 190 NEW YORK ENTOMOLOGICAL SOCIETY of family status for the cyrtocorids but did not formally elevate the taxon from subfamily rank. We recognize 5 subfamilies of Pentatomidae in the Western Hemisphere; Asopinae, Discocephalinae, Edessinae, Pentatominae, and Podopinae. The conspicuous departure from the usual classification is the elevation of the Edessinae from tribal status. The genus Pantochlora Stal, ejected by Kumar (1969) from the Tessaratomidae, is included in the Edessinae. The Pentatominae of the Western Hemisphere have long been distributed among the tribes Halyini, Mecideini, Pentatomini, and Sciocorini. No rea- son appears for disturbing this arrangement. We are not in accord with a proposal by Leston (1957) to raise the Mecideini to subfamily rank. Key to Families of American Pentatomoidea 1. Scutellum covering most of fore wings (Fig. 1) 9 - Scutellum leaving most of fore wings exposed even when scutellum attains apex of abdomen (Fig. 2) 2 2. Scutellum bearing large mesial spine or vertical plate Cyrtocoridae - Scutellum not so armed 3 3. Metathoracic scent gland orifice near lateral margin of pleuron (Fig. 3); antennae 3-segmented Phloeidae | - Metathoracic scent gland orifice distant from lateral margin of pleu- ron; antennae 4- or 5-segmented 4 4. Each pair of trichobothria on sternites iii-vii* on large callus lo- cated mesad of adjacent spiracle Dinidoridae - Trichobothria not on large callus, both (if paired) rarely mesad of , spiracles on all sternites 5 i 5. Pronotum extending over base of scutellum Tessaratomidae j - Pronotum ending at base of scutellum 6 i 6. Tibial spines if present confined to apex of tibiae 7 \ - Tibiae bearing many spines in addition to those at apex of tibiae and in addition to setae 8 i Fig. 1. Chelysoma scurrilis. Dorsum. Fig. 2. Moncus obscurus. Dorsum. Fig. 3. Phloea subquadrata. Scent gland orifice (s.a). Fig. 4. Ditomotarsus punctiventris. Terminal segments of male abdominal venter: sternite viii; pygophore (py). Fig. 5. Planois gayi. Pendergrast’s organs (pe). Fig. 6. Canopus burmeisteri. Spermatheca: spermathecal dilation (d). Fig. 7. Canopis orbicularis. Aedeagus: median penial lobe (m.p); ejaculatory duct (ej.d); conjunctival appendage (c.a). Fig. 8. Cyrtocoris sp. Spermatheca: spermathecal dilation (d). Sternite ii is the first visible sternite. VOLUME LXXXVII, NUMBER 3 191 viii 192 NEW YORK ENTOMOLOGICAL SOCIETY 7. Sternite viii exposed in males (Fig. 4); Pendergrast’s organs usually present in females (Fig. 5); tarsi 2-segmented Acanthosomatidae - Sternite viii concealed in males; Pendergrast’s organs absent in females; tarsi usually 3-segmented Pentatomidae 8. Fore tibiae usually expanded, sometimes cultrate with tarsi inserted midway of length, bearing lateral row of especially stout spines unless cultrate; apex of scutellum usually narrowly rounded Cydnidae - Fore tibiae subcylindrical, spines on lateral margin not notably larg- er than others; apex of scutellum broadly rounded Corimelaenidae 9. Tarsi 2-segmented Megarididae - Tarsi 3-segmented 10 10. Tibiae bearing many spines in addition to those at apex and in addition to setae, if weakly spined lateral margins of pronotum fringed with long setae Corimelaenidae - Tibial spines if present confined to apex 1 1 11. Sutures of abdominal venter complete, reaching lateral margins; second antennal segment much longer than diameter 12 - Sutures of abdominal venter obsolete laterad of spiracles; length of second antennal segment subequal to diameter Canopidae 12. Trichobothria paired; frena lacking Scutelleridae - Trichobothria single; short frena present Pentatomidae Acanthosomatidae Signoret, 1863 Acanthosomites Signoret, 1863, p. 549. Tarsi 2-segmented. Sternite viii exposed in males (Fig. 4). One or 2 pairs of Pendergrast’s organs usually present in females, these depressed, round or elongate, located laterally on sternites v-vii, vi-vii or sternite vii only, a single depression often extending across more than one sternite (Fig. 5). Scutellum subtriangular, about half or less than half as long as abdomen; frena extending along at least basal 7 tenths of lateral margins of scutellum. The genera of the Western Hemisphere were keyed by Rolston and Ku- mar (1974) and the family was revised by Kumar (1974). Canopidae Amyot and Serville, 1843 Canopides Amyot and Serville, 1843, p. 70. Canoparia Horvath, 1919, p. 205. Scutellum covering most of fore wings, leaving little more than exocorium exposed except at base. Tibiae setose but lacking stout spines. Sutures between abdominal sternites obsolete laterad of spiracles. Tarsi 3-segment- ed. Antennae 5-segmented; second segment subequal to its diameter. Body obovate, strongly convex, shiny black. VOLUME LXXXVII, NUMBER 3 193 Dilation of spermathecal duct discoidal. Spermathecal pump well devel- oped (Fig. 6). Conjunctival appendages of aedeagus well developed. Median penial lobes fused into trough beneath ejaculatory duct (Fig. 7). Canopus Fabricius, the only genus in this entirely American family, was revised by McAtee and Malloch (1928). Corimelaenidae Uhler, 1872 Corimelaenidae Uhler, 1872, p. 471. Thyreocoridae Van Duzee, 1907, p. 5. Eucoriinae Sailer, 1945, p. 133. Scutellum covering most of fore wings, leaving little more than exocoria exposed except basally, or U-shaped, broadly rounded apically, leaving most of fore wings exposed. Fore tibiae neither notably expanded nor cul- trate; tibiae bearing in addition to setae numerous spines along length; if only hind tibiae spined and weakly so, then coria and lateral margins of pronotum fringed with long setae. Anterior margins of middle and hind coxae bordered with dense fringe of short setae. Reticulation of eyes not attaining ventral surface of head. Tarsi 3-segmented. Antennae 5-seg- mented. In their revision of the corimelaenids, McAtee and Malloch (1933) rec- ognized 9 genera, all from the Western Hemisphere except one monotypic genus. The structure of the spermatheca and male genitalia varies considerably among genera of corimelaenids, suggesting that further consideration should be given to the suprageneric classification of this family. Family name. — There has been no concensus of opinion on the name of this family, and both Corimelaenidae and Thyreocoridae have been widely used. We believe that Corimelaenidae is the appropriate family name. Thyreocorides was proposed by Amyot and Serville (1843) as a supra- generic name for 6 genera {Chlaenocoris, Coptosoma, Heterocratis, Pla- taspis, Strombosoma and Thyreocoris). However, 7 of the 9 species listed under these genera are plataspids, including the 2 species assigned by Amyot and Serville to Thyreocoris: T. coccinelloides (Lap.) and T. punctatus (Leach). Another species is a canopid {Canopus impressus (Fabricius) = Chlaenocoris impressus). McAtee and Malloch (1933) left the remaining species, Strobosoma unipunctatum Amyot and Serville, unclassified although admitting its affinity to “Thyreocorinae.” The principal corimelaenid gen- era, Corimelaena and Galgupha, were placed by Amyot and Serville with the scutellerid genus Odontoscelis under their supra-generic name Odon- toscelides. It would seem therefore that Thyreocorides Amyot and Serville should be regarded as a synonym of Plataspidae since the majority of the 194 NEW YORK ENTOMOLOGICAL SOCIETY included species are of this family. Certainly it has little or no relevance to the corimelaenids. Corimelaenidae was introduced casually by Uhler (1872) in connection with notes on Galgupha nitiduloides (Wolff) and Corimelaena extensa Uhl- er, both of which he placed in the latter genus. Nevertheless, Uhler’s con- cept of the family clearly included its 2 large genera, and there is no evidence that it extended beyond the present concept of the family. Corimelaenidae appears to be the first family-group name relevant to this taxon that has been widely accepted. The argument for using Corimelaenidae for this taxon pivots on whether or not Thyreocoridae has been “generally accepted by zoologists interested in the group concerned,’’ a necessary requisite for availability (under Article 1 1 (e) iii of the International Code of Zoological Nomenclature) of family- group names originally published before 1900 and subsequently fully latin- ized. The persistence of a dual nomenclature for this family indicates to us the lack of general acceptance of Thyreocoridae, and the reason that this name should not become preeminent we believe to be compelling. Cydnidae Billberg, 1820 Cydnides Billberg, 1820, p. 70. Fore tibiae with row of stout spines along lateral margins, usually ex- panded, or cultrate with tibiae inserted midway of length. Middle and hind coxae thickly fringed with short setae along anterior margins. Reticulation of eyes reaching ventral surface of head. Scutellum exposing nearly all of fore wings. Tarsi 3-segmented, sometimes absent on hind legs. Antenna 4- or 5-segmented. Froeschner (1960) recognized 15 genera and 5 subfamilies in his revision of Western Hemisphere representatives of this family. Fig. 9. Cyrtocoris sp. Aedeagus, lateral view; conjunctival appendage (c.a); ejaculatory reservoir (ej.r); ejaculatory duct (ej.d). Fig. 10. Cyrtocoris sp. Aedeagus, ventral view: theca (t); conjunctival appendage (c.a); ejaculatory duct (ej.d). Fig. 11. Dinidor sp. Spermatheca: accessory spermathecal dilation (a.d). Fig. 12. Dinidor sp. Aedeagus, retracted: ejaculatory reservoir (ej.r); median penial lobe (m.p); ejaculatory duct (ej.d). Fig. 13. Dinidor sp. Aedeagus, expanded: theca (t); ejaculatory reservoir (ej.r); ejaculatory duct (ej.d). Fig. 14. Megaris laevicollis. Spermatheca. Fig. 15. Megaris constricta. Aedeagus. Fig. 16. Alitocoris parvus. Spermatheca: spermathecal dilation (d). Fig. 17. Edessa sp. Spermatheca: spermathecal dilation (d). VOLUME LXXXVII, NUMBER 3 195 I 196 NEW YORK ENTOMOLOGICAL SOCIETY Cyrtocoridae Distant, 1880 Oxynotides Amyot and Serville, 1843, p. 58. (part) Cyrtocorinae Distant, 1880, p. 43. Body covered with fine setae and waxy granules, the latter often fused. Scutellum extending to apex of abdomen, bearing mesial spine or vertical plate arising on basal half of scutellum and usually sloping dorsocaudad. Abdominal segments iii-vi projecting laterad well beyond coria, ii-v sepa- rated at lateral margins; sternites vi-vii laterally and genitalia nearly verti- cal. Tarsi 2-segmented. Antennae 5-segmented. Spermatheca terminating in C-shaped bulb, without flanges or obvious pumping mechanism (Fig. 8). Ejaculatory duct of aedeagus large; channel small, sinuous, enlarged distally (Figs. 9, 10). Ejaculatory reservoir ever- sible. Conjunctival appendages present. This wholly American family consists of 3 genera and 1 1 nominal species. Horvath (1916) revised the family and Kormilev (1955) added to this base in his treatment of the representatives occurring in Argentina. Dinidoridae Stal, 1870 Dinidorina StM, 1870, p. 79. Coridiinae Schumacher, 1924, p. 335. Each pair of trichobothria located on large callus mesad of spiracular line. Scutellum relatively small, about 4 tenths as long as abdomen, rounded apically. Antennae 4-segmented, distal 3 segments compressed, superior surface of segments 2 and 3 longitudinally impressed. Spermathecal duct with globular side dilation basally. Spermathecal pump well developed (Fig. 11). Ejaculatory duct of aedeagus fairly voluminous, membranous, surrounded basally by median penial lobes (Figs. 12, 13). Conjunctival lobes apparently absent. Base of ejaculatory duct together with complex spermathecal reservoir completely eversible. The above diagnosis applies only to the one American genus, Dinidor Latreille. Schouteden (1913), in his review of the world genera, listed 6 nominal species in Dinidor, one purportedly from Africa and the remainder from the Western Hemisphere. Megarididae McAtee and Malloch, 1928 Megaridinae McAtee and Malloch, 1928, p. 1. Body strongly convex, less than 5 mm long, usually less than 2 mm in length. Scutellum covering most of fore wings, coria little exposed except at base. Tarsi 2-segmented. Antennae apparently 4-segmented, with many setae as long as diameter of segments in female and much longer in male. VOLUME LXXXVIl, NUMBER 3 197 Spermatheca simple, saccular or globular (Fig. 14). Flanges and obvious pumping mechanism lacking. Aedeagus with membranous conjunctival lobes (Fig. 15). Ejaculatory duct tubular, simple. McAtee and Malloch (1928) revised the single genus, Megaris Stal, of this wholly American family. McDonald (1979) has reviewed the genitalia. Pentatomidae Leach, 1815 Pentatomidae Leach, 1815, p. 121. Scutellum leaving most of fore wings exposed, usually subtriangular, sometimes elongated and reaching apex of abdomen; disk flat, convex or gibbose basally. Pronotum terminating at base of scutellum. Tibial spines if present confined to apex. Tarsi usually 3-segmented, rarely 2-segmented. Trichobothria rarely located mesad of adjacent spiracle, if so not on large callus, usually paired but sometimes single. Metathoracic scent gland orifice distant from lateral margin of pleuron. Dilation of spermathecal duct fusiform, membranous, with central inva- gination around sclerotized tube (Figs. 16, 17). Spermathecal pump usually well developed, with distal and proximal flanges and variously shaped bulb. Ejaculatory reservoir of aedeagus not eversible. Median penial lobes pres- ent, sometimes much expanded (Figs. 18, 19, 20). Conjunctival appendages [ usually present. Key to Subfamilies of American Pentatomidae 1. Either first labial segment stout and extending well beyond bucculae, or fore tibiae foliate; pygophoral plate located entad of each para- mere (Fig. 25) Asopinae - First labial segment little enlarged, lying between bucculae (although often projecting beyond bucculae); fore tibiae not greatly expanded; pygophoral plate absent 2 2. Metasternum produced anteriorly onto mesosternum or rarely onto prosternum; rostrum not surpassing mesocoxae Edessinae I - Metasternum rarely produced anteriorly onto mesosternum, rostrum then extending onto abdomen; rostrum usually reaching at least to metacoxae 3 3. Trichobothrium nearest spiracle on sternite vii laterad of imaginary line tangential to spiracular openings on sternites vi and vii by dis- tance at least equal to greatest diameter of spiracular opening (Fig. 26) 4 - At least one trichobothrium on sternite vii in or near imaginary band connecting spiracles and projected caudad of spiracle on sternite vii, or mesad of band 5 198 NEW YORK ENTOMOLOGICAL SOCIETY Eig. 18. Uncinala tau. Aedeagus: median penial lobe (m.p); ejaculatory duct (ej.d); con- junctival appendage (c.a). Fig. 19. Edessa sp. Aedeagus: ejaculatory duct (ej.d); median penial lobe (m.p). VOLUME LXXXVII, NUMBER 3 199 4. Base of abdominal venter with mesial tubercle and metasternum produced, flattened (in part) Pentatominae - Abdominal venter rarely tuberculate at base, then metasternum thin- ly carinate mesially (in part) Discocephalinae 5. Labium arising on or behind imaginary line traversing head at an- terior limit of eyes and/or superior surface of 3rd tarsal segment of hind legs shallowly excavated in females (in part) Discocephalinae - Labium arising before such a line; superior surface of tarsal seg- ments convex or flattened 6 6. Trichobothria single; frena short, less than one-third length of scu- tellum; scutellum reaching apex of abdomen Podopinae - Trichobothria paired; frena one-third or more length of scutellum; scutellum not reaching apex of abdomen Pentatominae Asopinae Spinola, 1850 ' Spissirostres Amyot and Serville, 1843, p. 74. Asopoidae Spinola, 1850, p. 69 (1852). (Asopoideae intended.) Amyotinae Schouteden, 1906, p. 1. Arminae Bergroth, 1908, p. 180. Cimicinae Kirkaldy, 1909, p. 1. Pygophoral plate located entad of each paramere (Fig. 25). First labial segment usually stout and extending well beyond bucculae; if first labial segment of normal size and lying between bucculae for entire length, fore tibiae foliate. Males often bear pair of large pilose sensory patches, these usually extending across all or part of last 3 abdominal sternites. Tarsi 3- segmented. Antennae 5-segmented. Schouteden (1906) revised the world genera and listed the nominal species then known of each. I Fig. 20. Brochymena cariosa. Aedeagus: conjunctival appendage (c.a); ejaculatory duct ' (ej.d); median penial lobe (m.p). Fig. 21. Halys neelgirensis. Aedeagus: median penial lobe (m.p); conjunctival appendage i (c.a); ejaculatory duct (ej.d). : Fig. 22. Phloeci subquadrata. Genital plates; 9th paratergite (pt9); 2nd gonocoxae (2gx); 1st ; gonocoxae ( Igx). j Fig. 23. Phloea subquadrata. Spermatheca: spermathecal dilation (d); ring sclerites (r.s); ramus (r). Fig. 24. Phloea subquadrata. Aedeagus: ejaculatory reservoir (ej.r); conjunctival lobes I (c.a); ejaculatory duct (ej.d). 200 NEW YORK ENTOMOLOGICAL SOCIETY Discocephalinae Fieber, 1860 Discocephalidae Fieber, 1860, p. 26. Discocephalidum StM, 1867, p. 499. Labium usually arising on or posterior to imaginary line traversing head at anterior limit of eyes (Fig. 27). Trichobothria paired, one nearest spiracle on sternite vii usually laterad of imaginary band connecting spiracles and projected caudad of spiracle on sternite vii (Fig. 26). When labium arises before anterior limit of eyes, trichobothrium nearest spiracle on sternite vii laterad of spiracle by distance at least equal to greatest diameter of spirac- ular opening and/or superior surface of third tarsal segment of hind legs excavated in females. Metasternum not produced anteriorly onto mesoster- num. Tarsi 3 segmented. Antennae 4- or 5-segmented. Spermatheca typically pentatomid with central sclerotized tube in dilation of spermathecal duct and well developed pump (Fig. 16). Theca of aedeagus, ejaculatory duct, median penial lobes and conjunctival appendages (if pres- ent) heavily sclerotized, latter fused to margin of theca and permanently exserted (Fig. 18). All American genera formerly included in the Halyini are removed from this tribe except Brochymena. Most of these genera will form a tribe of the Discocephalinae, but Caracia StM, 1872, Marghita Ruckes, 1964, and Ja- neirona Distant, 1911 belong in the Pentatomini. The latter generic name is a senior synonym of Zimmerana Ruckes, 1962 (=Zimmeria Ruckes, 1958). Edessinae Kirkaldy, 1909 Edessini Kirkaldy, 1909, p. 153. Metasternum produced anteriorly onto mesosternum (onto prosternum in Pantochlora) and laterad between mesocoxae and metacoxae; posterior margin notched, receiving mesial tubercle of abdomen; anterior projection bifid (entire in Pantochlora). Rostrum terminating in anterior notch of me- tasternal projection (lying along projection in Pantochlora), reaching no farther than mesocoxae. Antennae 4- or 5-segmented. Tarsi 3-segmented. Spermatheca typically pentatomid with central sclerotized tube in dilation of duct and well developed pump (Fig. 17). Theca elongated, slightly curved dorsoventrally (Fig. 19). Median penial lobes small, heavily sclerotized, surrounding short ejaculatory duct except ventrally. Conjunctive inconspic- uous, sometimes apparent dorsally as small cone. Pantochlora vivida Distant, the sole species in the genus, is not remark- ably different from other members of the subfamily except in the form of the anterior metasternal projection. The genitalia, which Kumar (1969) has figured, conform to the diagnosis above. VOLUME LXXXVII, NUMBER 3 201 0’5m m 1-5 m m Fig. 25. Podisus maculiventris. Pygophore: ventral border (v.b); pygophoral plate (py.pl). Fig. 26. Dryptocephala obtusiceps. Abdominal sternite iv: trichrobothria (tr); spiracles (s). Fig. 27. Discocephalessa humilis. Ventral surface of head. Fig. 28. Euschistus tristigmus. Ventral surface of head. Fig. 29. Brochytnena quadripustulata. Membrane of hemelytra. Fig. 30. Brochymena parva. Head, dorsal view. 202 NEW YORK ENTOMOLOGICAL SOCIETY Pentatominae Leach, 1815 Pentatomida Leach, 1815, p. 121. Trichobothria paired, usually at least one of pair on each side of sternites iii-vii on or near imaginary band connecting spiracles on each side, pair rarely well mesad of spiracles. Metasternum rarely produced anteriorly onto mesosternum, if so rostrum reaching metacoxae. Basal segment of labium arising anterior to line traversing head at anterior margin of eyes, not es- pecially stout, lying entirely between bucculae or distal end surpassing buc- culae. Scutellum not attaining apex of abdomen, frena extending 4 tenths or more length. Dilation of spermathecal duct fusiform, membranous excepting sclero- tized rod around which dilation invaginates. Pump with proximal and distal flanges. Bulb spherical, digitiform or elongated and diverticulate. Conjunc- tival lobes usually large, usually bearing membranous appendages and sometimes sclerotized plates, in part or whole retractable into theca. Eja- culatory duct often extending beyond conjunctiva as sclerotized tube (pen- isfilum), this often looped or coiled. Key to Tribes of Pentatominae 1. Abdominal venter with a longitudinal band of striations on each side extending across basal three or more segments Mecideini - Striations absent on abdominal venter 2 2. Membrane of hemelytra bearing arborescent dark markings (Fig. 29); lateral jugal margins toothed preapically (Fig. 30) Halyini - Membrane of hemelytra immaculate or with streaks or dots; jugal margins unarmed or with several small teeth 3 3. Lateral margins of pronotum explanate and second antennal segment at least 1.5 times length of third segment Sciocorini - Lateral margins of pronotum rounded or carinate, if sublaminate second antennal segment less than 1.5 times length of third seg- ment Pentatomini Halyini Dallas, 1851 Halydidae Dallas, 1851, p. 150. Membrane of hemelytra with aborescent dark markings on frosty back- ground (Fig. 29). Preapical tooth present on lateral margin of juga (Fig. 30). Patches of waxy secretion evident on venter. Mesial sulcus extending over several basal sternites on abdomen broad, shallow. Antennae 5-segmented. In the Western Hemisphere only Brochymena Amyot and Serville is closely allied to Halys Fabricius. These two genera have similar features. I VOLUME LXXXVII, NUMBER 3 I 203 I including the genitalia (Figs. 20, 21). Leston (1953a) suggested that the tribe Halyini might be defined on the basis of wax glands. Wax glands are not unique to the halyines, although they may be a common characteristic of ' the tribe. j Brochymena has been revised by Ruckes (1946). * Mecideini Distant, 1902 Mecidaria Distant, 1902, p. 140. , Abdominal venter with a longitudinal band of striations on each side ex- I tending across basal 3 or more segments. Body elongate, linear. Antennae I 5-segmented, second segment triangular in cross-section. The only mecideine genus in the Western Hemisphere, Mecidea Dallas, ' has been reviewed by Sailer (1952). Pentatomini Leach, 1815 Pentatomidae Leach, 1815, p. 121. Lateral margins of pronotum usually rounded or carinate from lateral view, if explanate second antennal segment less than 1.5 times length of third segment, usually subequal or shorter. Orifice of metathoracic scent [ gland often attended by elongate ruga, if auriculate auricle and evaporative area not sunken below metapleural surface. Antennae usually 5-segmented, occasionally 4-segmented. Sciocorini Amyot and Serville, 1843 Sciocorides Amyot and Serville, 1843, p. 118. Lateral margins of pronotum explanate. Second antennal segment at least 1.5 times length of third segment. Metathoracic scent gland opening auric- ulate, auricle and small evaporative area often sunken below surface of pleuron. Antennae 5-segmented. Podopinae Amyot and Serville, 1843 Podopides Amyot and Serville, 1843, p. 56. Graphosomini Jakovlev, 1884, p. 204. Trichobothria single, one behind each spiracle in or near imaginary band connecting spiracles. Frena extending less than one-third length of scutel- lum. Scutellum reaching apex of abdomen, covering most of fore wings. Antennae 4- or 5-segmented. Tarsi 3-segmented. Pygophoral appendage (hypopygial appendage) often present, attached to posterolateral margin of pygophore. Barber and Sailer (1953) revised the North American representatives of this subfamily. 204 NEW YORK ENTOMOLOGICAL SOCIETY Phloeidae Amyot and Serville, 1843 Phleides Amyot and Serville, 1843, p. 115. Body extremely depressed; outer margins of juga, pronotum, coria basally and abdomen broadly foliate. Antennae and tarsi 3-segmented. Orifice of metathoracic scent gland near margin of pleuron (Fig. 3). Ninth paratergites greatly elongated (Fig. 22). Dilation of spermathecal duct sclerotized, cylindrical, striated (Fig. 23). Pump well developed, bulb sinuous. Ring sclerites and sclerotized rami present within vulva. Ejacula- tory reservoir of aedeagus eversible (Fig. 24). Channel of ejaculatory duct convoluted. Median penial lobes absent. The family consists of two American genera, Phloeophana Kirkaldy and Phloea Lepeletier and Serville, and 3 species. Leston (1953b) monographed the family. Scutelleridae Leach, 1815 Scutellerida Leach, 1815, p. 121. Odontoscelidae Douglas and Scott, 1865, p. 13. Eurygastridae Douglas and Scott, 1865, p. 13. Scutellum leaving little of fore wings exposed except basally. Erena ab- sent. Tibiae setose but without spines along length. Suture of abdominal venter extending to lateral margins. Antennae 5-segmented. Tarsi 3-seg- mented. Schouteden (1904, 1906) revised the world genera and enumerated the nominal species then known of each. Tessaratomidae StM, 1865 Tessaratominae StM, 1865, p. 33. Pronotum projecting over base of scutellum. Metasternum produced lat- erad between coxae and cephalad onto mesosternum, most strongly pro- duced as anterior wedge reaching nearly to procoxae, posterior margin transverse at junction with abdominal sternite. Six pairs of spiracles usually visible. The only genus represented in the Western Hemisphere, Piezosternum Amyot and Serville, contains 3 species. The genitalia of this genus have been figured by Leston (1954), McDonald (1966) and Kumar (1969). Acknowledgments H. Dodge Engleman, M.D., Dr. R. C. Froeschner (U.S. National Mu- seum) and Dr. Randall T. Schuh (American Museum of Natural History) contributed to this work by graciously loaning specimens. VOLUME LXXXVII, NUMBER 3 205 Dr. W. R. Dolling (British Museum (Natural History)) and Dr. Randall T. Schuh verified literature citations of papers unavailable to the authors. We are grateful for the constructive criticism offered by Dr. P. H. van Doesburg (Rijksmuseum van Natuurlijke Historie); H. Dodge Engleman, M.D., Dr. Jocelia Grazia (Universidade Estadual de Campinas), Dr. C. W. Schaefer (University of Connecticut), Dr. Randall T. Schuh, Prof. G. G. E. Scudder (University of British Colombia), and Prof. T. E. Woodward (Uni- versity of Queensland). I Literature Cited Amyot, C. J. B. and J. G. Audinet-Serville. 1843. Histoire naturelle des insects. Hemipteres. Paris. Barber, H. G. and R. I. Sailer. 1953. A revision of the turtle bugs of North America. J. Wash. Acad. Sci. 43(51:150-162. i Bergroth, E. 1908. Enumeratio Pentatomidarum post Catalogum Bruxellensem descriptarum. , Mem. Soc. Entomol. Belg. 15:131-200. I Billberg, G. J. 1820. Enumeratio insectorum in Museo G. J. Billberg. Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of the British Museum. London. Distant, W. L. 1880. Insects. Rhynchota, Hemiptera-Heteroptera. In Godman, F. D. and O. Salvin, Biologia Centrali-Americana. Vol. 1. London. . 1902. The fauna of British India, including Ceylon and Burma. Rhynchota, Vol. 1. Heteroptera. London. ' Douglas, J. W. and J. Scott. 1865. The British Hemiptera. I Fieber, F. X. 1860-61. Die europaischen Hemiptera. Vienna. ; Froeschner, R. C. 1960. Cynidae of the Western Hemisphere. Proc. U.S. Nat. Mus. Ill (no. ' 34301:337-680. Horvath, G. 1916. Revision Cyrtocorinarum. Ann. Mus. Natl. Hung. 14(1916):219-224. . 1919. Analecta ad cognitionem Cydnidarum. Ann. Mus. Natl. Hung. 17(1919):205- 273. Jakovlev, V. E. 1884. Hemiptera Heteroptera des astrachanischen Gebietes. Horae Societatis Entomologicae Rossicae. 18:141-243. Kirkaldy, G. W. 1909. Catalogue'of the Hemiptera (Heteroptera). Vol. 1. Cimicidae. Berlin. Kormilev, N. A. 1955. La subfamilia Cyrtocorinae Distant en la Argentina (Hemiptera, Pen- tatomoidea) Rev. Ecuat. Entomol. Par. 2(3/4):321-334, 3 pis. Kumar, R. 1969. Morphology and relationships of the Pentatomoidea (Heteroptera), III. Na- talicolinae and some Tessaratomidae of uncertain position. Ann. Entomol. Soc. Amer. 62(41:681-695. . 1974. A revision of world Acanthosomatidae (Heteroptera: Pentatomoidea): Keys to and descriptions of subfamilies, tribes and genera, with designation of types. Austral. J. Zool., Suppl. Ser. no. 34:1-60. Leach, W. E. 1815. Hemiptera. Brewster’s Edinburgh encyclopaedia. Vol. 9. Leston, D. 1953a. Notes on the Ethiopian Pentatomoidea (Hem.): XIII, On a waxy secretion by Halyini (Pentatomidae). Entomol. Monthly Mag. 89:147. . 1953b. “Phloeidae” Dallas: systematics and morphology, with remarks on the phy- togeny of “Pentatomoidea” Leach and upon the position of “Serbana” Distant. Rev. Brasil. Biol. 13:121-140. 206 NEW YORK ENTOMOLOGICAL SOCIETY . 1954. Notes on the Ethiopian Pentatomoidea (Hemiptera): XVII, Tessaratominae, Dinidorinae and Phyllocephalinae of Angola. Publ. Cult. Cia. Diam. Angola. 24:11-22. . 1957. Nomenclatural changes in Mecideinae (Hem., Pentatomidae). Entomol. Month- ly Mag. 93: 179. McAtee, W. L. and J. R. Malloch. 1928. Synopsis of pentatomid bugs of the subfamilies Megaridinae and Canopinae. Proc. U.S. Nat. Mus. 72 (art. 25): 1-21, 2 pis. . 1933. Revision of the subfamily Thyreocorinae of the Pentatomidae (Hemiptera-Het- eroptera). Ann. Carnegie Mus. 21:191-411, pis. 4-17. McDonald, E. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera-Het- eroptera). Quaest. Entomol. 2:7-150. . 1979. A new species of Megaris and the status of the Megarididae McAtee & Malloch and Canopidae Amyot & Serville (Hemiptera: Pentatomoidea). J. N. Y. Entomol. Soc. 87(1): 42-54. Rolston, L. H. and R. Kumar. 1974. Two new genera and two new species of Acanthoso- matidae (Hemiptera) from South America, with a key to the genera of the Western Hemisphere. J. N. Y. Entomol. Soc. 82(4):271-278. Ruckes, H. 1946. Notes and keys on the genus Brochymena (Pentatomidae, Heteroptera). Entomol. Amer. 26(4): 143-239. . 1958. New genera and species of neotropical discocephaline and halyine pentatomids. (Heteroptera, Pentatomidae). Amer. Mus. Nov. no. 1868:1-27. Sailer, R. I. 1945. The status of Corimelaena White, 1839, Eucoria Mulsant & Rey, 1865, and AUocoris McAtee and Malloch, 1933. Proc. Entomol. Soc. Wash. 47:129-135. . 1952. A review of the stink bugs of the genus Mecidea. Proc. U.S. Nat. Mus. 102 (no. 3309):47 1-505. Schouteden, H. 1904, 1906. Heteroptera. Earn. Pentatomidae. Subfam. Scutellerinae. Wyts. Gen. Ins., Ease. 24. pp. 1-98, 5 pis. Addenda et corrigenda (1906), pp. 99-100. . 1906. Heteroptera. Earn. Pentatomidae. Subfam. Asopinae (Amyoteinae). Wyts. Gen. Ins., Ease. 52. pp. 1-82, 5 pis. . 1913. Heteroptera. Earn. Pentatomidae. Subfam. Dinidorinae. Wyts. Gen. Ins., Ease. 153. pp. 1-19, 2 pis. Schumacher, F. 1924. Zwei ubersehene Hemipteren-Gattungen. Deutsch. Entomol. Zeitschr. (1924):335-337. Signoret, V. 1863. Revision des Hemipteres du chili. Ann. Soc. Entomol. France (4)3:541- 588, pis. 11-13. Spinola, M. 1850. Tavola sinottica dei genera spettanti alia classe degli insetti arthrodignati. Hemiptera Linn., Latr., Rhyngota Fabr. Rhynchota Burm. Modena, (not seen) reprinted 1852 Mem. Mat. Fis. Soc. Ital. Modena. 25:43-100, 1 plate. Stal, C. 1865. Hemiptera Africana. Vol. 1. Stockholm. . 1867. Bidrag till Hemipterernas Sytematik. Conspectus generum Discocephalidum. Ofver. K. Svenska Vet.-Akad. Forh. 24(7): 499-501. . 1870. Enumeratio Hemipterorum. Bidrag till en foreteckning ofver alia hittels kanda Hemiptera, jemte systematiska meddelanden. Enumeratio Dinidorinorum. K. Svenska Vet.-Akad. Handl. 9(l):79-89. Uhler, P. 1972. Notices of the Hemiptera of the Western Territories of the United States, chiefly from the surveys of Dr. F. V. Hayden. In Hayden, Preliminary report of the United States Geological Survey of Montana . . . pp. 392-423. Van Duzee, E. P. 1907. Notes on Jamaican Hemiptera: A report on a collection of Hemiptera made on the island of Jamaica in the spring of 1906. Bull. Buffalo Soc. Nat. Sci. 8:3-79. (LHR) Department of Entomology, Louisiana State University, Louisiana Agricultural Experiment Station, Baton Rouge, Louisiana 70803; and VOLUME LXXXVII, NUMBER 3 207 (FJDM) Department of Plant Pathology and Agricultural Entomology, Uni- versity of Sydney, Sydney, N.S.W., Australia 2006. Received for publication February 13, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIIO), 1979, pp. 208-215 POSITIONAL VARIATIONS AND MODIFICATIONS RELATING TO THE PROTERGUM IN HYMENOPTERA Malkiat S. Saini, Suijit S. Dhillon and Tarlok Singh Introduction The present work is a part of a general research project concerned with the tracing of evolutionary trends related to various body features within the entire range of the order Hymenoptera. This work specifically includes observations dealing with the gradual changes in the position of the proter- gum, which in higher Hymenoptera shifts backwards to become closely associated with the anterior margin of the mesothorax. The extent and mag- nitude of this association varies, and these variations form a systematic graded pattern which cannot be without evolutionary significance. This study also throws some light on phylogenetic relations among the different families of this insect order. Thus far no comprehensive study has been undertaken indicating the succession of these changes which affects the thoracic topography in Hymenoptera. In 1910 Snodgrass recorded some observations pertaining to the changes in the hymenopterous thorax as a whole, but his observations were based on only eight families. The following authors also referred to this thoracic component, but without discussing the magnitude of its shift from any comparative angle: Snodgrass (1910, 1925), Bird (1926), Reeks (1937), Duncan (1939), Alam ( 1951), Arora (1953), Rivard (1955), Bracken (1961), Tait (1962), Wong (1963), Dhillon (1966) and Mat- suda (1970). However, the present study which is based on observations made on the various hymenopterans belonging to about 20 different families of this insect order, does exactly this. Summary In Pamphiliidae and Xyelidae the protergum with its small antero-lateral angles is loosely attached to the mesotergum and the membranous areas prominently intervene between its posterolateral sides and the mesoepi- sternum. The protergum, however, in Tenthredinidae, Diprionidae, Argi- dae, Cimbicidae, Cephidae, Xiphydridae and Siricidae is extended at its anterolateral angles and is rather more closely attached to the mesotergum. The above mentioned membranous areas are also considerably reduced. The chalcidoidea present further advancement in this trend, in which case i the protergum is closely applied to the mesotergum and the membranous areas are mere interposing spots. In Ichneumonoidea the antero-lateral an- i gles of the protergum are further extended antero-ventrally giving the pro- < VOLUME LXXXVII, NUMBER 3 209 tergum a collar-like appearance which on its postero-lateral sides abuts against the adjoining margin of the mesopectus thus leaving no trace of intervening membranous areas. A similar condition prevails in the males of Formicidae and Mutillidae, but in the females the protergum is secondarily fused with the mesotergum and mesopectus. Another line of modification is represented by Chrysididae, Scoliidae, Vespidae and Eumenidae, where, besides the considerable extension of the antero-lateral angles of the pro- tergum, its postero-lateral angles also extend laterally in the form of spirac- ular lobes to cover the first pair of the spiracles. These modifications are further magnified in Sphecidae, in which the further extended antero-lateral angles of the protergum imparts to the latter an appearance of a pronounced collar. However, here the two opposite ends of the collar just touch but do not fuse. In Apoidea the opposite ends of the collar-shaped protergum fuse with each other and exist in the form of an entire annulus completely girdling the anterior margin of the mesothorax. Material and Methods To carry out the present study, most of the specimens of Apocrita were collected from the Punjab and Himachal Pradesh during the months of Sep- tember and October 1975 and preserved in 80% alcohol. Symphyta, with the exception of Megalodontidae and Orussidae, were supplied by the Bio- systematic Research Institute, Canada, and the Zoological Survey of India. As the specimens provided by them were dry, they were softened by im- mersion in 2% KOH for about 6 days. Diagrams were drawn with the help of a Stereoscopic microscope fitted with a graph eye-piece. OBSERVATIONS AND DISCUSSION (Figs. Plate I, II) The sequential separation of the protergum from the propectus and its complete integration with the anterior margin of the mesothorax is a char- acteristic feature of the order Hymenoptera. This disassociation probably was initiated among the ancestors of the suborder Symphyta but it reached its climax in the higher apocritans where the protergum seems to be more an integral part of the mesothorax than of the prothorax. In the females of the families Formicidae and Mutillidae, the protergum is completely fused with the mesothorax and even the line of fusion is quite obscure. This fact reveals a trend of considerable evolutionary significance. An effort has been made to work out in sequence all the stages of the shift as represented in the position of the protergum vis-a-vis the propectus and the mesotergum in extant families. The most primitive known condition is observed in the members of the families Pamphiliidae and Xyelidae. In Acantholyda maculiventris (Fig. 1) 210 NEW YORK ENTOMOLOGICAL SOCIETY Side view of the protergum and its association with the mesothorax. 1. Acantholyda maculi- ventris', 2. Xyela bakeri\ 3. Neodiprion abietis', 4. Arge clavicornis\ 5. Pristiphora cinctcr, 6. Cimbex americana americana ; 1. Cephiis (Cephus) cinctus \ 8. Xiphydria melUpes \ 9. Sirex cyaneus; association of the protergum with mesothorax and the front view of the protergum; VOLUME LXXXVII, NUMBER 3 211 10, 11. Sycoscapter stabilis', 12, 13. Netelia kashmirensis', 14, 15. Trachysphyrus sp.; 16, 17. Chrysis indogotea', 19. Scolia quadripustulata; 20,21. Vespa orientalis', 22, 23. Eumenes dimidiatepennis', 24, 25. Dorylus labiatus; 26, 27. Sima rufonigra', 28, 29. Camponotus ca- rnelinus', 30, 31. Scelephron intriidens', 32, 33. Slizus vespiformis . Side view of the protergum and its association with the mesothorax; 34. female Dorylus labiatus; 35. female Mutilla sp.; association of the protergum with mesothorax and the front view of the protergum. 36, 37. Xylocopa lemuiscapa . Abbreviations EPST — Episternum; INF — Inflection; MSAN — Mesalinotum; MR — Membrane or (Membra- nous); PCX — Procoxa; PPCT — Prepectus; PRPL — Propleuron; PT — Protergum; S — Spiracle; SL — Spiracular lobe; VB — Ventral bridge. 212 NEW YORK ENTOMOLOGICAL SOCIETY (Pamphiliidae) and Xyela baked (Fig. 2) (Xyelidae) the protergum is weakly attached to the anterior margin of the mesonotum and also there are present membranous areas between its sides and the mesoepisternum. On the whole, the association of the pronotum with the mesothorax is not rigid and in this case it does not seem to be an integral part of the latter. A similar condition has also been observed in Pamphilius luteicornis and Cephalcki provanchen (Pamphiliidae). These observations are further substantiated by the similar work of Rivard (1955) on Cephalcki marginata. Matsuda (1970) has also described an equivalent condition in Pamphilius luteicornis but with the difference that he has shown the presence of a narrow bridge which connects the protergum and the mesoepisternum. However, close obser- vation reveals that this bridge is not a sclerotic connection but is a forward extension of the mesoephisternum and which is distally in membranous connection with the protergum. Conditions are different in the rest of the higher symphytans. In Pristi- phora cincta (Fig. 5), Pachyprotasis brunetti and Tomostethus {Eutonios- lethus) assomensis (Tenthredinidae) Neodiprion abietis (Fig. 3) (Diprioni- dae), Arge clavicornis (Fig. 4) (Argidae), Cimbex americana americana (Fig. 6) (Cimbicidae), Cephas {Cephas) cinctus (Fig. 7) (Cephidae), Xiph- ydria mellipes (Fig. 8) (Xiphydridae) and Sirex cyaneus (Fig. 9) (Siricidae), the protergum is closely attached to the mesotergum and the membranous area between it and the mesoepisternum is reduced further. The antero-lateral angles of the protergum are drawn out in the form of a triangular extension reaching far up to the base of the procoxa. As such it seems to be a part of the mesothorax girdling the latter anteriorly. An almost similar condition has also been shown by Snodgrass (1910) in Arge sp., Tremex columba, Lygaenematus erichsonii. Bird (1926) in Hoplocam- pa halcyon, Reeks (1937) in Diprion polytomum, Arora (1953) in Diprion pini. Bracken (1961) in Anoplonyx luteipes, Tait (1962) in Perga affinis affinis, Wong (1963) in Pristiphora erichsonii and Dhillon (1966) in Athalia proximo . Members of the group Hymenoptera Parasitica show a further modifica- tion over the condition described up to this point. In Sycoscapter stabilis (Figs. 10, 11) (Torymidae-Chalcidoidea) the posterior margin of the proter- gum abuts against the mesotergum, thereby forging a tighter connection, and the membranous area between it and the episternum is further reduced. However, the ventral extensions of the antero-lateral angles of the pronotum are of the similar pattern as above. Similar observations have been made in the other chalcids studied including Walkerella temeraria and Micranisa pteromaloides (Torymidae), Sycophila decatomoides (Eurytomidae) and Blastophaga masoni (Agaonidae). Snodgrass (1910) has also shown a similar condition in the chalcids under his observation. In the members of the Su- perfamily Ichneomonoidea such as Netelia kashmirensis (Figs. 12, 13) and VOLUME LXXXVII, NUMBER 3 213 Tnichysphyrus sp. (Figs. 14, 15), further modifications to this effect are traceable. In them the ventral extension of the antero-lateral corners of the protergum is more pronounced. They reach far down on the sides, fitting into the space between the base of the procoxa and mesopleuron. Moreover the protergum is more firmly attached with the mesonotum than even in the case of the chalcids. Snodgrass (1910), Alam (1951) and Jonathan and Gupta (1973) have made similar observations. In the male representatives of the families Formicidae (Figs. 24, 25 and 26 to 29) and Mutillidae, conditions are similar to those which prevail in ichneumonids but in the females of Dorylus labiatus (Fig. 34) (Formicidae) and Miitilla sp. (Fig. 35) (Mutillidae), conditions are further modified. The posterior and postero-lateral angles of the protergum have become com- pletely fused with the corresponding anterior margin of the mesonotum, and this fusion is such that even the lines of fusion are not traceable. Studying only these insects, one can easily be misled to believe the protergum to be a part of the mesothorax. However, after comparing the female represen- tatives with the males of these species, it becomes quite clear that in females the fusion of the protergum with the mesonotum is only a secondarily ac- quired character. The propectus lies much ahead and looks entirely disas- sociated from the protergum. The second line of modification covers the rest of the families of suborder Apocrita. In Chrysis inclogotea (Figs. 16, 17) (Chrysididae), Scolia quad- ripustidata (Figs. 18, 19) (Scoliidae), Vespa orientalis (Figs. 20, 21) (Ves- pidae), Eamenes dimidiatepennis (Figs. 22, 23) (Eumenidae), the conditions resemble very much those of the ichneumons. In them the postero- lateral angles of the protergum extend to cover the spiracles and these ex- tensions can be compared with the spiracular lobe of the honeybee (Snod- grass, 1925). On the other hand, the antero-lateral angles of the protergum are almost equal to those seen in the ichneumons. Conditions are further modified in the members of family Sphecidae. In Scelephron intrudens (Figs. 30, 31), Stilus vespifonnis (Figs. 32, 33), the ventral extensions of the antero-lateral angles of the protergum are much more pronounced. These extensions, after flanking the sides, also extend on the ventral aspect of the prothorax so as to come close to one another along the mid-ventral line without actually fusing. Thus the entire pronotum ap- pears to form a collar skirting the mesothorax along its anterior boundary. Prothoracic spiracles in these cases are completely covered over by the well developed spiracular lobes which extend laterally in the postero-lateral an- gles of the protergum. In the case of Xylocopa lemuiscapa (Figs. 36, 37), a member of superfamily Apoidea, ventral extension of the protergum, after flanking the sides of the segment, not only lie close to one another but they actually fuse to form a ventral plate between the prosternum and the meso- sternum. In this way, the protergum forms an entire annulus which encircles 214 NEW YORK ENTOMOLOGICAL SOCIETY the mesothorax along its anterior margin. In this process it gets far removed from the propectus as well, with which it has only membranous connection. The propectus occurs in a much forward position with only its posterior tip lying just in the enclosure of the protergum. Similar conditions have also been noted in some other unidentified specimens of this superfamily. This observation is further substantiated by similar studies made by Snodgrass (1910) in honeybee and in Proctotrypes candatus. This stage probably rep- resents the most advanced stage of evolution as far as the disassociation of protergum from the propectus and the complete transformation of the for- mer from a dorsal plate into an annular sclerite are concerned. This evo- lutionary stage is also seen in the association of the pronotum with its lateral extensions covering the two first spiracles in the form of lobes. Acknowledgments The authors acknowledge the cooperation of the Biosystematic Research Institute, Canada, and the Zoological Survey of India, Calcutta, who sup- plied some of the insect specimens. Literature Cited Alam, S. M. 1951. The skeleto-muscular mechanism of Steenobracon deesae cam. (Bracon- idae, Hymenoptera) — an ectoparasite of sugarcane and juar borers of India. Part I, Head and thorax. Alig. Musi. Uni. Publ. Zool. (Ser.) Ind. Ins. Typ. 3(1);74 pp. Arora, G. L. 1953. The external morphology of Diprion pint L. (Symphyta, Hymenoptera). Res. Bull. E. Punjab. Uni., Hoshiarpur, Zool. 25:1-21. Bird, R. D. 1926. The external anatomy of the adult of Hoplocampa halcyon, Nort (Hym.- Tenthrid.). Ann. Ent. Soc. America 19:268-277. Bracken, D. F. 1961 . The external morphology of two eastern species of the genus Anoplonyx (Hymenoptera: Tenthredinidae) with special reference to Anoplonyx luteipes (Cresson). Can. Ent. 93:573-593. Dhillon, S. S. 1966. The morphology and biology of Athalia proximo (Tenthredinidae, Hy- menoptera). Alig. Musi. Uni. Publ. (Zool. Ser.) Ind. Ins. Type 7:165 pp. Duncan, C. D. 1939. A contribution to the biology of North American vespine wasps. Stanford Uni. Publ. Biol. Sci. 8(1):272 pp. Jonathan, J. K. and Gupta, V. K. 1973. Ichneumonologia orientalis. Oriental Insects Mono- graph No. 3. The association for the study of oriental insects % Department of Zoology, University of Delhi. Matsuda, R. 1970. Morphology and evolution of the insect thorax. Mem. Ent. Soc. Canada No. 76:431 pp. Reeks, W. A. 1937. The morphology of the adult of Diprion polytomum (Hartig). Canad. Ent. Orillia 69:257-264. Rivard, I. 1955. Contribution to the morphology of the pine web-spinning sawfly Cephalcia marginata Middlekauff (Hymenoptera: Pamphiliidae). Can. Ent. 87:382-399. Snodgrass, R. E. 1910. The thorax of the Hymenoptera. Proc. U.S. Nat. Mus. 39:37-91. . 1925. Anatomy and Physiology of Honeybee . McGraw-Hill Book Co., New York. Tait, N. N. 1962. The anatomy of the sawfly Perga affinis affinis Kirby (Hymenoptera: Symphyta). Aust. J. Zool. 10:652-683. VOLUME LXXXVIl, NUMBER 3 215 Wong, H. R. 1963. The external morphology of the adult and ultimate larval instar of the larch sawfiy, Pristiphora erichsonii (Htg.) (Hymenoptera: Tenthredinidae). Can. Ent. 95:897- 921. Zoology Department, Punjabi University, Patiala- 147002 (India). Received for publication February 27, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK3), 1979, pp. 216-222 INSECTS ASSOCIATED WITH WEEDS IN THE NORTHEASTERN UNITED STATES. II. CINQUEEOILS, POTENTILLA NORVEGICA AND P. RECTA (ROSACEAE)' S. W. T. Batra Abstract. — Eighty species of insects, including 15 crop pests and 8 pol- linators, are associated with Potentilla norvegica L. and P. recta L. in the northeastern United States. Among the pests are the strawberry root weevil, Otiorhynchus ovatiis (L.), and the strawberry aphid, Chaetosiphon fragae- folii (Cockerell). A biological control program using insects against these weeds would be difficult due to the close genetic relationship between straw- berries and cinquefoils, and probable consequent attractiveness of Fragaria to phytophagous insects that attack Potentilla. Introduction A survey of phytophagous insects and pollinators of the rough cinquefoil, Potentilla norvegica L. and the sulfur cinquefoil, P. recta L., was under- taken to determine the trophic niches occupied by North American insects. According to Werner and Soule (1976), there is no readily available infor- mation regarding insects affecting cinquefoils. The genus Potentilla consists of about 300 primarily Holarctic species with a wide range of polyploidy (Kohli and Denford 1977), of which there are over 50 species in North America, but only about five of these are weedy. Potentilla norvegica (2N = 70) is an annual, biennial or perennial of Hol- arctic origin with two subspecies: norvegica, being native to northern Eur- asia; and monspeliensis (L.) Asch. and Gr., which originates in North America (Hulten 1974). Potentilla recta (2N = 28, 42; Darlington and Wylie 1956) is a perennial of Eurasian origin that has become established primarily in northeastern North America (Erankton and Mulligan 1970). There are three North American varieties: sulphurea (Lam. and DC.) Peyr., obscura (Nestler) Koch., and pilosa (W.) Led. (Werner and Soule 1976). Both species spread by dispersal of achenes. In the northeastern and north central United States, cinquefoils are among the most important weeds in forage crops, lawns, pastures and hay (Dan- ielson et al. 1965). Although cinquefoils may be controlled by cultivation or ' This is the second publication in a series on native insects associated with introduced weeds in the northeastern United States. Other genera investigated are; I, Galinsoga (Environ. Entomol., in press); III, Stellaria (J. New York Entomol. Soc., in press); Hieracium , Galium, Galeopsis, Lychnis {=Melandrium), Cerastium, Sonchus, and Matricaria (in preparation). VOLUME LXXXVIl, NUMBER 3 217 application of herbicides such as 2, 4-D or Silvex (the mention of a pesticide in this paper does not constitute a recommendation of this product by the USDA) (Werner and Soule 1976), control of these weeds in strawberries is difficult (Anonymous 1976). Cinquefoils are closely related to strawberries and some species have recently been hybridized with them by breeders for strawberry improvement (Scott and Lawrence 1976; Barrientos-Perez 1976). Other beneficial uses of cinquefoils are as ornamentals, including a variety of P. recta, "Warrensii”; for tannins (Werner and Soule 1976); as forages (Chevtaeva 1975); and as a source of anti-bacterial chemicals (Makarenko and Chaika 1974). Materials and Methods Phytophagous insects and pollinators of P . norvegica and P . recta were collected in Virginia, Maryland, Pennsylvania, New Jersey and New York during three years (1975-1977), at 15 locations for each species. The insects were observed, then hand-picked, aspirated or netted from plants in the field, and any feeding damage was noted. These plants were cut or uprooted, placed in large plastic bags, removed in the laboratory, and examined and beaten against a white oilcloth to loosen clinging insects. The plants were then placed in large clean plastic bags with netting caps for development or emergence of additional insects. The bagged plants were kept in the labo- ratory for about a month or until they decomposed and insects ceased to emerge. Most surveyed cinquefoil was collected in vacant lots or weedy pastures with a mixed plant population, but some plants were in cultivated fields. Results and Discussion Phytophagous insects and pollinators associated with P. norvegica and P. recta are listed in Table 1. Some additional insects associated with the introduced species P. intermedia L. in New York are as follows: Pseudoc- cidae, the lampyrid beetle Pyropyga mimita Le Compte, the weevils, An- thonomus sp. nr. consimilis Say and Centorhynchus sp., and the leafhopper Graphocephala sp. Unidentified tortricid, geometrid, and noctuid (Hermi- niinae) larvae were collected on the native P. canadensis L. in Maryland. Cinquefoils benefit from cross pollination to set the abundant seed necessary for their propagation, but some species, including P. recta, may reproduce agamospermously (Werner and Soule 1976). In a survey of insects visiting flowers of various weeds. Mulligan and Kevan (1973) found that flowers of Potentilla are unattractive. However, Mitchell (1960, 1962) lists 45 species of Apoidea in 21 genera as visiting cinquefoil flowers. Table 1 includes ants, bees and syrphid flies as pollinators. Cinquefoils are hosts of yellows viruses (Surgucheva 1976), that may be transmitted to crops by some of the Ho- 218 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Insects associated with cinquefoil. Relative frequency: C, commonly collected at most locations; M, moderate abundance, collected at 3-5 locations; R, rare, only 1 or 2 spec- imens, or found at less than 3 locations; — , not collected. Plant parts affected: F, flower, L, leaf, S, stem, Rt, root. Remarks: P, pollen feeder; N, nectar feeder; V, probable vector to crops of yellows virus occurring in Potentilla', numbers refer to months when insects were collected. Relative frequency P. nor- P. Plant part vegicQ recta affected Remarks COLEOPTERA Bruchidae Bruchus brachialis F. R — F 8, Pest Cantharidae Chauliognathus sp. — M L, S 7 Chrysomelidae Alticine larvae R — L 8 Chrysomelid larvae R — L, S 7 Longitarsus sp. — R L, S 7 Siimitrosis ancoroides — R L, S 7 Curculionidae Calomycterus setarius Roelofs — R Rt 7, introduced Gymnetron pascuoruin (Gyllenhal) — R L, S 7 Hypera nigrirostris (F.) — R L, S 7 Oedophyrus hilleri (Faust) C — L, S 7, 8, introduced Otiorhynchus ovatus (L.) R — Rt 11, Pest Tychius picirostris (F.) C — L, S 7, 9, Pest Nitidulidae Brachypterolus pidicarius L. — R F 6, introduced also on strawberry Meligethes nigrescens Stephen R — F 7, Pest Scarabaeidae Popillia japonica Newman C — L 7, Pest, introduced DIPTERA Cecidomyiidae Mycodiplosis inimica (Fitch) — C L, S, F 8, feeds on rust spores Mycodiplosis thoracica (Fitch) — C L, S, F 8, feeds on rust spores Syrphidae Sphaerophoria contigua (Macquart) — M F 6, N, also predaceous larvae Sphaerophoria philanthus (Meigen) — R F 6, N, also predaceous larvae Toxomerus geminatus (Say) C F 6, 8, N, also predaceous larvae VOLUME LXXXVII, NUMBER 3 219 Table 1. Continued. Relative frequency P. nor- vegica P. recta Plant part affected Remarks Toxomerus marginatus (Say) — c F 6, 7, N, also predaceous larvae Berytidae Jalysus spinosiis (Say) nymphs — M L, F 6, Pest Miridae Lygus lineolaris C C F 7, Pest (Palisot de Beauvois) Psalliis sp. — R F 8 Tingidae Corythucha marmorata (Uhler) — R L, S 6 HOMOPTERA Aleyrodidae Aleurodicine sp. R — L 9 Aphididae Acyrthosiphon sp. R C L, S 6, 7 Aphid spp. nymphs R C L, S 6 Aphis sp. — R L, S 7, 8 Chaetosiphon fragaefolii C — L, S 7, 8, Pest (Cockerell) Macrosiphina sp. — C L, S 6, 8 Macrosiphum euphorbiae c c L, S 6, 7, 11, Pest (Thomas) Myzus persicae (Sulzer) M — L, S 1 1 , V, Pest Rhopalosiphum maidis (Fitch) C — L, S 8,11, Pest Schizaphis gmmimtm (Rondani) R — L, S 1 1 , Pest Cercopidae Cercopid nymphs M — L, S 6 Philaenus spumarius (L.) — c L, S 6, V, Pest Cicadellidae Agallia constricta Van Duzee M R L, S 6 Aphrodes bid net us (Schrank) — M L, S 7 Cicadellid nymphs M — L, S 7, 8 Deltocephaline nymphs M — L, S 8 Graphocephala versuta (Say) C — L, S 7, 8 Gyponana sp. nymphs M — L, S 6 Macrosteles fasdfrons (Stal) R — L, S 9, V, Pest Flatidae Anormenis sp. nymphs — R L 7 Membracidae Membracid nymphs Pseudococcidae — M L, S 6 Pseudoccocid nymphs M C Rt 7, 8 220 NEW YORK ENTOMOLOGICAL SOCIETY Table I. Continued. Relative frequency P. nor^ P. vegica recta Plant part affected Remarks HYMENOPTERA Apidae Apis tneUifera L. M F 8, N Eormicidae Lasiiis neoniger Emery _ C F 7, N Leptothorax sp. — R F 6, N Monomorium minimum (Buckley) C C F 6, N Halictidae Dialictus mitatus (Smith) M M F 8, N, P Dialictus uncinus (Sandhouse) — M F 7, N Halictus confusus Smith R — F 8, N Halictiis ligatus Smith R R F 6, 8, N Tenthredinidae Fenusini larvae R L 7 LEPIDOPTERA Blastobasidae Blastobasid sp. R Rt 6 Geometridae Eiipathecia sp. larvae R L, S 7, 8 Geometrid larvae — R L, S 7 Lycaenidae Lycaena sp. larvae R L, S 7 Microlepidoptera sp. R — Rt 8 Noctuidae Lacinipolia sp. larvae _ R Rt 7 Noctuid larvae M M L, S 6 Plathypena scabra (Fabricius) R — L, S 7 larvae Plusiine larvae R L, S 8 Pyrrhia umbria (Hufnagel) — M L, S 8 larvae Pyralidae Pyrausta sp. larvae R L, S 11 Tortricidae Platynota sp. larvae M R L, S 6, 8 Sparganothis sulphurana (F.) M C L, S 6, 7 larvae, adults Tortricid larvae M — L, S 11 ORTHOPTERA Gryllidae Oecanthus sp. nymphs R L 6 PSOCOPTERA Ectopsocidae Ectopsocopsis cryptomeriae R L 7 (Enderlin) VOLUME LXXXVII, NUMBER 3 221 Table 1. Continued. Relative frequency P. nor- P. vegica recta Plant part affected Remarks THYSANOPTERA Idolothripinae Apterous Winged form R R L, F L, F 7 6 Thripidae Frankliniella fusca (Hinds) Frankliniella tritici (Fitch) Sericothrips variabilis (Beach) Taeniothrips atratus (Haliday) Thripidae sp. Thrips tabaci Lindeman C C R R R C C C L, F L, F L L, F L, F L, F 6, 7, 8, Pest 6, 7, 8, Pest 11 8 7 6, Pest moptera listed in Table 1. They were also hosts of 15 crop pests, including two species that attack strawberries. These weeds were not severely dam- aged by the insects listed here. Cinquefoils were frequently attacked and damaged by rust fungi, which were eaten by cecidomyiid larvae. Due to the close genetic and physiological similarity between cinquefoils and strawberries, it may be difficult to locate specific biological control agents that will not also attack Fragaria . The initiation of a biological con- trol program for these weeds is therefore not highly recommended. I thank the following specialists for identifying many of the insects as- sociated with cinquefoils: D. M. Anderson, W. A. Connell, R. J. Gagne, J. L. Herring, J. P. Kramer, D. R. Miller, E. L. Mockford, K. O’Neill, D. R. Smith, M. B. Stoetzel, F. C. Thompson, D. M. Weisman, R. E. White and D. R. Whitehead, Systematic Entomology Laboratory, and cooperating sci- entists, AR/SEA, USDA, and R. C. Froeschner, U.S. National Museum, Washington, D.C. Anonymous. 1976. Strawberry Weed Survey. Review of Development Work in the Southwest Region. London, 149 pp. Barrientos-Perez, F. 1976. Development and exploitation of interspecific Fragaria-Potentilla amphiploids in strawberry breeding. Ph.D. thesis, Univ. Calif., Davis. 126 pp. Chevtaeva, V. A. 1975. Nutritive value of some species of Potentilla. Dokl. Akad. Tadzh. SSR 18:60-62. Danielson, L. L., W. B. Ennis, Jr., D. L. Klingman, W. C. Shaw, F. L. Timmons, J. E. Acknowledgments Literature Cited 222 NEW YORK ENTOMOLOGICAL SOCIETY Jernigan, J. R. Paulling and P. E. Strickler. 1965. A survey of extent and cost of weed control and specific weed problems. USDA, ARS 34-23-1:78 pp. Darlington, C. D. and A. P. Wylie. 1956. Chromosome Atlas of Flowering Plants. George Allen & Unwin, London. 519 pp. Frankton, C. and G. A. Mulligan. 1970. Weeds of Canada. Canada Dept. Agr. Publ. 948. 217 pp. Hulten, E. 1974. The Circumpolar Plants. II. Dicotyledons. K. Sven. Vetenskapsakad. Handl. 13(1):463 pp. Kohli, B. L. and K. E. Denford. 1977. A study of the flavonoids of the Potentilla pensylvanica complex in North America. Can. J. Bot. 55:476-479. Mararenko, N. G. and V. M. Chaika. 1974. Antibacterial activity of some members of the genus Potentilla of southeastern Altai in connection with their taxonomy and ecology. Rastit. Resur. 10:180-187. Mitchell, T. B. 1962, 1966. Bees of the Eastern United States. Vol. I, II. N. Carolina Agric. Exp. Stn. Tech. Bull. 141, 152:538 pp.; 557 pp. Mulligan, G. A. and P. G. Kevan. 1973. Color, brightness and other floral characteristics attracting insects to the blossoms of some Canadian weeds. Can. J. Bot. 51:1939-1952. ' Scott, D. H. and F. J. Lawrence. 1976. Strawberries. Pp. 71-97 in J. Janick and J. N. Morey (ed.). Advances in Fruit Breeding. Purdue Univ. Press. I Surgucheva, N. A. 1976. Etiology of yellow diseases in plants of wild flora. Dokl. Akad. Nauk. SSSR 226:225-226. Werner, P. A. and J. D. Soule. 1976. The biology of Canadian weeds. 18. Potentilla recta L., P. norvegica L. and P. argentea L. Can. J. Plant Sci. 56:591-603. Beneficial Insect Introduction Laboratory, Insect Identification and Ben- eficial Insects Introduction Institute, Agricultural Research, SEA, USDA, Beltsville, MD 20705. Received for publication April 26, 1979. I I I NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK3), 1979, pp. 223-235 INSECTS ASSOCIATED WITH WEEDS IN THE NORTHEASTERN UNITED STATES. III. CHICKWEED, STELLARIA MEDIA, AND STITCH WORT, 5. GRAMINEA (CARYOPHYLLACEAE) S. W. T. Batra Abstract. — 156 species of phytophagous and pollinating insects and 3 phy- tophagous mites are associated with the introduced Eurasian chickweeds, Stellaria media L. (Cyrillo) and S. graminea L. in the northeastern United States. Among these are 16 crop pests, including three known vectors of crop viruses that also infect 5. media. Pollinators are primarily Apoidea, Syrphidae and Formicidae. The collection of Tmetothrips suhapterus (Hal- iday) on 5. graminea represents the first Western Hemisphere record. Stel- laria media ranks among the ten most important weeds in the United States. A biological control program is possible, although it may be difficult due to the habit and habitat of this winter annual. A survey of phytophagous insects of the introduced common chickweed, Stellaria media (L.) Cyrillo, and stitchwort or little starwort, 5. graminea L. was undertaken to determine the trophic niches occupied by North American insects before initiating foreign exploration for possible biological control agents. Stellaria media is a weed associated with people in Europe since prehis- toric times (King 1966). It is now ubiquitous (Coquillat 1951), occurring in moist, fertile, disturbed soils throughout much of the world including the tropics (Hulten 1970), where it may occur as a winter annual during the cool season or at high elevations (Chopra et al. 1956; Cardenas et al. 1970). Chickweed is a very variable, polymorphic species (2n = 28, 40, 42, 44; Darlington and Wylie 1956; Blackburn and Morton 1957) with several eco- types (King 1966), subspecies and varieties (Hulten 1970). In the temperate zones it is a winter annual or biennial. The plants germinate in late summer, grow during fall, and remain green throughout winter, while surviving — 12‘^C temperature (due to high cell sugar content), and producing fertile seeds in the cleistogamous flowers (King 1966). Rapid growth and open pollination begin in early spring; most plants die before midsummer. The winter growth of S. media resembles that of arctic Stellaria spp. (Bell and Bliss 1977). This suggests that chickweed may be a post-glacial relict, perhaps originally occurring on nutrient-rich moraine deposits, later becoming associated with human refuse dumps, and as agriculture developed, invading fertile fields. Chickweed requires neutral soil pH (Buchanan et al. 1975), a high nitrogen level (King 1966), and is sensitive to phosphorous deficiency (Hoveland et 224 NEW YORK ENTOMOLOGICAL SOCIETY al. 1976). The small seeds are transported by ants (King 1966). They can remain viable while buried under 27.0 or 7.5 cm of soil for 32 or 58 years, respectively (Harper 1960). Optimal seed depth for germination is 0.5 to 1 cm; maximum is 2 cm; alternating temperature (20-30°C) is beneficial (An- dersen 1968; Thompson et al. 1977). Viable seeds in soil may reach densities of 12.5 million per hectare (4 Kg.) in pastures, and 24.6 million per hectare in arable areas (King 1966); chickweed seed is also abundant in marshland soils (Hunyadi and Pathy 1976). Individual plants may produce 2,200-2,700 seeds (Kavanagh 1974). Although Mulligan and Kevan (1973) found no in- sects visiting the small, white autogamous flowers of 5. media, Mitchell (1962, 1966) lists 34 bee species in 10 genera on Stellaria. Philipp (1975) found that seed setting in the self-fertile, arctic 5. longipes Goldie was enhanced by frequent visits by insects; the alpine species, 5. cerastioides L., is pollinated by empid, muscid and syrphid flies (Miiller 1881). Stellaria media flowers in early spring are an important food source for many bees, wasps and flies (Table 1); S. graminea flowers in summer also attract many insects (Table 1; Muller 1881). Common chickweed ranks among the ten most important weeds in the United States (Jansen et al. 1972), where it is widespread (Fernald 1970). It is a major pest in wheat, small grains, legume seed crops, and potatoes; in vegetables such as asparagus, legumes, root crops, greens, salad crops and cole crops; also in stone fruits, ornamentals, lawns, turf, hay and pas- tures (Vengris 1953; Jansen et al. 1972). Chickweed invades 26 crops in 40 states, particularly in the northeast and south; and it occupies over 2.8 million acres of cultivated cropland (Jansen et al. 1972). The rapidly growing plants in early spring effectively compete with crop seedlings for nutrients, water and light (Welbank 1963; Gibson and Courtney 1977); however, chick- weed may be used to suppress the growth of bindweeds in vineyards (Stalder et al. 1977). In Europe it invades overgrazed pasture (Haggar 1974), domi- nates recently uncultivated land (Covarelli 1976), and is abundant in crops such as winter and spring cereals, barley, wheat, oats, pulses, linseed and carrots (Granstrom and Almgard 1955). Common chickweed in row crops is controlled by various herbicides (Aldrich 1957; Gummesson 1976; Paro- chetti and Bell 1975). Stellaria graminea, a perennial and also native to Eurasia, occurs in grass- lands in north central and northeastern North America (Eernald 1970). In Quebec it occurs in cultivated fields, but it is more abundant in cereals, young and old meadows (Hamel and Dansereau 1949). In eastern Europe it is common in prairie (Hruska-Dell’Uomo 1976) and floodplain meadows (Shcherbach 1977). There are three European cytotypes: diploids (2N = 26); triploids (2N = 39) and tetraploids (2N = 52; Gadella 1977). The trip- loid plants are male-sterile and do not produce viable seed; although not previously reported outside the Netherlands (Gadella 1977), I often found VOLUME LXXXVII, NUMBER 3 225 Table 1. Insects and mites associated with StelUma. Relative frequency: C, commonly col- lected at most locations; M, moderate abundance, collected at 3-5 locations; R, rare, only I or 2 specimens or found at less than 3 locations; — , not collected. Plant parts affected: F, flower; L, leaf; S, stem; Rt, root. Remarks: P, pollen feeder; N, nectar feeder; V, vector of crop viruses. Numbers refer to months of collection. Relative frequency on S. S. gra- media minea Plant part affected Remarks ACARINA Tetranychidae Bryobia praetiosia Koch c L, S 4, 5, 10 Tetranychus sp. immatures R — L, S II Tetranychiis iirticae Koch R — L, S 1 1 , Pest COLEOPTERA Cantharidae Cantharis sp. R L, S 5 Chrysomelidae Disonycha sp. larvae c L, S 6, defoliates Oedionychis sp. larvae — M L, S 7 Phyllotreta sinuata R — L, S 6 (Stephens) Curculionidae Hypera sp. larvae M S 7 Idiostethus sp. — R S 7 Odontocorynus scutellum- — M F 6, P album (Say) Dermestidae Anthrenus scrophu- R F 7, P lariae (L.) Elateridae Conoderus bellus (Say) R Rt 5 Nitidulidae Glischrochilus quadri- R F 5 signatus (Say) Meligethes nigrescens M F 5, Pest (Stephens) COLLEMBOLA Entomobryidae Lepidocyrtus allegha- M S, Rt 4 neyensis Maynard Isotomidae Isotorna viridis Bourlet R S, Rt 4 Proisotoma minuta R — S, Rt 4 Tullberg Poduridae Xenylla grisea Axelson M S, Rt 4 DIPTERA Acalyptratae Acalyptrate sp. R L, S 7 226 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Continued. Relative frequency on S. S. gra- media minea Plant part affected Remarks Agromyzidae Agromyzid larva R L 7 Melanagromyza — R E 7 buccalis Spencer Anthomyiidae Hylemya platura M F 3, 4, N (Meigen) Pegotnya sp. larvae R L, S 10 Anthomyzidae Anthomyza sp. R F 6 Mumetopia occipitalis C — F 4, 6, N Melsheimer Calliphoridae Phormia regina R F 4, N (Meigen) Chloropidae Elachiptera erythro- M S, Rt 6 pleura Sabrosky Elachiptera nigriceps R S, Rt 6 (Loew) Hippelates dissidens M S, Rt 4 (Tucker) Monochaetoscinella R S 6 nigricornis (Loew) Olcella trigramma C S, Rt 6 (Loew) Oscinella carbonaria C S, Rt 4, 6, Pest (Loew) Oscinella melancholica M S, Rt 6 Beck Oscinella soror C R S, Rt 6, Pest (Macquart) Oscinella umbrosa R S, Rt 6 (Loew) Drosophilidae Drosophila busckii R S, Rt 10 (Coquillett) Orthocladiine larvae C S, Rt 6 Scaptomyza adusta M — S, Rt 3, 4, 6, 10 (Loew) Scaptomyza pallida C S, Rt 3, 4, 6, 10 (Zetterstedt) Otitidae Otitid larvae M — S 6 VOLUME LXXXVII, NUMBER 3 227 Table 1. Continued. Relative frequency on 5. S. gra- media minea Plant part affected Remarks Stratiomyidae Stratiomyid larvae Syrphidae M — s 6 Carposcalis obscurum (Say) C — F 4, N Eristalis arbiistorum (L.) — R F 6, N Eristalis dirnidiatus Wiedemann — C F 6, N Eristalis tenax (L.) — R F 6, N Helophilus fasciatus Walker — R F 6, N Helophilus latifrons Loew — R F 6, N Megasyrphus laxus (Osten Sacken) M F 6, N Metasyrphus ameri- canus (Weidemann) R — F 3, N Metasyrphus lapponicus (Zetterstedt) — M F 6, N Parhelophilus laetus (Loew) — R F 6, N Platycheirus quadratus Say R — F 3, N Sphaerophoria contigua (Macquart) C — F 11, N Sphaerophoria philan- thus Meigen R C F 4, 6, N Syritta pipiens (L.) — M F 6, N Syrphus rectus Osten Sacken — M F 6, N Syrphus torvus Osten Sacken — M F 6, N Syrphus vitripennis Meigen — R F 6, N Toxomerus gennnatus (Say) — R F 6, N Toxomerus rnarginatus (Say) C R F 4, 6, N Xylota hinei (Curran) Tachinidae — R F 6, N Epalpus signifer (Walker) M — F 3 Gonia sp. R — F 4 Gymnoclytia occidua (Walker) ■ R F 6 228 NEW YORK ENTOMOLOGICAL SOCIETY Table I. Continued. Relative frequency on 5. S. gra- Plant part media rninea affected Remarks HEMIPTERA Lygus lineolaris — (Palisot de Beauvois) Mirid nymphs C Pentatomidae Cosmopepla bimaculata — (Thomas) Pentatomid nymphs C HOMOPTERA Aphididae Acyrthosiphon sp. — Acyrthosiphum siber- M icum (Mordv.) Aphidius sp. C Aphidini nymphs M Aphis sp. C Aphis gossypii R (Glover) Dactynotus sp. — Hyalopterus pruni R (Geoffroy) Macrosiphini nymphs M Macrosiphum euphorbiae C (Thomas) Myzus sp. M Myzus persicae (Sulzer) C Rhopalosiphum maidis R (Fitch) Rhopalosiphum padi (L.) C Schizaphis graminum C (Rondani) Cercopidae Cercopid nymph R Philaenus spumarius R (L.) Cicadellidae Agallia sp. M Agallia constricta M (Provancher) Aceratogallia sangui- M nelenta Van Duzee Cicadellid sp. nymphs C Deltocephaline nymphs M R L, S, F 6 R L, S, F 6, 7, damage meristem R F 6 R L, S 7 R L, S 6, 7 — L, S 4, 6 L, S 4, 6 M L, S 6 — L, S 2 — L, S 9, Pest R L, S 7 — L, S 4 R L, S 6, 11 — L, S 4, 6, 9, Pest L, S 5 — L, S 2, 4, 5, 6, 7, 9, 10, 11, V, Pest — L, S 1 1 , Pest L, S 10, 11 — L, S 10, 11 R L, S 4, 9 C S 6, Pest L, S 6 — L, S 2, 4 — L, S 9 R L, S 6, 7, 9, 11 — L, S 7 VOLUME LXXXVII, NUMBER 3 229 Table 1. Continued. Relative frequency on 5. S. gra- media minea Plant part affected Remarks Doratura stylata (Boh.) — M L, S 7 Empoasca sp. R M L, S 9 Empoasca erigeron DeLong R — L, S 6 Gyponine nymphs HYMENOPTERA Andrenidae R L, S 6 Andrena spp. M M F 3, 4, 5, 6, 7, N Andrena carlini Cockerell C — F 4, N, P Andrena dunningi Cockerell — R F 6, N, P Andrena sigmundi Cockerell R — F 4, N, P Andrena viburnella Graenicher R — F 4, N Andrena vicina Smith C — F 4, N Calliopsis andreni- forrnis Smith Anthophoridae M F 7, N, P Xylocopa virginica (L.) Apidae M — F 3, N Apis mellifera L. M — F 4, N, P Bombas bimUculatus Cresson R — F 3, N Bombas ternarias Say — M F 7, N Bombas terricola Kirby Braconidae — M F 7, N Chelonas sp. Formicidae — R F 6, N Camponotas novebora- censis (Fitch) — R F 6, N Formica sabsericea Say — R F 6, N Leptothorax mascoram (Nylander) — M F 7. N Myrmica lobicornis fracticornis Emery — R F 6, N Prenolepis imparis (Say) M — F 4, N Tapinoma sessile (Say) Halictidae M M F 6, 7, N Dialictus lineatalus (Crawford) — M F 7, N Dialictas versatas (Robertson) R M F 4, N 230 NEW YORK ENTOMOLOGICAL SOCIETY Table 1. Continued. Relative frequency on 5. S. gra~ media minea Plant part affected Remarks Haliclus confusus Smith — C F 7. N Halictus rubicundus (Christ) — R F 7, N Lasioglossum forbesii (Robertson) Ichneumonidae R F 7, N, P Banchus flavescens Cresson — R F 6, N Exyston chlavatum (Cresson) Megachilidae R F 6, N Osmia cornifrons (Radoszkowskii) Pompilidae M F 4, N, introduced Anoplius sp. Vespidae — M F 6, 7, N Polistes fuscatus (F.) LEPIDOPTERA Arctiidae R F 3, N Diacrisia sp. larvae Coleophoridae M — L, S 9 Coleophora sp. larvae Gelechiidae — R L, S 6 Stomopteryx sp. Geometridae R — Rt 4 Eupathecia larvae R — L, S 6 Geometrid larvae R — L, S 9 Microlepidoptera adult Noctuidae — R S 7 Amphipyra pyramidoides Guenee R — L 5 Amathes badinodis (Grote) R — L, S 4 Eupsilia sp. larvae R — L, S 4 Euxoa sp. larvae R — L, S 5 Lacinipolia sp. larvae R — L, S 4 Noctuid larvae (1st instar) C — L, S 4, 5 Plathypena scabra (F.) larvae M — L, S 9 Plusiine larvae (1st instar) R — L, S 4 VOLUME LXXXVII, NUMBER 3 231 Table 1. Continued. Relative frequency on S. S. gra- media minea Plant part affected Remarks Pseudaplusia sp. larvae Pyralidae R — L, S 10 Udea rubigalis (Guenee) larvae Tortricidae M L, S 6, 9, Pest Sparganothis reticulana (Clemens) R — L, S 9 Sparganothis sulph- urana (F) R R L, S 7 Tortricid larvae ORTHOPTERA Gryllidae R L 6 Anaxipha nymph THYSANOPTERA Phlaeothripidae R L 4 Haplothrips leucan- therni (Schrank) Thripidae M F 5, Pest Anaphothrips obscurus (Mueller) M — L, S 4, 5, 6 Aptinothrips rufiis (Mueller) M — L, S 6 Aptinothrips stylifer Trybom — R L, S 6, Pest Chirothrips sp. — R L, S 6 Frankliniella fusca (Hinds) C — L, S 4, 6, 9, 10, 11 V, Pest Frankliniella tenui- cornis (Uzel) R — L, S 10, V, Pest Frankliniella tritici (Fitch) M M L, S 6, Pest Limothrips cerealium (Haliday) R — L, S 5 Taeniothrips atratus (Haliday) M — L, S 6 Thrips physapus L. R — L, S 6 Thrips tabaci Lindeman M — L, S 4, 10, Pest Tmetothrips sub- apt eras (Haliday) C L, S 6, European species, new record for New World 232 NEW YORK ENTOMOLOGICAL SOCIETY male-sterile plants (perhaps triploids) growing among normal plants at Tru- deau, N.Y. Light or temperature (10 to 30°C) alternation benefits germina- tion (Andersen 1968). This species, although common in meadows in upstate New York and New England, is much less weedy and invasive than S. media . Materials and Methods Phytophagous insects, mites and pollinators of S. media and S. graminea were collected at 41 and 13 locations, respectively, in Maryland, Pennsyl- vania, New York and Vermont during four years (1975-1978). The insects were observed, then they were hand-picked, aspirated or netted from the plants in the field, and any feeding damage was noted. The plants were then cut or uprooted, placed in large plastic bags, removed from the bags in the laboratory, examined, and beaten against a white oilcloth to loosen clinging insects. The plants were then placed in large, clean, clear plastic bags with netting caps for development and emergence of additional insects. The bagged plants were kept in the laboratory for about a month or until they decomposed and insects ceased to emerge. Stellaria media was collected from pastures, lawns, dumps, fallow land, forest edges, roadsides, and at edges of fields planted to soybeans, tomatoes and alfalfa. Stellaria graminea was collected from pastures, meadows, hayfields and roadsides. Results and Discussion Phytophagous insects and mites, and pollinators associated with chick- weed and stichwort are listed in Table 1. Relatively few insects were com- mon to both species of Stellaria ', this may be largely due to differences in plant habitat and seasonal occurrence, since most S. media was collected in spring to early summer in southern Maryland, and most 5. graminea was collected in mid- to late summer in northern New York. Stellaria media is an important reservoir of crop viruses, such as beet curly top, tomato spotted wilt (Miller et al. 1960), turnip mosaic (Citir and Varney 1974), beet mild yellows (Hartleb and Bauer 1977), nepoviruses (Hanada and Harrison 1977), carnation ringspot (Rudel et al. 1977), cu- cumber mosaic (Bruckart and Lorbeer 1976; Kazda and Hervert 1977) and others (Kavanagh 1974). It is probable that viruses and other pathogens overwinter in this plant, and in its seeds, to be transmitted to crops in the spring (Kavanagh 1974). Insects that are known to transmit these crop vi- ruses (Carter 1962), and that were collected on Stellaria are indicated in Table 1. Included in Table 1 are 16 species of crop pests harbored by S. media and 5. graminea and 57 pollinators, belonging primarily to the Apoi- dea, Syrphidae and Formicidae. Tmetothrips subapterus (Haliday), collect- ed on 5. graminea at Rew, Pennsylvania, represents the first record of this VOLUME LXXXVII, NUMBER 3 233 European monotypic genus in the Western Hemisphere. It lives in Stellaria galls and may be worthy of further investigation as a biological control agent (K. O’Neill, pers. comm.). Stellaria media is eaten to some extent by 35 species of North American wildlife (mainly birds, Martin et al. 1951), and it is palatable to livestock, with good caloric content in winter (Caspers 1977). However, in view of its importance as a major weed, its lack of close relationship to valuable plants, its exotic origin and consequent lack of stenophagy or dependence by North American wildlife, it may be worthwhile to begin a search for potential biological control agents in Eurasia. However, its winter annual growth habit and usual predominance in cultivated areas do not favor the applica- tion of biological control methods. Insects and pathogens that destroy the flowers and seeds of chickweed would probably be the most effective con- trol agents. Except for numerous Dysonycha sp. larvae that totally defoliated plants at one Beltsville location, and mirid nymphs that damaged the meristem (Table 1) native North American insects did not appreciably affect S. media, as was expected. This weed seems to be an important overwintering res- ervoir or winter food source for crop viruses and insect pests such as Myzus persicae (Sulzer). The flowers provide food for beneficial insects such as bees and syrphid flies during late fall and early spring when few other flow- ers are blooming. Acknowledgments I thank the following specialists for identifying many of the arthropods associated with Stellaria: D. M. Anderson, E. W. Baker, R. W. Carlson, W. A. Connell, R. J. Gagne, R. D. Gordon, A. B. Gurney, J. L. Herring, R. W. Hodges, J. M. Kingsolver, J. P. Kramer, P. M. Marsh, A. S. Menke, K. O’Neill, C. W. Sabrosky, D. R. Smith, T. J. Spilman, G. Steyskal, M. B. Stoetzel, F. C. Thompson, D. M. Weisman, R. E. White, D. R. White- head, W. W. Wirth, and D. L. Wray, Systematic Entomology Laboratory and cooperating scientists, AR7SEA, USDA, and P. J. Spangler, U.S. Na- tional Museum, Washington, D.C. Literature Cited Aldrich, R. J. 1957. Chickweed control in alfalfa. USDA Tech. Bull. 1174; 1-14. Andersen, R. N. 1968. Germination and establishment of weeds for experimental purposes. W. F. Humphrey Press, Geneva, N.Y. 1-236. Bell, K. L. and L. C. Bliss. 1977. Overwinter phenology of plants in a polar semi-desert. Arctic 30:118-121. Blackburn, K. B. and J. K. Morton. 1957. The incidence of polyploidy in the Caryophyllaceae of Britain and of Portugal. New Phytol. 56:344-351. 234 NEW YORK ENTOMOLOGICAL SOCIETY Bruckart, W. L. and J. W. Lorbeer. 1976. Cucumber mosaic virus in weed hosts near com- mercial fields of lettuce and celery. Phytopathology 66:253-259. Buchanan, G. A., C. S. Hoveland and M. C. Harris. 1975. Response of weeds to soil pH. Weed Sci. 23:473-477. Cardenas, J., O. Franco, C. Romero and D. Vargas. 1970. Malezas de Clima Frio. Internat. Plant Prot. Center, Oregon State Univ., Corvallis. 1-127. Carter, W. 1962. Insects in relation to plant disease. J. Wiley Publ. N.Y. 1-705. Caspers, N. 1977. Seasonal variations of caloric values in herbaceous plants. Oecologia 26:379-383. Chopra, R. N., S. L. Nayar and I. C. Chopra. 1956. Glossary of Indian Medicinal Plants. C.S.I.R. New Delhi. 1-330. Citir, A. and E. H. Varney. 1974. Common chickweed a major weed host of turnip mosaic virus in New Jersey. Proc. Am. Phytopathol. Soc. 1:134. Coquillat, M. 1951. Sur les plantes les plus comunes a la surface du globe. Bull. Mens. Soc. Linneene, Lyon (France) 20:165-170. Covarelli, G. 1976. Phytosociological study of vegetation on uncultivated hill land in central Italy. Riv. Agron. 10:249-260. Darlington, C. D. and A. P. Wylie. 1956. Chromosome Atlas of Flowering Plants. George Allen and Unwin, London. 1-519. Fernald, M. L. 1950. Gray’s Manual of Botany. D. Van Nostrand Co., N.Y.: 1-1632. Gadella, T. W. J. 1977. Cytotaxonomic studies in Stellaria graminea in the Netherlands. Proc. K. Ned. Akad. Wet. Ser. C Biol. Med. Sci. 80:161-170. Gibson, D. I. and A. D. Courtney. 1977. Effects of Poa trivialis, Stellaria media and Rumex obtusifolius on the growth of Lolium perenne in the glass house. Ann. Appl. Biol. 86:105-110. Granstrom, A. B. and G. Almgard. 1955. Studies on the weed flora in Sweden. Medd. Statens Jordbruksforsok 56:187-209. Gummesson, G. 1976. Weed control in peas and field beans. Swedish Weed Conf. 17:D28- D29. Haggar, R. J. 1974. Current research on weed problems associated with grassland intensifi- cation. J. British Grassland Soc. 29:77. Hamel, A. and P. Dansereau. 1949. L’aspect ecologique du probleme des mauvaises herbes. Bull. Ser. Biogeographie Univ. Montreal 5:1-41. Hanada, K. and B. D. Harrison. 1977. Effects of virus genotypes and temperature on seed transmission of nepoviruses. Ann. Appl. Biol. 85:79-92. Harper, J. L., ed. 1960. The Biology of Weeds. Blackwell Sci. Publ. Oxford. 1-256. Hartleb, H. and E. Bauer. 1977. The importance of field weeds for overwintering of the virus of mild yellowing of beet in the German Democratic Republic in the years 1972 to 1976. Arch. Phytopath. Pflanzenschutz 13:153-162. Hoveland, C. S., G. A. Buchanan and M. C. Harris. 1976. Response of weeds to soil phos- phorus and potassium. Weed Sci. 24:194-201. Hruska-Dell’uomo, K. 1976. The prairie vegetation in Moslavinia, Croatia, Yugoslavia. Acta Bot. Croat. 35:135-142. Hulten, E. 1970. The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vetenskapsakad. Handlingar 13:1—463. Hunyadi, K. and Z. Pathy. 1976. Weed seed infestation level in the marshland soils of the Keszthely area. Novenyvedelem 12:391-396. Jansen, L. L., L. L. Danielson, W. B. Ennis, Jr., P. A. Frank, J. T. Holstun, Jr., D. L. Klingman, J. R. Paulling, R. A. Wearne and A. S. Fox. 1972. Extent and cost of weed control with herbicides and an evaluation of important weeds, 1968. USDA-ARS Pub- lication H-1: 1-227. VOLUME LXXXVII, NUMBER 3 235 Kavanagh, T. 1974. The influence of herbicides on plant disease. II. Vegetables, root crops and potatoes. Sci. Proc. R. Dublin Soc. Ser. B 3:251-265. Kazda, V. and V. Hervert. 1977. Epidemiology of cucumber mosaic virus of glasshouse cucumber. Ochrana Rostlin 13:169-176. King, L. J. 1966. Weeds of the World. Interscience Publ., N.Y. 1-526. Martin, A., H. S. Zim and A. L. Nelson. 1951. American Wildlife and Plants. Dover Publ., N.Y. 1-500. Miller, P. R., F. Weiss and M. J. O’Brien. 1960. Index of plant diseases in the United States. U.S.D.A. Agriculture Handbook 165:1-531. Mitchell, T. B. 1962, 1966. Bees of the Eastern United States, Vols. I, II. N. Carolina Agr. Expt. Sta. Tech. Bull. 141:1-538; 152:1-557. Muller, H. 1881. Alpenblumen, ihre Befruchtung durch Insekten. Verl. W. Engelmann, Leip- zig. 1-522. Mulligan, G. A. and P. G. Kevan. 1973. Color, brightness, and other floral characteristics attracting insects to the blossoms of some Canadian weeds. Can. J. Bot. 51:1939-1952. ^ Parochetti, J. V., and A. W. Beil. 1975. Carbetamide on common chickweed in semi-dormant alfalfa. Proc. Northeastern Weed Sci. Soc. 61-62. Philip, M. 1975. Flower biology of Stellaria longipes. Canad. J. Genet. Cytol. 16:499-514. Rudel, M., G. Querfurth and H. L. Paul. 1977. Natural occurrence of carnation ringspot virus in Stellaria media (L.) Cyrill. from vineyards detected for the first time. Nachrichtenbl. j Deutsch. Pflanzenschutz. 29:59-60. Shcherbach, S. R. 1977. Ecological and phytocenotic analysis of the flora of meadow phy- tocenoses of the upper Berezina, USSR, floodplain. Vyestsi Akad. Navuk BSSR Syer Biyal Navuk 2:9-16. I Stalder, L., C. A. Potter and E. Barben. 1977. Recent experiences with integrated measures for the control of field and hedge bindweed in vineyards. Proc. EWRS Symp. Weed I Contr., Uppsala. 221-228. I Thompson, K., J. P. Grime and G. Mason. 1977. Seed germination in response to diurnal ! fluctuations of temperature. Nature 267:147-149. Vengris, J. 1953. Weed populations as related to certain cultivated crops in the Connecticut River Valley, Mass. Weeds 2:125-134. Welbank. P. J. 1963. A comparison of competitive effects of some common weed species. Ann. Appl. Biol. 51:107-125. Beneficial Insect Introduction Laboratory, Insect Identification and Ben- eficial Insect Introduction Institute, Agricultural Research, SEA, USDA, Beltsville, MD 20705. Received for publication May 8, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK3), 1979, pp. 236-255 THE LIFE HISTORIES OF THE AUTODICE AND STERODICE SPECIES-GROUPS OF TATOCHILA (LEPIDOPTERA; PIERIDAE) Arthur M. Shapiro Abstract. — The egg, larva, pupa and host relations of Tatochila autodice, blanchardii, sterodice macrodice, sterodice sterodice, vanvol.xemii and mercedis are described. These species occur in temperate to subantarctic Argentina and Chile. The sterodice species-group is very homogeneous and seems to be closely related to T. xanthodice from Colombia. The autodice group shows many divergences in the larva and pupa and is convergent in various characters with a number of Holarctic taxa. T. mercedis is some- what intermediate to the autodice group in certain characters. This is the third in a series of papers describing the preparatory stages of the Pierini of the Andean region. This tribe has undergone an extraordinary adaptive radiation, with eight endemic genera and about 40 species in non- tropical habitats, from northeastern Colombia to Tierra del Fuego. The biology of the genus Tatochila, sens, lat., was reviewed by Giaco- melli (1915) and Jorgensen (1916). The former prepared a hypothetical phy- logenetic “tree” based on adult characters, mostly superficial. At that time the early stages of only one species, T. autodice (Hbn.) had been described, and even these had not been illustrated. When the genus was revised by Herrera and Field ( 1959) little more was known of its biology. T. blanchardii Butl. and T. mercedis (Esch.), common Chilean species, were reared but not published; T. sterodice arctodice Stgr. was studied as a truck-garden pest in an unpublished Colombian dissertation (Alvarez and Delgado 1969; Bravo pers. comm.). T. autodice has been redescribed several times, al- ways unfigured, most recently by Margheritis and Rizzo (1965). Recent studies (Shapiro 1978a, b and unpublished) indicate that the An- dean Pierines are an important group for interpreting the historical biogeog- raphy and evolution of the regional biota. It thus becomes imperative to augment our knowledge of their biology. The descriptions which follow are based on field work in northern and central Argentina in 1977 and in south- ern Argentina in 1979 by the author, and in Chile in 1978 by Mr. C. V. Kellner. Alcoholic material will be deposited in the U.S. National Museum, Washington, D.C. and the University of California, Davis collections. Rear- ing was on continuous illumination at 25°C. Color descriptions in parenthe- ses refer to the system of Kornerup and Wanscher (1978). VOLUME LXXXVII, NUMBER 3 237 The T. autodice Species-Group As defined by Herrera and Field this group contains two species and one subspecies. The adult characters make it a natural and very distinct group within the genus. Both species occur at low to moderate elevations. Tatochila autodice (Hiibner) (Fig. 1) This is the common vacant-lot mariposa blanca of low elevations in northern and central Argentina. It colonizes in summer above 2000 m but does not seem to overwinter there. It is an occasional truck-garden pest, the gusano rayado de las cruciferas of Margheritis and Rizzo (1965). It was studied afield in the provinces of Tucuman, Cordoba, La Rioja, and Buenos Aires and in the Capital Federal (city of Buenos Aires) (26°49'S to 38°44'S). No geographic differences in morphology or development were observed. Ecologically, T. autodice is largely confined to highly disturbed situations within regions supporting desert, subtropical forest, and temperate subhu- mid to subarid grassland biomes from at least 17°S to 40°S. Its occurrence in the desert is limited to river bottoms and washes. It is only an occasional stray in the humid Andes in Rio Negro and Neuquen and does not normally breed at Bariloche. It is multivoltine throughout its range, with 5-6 broods in the northwestern provinces, about 4 at Buenos Aires, and perhaps 3-4 at Bahia Blanca. Winter is spent as a diapausing pupa. Diapause induction is under photoperiod-temperature control. Eggs or larvae have been collected from the following wild crucifers in various locations: Rapistrum rugosum (L.), Coronopus didymus (L.) Smith, I Cardaria draba (L.) Desv., Cardamine hirsuta L., Diplotaxis muralis (L.) DC., Rophanus sativus L., Brassica geniculata (Desf.) Ball (the most im- portant host in Cordoba), B. campestris L., 5. napus L., Sisymbrium of- ficinale (L.) Scop., S. irio L., Eruca sativa Hill, Conringia orientalis (L.) Dumort, and Nasturtium officinale R. Br. (all introduced from the Palaearc- tic Region); and Lepidium spicatum Desv., L. bonariense L., L. aletes Macb., Descurainia appendiculata (Griseb.) Schulz and D. argentina ' Schulz (all native but weedy). Giacomelli (1915), citing Berg and Burmeister, recorded T. autodice on the genera Cestrum (Solanaceae) and Medicago \ (Leguminosae). Neither has been confirmed, and captive larvae rejected Medicago sativa L. and M. hispida Gaertn. Adults are common in alfalfa fields and often visit the flowers in the company of the alfalfa pest Colias < lesbia (F.) and of Tatochila vanvolxemii (Capr.). Captive larvae do accept garden nasturtium, Tropaeolum majus L., and almost certainly use this indigenous genus (Tropaeolaceae) afield. 238 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 1. Tatochila autodice from central Argentina: 1, egg; 2, mature larva, dorsal view; 3, mature larva, lateral view; 4, mature larva, head capsule; 5, pupa, lateral view; 6, pupa, ventral view; 7, pupa, dorsal view. Descriptions Egg. — Erect, milk-bottle shaped, 1.20 x 0.35 mm, the chorion sculptured as figured, with 8-10 vertical and about 32 horizontal ribs. Laid singly on leaves, stems, or inflorescences; when on leaves about equally on upper or lower surface. Often laid on small rosettes or on isolated plants along walls, hedgerows, or roadsides. Light orange when laid, darkening to deep orange after 12-18 h, translucent blackish 12 h before hatching. Time to hatch, 3- 4 days. The larvae eat more or less of the chorion. VOLUME LXXXVII, NUMBER 3 239 Larva: first instar. — At hatch 1.6 mm long at rest. Dull yellow with a black head, becoming grayish-green with full pattern (see below) of longi- tudinal pale yellow stripes after feeding. Numerous black tubercles bearing pale hairs. Begins by excavating pits in buds, flowers, green fruit, or leaves. Length of instar, 2 days. Second instar. — After molt 5 mm. Slate gray with yellow stripes as fol- I lows: a faint mid-dorsal line; stronger subdorsal; and stigmatal. Head slate gray. Ocelli and true legs black; venter and prolegs greenish-gray. Head and ; body with many black tubercles falling into 3 size classes, surmounted by whitish hairs. Length of instar, 2-3 days. Third instar. — After molt 9.4 mm. Similar, the yellow dorsal line tending I to disappear; feeds openly near top of plant on inflorescences and leaves. 1 3-4 days. Fourth instar. — After molt 15.3 mm. As before, the mid-dorsal line often completely lost. 3^ days. Fifth instar. — After molt 25.4 mm, when full grown 34 mm. Head gray with many black tubercles, each bearing a whitish hair, and faintly mottled in orange. Body with the yellow dorsal line barely if at all visible within a broad dorsal stripe of gray (“grayish turquoise,” 24E3); below this are greenish-yellow (“primrose yellow,” 1A6) subdorsal stripes including two orange (6A6) dots and two large black tubercles on each segment; below I this a gray (24E3) stripe and below it a yellow (1A6) stigmatal stripe con- , taining two orange (6A6) dots, one anterior and one posterior to the spiracle; shading into blue-green (“greyish green,” 25D5) which continues across the venter, including the prolegs. Body with numerous black tubercles as shown, some conical, each bearing a short whitish hair. The anterior ends of all of the yellow stripes on the prothorax are orange and expand to form a narrow orange “collar” directly behind the head. Occasional aberrant larvae have been found in which the dark stripes are dark bluish green (“opaline green,” 25C6) or else purplish gray (“dull vi- olet,” 16D3) with the light markings all orange (6A6). The former seems to be genetically determined, while the latter is associated with parasitism by the Braconid Apanteles ayerzai Brethes. Mature larvae “stem” the hosts and sit on the bare stalk, where the striped pattern is highly cryptic. The final one or two fecal pellets are red. Length of instar, 6-7 days. Prepupa. — Vertical, head up, attached in usual Pierid fashion by the anal prolegs and a silken girdle around the thorax. The color does not change conspicuously (unlike the genus Euchloe of the Holarctic, with a similar larva). Unlike most Pierines, the mature larva does not wander before pu- pation and usually attaches itself to the host, either on the bare stem or (if the host is quite leafy) on the underside of a leaf. 18-30 hours. Pupa. — Length 24-25.5 mm, the female noticeably larger. Width at girdle 240 NEW YORK ENTOMOLOGICAL SOCIETY 5- 5.5 mm. Initially colored like the larva, assuming its final coloration in 4- 6 h. Qround light pinkish gray (“pinkish white,” 9A2) with numerous but inconspicuous black tubercles corresponding to those of the larva; dorsal midline strongly carinate on the thorax, with a series of black-tipped smaller keels anteriorly on the abdominal segments; frontal and supraocular prom- inences all the same size, black, 3 dorsolateral flaring prominences above the spiracles moderately developed, not marked with black; two rows of light orange dots, each with two to a segment, on either side of the dorsal midline and in line with the spiracles, corresponding to their larval positions although the yellow stripes containing them are essentially lost; eyes and appendage cases, including wings, light gray (7B1), unmarked except for a black spot at the end of the discal cell and tiny black dots along the outer wing margin. Spiracles black. Cremaster light gray. Thorax mottled dorsally with olivaceous gray (“brownish grey,” 6C2); frons and vertex creamy white, contrasting. The color scheme of the pupa is invariant and reminiscent of Pieris beckeri Edwards of western North America. It is probably a bird- dropping mimic. The shape is more slender and angular than T. xanthodice Lucas (Shapiro 1978b). Before eclosion the eyes, wings, and body are pigmented in that order. White is laid down 12-18 h before black. Meconium red. Length of instar 6- 10 days. Tatochila blanchardii Butler (Fig. 2) T. blanchardii is the ecological equivalent of autodice in Chile, from Tarapaca (20°S) to Cautin (39°S) and crossing the Andes into the Argentine provinces of Chubut and Rio Negro, including the Nahuel Huapi National Park and the Bariloche district, where autodice is a rare casual. In the Central Valley of Chile it is common in disturbed urban and agricultural habitats and breeds on the introduced Crucifers Rapistrum rugosum, Sis- ymbrium officinale, and S. orientate L. Herrera and Field (1959) and Pena (1975) suggest Tropaeolaceae as hosts, perhaps following Giacomelli (1915). Tropaeolum majus is accepted by the larvae. The species is multivoltine, with about four generations per year at San- tiago (September-May). At Bariloche, where it breeds on Brassica genic- ulata and Sisymbrium officinale, it has at least two generations with perhaps a partial third. Winter is spent as a diapausing pupa. The following descriptions are based on a composite of material from the Chilean Central Valley {leg. Kellner) and bred ex ovo from Bariloche. |j Descriptions Egg. — Similar to T. autodice, slightly smaller, 1.10 x 0.30 mm, with 8- | 9 vertical and about 32 horizontal ribs. Laid singly on stems, fruits, flowers, 1 VOLUME LXXXVII, NUMBER 3 241 Eig. 2. Tatochila blanchardii from Santiago, Chile: 1, egg; 2, mature larva, dorsal view; 3, mature larva, lateral view; 4, lateral view of seventh segment, showing major tubercles; 5, pupa, ventral view; 6, pupa, dorsal view; 7, pupa, lateral view. and indiscriminately on upper and lower surfaces of leaves. Usually laid on plants in at least partial shade. Light orange when laid, darkening to deep orange after 12-18 h, and to translucent blackish 12 h before hatch. 3 days. Larva: first instar. — At hatch 1.5 mm at rest. Yellow, with black head; becoming dark gray-green with full Tatochila pattern after feeding. Length ' of instar, 2-3 days. Second instar. — After molt 4.5 mm. Black, with a barely discernible yel- low mid-dorsal line; two strong subdorsal yellow stripes; shading into gray from the spiracles across the venter, including the prolegs. Head dark gray, ocelli black. Tubercles as in autodice but more prominent, bearing whitish hairs; the largest with dark hairs. Length of instar, 3 days. 242 NEW YORK ENTOMOLOGICAL SOCIETY Third instar. — After molt 7 mm. Similar; resting for long periods on the stem, where highly cryptic. Length of instar, 3 days. Fourth instar. — After molt 12.0 mm. As before. 3-4 days. Fifth instar. — After molt 19 mm, reaching 29.5 mm when full-fed. The most slender Tatochila larva yet reared. Mustard yellow (2A6) with very prominent large tubercles as shown, bearing dark hairs; smaller tubercles in two sizes, less conical, bearing light hairs. With a narrow but well-defined mid-dorsal yellow line and subdorsal stripes, about three times as wide and of the same color (2A6); dark stripes purplish-gray (13D2), containing a row of very large tubercles directly above the spiracles; pleural and ventral areas light grayish yellow, the venter slightly green-tinged, separated by a diffuse purplish-gray line. Spiracles black. Head gray with irregular yellowish mot- tling and many black tubercles; ocelli black. As in T. autodice, the larva eats all parts of the plant, “stemming” it. It rests low on the bare stem. The final one or two fecal pellets are red. Length of instar, 3-5 days. Prepupa. — Vertical, head up, as in T. autodice', at Bariloche commonly on the host. Length of prepupal period 18-30 h. Pupa. — Length 21 mm, width at girdle 6 mm (reared T. b. ernestae Her- rera, from extreme northern Chile, average only 19 x 4.9 mm but do not differ in color and pattern). Similar to T. autodice in color and pattern, but slightly stockier; with broad olivaceous gray (6C2) stripes above the spira- cles, dorsad of the flaring prominences, concolorous with the dorsal thoracic mottling. Meconium red. Length of instar, 7-12 days. The T. sterodice Species-Group This is a very homogeneous group of six named entities: T. sterodice with four subspecies, T. vanvolxemii, and T. mercedis. All of these are apparently allopatric (or in the case of T. s. macrodice and T. vanvolxemii, isolated by altitude). T. sterodice has the widest latitudinal range of any South American butterfly, from central Colombia (3°N) to Tierra del Fuego (55°S). The other species are confined to the temperate mid-latitudes. All lay orange eggs, which differ among themselves in the sculpturing of the chorion; all are more fusiform than in the autodice group. All have stocky larvae and pupae, relative to that group. The larvae are striped lengthwise in black and yellow, generally with complete Tatochila pattern (yellow mid-dorsal line, subdorsal stripes and a broken stigmatal line which may be reduced to pairs of spots, one anterior to and one posterior to, each spiracle). When mature they are somewhat lighter in ground color and have a pronounced resemblance to some Noctuid “cutworm” larvae, from which they differ in having many black tubercles in three sizes, bearing light hairs. The pupae are smooth, without the abdominal keels and with only rudi- VOLUME LXXXVII, NUMBER 3 243 mentary dorsolateral prominences, and resemble the pupa of T. xanthodice I except for the much longer proboscis case in this group. All the species are associated with weedy Crucifers in highly disturbed habitats, and are today mainly anthropophilic. No member of this species- group has been recorded on Tropaeolaceae to date. All known populations, except perhaps T. s. fueguensis from Tierra del Fuego, are at least bivoltine and overwinter as a pupa under photoperiod-temperature induced diapause (except T. s. arctodice, which breeds continuously). Tatochila sterodice Staudinger The four races of this Andean insect replace one another latitudinally and are associated with major physiographic-ecological regions: T. s. arc- todice in the northern Andean altiplano, T. s. macrodice in the puna, T. s. sterodice in the Patagonian Andes, and T. s. fueguensis in the subant- arctic forest region. The last two are probably clinal across southern Pata- gonia and the Isla Grande de Tierra del Fuego. The most distinct of the subspecies is arctodice Stgr., which is circumequatorial and uniquely in the group does not appear to diapause. Its biology was studied by Alvarez and Delgado (1969) and will not be repeated here. The other three subspecies all occur in Argentina; their early stages show no more differentiation than the adults. Tatochila sterodice macrodice Staudinger (Fig. 3) This subspecies was studied around Tafi del Valle, Province of Tucuman, in a dry-montane environment in one of the isolated ranges east of the Andes proper (27°S). It was found in vacant lot and roadside habitats, exploiting weeds such as Descurainia argentina, Brassica geniculata, and Lepidium sp. (? ruderale L.). In Salta and Jujuy it occurs above 3000 m (B. Mac- Pherson, R. Eisele, pers. comm.) and perhaps has native non-weedy hosts. The range extends to southern Peru, extreme northern Chile, east into Bo- livia and south in Argentina to Mendoza, where it may intergrade to the next. All records known to me are from above 1500 m. Males are vigorous “hilltoppers.” The behavior of this subspecies was described in detail by Jorgensen (1916). In northwestern Argentina its flight season is November to April, with two or three broods. Descriptions Egg. — Erect, fusiform, 1.15 x 0.40 mm, with about 16 vertical and 35 horizontal ribs. Bright orange when laid, scarcely darkening until 12 h before hatch when it becomes translucent blackish. Time to hatch, 5-6 days. The eggs are laid singly on stems, inflorescences, and cauline leaves of larger 244 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 3. Tatochila sterodice macrodice from Tafi, Tucuman, Argentina: 1, fifth instar larva, dorsal view; 2, fifth instar larva, lateral view; 3, mature larva, head capsule; 4, pupa, dorsal view; 5, pupa, ventral view; 6, pupa, lateral view. plants, and also on the upper surfaces of small rosettes. More than half the eggs seen at Tafi were laid on apical buds. Larva: first instar. — At hatch 1.4 mm. Pale yellow with dark brown head. After feeding olive green with the usual yellow pattern at first weakly, then more strongly indicated, with many black tubercles. The larva eats pits in soft tissue, preferably of buds. It eats its entire chorion. Time to molt, 2 days. Second instar. — After molt 4 mm. Black, with lemon yellow stripes in the VOLUME LXXXVII, NUMBER 3 245 I I full Tatocliila pattern; tubercles, ocelli and spiracles black; venter gray, i Length of instar, 2-3 days. I Third instar. — After molt 6.5 mm. Same pattern, with indistinct orange j spots at the anterior end of each subdorsal yellow stripe on each segment, and on either side of each spiracle. Mid-dorsal yellow line strong. When not feeding the larva sits along the stem. 3 days. Fourth instar. — Length after molt 10.5 mm. Subdorsal stripes less bold, the mid-dorsal one persistent. Length of instar, 4 days. Fifth instar. — After molt 18.5 mm, reaching 30-32 mm at maturity. Body I blackish gray (“medium grey,” lEl); mid-dorsal yellow line distinct, the I subdorsal stripes less so but containing an orange spot at the anterior, and I sometimes the posterior, end on each segment; a distinct orange spot an- j terior, and a less distinct one posterior to each spiracle; venter dull gray I (“pastel grey,” ICl). Tubercles in 3 size classes, black, bearing light hairs, j The stripes of ground color contain large squarish black spots anteriorly as I figured, on the dorsum of each segment; they occur throughout the species- I group. Larvae “stem” the host, sitting low On the leafless stalk, usually I head down. The last fecal pellets are red. Length of instar, 6 days. I Prepupa. — Formed at some distance from the host, the larva wandering I without feeding for 2-6 h. Time to molt, 14-27 h. ' Pupa. — 22.5 X 5.5 mm. Initially deep slate-green (“greyish green,” 25D5); within 8 h turning light pearly gray (“reddish grey,” 9B2); with numerous small black tubercles corresponding to their larval positioris, but inconspicuous; dorsal and subdorsal lines as in the larva, whitish, the latter j containing faint orange dots anteriorly on each segment; the orange am- phispiracular spots also faintly indicated on the abdomen. Head and ap- pendages— including wing-cases — ochreous (“greyish yellow,” 4B4) with variable but rarely extensive black filling between veins on the outer half, and a black discocellular dot. Attached vertically, head up, on dry weeds, fence-posts, walls, etc. Pigment sequence as usual; meconium red; time to hatch, 8-13 days. Some pupae of T. s. macrodice have a series of black dots anteriorly on the dorsal abdominal midline, corresponding to those in the autodice group. Tatochila sterodice sterodice Staudinger (Fig. 4) Cultures were established from localities near the northern and southern I extremities of the range of this subspecies: San Carlos de Bariloche, Rio ‘ Negro (41°08'S) and Rio Gallegos, Santa Cruz (51°37'S), the latter transi- ! tional lo T. s. fueguensis Field. No morphological differences were noted, ^ but Rio Gallegos stock grows more rapidly, its mean time from oviposition I to eclosion being 3-4 days shorter. Data given below are for Bariloche, I where the population is partially triple-brooded (October to March). At Rio 246 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 4. Tatochila sterodice sterodice from Bariloche, Argentina; 1, egg; 2, newly hatched larva showing primary tubercles and setae; 3, mature larva, dorsal view; 4, mature larva, lateral view; 5, lateral view of seventh segment, showing major tubercles; 6, pupa, dorsal view; 7, pupa, ventral view; 8, pupa, lateral view. Gallegos it is probably at least partially double-brooded (November-Feb- ruary). Winter is spent as a diapausing pupa. Although males “hilltop,” this subspecies is often found in areas of uniform low relief. The recorded hosts are numerous and all of Palaearctic origin: at Bari- loche: Rapistrum rugosum, Coronopiis didymus, Lepidium ruderale, L. perfoliatum, Raphanus sativus, Brassica genicidata, B. campestris, Sis- ymbrium officinale, and 5. irio. At Rio Gallegos: Cardaria draba, L. per- foliatum, R. sativus, B. genicidata, B. campestris, B. kaber (DC.) Wheel- er, 5. officinale, S. altissimum L., and 5. irio. There are no native Crucifers on the Patagonian coast, where sterodice is common in all the larger towns. VOLUME LXXXVII, NUMBER 3 247 Descriptions Egg. — As in the preceding, 1.12 x 0.37 mm, with about 18 vertical and 35 horizontal ribs, dull orange when laid, darkening slightly after 6-12 h, becoming blackish-translucent 8-12 h before hatch. Laid singly on buds and stems of flowering plants and commonly on leaves of rosettes. Time to hatch, 6 days. Larva: first instar. — At hatch 1.4 mm long. Pale yellow with brown head; after feeding olive green with the full yellow Tatochila pattern. Tubercles black. On flowering plants the larva consumes the buds; on rosettes it ex- cavates pits. Time to molt, 4 days. Second instar. — Length after molt, 3.9 mm. Olive green, the usual pattern yellow, with strong mid-dorsal and subdorsal lines and numerous black tu- bercles. Feeds by preference on flowers and young, green fruit. 3 days. Third instar. — After molt 7.2 mm. Blackish olive (“greyish green,” 28E5) I with the full yellow pattern, including distinct amphispiracular spots. 4-4.5 days. Fourth instar. — After molt 10.5 mm. As before, the dark blotches on the anterior portion of each dorsum more conspicuous; orange spots at the anterior end of each subdorsal stripe on each segment, and on both sides of the spiracles. Time to molt, 5 days. j Fifth instar. — After molt 17 mm, reaching 28 mm. Olivaceous gray-green I (“olive,” 3E6) with a well-defined mid-dorsal and pair of subdorsal lines, I all of about equal width, yellow; an orange dot within each subdorsal line ! at the anterior end of each segment; orange dots anterior and posterior to each spiracle (in about 40% of larvae these dots are yellow); subspiracular area and venter, including prolegs, greenish gray (1D2); head brownish-gray mottled with orange, tubercles and ocelli black. Body with many tubercles falling in three size classes, surmounted by fine pale hairs. The larva “stems” the plant and as usual sits lengthwise on the stem, head up or down. Last fecal pellets red. Time to molt, 7-8 days. Prepupa. — Wandering for 3-8 h before spinning. Vertical, head up. 18- 36 h. Pupa. — Length 20.5-22 mm, width at girdle 5-6 mm. Initially slate green (“greyish green,” 25D7), assuming its final coloration in 6 h. Colored ex- actly as in T. s. macrodice, generally with little black filling on the wing- cases. Dorsolateral prominences very weak. Frontal and supraocular prom- inences equal, black. Pre-hatch pigmentation sequence normal, meconium red. 8-15 days. This is the slowest-developing Tatochila yet reared. Tatochila vanvolxemii (Capronnier) (Fig. 5) Distributed in Argentina from the Province of Tucuman (27°S) to Rfo Negro and southern Buenos Aires (40°S) in subhumid to subarid grassland. 248 NEW YORK ENTOMOLOGICAL SOCIETY Fig. 5. Tatochila vanvolxemii from central Argentina: 1, egg; 2, mature larva, dorsal view; 3, mature larva, lateral view; 4, mature larva, head capsule; 5, pupa, lateral view; 6, pupa, ventral view; 7, pupa, dorsal view. its ecological range almost exactly matching that of the North American Pieris protodice Bdv. & LeC. which it also resembles phenotypically. It is very abundant in disturbed sites and has been recorded on both native and introduced Cruciferous weeds: Rapistrum rugosum, Coronopus didymus, Cardaria draba, Lepidium (several species), Descurainia appendiculata, Raphanus sativus, Brassica geniculata, Sisymbrium officinale, S. altissi- mum, and Enica sativa. Of these B. geniculata and the Lepidiums seem to be most often used. The species is 3-5-brooded in various parts of its range, VOLUME LXXXVII, NUMBER 3 249 overwintering as a pupa in photoperiod-and-temperature-induced diapause. , Jorgensen (1916) records it from early October to mid-May in Mendoza at 937 m. Primarily a lowland species, it does not appear to “hilltop.” Stocks were obtained from Cordoba and Capilla del Monte, Province of Cordoba (31°S) and Bahia Blanca, Province of Buenos Aires (38°44'S), near its northern and southern range extremities. No morphological or devel- opmental differences were found. Both stocks are polymorphic for a number i of adult and larval characters. Descriptions j Egg. — Usual form, 1.2 x 0.33 mm, when laid light yellow to yellowish- orange, usually but not always darkening to deeper orange after 6 h (the difference appears to be hereditary through the female line); with 12-15 vertical and 34-37 horizontal ribs. Becomes opaque and hyaline 12 h before hatch. Laid singly on buds, fruit, stems or leaves. This species produces ! very large numbers of eggs per female, sometimes over 600 (Margheritis i and Rizzo, 1965 quote a figure of 4-500 for T. autodice). Time to hatch, 3- ' 4 days. Larva: first instar. — At hatch 1.6 mm. Dull olivaceous-yellow, head ; black. Feeds preferentially on flowers, but will eat pits in leaves; becoming j gray-green with full yellow pattern. 2 days. , Second instar. — 3.3 mm after molt. Slate gray, with full yellow pattern , and numerous black tubercles. Time to molt, 2-3 days. Third instar. — Length after molt 9.5 mm. Slate gray with the usual yellow pattern of a mid-dorsal line and subdorsal lines, about twice as broad; a I yellow spot anterior and posterior to each spiracle, the size and shape of i these spots variable; four blackish blotches on the anterior dorsum on each I segment, as illustrated; head gray, mottled with dull yellow, ocelli and tu- I bercles black. Venter and prolegs gray. True legs and tubercles black, the t latter in three size classes, bearing light hairs. Feeds on all parts of the [ plant. 3-4 days. Fourth instar. — 15.5 mm. Head gray, with a yellow cast. Yellow amphi- spiracular spots frequently connected to form a zigzag stripe. Faint orange spots occasionally present anteriorly in the subdorsal lines on each segment. The mid-dorsal line weak but still present. 3-4 days. Fifth instar. — After molt 25 mm, reaching 33.5 mm. As before, the ground color becoming dingy mouse-brown (“greyish brown,” 5D3), producing a very cutworm-like effect, with the black dorsal blotches strongly contrast- ing; the spiracular spots joined in a line in about 65% of the larvae, otherwise variously shaped, sometimes partly or all orange. As usual the larvae “stem” the host and sit on the bare stalk. The last fecal pellet or two are red. Time to molt, 5-7 days. 250 NEW YORK ENTOMOLOGICAL SOCIETY Prepupa. — Attached in the usual manner, vertical, head up, after a wan- dering period of 4-6 h or more. Duration of prepupal period, 12-20 h. |; Pupa. — 21.5 X 5.6 mm. At first slate green (“greyish green,” 25D7), with- in 6 h colored exactly as in T. s. macrodice \ black filling on wing cases variable, rarely pronounced. Black dots on dorsal abdominal midline almost always present and conspicuous. Dorsolateral prominences very weak. Frontal and supraocular prominences equal, black. Eyes, wings and body become pigmented in that order; white is laid down 30 h, black 12 h, before eclosion, which is almost always within 30 min of sunrise. Meconium red. Length of instar 6-9 days. Wild pupae occur on walls, fence posts, rubbish, and dry weeds and have not been found on the host. If T. vanvolxemii breeds on any of the Cap- paridaceous shrubs which occur in its range it might pupate on them. Tatochila mercedis (Eschholtz) (Fig 6) This small species is the most aberrant in the group. It occurs in Medi- terranean Chile from Atacama to Llanquihue (27°-41°S), generally in highly disturbed habitats in agricultural or waste ground on valley floors and in foothill canyon bottoms. It is multiple-brooded (August-April at Santiago; five broods?) and winters as a diapausing pupa. Its recorded hosts are Rap- istrum rugosum, Raphanus sativus, Brassica campestris. Sisymbrium offi- cinale and 5. orientate. All of these are European, and Pena (1975) predicts that a native Chilean host will be found. T. mercedis is occasionally recorded at Bariloche, Argentina (Herrera and Field 1959) and — obviously in error — in subtropical Misiones, north- east Argentina, by Breyer (1938). Giacomelli (1915) recorded it also from Bolivia, but there are no recent records from there. | Descriptions | Egg. — Similar to the others in this species-group but slightly smaller, ij 1.05 X 0.27 mm, with about 16 vertical and 33 horizontal ribs. Dull orange j when laid, deepening in color after about 6 h; translucent blackish 12 h 1 before hatch. Laid singly, mostly in inflorescences, but also on leaves of i small rosettes. Time to hatch, 3^ days. Larva: first instar. — Dull yellow with black head; 1 .05 mm at hatch. Feeds i preferentially on buds, flowers or young fruit, but will excavate pits in leaves. Becoming gray-green with the usual yellow pattern after feeding; time to molt 2 days. Second instar. — After molt 2.3 mm. Black, with the following yellow markings: a weak mid-dorsal line; a stronger, wider subdorsal stripe on each side; spots anterior and posterior to the spiracles. Head dark gray. Venter and prolegs dull greenish gray. Tubercles, prolegs and ocelli black. Length of instar, 3 days. (The spiracular spots may be orange.) VOLUME LXXXVII, NUMBER 3 251 Fig. 6. Tatochila mercedis from central Chile: 1, newly hatched larva showing primary tubercles and setae; 2, mature larva, dorsal view; 3, mature larva, lateral view; 4, lateral view of seventh segment, showing major tubercles; 5, pupa, ventral view; 6, pupa, dorsal view; 7, pupa, lateral view. Third instar. — After molt 4.2 mm. Pattern as before. Feeds in the open on the upper portion of the plant. Length of instar, 3-4 days. Fourth instar. — Length 8.2 mm. Pattern as before, the dark blotches on the anterior dorsum of each segment becoming distinct. Tubercles conspic- uous, black. 2-3 days. Fifth instar. — 15.5 mm, reaching 23 mm. Somewhat lighter, body gray- black (“medium grey,” lEl) with distinct black dorsal blotches as shown; usual pattern primrose yellow (1A6), the amphispiracular spots variable, sometimes connected in a zigzag line as in T. vanvolxemii, the dorsal midline 252 NEW YORK ENTOMOLOGICAL SOCIETY Still visible but narrow. Venter dull gray (“pastel grey”, ICl). Head brown- ish-gray with a yellow cast. Tubercles, ocelli, and true legs black. The large tubercles are more conical in this species than in the others of the group, suggesting T. blanchardii. Habits as in the other species, “stemming” the plant. Last fecal pellets red. Time to the prepupa, 6 days. This description applies to larvae reared on slender Crucifers. When reared on heart leaves of cabbage (Brassica oleracea L.) lighter in color: ground brownish gray (7D3), the yellow markings paler and more extensive, making the tuberculation more contrasting. Prepupa. — Head up, attached in the usual manner. Length of prepupal period, 14-20 h. Captive larvae wander for 3-6 h before spinning. Pupa. — 20 X 4.5 mm. Distinctively slender, but the dorsolateral promi- nences still weak. Frontal prominence often less blackened than the supra- oculars, but of similar size. At first slate green (“greyish green,” 25D7), but within 6 h turning buffy gray (“greyish orange,” 5B3); wing-cases scarcely contrasting but sometimes slightly ochreous, without black filling between the veins; dorsal and subdorsal yellow lines preserved, along with numerous but inconspicuous black tubercles; spiracles black; a few black points along veins and wing margins, but little or no dark discal spot. Eyes, wings, and body pigmented in that order, white appearing 20 h, black 8-12 h, before eclosion. Meconium red. Length of instar, 8-11 days. Discussion Both species-groups recognized by Herrera and Field ( 1959) on adult char- acters are cohesive in their immature stages, as well. The larval coloration of Talochila seems very conservative. Within the sterodice group the differences among species are scarcely greater than those between subspecies. The most distinctive member of the group is mercedis, and its characters seem to bear on the evolutionary affinities between the autodice and sterodice groups. Giacomelli (1915) derived the entire genus from autodice, making "dheod- ice" (actually blanchardii) a very early offshoot, "’volxemi" (yanvolxemii) next, then the sterodice group, even while admitting that “one must say in truth that volxerni is of doubtful placement and, more than all the other Tatochila, is of uncertain affinities.” Giacomelli did not know the genitalic characters used by Herrera and Field (1959) in placing vanvolxemii solidly into the sterodice group (they found that the male genitalia of sterodice and vanvolxemii were indistinguishable). The early stages unambiguously support this judgment and at the same time suggest that the member of the group closest to autodice and blanchardii is not vanvolxemii but mercedis. The dorsal pattern of both sexes of autodice and of the male blanchardii is a reduction of the full Tatochila pattern (still expressed in female blan- VOLUME LXXXVIl, NUMBER 3 253 chardii, and in males of ssp. ernestae). Of the other Tatochila, mercedis and some individual sterodice show the closest approximation to autodice dorsally. On the ventral hindwing, autodice and blanchardii almost always have an isolated spot in interspace Sc + R,. Elsewhere in the genus this spot appears frequently in mercedis and occasionally in sterodice and no- where else. In both it is usually connected to the vein. It represents pigment deposition in an aborted vein trace and may imply a phylogenetic connection between the autodice group and mercedis-sterodice . (This must be inter- preted with caution; the same spot appears, obviously independently, in some stocks of Nearctic Pieris napi (L.)!) This hint is borne out in the larvae and pupae of mercedis — in the prominence of conical subdorsal tubercles on the larva, suggestive of blanchardii, and in the slender pupal shape. T. mercedis is confined to Chile, a fact which bears on the direction of evolution in these groups. Due to its isolation Chile has a depauperate but- terfly fauna (Pena, 1975) mostly derived from the Andes. With sterodice so widely distributed in the Andes it would more likely be ancestral to mercedis than the reverse; if the mercedis -autodice group connection is real, the likelihood that the sterodice complex in toto is derived from the autodice group is very low. Thus evolution is more likely to have proceeded from the sterodice to the autodice group than the reverse (as postulated by Gia- comelli). T. s. arctodice of the northern Andes is phenotypically very close to the sympatric T. xanthodice, which on genitalic grounds is grouped by Herrera and Field ( 1959) with T. distincta Jorg. in a separate species-group. The early stages of xanthodice and of the sterodice group are also very similar, differing primarily in the number of ribs in the egg and the length of the pupal proboscis-case. Tentatively these two species-groups may be considered close, and primitive relative to the autodice group. Parallel and convergent adaptations occur repeatedly in the Holarctic and Andean Pierid faunas. The rounded, stocky pupae of the sterodice group are approached by the montane and alpine members of the Holarctic Pieris (Synchloe) callidice Hbn. complex. The striped pattern is a very widespread one in the family, but there is an almost uncanny resemblance among the mature larvae of Tatochila autodice, Pieris protodice, and the Euchloine Euchloe ausonides Lucas. The angular pupa of the autodice group is similar in shape to the Holarctic Pieris rapae (L.) and P. napi (L.) species-groups, but less extreme than the latter. In South America this trend is carried to its apex in Ascia monuste (L.), in which the dorsolateral prominences are prolonged into curving black spines. The angularity of the pupa in Pierini seems very broadly correlated with environmental humidity, but there are too many exceptions to make this a cause-and-effect argument. “Bird drop- ping mimicry” occurs in the pupae of Pieris beckerii Edw. in western North America, and in A. monuste, which like T. autodice often pupates on the host. In subsequent phenetic or cladistic analysis of Pierini, including life 254 NEW YORK ENTOMOLOGICAL SOCIETY history traits, it will be necessary to keep in mind that bewildering, non- concordant parallelisms occur again and again in Pierine evolution (as noted for adult characters by Field (1958).) Acknowledgments This research was supported by the National Science Foundation (USA) under grant DEB-76-18611. The illustrations are by Ms. Lynn S. Kimsey. Research in Argentina was executed under the patronage of the Servicio Nacional de Parques Nacionales, Dr. Felipe Lariviere, Presidente del Di- rectorio, and with the particular aid of Dr. Norberto Ovando (SNPN) and Dr. Alberto Anziano, director of the Museo de la Patagonia, Bariloche. Work in northern Argentina was assisted by Sr. Eduardo Dominguez of San Miguel de Tucuman and by the staff of the Instituto Miguel Lillo. Research in Chile was carried out by Mr. Clinton V. Kellner with the gracious assis- tance of Dr. Jose Herrera G. and of the Departamento de Biologi'a de la Universidad de Chile. Ms. Adrienne R. Shapiro was a devoted field assistant on both Argentine voyages. Some rearing was accomplished in the U.S. under two permits granted by the Department of Agriculture — APHIS. Un- published data were supplied by Dr. Gilberto Bravo V. of the Universidad de Narino, Colombia and by Messrs. B. MacPherson and R. Eisele. Various Crucifers were determined in the U.C. Davis herbarium by Ms. June McCaskill. To all I am deeply grateful. Literature Cited Alvarez, J. and C. H. Delgado. 1969. Ciclo biologico del gusano rayado de las Cruciferas, Tatochila arctodice Staudinger (Lepidoptera; Pieridae) en algunas zonas del altiplano de Pasto, Narino, Colombia. Unpublished thesis, Universidad de Narino. Breyer, A. 1938. Uber die Argentinischen Pieriden (Lepidoptera, Rhopalocera). Proc. VII Int. Congr. Ent. Berlin 1:26-55. Field, W. D. 1958. A redefinition of the butterfly genera Tatochila, Phidia, Piercolias, and Baltia, with descriptions of related genera and subgenera. Proc. U.S. Nat. Mus. 108:103-131. Giacomelli, E. 1915. El genero Tatochila Butl.: lo que sabemos y lo que ignoramos de el. Anales del Museo Nacional de Historia Natural, Bs. As. 26:403-415. Herrera, J. and W. D. Field. 1959. A revision of the butterfly genera Theochila and Tatochila (Lepidoptera: Pieridae). Proc. U.S. Nat. Mus. 108:467-514. Jorgensen, P. 1916. Las mariposas Argentinas (Lepidoptera): familia Pieridae. Anales del Museo Nacional de Historia Natural, Bs. As. 28:427-520. Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook of Colour, third edition. London: Methuen. 252 pp. Margheritis, A. E. and H. F. E. Rizzo. 1965. Lepidopteros de Interes Agricola. Bs. As.: Editorial Sudamericana. 195 pp. Pena G., L. E. 1975. Guia para reconocer mariposas. Santiago: Editora Nacional Gabriela Mistral. 120 pp. VOLUME LXXXVII, NUMBER 3 255 Shapiro, A. M. 1978a. The life history of Reliquia santamarta, a neotropical alpine Pierine butterfly (Lepidoptera; Pieridae). J. N.Y. Ent. Soc. 86:45-50. . 1978b. The life history of an equatorial montane butterfly, Tatochila xanthodice (Lep- idoptera: Pieridae). Ibid. 86:51-55. Department of Zoology, University of California, Davis, California 95616. Received for publication April 30, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(3), 1979, pp. 256-266 A RECONSIDERATION OF THE GENUS BAKERIELLA (HYMENOPTERA: BETHYLIDAE) Howard E. Evans Abstract. — The concept of the neotropical genus Bakeriella is broadened to include several species formerly included in Epyris. Three species are described as new: erythrogaster (Costa Rica), quadriceps (Colombia), and grandis (Costa Rica). Females of polita Evans and inconspicua Evans are described for the first time. Range extensions for several other species are given. The 18 known species collectively range from Florida and Central Mexico to Brazil and Argentina. Introduction The genus Bakeriella was described in 1910 by Kieffer to include two South American bethylids related to Epyris but having pronotal carinae laterally, medially, and anteriorly. In 1964 1 pointed out that there are similar species having the lateral and median carinae evanescent or even absent, and I redefined the genus to include these species. In my revision of Epyris (1969) I pointed out the close resemblance of members of the montivagus group of that genus to the species of Bakeriella. In fact, I suggested that the montivagus group might more properly be shifted to Bakeriella despite the lack of even the anterior pronotal carina in that group. Study of further material has convinced me that this step is inevitable, and furthermore that Bakeriella has evolved from a Rhabdepyris -like stock quite independently of Epyris. One species, described below as erythrogaster, in fact lacks a dividing septum in the scutellar groove, and thus will key to Rhabdepyris in most keys, although in all other features it is a true Bakeriella. I believe that Bakeriella so redefined is a natural group, now known to contain 18 species collectively ranging from Florida and central Mexico to Brazil and Argentina. I present here a brief diagnosis of the genus as now conceived, a key to species, and remarks on each of the species, including a description of three new species and of the first known females of two others. Generic features . — Epyrini fully winged in both sexes and with venation similar to that of Epyris. Antennal scrobes of female not carinate, but males ' Department of Zoology and Entomology, Colorado State University, Fort Collins, Colo- rado 80523. This is part of a study of the world fauna of Bethylidae, supported by the National Science Foundation, grant no. DEB 75-17142. VOLUME LXXXVII, NUMBER 3 257 with a delicate carina crossing above the antennal insertions and extending nearly to the lower eye margins; male antennae with dense erect or suberect setulae, third segment at least nearly as long as second. Eyes of female sparsely hairy, those of male barely or not at all hairy. Pronotum with a punctate groove paralleling the posterior margin, the disc prominent an- terolaterally and often with a transverse carina connecting the anterior an- gles; sides of pronotal disc usually abrupt, often subcarinate or carinate; midline also sometimes carinate. Notauli strong on posterior half of meso- scutum; scutellum with a transverse basal groove which is usually divided by a thin septum, but sometimes undivided or divided by a flat-topped ridge. Propodeal disc elongate, tricarinate, the lateral discal carinae somewhat bowed, space between the carinae often transversely ridged; posterior an- gles of propodeum foveolate. Mesopleura with a single large fovea, the upper margin of which crosses above the pit. Middle tibiae of female spinose above. Grouping of the species. — The following represents what I believe to be a natural grouping of the known species. I regard group I as most Rhab- depyris-\\ke, group IV the most highly evolved in terms of thoracic sculp- turing. In each case I have indicated which sexes have been described; at present 4 species are known only from females, 7 only from males. Group I. Pronotum ecarinate. Scutellar groove simple. 1. erythrogaster n. sp. 9 <5 Group II. Pronotum ecarinate. Scutellar groove divided. 2. montivagus (Kieffer) 9 6 3. reclusus (Evans) 9 c? 4. quinqiiepartitus (Kieffer) 9 5. quadriceps n. sp. 9 6. grandis n. sp. S Group III. Pronotum with transverse carina. Scutellar groove divided. 7. polita Evans 9 d 8. rufocaudata Evans S 9. rossi Evans S 10. brasiliana Evans 9 1 1 . floridana Evans 9 S 12. inconspicua Evans 9 S 13. olmeca Evans 9 6 (Male has a weak median pronotal carina) Group IV. Pronotum with transverse and median carinae. Scutellar groove divided. 14. inca Evans 9 15. azteca Evans 6 16. depressa Kieffer S 17. flavicornis Kieffer 6 18. cristata Evans 9 S 258 NEW YORK ENTOMOLOGICAL SOCIETY Key to Species Females 1. Pronotum without a transverse Carina 2 - Pronotum with a transverse anterior carina 6 2. Scutellar groove not divided by a septum; claws strongly curved, trifid 1 erythrogaster n. sp. - Scutellar groove divided by a septum; claws moderately curved, dentate 3 3. Clypeus very short, subtruncate; posterior angles of head promi- nent and with a welt-like elevation 5 quadriceps n. sp. - Clypeus angulate or subangulate medially ; head rounded above eyes 4 4. Pronotal disc alutaceous, weakly punctate; scutellar pits only slightly wider than long 3 reclusus (Evans) - Pronotal disc with weak surface sculpturing and rather strong punc- tures; scutellar pits much wider than long 5 5. Pronotal disc moderately shining, more strongly alutaceous than front and vertex, somewhat parallel-sided; propodeal disc slightly longer than wide 4 qidnquepartitus (Kieffer) - Pronotal disc shining, not or barely more alutaceous than front and vertex, shorter and with the sides more divergent behind; propo- deal disc as wide as or slightly wider than long 2 monlivagus (Kieffer) 6. Pronotum without a median carina; head black 7 - Pronotum with a complete median carina set off by linear grooves; or with a partially developed median carina, in this case the head dark green 1 1 7. Legs wholly testaceous; propodeal disc covered with weak trans- verse striations 1 1 floridana Evans - Coxae and femora largely fuscous; sides of propodeal disc shining and with at most very weak sculpturing 8 8. Pronotal carina weakly angled forward medially; median area of propodeum rather weakly sculptured 9 \ - Pronotal carina evenly arched; median area of propodeum with ; strong transverse ridges 10 i 9. Mandibles with 5 teeth, the basal 3 small; front with small punctures which are separated by 1.5-3. Ox their own diameters 13 olnieca Evans - Mandibles with only two large apical teeth; front with somewhat stronger and more widely spaced punctures 10 brasiliana Evans 10. Length of fore wing about 3.8 mm; sides of pronotal disc subcari- nate anteriorly 7 polita Evans - Length of fore wing about 3.2 mm; sides of pronotal disc carinate on anterior half 12 inconspicua Evans VOLUME LXXXVII, NUMBER 3 259 11. 1. 2. 3. 4. i 6. 7. 8. 9. 10. Median carina of pronotum complete; head black; scutellar pits separated by a thin septum 14 inca Evans Median carina weakly developed on posterior third of pronotum; head with dark, metallic green reflections; scutellar pits separated by a flat-topped ridge 18 cristata Evans Males Pronotum without a transverse carina 2 Pronotum with a transverse anterior carina 5 Scutellar groove not divided by a septum; claws trifid 1 erythrogaster n. sp. Scutellar groove divided by a septum; claws dentate 3 Abdomen rufous except basal segment largely black; length of fore wing 5.6 mm; propodeal disc slightly longer than wide 6 grandis n. sp. Abdomen black except extreme tip sometimes rufous; fore wing 2. 5-4.0 mm; propodeal disc slightly wider than long 4 Eront polished, very obscurely alutaceous and weakly punctate; notauli somewhat widened behind, but much more widely separat- ed than their own greatest widths 2 montivagus (Kieffer) Eront moderately alutaceous, rather weakly shining; notauli wide behind, separated by about their own greatest widths 3 reclusus (Evans) Pronotum without a median carina 6 Pronotum with a median carina (sometimes rather weak) 10 Front and pronotal disc polished, barely if at all alutaceous; larger species, fore wing 2. 8-4.0 mm 7 Front somewhat alutaceous, pronotal disc weakly shining, strongly alutaceous; smaller species, fore wing less than 2.5 mm in length 9 Antennae elongate, segment 1 1 at least twice as long as wide; apical 0.4 of abdomen rufous 8 riifocaudata Evans Antennae somewhat shorter, segment 11 not more than 1.7x as long as wide; tip of abdomen at most weakly suffused with reddish brown 8 Vertex rather narrowly rounded off far above eye tops; median lobe of clypeus roundly subangular 7 polita Evans Vertex more broadly rounded off, not so strongly angular 9 rossi Evans Sides of pronotal disc subparallel, carinate anteriorly; coxae and femora fuscous 12 inconspiciia Evans Sides of pronotal disc diverging behind, not at all carinate; legs bright testaceous except front coxae infuscated 1 1 floridana Evans Temples not carinate 1 1 Temples with a strong carina extending from vertex to near man- dibular bases 12 260 NEW YORK ENTOMOLOGICAL SOCIETY 11. Scutellar pits separated by a thin septum; pronotal disc subcarinate laterally, with a strong median carina 15 azteca Evans - Scutellar pits separated by a flat-topped ridge; pronotum rounded laterally, with a weak median carina 18 cristata Evans 12. Median carina of pronotum very weak; minimum width of front about 1.2x eye height 13 olmeca Evans - Median carina of pronotum strong; minimum width of front barely if at all exceeding eye height 13 13. Transverse pronotal carina with the crest somewhat sinuate as seen obliquely from behind, subdentate on the sides 16 depressa Kieffer - Transverse pronotal carina with the crest rather even, not subden- tata 17 flavicornis Kieffer 1. Bakeriella erythrogaster n. sp. Holotype. — $, COSTA RICA: Monte Verde, 13-15 Eeb. 1972 (H. M. Powell) [Univ. Calif. Davis]. Description of female type. — Length 5.0 mm; fore wing 3.8 mm. Head 1 and thorax black; abdomen bright rufous except basal segment partially suffused with black; mandibles castaneous; antennae light brown except darker on upper surface of flagellum; coxae and femora fuscous, remainder of legs medium brown; front wings uniformly tinged with brownish. Entire body strongly shining, with scattered short, brownish hairs, somewhat dense on head, pronotum, and femora. Mandibles with 4 teeth in an oblique series, the basal two teeth smaller and more rounded than the others. Me- dian lobe of clypeus obtusely angulate, with a median carina which is 1 straight in profile. Head 0.86x as wide as high; width of front 1.4x eye \ height; ocelli in a compact triangle far above eye tops, ocello-ocular line i 2.2x width of ocellar triangle. Eyes somewhat bulging from sides of head; vertex strongly arched, distance from eye tops to vertex crest (lateral view) subequal to eye height. Front very weakly alutaceous, with strong punctures which are separated by 2-5 x their own widths. Pronotum elongate, disc somewhat prominent anterolaterally; scutellar groove undivided, slightly widened on each side. Propodeal disc very slightly longer than wide, with a strong median carina that extends down the declivity; disc with two ad- ditional shorter carinae which are bowed in the form of a lyre, surface between these weakly and irregularly transversely striate, disc elsewhere with only very weak surface sculpturing. Claws strongly curved, trifid. Allotype. — S , COSTA RICA: Same data as type but dated 29 June 1972 [Univ. Calif. Davis]. Description of male allotype. — Length 4.0 mm; fore wing 3.5 mm. Color as in female except basal third of abdomen extensively suffused with black, antennae rather uniformly medium brown; body shining and with short, light brown hairs as in female. Mandibles with a strong apical tooth and 4 much VOLUME LXXXVII, NUMBER 3 261 smaller teeth in a nearly straight series based at this tooth; clypeus as in female. Head 0.89x as wide as high; width of front l.lx eye height; ocelli rather large, ocello-ocular line 1.3x width of ocellar triangle. Antennae elon- gate, first 4 segments in a ratio of 12:7:8:10, segment 3 about 1.5x as long as wide, segment eleven 3x as long as wide. Eyes strongly bulging, not hairy, vertex broadly rounded off far above eye tops. Front very weakly alutaceous, its punctures smaller and sparser than in female. Features of thorax and propodeum as in female; claws trifid as in that sex. Paratypes. — 12 9$, 2 6 6. COSTA RICA: 1 5, same data as allotype [Univ. Calif. Davis]; COLUMBIA: 11 9 9,266, Penas Blancas, Dept. Valle, 24 March 1975 (except 1 9 22 Nov. 1974) (R. Wilkerson, Malaise trap) [Fla. State Coll. Arthropods; U.S. National Museum; Mus. Comp. Zool.j. Variation. — The Costa Rica paratype is closely similar to the type. The females from Colombia consistently have the basal two abdominal segments black, while the males have the greater part of the abdomen black, only the apical 2-3 segments rufous. In some of the females the basal mandibular tooth is partially divided, such that the mandibles are indistinctly 5-toothed. Some of the Colombia females are somewhat larger than those from Costa Rica, fore wing up to 4.5 mm. The head is consistently shorter in Colombia specimens than in those from Costa Rica, the head being from 0.88 to 0.92 x as wide as high in the females, from 0.94 to 0.96 x as wide as high in the males. However, there seems no basis for regarding these two series as representing different species or, in the absence of material from other lo- calities, different subspecies. Remarks. — This species will run to Rhabdepyris in existing keys because of the undivided scutellar groove. However, it fits poorly in that genus on other features and is quite clearly related to the more generalized species of Bakeriella . 2. Bakeriella montivaga (Kieffer) new combination Epyris montivagus Kieffer, 1910; Evans, 1969. This is possibly the most common and widely distributed member of the genus, ranging from Bolivia to Venezuela and to southern Mexico. Synon- ymy and a redescription are provided by Evans (1969). 3. Bakeriella reclusa (Evans) new combination Epyris reclusus Evans, 1969. This species was described from Costa Rica and El Salvador. I have since seen a female from ECUADOR; 11 km SE San Lorenzo. Prov. Esmeraldas, 1975 (S. & J. Peck) [Mus. Comp. Zool.]. 262 NEW YORK ENTOMOLOGICAL SOCIETY 4. Bakeriella qidnquepartita (Kieffer) new combination Epyris qidnquepartitus Kieffer, 1910; Evans, 1969. A redescription of this species was presented in 1969. Only the type fe- male, from Marcapata, Peru, is known. 5. Bakeriella quadriceps n. sp. Holotype. — $, COLOMBIA: Finca los Guaduales, near San Jose del Pal- mar, Choco, 730-800 m, 1 June 1978 (C. Kugler) [Mus. Comp. Zool.]. Description of female type. — Length 5.2 mm; fore wing 3.5 mm. Head and thorax black; basal two abdominal segments black, third segment black except rufous along posterior margin, fourth segment partially suffused with black dorsally, remainder of abdomen bright rufous; mandibles dull ferru- ginous, black at base; antennae uniformly dull ferruginous; legs black except trochanters, tibiae and tarsi dusky ferruginous; wings subhyaline. Front with numerous short, golden brown setae, temples and thoracic dorsum and pleu- ra more sparsely setose. Mandibles much broadened apically, with 4 large teeth, most basal teeth partially subdivided. Clypeus with a very short, broad truncate median lobe, its median line weakly carinate. Head quadrate, its posterior angles prominent and with a rounded welt; head very slightly higher than wide; width of front 1.4x eye height; front angle of ocellar triangle less than a right angle, ocelli far below vertex crest, ocello-ocular line 2.3 X width of ocellar triangle. First 4 antennal segments in a ratio of 25:6;5:6, segment 3 about as wide as long. Eyes weakly hairy; distance from eye tops to vertex crest exceeding eye height; vertex nearly straight in anterior view for a considerable distance. Front alutaceous, moderately shining, with shallow punctures which are separated by 3-7x their own diameters. Pronotal disc somewhat flat, prominent anterolaterally, surface somewhat more alutaceous and weakly punctate than that of front; scutellar pits slightly wider than long, separated by a thin septum medially. Propodeal disc distinctly widened posteriorly, length slightly exceeding its maximum width; median area with delicate, irregular transverse striae. Claws dentate. Remarks. — This species is known only from the type. 6. Bakeriella grandis n. sp. Holotype. — S , COSTA RICA; Monte Verde, 29 June 1972 (H. M. Powell) [Univ. Calif. Davis]. Description of male type. — Length 8.0 mm; fore wing 5.6 mm. Head and thorax black; abdomen bright rufous except first segment largely suffused with black; mandibles rufous above and on apical half; antennae rufous except scape somewhat infuscated and apical third of flagellum fuscous; coxae and femora black, legs otherwise rufotestaceous except trochanters VOLUME LXXXVII, NUMBER 3 263 and tibiae slightly infuscated; wings lightly and uniformly tinged with brown. Body and legs with fairly abundant short, light brown hairs. Mandibles with a large apical tooth and 4 very small teeth basad of this. Median lobe of clypeus angulate, tectiform. Head 0.95 x as wide as high; width of front 1.3x eye height; ocelli in a compact triangle far above eye tops, ocello- ocular distance 1.55x width of ocellar triangle. First 4 antennal segments in a ratio of 14:6:7:9, segment three 1.7x as long as wide. Distance from eye tops to vertex crest subequal to eye height; vertex broadly arched. Front polished, very weakly alutaceous, punctures very small, separated by 4-6 X their own diameters. Pronotal disc slightly duller and more alutaceous than front, but similarly punctate; pronotum prominent anterolaterally but without carinae. Notauli strong, extending for length of mesoscutum; scu- tellar groove rather wide, with a thin septum medially. Propodeal disc 0.95 x as wide as long, surface rather shining and with very weak surface sculpturing even between the carinae. Mesopleura punctate and with somewhat irreg- ular sculpturing. Claws dentate. Remarks. — This striking species is known only from the type. 7. Bakeriella polita Evans Bakeriella polita Evans, 1964. I described this species from males from Bolivia and Peru. I take this , opportunity to describe a female from PERU: Monson Valley, Tingo Maria, ; 26 October 1954 (E. I. Schlinger & E. S. Ross) [Calif. Acad. Sci.]. I Description of female. — Length 6.0 mm; fore wing 4.8 mm. Black, except apical third of abdomen suffused with dull rufous; mandibles and antennae i ferruginous; coxae and femora fuscous, legs otherwise dull ferruginous; wings ' subhyaline. Mandibles 5-toothed; clypeus obtusely angulate, with a sharp median carina. Width of head 0.90x length of head; width of front 1.25x I height of eye; ocello-ocular line 1.9x width of ocellar triangle. Third anten- nal segment about as wide as long. Vertex broadly rounded off well above ; eye tops, top of occipital carina visible in full frontal view. Eront polished, very weakly alutaceous, with strong punctures which are separated by 3- 5x their own diameters. Pronotal disc similarly punctate but more distinctly alutaceous; front margin with a strong, evenly arched carina, side margins sharp but not carinate. Scutellar pits only slightly wider than long, septum between them round-topped, somewhat wider than in related species. Pro- podeal disc 1. 15x as wide as long, the median area with rather strong trans- verse rugae. Claws dentate. 8. Bakeriella rufocandata Evans Bakeriella rufocandata Evans, 1964. This species is known from a single male from the mountains of Colombia. 264 NEW YORK ENTOMOLOGICAL SOCIETY 9. Bakeriella rossi Evans Bakeriella rossi Evans, 1964. I have seen no additional specimens of this species, described from males from two localities in Colombia. 10. Bakeriella brasiliana Evans Bakeriella brasiliana Evans, 1964. This species is evidently widely distributed in the neotropics. A female from the Chagres River, PANAMA, collected 14 July 1918 by Dietz & Zetek [U.S. Nat. Mus.] represents a considerable range extension. 1 1 . Bakeriella floridana Evans Bakeriella floridana Evans, 1964; Evans, 1970. This small but distinctive species was described from Dade Co., Florida, and later reported from Good Hope, Jamaica. Presumably it will be found to occur elsewhere in the West Indies. 12. Bakeriella inconspicua Evans Bakeriella inconspicua Evans, 1964. I have seen several additional specimens of this species since the original description. A male from Tamazunchale, San Luis Potosi, MEXICO, col- lected 6 July 1965 (H. E. Evans) [Mus. Comp. Zool.] represents a small northward extension of the range. I have also seen a series of 1 9 and 6 6 6 from Merida, VENEZUELA, collected 17 November 1972 (G. E. Bo- hart) [Utah State Univ.]. Since this is the first female known, a short de- scription follows. Description of female. — Length 4.4 mm; fore wing 3.2 mm. Black, except apical fifth of abdomen bright rufotestaceous; apical half of mandibles fer- ruginous, antennae wholly of this color except apical segment dusky; legs fuscous except tarsi and parts of tibiae testaceous; wings hyaline. Mandibles with 3 strong teeth (may be 5-toothed); clypeus obtusely angulate, carinate medially. Width of head 0.87 x height of head; width of front 1.35x eye height; ocello-ocular line 1.7x width of ocellar triangle. Third antennal seg- ment about as wide as long. Eyes slightly bulging, quite strongly hairy; vertex rounded off far above eye tops. Front shining, weakly alutaceous, punctures strong, separated by 2-4 x their own diameters. Pronotal disc more strongly alutaceous than front and with sparser punctures; disc with a strong, arching carina in front and the sides carinate on the anterior half. Scutellar pits ovoid, separated by a thin septum. Propodeal disc very slightly VOLUME LXXXVII, NUMBER 3 265 longer than wide, its median area with irregular transverse rugae. Claws simple. 13. Bakeriella olmecci Evans Bakeriella olineca Evans, 1964. This species was described from Veracruz, MEXICO. I am now able to present several additional records: BELIZE: 2 9 9, Middlesex, March 1965 (E. C. Welling) [Canad. Nat. Coll.]; NICARAGUA: 1 9, Musawas, Waspuc R., 23 October 1965 (B. Malkin) [Univ. Calif. Berkeley]; COSTA RICA: Osa Peninsula, Puntarenas (R. W. Matthews) [Mus. Comp. Zool.]. 14. Bakeriella Inca Evans Bakeriella inca Evans, 1964. This species was described from Peru, and can now be recorded from adjacent countries, as follows: ECUADOR: 1 9, Chimbaratzo, Zamora, 3, March 1965 (L. Pena) [Amer. Mus. Nat. Hist.]; BOLIVIA: 18 9 9, Rio Itenez, Beni, 1964 (Bouseman & Lussenhop) [Amer. Mus. Nat. Hist.]; 1 9, 20 km W Laranjeiras, August 1964 (Bouseman & Lussenhop) [Amer. Mus. Nat. Hist.]. 15. Bakeriella azteca Evans Bakeriella azteca Evans, 1964. I have seen no additional specimens of this species described from a male from Morelos, MEXICO. 16. Bakeriella depressa Kieffer Bakeriella depressa Kieffer, 1910; Evans, 1964. I am unfamiliar with the female of this species, which was described from Peru and later reported from Panama and Costa Rica. I have also seen a male from ECUADOR: Puyo, 18 April 1958 (R. Hodges) [Mich. State Univ.]. 17. Bakeriella flavicornis Kieffer Bakeriella flavicornis Kieffer, 1910; Evans, 1964. This species is known only from the type from Para, Brazil. It is doubt- fully distinct from the preceding species. 266 NEW YORK ENTOMOLOGICAL SOCIETY 18. Bakeriella cristata Evans Bakeriella cristata Evans, 1964; Evans, 1973. I described this species from males from Brazil and Bolivia, and subse- quently reported males from two localities in Argentina (in Salta and Mi- siones); I also described a female from Santa Catarina, Brazil. Evidently this distinctive species is widely distributed in South America. Literature Cited Evans, H. E. 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bull. Mus. Comp. Zool. Harvard 132:1-222. . 1969. A revision of the genus Epyris in the Americas (Hymenoptera: Bethylidae). Trans. Amer. Ent. Soc. 95:181-352. . 1970. West Indian wasps of the subfamilies Epyrinae and Bethylinae. Proc. Ent. Soc. Wash. 72:340-356. -. 1973. Further studies on South American Bethylidae (Hymenoptera). Proc. Ent. Soc. Wash. 75:194-204. Kieffer, J. J. 1910. Description de nouveaux bethylides [Hymen.]. Ann. Soc. Ent. France 79:31-56. Received for publication May 15, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK3), 1979, p. 267 BOOK REVIEW Diseases, Pests and Weeds in Tropical Crops. Jurgen Kranz, Heinz Schmutterer and Werner Koch, eds. John Wiley & Sons, 1978. 666 p. $61. This book has been prepared by more than 150 experts, but it appears homogenous because the contributors rigidly followed the same general out- line. The volume contains an exhaustive number of fine illustrations, in- cluding numerous excellent color plates. The authors have succeeded in providing a text that is brief, readable, and authoritative. The information will be especially useful for those working in tropical Third World nations. Plant protection is emphasized throughout. The important pests and plant diseases are described in a way understandable not only by entomologists and plant pathologists, but also by students in agricultural colleges and lay persons. The work of many contributors has been sponsored by the German Government and a large number of contributors are from Central Europe. The paper as well as the printing and color reproduction, by Saledril Stein- kopf & Son in Berlin, Germany, are of high quality. The book provides an important reference source and it belongs in every library in agricultural colleges and universities. There is an adequate subject index. Although the cited references indicate that it took 3-4 years to produce this book from submitted manuscripts, it is nevertheless up-to-date in most instances. The volume constitutes a major contribution to the crop protection literature. K. Maramorosch, Waksman Institute of Microbiology, Rutgers — The State University. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK3), 1979, p. 268 BOOK REVIEW American Spiders. (Second ed.) Willis J. Gertsch. Van Nostrand Reinhold Co. 1979. 274 p. $24.95. The first edition of this book appeared in 1949, with the description of nearly 3,000 species of American spiders. After 30 years the author revised his fascinating account, adding numerous new black and white photographs, 32 fine color illustrations, as well as descriptions of certain spiders living outside the New World. The book starts with an overview of the life of spiders, their silk spinning, courtship and mating, as well as the evolution of spiders and their relationship to other animals. The major groups of spi- ders— the mygalomorphs, to which the tarantulas belong, the Dribellate, aerial, and hunting spiders are all described. There is a very interesting chapter on the economic and medical importance of spiders. The book is not only authoritative but also quite readable and entertaining, thanks to anecdotes, myths, legends and beliefs. In addition to entomologists and students of spiders, everyone interested in natural history will find this vol- ume stimulating and useful. The glossary and index are short but sufficiently detailed. Gertsch is meticulous about his facts and data and his contribution will be a welcome addition to college libraries and to personal libraries. It will appeal to lay persons as well as to science students, young and old, learned amateurs and professionals, as a fascinating account of the largely ignored, often feared, strange and complex spiders. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers — The State University. Journal of the New York Entomological Society VOLUME LXXXVII DECEMBER 1979 NO. 4 I , CONTENTS Variation in the embolus of Metaphidippus insignis (Banks) (Araneae: Salticidae) Bruce Cutler 270-274 (Bionomics and behavior of Alloxysta tnegourae, an aphid hyperparasitoid (Hyme- I noptera: Alloxystidae) Irene Matejko and Daniel J. Sullivan, S. J. 275-282 iThe larva of Calopteron terminale (Say) with additional notes on adult behavior : (Coleoptera: Lycidae) Tim L. McCabe and Linnea M. Johnson 283-288 Notes on a population outbreak of the beetle Uroplata sp. (Coleoptera: Chrysomeli- ; dae) on the tree /7//oiu (Malpighiaceae) in Costa Rica Allen M. Young 289-298 [Field studies and parasites of Liriomyza trifoUearum (Diptera: Agromyzidae) in ^ northeastern USA R. M. Hendrickson, Jr. 299-303 Three new Middle American species of aquatic beetles in the genus Notionotus I Spangler (Hydrophilidae: Hydrobiinae) Philip D. Perkins 304-311 Review; Tabanidae of the East Coast as an economic problem Elton J. Hansens 3 12-3 18 Acknowledgment 319 [Book Reviews 319-324 Honorary Life and Sustaining Members 324 Index of Scientific Names of Animals and Plants for Volume LXXXVII 325-329 Index of Authors for Volume LXXXVII i_iii NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4), 1979, pp. 270-27^ \ VARIATION IN THE EMBOLUS OE METAPHIDIPPUS INSIGNIS (BANKS) (ARANEAE: SALTICIDAE) Bruce Cutler Abstract. — Abnormal morphological variation in the emboli of male palpi of M. insignis from Minnesota and Montana is illustrated. Some of these emboli resemble normal emboli of M. montanus, a closely related species. This resemblance may lead to incorrect species determinations. The palpus of the mature male spider is considered to offer the most consistent morphological elements for species discrimination. In some spi- der species parts of the palpus remain attached to the female epigynum as a normal consequence of mating (Exline and Whitcomb, 1965; Levi, 1970, 1973; McCrone and Levi, 1964). It is suspected in these species, that an individual female mates once or twice, and the loss of the male palpal ele- ments prevents other males from mating with the same females. Since strik- ing differences can occur in morphology between virgin and mated males, taxonomists must take this source of variation into account. However, since these parts break at the same points, and may be recovered from the epi- gyna, once this process is recognized little difficulty will be encountered in interpreting such variation taxonomically. This paper describes a variation in palpal structure which is more difficult ] to explain, and may be taxonomically misleading. The embolus is the in- tromittent sclerite in male spiders. In the genus Metaphidippus it offers a i reliable morphology for distinguishing species. Indeed it is almost the only i criterion used to separate preserved male specimens, although differences ) may be subtle (Kaston, 1973). Figure lA depicts a ventral view of the cym- j bium, bulb and embolus of a typical male M. insignis from Minnesota. This 1 differs only slightly from the embolus illustrated by Kaston (1973). The' I embolus of M. montanus (Emerton), a closely related species, is shown in i Figure IB. The remaining figures are of aberrant emboli of specimens from i Minnesota and Montana. The embolus illustrated in Figure 1C resembles * that of M. montanus closely, yet the other palpus, body pattern, and body | size are those of typical M. insignis. , ’ Out of 39 male M. insignis taken in Minnesota, 9 exhibited at least one, i abnormal palpus. One example (not illustrated) had 2 extremely abnormal ( palpi in which all elements of the palpus were deformed. This may represent i a true teratology, or may be the result of abnormal conditions during the final molt. There is no geographic clustering evident in these specimens, they come from all over the range of the species in the state. One gets the, VOLUME LXXXVIl, NUMBER 4 271 I Fig. 1. A = Ventral view of typical left palpal tarsus of male Metaphidippus insignis from Minnesota, E is the embolus. B = Ventral view of left embolus of Metaphidippus montanus • Tom Manitoba. C = Ventral view of aberrant left embolus of Metaphidippus insignis from I 'Petroleum County, Montana. I 'impression that the typical rounded embolus tip has broken off to a variable I degree producing the angular aberrant emboli. Over 50 females, including j some collected at the same time as the males with atypical emboli, were J examined. None had embolic remnants in the epigyna. I have also examined ' t males from Wyoming and Montana, and one of these has an abnormal imbolus (Fig. 1C). Kaston ( 1973) did not indicate any range of morphological variation in the emboli of Metaphidippus covered in his study. Galiano (1963) encountered 272 I NEW YORK ENTOMOLOGICAL SOCIETY iC- i i Left embolus of a male from Ramsey County. B = Right embolus of the same male as in A : C = Left embolus of a different male from Ramsey County. D = Right embolus of male fron [-1 Ottertail County. E = Left embolus of another male from Ramsey County. Nonillustratei males similar to the illustrated specimens were from Norman, Pipestone, Stearns and Winon: i Counties. an extreme range of variation in palpal morphology in male Euophrys (Sal-' ticidae) reared from one eggsac. Possibly some of this variation was th« result of rearing. Male spiders often have difficulty in freeing the palpi during h the final molt under laboratory conditions, although this usually affects the i I VOLUME LXXXVIl. NUMBER 4 273 I ! Structure of the other palpal elements rather than the embolus (personal observations). The few male M. insignis I have reared from wild caught limmatures had typical palpi when mature. Perusal of the literature uncov- ered no species of jumping spider where a palpal element is broken off during mating. 1 cannot explain the variations seen in M. insignis. I have examined hundreds of specimens of other species of Metaphidippus from Minnesota, iand have never found any variation of the magnitude shown here in the emboli of these other species. It is possible that Minnesota and states to the west might lie in a hybrid zone between M. insignis and M. montanns. 1 Unfortunately, few specimens are available from surrounding states and ^provinces to adequately study the problem. M. insignis is found from New England and Ontario south to New Jersey, and west in the northern tier of states to the Rocky Mountains. In Minne- sota, it is a grassland and meadow species, not found on primarily forested , sites. M. montanns is found in the boreal forest from Newfoundland to the District of Mackenzie, and south in the high mountains of New England and New York. Many specimens determined as this species in collections, es- pecially from the Rocky Mountains, belong elsewhere. It is a larger species than M. insignis with a slightly different body pattern. All known eastern records indicate that the species are allopatric. I As a precautionary measure, both palpi should be examined during de- terminations in this species group. Bilateral asymmetries and abnormal palpi .will become apparent, and be less misleading. Acknowledgments I wish to thank Dr. R. Carter, Manitoba, and Mr. R. Huber, Minnesota, for collecting sometof the specimens used in this study. Literature Cited 'Exline, H. and W. H. Whitcomb. 1965. Clarification of the mating procedure of Peucetici viridans (Araneida; Oxyopidae) by a microscopic examination of the epigynal plug. ' Florida Entomol. 48: 169-171. Galiano, M. E. 1963. Las variaciones individuales en Euophrys siitri.x Holmberg, 1874 (Ara- neae, Salticidae). Rev. Soc. Entomol. Argentina 24 (1961):23-28. iKaston, B. J. 1973. Four new species of Metaphidippus, with notes on related jumping spiders (Araneae: Salticidae) from the eastern and central United States. Trans. Am. Micros. Soc. 92:106-122. ILevi, H. W. 1970. Problems in the reproductive physiology of the spider palpus. Bull. Mus. Natl. Hist. Natur. Paris 2nd ser., 41, suppl. 1:108-111. 274 NEW YORK ENTOMOLOGICAL SOCIET’i . 1973. Observations on the reproductive physiology of the spider Singa (Araneidae). Proc. 5th Int. Arachnol. Cong. (Brno, 1971): 189-192. McCrone, J. D. and H. W. Levi. 1964. North American widow spiders of the Latrodectus curicaviensis group. Psyche 71:12-27. 1747 Eustis Street, St. Paul, Minnesota 55113. Received for publication March 1, 1979. 'i I 1 NEW YORK ENTOMOLOGICAL SOCIETY LXXXV1K4), 1979, pp. 275-282 BIONOMICS AND BEHAVIOR OF ALLOXYSTA MEGOURAE, AN APHID HYPERPARASITOID (HYMENOPTERA: ALLOXYSTIDAE) Irene Matejko* and Daniel J. Sullivan, S. Abstract. — The authors studied Alloxysta megourae (Ashmead), a hy- perparasitoid of the pea aphid, Acyrthosiphon pisum (Harris). Larval, pre- pupal, and pupal stages of the primary parasitoid {Aphidius smithi) found in non-mummified aphids are attacked by A. megourae. Several aspects of A. megourae development within its host differ from those described in other species of Alloxysta. The sensory structures on the tip of the ovipos- itor of A. megourae are described. Introduction Several hymenopteran families are aphid hyperparasitoids. These aphid hyperparasitoids can be divided into two categories based on their attack behavior. (1) Endoparasitoids: the female oviposits in the primary parasitoid larva and then the hyperparasitoid larva feeds internally. (2) Ectoparasi- toids: the female oviposits on the surface of the primary parasitoid and then the hyperparasitoid larva feeds externally. We studied Alloxysta megourae (Ashmead), an endoparasitoid of Aphidius smithi. The classification of Alloxysta was studied by Andrews (1978). Haviland (1921) was the first to comment on the relationship among three species of Alloxystidae (given as Cynipidae) and their primary parasitoid host. The embryology and larval development of Alloxysta (given as Charips) was described by Haviland (1921). Gutierrez and van den Bosch (1970a, b), Gutierrez (1970a-d), Sullivan and van den Bosch (1971) and Sullivan (1972) studied Alloxysta (Charips) victrix in the field and laboratory. The bio- nomics and behavior of Alloxysta megourae have not been reported before. The sensory apparatus on the ovipositor of A. megourae is described. Materials and Methods The pea aphid, Acyrthosiphon pisum (Harris), served as the host in this study, and was reared on broad bean, Vida fava L. (Windsor variety). ' Present address: Department of Biological Sciences, Stratton Hall, Drexel University, Philadelphia, Pennsylvania 19104. Manuscript is a portion of a dissertation submitted by the first author in partial fulfillment of the requirements for the Ph.D. degree in the Department of Biological Sciences, Fordham University, Bronx, New York 10458. ^ Presently at Fordham University as an Associate Professor in the Department of Biological Sciences, Bronx, New York 10458. 276 NEW YORK ENTOMOLOGICAL SOCIETY, Table 1. Composite life cycles of the primary parasitoid, Aphidius smithi, and the hyper- parasitoid, Allo.xysta megourae, under laboratory conditions. Age in days Aphidius smithi Age in days Alloxysta megourae 2 3 4 5 6 7 8 9 10 11 12 Egg deposited in aphid 1st larval instar 2nd larval instar 3rd larval instar Host aphid mummified® Prepupa (meconium voided) Pupa Adult emerges 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Egg deposited in Aphidius Egg hatches (host aphid mummified) 1st larval instar 2nd larval instar 3rd larval instar Mature larva feeds externally Prepupa (meconium voided) Pupa Adult emerges SI !le When hyperparasitized by Alloxysta, Aphidius ceases development. Aphidius smithi Sharma & Subba Rao, the primary parasitoid, and Alloxystdi megourae (Ashmead), the hyperparasitoid, were reared with the plants andj aphids in a bioclimatic chamber (Percival Environator, Model E-54U) ac- cording to the method described by Bennett and Sullivan (1978). The daytime (16 hr) temperature was 21.1 ± 0.6°C at 75 ± 5% RH. At night the temperature was 15.5 ± 0.6°C at 85 ± 5% RH. The primary parasitoid, Aphidius smithi, was reared by placing 2-A mated -t females in a glass tube with cut broad bean stems and 20 4th instar pea tii aphids. The tube was kept in the bioclimatic chamber for 6 hr. Then the i . wasps were removed and the parasitized aphids replaced on the broad bean 4 leaves and returned to the temperature chamber. After 12 days, the adult L Aphidius emerged from the aphids. j ^ To observe the process of courtship and mating, the adult Alloxysta male} si and female were left undisturbed for 6 hr in the Dixie container. We ob-^ ite VOLUME LXXXVll. NUMBER 4 277 served oviposition by placing 5-6 adult AUoxysta females in a 60 x 15 mm plastic Petri dish with 15 4th instar aphids. The life cycle of A. megourae ,;Table 1) was studied by a technique allowing continuous observation of all jstages of development within the aphid mummy (Keller & Sullivan, 1976). To determine whether A. megourae females attack both parasitized and unparasitized aphids, we placed a female AUoxysta in a Petri dish for 3 hr with a parasitized aphid having a 6-7-day-old Aphidius larva and an unpar- asitized aphid. All aphids used in 50 replicates were in the 4th instar. After the AUoxysta was removed, we dissected and recorded the eggs in each aphid. Results and Discussion Courtship and Mating AUoxysta adults were able to mate soon after emergence. The courtship begins when the male follows the female around the observation chamber. His behavior was predictable; he positioned himself behind the female and tapped her abdomen with his antennae, while she remained motionless. This phase lasted from 2-7 minutes. More intensified courtship began when the Imale mounted the female. He climbed on her back with his legs firmly [grasping the sides of her body. Pairs were frequently observed mounted, ,with the male on top of the female and being carried about by the female |for as long 30 seconds to 1 minute. Copulation lasted from 10-20 seconds. lOviposition Behavior A. megourae behaves as an endoparasitoid. It attacks the Aphidius larva, iwhich has been developing inside a live aphid, and deposits the egg within ithe primary parasitoid larva. j The egg-laying behavior of A. megourae has the following pattern: the jfemale approaches the live parasitized aphid and rapidly antennates its sur- iface. She then mounts the back of the aphid and assumes a squatting po- Isition with her abdomen slightly bent (Fig. 1). Once the AUoxysta female isecures herself tightly on the dorsum of the aphid, the aphid moves violently as if trying to dislodge the wasp. She then inserts her ovipositor through the thin exoskeleton of the aphid and probes the aphid’s hemocoel in search of the primary parasitoid larva (Fig. 2). If unsuccessful at that site, the AUoxy- sta female withdraws the ovipositor, moves to a new position on the aphid’s dorsum and begins to probe again for the host. A. megourae females have not been seen to host-feed, neither before nor after oviposition nor do they paralyze their hosts. 1 It was not unusual to observe 4-5 AUoxysta females ovipositing at the same time on the same live, parasitized aphid. They were not easily dis- 278 NEW YORK ENTOMOLOGICAL SOCIET i ?! VOLUME LXXXVII, NUMBER 4 279 urbed when ovipositing, even when some Alloxysta females climbed over ind under each other while others were ovipositing. Once situated on the iphid, all 4 or 5 Alloxysta females probe simultaneously for the primary barasitoid within the aphid. Invariably, although several eggs may be laid, |)nly one adult developed from each host. Once oviposition is completed, he Alloxysta female withdraws her ovipositor and abruptly leaves the iphid. knsory Structures on Ovipositor ! Scanning electron micrographs of the sensory structures on the ovipositor )f A. megoiirae are seen in Figs. 3 and 4. The ovipositor is composed of |)aired 1st valvulae, and a fused pair of ventral 2nd valvulae. Each of the ; St valvulae has three sensory pits. The 2nd valvulae have barbs on the ')uter surface, but no pits. development within the Mummy :i Eclosion of the Alloxysta larva from the egg occurs about 72 hr after mummy formation and is encased within a white, opaque mass. The larva Is translucent white with visible segmentation on the surface. It undergoes leveral instars, and as the larva increases in size, a color change is evident. It becomes bright yellow, almost indistinguishable from the yellow Aphidius larva. j By the 9th day after Alloxysta egg deposition, the surface of the Aphidius 'jarva begins to change to a dark brown color, it appears wrinkled and slowly ;;!eteriorates. On the 10th day, the Alloxysta larva emerges from the Aphi- \\'ius larva and begins to feed externally on it (Fig. 5). By the end of the 1 1th ijiay, the entire Aphidius larva is consumed with the exception of the head Ijapsule which is pushed to one side of the mummy. On the 12th day, the I inidgut and hindgut of Alloxysta are connected and the meconium is voided ind is pushed against one side of the mummy. This begins the prepupal j Figs. 1-6. Ovipositional behavior and developmental stages of Alloxysta megourae. 1. I \dult Alloxysta female exhibiting a typical drilling position (a) on the parasitized pea aphid . b). The head is down and the legs are firmly attached on the aphid’s dorsum. (25 x) 2. Adult emale Alloxysta ovipositing with antennae extended straight back. (25x) 3. Scanning electron nicrograph of the abdomen of an Alloxysta female with the ovipositor evident (a). (200 x) 4. scanning electron micrograph showing details of the structures located on the tip of an Alloxy- ta ovipositor. Three sensory pits (a) on the 1st valvulae and a row of barbs (b) on the fused ' j’.nd valvulae are shown. (lOOOx) 5. A 10-day-old Alloxysta larva (a) emerging from its host, \phidius (b), which it consumes within a few hours. (80x) 6. The characteristically irregular ;xit hole of Alloxysta located on the posteriodorsal side of the aphid mummy. (50x) NEW YORK ENTOMOLOGICAL SOClET\ 280 Table 2. Discrimination between parasitized and unparasitized aphids by the hyperparasi-i toid female Alloxysta megourae for oviposition (based on 50 replicates). Time in min. probing Sid. Dev. No. of reinsertions Sid. Dev. No. eggs laid Sid. Dev. 1^ Parasitized 8.0 2.80 4.0 0.21 2.7 0.23 Unparasitized 5.6 2.71 3.4 0.25 0 0 ' Stage which lasts about 24 hr. On the 13th day, the pupa is formed. It is' bright yellow, but within 4-5 days, it gradually becomes dark brown. During this pupal stage, the hyperparasitoid was completely motionless. However, it did exhibit abrupt jerking movements if touched with a probe. The adult Alloxysta emerges from the aphid mummy about 19 days after oviposition. The irregular-shaped emergence hole made by the Alloxysta adult was almost always located on the dorsal side of the aphid mummy! (Fig. 6). Once emerged, the Alloxysta adult cleaned itself, fed on the honey- water mixture, and then began the courtship and mating behavior. ; Discrimination between Parasitized and Unparasitized Aphids The parasitoid attack behavior of Charips (Alloxysta) (Haviland 1921; Gutierrez and van den Bosch 1970b) is similar to that of A. megourae. \ However, Haviland (1921) reported that Charips selected only aphids con- taining a primary parasitoid whereas unparasitized aphids were ignored. Gutierrez (1970a) reported that Charips victrix (Westwood) probed unpar-^ asitized aphids. We found that A. megourae females always attacked and probed live aphids, both parasitized and unparasitized. These aphids were then dissected and in the 50 replicates, an average of 2.7 eggs were found in the Aphidius larvae, while no eggs were found in the unparasitized aphids (Table 2). Hence, discrimination appears to be accomplished during probing of the aphid with the ovipositor. : These data indicate several points: (1) Alloxysta females attacked unpar-jj asitized aphids but did not oviposit. (2) Alloxysta females attacked parasit- i ized live aphids containing the Aphidius larva and would readily oviposil j" within the primary parasitoid host. However, the host larva may not always ^ be detected since it may be in an inaccessible part of the aphid’s hemocoeft- In these situations, A. megourae changes position on the aphid’s dorsurr i and probes and searches the hemocoel again (Gutierrez 1970a). Thus, nol i' only does Alloxysta attack the same parasitized aphid more than once (Sul- * livan 1972) but a variable number of eggs are laid in the Aphidius larva « within the aphid as a result of these attacks. A study of Table 2 would seenr . to indicate that there is no direct correlation between the number of ovi- . positional attempts and the actual number of eggs deposited. I 281 iVOLUME LXXXVll. NUMBER 4 iSensory Structures on Ovipositor It has been known that sensory structures are important in host selection and host discrimination. Muesebeck and Dohanian (1927) stated that hy- ■perparasitoids are less discriminatory than primary parasitoids in host se- lection. But, Gutierrez (1970d) showed that the ovipositor of A. victrix has sensory structures near the tip which apparently are used by this hyperpar- jasitoid to discriminate among different species of primary parasitoid host larvae. In our study, A. megourae showed a similar morphology of the ovipositor. The 3 sensory pits are sensilla coeloconica as described by Snod- jgrass (1935) and Chapman (1969). However, in addition to Gutierrez’ men- tion of the sensory pits on the paired 1st valvulae which probably have a ichemosensory function, our photomicrographs also show 8 or more barbs on the outer surface of the 2nd valvulae. Probably these barbs anchor the |2nd valvulae during oviposition, while the 1st valvulae are inserted into the japhid’s hemocoel in order to search for the Aphidius larva. Acknowledgments Special appreciation is expressed to Dr. F. G. Andrews (USDFA, Sac- ramento, California) for taxonomic determination of Alloxysta megourae (Ashmead). Literature Cited Andrews, F. G. 1978. Taxonomy and host specificity of Nearctic Alloxystinae with a catalog of the world species. Occ. Pap. Ent., California Dept, of Food Agric., #25, 128 p. I Bennett, A. W. and D. J. Sullivan, S. J. 1978. Defensive behavior against tertiary parasitism by the larva of Dendrocerus carpenteri an aphid hyperparasitoid. Jour. New York Ento- mol. Soc. 86:153-160. Chapman, R. F. 1969. The Insects. American Elsevier F*ublishing Co. Inc., New York. 819 p. Gutierrez, A. P. 1970a. Studies on host selection and host specificity of the aphid hyperparasite , Charips victrix (Hymenoptera: Cynipidae). 3. Host suitability studies. Ann. Entomol. ; Soc. Amer. 63:1485-91. . 1970b. Studies on host selection and host specificity of the aphid hyperparasite Char- ips victrix (Hymenoptera: Cynipidae). 4. The effect of age of host on host selection. Ibid. 63:1491^. . 1970c. Studies on host selection and host specificity of the aphid hyperparasite Charips , victrix (Hymenoptera: Cynipidae). 5. Host selection. Ibid. 63:1495-8. . 1970d. Studies on host selection and host specificity of the aphid hyperparasite Char- ips victrix (Hymenoptera: Cynipidae). 6. Description of sensory structures and a syn- opsis of host selection and host specificity. Ibid. 63:1705-09. , and R. van den Bosch. 1970a. Studies on host selection and host specificity of the aphid hyperparasite Charips victrix (Hymenoptera: Cynipidae). I. Review of hyperpar- asitism and the field ecology of Charips victrix. Ibid. 63:1345-54. . 1970b. Studies on host selection and host specificity of the aphid hyperparasite Char- 282 NEW YORK ENTOMOLOGICAL SOCIET'i Ibid ips victrix (Hymenoptera: Cynipidae). 2. The bionomics of Charips victrix 63:1355-60. Haviland, M. D. 1921. On the bionomics and development of Lygocerus testaceimaniis Kief- fer and Lygocerus cameroni Kieffer (Proctotrupoidea: Ceraphronidae), parasite ol Aphidius. Quart. Jour. Micros. Sci. 65:101-127. Keller, L. J., and D. J. Sullivan, S. J. 1976. Oviposition behavior and host feeding of Asaphes lucens an aphid hyperparasitoid. Jour. New York Entomol. Soc. 84:206-211 Muesebeck, C. F. S., and S. M. Dohanian. 1927. A study of hyperparasitism with particular reference to the parasites of Apanteles melanoscelus (Ratzeburg). U. S. Dept. Agric Bull. 1487:1-35. Snodgrass, R. E. 1935. Principles of Insect Morphology. McGraw-Hill, New York and Lon- don. 667 p. Sullivan, D. J. 1972. Comparative behavior and competition between two aphid hyperparas ites: Alloxysta victrix and Asaphes californicus (Hymenoptera: Cynipidae; Pteromali dae). Environ. Entomol. 1:234-244. , and R. van den Bosch. 1971. Field ecology of the primary parasites and hyperparasites of the potato aphid, Macrosiphum euphorbiae, in the East San Francisco Bay Area Ann. Entomol. Soc. Amer. 64:389-394. ll B Department of Biological Sciences, Fordham University, Bronx, New York 10458. Received for publication April 17, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4). 1979, pp. 283-288 LARVA OF CALOPTERON TERMINALE (SAY) WITH ADDITIONAL NOTES ON ADULT BEHAVIOR (COLEOPTERA: LYCIDAE)' Tim L. McCabe and Linnea M. Johnson Abstract. — Gregarious behavior was observed in the pupae and adults of ' Calopteron tenninale (Say) (Coleoptera: Lycidae). Predatory habits in the adults were not confirmed by experiments. The larva and pupa are described and illustrated. Young & Fischer (1972) observed ultimate instar larvae of Calopteron terminate (Say) that were apparently seeking a pupation site. A description of the pupa and details on the pupation were given, based on captive spec- I imens, although the larva was not described. The larval skin is not shed, I but splits on the side and remains attached at the base similar to Calopteron ifasciatum Fab. (Withycombe, 1926). According to Withycombe (1926): “When full grown, the larvae of C . fasciatum congregate in masses on the i underside of the trunk for pupation. From a nucleus of early individuals a mass of several hundreds may radiate.” Young & Fischer (1972) did not report gregarious pupation in C. terminate, but they were observing captive specimens. On August 15, 1971, mass pupation of C. terminate was ob- served near North Twin Lake, Becker County, Minnesota. As with C. fasciatum, C. terminate would pupate in an expanding circle from a point of origin; this point being a hole in the bark from which the larvae were emerging. Thirty larvae and pupae were on the surface of the bark. The site was not revisited and more individuals may have emerged from beneath the bark. Several adults were observed eclosing at midday. Mating did not occur at the pupation site and unmated adults were observed taking their maiden flight. Mated pairs have been observed dropping from canopy aggregations of C. terminate in Minnesota and North Dakota and in canopy aggregations of Lycus loripes (Chevrolat) and Lyciis arizonensis Green in Arizona. The weak flight of lycids is not capable of supporting two individuals and mated pairs frequently become dislodged and fall to the ground. Burke (1976) gave a detailed account of pre-copulatory behavior in C. terminate. Eisner & Kafatos (1962) proved that the gregarious activities of Lycus loripes were facilitated by an unidentified pheromone emitted by the males. ’ Published by Permission of the Director, New York State Museum, State Education De- partment, as Journal Series No. 724. 284 NEW YORK ENTOMOLOGICAL SOCIETY Figs. 1-2. Calopteron terminate (Say), mature larva, dorsal and ventral aspects (insert: abdominal spiracle). North Twin Lake, Becker County, Minnesota. Total length 14.0 mm. It is not known if C. terminale employ a pheromone. The senior author and Robert Dregseth encountered an aggregation of several hundred C. termi- nale on June 22, 1972, near the Walcott Dunes, Richland County, North Dakota. The beetles were abundant in the uppermost leaves of a boxelder (Acer negundo L.). The tree was in a sunny location and was infested with aphids. A sample of the beetles was collected with an eight foot pole-and- net assembly from the roof of a van. Young & Fischer ( 1972) and Britton ( 1967) speculated that the adults were predaceous. Four C. terminale adults from the Catskill Mountains, Greene County, New York, were deprived of food and water for three days. After this time, they were offered droplets of water upon which they drank until VOLUME LXXXVll, NUMBER 4 285 Eigs. 3^. Calopteron terminate (Say), larval head capsule, dorsal and ventral aspects. North Twin Lake, Becker County, Minnesota. 286 NEW YORK ENTOMOLOGICAL SOCIET'^ satiated, then they were offered de-winged flies, chrysomelid larvae, mirid immatures, and a noctuid larva. They would not attack any of these and; would actually make escape movements when one of the “prey” was en- countered. Burke (1976) offered adults of C. terminate a small weevil larva, Cossoniis sp., but could not get them to feed. The Catskill beetles were also I offered aphids. Again, escape movements were noted when the prey were' detected. Finally, honey dew from the aphids was provided. The beetles, even though satiated for water, fed readily on the honey dew. However, sugar solutions are readily taken in by beetles (Thorsteinson (1960); Gotts- chalk (1957)). Nonetheless, the presence of honey dew may be important for site selection in C. terminate. Linsley, et al. (1961) reported nectar and pollen feeding in Lyciis toripes. In addition, Lycus minutus Green has been observed feeding on the staminate cones of Satix sp. in Miller Canyon, Huachuca Mountains, Arizona. The pupa (Figs. 5 & 6) has been previously described by Young & Fischer VOLUME LXXXVIl, NUMBER 4 287 (1972) and a photograph of the pupa was reproduced. The dorsal tubercles found in the pupal stage are reminiscent of the larval tubercles of Caenia dimidiata Fab. (figured by Boving & Craighead, 1931) and suggests the close relationship of the two genera. The larva of C. terminale lives beneath as well as upon the bark of dead ' trees, apparently showing a preference to erect trunks. Lycid beetle larvae I of the tribe Lygistopterini also occur under bark or in rotten wood, but I many Lycini larvae can be taken on open ground at night. Larvae of C. terminale are probably lignivorous as is C. fasciatum (Withycombe, 1926). , The description is based on an ultimate instar larva. The Larva of Calopteron terminale (Say) General. — Onisciform; 14 mm long, 5 mm wide; body depressed, curved in lateral view, with dark brown markings on a brown body, glabrous. Head. — Prognathous, depressed, subquadrate, partially hidden by pro- thorax; frontal and epicranial sutures absent; frons, clypeus, and labrum fused; a single large ocellus on lateral margin posterior to each antenna; antennae prominent, two-segmented, basal segment a narrow ring, terminal segment short, blunt, “dome shaped,” with membranous tip; mandibles ' falciform, each bearing single, short seta, in two parts, inner ensheathed by outer shell, curved at tip, opposed at base; maxillary palpi conical, each 4-segmented; galea subequal to palpus in length, bearing 4 short setae; stipes and cardo fused to enlarged mentum; labial palpi small, one-third length of ! maxillary palpi, 2-segmented, each arising from membranous basal segment, approximate to each other but distant from base of maxillary palpus; men- I turn and submentum fused. Thorax. — Prothorax longest of three sequentially decreasing segments; 1 prothoracic spiracle located near anterior margin of mesothorax within lat- eral sclerotized projection; meso- and metathorax with somewhat circular dorsal shield; legs moderately long, tibia bearing tiny hairs, tapering to single tarsal claw. Abdomen. — Depressed, widest and thickest at mid-abdominal region, seg- ments 1-8 with elevated subquadrate dorsal shields, distinct lateral flanges sclerotized dorsally; abdominal spiracles (Fig. 1) apparently of an annular biforous type with annular portion inconspicuous, borne on tubercle; ventral abdominal aspects with circular markings each bearing two extremely mi- nute setae; segment 9 flat, posterior margin concave bearing 4 short setae ventrally; urogomphi absent. Material examined. — One mature larva. North Twin Lake, Becker Coun- ty Minnesota, August 15, 1971, collected and determined by association with reared adults by T. L. McCabe. 288 NEW YORK ENTOMOLOGICAL SOCIETY Literature Cited Boving, A. G. and F. C. Craighead. 1931. An illustrated synopsis of the principal [sic] larval forms of the order of Coleoptera. Ent. Amer. 1 1(3): 161-256. Britton, E. B. 1970. Coleoptera. In CSIRO, The Insects of Australia. Melbourne Univ. Press, Canberra, pp. 495-621. Burke, H. R. 1976. Observations on adult behavior of the lycid beetle Calopteron terminate (Coleoptera: Lycidae). Ent. News 87:229-232. Eisner, T. and F. C. Kafatos. 1962. Defense mechanisms of arthropods. X. A pheromone promoting aggregation in an aposematic distasteful insect. Psyche 69(2):53-61. Gottschalk, C. 1957. Untersuchungan fiber die Wirkung verschiedener Substanzen als Lock- oder Frassstoff auf einige Insektenarten. Beitrage zur Entomologie 7:177-179. Linsley, E. G., T. Eisner, and A. B. Klots. 1961. Mimetic assemblages of sibling species ol lycid beetles. Evolution 15:15-29. Young, D. K. and R. L. Fischer. 1972. The pupation of Calopteron terminate (Say) (Cole-| Withycombe, C. L. 1926. The biology of lycid beetles in Trinidad. Proc. Ent. Soc. Lond (TLM) New York State Museum, State Education Department, Albany, New York 12234, and (LMJ) Boyce Thompson Institute, Ithaca, New York 14853. Thorsteinson, A. J. 1960. Host selection in phytophagous insects. Ann. Rev. Entomol. 5: 193-1 218. ' B optera: Lycidae). Coleopt. Bull. 26:17-18. lit 1:32. Received for publication May 15, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4). 1979, pp. 289-298 NOTES ON A POPULATION OUTBREAK OE THE BEETLE UROPLATA SP. (COLEOPTERA: CHRYSOMELIDAE) ON THE TREE BUNCHOSIA PILOSA (MALPIGHIACEAE) IN COSTA RICA Allen M. Young Abstract. — An outbreak of the herbivorous beetle Uroplata sp. (Chrys- omelidae) on the tree Bunchosia pilosa H.B.K. (Malpighiaceae) in the cen- tral highlands of Costa Rica is described. During the early phase (June) of the rainy season, adult beetles were abundant on leaves of lower branches, 'where they feed by stripping away tissue from the upper side of leaves. Immature stages of the beetle were not seen, nor was any mating activity iobserved during the morning hours. Beetles do not feed on the very pilose undersides of the leaves. No other herbivores were seen. As the rainy sea- son advances, the Uroplata infestation spreads into the upper portions of [ the tree, perhaps as a response to increasing adult density and depletion of food supply in the lower region. Infestations were large during June and July (1974) but diminished by August. The phenomenon of population out- breaks in tropical herbivorous insects is discussed. ; In the American tropics, the traditional view of insect-plant interactions l,in relation to community structure has been one of biotically-controlled [herbivore populations seldom attaining population outbreak conditions jj(Pianka 1966). Therefore, population outbreaks of herbivorous insects should be documented and follow-up experimental studies conducted when- ever possible. The purpose of this paper is to call attention to a population outbreak of the beetle Uroplata sp.* (Coleoptera: Chrysomelidae) on the tree Bunchosia pilosa H.B.K. (Malpighiaceae) in the central highlands of Costa Rica. Owing primarily to the highly specialized and coevolved feeding relationships be- tween herbivorous insects and their host plants in the tropics (Janzen 1973) and the apparent maintenance of rather stable insect populations that fluc- tuate primarily with seasons in the tropics (e.g., Janzen and Schoener, 1968; Wolda, 1978a, b), the frequency of outbreaks of herbivores associated with any one host plant species is expected to be low. The documentation of ^ such outbreaks in the tropics provides information that can be used for further studies on the regulation, or lack thereof, of insect populations in * The taxonomy of Neotropical Uroplata is presently too unclear to assign a correct specific name to the species studied in this paper. 290 NEW YORK ENTOMOLOGICAL SOCIETY. the tropics. This paper describes some of the major features of the Buncho- sia-Uroplata interaction. I use the term “outbreak” in this paper to refer to a large population increase of a phytophagous insect species on a single individual of a tree species. In the traditional sense, outbreak refers to large numbers of insects on many plants in an area. In the tropics, given the very patchy spatial distribution of many tree species, outbreaks may be confined to only certain individuals of a certain species. Locality and Study Methods The interaction of Uroplata beetles with Biinchosia was studied June- August 1974 at “San Rafael de Ojo de Agua” (1,000 m elev.), Alajuela Province, Costa Rica, a region described as “tropical moist forest” (Hold- ridge, 1967). The study site was a single individual adult Bunchosia tree (about 10 m tall) growing along the Rio Segundo. At this locality the river is lined predominantly with Zygia longifolia (H.&B.) Britton & Rose (Le- guminosae) trees. The single Bunchosia was the only one of this species encountered in a 0.25 km river-edge survey (both sides) and no individuals were found in nearby pastures (back to 500 m from either side of the river). On June 26, at least several hundred adults of Uroplata were discovered on this tree during a census of emerging adult cicadas (Young, 1979). The flattened, almost rectangular black beetles (10 mm long) were easy to see on the leaves. This locality is highly seasonal in terms of the annual monthly pattern of rainfall; a distinct dry season, with little or no rain, occurs be- tween January and April and therefore the observations on Uroplata were made in the early rainy season. Observations were made on the Bunchosia-Uroplata interaction over a three-day period in June. The following types of information were recorded:! (1) description of damaged parts of the tree, (2) presence of beetles on other trees within 40 m to either side of the infested Bunchosia tree, (3) vertical distribution of the infestation on the tree including an examination for im-. mature stages, (4) intensity of the infestation in shady versus sunlight por-! ■ tions of the tree, (5) density of the beetles in sections of high and low infestation, and (6) percentages of leaves destroyed by the beetles on branches of high and low infestation. In addition, the infestation was ex-I amined during July and August. The presence of other feeding insects on the Bunchosia tree was also noted. To measure the intensity and vertical pattern of leaf damage from Uro-. plata, several branches were selected from both lower and upper portions of the tree; lower branches were those below 3 meters from the ground. Thei number of both healthy and damaged leaves on each of several branches; from the two regions was recorded as was the number of feeding beetles* per leaf. In areas of high and low infestation, an area 2 meters long by 1.5| VOLUME LXXXVll, NUMBER 4 291 Fig. 1. A branch of Bunchosia pilosa showing leaves damaged by feeding Uroplata beetles, i The light areas on the leaves are where the beetles have destroyed leaf tissue. I meters high was used to count the number of beetles present and feeding. As estimate of the vertical distribution of the infestation was made by count- ing the number of light brown patches from top to bottom of the tree. Owing to the large size of the leaves (120 cm^) and the conspicuousness of the brown patches against the dark green color of healthy tissue, it was easy to estimate the distribution of attack by counting damaged leaves with the naked eye and binoculars. Results Uroplata beetles were found only on the Bunchosia tree and not on other trees and herbaceous plants in the area. Only adult beetles were found on the tree and these fed in small groups (5-40 beetles) in neat rows. Adults scraped away tissue from the smooth upper side of a leaf. No beetles were seen feeding from the very pilose undersides. The result of this feeding behavior was that leaves damaged by the beetles had large areas of light brown dead tissue (Fig. 1) as beetles did not chew all the way through the 292 NEW YORK ENTOMOLOGICAL SOCIETY (01 Table 1. Some summary statistics describing the infestation by Uroplata beetles as her- | bivores of the tree Bunchosia pilosa. (1) Branch position (and leaves) and pattern of leaf destruction for 10 lower and 8 upper branches Mean no. leaves per branch (x ± S.D.) Mean no. leaves with >33% destroyed surface (X ± S.D.) Mean % damage (X) Lower 48.3 ± 22.7 20.9 ± 10.2 44.7 ± 9.62 Upper 45.9 ± 26.3 8.4 ± 8.1 12.9 ± 11.7 (2) Beetle density and approx. % destruction of leaves in 2 patches of high infestation Mean no. beetles per leaf (x ± S.D.) Mean % leaves destroyed, and range Patch no. 1 (12 leaves) Patch no. 2 (12 leaves) 8.3 ± 11.1 9.3 ± 4.5 57.5 ± 24.7 50.0 ± 10.0 (3) Beetle density and approx. % destruction of leaves in 2 patches of low infestation Mean no. beetles per leaf (x ± S.D.) Mean % leaves destroyed, and range Patch no. 1 (10 leaves) Patch no. 2 (15 leaves) 0.6 ± 1.5 1.5 ± 2.3 4.0 ± 4.6 1.3 ± 2.9 [ leaves. Feeding beetles were found during both morning and afternoon i hours. Nocturnal feeding was not determined. Using 33% of more destroyed leaf surface as an indicator of “intense herbivore damage” in this system, leaves on the lower branches were found to be more damaged than leaves on upper branches (Table 1). Leaves of the lower branches had many large brown spots (Fig. 2). In two different patches of intense feeding, the Uroplata density was between 8 and 9 bee- tles per leaf, and with 50% or more of the leaves destroyed (33% or more damage of leaf surface area) in the two samples (Table 1). In sharp contrast, in two patches of low predation, there was about one beetle per leaf (0-2 beetles) and less than 5% of the leaves were severely damaged (Table 1). The leaves of lower branches were more severely damaged than leaves on upper branches (Table 2). On lower branches in patches of intense preda- tion, the actual range in percentage of surface area destroyed per leaf was 30% to 90% even though beetle abundance per leaf was more variable (Table 3). No other insects were observed feeding on the leaves and no immature stages of Uroplata were present. No copulating pairs of beetles were seen. The greatest numbers of beetles were found on leaves in the shade. When a branch bearing leaves being attacked by the beetles was pulled into the I I ! VOLUME LXXXVII. NUMBER 4 293 1 Fig. 2. The lower branches of this Bunchosia pilosa tree had many leaves severely damaged by Uroplata beetles during the rainy season. direct sunlight, beetles crawled away within 2-8 minutes. During the morn- 1 ing hours, the beetles occurred in greatest numbers on the upper sides of ' leaves, and by 1:00 P.M. on sunny days, most beetles moved to the ventral ' sides of the leaves. Large aggregations (8-40 beetles) were found resting on the undersides of partially or completely destroyed (brown) leaves in the afternoon. Although the Uroplata infestation was high during June (June 26-28), the infestation was even greater on July 14 and 30 and at this time it had spread to the upper portion of the tree. By August 11 no beetles were present on the tree. Although, as reported above, the infestation of adult beetles on the tree was very large during the 1974 wet season, no outbreaks were seen on this tree over several other years, namely from 1971-1973 and again in 1975. The study site was visited several times in both wet and dry seasons in these years, and no outbreaks of Uroplata were seen. There were no out- ' breaks in three successive years preceding the outbreak year, or in the year following it. Furthermore, the appearance of outbreak numbers of adult 294 NEW YORK ENTOMOLOGICAL SOCIETY «1 Table 2. The distribution of the infestation of Uroplata beetles (Chrysomelidae) on a mature individual of the tree Bunchosia pilosa (Malpighiaceae) in central Costa Rica. Branch positions Branch no. No. leaves per branch No. leaves 33% destroyed % Destroyed “Low”* 1 19 8 42% 2 36 20 56% 3 27 14 52% 4 77 35 45% 5 83 42 51% 6 41 15 36% 7 43 19 44% 8 64 19 30% 9 26 15 58% 10 67 22 33% “High” 1 94 21 22% 2 72 15 21% 3 55 17 31% 4 41 7 17% 5 35 4 11% 6 30 1 0.3% 7 23 1 0.4% 8 17 1 0.6% * “Low” branches are those 3 meters or less above the ground; “high” branches are above 3 meters to top of canopy. beetles on the tree was seen only in the 1974 wet season, despite the fact that a distinct dry season occurs in the region. Discussion From studying herbivorous insects in the Central American tropics over the past 1 1 years, I attach the term “population outbreak” to the infestation of Uroplata beetles on Bunchosia since the numbers found were far greater than I have seen for Chrysomelidae on many plant species, and generally for other herbivorous insects in general. For the Chrysomelidae, in herba- ceous to slightly woody plants with a canopy of 2 meters or less (in sec- ondary succession), beetle numbers per plant generally ranged from 1-20 (pers. obs.). During the early rainy season in Costa Rica, new flushes of vegetative growth on many plant species promotes the growth of insect populations (Janzen and Schoener, 1968; Janzen, 1973) and herbivorous insects may synchronize emergence and peak adult abundance with periods of high availability of plant tissue utilized as food. In natural communities, predators and parasites on herbivorous insect species are also expected to have population cycles synchronized with their hosts (Allee et al., 1949). In plant communities where human or catastrophic I VOLUME LXXXVll, NUMBER 4 295 Table 3. The distribution of Uroplata beetles in two patches* of lower branches of the tree Bunchosia pilosa where infestations were high.** Bunchosia Leaf code No. beetles leaf patch no. present % Leaf area destroyed 1 1 3 90% 2 39 75% 3 3 90% 4 0 30% 5 9 30% 6 0 40% 7 20 75% 8 2 20% 9 6 45% 10 6 45% II 9 50% 12 2 90% 2 1 II 60% 2 9 50% 3 3 40% 4 4 60% 5 4 50% 6 12 50% 7 II 70% 8 7 40% 9 8 30% 10 8 50% 11 12 50% 12 9 60% 13 7 40% 14 14 50% 15 21 50% * Each “patch” has a one-by-one meter area of dense leaves within the lower branch region of the tree. ** The areas of high beetle densities were selected in the lower branch region at the time of census. Censuses in both patches were conducted between 9:50 A.M. and 1:00 P.M. on the same day, with about equal time spent counting beetles and leaf damage in each one. I disturbance has resulted in a reduction of plant species diversity locally, the I probability of outbreaks of herbivorous insects is greater than in commu- inities characterized by higher plant diversity (Pimentel, 1961). Climatic ex- planations of insect outbreaks are generally confined to regions character- ized by harsh environmental conditions with large fluctuations annually (Elton 1927). Such effects are generally not expected in the tropics, although the interaction of (1) highly synchronized population cycles in seasonal en- vironments and (2) the long-term reduction in host plant diversity are ex- ' pected to result in population outbreaks of herbivorous insects in the trop- 296 NEW YORK ENTOMOLOGICAL SOCIETY ics. Is this what is occurring the the Biinchosia-U roplata system in central Costa Rica in the early rainy season? With more data from several trees, I believe that the answer to this question would be “yes.” Regions of the world with considerable variations in annual rainfall pro- mote large fluctuations in insect abundance (Birch, 1957; Wolda, 1978b). ' The central highlands of Costa Rica represent seasonal environments in terms of rainfall, and year-to-year variations in rainfall may contribute to \ fluctuations in insect populations, perhaps in some years pushing some ' species into the population abundance zone of outbreak conditions. Con- sistent and comparable annual rainfall data for San Rafael de Ojo de Agua are not available for such a correlation for the Bunchosia-Uroplata inter- ' action. Sixty genera and approximately 800 species of woody vines, shrubs , and trees comprise the Neotropical Malpighiaceae, and Bunchosia is one , of the largest genera (Hartshorn 1971). Bunchosia pilosa, known in Costa* Rica as “Cerezo” or “Orquetilla,” occurs most commonly between 110-^ 1 ,800 meters above sea level and ranging from Costa Rica to Colombia (Standley 1937). Hartshorn (1971) mentions that B. pilosa is rare at “Finca La Selva” in the northeastern lowland rain forest (98 m elev.) region of' Costa Rica. This tree generally has a patchy spatial distribution in highland forest environments and as such, it is probably a highly dispersed and in- ; conspicuous resource for host-specific herbivores. Furthermore, in the cen- ^ tral highlands of Costa Rica, a large percentage of natural forest has been : removed as the result of human activity, contributing to the scarcity of this tree at localities such as San Rafael de Ojo de Agua. At this locality, forest trees are presently limited to the margins of rivers and streams, sometimes i in steep gulleys. If this particular species of Uroplata is host-specific for 1 Bunchosia, infestations of this herbivore will also be patchy. In highly dis- i turbed habitats, the interactions between Buchosia and Uroplata could re- ; r suit in outbreaks of the beetle if reduction in plant species diversity has alsoljl resulted in a reduction in the availability of predators and parasites of the o beetle (see Pimentel, 1961 for a general discussion). ' The data suggest that the Uroplata infestation begins and mushrooms in the lower region of the Bunchosia tree. The spread of the infestation into n upper portions of the tree later could be a density-related response to in-|r creased adult densities and decreasing food supply in the lower region during the rapid growth of the adult beetle population. Although the outbreak numbers of adult beetles occurred during the early phase of the 1974 wet season, a large larval population must have been present prior to this period. The life cycle of Uroplata beetles is not known, and as pointed out by Arnett (1963), most of the species in the tribe Uro- ■ platini of the subfamily Hispinae (about 1,500 species world-wide) are trop- ical and very poorly described and far less understood. Whether or not there . is an actively growing larval population in the dry season, or whether the 1 VOLUME LXXXVll, NUMBER 4 297 I dry season is passed as ai diapausing egg, larva, or pupa, remains to be studied. In some species of phytophagous beetles, adult numbers are reg- I ulated by interspecific competition, and sometimes the intensity of such competition is influenced by local climatic factors (Utida, 1961). Populations ' of a host-specific temperate-zone weevil Sitona regensteinensis are regu- lated by heavy mortality of eggs and larvae from parasites and predators (Danthanarayana, 1969). It is sometimes the case that both density-depen- i dent and density-independent factors regulate populations of phytophagous I beetles (Parnell, 1966; Beaver, 1967). Part of the explanation of outbreaks I of locusts is related to annual variations in rainfall patterns in target areas (Gunn and Symmons, 1959). A sudden outbreak of Uroplata beetles on i Bunchosia could result from very favorable climatic conditions affecting larval populations. Furthermore, given the highly disturbed condition of the habitat, predators and parasites of Uroplata may no longer be present, reducing the operation of density-dependent mortality factors and increasing ! the role of density-independent factors such as rainfall or lack thereof. In the Chrysomelidae, the larvae often feed on the same host plants as the I adults (Huffaker, 1953) and can sometimes become severe defoliators when [ regulatory factors are absent. Given the high densities of Uroplata adults, ! and although the sample is small, larval survival must have been very high, ! suggesting an absence of density dependent control or unfavorable climatic I conditions in this particular year. As the outbreak was not present in other , years, variations in climatic conditions may affect predators and parasites I of Uroplata on this tree. Acknowledgments This research was a by-product of National Science Foundation grant 1 GB-33060. Logistical support was provided by Dr. J. Robert Hunter, former I Director of the Costa Rican field studies program of the Associated Colleges , of the Midwest. David Ewart and Karen Kinsell assisted with the field work. ! The beetle was identified by Dr. Richard E. White of the Systematic Ento- mology Laboratory of the U.S. National Museum, and the tree was iden- tified by Luis Poveda of the Herbarium at the National Museum of Costa Rica. Literature Cited Alice, W. C., A. E. Emerson, O. Park, T. Park, and K. P. Schmidt. 1949. Principles of Animal Ecology. Philadelphia: W. B. Saunders Co., 837 pp. Arnett, R. H. 1963. The beetles of the United States. Washington, D.C.: Catholic Univ. America Press, 1 1 12 pp. Beaver, R. A. 1967. The regulation of population density in the bark beetle Scolytiis scolytus (F.). J. Anim. Ecol. 36:435^51. 298 NEW YORK ENTOMOLOGICAL SOCIETY Birch, L. C. 1957. The role of weather in determining the distribution and abundance of animals. Cold Spring Harb. Symp. Quant. Biol. 22:203-218. Danthanarayana, W. 1969. Population dynamics of the weevil Sitona regensteinensis (Hbst.) on broom. J. Anim. Ecol. 38:1-18. Elton, C. 1927. Animal Ecology. London: Sidgwick and Jackson, Ltd., 209 pp. Gunn, D. L. and P. M. Symmons. 1959. Forecasting locust outbreaks. Nature 184:1425-1427. Hartshorn, G. S. 1971. Species list of trees. La Selva, Heredia, Costa Rica. /n Handbook for Tropical Biology in Costa Rica. San Jose, Costa Rica: Organ. Tropical Studies. Holdridge, L. R. 1967. Life zone ecology. San Jose, Costa Rica: Tropical Sci. Center. Huffaker, C. B. 1953. Quantitative studies on the biological control of St. John’s wort (Kla- f math weed) in California. Proc. Seventh Pacif. Sci. Congress 4:303-313. Janzen, D. H. 1973. Comments on host-specificity of tropical herbivores and its relevance to •• species richness, pp. 201-211. In Taxonomy and Ecology (V.H. Heywood, ed.). Syst. jl Assoc. Spec. Vol. 5. New York: Academic Press. and T. W. Schoener. 1968. Differences in insect abundance and diversity between wetter and drier sites during a tropical dry season. Ecology 49:96-110. Parnell, J. R. 1966. Observations on the population fluctuations and life histories of the beetles Bruchidius ater (Bruchidae) and Apion fuscirostre (Curculionidae) on broom (Saro- thamnus scoparius). J. Anim. Ecol. 35:157-188. Pianka, E. R. 1966. Latitudinal gradients in species diversity: a review of concepts. Amer. Nat. 100:33^6. Pimentel, D. 1961. Species diversity and insect population outbreaks. Ann. Entomol. Soc. Amer. 54:76-86. Standley, P. C. 1937. Flora of Costa Rica. Vol. XVIII. Chicago: Field Museum Nat. Hist. Bull., 780 pp. Utida, S. 1961. Experimental studies on the interaction between the bean weevils and their, parasitic wasps. XI Internat. Kongr. Entomol., Wien 1:731-734. ^ |l Wolda, H. 1978a. Seasonal fluctuations in rainfall, food and abundance of tropical insects. J. i, Anim. Ecol. 47:369-381. ‘J . 1978b. Fluctuations in abundance of tropical insects. Amer. Nat. 1 12: 1017-1045. ^ Young, A. M. 1979. Habitat and seasonal relationships of some cicadas (Homoptera: Cicad- ^ idae) in central Costa Rica. Amer. Mid. Nat. 98: in press. jl Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin H 53233. Received for publication June 18, 1979. ! p; NEW YORK ENTOMOLOGICAL SOCIETY LXXXV1K4), 1979, pp. 299-303 FIELD STUDIES AND PARASITES OF LIRIOMYZA TRIFOLIEARUM (DIPTERA: AGROMYZIDAE) IN NORTHEASTERN USA R. M. Hendrickson, Jr. Abstract. — Liriomyza trifoliearum Spencer, a native serpentine leafminer of alfalfa in the USA and Canada exists at low population densities. In 1978 the density averaged 0.2 mines per stem with a maximum of 1.3 mines per stem. It is controlled biologically (63% parasitism over a 4-year period) by 17 native parasite species plus 2 from Europe that recently were established against Agromyza frontella (Rondani). A list is provided of recovered par- asite species and numbers of hosts parasitized by each species. Liriomyza trifoliearum Spencer, a native serpentine leafminer that is not of economic importance, has been reported from Florida, California, and Prince Edward Island, Canada (Spencer, 1973). We found it on alfalfa throughout the northeastern USA and in Ontario, Canada; thus, its distri- bution is probably Nearctic on alfalfa. It also has been recovered from Pisum sativum L., Trifolium incarnatum L., and T. repens L. (Spencer, 1973). The present study was conducted in Delaware, south-central New Jersey, and southeastern Pennsylvania, coincidentally with studies of the alfalfa blotch leafminer (ABL), Agromyza frontella (Rondani), an imported European pest of alfalfa. The purpose of our study was to present basic observations on field populations of L. trifoliearum on alfalfa and compile a list of species of recovered parasites with numbers of parasitized hosts. The order, family, and identifier of species mentioned in this paper appear in Table 1. In studying L. trifoliearum, we had difficulty separating its larvae or pu- paria from those of other agromyzid species infesting alfalfa: ABL, L. tri- folii, and the vegetable leafminer, L. sativae. (Liriomyza sativae was never collected by us in the northeastern USA, but it was reported from Ohio (Spencer, 1973) and from greenhouses in Canada (McClanahan, 1978), from which it might escape.) We made no effort to separate these species by larval morphology, since many larvae were completely consumed by de- veloping parasites. Agromyzid species were distinguished by mine type, frass distribution, and pupation habit (Table 2). In both field and laboratory studies, we observed that L. trifoliearum always pupated in the leaflet. When isolated leaflets containing puparia were placed in tight-fitting petri dishes, the epidermis of the drying leaflet occa- sionally ruptured, and the puparium dropped out after formation. This is 300 NEW YORK ENTOMOLOGICAL SOCIETY Table I. Order, family, and identifier of species mentioned in this paper. Species Species HYMENOPTERA: BRACONIDAE Dacnusa dryas (Nixon)"* Opius dimidiatus (Ashmead)® HYMENOPTERA: PTEROMALIDAE Bubekia fallax Gahan** Halticoptera circiilus (Walker)' Halticoptera laevigata Thomson® DIPTERA: AGROMYZIDAE Agromyza frontella (Rondani)"’ Liriomyza sativae Blanchard"* Liriomyza trifoUearum Spencer"* Liriomyza trifolii (Burgess)"* HYMENOPTERA: EULOPHIDAE Achrysocharella formosa (Westwood)® Chrysocharis clarkae Yoshimoto® Chrysocharis giraulti Yoshimoto® Chrysocharis pubicornis (Zetterstedt)® Chrysocharis punctifacies Delucchi® Closterocerus cinctipennis Ashmead** Closterocerus tricinctus (Ashmead)** Diaulinopsis callichroma Crawford* Diglyphus intermedins (Girault)** j Diglyphus pulchripes (Crawford)** j Diglyphus websteri (Crawford)' Pediobius bucculatricis (Gahan)** Pnigalio minio (Walker)® Zagrammosoma midtilineatum (Ashmead)** | ® P. M. Marsh, ** G. Gordh, ® E. E. Grissell, and "* G. C. Steyskal, Systematic Entomology n Laboratory, Agricultural Research, Science and Education Administration, USDA, c/o U.S. . National Museum, Washington, DC 20560. ® C. M. Yoshimoto, Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario ■ ’i KIA 0C6. ' R. M. Hendrickson, Jr. perhaps what Spencer (1973, p. 99) referred to when he said, “Pupation appears to be variable, either in or outside the mine.” During 3 seasons (1975-77), we collected mined leaflets and recovered parasites at 7 fields (1 at Newark, DE; 3 near Oxford, PA; and 3 near Rancocas, NJ) by the method of Hendrickson and Barth (1979). The same procedure was followed in 1978 except the collection of 50 mined leaflets from each field was limited to Liriomyza spp. only, which were separated to species at the laboratory. In 1978, we also collected 20 alfalfa stems from each field and kept them fresh in a plastic bag in a car refrigerator. At the laboratory, the mines of ABL, L. trifoUearum, and L. trifolii were counted, disregarding the condition of the mining larvae. The ratio of mines of ABL:L. trifoliearurrv.L. trifolii was ca. 1800:40:1. Liriomyza trifolii was first found in mid-July 1978, on late 2nd-cutting alfalfa, and thereafter infre- quently through October. It was probably present before July, but we did not encounter it, perhaps because the density was extremely low. The average season-long density of L. trifoUearum was 0.2 mines/stem; maximum density was 1.3 mines/stem. If a leaflet was mined, we usually i VOLUME LXXXVII, NUMBER 4 301 Table 2. Some characteristics for distinguishing species of Agromyzidae found on alfalfa in northeastern USA. Species Mine type Frass distribution Pupation site Agromyza frontella 1st and 2nd instars linear broad strips 3rd instar blotch broad strips soil Liriomyza trifoliearum serpentine isolated lumps leaflet Liriomyza trifolii serpentine narrow strips soil Liriomyza sativae^ serpentine narrow strips soil ® A species which was potentially collectible, but that we never found in this area. found 1 mine/leaflet; the maximum was 4 mines/leaflet. We found L. trifol- ieanim first in late May ca. 7-10 days after the first appearance of ABL mines and often in the same leaflet as ABL larvae. This was not surprising, since ABL larvae may mine >50% of the leaflets in a field at peak infesta- tions. Liriomyzci trifoliearum makes feeding perforations through either the up- per or lower leaflet epidermis. These perforations are angled rather than perpendicular, and do not penetrate to the opposite epidermis. When viewed against the sky, they appear light green. In contrast, ABL feeding perfo- rations are always made through the lower leaflet epidermis, are perpendic- ular, and penetrate to or through the upper leaflet epidermis, giving the impression of clear “pinholes” when viewed against the sky. Liriomyzo trifoliearum pupates in the alfalfa leaflet. Thus at each harvest, some puparia are removed from the field. However, we think that most of the puparia remain in the field on the soil in dehisced leaflets. Nearly all leaflets have dropped from the lower half of stems by harvest because of senescence. Any puparia in these leaflets are on the soil. Some puparia in leaflets on the upper half of the stem are also on the soil, according to our observation that leaflets mined by agromyzids dehisce before adjacent un- damaged leaflets. This inclination is exacerbated during rainy weather, when moisture accelerates development of secondary plant pathogens in mines, and rain and wind cause mechanical dehiscence. Parasitism of L. trifoliearum in 1978 was: 1st cutting, 28% (131 live forms); 2nd cutting, 38% (373 live forms); 3rd cutting, 74% (188 live forms); and post-3rd cutting, 79% (577 live forms). (Live forms are leafminer larvae or puparia that produce either adult L. trifoliearum or parasites.) For the 4 sampling periods, parasitism averaged 55% and totaled 62% (1,269 live forms). For the 4 seasons (1975-78), total parasitism was 63% (2,291 live forms); the parasite species and numbers of parasitized hosts appear in Table 3. To our knowledge, these were all primary parasites. 302 NEW YORK ENTOMOLOGICAL SOCIETY ,% Table 3. Parasite species and numbers reared from Liriomyza trifoliearum collected in i Delaware, southern New Jersey, and southeastern Pennsylvania, 1975-78. Recovered from Number of hosts® Species Larva Pupa Diglyphus intermedius X 372 Opius dimidiatus X 266 Chrysocharis clarkae X X 202 Chrysocharis giraulti X X 183 Diglyphus pulchripes X 91 Halticoptera circulus X 40 Halticoptera laevigata X 30 Pnigalio minio X 30 Closterocerus tricinctus X 28 Diaulinopsis callichroma X 28 Achrysocharella formosa X 22 Closterocerus cinctipennis X 21 Diglyphus websteri X 14 Pebiobius bucculatricis X 6 Chrysocharis punctifacies^ X 4 Dacnusa dryas'’ X 3 Bubekia fallax X 2 Chrysocharis pubicornis X 2 Zagrammosoma multilineatum Unidentifiable‘S X Total 1 94 1,439 ^ Occasionally more than 1 parasite emerged from a host. ’’ Introduced European species released against Agromyza front eUa. ' Adults were damaged or lost, or larvae were in diapause. The European parasites Dacnusa dryas and Chrysocharis piinctif tides, » established in 1978 against ABL in Delaware (Hendrickson, 1978), were k both recovered from L. trifoliearum in 1978. Thus this serpentine leafminer, 1. found throughout the range of ABL, may serve as an effective alternate h host in the biological control of ABL. Although we never observed predation, we found indirect evidence of it. 1. One or more small, round punctures through the leaflet epidermis over the t1 integumental remains of L. trifoliearum larvae suggested that one or more t of the hemipteran predators that are found on alfalfa had fed on the larvae, i These punctures were distinctly different from feeding perforations made L by adult female agromyzids. Hendrickson and Barth (1979) reported that 12 of the 14 species of native parasites that attack ABL were derived from native Liriomyza spp. on al- ; falfa. These 12 species were all recovered from L. trifoliearum, and some r were recovered from L. trifolii. In 1978, we recovered a single individual * of a 13th species, Zagrammosoma multilineatum, from L. trifoliearum. The I VOLUME LXXXVII, NUMBER 4 303 finding that in 1978 the larval population of L. trifoliearum was ca. 40 times that of L. trifoHi indicated that the native parasite complex that attacked ABL was derived almost entirely from the parasites of L. trifoliearum. Our data indicated that L. trifoliearum is not of economic importance on alfalfa in the northeastern USA and Canada because it is kept at low pop- ulation densities by 19 species of parasites and one or more species of predator. Acknowledgment This research was conducted with the technical assistance of Susan E. Barth. Literature Cited Hendrickson, R. M., Jr. 1978. Establishment of Dacnusa dryas (Nixon) (Hymenoptera: Bra- conidae) and Chrysocharis punctifacies Delucchi (Hymenoptera: Eulophidae), parasites of Agrornyza frontella (Rondani) (Diptera: Agromyzidae) in Delaware. J. N.Y. Entomol. Soc. 86:295. (Abstract) and S. E. Barth. 1979. Effectiveness of native parasites against Agrornyza frontella, an introduced pest of alfalfa. J. N.Y. Entomol. Soc. 87:85-90. McClanahan, R. J. 1978. Biological control of leafminers. Can. Agr. 24:27-28. Spencer, K. A. 1973. Agromyzidae (Diptera) of economic importance. In Series Entomolo- gica, Vol. 9, E. Schimitschek [ed.], W. Junk, The Hague, xi + 418 pp. Beneficial Insects Research Laboratory, Agricultural Research, Science and Education Administration, USDA, Newark, Delaware, 19713. Received for publication July 11, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4), 1979, pp. 304-311 THREE NEW MIDDLE AMERICAN SPECIES OE AQUATIC BEETLES IN THE GENUS NOTIONOTUS SPANGLER (HYDROPHILIDAE: HYDROBIINAE) Philip D. Perkins Abstract. — Three new species of the aquatic beetle genus Notionotus Spangler (Hydrophilidae) are described, one each from Mexico, Guatemala and Panama. Shared characteristics of these species which differ from those of the two previously described species from Venezuela are discussed and illustrated with scanning electron micrographs. Aedeagi of the new species, N. mexicanus, N. nucleus and N. tricarinatus are illustrated. Habitat pref- erences of N. mexicanus and N. nucleus are discussed, illustrated and contrasted with those of Venezuelan species. Introduction The aquatic beetle genus Notionotus was erected by Spangler (1972) for two species of Hydrophilidae from Venezuela. Described herein are three new species of Notionotus from Middle America, including one species each from Mexico, Guatemala and Panama. Based upon external similari- ties, these Middle American species form a monophyletic group and con- stitute the first known members of a lineage which possibly has a sister- group relationship with the Venezuelan species. The three Middle American species have the mesosternum longitudinally carinate in the midline, where- as in the Venezuelan species the median region is flat (cf. Figures 1, 4). The hind femur of Middle American species is more extensively pubescent, hav- ing only the apical V4 lacking pubescence, whereas the hind femur of Ven- ezuelan species lacks hydrofuge pubescence in the lower V2 of the midregion as well as apically (Figure 2; see also figure 6 in Spangler, 1972). The front femur of all 5 species lacks hydrofuge pubescence on the upper surface basally, although this region is alutaceous in some (Figure 2, cf. figure 4 in Spangler, 1972). Additionally, the known Middle American species have testaceous elytra in contrast to the Venezuelan species which are either banded basally with reddish brown {rosalesi Spangler) or completely piceus (liparus Spangler). There also appears to be differences in habitat preferences of the two putative sister-groups. Spangler (1972) collected the Venezuelan species in madicolous habitats such as “on the wet surfaces of rocks, in crevices, and on leaves in spring seepage areas in road cuts.” However, Notionotus that my wife Maureen and I collected in Mexico and Guatemala were not found VOLUME LXXXVIl, NUMBER 4 305 Figs. 1-4. 1, Notionotus liparus Spangler, ventral aspect (llOx); 2, as above (55 x); 3, N. nucleus new species, lateral aspect of meso- and metathorax (230x); 4, as above, ventral aspect (I95x). in madicolous habitats. Specimens of mexicanus, new species, were col- lected from plant debris which had become trapped between stones in a rapid stream (Figure 8); specimens of nucleus, new species, were collected by stirring small stones, gravel and plant debris at the margin of a very small, slowly moving brook in dense vegetation, allowing the disturbed ma- terial to drift into an aquatic net, and carefully sorting the contents. My suspicions that Notionotus species from Middle America are not strictly I 306 NEW YORK ENTOMOLOGICAL SOCIETY \ hygropetric are reinforced by the fact that my wife and I collected many r specimens of the strictly madicolous hydrophilid Oocyclus in several local- t ities in Guatemala and Mexico, but were unable to find a single specimen 5 of Notionotus in such habitats. Key to Species of Notionotus 1. Mesosternal process flat or weakly rounded (Figs. 1, 2); pubescence of hind femur less developed, absent from lower Vi of midregion as well as apically (Fig. 2); elytra darkly colored, at least basally; Ven- ezuela 2 - Mesosternal process carinate (Fig. 4); pubescence of hind femur more developed, absent only in apical 14; elytra testaceous; Middle America 3 j 2. Elytra piceus liparus Spangler | - Elytra with reddish brown fascia at base, remainder testaceous | rosalesi Spangler 3. Head punctation finer and sparser, punctures separated by about 5 times their width; size larger, about 1.10 mm; aedeagus as illustrated (Eig. 5); Mexico mexicanus n. sp. - Head punctation coarser and denser, punctures near eyes separated by their width, others by 3-4 times their width; size smaller, about 1.00 mm; Guatemala and Panama 4 4. Aedeagus as illustrated (Eig. 6); Guatemala nucleus n. sp. - Aedeagus as illustrated (Eig. 7); Panama tricarinatus n. sp. ' Notionotus mexicanus Perkins, n. sp. (Eigure 5) Type~data. — Holotype (male). Mexico, Oaxaca, 8 miles E. Tapanatepec, ; tropical stream with large boulders, 3-VII-1974, M. E. and P. D. Perkins. ' Deposited in the National Museum of Natural History, Smithsonian Insti- tution. Paratype. — (1 female). Same data as holotype. • Diagnosis. — This species is slightly larger than nucleus and tricarinatus i ( 1 . 10 vs. 1.00 mm), has the head punctation distinctly finer and sparser than | those species, and differs in aedeagal form (Eigure 5). | Description of holotype. — Length 1.10 mm, greatest width 0.65 mm at I midlength. Color testaceous dorsally except for brown between eyes and | two small black spots near hind margin of pronotum, separated by about Vs I width of pronotum; mouthparts, antennae and most of legs testaceous, re- ( mained of venter reddish brown. Head shining, finely sparsely punctate, punctures separated by 5 times n their width. Clypeus expanded and shelflike in front of eyes, covering most VOLUME LXXXVIl, NUMBER 4 307 Figs. 5-7, Notionotus aedeagi, holotypes (dorsal and lateral views). 5, mexicanus \ 6, nu- cleus', 7, tricurimitus. of labrum, very finely alutaceous along anterior margin. Labrum shallowly emarginate medially. Ventral surface of head finely alutaceous behind eyes and in gular region; mentum shiny, finely punctate. Pronotum almost 2.5 times as wide as long; punctures finer and sparser than those on head; narrowly margined laterally; anterolateral and postero- lateral angles rounded. Prosternum longitudinally carinate in midline, acute- ly angulate at apex. Prosternal process distinctly produced posteriorly, dis- tinctly separating front coxae; apex concave for reception of mesosternal protuberance. Elytra convex, narrowly margined laterally; widest at anterior 14, ex- tremely finely sparsely punctate similar to pronotum. Sutural stria absent. Scutellum a small equal-sided triangle. Epipleura almost vertical. Mesosternum with prominent triangular protuberance whose midline is carinate and on same plane as metasternum. Metasternum smooth and im- punctate on swollen medial region, medial region lacking pubescence, ad- jacent lateral areas with sparse, rather long pubescence; apex of medial region broad between mesocoxae. Abdominal sterna finely alutaceous and moderately densely covered with short pubescence; last segment with a group of tiny golden setae set in small apicomedial emargination. Eront legs with femora pubescent along lower margin in basal %, upper margin alutaceous in basal %, apical 14 smooth and shiny. Middle and hind legs alutaceous and pubescent in basal V.5, apical '/s smooth and shiny. Variation. — The single female known has the pubescence at the borders of the swollen medial area of the metasternum shorter and sparser than the holotype. 308 NEW YORK ENTOMOLOGICAL SOCIETY Distribution. — Currently known only from the type-locality in southern- most Oaxaca, Mexico. Notionotus nucleus Perkins, n. sp. (Figures 3, 4, 6) Type-data. — Holotype (male). Guatemala, Alta Verapaz, 5 miles W. La Tinta, small tropical brook, 6-VI-1974, M. E. and P. D. Perkins. Deposited in the National Museum of Natural History, Smithsonian Institution. Paratypes. — (4 males, 7 females). Same data as holotype, deposited in NMNH and the author’s collection. Diagnosis. — Distinguished from mexicanus by its slightly smaller size, more strongly punctate head, and aedeagus (Figure 6). Aedeagal form must be used to differentiate nucleus and tricarinatus. Description of holotype. — Length 1.00 mm, greatest width 0.60 mm at midlength. Color testaceous dorsally except for brown between eyes and two small black spots near hind margin of pronotum, separated by about Vi width of pronotum; mouthparts, antennae and most of legs testaceous, re- mainder of venter reddish brown. Head distinctly punctate, some punctures near eyes separated by only their widths, others by 3-4 times their width. Clypeus expanded and shelf- like in front of eyes, covering most of labrum, very finely alutaceous along anterior margin. Labrum shallowly emarginate medially. Ventral surface of head finely alutaceous behind eyes and in gular region; mentum shiny, finely punctate. Pronotum almost 2.5 times as wide as long; punctures finer and sparser than those on head; narrowly margined laterally; anterolateral and postero- lateral angles rounded. Prosternum longitudinally carinate in midline, acute- ly angulate at apex. Prosternal process distinctly produced posteriorly, dis- tinctly separating front coxae; apex concave for reception of mesosternal protuberance. Elytra convex, narrowly margined laterally; widest at anterior 14, ex- tremely finely sparsely punctate similar to pronotum. Sutural stria absent. Scutellum a small equal-sided triangle. Epipleura almost vertical. Mesosternum with prominent triangular protuberance whose midline is carinate and on same plane as metasternum. Metasternum smooth and im- punctate on swollen medial region, medial region lacking pubescence, ad- jacent lateral areas with sparse, rather long pubescence; apex of medial region broad between mesocoxae. Abdominal sterna finely alutaceous and moderately densely covered with short pubescence; last segment with a group of tiny golden setae set in small apicomedial emargination. Front legs with femora pubescent along lower margin in basal %, upper VOLUME LXXXVIl, NUMBER 4 309 Fig. 8. Biotope of Notionotus mexicanus, Mexico, Oaxaca, 8 miles E. Tapanatepec. margin alutaceous in basal %, apical V3 smooth and shiny. Middle and hind legs alutaceous and pubescent in basal apical 'k smooth and shiny. Variation. — The 12 specimens studied were quite homogeneous. Distribution. — Currently known only from the type-locality in southeast- ern Guatemala. Etymology. — Latin, nucleus, kernel. I refer to the small size, smooth convex dorsum and testaceous color of this species, and also to its distri- bution in the region known biogeographically as “nuclear” Middle America. Notionotus tricarinatus Perkins, n. sp. (Figure 7) Type-data. — Holotype (male). Panama, Canal Zone, Albrook Forest Site, 22-III-1968, R. S. Hutton. Deposited in the National Museum of Natural History, Smithsonian Institution. 310 NEW YORK ENTOMOLOGICAL SOCIETY V Paratypes. — (6 males, 10 females). Same data as holotype, deposited in collections of NMNH, David C. Miller and author. Diagnosis. — Smaller and with punctation of the head more developed than mexicaniis. Aedeagal form (Figure 7) must be used to reliably differ- 1 entiate tricarinatus and nucleus. Description of holotype. — Length 0.98 mm, greatest width 0.55 mm at midlength. Color testaceous dorsally except for brown between eyes and two small black spots near hind margin of pronotum, separated by about Vs width of pronotum; mouthparts, antennae and most of legs testaceous, re- mainder of venter reddish brown. Head distinctly punctate, some punctures near eyes separated by only their width, punctures near midline separated by 3-4 times their width. Clypeus expanded and shelflike in front of eyes, covering most of labrum, very finely alutaceous along anterior margin. Labrum shallowly emarginate medially. Ventral surface of head finely alutaceous behind eyes and in gular region; mentum shiny, finely punctate. Pronotum almost 2.5 times as wide as long; punctures finer and sparser than those on head; narrowly margined laterally; anterolateral and postero- lateral angles rounded. Prosternum longitudinally carinate in midline, acute- ly angulate at apex. Prosternal process distinctly produced posteriorly, dis- tinctly separating front coxae; apex concave for reception of mesosternal protuberance. Elytra convex, narrowly margined laterally; widest at anterior !4, ex- tremely finely sparsely punctate similar to pronotum. Sutural stria absent. Scutellum a small equal-sided triangle. Epipleura almost vertical. Mesosternum with prominent triangular protuberance whose midline is carinate and on same plane as metasternum. Metasternum smooth and im- punctate on swollen medial region, medial region lacking pubescence, ad- jacent lateral areas with sparse, rather long pubescence; apex of medial region broad between mesocoxae. Abdominal sterna finely alutaceous and moderately densely covered with short pubescence; last segment with a group of tiny golden setae set in small apicomedial emargination. Eront legs with femora pubescent along lower margin in basal Vs, upper margin alutaceous in basal Vs, apical Vs smooth and shiny. Middle and hind legs alutaceous and pubescent in basal ‘‘/s, apical Vs smooth and shiny. Variation. — Some specimens are brownish dorsally. Distribution. — Currently known only from the Canal Zone, Panama. Etymology. — Latin, tricarinatus, in reference to the tricarinate mesoster- nal protuberance. Acknowledgments I thank my wife Maureen for assistance with fieldwork in Mexico and VOLUME LXXXVII. NUMBER 4 311 Guatemala. I am grateful to David C. Miller for the opportunity to study specimens from Panama. Thanks are due technicians of the Scanning Elec- tron Microscope Laboratory, Smithsonian Institution, for taking the micro- graphs. Paul J. Spangler kindly reviewed the manuscript. Travel funds for fieldwork in Middle America were provided, in part, by the Smithsonian Institution. Literature Cited Spangler, P. J. 1972. A new genus and two new species of madicolous beetles from Venezuela (Coleoptera: Hydrophilidae). Proc. Biol. Soc. Wash. 85:139-146. Department of Entomology, Smithsonian Institution, Washington, D.C. 20560 U.S.A. Received for publication July 20, 1979. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4), 1979, pp. 312-318 REVIEW: TABANIDAE OF THE EAST COAST AS AN ECONOMIC PROBLEM Elton J. Hansens*’^ Abstract. — Tabanidae are pests of man and animals in many areas of the coastal states but especially near salt marshes. The major species, Tahanus nigrovittatus and Chrysops atlanticus, move from the marshes to nearby beaches, camp grounds, golf courses, and other recreational areas and onto boats in the bays and estuaries. Chrysops congregate in dense vegetation and attack when humans or animals move into such places. Both Tahanus and Chrysops are severe problems to agricultural workers when the flies are numerous. Livestock are readily attacked by Tabanidae with consequent effects on thriftiness, weight gains and milk production and possible trans- mission of causal agents of disease. Biology and habits of both salt marsh and upland species are poorly known. Probably T. nigrovittatus is a species complex. Controls are inadequate though traps and vegetative barriers have been shown useful against Tahanus and some insecticides have given re- duction but not adequate control of both Tahanus and Chrysops. I When one thinks of the blood-sucking Diptera of the east coast one thinks first of mosquitoes as the major problem. However, in many areas 50 to 75 years of organized control programs have resulted in only sporadic mosquito annoyance. Other biting flies are then regarded as greater pests such as Tabanidae, both greenheads (Tahanus) and deer flies (Chrysops). Tabanidae as Pests tdi The Tabanidae are selective feeders and only a few are important pests of humans. Especially the larger Tahanus seem to limit their attacks to large mammals. Only female Tabanidae take blood meals needed for maturation of eggs. Some species of both Tahanus and Chrysops are autogenous, the females depositing the first egg mass before seeking a blood meal. Nutrients for normal fly activities are obtained from flowers and other plant sources or from food reserves stored in the larval stage. The relatively few species of Tabanidae that bite man often occur in large )c ml) Ool ' Diptera: Tabanidae ^ Paper of the Journal Series, New Jersey Agricultural Experiment Station, Cook College, Rutgers University, New Brunswick, N.J. This work was performed as part of NJAES Project No. 409, supported by the New Jersey Agricultural Experiment Station. VOLUME LXXXVIl, NUMBER 4 313 numbers and make life miserable. People feel the bite when the female pierces the skin or probes deeply for an adequate flow of blood. Usually no swelling, reddening or irritation occurs after the bite but some people have moderate reactions which last from a few minutes to several hours. A small segment of the population respond with allergic reactions involving exten- I sive swelling, erythema, itching, and related effects. In extreme cases hos- i pitalization is necessary and emergency measures to counteract allergic ef- i fects or anaphylactic shock are essential. ' This tabanid problem is primarily associated with salt marshes, especially ' portions of Spartina alterniflora marshes. The most important fly in this ' area is the greenhead, Tabanus nigrovittatus, which often makes up 90 to ' 95% of the adult pest population. The salt marsh tabanid problem extends from Nova Scotia to Florida and along the Gulf Coast to eastern Texas in areas of the salt marsh suitable to the various species or strains. Apparently suitable larval habitat for one strain of T. nigrovittatus exists only along banks of natural and mosquito control ditches (Freeman and Hansens, 1972). Tabanus nigrovittatus females readily bite man and range quite widely from the salt marsh into nearby areas. Prevailing winds often favor infes- I tation of beaches. Greenheads tend to fly within a foot or two of the ground, and thus on bathing beaches exposure of sun bathers results in maximum annoyance. A single bite often causes human response all out of proportion jto the seriousness of the injury. Once the female fly settles down to feed, if one can wait that long, swatting the fly is easy. The wounds caused by tabanids are larger than those of other Diptera and blood often oozes from ithe wound after the fly leaves. If a larger blood vessel is pierced, bleeding may be profuse. Occupants of fishing and pleasure boats are often attacked by greenheads. iThe flies readily move into the boat cabins as the vessels traverse the creeks I and thoroughfares. These flies may then seek a host when the craft has moved ra considerable distance from land. Flies may be attracted by petroleum I products or the exhaust from gas engines. This accounts in part for the large numbers of greenheads which accumulate in power boats. Both sexes have , recently been taken on oil drilling rigs in the Gulf of Mexico many miles I from land and probably by direct flight from land (personal communication, ! John Burger, U. New Hampshire). \ Golf courses in a number of localities have been constructed close to salt I marshes. Vegetation may be cleared between the course and the marsh to give players a view of the wide expanse of the marsh. Observations show that such openings to the marsh greatly facilitate movement of flies to the course and subsequent annoyance of the players. Campgrounds are a rapidly growing industry in coastal areas, especially ' in New Jersey. In Cape May County alone there are some 75 campgrounds. 314 NEW YORK ENTOMOLOGICAL SOCIETY t Most of these are close to salt marshes and greenhead and deerfly annoyance i is very severe. At one location in 1976 at least two children required hospital i treatment for greenhead bites. At a number of locations, campers pulled ^ stakes and left after a day or less of enduring greenheads and deer flies. i This also happened on Grand Manan Island, New Brunswick, Canada (per- n sonal communication, L. L. Pechuman, Cornell Univ.). An important problem with Tabanidae was apparent in South Carolina in , the planning for the Charles Towne Landing Park in 1970 and a suitable, control program was implemented (Adkins, 1974). Greenheads have an impact on crop production in vegetable growing areas of New Jersey along Delaware Bay. In some locations intensive agriculture is surrounded on three sides by salt marsh and greenheads cause great' annoyance to harvesting crews and other farm labor during much of July and August. ' In other areas salt marsh has been reclaimed for housing by digging chan- nels for boat moorings and raising adjacent land above high tide level with the spoil. Such marsh destruction removes greenhead breeding areas, but plenty of nearby marsh produces flies which easily infiltrate the develop- ments. Similarly the salt marsh deer flies, Clirysops fuliginosus and C. atlanticus are important pests of man. C. atlanticus is an avid feeder on man, exists in large numbers for several weeks, and reaches peak numbers when more i people are in resorts and when crop harvesting is in progress. Along the I South Atlantic coast other species are important especially in the George- t town area of South Carolina where C. pudicus and C. niger taylori are pests. * (Adkins, 1974). Human annoyance from Chrysops is restricted to smaller geographical. . areas than Tabanus. Deer flies from the salt marsh move into adjacent i woods and other vegetation but not far into open fields. Typically they are.), not a problem on beaches or in boats but may be more serious than green- i heads on golf courses, in camp grounds, parks, and along the wooded mar- c gins of cultivated fields. Chrysops tend to feed on the head and arms and ^ are much more attracted to a moving host than one standing still. In the i summer of 1976 at the margin of cultivated fields near Cedarville, N.J. as. many as 180 C. atlanticus were taken in 10 figure-8 sweeps of an insect net * over the head. With such a population, 30 actual bites were counted in 90 ; seconds, and observation for longer periods was unbearable. So far reference has been made primarily to a small number of salt marsh species. There is a much larger fauna of Tabanidae in upland areas. Chiy- sops become pests in many local areas. Chrysops vittatus is probably the most generally distributed and hence the most annoying of the freshwater deer flies. Flies of the genus Diachlorus are avid biters and occasionally are i important pests. Only a few species of upland Tabanus cause appreciable i annoyance to humans. I VOLUME LXXXVII, NUMBER 4 315 In the Atlantic salt marsh areas, livestock production is not an important part of agriculture. The biting fly complex of mosquitoes, horse flies, deer flies, stable flies, and others greatly affects livestock thriftiness, weight gains and milk production. Granett and Hansens (1956) showed the cost of sprays on dairy animals was exceeded by increased return from milk production when blood-sucking Diptera were sharply reduced. The amount of blood taken from livestock when Tabanus are present is considerable but depends on the species involved as well as the size of the population. Philip (1931) estimated a blood loss of 300 ml from a constant population of 50 flies feeding over a 10 hour period. Tashiro and Schwardt (1949 and 1953) similarly weighed engorged flies and calculated daily blood loss of 59 to 352 ml per day. Often even more blood is lost from the wounds the flies cause. In coastal areas, especially in the south, animals suffer large blood loss over several months because the fly population is very large and the fly season is prolonged. Soboleva (1956) reported loss of 40 to 200 mg of blood from feeding by a single fly and qualitative changes in blood with feeding of numerous flies, i.e. a decrease in haemoglobin and erythrocytes and an increase in leucocytes. Disease Transmission Active transmission of human disease agents by Tabanidae is not known to occur in the Atlantic coastal states. Possible exceptions are tularemia and the viruses of the California encephalitis group. Laboratory tests have re- cently demonstrated that C. atlanticus is an effective vector of the filarial worm Loci loa of Africa (Orihel and Lowrie, 1975) but introduction of the disease agent to the east coast seems unlikely. An additional factor in livestock production is the known potential of Tabanus as mechanical vectors of bovine anaplasmosis, equine infectious anemia, and vesicular stomatitis (not proved) (Krinsky, 1976). Hog cholera transmission was shown by Tidwell et al. (1972). Research in New Jersey showed the populations of Tabanidae on hogs are small but the number of species which visit hogs is quite large (Weiner and Hansens, 1975). Hog cholera transmission by tabanids is believed to be insignificant but such rare occurrences may be very important in reaching our national goal of complete hog cholera eradication. Wildlife of several species are subject to trypano- somes and filarial worms transmitted by tabanids. Control Efforts Control of Tabanidae has yet to be achieved. A variety of measures now in use give partial control. For preventing biting of man and animals no satisfactory repellents exist though DEBT and ethyl hexanediol give some reduction in Chrysops biting. Catts (1968) advocates use of repellent im- 316 NEW YORK ENTOMOLOGICAL SOCIETY pregnated shirts to reduce biting annoyance. Light colored clothing also helps reduce attacks from greenheads and deer flies (Hansens, 1947). Insecticide applications have met with only limited success. The areas to be treated are large because large expanses of salt marsh are breeding areas for these flies and the adult flies move considerable distances. The vegeta- tion where Chrysops concentrate is often difficult to penetrate with air ap- plication and is inaccessible from the ground. Concentrations of insecticide needed often exceed the amounts which are environmentally safe. Large scale control of T. nigrovittatus by chemicals is unlikely to be acceptable except in emergency situations. In the case of Chrysops, control in localized areas where flies concentrate is feasible. Synthetic pyrethroids with short residual activity may be useful. In 1976 in New Jersey treatment along the edges of fields with resmethrin sprays resulted in relief from deer flies for only a day or two (unpublished, Hansens). In recreational areas such ap- plications might also reduce annoyance to tolerable levels. Use of box and canopy traps for control of T. nigrovittatus has met with considerable success in Maine, Massachusetts, New Jersey and Delaware. Large numbers of box traps have been used in Massachusetts since 1967 to protect beach areas (Spencer, 1971), in New Jersey to reduce fly annoyance on a golf course adjacent to salt marsh, and in Delaware to protect a small community from fly annoyance. In New Jersey the traps were successful enough that one golf course now includes traps as part of their pest man- agement program. In Massachusetts traps are operated by mosquito control agencies and are important in reducing annoyance on beaches, in marinas, on golf courses, and at horse shows. In Delaware box and canopy traps placed in flyways (openings through the barrier of vegetation along the marsh) prevented large numbers of flies from moving into inhabited areas. In all of these efforts traps do not eliminate flies but reduce them to tolerable levels. Canopy traps (personal communication, L. L. Pechuman) are being used in the Hudson Valley, N.Y. around paddocks where valuable stud horses are kept and where EIA is a problem. Horse breeders say they are a great help in reducing populations. Flooding of breeding areas was shown to control Chrysops larvae in Con- necticut (Anderson and Kneen, 1969) but extensive area control would de- stroy too much desirable salt marsh. Research Needs Effective control of the various tabanids on the Atlantic seaboard will be difficult to achieve without much more knowledge of the life history and habits of the flies and development of laboratory rearing technique. In the case of the salt marsh species, oviposition habits of C. fuliginosus are un- known. Oviposition and larval and pupal habits need more study with T. VOLUME LXXXVIl, NUMBER 4 317 nigrovittatus and C. atkmticiis. A recent paper by Magnarelli & Anderson (1977) adds considerable knowledge relative to feeding and gonotrophic ac- tivity. Taxonomic and biological research is needed to clarify the T. nigrovit- tatits complex. The recognition by Freeman and Hansens (1972) of two distinct larvae in two distinct habitats on the salt marsh led to the reasoning that T. nigrovittatus populations may be of two species which are difficult to separate as adults. The second species is probably T. simulans and the two species overlap in Delaware and New Jersey. We also know that many fewer of the so-called T. nigrovittatus are taken in box traps in North and South Carolina than in Delaware and New Jersey. T. nigrovittatus also is not as serious a pest of man in the Carolinas as it is farther north. All of this gives credence to the idea that a species complex exists. In general, adequate regional keys to adults are available for both salt marsh and upland species. Keys to larvae are less complete. Our big gaps in knowledge are in biology and habits and in establishment of a laboratory colony of any species. When these are known, new approaches to control will follow. Development of adequate controls will then make life much more enjoyable in many resort and agricultural areas of the east coast. Literature Cited Adkins, T. R., Jr. 1974. Biology, distribution importance and control of deer flies and horse flies (Diptera: Tabanidae) in water-oriented recreational areas. Water Resources Insti- tute, Clemson Univ. Report No. 42. 172 pp. Anderson, J. R. and F. R. Kneen. 1969. The temporary impoundment of salt marshes for the control of coastal deer flies. Mosq. News. 29:239-42. Catts, E. P. 1968. DEET-impregnated net shirt repels biting flies. Jour. Econ. Entomol. 61:1765. Freeman, J. and E. J. Hansens. 1972. Collecting larvae of the salt marsh greenhead Tahanus nigrovittatus and related species in New Jersey: comparison of methods. Envir. Ento- mol. l(5):653-658. Granett, P. and E. J. Hansens. 1956. The effect of biting fly control on milk production. Jour. Econ. Entomol. 49(4):465^67. Hansens, E. J. 1947. Greenhead flies like dark colors. N.J. Agriculture 29(4):3-4. Krinsky, W. L. 1976. Animal disease agents transmitted by horse flies and deer flies (Diptera: Tabanidae). Jour. Med. Entomol. 13(3):225-275. Magnarelli, L. A. and J. R. Anderson. 1977. Follicular development in saltmarsh Tabanidae (Diptera) and incidence of nectar feeding with relation to gonotropic activity. Ann. Entomol. Soc. Amer. 70:529-533. Orihel, T. C. and R. C. Lourie, Jr. 1975. Loa loa: development to the infective stage in an American deerfly, Chrysops atlanticus. Amer. Jour. Trop. Med. Hyg. 24(4):610-6I5. Philip, C. B. 1931. The Tabanidae (Horseflies) of Minnesota, with special references to their biologies and taxonomy. Univ. Minn. Tech. Bull. 80. 132 pp. Soboleva, R. G. 1956. Tabanids as ectoparasites of domestic animals. Veterinariya 33(4)' 71-77. 318 NEW YORK ENTOMOLOGICAL SOCIETY Spencer, R. M. 1971. A mechanical approach to the abatement of the greenhead fly, Tabanus nigrovittatus. Proc. N.J. Mosq. Exterm. Assoc. 58:71-77. Tashiro, H. and H. H. Schwardt. 1949. Biology of the major species of horse flies of central New York. Jour. Econ. Entomol. 42(2):269-272. and H. H. Schwardt. 1953. Biological studies of horseflies in New York. Jour. Econ. Entomol. 46(5):813-822. Tidwell, M. A., W. D. Dean, G. P. Combs, D. W. Anderson, W. O. Cowart, and R. C. Axtell. 1972. Transmission of hog cholera virus by horseflies (Tabanidae: Diptera). Amer. Jour. Vet. Res. 33(3):615-622. Weiner, T. J. and E. J. Hansens. 1975. Species and numbers of bloodsucking flies feeding on hogs and other animals in southern New Jersey. Jour. N.Y. Entomol. Soc. 83(3): 198- 202. Received for publication August 29, 1979. Department of Entomology and Economic Zoology, Cook College, Rut- gers University, New Brunswick, New Jersey 08903. VOLUME LXXXVIl, NUMBER 4 319 ACKNOWLEDGMENT The Editors wish to express their appreciation to all those who have helped in reviewing the manuscripts submitted during 1979 for publication in the Journal: T. A. Adkins, L. A. Andres, Richard Axtell, Pedro Barbosa, James V. Bell, Robert Byers, E. Paul Catts, Robert F. Denno, J. J. Drea, Henry S. Dybas, Richard J. Dysart, Paul P. Feeny, B. A. Foote, R. H. Foote, Harry G. Fowler, A. Gupta, Lee Herman, F. O. Holmes, Ivan Hub- er, Alexander B. Klots, Timothy J. Kurtti, James Lashomb, D. M. Maddox, David Miller, Neil Morgan, William L. Murphy, L. L. Pechuman, Jonathan Reiskind, R. B. Roberts, Reece I. Sailer, John B. Schmitt, T. J. Spilman, Jane D. Wall, Pedro Wygodzinsky. NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4), 1979, p. 319 BOOK REVIEW Viroids and viroid diseases. T. O. Diener. John Wiley & Sons. 252 pp. 1979. $19.95. What are viroids? The subject, and the name, might be unfamiliar to many, because the study of viroids began in the present decade. Quoting from the preface of this volume: “Viroids are the smallest known agents of infectious disease. They are responsible for a number of destructive diseases of cultivated plants, but may also occur and cause disease in animals. Al- though some of the diseases that viroids cause had been known for decades, these diseases were generally believed to be due to infection by conventional viruses. The unique nature of their causative agents came to light only in 1971 . . . The author of this volume is the foremost authority and the actual discoverer of viroids, who has coined the name “viroid” for these small nucleic acid molecules. The interest in viroids is not limited to plant pathologists, and it seems likely that diseases of insects and of higher ani- mals will be recognized as caused by these agents, once the techniques, clearly described in this volume, will be applied more widely to the study of “uncertain etiology” ailments. Diener has provided a remarkably com- prehensive, unified, and well-written volume on all aspects of viroids. There is no other book available that successfully covers this range of material. The introductory chapter presents the chronology of discoveries that re- sulted in the convincing evidence that viroids consist of nonencapsulated nucleic acid, that the infectious process is not caused by a contaminating virus, that viroids contain a low molecular weight nucleic acid that replicates autonomously without a helper virus, and consists of one molecular species only. At present (1979) seven viroids have been recognized, all causing plant 320 NEW YORK ENTOMOLOGICAL SOCIETY diseases of economic importance. The author discusses the widespread na- ture of established viroids and their serious potential threat to agriculture. He also provides a very stimulating discussion of future research problems and approaches, including the intriguing possibility that certain “slow vi- ruses” causing human diseases might actually be caused by viroids. Al- ^ though the primary audience for this book will consist of plant pathologists, medical and veterinary researchers, virologists, and molecular biologists, many entomologists, particularly insect pathologists will find this subject of considerable interest. The exhaustive bibliography and subject index, to- taling 27 pages, add to the value of this definitive treatise. The book is a joy to read and it will be valuable for many years to come both as a text and j a reference. I Karl Maramorosch, Waksman Institute of Microbiology, Rutgers — The '' State University ' * ' [ NEW YORK ENTOMOLOGICAL SOCIETY i LXXXVIK4), 1979, p. 320 | BOOK REVIEW Twelve Little Housemates. Karl von Frisch. Pergamon Press, Oxford, New York. 1979. 155 pp. $12. Hard cover; $6. Paperback. The 4th (1955) German edition of the popular book by Nobel laureate Karl von Frisch has been translated into English in 1960 and reprinted sev- eral times. Now the enlarged and revised English edition has been published for the Pergamon International Library of Science. The book is written for lay people in a non-technical, humorous style and it is intended for the 1 general public. Although not specifically aimed at entomologists, it never- | theless belongs to the personal library of every entomologist. The 12 inver- | tebrate “housemates” are the housefly, gnats, the flea, the bed bug, lice, J the clothes moth, the cockroach, aphids, ants, silverfish, spiders and ticks. \ The book was aimed primarily at the European reader and the author did 3 not include termites, practically unknown in Central Europe, but of consid- . erable importance in other parts of the world. Here are samples of some \ subtitles: “How to recognize a fly”; “How flies help doctors”; “How one ( can get lice, and how to get rid of them by methods other than those of gas n warfare”; and on cockroaches: “Mouths that bite, lick, suck and sting.” The book is enjoyable and educational not only for children but also for , adults, including scientists. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers — The State University VOLUME LXXXVII, NUMBER 4 321 NEW YORK ENTOMOLOGICAL SOCIETY LXXXV1K4), 1979. pp. 321-322 I BOOK REVIEW I Pern, My Unpromised Land. Eelix Woytkowski. 232 pp. Published for The Smithsonian Institution and NSF by National Center for Scientific, Tech- nical and Economic Information, Warsaw, Poland, 1978. Available from U.S. Department of Commerce, NTIS, Springfield, VA 22161. (PB295870T) $12.00. i[ This is a fascinating account of entomological and botanical explorations I made in Peru by the Polish collector Felix Woytkowski during the years 1929-1964. The book was published first in Polish in 1974 and it was now I translated into English. The highly readable volume opens with a description i of the author’s early memories of his childhood and life in Southeastern j| Poland (now Soviet Western Ukraine) and his educational and family back- ground. As a son of a physician, Woytkowski obtained a thorough training I in Poland and Western Europe, including a period spent in Oxford, England. * In 1929 he was lured by an advertisement to leave a rather well paid position and to go to Peru, where he was promised free passage, tools, and a home- stead farm. Upon arrival, he found none of the promised benefits and was placed in a shanty immigrant camp with hundreds of French, Belgian and j German immigrants who had been waiting for many months to be trans- I ported to the “free farmland.’’ Finally many of the immigrants were able ! to return to their countries of origin, but Woytkowski decided to stick it out and to remain in Peru. He began to collect insects and medicinal plants, and to ship them to taxonomists in the United States. The second part of the book covers the years 1937-1951, a period during which he suffered hunger and pain, lost his only son and was abandoned by his wife, but did not give up the profession that gave him the only satisfaction in life. His unusual courage and perseverance helped him to overcome the incredible obstacles facing a stranger in the remote areas of Peru. Numerous, life-long friendships with U.S. entomologists sustained Woyt- kowski through the most difficult years of deprivation. Many of his J “clients’’ were members of the N.Y. Entomological Society and among his j best friends were Charles P. Alexander, F. Martin Brown, J. Douglas Hood, * Leonora W. Gloyd and Herbert B. Hungerford. His fondness for traveling I and enthusiasm guided him throughout his life. Interested in nature and I people, he struggled to survive in a hostile environment, finding new and j rare species of insects and supported morally and financially by prominent I entomologists. His family life was an unhappy one and his dream to discover I the mysterious “Callanga” of Peru was never fulfilled. The third part of the volume describes exportions from 1951 to 1965 in the valley of the Costripata River, the jungle of the Madre de Dios basin, the search for medicinal plants, and encounters with hostile Indians in the 322 NEW YORK ENTOMOLOGICAL SOCIETY little-explored interior of Peru. Several years ago F. Martin Brown wrote: “Woytkowski had been a hero in the domain of natural science, about whom no books were written so far . . . The present account, in the style of a narrative by Woytkowski, actually has been written by M. Salomea Wie- lopolska, who was able to assemble not only his letters, photographs and notes, but also obtain additional information from entomologists who had close contacts with Woytkowski. Consequently, the account is vivid, often frightening, describing the dangers and deprivations of one of the most de- voted collectors of recent years. The book ends with an epilogue and a chronological list of expeditions. After 36 years in Peru, Woytkowski, by then an old and sick man, found that he could not qualify for a small pension from the Botanic Garden in Lima, because he never became a Peruvian citizen. He decided to return to his native Poland, now a different and strange country to him, where he suffered not only because of the cooler climate and conditions, but, as he wrote, “because I found myself in a situation in which I could be useful and constructive no longer.” He died in Krakow, Poland, in 1966. The English translation of the Polish text is good; numerous black and white photographs taken by the author illustrate the localities and people of Peru’s seldom visited mountains and valleys. There are 6 charts of the areas explored in the years 1930-1964. The book is much more than a travelog — it is the account of a naturalist’s life and hardships and of a devotion to entomology that will appeal to professional and amateur collectors alike. The information will be of special interest to anyone concerned with the culture and the people of Peru. The author’s insight and the description of problems encountered during his explorations might help future collectors. Karl Maramorosch, Waksman Institute of Microbiology Rutgers — The State University NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4), 1979, pp. 322-323 BOOK REVIEW Ecological Methods, With Particular Reference to the Study of Insect Pop- ulations. T. R. E. Southwood. Second ed. Chapman & Hall-Halsted Press-John Wiley and Sons, New York. 524 pp. 1979. $25. The popularity of this book is evidenced by the fact that, after its publi- cation in 1966 and reprinting in 1968, it has been reprinted in 1971, 1975 and 1976, before the present, revised edition first appeared in Cambridge, En- gland in 1978. The reason for the popularity of this scholarly treatise can be found in the remarkable developments in the science of ecology. Ecologists need reliable quantitative data which this study contains. Written in a charming style, it gives a complete account of the theory and applications VOLUME LXXXVll, NUMBER 4 323 of the subject. The chapters deal with the study of animal populations, the li sampling, description of dispersion, absolute population estimates using marking techniques, sampling a unit of habitat, such as air, plants, plant products and vertebrate hosts, soil, litter and fresh-water habitats. Methods of population measurement, estimation of natality, mortality, and dispersal, 1 are described in depth. Entomologists will be particularly interested in the ' discussion of age-grouping of insects, time-specific life tables and predictive population models. An author index and a general index are provided. Need- less to say that this is a most useful book which will reward the individual reader and will also serve as a textbook. The book is technically excellent ’ and its wide scope and lucidity make this new revised editin well worth having wherever ecological problems are being studied. Karl Maramorosch, Waksman Institute of Microhiology, Rutgers — The I State University NEW YORK ENTOMOLOGICAL SOCIETY LXXXVII(4), 1979, p. 323 BOOK REVIEW Arthropod Phytogeny. A. P. Gupta, editor. Van Nostrand Reinhold. 762 pp. 1979. $32.50. This book provides a much needed up-to-date review of the growing body of information on the phylogeny of arthropods. Written by 13 scientists who have made many significant contributions to our knowledge of arthropods, this is a well organized, well illustrated, and clearly written book. The chap- ters deal with the morphology of fossil arthropods, early and late embryonic I stages, abnormal metamorphosis, evolution of antennae and scent detection ■ mechanism, eye structure, functional morphology and evolution of hexa- ' pods, arthropod visceral anatomy, intersegmental tendon system, sperm transfer and ultrastructure, neuroendocrine structures, and hemocytes. Sev- I eral chapters are of special interest because of the well-organized and thor- ' ough treatment of their topics. Gupta’s chapter on types of arthropod he- mocytes in various arthropod groups and his penetrating analysis how they relate to arthropod phylogeny and evolution in general, as well as K. V. ! Clarke’s presentation of the visceral anatomy and arthropod phylogeny are real masterpieces. Other topics are also treated exceedingly well. A taxo- nomic and a subject index are provided. I am convinced that the quality of this treatise will establish this book as a standard reference work and an essential addition to all scientific and biological libraries. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers — The State University 324 NEW YORK ENTOMOLOGICAL SOCIETY NEW YORK ENTOMOLOGICAL SOCIETY LXXXVIK4). 1979, p. 324 BOOK REVIEW Recent Advances in Acarology. I. J. G. Rodriguez, editor. Academic Press. 631 pp. 1979. $35.00. This volume (I) of the proceedings of the V International Congress of Acarology contains 80 reports given at the congress in August 1978. The book is divided into 6 parts: pest management of agricultural mites; biology of spider mites; stored product acarology; physiology, biochemistry and taxicology; ecology and bionomics; and recent advances in soil mite biology. The volume fills a gap and its prompt publication will be most welcomed by all interested in mite and tick research. The book has contributions from an international group of congress participants, including world authorities as well as young researchers. Careful editing resulted in a very readable vol- ume. The book will be of special interest to mite specialists, entomologists, graduate students, teachers, researchers and practically oriented agricultur- al experts. The volume lacks a subject index. Karl Maramorosch, Waksman Institute of Microbiology, Rutgers — The State University HONORARY LIFE AND SUSTAINING MEMBERS Charles P. Alexander Joseph Bequaert Raymond Brush F. S. Chew Lucy Clausen Howard E. Evans S. W. Frost Irving Granek Mark Indenbaum Alexander B. Klots Nellie Payne L. L. Pechuman John D. Plakidas Ruth R. Ralston Richard P. Seifert Edwin Way Teale Hubert J. Thelen George Townes Asher Treat Roman Vishniac INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS VOLUME LXXXVII Generic names begin with capital letters. New genera, species, subspecies and varieties are printed in italics. The following items and articles are not indexed: 1. Table I. Order, family, and identifier of species mentioned in this paper, p. 86. 2. Table 2. Parasite species and numbers reared from 13,351 alfalfa blotch leafminers col- lected in Delaware, southern New Jersey, and southeastern Pennsylvania in 1975-1977. p. 88. 3. Table 3. Parasite species and numbers reared from 33,573 alfalfa blotch leafminer puparia collected in Europe in 1976-1977 by personnel from the USDA European Parasite Laboratory, Paris, France, p. 89. 4. Key to Species of Cicindela of Argentina, pp. 99-101. ' 5. Table I. Growth and behavioral responses of Pieris ncipi macdiinnoiighii to crucifer species growing in a montane Colorado community, p. 130. 6. Table 2. Results of qualitative analysis of crucifer leaves for glucosinolates. p. 131. 7. Table I. Order, family, and determiner of species mentioned in this paper, p. 168. 8. Table 2. Releases of European parasites against the alfalfa blotch leafminer in the USA. pp. 169-171. j! 9. Tablet. Insects associated with cinquefoil, pp. 218-221 . !| 10. Table I. Insects and mites associated W\\\\ StelUiria. pp. 225-231. ' II. Grouping of the species- BrtAer/>//n. p. 257. I 12. Key to species-Bakeriella. pp. 258-260. i 13. Key to Families of American Pentatomoidea. pp. 190-192. I 14. Table I. Order, family, and identity of species mentioned in this paper, p. 300. 15. Table 3. Parasite species and numbers reared from Liriomyza trifoliecmun collected in j Delaware, southern New Jersey, and southeastern Pennsylvania, 1975-1978. p. 302. I Acantholyda maculiventris 209 Acanthomyops plumipilosis 92 1 Acer negundo 284 I Aculus schlechtendali 15 i Acyrthosiphon pisum 275 Adelpha 68 Agistemus fleschneri 19 Agromyza frontella 85, 167, 299 parvicornis 172 Alloxysta megourae 275 ' Amblyseius fallacis 15 Ambrosia artemisiifolia 118 psilostachya 118 trifida 118 Anaea 68 Anoplonyx luteipes 212 ' Anthonomus sp. nr. consimilis 217 Antirrhea 73 Anurogryllus celerinictus 126 I Apanteles ayerzai 239 I Aphidius smithi 275 Arabis drummondii 129 I Arge 212 clavicornis 212 Ascia monuste 253 Athalia proxima 212 Autographa californica 55 Bakeriella 256 azteca 265 brasiliana 264 cristata 266 depressa 265 erythrogaster 260 flavicornis 265 floridana 264 grandis 262 inca 265 inconspicua 264 monlivaga 261 olmeca 265 polita 263 quadriceps 262 quinquepartita 262 reclusa 26 1 rossi 264 rufocaudata 263 Bathyplectes 9 325 326 anurus 9 curculionis 9 stenostigma 10 Blastophaga masoni 212 Brachymeria intermedia 175 Brassica geniculata 237, 243 campestris 237 kaber 246 napus 237 oleracea 252 Brassolis 68 Brochymena 189 Bryobia praetiosa 15 Bunchosia pilosa 289 Caenia dimidiata 287 Caerois 73 Caligo 68 Calliphora 140 Calopteron fasciatum 283 terminale 283 Canopus 42, 193 burmeisteri 42 caesus 42 fabricii 42 germari 48 impressus 42, 193 orbicularis 42 Caracia 189 Cardamine cordifolia 129 hirsuta 237 Cardaria draba 237 Cephalcia marginata 212 provancheri 212 Cephus (Cephas) cinctus 212 Cestrum 237 Ceutorhynchus 217 Chaetosiphon fragaefolii 216 Charpis 275 Cheiranthus 130 Chlaenocoris 193 impressus 193 Chrysis indogotea 213 Chrysocharis punctifacies 90, Chrysops atlanticus 312 fuliginosus 314 niger taylori 314 pudicus 314 vittatus 314 Cicindela 98 argentinica 98 cribrata 98 drakei latifascia 98 halophila 98 hirsutifrons 98 obsoletesignata 98 NEW YORK ENTOMOLOGICAL SOCIETY siccalacicola 98 stamatovi 98 Cimbex americana americana 212 Closterocerus utahensis 89 Clypearia angustior 78 apicipennis 78 weyrauchi 78 Coclebotys nmtuuri 2 Coleomegilla maculata 154 Colias lesbia 237 Conocephalus 39 Conringia orientalis 237 Coptosoma 193 Corimelaena 193 extensa 194 Coronopus didymus 237 Cossonus 286 Coumarouna oleifera 67 Cryphula 160 abort! va 163 apicata 163 australis 163 bennetti 163 dubia 160 nitens 164 parallelogramma 160 Cyrtogaster 88 Dacnusa dryas 90, 167, 302 dorsalis 123 Descurainia appendiculata 237 argentina 237 richardsonii 129 Diachlorus 314 Diglyphus begin! 88 intermedins 88, 173 isaea 172 Dinidor 196 Diplotaxis muralis 237 Diprion pini 212 polytomum 212 Dorylus labiatus 213 Draba aurea 129 167, 302 Drosophila 120 Dysonycha 233 Entomophthora bullata 135 Epyris 256 montivagus 256 Eruca sativa 237 Erysimum asperum 129 Euaresta bella 118 festiva 118 Euarestoides acutangulus 119, 120 Euchloe 239 ausonides 253 VOLUME LXXXVII, NUMBER 4 327 Eumenes dimidiatepennis 213 Euophrys 272 Florida coerulea 126 Fragaria 216 Galgupha 193 nitiduloides 194 Galleria mellonella 176 Graphocephala 217 Guazuma ulmifolia 73 Halys 202 Heterocratis 193 Hippodamia convergens 155 Historis 68 Hoplocampa halcyon 212 Hyaloides vitripennis 15 Hypera postica 9 Janeirona 189 Laccobius minutoides 64 reflexipenis 59 spangleri 59 Lepidium 243 aletes 237 bonariense 237 perfoliatum 246 ruderale 246 spicatum 237 Leptophylemyia coarctata 138 Liriomyza sativae 299 trifoliearum 85, 168, 299 trifolii 85, 299 Loa loa 315 Lotus corniculatus 144 Lycus arizonensis 283 loripes 283 minutus 286 Lygaenematus erichsonii 212 Mangifera 73 Marghita 189 Marpesia 68 Mecidea 203 Meconema thalassinum 38 Medicago 237 hispida 237 sativa 237 Megaris 42, 197 atratula 42 constricta 42 laevicollis 42 puertoricensis 42 rotunda 42 stalii 42 vianai 42 Metaphidippus insignis 270 montanus 270 Metapolybia cingulata 84 Micranisa pteromaloides 212 Miscogaster hortensis 173 maculata 173 Morpho 68 amathonte 73 peleides 72 Musa 73 Mutilla 213 Myrmecia 92 Myzus persicae 233 Nasturtium officinale 237 Neodiprion abietis 212 Netelia kashmirensis 212 Notionotus 304 liparus 304 mexicanus 304 nucleus 304 rosalesi 304 tricarinatus 304 Odontoscelis 193 Onoclea sensibilis 154 Oocyclus 306 Opsiphanes 68 Orchelimum 39 Orius insidiosus 15 Otiorhynchus ovatus 216 Pachyprotasis brunetti 212 Pamphilius luteicornis 212 Panonychus ulmi 15 Pantochlora 189 vivida 200 Papilio polyxenes 148 zelicaon 148 Pemphigus 21 populicaulis 21 populitransversus 21 Pentacomia 98 Perga affinis affinis 212 Phaenicia sericata 135 Phanomeris braconius 172 Phloea 204 Phloeophana 204 Phormia regina 135 Pieris 128 beckeri 240 brassicae 128 (Synchloe) callicide 253 napi 253 328 NEW YORK ENTOMOLOGICAL SOCIETY napi macdunnoughii 128 occidentalis 129 protodice 248 rapae 128. 143, 253 Piezosternum 204 Piriella 73 Pisum sativum 299 Plataspis 193 Populus 21 deltoides 21, 155 Potentilla 216 canadensis 217 intermedia 217 monspeliensis 216 norvegica 216 obscura 216 pilosa 216 recta 216 sulphurea 216 Prepona 68 Pristiphora cincta 212 erichsonii 212 Proctotrypes candatus 214 Protophormia terraenovae 135 Psidium 73 Pyrausta 2 euryphaea 2 Pyropyga minuta 217 Raphanus sativus 237 Rapistrum rugosum 237 Rattana 2 Rhabdepyris 256 Rhagoletis 123 Rubus occidentalis 154 Salix 286 nigra 154 Samanea saman 73 Sarcophaga aldrichi 140 Scelephron intrudens 213 Scolia quadripustulata 213 Sirex cyaneus 212 Sisymbrium altissimum 246 irio 237 officinale 237 orientale 240 Sitona regensteinensis 297 Spartina alterniflora 313 Spodoptera eridania 141 Stellaria cerastioides 224 graminea 223 longipes 224 media 223 Stethorus punctum 15 Stizus vespiformis 213 Strombosoma unipunctatum 193 Svastra 91 Sycophila decatomoides 212 Sycoscapter stabilis 212 Syngenicus 160 Tabanus nigrovittatus 312 simulans 317 Tatochila 236 autodice 236 blanchardii 236 blanchardii ernestae 242 mercedis 236 sterodice arctodice 236 sterodice fueguensis 243 sterodice macrodice 236 sterodice sterodice 236 vanvolxemii 236 xanthodice 236 Taygetis 73 Tephritis dilacerata 120 stigmatica 121 Tetranychus mcdanieli 15 urticae 15 Tetrastichus incertus 9 Thlaspi arvense 129 montanum 129 Thyreocoris 193 coccinelloides 193 punctatus 193 Tomostethus (Eutomostethus) assomensis 212 Tmetothrips subapterus 223 Trachysphyrus 213 Tremex columba 212 Trifolium incarnatum 299 repens 299 Triglyphothrix 92 Tropaeolum majus 237 Tropistethus 160 australis 160 dubius 160 Typhlodromus 15 longipilus 19 pomi 15 pyri 19 Urophora jaceana 119 Uroplata 289 Valentibulla 119 Valtissius 163 Vespa orientalis 213 Vida fava 275 Walkerella temeraria 212 VOLUME LXXXVII, NUMBER 4 329 Xiphydria mellipes 212 Xyela bakeri 212 Xylocopa lemuiscapa 213 Zagrammosoma miiltilineatum 89, 302 Zetzellia mail 15 Zimmerana 189 Zimmeria 200 Zygia longifolia 290 ^ H 4hiiu»/WiA(^ , ,,. I ♦, ^IC, t(a4'.iifi' -« -Hnui'V. •• 'fc.:.- ,ii», .*, ''Ti'j'Hof dBiJt'.-; -"fTA -f u.r^ C .IN' »•.' rt^iUi-n'.f'iUin. I "■ . Vv - ■p ,«',^-,-- -:V a Wr'- •*y*r r 41 i I I m a « I I I l »■ r' ^mx■‘^lMv\aK^ -j(?l ^ i !««• .rtt r - ■ •.>/. l>^ ,4 ~ « >i4ri«> 4t^ • >4M %!>0Vkme* , .' -' •S>-!Mtifl^ iU r* ^ ■ ^ « - <>.«»..»■ »ii<'«»» * <»•*'#♦ 'Vi* «■ i- ' >»»-- W-1<# '_'■*■ •!*t j , :*' i'' . ^ _ M» t-4« (* ■ . '■ . -r-r ■>. * '•,. >f^'' «; iu.*4.^i*. 4JL - "•■■amt- r- .,> ,j’ .(. r . -9S Journal of the New York ENTOMOLOGICAL SOCIETY Devoted to Entomology in General VOLUME LXXXVII Published by the Society New York. N.Y. INDEX OF AUTHORS BARBOSA. P. and E. A. FRONGILLO. JR. Photoperiod and temperature influences on egg number in Brachymeria intermedia (Hymenoptera: Chalcididae). a pupal parasitoid of Lymantria dispar (Lepidoptera: Lymantriidae) BATRA, S. W. T. Reproductive behavior of Enresta Bella and E. (estiva (Diptera: Tephritidae). potential agents for the biological control of adventive North American ragweeds [Ambrosia spp.) in Eurasia BATRA, S. W. T. Insects associated with weeds in the northeastern United States. II. Cinquefoils, PotentiUa norvegica and P. recta (Rosaceae) BATRA, S. W. T. Insects associated with weeds in the northeastern United States. III. Chickweed. Stellaria media, and Stitchwort, 5. graminea (Caryophyllaceae) BELL, PAUL D. Acoustic attraction of herons by crickets BENTON, ALLEN H. and ANDREW J. CRUMP. Observations on aggregation and overwintering in the coccinellid beetle Coleomegilla macidata (De Geer) CHEW, F. S. Community ecology and Pieris-Crncifer coevolution CUTLER, BRUCE. Variation in the embolus of Metaphidippus insignis (Banks) (Araneae: Salticidae) DOWELL. ROBERT V. Effect of low host density on oviposition by larval parasitoids of the alfalfa weevil EVANS, HOWARD E. A reconsideration of the genus Bakeriella (Hymenoptera: Bethylidae) FAITH, DANIEL P. Strategies of gall formation in Pemphigus aphids HANSENS, ELTON J. Review: Tabanidae of the East Coast as an economic problem HENDRICKSON, R. M., JR. Field studies and parasites of Liriomyza trifolieariim (Diptera: Agromyzidae) in northeastern USA HENDRICKSON. R. M.. JR. and S. E. BARTH. Effectiveness of native parasites against Agramyza frontella (Rondani) (Diptera: Agromyzidae), an introduced pest of alfalfa HENDRICKSON. R. M., JR. and S. E. BARTH. Introduced parasites of Agramyza frontella (Rondani) in the USA JEANNE. ROBERT L. Nest of the wasp Clypearia weyranehi (Hymenoptera: Ves- pidae) 17.5 118 216 223 126 L54 128 270 9 2.56 21 312 299 85 167 78 I 186 KNIZESKI, HENRY M.. JR. Dr. Charles Paul Alexander KRAMER. JOHN PAUL. Interactions between blow flies (Calliphoridae) and Ento- nioplitliora hidlata (Phycomycetes: Entomophthorales) LANHAM, U. N. Possible phylogenetic significance of complex hairs in bees and ants MALCOLM, STANLEY E. Two new species of Lacvohius from eastern North America (Coleoptera: Hydrophilidae) MATEJKO, IRENE and DANIEL J. SULLIVAN, S.J. Bionomics and behavior of AUaxystci mepourae. an aphid hyperparasitoid (Hymenoptera: Alloxystidae) McCABE, TIM L. and LINNEA M. JOHNSON. The larva of Ccilopiemn tenniniile (Say) with additional notes on adult behavior (Coleoptera: Lycidae) McDonald, F. j. D. a new species of Megaris and the status of the Megarididae McAtee and Malloch and Canopidae Amyot and Serville (Hemiptera: Pentatomoidea) McIntosh, Arthur h., karl maramorosch and russell riscoe. auw- graplia ccilifoniica nuclear polyhedrosis virus (NPV) in a vertebrate cell line: local- ization by electron microscopy PERKINS, PHILIP D. Three new Middle American species of aquatic beetles in the genufi Notionoiiis Spangler (Hydrophilidae: Hydrobiinae) ROLSTON, L. H. and F. J. D. MCDONALD. Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera) ROSE, H. S. and H. R. PAJNl. The taxonomic position of three North-West Indian species commonly referred to the genus Pyniiista Schrank (Lepidoptera: Pyrali- dae) SAINL MALKIAT S., SURJIT S. DHILLON and TARLOK SINGH. Positional vari- ation and modification relating to the protergum in Hymenoptera SCRIBER. J. MARK. Post-ingestive utilization of plant biomass and nitrogen by Lepidoptera: legume feeding by the southern armyworm SHAPIRO, ARTHLIR M. The life histories of the autodice and sterodice species- groups of Tatochila (Lepidoptera: Pieridae) SLATER, JAMES A. On the identity of two species of Rhyparochrominae from Ar- gentina (Hemiptera: Lygaeidae) SMITH, BURKE, JR. European katydid Meconenui thalassinum (DeGeer) recorded from new location on Long Island. New York (Orthoptera: Tettigonidae) SUMLIN, WILLIAM D., HI. A brief review of the genus Cicindela of Argentina (Coleoptera: Cicindelidae) WEIRES. R. W. and G. L. SMITH. Mite predators in Eastern New York commer- cial apple orchards YOUNG. ALLEN M. The evolution of eyespots in tropical butterflies in response to feeding on rotting fruit: an hypothesis YOUNG. ALLEN M. Notes on a population outbreak of the beetle Uroplata sp. (Coleoptera: Chrysomelidae) on the tree Bunchosia pilosa (Malpighiaceae) in Costa Rica 135 L)R 91 59 275 fin 283 42 55 304 189 208 141 236 160 38 98 15 66 289 Book Reviews MARAMOROSCH. KARL, The Year of the Ant. George Ordish 95 SIMOVA-T(^SIC. D. Tabanini of Thailand above the Isthmus of Kra (Diptera: Ta- banidae). John J. S. Burton 96 MARAMOROSCH, KARL. Introduction to Plant Viruses. C. L. Mandahar 181 MARAMOROSCH, KARL. Biochemistry of Insects. Morris Rockstein. ed. 182 MARAMOROSCH. KARL. The Life that Lives on Man. Michael Andrews 182 MARAMOROSCH, KARL. Introduction to Insect Biology and Diversity. Howell V. Daly, John T. Doyen. Paul R. Ehrlich 183 I MARAMOROSCH, KARL. Diseases, Pests and Weeds in Tropical Crops. Jurgen Kranz, Heinz Schmutterer and Werner Koch, eds. 267 I MARAMOROSCH, KARL. American Spiders. Willis J. Gertsch 268 i MARAMOROSCH, KARL. Viroids and Viroid Diseases. T. O. Diener 319 I MARAMOROSCH, KARL. Twelve Little Housemates. Karl von Frisch 320 I MARAMOROSCH, KARL. Peru, My Unpromised Land. Felix Woytkowski 321 I MARAMOROSCH, KARL. Ecological Methods, with Particular Reference to the Study of Insect Populations. T. R. E. Southwood 322 MARAMOROSCH, KARL. Arthropod Phylogeny. A. P. Gupta, ed. 323 li MARAMOROSCH, KARL. Recent Advances in Acarology. 1. J. G. Rodriguez, ed. 324 iii • r 4. .1 E .r » «»> 5^ if > » ■ fs o *3 ?» I O « Q. CXI I 00 00 Ln o o CD r+ << Journal of the New York Entomological Society V. 87-88 1979-80 59.57.06(747) Journal of the Ne