The
PAN-PACIFIC
ENTOMOLOGIST
Volume 77 January 2001 Number 1
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603)
The Pan-Pacific Entomologist
EDITORIAL BOARD
J. Y. Honda, Editor R. M. Bohart
Z. Nisani, Assistant to the Editor J. T. Doyen
R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta F. Parinas, Treasurer Warren E. Savary
Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary; and financial communications should be addressed to: Julieta F. Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Application for membership in the Society and changes of address should be addressed to: William Hamersky, Membership Committee chair, Pacific Coast Entomological Society, Dept. of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. Robert V. Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture, 1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con- tributors, and volume 73(4): 248—255, October 1997, for more detailed informa- tion. Information on format for taxonomic manuscripts can be found in volume 69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.
The annual dues, paid in advance, are $25.00 for regular members of the So- ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for institutional subscriptions or sponsoring members. Members of the Society receive The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes are available; see 72(4): 247 for current prices. Make checks payable to the Pacific Coast Entomological Society.
Pacific Coast Entomological Society
OFFICERS FOR 2000
David T. Wyatt, President Vincent F. Lee, Managing Secretary Julieta F. Parinas, Treasurer Joshua J. Fairbanks, Recording Secretary
THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and addition- al mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
This issue mailed 2 January 2001
The Pan-Pacific Entomologist (SSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.
The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PUBLICATIONS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY
PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.
Vol. 1 (16 numbers, 179 pages) and vol. 2 (9 numbers, 131 pages). 1901- 1930. Price $5.00 per volume.
THE PAN-PACIFIC ENTOMOLOGIST.
Vol. 1 (1924) to vol. 51 (1975), price $10.00 per volume of 4 numbers, or $2.50 per single issue. Vol. 52 (1976) to vol. 57 (1981), price $15.00 per volume or $3.75 per single issue, except for vol. 57, no. 1, $10.00. Vol. 58 (1982) to vol. 66 (1990), $20.00 per volume or $5.00 per single issue. Vol. 67 (1991) to vol. 69 (1993), $30.00 per volume or $7.50 per single issue. Vol. 70 (1994) and subsequent issues, $40.00 per volume or $10.00 per single issue. Since some issues are out of print, please ask about availability before purchasing.
MEMOIRS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.
Volume 1. The Sucking Lice by G. F. Ferris. 320 pages. Published October 1951. Price $10.00 (plus $1.00 postage and handling).
Volume 2. A Revision of the Spider Mite Family Tetranychidae by A. Earl Pritchard and Edward W. Baker. 472 pages. Published July 1955. OUT-OF-PRINT.
Volume 3. Revisionary Studies in the Nearctic Decticinae by David C. Rentz and James D. Birchim. 173 pages. Published July 1968. Price $4.00* (plus $0.75 postage and handling).
Volume 4. Autobiography of an Entomologist by Robert L. Usinger. 343 pages. Published August 1972. SPECIAL PRICE $5.00 (plus $1.00 tax, postage, and handling for California orders; $0.70 postage and handling for non-California U.S. orders, or $1.70 for foreign orders). No members discount at this special price.
Volume 5. Revision of the Millipede Family Andrognathidae in the Nearctic Region by Michael R. Gardner. 61 pages. Published January 21, 1975. Price $3.00* (plus $0.75 postage and handling).
*For California orders only, add appropriate sales tax currently in effect in your county. If requested, members of the Society can receive a 20% discount on the price of the memoirs, except for vol. 4.
Send orders to:
Pacific Coast Entomological Society c/o California Academy of Sciences Golden Gate Park
San Francisco, California 94118-4599 U.S.A.
PAN-PACIFIC ENTOMOLOGIST 77(1): 2-8, (2001)
THE GENUS NOTOBITOPSIS BLOTE WITH THE DESCRIPTION OF TWO NEW SPECIES (HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE: MICTINI)
HARRY BRAILOVSKY AND ERNESTO BARRERA
Departamento de Zoologia, Instituto de Biologia Universidad Nacional Aut6noma de México Apdo Postal 70153, México 04510 D.F, México
Abstract—Two new species of Notobitopsis, N. novoguinensis and N. sandaracinus, collected in New Guinea are described and illustrated. New records for N. limbativentris and N. militaris are given and a key to the five known species is provided.
Key Words.—Insecta, Hemiptera, Heteroptera, Coreidae, Mictini, Notobitopsis, new species, New Guinea.
Stal (1863) and Distant (1911a, b) described the three previously known species of Notobitopsis and included them in the genus Mictis: M. aruana Distant, M. limbativentris Stal and M. militaris Distant. Bléte (1938) placed limbativentris and militaris in a new subgenus of Mictis (Notobitopsis). O’Shea and Schaefer (1980) revised the tribe Mictini in the Asian and Australian region, recognizing nineteen genera, and including Notobitopsis which they elevated to generic status and redescribed.
Within the tribe Mictini, the genus Notobitopsis is characterized by the lack of a tubercle at the apex of scutellum; the cylindrical fore tibia; pronotum not steeply declivent with humeral angles rounded, not angulate; middle third of posterior margins of abdominal sternites III and IV not produced posteriorly; abdominal stemite III with large tubercle on each side; and posterior margin of abdominal stemite VI without a median tubercle.
This contribution adds two noteworthy records, two new species and a key to the five known species of Notobitopsis.
NOTOBITOPSIS ARUANA (Distant) Mictis aruana Distant 1911b: 389-390.
This is the largest known species of Notobitopsis (length over 34 mm) and readily distinguishable by the following combination of characters: antennal segment IV and clavus black, and apical margin of corium yellow to orange yellow.
Distribution.—This species was described from the Aru Islands and is only known from the type specimen.
Material Examined—1 male; data: Aru Islands. Type deposited in The Natural History Museum, London.
NOTOBITOPSIS LIMBATIVENTRIS (Stal) (Figs. 5 and 6)
Mictis limbativentris Stal 1863: 603.
This species is easily identified within the genus by having the clavus and apical margin of corium black, and the paramere tip stout and almost straight (Figs. 5
2001 BRAILOVSKY & BARRERA: THE GENUS NOTOBITOPSIS 3
and 6). In N. aruana and N. militaris the corial apical margin is yellow to orange yellow, and the clavus orange yellow or black with reddish reflections. Notobi- topsis limbativentris is much smaller than the two other species with a total body length of less than 30.00 mm.
Distribution.—This species was originally described from Dory, New Guinea (Stal), and subsequently reported from Salawatti, New Guinea (Blote 1938).
Material Examined.—1 female; data: NE New Guinea. Adelbert Mts., Wanuma, 800-1000 m, 23 October 1958, J. L. Gressitt. Deposited in Bernice P. Bishop Museum, Honolulu, Hawaii. 12 males, 11 females; data: Indonesia Irian Jaya. Baliem Valley nr., Jiwika, 5800-6000’, 24 October 1990, P. J. Clausen. Deposited in University of Minnesota, Insect Collection.
NOTOBITOPSIS MILITARIS (Distant) (Figs. 7 and 8)
Mictis militaris Distant 191la: 595-596.
This species is similar to N. aruana (Distant) but slightly smaller, less than 33.00 mm, with clavus orange yellow to dull sanguineous. Notobitopsis aruana is longer than 34.00 mm, with clavus black.
Distribution.—This species previously was known from Wataikwa River and Hollandia in New Guinea (Distant 1911la, Blote 1938).
Material Examined.—1 female; data: New Guinea. Saidor Apo 321, March—May 1944, O. H. Gra- ham. Deposited in Bernice P. Bishop Museum, Honolulu, Hawaii. | male, 2 females; datas NE New Guinea. Kar Kar Is., Kurum Bagiai Crater trail, 0-100 m, August 1968, N. L. H. Krauss. Deposited Instituto de Biologia, UNAM, México.
NOTOBITOPSIS NOVOGUINENSIS BRAILOVSKY AND BARRERA, NEW SPECIES (Figs. 1, 3 and 4, 11)
Types.—Holotype: male; data: Papua New Guinea. Eastern Highlands, Uba- gubi, 20 mi S Goroka, 6°22’ S, 145°11’ E, June 1986, G. Dodge. Deposited in Cornell University, Insect Collection, Ithaca, New York. Paratype: 1 male: same data as holotype. Deposited in the Coleccién Entomolégica del Instituto de Biol- ogia, UNAM, México.
Description.—Male (holotype). Dorsal coloration: Head, antennal segments I-IV, pronotum, scu- tellum, and abdominal segments black; head below each postocular tubercle with small yellow dis- coidal spot; clavus shiny reddish dark; corium shiny reddish dark, with costal margin black; hemelytral membrane shining metallic blue; connexival segments IJI—VI and anterior half of VII shiny orange; posterior half of connexival segment VII black. Ventral coloration. Ground color including rostral segments and legs black; propleura, mesopleura, and metapleura with large carmine-red spot, one on each segment; anterior and posterior lobe of metathoracic peritreme yellow; pleural margin of abdom- inal sterna III—-VII orange, with spiracles black. Structure. Head: Rostrum reaching posterior margin of mesosternum. Pronotum: Lateral margins finely dentate; humeral angles slightly produced. Legs: Fore and middle femora with two distinct spines near apex; hind femur longer, incrassate, slightly curved, attenuated at base, densely granulate, without apical spine; fore and middle tibiae sulcate, cylindrical; hind tibia sulcate, moderately dilated ventrally, with strong spine at inner face before apex. Abdomen: Abdominal sternite III with small tubercle on each side. Genitalia (Figs. 1, 3 and 4).— Genital capsule (Fig. 1): Posteroventral edge thickest, scarcely cordiform; body without tubercles. Parameres (Figs. 3 and 4): Stout, with elongate and almost straight tip. Measurements. Total body length: 26.90 mm. Head length: 2.04 mm; width across eyes: 2.66 mm; interocular space: 1.36 mm; interocellar space: 0.68 mm. Antennal segments length: I, 5.70 mm, II, 5.16 mm, III, 4.18 mm, IV, 4.18 mm. Pronotal length: 5.47 mm; width across frontal angles: 2.43 mm; width across humeral angles: 7.52 mm. Scutellar length: 3.11 mm; width: 2.96 mm.
4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Figures 1 and 2. Male genital capsule of Notobitopsis spp. Figure 1. N. novoguinensis Brailovsky and Barrera, NEW SPECIES. Figure 2. N. sandaracinus Brailovsky and Barrera, NEW SPECIES. Figures 3-10. Parameres of Notobitopsis spp. Figures 3 and 4. N. novoguinensis Brailovsky and Barrera, NEW SPECIES. Figures 5 and 6. N. limbativentris (Stal). Figures 7 and 8. N. militaris (Distant). Figures 9 and 10. N. sandaracinus Brailovsky and Barrera, NEW SPECIES.
Female.—Unknown.
Discussion—This handsome species is readily separable by having the clavus and corium shiny reddish dark, and the hemelytral membrane metallic blue. The other known species have black or orange yellow clavus, the corium black, and the hemelytral membrane shiny dark olivaceous or black. The parameres are dis- tinct (Figs. 3-10).
Distribution.—Only known from New Guinea.
Etymology.—Named for its occurrence in New Guinea.
2001 BRAILOVSKY & BARRERA: THE GENUS NOTOBITOPSIS 5
Figure 11. Dorsal view of Notobitopsis novoguinensis Brailovsky and Barrera, NEW SPECIES.
6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
NOTOBITOPSIS SANDARACINUS BRAILOVSKY AND BARRERA, NEW SPECIES (Figs. 2, 9 and 10, 12)
Types.—Holotype: male; data: New Guinea. Finschhafen, 16—21 April 1944, E. S. Ross. Deposited in California Academy of Sciences, San Francisco, Cali- fomia (CAS). Paratypes: 9 males, 9 females; data: same locality and date as holotype. Deposited in California Academy of Sciences, San Francisco California and Coleccién Entomolégica del Instituto de Biologia, UNAM, México.
Description.—Male (holotype). Dorsal coloration: Head black; antennal segments I-III black red- dish brown, IV shiny orange with basal joint black; pronotum chestnut orange with anterior margin, lateral margins, and humeral angles black, and lateral border orange; scutellum chestnut orange with apex orange; clavus chestnut orange; corium chestnut orange with costal margin black, and costal border yellow; hemelytral membrane dark shining olivaceous, with blue, green and pink metalic iri- discence; connexival segments dark chestnut orange with upper margin densely granulate with black; dorsal abdominal segments dark chestnut orange. Ventral coloration. Head and bucculae shiny orange with the space close to eyes black; rostral segments I-IV black with chestnut orange reflections; prosternum, mesosternum, and metasternum chestnut orange; propleura, mesopleura, and metapleura with large and wide carmine-red spots, one on each segment; upper margin of propleura and meso- pleura black, and upper margin of metapleura reddish orange; acetabulae reddish brown to shiny orange; anterior and posterior lobe of metathoracic peritreme shiny orange; coxae with inner face chestnut orange and outer face reddish brown; trochanters reddish brown; femora chestnut orange with apical joint black; fore and middle tibiae dark reddish brown, with chestnut orange reflections; hind tibia chestnut orange with subapical spine and apical third reddish brown; tarsi reddish brown; ab- dominal sterna dark chestnut orange with spiracles, apex of the tubercle of abdominal sternite III, and posterior margin of abdominal sternite VII black; pleural margin of abdominal sterna II-VI dirty yellow; genital capsule black. Structure. Head: Rostrum reaching posterior margin of mesosternum. Pronotum: Lateral margins densely crenulate; humeral angles obtuse, not exposed. Legs: Similar to N. novoguinensis. Abdomen: Abdominal sternite II with large and prominent tubercle on each side; upper margin of connexival segments densely granulate. Genitalia (Figs. 2, 9 and 10).—Genital cap- sule (Fig. 2): Posteroventral edge thickest, simple, with small tubercles close to middle third and sinuate. Parameres (Figs. 9 and 10): Stout with broad curved tip. Measurements. Total body length: 30.80 mm. Head length: 2.26 mm; width across eyes: 2.96 mm; interocular space: 1.52 mm; intero- cellar space: 0.82 mm. Antennal segments length: I, 6.30 mm, II, 5.92 mm, III, 4.71 mm, IV, 4.86 mm. Pronotal length: 6.08 mm; width across frontal angles: 2.66 mm; width across humeral angles: 7.82 mm. Scutellar length: 3.42 mm; width: 3.02 mm.
Female——Coloration: Similar to male. Connexival segments VIII and IX dark chestnut orange; dorsal abdominal segments VIII and IX, and genital plates black. Structure. Legs: Hind femur slightly incrassate, granulate with distinct spine near apex; hind tibia sulcate, moderately dilated, unarmed. Measurements. Total body length: 31.30 mm. Head length: 2.35 mm; width across eyes: 2.99 mm; interocular space: 1.53 mm; interocellar space: 0.91 mm. Antennal segments length: I, 6.46 mm, II, 5.39 mm, III, 4.25 mm, IV, 4.94 mm. Pronotal length: 6.46 mm; width across frontal angles: 2.87 mm; width across humeral angles: 8.05 mm. Scutellar length: 3.64 mm; width: 3.19 mm.
Discussion.—Antennal segment IV of N. sandaracinus, is shiny orange with basal joint black, and general color of body chestnut orange. In other species, antennal segment IV is always black, and the general color of the body black or black with yellow stripes. Additionally, the shape of the parameres of N. sandar- acinus, N. limbativentris (Stal), and N. militaris (Distant) are distinct (Figs. 5— 10).
Distribution.—Only known from New Guinea.
Etymology.—From the greek sandaracinos, orange colored.
: THE GENUS NOTOBITOPSIS
BRAILOVSKY & BARRERA
2001
Dorsal view of Notobitopsis sandaracinus Brailovsky and Barrera, NEW SPECIES.
Figure 12.
8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
KEY TO SPECIES OF THE GENUS NOTOBITOPSIS
1. Antennal segment IV shiny orange; scutellum chestnut orange; fore and middle rémora chestnut orange... 6.5 foe. gh eats te teens ee ages oes se ee N. sandaracinus Brailovsky and Barrera, NEW SPECIES 1’. Antennal segment IV black; scutellum black; fore and middle femora IDIACKS 2 SE Shee eel ieee cas teien ce tet WELT ced ee Ss vit ee ee Ge ee 2 2. Clavus shiny reddish dark; corium shiny reddish dark with costal margin black; hemelytral membrane shining metallic blue .................... Pt, Oa ee Sa N. novoguinensis Brailovsky and Barrera, NEW SPECIES 2’. Clavus black or orange yellow; corium almost entirely black; hemelytral membrane dark shining olivaceous or black ...................005. 3 3. Apical margin of corium black; body length less than 30.00 mm. ........ FU ever OP exes PP eee ER, cs cs SAMS SS OPIS SS rees SUBS EO N. limbativentris (Stal) 3’. Apical margin of corium yellow to orange yellow; body length longer PietriS 0300 gritty Me hotles EN ER la cea Mri hela ee cig 4
epee aA SOS ACT Pg ee Er UN Cte ne ee nts See ee Ce Pe N. militaris (Distant) 4’. Clavus black with reddish reflections; body length longer than 34.00 mm. As os AEN, A PR ean ORLA Re Hm ins ok oe LRN Pee N. aruana (Distant)
ACKNOWLEDGMENT
We thank the following colleagues and institutions for loaning us specimens and giving assistance relevant to this study: Gordon Nishida (Bemice P. Bishop Museum, Honolulu, Hawaii); Mick Webb (The Natural History Museum, London, U.K.); Norman D. Penny and Vincent Lee (Califomia Academy of Sciences, San Francisco, Califomia); Richard C. Hoebeke (Cornell University, Insect Collection, Ithaca, New York); P. J. Clausen (University of Minnesota, Insect Collection). Special thanks to Albino Luna for dorsal view illustrations.
LITERATURE CITED
Blote, H. C. 1938. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie Part IV. Coreinae, Third Part. Zool. Meded., 20: 275-508.
Distant, W. L. 1911a. An Enumeration of the Rhynchota collected during the Expedition of the British Omnithologists Union to Central Dutch New Guinea. Trans. Ent. Soc. London, 3: 591-604.
Distant, W. L. 1911b. Rhynchota from the Aru Islands. Ann. Mag. Nat. Hist. (Series 8), 8: 389-390.
O’Shea, R. & C. W. Schaefer. 1980. A generic revision of the Asian and Australian Mictini (Heter- optera: Coreidae). Oriental Insects, 14: 221-251.
Stal, C. 1863. Hemipterorum exoticorum generum et specierum nonnullarum novarum descriptiones. Trans. Ent. Soc. London, 1: 571—603.
Received 15 Nov 1999; Accepted May 31, 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 9-18, (2001)
THREE NEW SPECIES OF HELICOPSYCHE FROM VIETNAM (TRICHOPTERA: HELICOPSY CHIDAE)
PATRICIA W. SCHEFTER! AND KJELL ARNE JOHANSON? ‘Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario, Canada M5S 2C6 Swedish Museum of Natural History, PO. Box 50007, S-104 05 Stockholm, Sweden
Abstract.—Three new species of Helicopsyche are described from Vietnam: Helicopsyche khe- moiensis NEW SPECIES, Helicopsyche azunensis NEW SPECIES and Helicopsyche dackles- tensis NEW SPECIES. Helicopsyche khemoiensis is most closely related to H. coreana Mey, the two species comprising the subgenus Galeopsyche. Helico psyche khemoiensis is distinguished from H. coreana by the shape of the gonocoxite and armature of tergite X. Immature stages of H. khemoiensis are described. Helicopsyche azunensis and H. dacklestensis, subgenus Helicop- syche, are most closely related to Helicopsyche chrysothoe (Schmid), H. xenothoe (Schmid) and other species characterized by four-segmented maxillary palps, absence of mesoscutal warts and distally hooked gonocoxal secondary branch. Helicopsyche azunensis and H. dacklestensis are
distinguished from other species in that group by the presence on the phallus of large endothecal sclerites.
Key Words.—Insecta, Trichoptera, Helicopsychidae, Helicopsyche, Vietnam, NEW SPECIES.
Biologists from the Centre for Biodiversity and Conservation Biology at the Royal Ontario Museum, Toronto, are collecting insects, reptiles and amphibians, and small mammals from the forests of Vietnam in an effort to determine the diversity and uniqueness of their faunas. Deforestation is proceeding rapidly and environmentalists are racing to find areas of high endemicity which should be protected from further degradation. Collections brought back to the Museum for identification have yielded a number of species unknown to science. Terrestrial and aquatic insect collections, including Trichoptera, are particularly rich in un- described species. This paper describes three new species of Helicopsyche (Tri- choptera) from streams in Vietnam.
Originally described as a pulmonate snail (Say 1821), the larva of the caddisfly Helicopsyche builds a dextrally coiled sand grain case within which it grazes periphyton from rock surfaces. The case which bears an uncanny resemblance to that of a snail provides protection from predators and its shape enhances the larva’s ability to maintain position in the current of a freshwater lotic habitat; its sturdy construction resists crushing, and permits the larva to burrow deeply into the substrate (Williams et al. 1983).
The family Helicopsychidae occurs in all faunal regions and is most diverse in the Oriental and Neotropical regions. Of the nearly 180 species which have been described approximately 150 are in the genus Helicopsyche. Only two species of Helicopsyche, H. coreana Mey from North Korea and Helicopsyche azwudschgal Malicky from northern Vietnam were previously known as adults from the East Asian Subregion (sensu Banarescu 1992) (central and northern Vietnam, China, Korea, the Japanese archipelago, Sakhalin Island, Taiwan, Hainan); Helicopsyche yamadai Iwata from Japan was described from larval material and undescribed larvae of the family were also reported from Hong Kong (Dudgeon 1988). The
10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
three species described in this paper, Helicopsyche khemoiensis Schefter & Jo- hanson NEW SPECIES, Helicopsyche azunensis Schefter & Johanson NEW SPE- CIES and Helicopsyche dacklestensis Schefter & Johanson NEW SPECIES sub- stantially increase the known helicopsychid fauna of the subregion. The neigh- boring South Asian Subregion (sensu Banarescu 1992) (southwestern Vietnam, Thailand, Malaysia, Laos, Myanmar, the Indonesian archipelago, southern India and Sri Lanka) harbours approximately 50 species of Helicopsyche, sensu lato (Malicky & Chantaramongkol 1993; Schmid 1993; Malicky 1994, 1995; Johanson 1998).
Morphological terms used in this paper are adopted from Johanson (1998). Types are deposited in the Royal Ontario Museum, Toronto, Canada.
HELICOPSYCHE KHEMOIENSIS SCHEFTER AND JOHANSON, NEW SPECIES (Figs. 1-26)
Types.—Holotype, male; VIETNAM. NGHE AN: West of Con Cuong, Khe Moi Forestry Camp, 28 Oct 1994, D. C. Currie, ROM946112, coal oil lantern, tropical forest, Khe Moi River margin. Paratype: female, same data as holotype. Paratypes, immatures: VIETNAM. NGHE AN: ca. 25 km SW of Con Cuong, Khe Moi River Forestry Camp, 6 June 1995, Brad Hubley, ROM956170, small stream 100 m upriver of camp, tropical forest edge, 18°56'N, 104°49’E (2 pupae, 2 larvae).
Description —Male. Head (Fig. 1): postantennal warts small, ovoid; cephalic warts large, ovoid; eyes large, diameter equal to length of head (Fig. 1); antenna with scape swollen, four and one half times length of pedicel; pedicel and first flagellomere subequal, subsequent four segments graduated in length, remainder equal to fifth; tentorium (Fig. 2) with anterior arms subparallel, distally slightly expanded, short posterior arms and broad tentorial bridge; maxillary palps with segments 1 and 2 subequal, apical segment with long setae dorsally. Pronotum with one pair of setal warts. Legs: spur formula 1,2,4, apical tibial spur of anterior leg slightly longer than one half length of first tarsal segment (Fig. 3). Wings (Fig. 4): fore wing length 4.6 mm, forks 1,2 and 3 present; fork 1 originates on distal one third of Dc; R, subequal to Dc; A, and A, fused basally, A,,, fused with Cu, without reaching posterior wing margin; hind wing 3.7 mm, with 16 hamuli. Abdomen with small, pointed ViIth sternal process. Genitalia: segment IX, lateral view (Fig. 5), anteriorly nearly oval, with weakly developed apodemes; dorsally reduced to a sclerous transverse bridge (Fig. 7); in ventral view broad and deep (Fig. 9); gonocoxite divided into large primary and short secondary branch; in lateral view (Fig. 5) primary branch slender, arcuate; dorsal and ventral margins subparallel; secondary branches fused ventromesally forming shelf-like star shaped plate (Fig. 9); superior appendage inserted mesally, directed posteriad, clavate in lateral view (Fig. 5); Xth tergum depressed in lateral aspect, apices hooked dorsally; in dorsal view (Fig. 7) apices divergent, separated by V-shaped notch approximately one fourth length of segment, with short lobate lightly sclerous lateral processes inserted near base of tergum; macrosetae absent. Phallus slightly arcuate (Fig. 6), apically expanded, with bifurcate sclerous process inserted dorsally in endotheca (Figs. 6 and 8).
Female.—Head: eyes smaller than in male; cephalic warts small, round; antenna with scape not swollen, four and one half times length of pedicel, pecidel shorter than first and subsequent flagellom- eres. Spur formula 1,2,4; apical tibial spur of foreleg one third length of first tarsal segment. Wings (Fig. 10): fore wing length 5.6 mm; venation as in male, except Dc longer than M,; A,,, reaches posterior wing margin at some distance before Cu,; hind wing length 3.7 mm, with 19 hamuli; venation as in male, apex more acuminate than in male. Abdomen with short VIth sternal process, rounded in ventral view (Figs. 11 and 15). Sternite VIII with row of stiff, clear setae on apical margin. Genitalia: in lateral view (Fig. 12) segment IX with small group of ventrolateral setae, deep incision between the [Xth and Xth segments includes the posterior vaginal opening; Xth segment with setae arranged along dorsal margin of dorsal branch; dorsal branches broad and slightly curved ventrad; in ventral view (Fig. 14) ventral branch of segment X very small, external part of gonopods IX narrow, arcuate;
2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 11
superior
appendage
Figures 1-9. Helicopsyche khemoiensis (male). Figure 1. Head, dorsal view.
Figure 2. Tentorium, dorsal view.
Figure 3. Fore leg.
Figure 4. Wings.
Figure 5. Genitalia, lateral view.
Figure 6. Phallus, lateral view.
Figure 7. Genitalia, dorsal view.
Figure 8. Phallus, dorsal view.
Figure 9. Genitalia, ventral view.
spermathecal sclerite subquadrate; spermathecal duct with anterad-oriented microtrichia at midsection (Figs. 12 and 13).
Larva—Head (Figs. 16-18): light tan with six darker spots on lateral surface, three or four minute setae posterolaterally (Figs. 16 and 17, setae 19,20,21); dorsally with dark spots scattered on surface, and postocular spot with several long setae (setae 13-16); frontoclypeus with characteristically shaped dark figure on posterior surface; ventrally unicolorous with pair of dark spots near postoccipital margin (Fig. 18). Thorax (Fig. 19): pronotum with anterior margin bearing large sharp spine-like setae; dor- sally with scattered setae each surrounded by a dark spot; propleuron forming right angle, tipped by stout seta (Fig. 20). Legs: foreleg (Fig. 20) with long setae laterally on coxa, long ventral setae on trochanter and femur; mid leg (Fig. 21) and hind leg (Fig. 22) with thin setae laterally on coxa and
12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
13}.
vaginal sclerite
spermathecal duct
IXth segment
Xth segment
spermatheca
Figures 10-15. Helicopsyche khemoiensis (female). Figure 10. Wings.
Figure 11. VIth sternal process of abdomen, lateral view. Figure 12. Genitalia, lateral view.
Figure 13. Genitalia, dorsal view.
Figure 14. Genitalia, ventral view.
Figure 15. VIth sternal process of abdomen, ventral view.
ventrally on trochanter. Abdomen (Fig. 23) with small group of setae dorsolaterally on segment IX; long setae dorsad of anal prolegs on segment X; abdominal gills absent; anal gills form pair of expanded lobes; anal claw minute with many teeth. The larval case is similar to that illustrated by Wiggins (1996a).
Pupal case: anterior enclosure (Fig. 26) of densely woven silk for 75% of surface; the inner, ventral quadrant with several open bands bearing transverse strands of silk. Pupa: pupal mandibles and hook- plates as illustrated by Wiggins (1996b, fig. 17-87). Segment IX with long setae; two pairs dorsolateral, two pairs lateral and two pairs ventrolateral; anal appendages, lateral view (Fig. 24) short, angled dorsad, with two pairs long setae posteriorly and six pairs ventrally; in ventral view (Fig. 25) oriented posteriad; the ventral setae in two longitudinal rows.
Diagnosis.—Males of H. khemoiensis can be separated from closely related H. coreana by the star-shaped gonocoxal plate formed by the fused secondary
2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES
a
pronieroye |
Figures 16-23. Helicopsyche khemoiensis (larva). Figure 16. Head, lateral view.
Figure 17. Head, dorsal view.
Figure 18. Head, ventral view.
Figure 19. Thorax, dorsal view.
Figure 20. Propleuron and fore leg, anterior view. Figure 21. Mid leg, anterior view.
Figure 22. Hind leg, anterior view.
Figure 23. Abdominal segments IX and X, lateral view.
25
Figures 24—26. Helicopsyche khemoiensis (pupa).
Figure 24. Segment IX and anal appendages, lateral view. Figure 25. Segment IX and anal appendages, ventral view. Figure 26. Pupal case membrane.
13
14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
branches of the gonocoxite; by the Xth tergum which lacks megasetae, is shal-
lowly bifurcate and divergent apically, hooked dorsally and which bears short
baso-lateral processes. Females are recognized by the acuminate hind wing, the
short rounded VIth sternal process and by the large dorsal process of the Xth
segment which curves ventrad concealing the small ventral branch. Etymology.—khemoiensis, refers to river from which species was collected. Material examined.—see Types.
HELICOPSYCHE AZUNENSIS SCHEFTER AND JOHANSON, NEW SPECIES (Fics. 27—33)
Types.—Holotype, male: VIETNAM. GIA LAI: An Khe Dist; Tram Lap, Azun R., 2 km NW on trail from forestry building, 17 Jun 1996, D.C. Currie, J. Swann, ROM 961056, UV light, at rainforest edge/coffee plantation, 14°27’ N, 108°33' E.
Description —Male. Head (Fig. 27): antennae with lobate scape about 2X length of pedicel, 1st flagellomere as long as pedicel; eyes very large; interantennal warts as long as scape breadth, spherical, cephalic warts large, ovoid, laterad of elevated, triangular vertex; maxillary palps 4-segmented, the basal two segments together longer than labial palp. Legs: spur formula 1,2,4; fore leg spur longer than first tarsal segment. Pronotum with 2 pairs setal warts. Wings (Fig. 28): fore wing length 3.0 mm; forks 1,2,3 and 5 present; fork 1 originating on distal one third of Dc, R, subequal to Dc, A, and A, form basal loop, A,,, reaches wing margin close to Cu,: hind wing 2.3 mm, with 14 hamuli. Abdomen with small VIth sternal process. Genitalia: segment IX in lateral view (Fig. 29) subtriangular, with well developed lateral apodeme; narrow ventrally (Fig. 31); anterior margin ogival in dorsal view (Fig. 30); gonocoxite with large primary and small secondary branch; primary branch in lateral view (Fig. 29) with dorsal margin gently concave, apically hatchet-shaped; secondary branch slender, arcuate with slight apical hook (Fig. 31); superior appendage originating dorsally on segment nine, directed ventrad, clavate in lateral view; Xth tergum depressed in lateral view, in dorsal view (Fig. 30) bifurcate for one half length, divided into two tapering processes; macrosetae absent. Phallus thick, slightly curved ventrad (Fig. 32), dorsoapical membranous part of endotheca bifurcate, lobate, with a pair of slender sigmoid processes each with a small lateral tooth (Figs. 32 and 33).
Female and immature stages.—Unknown.
Diagnosis.—Helicopsyche azunensis is most closely related to H. chrysothoe (Schmid) and can be separated from it and other species with 4-segmented max- illary palps by the following characters: gonocoxite lacking an internal branch (present in H. chrysothoe); with hatchet-shaped primary branch longer than the tapered slightly notched secondary branch (in most other species of this group the secondary branch is strongly notched); lobate Xth tergum lacking macrosetae or notches (in other species the Xth tergum is acuminate and frequently notched apically); and phallus with a pair of stout sigmoid sclerous processes each with a subapical tooth (these do not occur in other species in this group).
Etymology.—azunensis, refers to river at collection site.
Material examined.—see Types.
HELICOPSYCHE DACKLESTENSIS SCHEFTER AND JOHANSON, NEW SPECIES (Figs. 34—39)
Types.—Holotype, male: VIETNAM: GIA LAI, An Khe Dist., Dacklest River, 5.2 km NE Tram Lap on forest road, 28 Jun 1996, D. C. Currie, J. Swann, ROM 961102, UV light, 200 m upstream bridge, 1° rainforest, 900 m, 14°24’ N, 108°33’ E.
Description.—Male. Head: as in Helicopsyche azunensis, except vertex forming a smaller triangle.
2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 15
Figures 27-33. Helicopsyche azunensis (male). Figure 27. Head, lateral view.
Figure 28. Wings.
Figure 29. Genitalia, lateral view.
Figure 30. Genitalia, dorsal view.
Figure 31. Genitalia, ventral view.
Figure 32. Phallus, lateral view.
Figure 33. Phallus, dorsal view.
Legs and pronotum as in H. azunensis. Wings (Fig. 34): fore wing length 2.7 mm, forks 1,2,3 and 5 present, fork 1 originating at distal one third of Dc, R, subequal to Dc, A,,, well separated from wing margin, does not reach wing margin; hind wing 2.1 mm; with 18 hamuli. Abdomen with well devel- oped VIth sternal process. Genitalia: segment IX, lateral view (Fig. 35), subtriangular and longer dorsally, with well developed lateral apodeme. Gonocoxite divided into large primary and small sec- ondary branch; primary branch in lateral view (Fig. 35) clavate with undulate apical margin; secondary branch slender, arcuate, notched at apex. Superior appendage originates medially on segment nine, directed ventrad, clavate in lateral view (Fig. 35); Xth tergum slender, slightly sinuate, strongly de- pressed in lateral view (Fig. 35), in dorsal view (Fig. 36) deeply bifurcate, each tapering lobe with a small ventrolateral process (Fig. 37); macrosetae absent. Phallus thick, slightly curved ventrally (Fig. 38) dorsoapical membranous endotheca expanded as posterior lobes with embedded paired slender sclerous processes curved apicolaterally (Figs. 38 and 39). Female and immature stages.—Unknown.
16
THE PAN-PACIFIC ENTOMOLOGIST
Figures 34-39. Helicopsyche dacklestensis (male).
Figure 34. Figure 35. Figure 36. Figure 37. Figure 38. Figure 39.
Wings.
Genitalia, lateral view. Genitalia, dorsal view. Genitalia, ventral view. Phallus, lateral view. Phallus, dorsal view.
Vol. 77(1)
Diagnosis.—Helicopsyche dacklestensis is most similar to H. xenothoe (Schmid) and can be separated from that and other species in this group by the slender and strongly arched Xth tergum, deeply bifurcate with small mesolateral processes, and the armature of the phallus.
Etymology.—dacklestensis, refers to river at collecting site.
Material Examined.—see Types.
DISCUSSION
Helicopsyche khemoiensis is most closely related to the North Korean H. co- reana Mey. Both are restricted to the East Asian subregion (sensu Banarescu 1992), the two species comprising the subgenus Galeopsyche (Johanson 1998). Characteristics supporting monophyly of this group are (plesiomorphic states in parenthesis): reduction of the lateral pronotal setal warts (present, unreduced);
2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 17
Figure 40. Collecting sites of Vietnamese Helicopsyche. A. H. awudschgal Malicky. B. H- khe- moiensis. C. H: azunensis. D. H. dacklenstensis.
fore wing crossvein R,,;—R,,; directed posterobasad (directed ventrad), and fore wing A,,, meeting Cu, or fusing with the posterior wing margin close to Cu, (distant to Cu,). East Asian subgenus Galeopsyche and the South American sub- genus Cochliopsyche are sister groups (Johanson 1998).
Species described in genus Cochliophylax (Schmid 1993) from northeastern India and Nepal were redesignated as a derived monophyletic clade within the subgenus Helicopsyche (Johanson 1998). Helicopsyche azunensis and H. dack- lestensis are closely related to the many Cochliophylax species described by Schmid, and together with them have 4-jointed maxillary palps, mesoscutal warts absent, narrow hind wings without crossvein M-Cu, and the gonocoxal secondary branch tapered and with an apical hook. Helicopsyche azunensis and H. dackles- tensis are considered sister species based on the presence of phallic endothecal sclerites. Their relationship to the remaining species in the group is as yet un- determined.
Distribution of the four known Vietnamese Helicopsyche species is shown in Fig. 40.
18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
ACKNOWLEDGMENT
We acknowledge the support of the Entomology division of the Centre for Biodiversity and Conservation Biology at the Royal Ontario Museum, Toronto, Ontario, Canada. This constitutes Contribution Number 158 of the Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario, Canada.
LITERATURE CITED
Banarescu, P. 1992. Zoogeography of fresh waters. Volume 2. Distribution and dispersal of freshwater animals in North America and Eurasia. AULAG-Verlag Wiesbaden.
Dudgeon, D. 1988. Preliminary investigations on the faunistic and ecology of Hong Kong Trichoptera. pp. 111-117 in Bournaud, M. & H. Tachet (eds), Proc. 5th Int. Symp. Trichoptera. Junk, The Hague.
Johanson, K. A. 1998. Phylogenetic and biogeographic analysis of the family Helicopsychidae (Insecta: Trichoptera). Ent. Scand. Suppl., 53: 1-176.
Malicky, H. 1994. Zwei neue Helicopsyche (Helicopsychidae) aus Perak, Malaysia. Braueria, 22: 4.
Malicky, H. 1995. Neue Kocherfliegen (Trichoptera, Insecta) aus Vietnam. Linzer Biol. Beitr. 27: 851- 885.
Malicky, H. & P. Chantaramongkol. 1993. Neue Trichopteren aus Thailand. Teil 2: Rhyacophilidae, Philopotamidae, Polycentropodidae, Ecnomidae, Psychomyidae, Xiphocentronidae, Helicopsy- chidae, Odontoceridae (Arbeiten iiber thailandische K6cherfliegen Nr. 12) (Fortsetzung). Linzer Biol. Beitr, 25: 1137-1187.
Say, T. 1821. Descriptions of univalve shells of the United States. J. Acad. Phil., 2, 1.
Schmid, E 1993. Considérations sur les Hélicopsychidés (Trichoptera, Integripalpia). Beaufortia, 43: 65-100.
Wiggins, G. B. 1996a. Larvae of the North American caddisfly genera (Trichoptera) (2nd ed). Uni- versity of Toronto Press, Toronto, Buffalo, London.
Wiggins, G. B. 1996b. Trichoptera families. In Merrit, R. W. & K. W. Cummins (eds). An introduction to the aquatic insects of America (3rd ed). Kendall/Hunt, Dubuque, Iowa.
Williams, D. D., A. T. Read & K. A. Moore. 1983. The biology and zoogeography of Helicopsyche borealis (Trichoptera: Helicopsychidae): a Nearctic representative of a tropical genus. Can. J. Zool., 61: 2288-2299.
Received 27 Apr 1999; Accepted 7 Feb 2000.
PAN-PACIFIC ENTOMOLOGIST 77(1): 19-27, (2001)
PREY OF THE SPIDER, DICTYNA COLORADENSIS, ON APPLE, PEAR, AND WEEDS IN CENTRAL WASHINGTON (ARANEAE: DICTYNIDAE)
EUGENE R. MILICZKY AND CARROL O. CALKINS
Yakima Agricultural Research Laboratory, United States Department of Agriculture—Agricultural Research Service, 5230 Konowac Pass Road, Wapato, Washington 98951
Abstract.—The-cribellate spider, Dictyna coloradensis Chamberlin, constructed webs on the up- per surface of apple and pear leaves (trees not treated with insecticide), and on weeds in adjacent, uncultivated ground, at a site in south central Washington. Prey found in D. coloradensis webs were assigned to one of three categories: pests, predators and parasitoids, or neutral in impact with respect to fruit trees. Pest taxa comprised 32%, predators and parasitoids 24%, and neutral groups 44% of 18,314 prey. Most prey were small, winged insects (length < 5 mm). Insects from 58 families in 10 orders were represented and small spiders in four families were occa- sionally trapped. Sciaridae and Chironomidae (Diptera) were the most numerous prey and made up 37% of the total. Most webs contained one or more of these flies and occasionally 25 or more were trapped. Alate aphids were the most frequently captured pest insects. Other pests included adults of the white apple leafhopper, the pear psylla, and thrips. Relatively non-mobile stages of the pests (leafhopper and pear psylla nymphs and apterous aphids) were less commonly found in the webs. Nineteen percent of all prey were parasitoid wasps, 14 families of which were identified. Known parasitoids of apple and pear pests were included. The only other pred- ator or parasitoid taxon that comprised more than 1% of total prey was the Empididae (3%).
Key Words.—Arachnida, Araneae, spider, Dictyna, prey use, apple, pear.
The cribellate spider genus Dictyna is represented in the Nearctic region by more than 100 species (Roth 1993). Dictyna construct irregular mesh webs in a variety of situations, at times in considerable numbers and high densities (Cham- berlin & Gertsch 1958, Heidger & Nentwig 1985). Species of Dictyna have fre- quently been reported from orchards where they are at times abundant. Muma (1975) found D. florens Ivie and Barrows common and widespread in Florida citrus where it constructed webs on leaves of orange and grapefruit. Putman (1967) reported D. annulipes Blackwall to be a common spider in Ontario, Canada peach orchards where it constructed webs on areas of rough bark. Also in Ontario, Hagley & Allen (1989) found D. annulipes to be the most abundant foliage- inhabiting spider in an apple orchard and they studied its prey utilization by examination of webs and assay of gut contents.
Dictyna coloradensis Chamberlin occurs throughout much of the central and northern United States and into the Northwest Territories of Canada. With females approaching 4 mm in length, it is among the larger species in the genus (Cham- berlin & Gertsch 1958). Dondale (1956) reported D. coloradensis from apple trees in Nova Scotia, Canada. During 1997, 1998, and 1999 this spider was very abun- dant on foliage of apple trees at the USDA-ARS research farm near Yakima, Washington. Webs were less abundant on pear foliage. Large numbers of D. co- loradensis also constructed webs on tall, dead stalks of annual weeds in adjacent, uncultivated ground in the spring, and later in the year utilized the current season’s growth.
Webs of D. coloradensis were collected during 1997, 1998, and 1999 and their
20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
prey contents identified. Because spiders are considered generalist predators (Wise 1993), we were interested in determining the taxonomic range of prey captured and the relative proportions that fell into three broad categories. 1) Pests: Included taxa are generally regarded as plant pests although not all specimens were nec- essarily pests of apple and pear. 2) Predators and parasitoids: All predatory and parasitic groups were included although not all were known predators and para- sitoids of apple and pear pests. 3) Neutral: Taxa in this category probably have little or no detrimental or beneficial impact on fruit trees.
MATERIALS AND METHODS
This study was conducted at the USDA-ARS research farm, 26 km east of Yakima, Yakima County, Washington. Several small blocks of fruit trees are plant- ed at the 130 ha farm. Other crops grown include asparagus and potatoes, but some ground has never been cultivated and native vegetation, dominated by big sagebrush, Artemisia tridentata Nuttall (Asteraceae), remains within and sur- rounds parts of the farm. Uncultivated ground with mixed native and introduced vegetation partly surrounded some of the tree fruit blocks. Fruit trees were not treated with insecticides.
Pear and apple leaves and weed stems with D. coloradensis webs were placed in plastic vials with tight fitting lids and refrigerated until examined. Webs were immersed in 70% isopropyl alcohol in a petri dish and examined under 6.5 X— 50 for prey identification. Prey were identified to family, if possible, using keys in Borror, Delong, & Triplehorn (1976) and Goulet & Huber (1993). Exceptions included the following: Chironomidae and Sciaridae, the most abundant Dipteran prey, were difficult to distinguish when large numbers were present and specimens were damaged and entangled in webbing. Many specimens were therefore cate- gorized as unidentified Nematocera. These two families made up the vast majority of prey in this category. Cyclorrhaphous Brachycera were categorized as medium size muscoid flies if larger than Drosophila but smaller than a housefly, and as small muscoid flies if Drosophila-size or smaller. Many chalcidoid wasps were identified only to superfamily (Chalcidoidea) and many small, non-chalcidoid, parasitoid wasps were classified as unidentified parasitoid wasps because small size, damaged specimens, and entanglement in webbing made identification dif- ficult.
Webs were collected from three apple varieties (“‘Fuji’’, ‘““Golden Delicious’’, and ‘Red Delicious’’), pears (mixed “‘Anjou”’ and “‘Bartlett’’ varieties), and dead weed stems in uncultivated land adjacent to the “‘Fuji’”’ apples. Weeds were pri- marily tumble mustard, Sisymbrium altissimum Linnaeus (Brassicaceae), an intro- duced species. A total of 984 webs was examined, distributed among the plant types as indicated in Table 1. Five to 15 webs were collected from a plant type per sample date at one to two week intervals. The sampling periods were 22 May to 16 Oct 1997, 30 Mar to 3 Nov 1998, and 20 Jul to 1 Oct 1999. Each plant type was not sampled throughout each sample period.
RESULTS
Web Placement.—Most webs on apple and pear were constructed on the upper, concave surfaces of leaves. The small webs of young spiders covered only 2 or 3 cm? and were usually near the leaf apex. Webs of older, larger individuals often
2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 21
Table 1. Number of sample dates and total number of Dictyna coloradensis webs examined from different plants during 1997, 1998, and 1999.
No. of samples dates—No. of webs examined
Plant 1997 1998 1999 Apple—“‘Fuji”’ 11-141 19-192 — Apple—“Golden”’ 10-57 10-56 — Apple—“‘Red”’ 9-59 — — Pear 5-17 9-91 11-155 Weeds —_ 18-216 —
covered most of the upper surface of a leaf. Leaves were up to 10 cm long. Prey accumulated in webs and older, larger webs contained up to 40 or more prey. Webs were occasionally found in the angle formed by two branches, among flower petioles, or between a leaf petiole and a branch. They were less visible in these locations and few were sampled. Webs on dead weed stalks were constructed among branches on the upper part of a stalk, 0.3—-1.0 m above ground.
Prey Utilization.—Fifty-eight families of insects in 10 orders and four families of spiders were identified from webs of D. coloradensis (Table 2). Some taxa were represented by few specimens (two Ephemeroptera among 18,314 prey) whereas others made up a high proportion of prey in webs from all sources all three years.
Insects ‘classified as neutral with respect to impact on fruit trees made up the largest proportion of prey items overall (7981 prey = 44%). Nematocerous Dip- tera, primarily Sciaridae (dark-winged fungus gnats) and Chironomidae (midges) were the most abundant prey of any kind (37% overall). Sciarids and chironomids were present throughout the season and were found in a majority of webs re- gardless of source. Webs occasionally contained 25 or more of these small insects. Other taxa of neutral prey rarely comprised more than 1% of the total from a plant in one year. Small muscoid flies, however, made up 5.4% of prey in webs from “‘Golden Delicious” in 1997.
Aphids were the most numerous pest insects found in D. coloradensis webs, and alates were generally much more abundant than apterous forms. Five to 30% of the total insects found in webs from each plant type each year were aphids. Aphids were not identified to species because of the large number captured and their often poor state of preservation (discoloration, damage, dehydration). Several Species are considered pests of apple in Washington and many appeared to be green apple aphid, Aphis pomi DeGeer, or the nearly identical spirea aphid, A. spireacola Patch. Apple/spirea aphid colonies were abundant on developing apple shoots all three years. Thrips (Thysanoptera) made up 5—11% of total prey in the samples from fruit trees but were more abundant in webs on weeds (22%). The western flower thrips, Frankliniella occidentalis (Pergande) is the only species listed by Beers et al. (1993) as a pest of tree fruit in Washington. A pale, yellowish insect, its host range includes several fruit trees, alfalfa, potatoes, and numerous species of weeds (Beers et al. 1993). Thrips were not identified to species due to small size, entanglement in webbing, and poor preservation. The vast majority, however, were pale, yellowish insects, in general resembling F. occidentalis. White apple leafhopper adults, Typhlocyba pomaria McAtee, were most abundant
22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Table 2. Total number of prey items in each taxon found in Dictyna coloradensis webs from apples, pears, and weeds.
Prey taxa Apples Pears Weeds
Neutral impact taxa
Chironomidae 195 163 745 Sciaridae 760 238 112 Bibionidae 25 0 94 Psychodidae 6 2 1 Scatopsidae 1 4 1 Simuliidae 1 3 0 Tipulidae 1 1 0 Nematocera—unidentified 2873 1022 590 Stratiomyidae 1 0 0 Therevidae 4 0 0 Bombyliidae 2 0 3 Drosophilidae 175 78 28 Phoridae 38 29 8 Conopidae 0 0 1 Small muscoid flies 375 100 87 Medium muscoid flies 30 26 10 Diptera—unidentified 0 0 5 Halictidae 12 7 0 Chrysididae 1 0 0 Tenthredinidae 0 0 1 Scarabaeidae 2 0 pi Coleoptera—unidentified 6 1 6 Aleyrodidae 1 0 0 Fulgoroidea 0 0 1 Lygaeidae 6 0 17 Hemiptera—unidentified 2 1 3 Psocoptera 42 18 9 Ephemeroptera 1 1 0 Pest taxa Aphididae—alate 1549 1011 431 Aphididae—apterous 230 67 5 Typhlocyba pomaria 289 ms) 2D Other Cicadellidae 18 1 2 Cacopsylla pyricola—adults 3 165 1 Cacopsylla pyricola—nymphs 0 50 0 Phyllonorycter elmaella 193 37 3 Lepidoptera—unidentified 3 0 3 Caterpillar 0 2 0 Mindae—Lygus sp. Z 0 3 Mindae—Campylomma sp.? 12 1 1 Thysanoptera 687 281 777 Acari 3 5 0 Predator and parasitoid taxa Cecidomyiidae 6 5 4 Empididae 387 132 6 Dolichopodidae 11 4 0 Pipunculidae 25 9 3 Tachinidae 6 6 1 Syrphidae 2 0 0 Braconidae 349 56 81 Ichneumonidae 76 16 18
2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 23
Table 2. Continued.
Prey taxa Apples Pears Weeds
Pnigalio flavipes 177 63 3 Trechnites insidiosus 0 50 0 Mymaridae 641 17 20 Encyrtidae 0 0 1 Chalcidoidea—unidentified 713 373 106 Platygastridae 40 16 21 Scelionidae 25 14 5 Ceraphronidae 73 14 16 Megaspilidae 2 2 1 Proctotrupidae 3 4 1 Dryinidae 1 1 2 Bethylidae 3 0 6 Diapriidae 1 0 3 Cynipoidea 12 3 1 Parasitoids—unidentified® 249 94 123 Sphecidae 8 8 0 Formicidae 15 7 133 Vespidae 1 1 0 Staphylinidae 39 4 35 Carabidae 2 0 3 Coccinellidae PL 1 0 Hemerobiidae 6 2 0 Chrysopidae ys 0 0 Anthocoridae—Orius 27 5 of Nabidae—Nabis 0 0 1 Lygaeidae—Geocoris 0 0 5 Miridae—Deraeocoris 2 2 0 Salticidae 7 1 1 Liny phiidae—Erigone 13 2. 2 Linyphiidae 10 9 17 Thomisidae 2 0 0 Oxyopidae 6 1 0 Araneae—unidentified 0 2 0 Prey totals 10,493 4243 3578
4 Campylomma also act as predators by feeding on such pests as aphids and mites. > Category includes only hymenopteran parasitoids.
in webs from “Fuji” apples during 1998 when they comprised nearly 5% of all prey. Typhlocyba comprised less than 2% of total prey in the other samples, and nymphs were rarely captured. Adult, western tentiform leafminer, Phyllonorycter elmaella Doglanar and Mutuura, made up 1—2% of the prey in each tree fruit sample, but only three of 3578 prey in webs from weeds. Pear psylla, Cacopsylla pyricola (Foerster), a serious pest of pear in the Pacific Northwest, made up 5.5% of the prey in D. coloradensis webs from pear during 1997, just under 1% in 1998, and 8% in 1999. Psylla populations at the research farm were low during 1998, probably accounting for their rarity as prey despite the greater number of webs examined compared to 1997. Psylla numbers were higher during 1999 when leaves and shoots were often sticky with honeydew, and this was reflected in the number captured in D. coloradensis webs. Most captured C. pyricola were adults, although a substantial number of nymphs fell victim in 1999 (50 nymphs, 134
24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
adults). However, it was difficult, at times, to distinguish psylla nymphs from their cast skins.
Small, hymenopterous parasitoids were the most abundant predatory and par- asitoid insects trapped in D. coloradensis webs. They comprised 12% to 28.6% of total prey in yearly samples from each of the plants. Hosts of many of the parasitoids were not determined or are unknown, some are probably hyperpar- asitoids, and some probably occurred only incidentally in the fruit trees. Known parasitoids of apple and pear pests were, however, captured. Pnigalio flavipes (Ashmead) (Eulophidae), the most common parasitoid of the western tentiform leafminer in the Pacific Northwest (Beers et al. 1993), made up 1%—2% of total prey in samples from fruit trees. It was rarely found in webs on weeds. Mymaridae (fairyflies) were captured in substantial numbers in webs on “Fuji”? (449 = 11.9%) and “‘Golden Delicious” (132 = 6.6%) during 1998. Both represented large increases over numbers found in 1997. Mymarids in the genus Anagrus are important egg parasitoids of the white apple leafhopper and parasitism rates of up to 70% have been reported in unsprayed orchards (Beers et al. 1993). Many mymarids found in webs on apple may have been leafhopper egg parasitoids. Two percent of the insects in webs from pear during 1999 were Trechnites insidiosus (Crawford) (Encyrtidae), the most important parasitoid of pear psylla in western North America (Beers et al. 1993).
Other taxa of insect predators and parasitoids rarely comprised more than 1% of the total prey in webs from a given plant type (Table 2). Empididae, however, made up 7.5% (233 flies) of the prey found in webs on “Fuji” apple in 1997. Spiders also were infrequently snared in D. coloradensis webs (< 1% of prey in samples from any of the plant types) and were small, either immatures or taxa of small body size.
DISCUSSION
The diversity of prey captured by D. coloradensis is in agreement with the idea of spiders as generalist predators (Wise 1993). Individual species, however, utilize a restricted range of available prey depending on factors such as spider size, hunting strategy, and web size and placement (Marc & Canard 1997). Spiders generally feed on prey smaller than themselves (Jackson 1977, Nyffeler et al. 1994). This was true of D. coloradensis, the vast majority of whose prey consisted of insects less than 5 mm in length. The predominant use of small prey has also been noted in D. segregata (Nyffeler et al. 1988), D. arundinacea (Heidger & Nentwig 1985), and 11 species, including D. coloradensis, studied by Jackson (1977). Large and dangerous prey were, however, captured occasionally. Two worker yellow-jackets (Vespula sp.) were found in 1997 webs and on 26 Apr 2000 a recently captured worker honeybee, Apis mellifera L., was noted in the web of a female on a dead weed. The spider was feeding on the bee at the time. Overall, however, few prey larger than a housefly were found in the webs. No codling moth, Cydia pomonella (L.) or leafrollers, Pandemis pyrusana Kearfott, were captured. Both are important apple pests in Washington (C. pomonella was very abundant at the farm in 1997-1998), but at adult lengths of 12 mm or more, they may be too large to be readily subdued by D. coloradensis webs.
A given species of spider may not utilize all stages in a prey species’ life cycle to equal degrees (Marc and Canard 1997). This was true for several prey species
2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 23
of D. coloradensis, most or all of whose life cycles are spent on the fruit trees. Adult white apple leafhopper, adult pear psylla, and alate aphids were common prey but immature leafhoppers and psylla and apterous aphids fell victim less frequently. This is probably related to the relative mobility of different stages of the prey and their locations on the plant. White apple leafhopper and pear psylla adults are mobile and fly readily when disturbed. They would appear much more likely to blunder into webs than the more sedentary nymphs. Also, white apple leafhopper nymphs generally feed on the lower surface of the leaf (Beers et al. 1993) whereas D. coloradensis webs are constructed almost exclusively on the upper surface. Psylla nymphs, with their flattened body form and leaf-surface hugging habits were often found alive beneath D. coloradensis webs—apparently able to avoid entanglement in the silk. Alate aphids, although not strong fliers, are more mobile than the apterous forms and again must be more likely to come into contact with webs. Green apple aphid colonies generally develop on succu- lent, young tissue and are found on growing shoot tips, shoot stems, and the undersides of leaves (Beers et al. 1993). Thus the sedentary, apterous forms are less likely to come into contact with D. coloradensis webs.
Dictyna coloradensis webs trapped many small (1—3 mm), hymenopterous par- asitoids (19.1% of all prey). Some parasitoids were observed crawling over leaf surfaces, which must often bring them into contact with webbing. This is true of P. flavipes and T. insidiosus females, both of which search leaf surfaces for hosts (Beers et al. 1993), and many became entangled in the webs. Nearly the same number of male P. flavipes (122) were captured as females (118). T. insidiosus is thelytokous in the western states (Unruh et al. 1995) and males were not found in the webs. Thirteen percent of the prey of D. arundinacea (L.) consisted of small parasitoid wasps (Heidger & Nentwig 1985).
Predatory insects, and parasitoids other than Hymenoptera, were infrequently found in D. coloradensis webs. Deraeocoris spp. (Miridae), for example, are important predators of pear psylla (Beers et al. 1993) and were abundant on the pears during 1999. Yet Deraeocoris accounted for only 0.09% of the prey in webs on pear during 1999. Aphid predators such as Coccinellidae and Chrysopidae, often abundant in unsprayed orchards with high aphid populations, made up sim- ilarly low percentages of prey in all samples. Such insects, perhaps because of size, behavior, and distribution on the plants may not be very vulnerable to en- tanglement in D. coloradensis webs.
Other spiders were infrequent prey in D. coloradensis webs. The low number taken (0.4% of total prey) is in accord with Nyffeler’s (1999) findings that web building spiders are 99% insectivorous whereas hunting types take a higher pro- portion of other spiders. Jackson (1977), for example, found that 27% of the prey of Phidippus johnsoni (Peckham and Peckham) (Salticidae) was other spiders.
Capture of beneficial insects and spiders by D. coloradensis (intraguild pre- dation in a broad sense) was substantial in terms of the overall numbers captured. However, the hosts of many of the parasitoids may not be orchard pests and their capture may have little negative impact on orchard ecology from a pest manage- ment standpoint. Greenstone (1999) noted that the net effect of intraguild pre- dation can only be determined by examining the system in the presence and absence of the predator. Although intraguild predation and competitive interac- tions among predators may in some cases disturb natural pest control, in others
26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
they may promote greater spider biodiversity and allow spiders to survive periods of low prey density (Sunderland 1999). |
Several studies have shown that small Diptera are important components in the diet of Dictyna spiders. Several families of small flies made up 50% of the prey captured by D. arundinacea in a meadow in Germany (Heidger & Nentwig 1985) and small Diptera dominated the prey of each of 11 Dictyna species studied by Jackson (1977). Chironomidae comprised 70.7% of the total prey of D. annulipes in an apple orchard in Ontario, Canada (Hagley & Allen 1989), and a diverse array of small Diptera were important components in the diet of D. coloradensis in Washington. The abundance of these small flies was likely important in sup- porting the high population of D. coloradensis observed during this study. Also, their presence during much of the season may help fill in gaps in availability of other types of prey, as noted by Sunderland (1999) with respect to intraguild predation.
Clearly, D. coloradensis is a polyphagous predator that includes a wide range of insects in its diet. Prey selectivity is based primarily on size and activity level— small, active insects are most heavily utilized. Small, active pests of apple and pear are taken in substantial numbers, but unfortunately, from a pest management standpoint, known parasitoids of some pests are also trapped, sometimes in con- siderable numbers.
ACKNOWLEDGMENT
We thank Matthew H. Greenstone, Thomas R. Unruh, and an anonymous re- viewer for their constructive comments on the manuscript.
LITERATURE CITED
Beers, E. H., J. EK Brunner, M. J. Willett & G. M. Warner. (eds.). 1993. Orchard pest management: A Resource book for the Pacific Northwest. Published by Good Fruit Grower, Yakima, Washing- ton.
Borror, D. J.. D. M. DeLong & C. A. Triplehorn. 1976. An introduction to the study of insects (4th ed.) Holt, Rinehart, and Winston, New York.
Chamberlin, R. V. & W. J. Gertsch. 1958. The spider family Dictynidae in America north of Mexico. Bull. Am. Mus. Nat. Hist., 116: 1-152.
Dondale, C. D. 1956. Annotated list of spiders (Araneae) from apple trees in Nova Scotia. Can. Entomol., 88: 697—700.
Greenstone, M. H. 1999. Spider predation: how and why we study it. J. Arachnol., 27: 333-342.
Goulet, H. & J. T. Huber (eds.). 1993. Hymenoptera of the world: An identification guide to families. Research Branch, Agriculture Canada. Publication 1894/E.
Hagley, E. A.C. & W.R. Allen. 1989. Prey of the cribellate spider, Dictyna annulipes (Araneae, Dictynidae), on apple tree foliage. J. Arachnol., 17: 366-367.
Heidger, C. & W. Nentwig. 1985. The prey of Dictyna arundinacea (Araneae: Dictynidae). Zool. Beitr. N. E 29: 185-192.
Jackson, R. R. 1977a. Comparative studies of Dictyna and Mallos (Araneae, Dictynidae): III. Prey and predatory behavior. Psyche, 83: 267-280.
Jackson, R. R. 1977b. Prey of the jumping spider Phidippus johnsoni (Araneae: Salticidae). J. Arach- nol., 5: 145-149.
Marc, P. & A. Canard. 1997. Maintaining spider biodiversity in agroecosystems as a tool in pest control. Agric. Ecosyst. Environ., 62: 229-235.
Muma, M. H. 1975. Spiders in Florida citrus groves. Fla. Entomol., 58: 83-90.
Nyffeler, M. 1999. Prey selection of spiders in the field. J. Arachnol., 27: 317-324.
Nyffeler, M., D. A. Dean & W. L. Sterling. 1988. Prey records of the web-building spiders Dictyna segregata, (Dictynidae), Theridion australe (Theridiidae), Tidarren haemorrhoidale (Theridi-
2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 27
idae), and Frontinella pyramitela (Linyphiidae) in a cotton agroecosystem. Southwestern Nat., 33: 215-218.
Nyffeler, M., W. L. Sterling & D. A. Dean. 1994. How spiders make a living. Environ. Entomol., 23: 1357-1367.
Putman, W. L. 1967. Prevalence of spiders and their importance as predators in Ontario peach orchards. Can. Entomol., 99: 160-170.
Roth, V. D. 1993. Spider genera of North America with keys to families and genera and a guide to literature (3rd ed.). (Distributed by American Arachnological Society). U. of Florida, Gaines- ville.
Sunderland, K. 1999. Mechanisms underlying the effects of spiders on pest populations. J. Arachnol., 27: 308-316.
Unruh, T. R., P. H. Westigard & K. S. Hagen. 1995. Pear psylla. Chapter 19. pp. 95-100. In Nechols, J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden, & C. G. Jackson. 1995. Biological control
in the western United States. Accomplishments and benefits of regional research project W-84, 1964-1989. Univ. Calif. Div. Agric. Nat. Res. Publ. 3361.
Wise, D. H. 1993. Spiders in ecological webs. Cambridge University Press, Cambridge, U.K. Received 29 Nov 1999; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 28-36, (2001)
OVERWINTERING POTENTIAL IN CALIFORNIA OF TWO RUSSIAN WHEAT APHID PARASITOIDS (HYMENOPTERA: APHELINIDAE ET APHIDITDAE) IMPORTED FROM CENTRAL ASIA
JULIO S. BERNAL!, D. GONZALEZ, & ERNESTO DAVID-DIMARINO
Department of Entomology, University of California, Riverside, California 92521-041
Abstract.—Aphelinus albipodus Hayat and Fatima and Diaeretiella rapae (M’Intosh) (Hyme- noptera: Aphelinidae and Aphidiidae) were imported to the USA for biological control of Russian wheat aphid (Diuraphis noxia Mordwilko) in 1992. Early laboratory studies identified potential limitations for their successful establishment in northern California where these parasitoids were extensively colonized and evaluated against Russian wheat aphid. Specifically, those studies showed that A. albipodus was particularly susceptible to moderately low temperatures (10° C). These results for A. albipodus were in contradiction with seasonal field temperatures prevalent in its collection site (Tahcheng, People’s Republic of China) where field temperatures are below freezing for several months each year. This study sought to reconcile these apparently contra- dictory results. We examined whether short daylength-induced diapause, i.e. winter diapause, occurs in A. albipodus and D. rapae, and thus could be used as an overwintering strategy by these parasitoids. Under laboratory conditions, > % of A. albipodus individuals entered diapause when daylength was reduced from 14 h to 12 h. In contrast, diapause was not detected in D. rapae under the same conditions. However, we suggested that diapause may occur in D. rapae under shorter daylengths because (1) this parasitoid’s development and survivorship are not af- fected by temperatures associated with 12 h daylengths at its collection site (Wuqia, PRC), and (11) other studies have demonstrated diapause in this parasitoid at daylengths < 12 h. We discuss our results in light of A. albipodus’ rapid establishment in northern California, and the use of our analytical procedure as a component of a process for screening natural enemies in importation biological control programs.
Key Words.—Insecta, Aphelinus albipodus, Diaeretiella rapae, diapause, biological control, Di- uraphis noxia, establishment, importation.
Numerous biological control campaigns involving exotic natural enemies have been successful since the introduction of the vedalia beetle to California more than 100 years ago (DeBach & Rosen 1991; Greathead & Greathead 1992). Ex- perience gained during that time and theoretical analyses have led to the identi- fication of traits that presumably make natural enemies effective biological control agents (Huffaker et al. 1974, 1977; Beddington et al. 1978; Murdoch et al. 1985; Luck 1990; Murdoch & Briggs 1996). However, at this time it is uncertain wheth- er these traits are useful or efficient for identifying the most promising natural enemies among a suite of available natural enemies (Luck 1990; Gonzalez & Gilstrap 1992). In contrast, identifying natural enemies that offer promise for successful establishment in a new area seems a more tenable goal. Characterizing an exotic natural enemy’s potential for establishment is important because lack of successful establishment precludes its eventual success in suppressing a pest pop- ulation.
Climate (excessively cold, hot, or dry seasonal weather) is the single most
‘Current Address: Biological Control Laboratory, Texas A&M University, College Station, Texas 77843-2475.
2001 BERNAL ET AL.: PARASITOID OVERWINTERING 29
frequently cited reason for natural enemy failure in biological control campaigns (Stiling 1993). Climate can early on limit a natural enemy’s potential for success by impeding its permanent establishment following colonization. Close matching between the climates of collection and colonization areas of natural enemies used in classical biological control campaigns, 1.e., climate-matching, is a long-standing conceptual “‘rule of thumb” for improving the chances of successful establishment of exotic natural enemies (Flanders 1940; Messenger 1959, 1971). It is unclear, however, how much emphasis is commonly placed on climate-matching in the planning phase of foreign exploration efforts. A comparatively information-inten- sive approach to climate-matching has been used successfully for forecasting range expansions of exotic pests (Messenger & Flitters 1954; Meats 1989; Hughes & Maywald 1990).
Shortly after the discovery of the Russian wheat aphid, Diuraphis noxia Mord- wilko (Homoptera: Aphididae), in the USA in 1986, a biological control campaign involving several state, federal, and university agencies was organized against this pest. Numerous Russian wheat aphid natural enemies were imported to the USA as a result of this combined effort (Gilstrap et al. 1994). Based on information available from their collection areas, a limited suite of parasitoids was selected from among these natural enemies for evaluation in California (Gonzélez et al., unpublished data). Among these parasitoids were Aphelinus albipodus Hayat and Fatima and Diaeretiella rapae (M’Intosh) (Hymenoptera: Aphelinidae and Aphi- diidae, respectively). A series of studies identified potential limitations for suc- cessful establishment of these parasitoids in California (Bernal & Gonzalez 1995, 1996, 1997; Bernal et al. 1997). In general, those studies suggested that A. albi- podus was less tolerant of low temperatures than D. rapae, but the reverse was true at high temperatures. The implications of these results were that establishment in California of A. albipodus could be restricted by winter temperatures, but es- tablishment of D. rapae could be restricted by summer temperatures. However, these results and their implications were in disagreement with available infor- mation concerning the climate of each of these parasitoid’s collection areas (ex- tremely cold and long winters, and moderate summers; see below). This was particularly true with regard to A. albipodus’ susceptibility to low temperatures in the laboratory (Bernal & Gonzalez 1996; Bernal et al. 1997), and its rapid establishment in northern California (Gonzalez et al., unpublished data).
Based on these observations, we examined whether A. albipodus and D. rapae enter diapause in response to a short daylength, 1.e., winter diapause. Entry into diapause under short daylength conditions would explain how these parasitoids survive the low temperatures prevalent during the late fall through early spring in their collection areas, and would influence their performance against Russian wheat aphid in California. Based on the results of this and previous studies we suggest that A. albipodus and D. rapae are able to survive the severe winters prevalent in their collection areas by entering a short daylength-induced diapause. In addition, we discuss how diapause and temperature-related developmental re- strictions can affect the potential for successful establishment and the population dynamics of these parasitoids under California conditions.
MATERIAL AND METHODS
Parasitoid and Host Cultures—Aphelinus albipodus and D. rapae were col- lected, respectively, near Tahcheng (46°42’ N, 83°00’ E, ca. 500 m elevation) and
30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Wugia (Ulugqat) (39°05’ N, 74°02’ E, ca. 3000 m elevation), Peoples Republic of China, by D. Gonzalez in 1992. The procedures for rearing Russian wheat aphid and both species of parasitoids were described earlier (Bernal & Gonzalez 1993).
Experiments.—Two sets of trials were conducted for each of A. albipodus and D. rapae. The first set was designed to provide baseline information concerming emergence rates from mummies, and the maximum egg to adult developmental times in each of the parasitoids. The second set was designed to detect diapause if it occurred in either parasitoid species. The methodology for both sets of trials, and for both parasitoids, was similar, except that the daylength used in the second set of trials was shorter than in the first set of trials.
Previous studies showed that emergence rates from mummies at 21° C, 50— 70% R. H., and 14 h daylengths were high (= 90%) in both A. albipodus and D. rapae. Thus, these environmental conditions (hereafter “‘non-diapause condi- tions’’) were used to obtain baseline information concerning emergence rates and developmental times in the first set of trials. Small pots of wheat, cv MIT, each covered with a small cage and containing ca. 40 Russian wheat aphids of instars If-adult (Bernal & Gonzalez 1997) were exposed to 20 previously mated and fed, 1—2 day old female A. albipodus or D. rapae. The aphids were exposed to the female parasitoids for 24 h, after which the parasitoids were removed and the aphids were incubated under the non-diapause conditions described above. When aphid mummies formed, they were transferred each to a gelatin capsule (size #0) and further incubated under non-diapause conditions to allow emergence of adult parasitoids. The mummies were monitored daily for emergence of adult parasit- oids, and the number of days from oviposition to emergence of individual adult parasitoids, the number of mummies formed, and the number of mummies yield- ing adult parasitoids from each pot were recorded. Fifteen pots with Russian wheat aphid were prepared for each of A. albipodus and D. rapae.
The second set of trials was similar to the first, except that the daylength was 12 h (hereafter “‘diapause conditions’’) rather than 14 h. In this case, 20 pots with Russian wheat aphid were used for A. albipodus and 15 pots for D. rapae. The results of the first set of trials showed that under non-diapause conditions, max- imum egg to adult developmental times were 28 days in A. albipodus (n = 344), and 23 days in D. rapae (n = 291). Thus, for the second set of trials, mummies of each species not yielding adults within their corresponding maximum egg to adult times were tentatively considered to be in diapause. Emergence rates from mummies under diapause versus non-diapause conditions were compared via log- likelihood ratios for 2 X 2 contingency tables, with Yate’s correction for conti- nuity, for each of A. albipodus and D. rapae (Zar 1996).
Mummies under diapause conditions yielded fewer adult parasitoids than mum- mies under non-diapause conditions in A. albipodus but not in D. rapae (see Results). Whether the A. albipodus mummies that did not yield adult parasitoids under diapause conditions indeed contained diapausing individuals was deter- mined via dissections. These mummies (n = 242) were divided into 2 groups of equal size at 28 days; one group was maintained under diapause conditions (i.e., 12 h daylength), the other was transferred to non-diapause conditions (i.e., 14 h daylength). Subsamples of mummies from each group were dissected at 48, 55, and 80 day from oviposition. Mummies containing live last instar parasitoid larvae
2001 BERNAL ET AL.: PARASITOID OVERWINTERING 31
Or pupae were considered to be in diapause. Both groups of mummies were further incubated if diapausing individuals were detected in the subsamples. The days to emergence, and the numbers of any parasitoids emerging from both groups of mummies were recorded.
Finally, the mean monthly temperatures and daylengths of each of the parasit- oid’s collection sites were compared versus a representative site from northern California, Tulelake (41°58' N, 121°28' W), where A. albipodus and D. rapae were colonized. Data for these comparisons were obtained from Walter & Lieth (1967), Pearce & Smith (1990), and Anonymous (1993).
RESULTS
The emergence rate of adult A. albipodus from mummies was significantly lower (P < 0.001) under the 12 h daylength relative to the 14 h daylength (Fig. 1). Less than % the proportion of A. albipodus adults emerging from mummies under the 14 h daylength emerged under the 12 h daylength. In contrast, emer- gence rates of D. rapae adults were similarly high and not significantly different (P = 0.439) under both daylengths (Fig. 1).
None of 121 mummies that were maintained at 12 h daylength for up to 84 days from the egg stage yielded A. albipodus adults. Dissection of these mummies revealed that they contained live last-instar larvae throughout this sampling pe- riod; 98% contained live last-instar larvae at 48 days from the egg stage (n = 65 mummies), 89% at 55 days (n = 28 mummies), and 93% at 80 days (n = 28 mummies). In contrast, ca. 12% of 121 mummies transferred from 12 h daylength to 14 h daylength at 28 days from the egg stage yielded A. albipodus adults within 47—75 days.
Dissection of five of the mummies transferred to 14 h daylength at each of 48, 55, and 80 days from the egg stage indicated that the remaining mummies con- tained living last-instar larvae (15/15 total mummies dissected contained live last- instar larvae). Although additional adult emergence from the mummies transferred to 14 h daylength was plausible given the prevalence of live last instar larvae inside these mummies after 80 days, these were accidentally destroyed at 84 days when a mechanical failure caused a sudden increase in the ambient temperature to >40° C for ca. 20 h.
DISCUSSION
Our results showed that A. albipodus entered diapause when it developed under a 12 h daylength. The incidence of diapause in A. albipodus at this daylength, discounting for mummy stage mortality evident under the 14 h daylength (ca. 8%), was ca. 68%, and it occurred in the last larval instar. In addition, our results showed that diapause could be broken in some A. albipodus individuals by ex- posing diapausing mummies to a 14 h daylength. In contrast to A. albipodus, D. rapae did not enter diapause when it developed under a 12 h daylength.
Our findings concerning A. albipodus were not unexpected, whereas those con- cerning D. rapae were initially puzzling given the climatic conditions prevalent in their areas of origin (Fig. 2). Mean monthly temperatures at Tahcheng, A. albipodus’ collection site, vary from —16° C in January to 22° C in July, and daylengths vary from 8.5 h in December to 15.9 h in June. Temperature conditions in Wuqia, D. rapae’s collection site, are less extreme than in Tahcheng, ranging
32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
—_ =) i=)
P << 0.001
Proportion Emerged (+ S.E.) o i) ol
= o o
A. albipodus D. rapae Parasitoid
Temperature (°C)
= , om ou
-15 ahcheng, PRC
8 9 10 11 12 13 14 15 16 17 Daylength (hours)
Figure 1. Emergence rates of Aphelinus albipodus and Diaeretiella rapae under two daylengths (21° C, 50-70% R.H.); hollow columns = 14 h daylength, stippled columns = 12 h daylength. Numbers above column pairs indicate significance of difference in emergence rates between 14 h and 12 h daylengths: A. albipodus, G = 306.470, 1 df, n = 685; D. rapae, G = 0.666, 1 df, n = 719.
Figure 2. Daylength (on the 15th of each month) and mean monthly temperature variation in: Tahcheng, People’s Republic of China, A phelinus albipodus’ collection site; Wugia, PRC, Diaeretiella rapae’s collection site, and; Tulelake, California, a representative northern California colonization site for A. albipodus and D. rapae.
2001 BERNAL ET AL.: PARASITOID OVERWINTERING 33
from —11° C in January to 13.5° C in July, while daylengths are similar, 9.4 h in December to 14.9 h in June. Despite differences in temperature, winter conditions in both Tahcheng and Wuqia would require a mechanism by which A. albipodus and D. rapae could survive during 5 months of subfreezing mean monthly tem- peratures.
Diapause provides a mechanism for winter survival in A. albipodus. Mean monthly temperatures and average daylengths in Tahcheng closely match tem- perature- and daylength-related developmental restrictions in A. albipodus that are evident from this and other studies (Bernal & Gonzdélez 1996; Bernal et al. 1997). Those studies showed that A. albipodus: (1) enters diapause at 12 h daylengths, and; (4i) cannot develop to adulthood if temperatures are 10° C or below. Aphel- inus albipodus mortality at 10° C is >80% during egg to mummy development, and adults do not emerge from mummies that develop at this temperature (Bernal & Gonzalez 1996; Bernal et al. 1997). Mean temperatures in Tahcheng decrease to 10° C and below in the fall, and are associated with 12 h and shorter daylengths. Mean temperatures then increase to 10° C and above in the spring and are asso- ciated with 14 h and longer daylengths. Thus, A. albipodus may avoid fatally low temperatures by entering diapause when daylengths decrease to 12 h in the fall, then resuming development when daylengths increase to 14 h in the spring.
In contrast to A. albipodus, diapause was not detected in D. rapae at 12 h daylength. Based on our results, however, we cannot discount the possibility of diapause in this parasitoid. Diaeretiella rapae populations in the Netherlands enter diapause during the winter (Hafez 1961). Moreover, diapause in these D. rapae populations peaks in late October (Hafez 1961) when daylengths are between 10.7 and 8.8 h. Thus, diapause may be induced at daylengths shorter than 12 h in the Wuqia D. rapae population. Average daylengths in Wugqia decrease to 12 h in the early fall and are associated with mean temperatures of ca. 7° C. Later in the fall, mean temperatures decrease to subfreezing levels, but daylengths by then decrease to 10.5 h and shorter. Previous studies show that developmental mortality in D. rapae is low at 10° C, and that its lower developmental threshold is in the range 2.5—3.9° C (Bernal & Gonzalez 1995, 1997). Thus, diapause at 12 h daylength in the Wuqia D. rapae population may not be necessary given the associated mean field temperatures (ca. 7 C) in the area. Hence, based on the results of previous studies (Hafez 1961; Bernal & Gonzdlez 1995, 1997) and on the lengthy and severe winters prevalent in Wugia, a likely scenario is that dia- pause in D. rapae is induced by daylengths shorter than 12 h, which are associated with near- or sub-freezing temperatures.
Our results concerning A. albipodus are consistent with previous reports of diapause in closely related Aphelinus spp. Yu (1992) found that >50% and >95% of A. nr. varipes (= A. varipes, see Bernal et al. 1997; J. B. Woolley, personal communication) collected in southern Alberta (Canada) entered diapause when they developed under 14 h and 12 h daylengths, respectively. The corresponding rates for A. varipes collected in Kazakhstan (= A. albipodus, see Bernal et al. 1997; J. B. Woolley, personnel communication) were 0% and >90%. Diapause was induced by short daylengths both in Yu’s and this study. Short daylengths in these cases are associated with low winter temperatures that may be particularly detrimental to Aphelinus species. Developmental thresholds and high mortality at temperatures between 5 and 10° C are common in Aphelinus spp. (e.g., Force &
34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Messenger 1964; Walker et al. 1988; Trimble et al. 1990; Asante & Danthanarayan 1992; Lajeunesse & Johnson 1992; Yu 1992; Bernal & Gonzdélez 1993, 1996; Bernal et al. 1997; Lee & Elliot 1998a, b). Thus, Aphelinus spp. appear to be highly susceptible to low temperatures and to employ diapause as an overwinter- ing strategy (Trimble et al. 1990; Yu 1992; this study).
Both A. albipodus and D. rapae were imported to Califomia for release against Russian wheat aphid (Gonzalez et al., unpublished data). Both were extensively released in northern California and A. albipodus rapidly became established. It is unclear at present whether D. rapae became established because this species al- ready occurred in the area. In either case, it is evident that the climate in northern California is milder during the winter relative to the collection areas of both parasitoids, and daylength variation during the year is similar between these areas, or less in the case of D. rapae (Fig. 2). Thus, although diapause may be induced by short winter daylengths in northern Califomia, it may not be required for winter survival of A. albipodus and D. rapae. Summer, rather than winter, temperatures may represent an obstacle for establishment of D. rapae, in particular, in northern California. Earlier studies showed that D. rapae is more susceptible to high tem- peratures than A. albipodus (Bernal & Gonzdlez 1995, 1996). This is suggested also by the seasonal variation in mean temperatures in each of these parasitoid’s collection sites. Thus, winter diapause and summer survival are two important factors that may affect the population dynamics of A. albipodus and D. rapae and their impact on Russian wheat aphid populations in California. Moreover, the timing of emergence of diapausing parasitoids in the spring will likely influence their impact on Russian wheat aphid populations. Previously it was suggested that in annual agroecosystems, early-appearing natural enemies are likely to have a greater impact on pest populations than those emerging later in the season (Bernal & Gonzalez 1993; Gilstrap 1997). Our results, and previous studies (Bernal & Gonzalez 1995, 1996), suggest that A. albipodus is likely to emerge later in the season than D. rapae.
Differences in climate between collection and colonization areas of exotic nat- ural areas are reported to contribute substantially to reducing the success rate of classical biological control (Stiling 1993). Specifically, they hamper biological control efforts early on in their development by precluding the successful estab- lishment of exotic natural enemies. A practice that could contribute substantially to reducing the number of biological control efforts that fail during the coloni- zation phase is to emphasize closer climate-matching during the planning phase of foreign exploration efforts. An approach similar to that taken to forecast po- tential range expansions of pest species (Meats 1989; Hughes & Maywald 1990) would be desirable, but may seldom be possible with exotic natural enemies. Unlike pest species, usually little or no information is available concerning an exotic natural enemy’s response to temperature and other biological parameters necessary for such an approach. Many times exotic natural enemies are undes- cribed species and biological information can only be inferred from related spe- cies; other times, the available information is limited to a taxonomic description. In both cases, pertinent biological information becomes available only after a series of studies are conducted. We believe that the analyses presented here, in- cluding the results from previous studies (Bernal & Gonzalez 1995, 1996, 1997; Bernal et al. 1997), provide an initial model for assessing the potential for suc-
2001 BERNAL ET AL.: PARASITOID OVERWINTERING 35
cessful establishment of exotic natural enemies early in the development of a biological control campaign, and a means for improving the success rate of clas- sical biological control. Ultimately, the success of a classical biological control
campaign is contingent upon the successful establishment of exotic natural ene- mies.
ACKNOWLEDGMENT
We are grateful to Dr. Bob Wharton and Janet Griset (both at Texas A&M University) for critically reviewing the manuscript and for suggestions for its improvement. Marcella Waggoner and her team provided the aphids and parasit- oids used in the experiments. Our Chinese collaborators, G. Zhang, J. Zhang, W. Zhang, Y. Min, R. Wang, Du Bing Ren, Wei Zhengming, Geng Shouguang, J. Wen, X. Liu, and Y. Zhong are gratefully acknowledged for their skilled assistance during foreign exploration in China.
LITERATURE CITED
Anonymous. 1993. Daily average temperatures and precipitation in California. Statewide IPM Project, University of California, Davis.
Asante, S. K. & W. Danthanarayana. 1992. Development of Aphelinus mali an endoparasitoid of wooly apple aphid Eviosoma lanigerum at different temperatures. Entomol. Exp. Appl., 65: 31-37.
Beddington, J. R., C. A. Free & J. H. Lawton. 1978. Characteristics of successful natural enemies in models of biological control of insect pests. Nature, 273: 513-519.
Bernal, J. S. & D. Gonzalez. 1993. Temperature requirements of four parasites of the Russian wheat aphid Diuraphis noxia. Entomol. Exp. Appl., 69: 173-182.
Bernal, J. S. & D. Gonzalez. 1995. Thermal requirements of Diaeretiella rapae (M’Intosh) on Russian wheat aphid (Diuraphis noxia Mordwilko, Hom., Aphididae) hosts. J. Appl. Ent., 119: 273- 277.
Bernal, J. S. & D. Gonzalez. 1996. Thermal requirements of Aphelinus albipodus (Hayat and Fatima) (Hym., Aphelinidae) on Diuraphis noxia (Mordwilko) (Hom., Aphididae) hosts. J. Appl. Ent., 120: 631-638.
Bernal, J. S. & D. Gonzalez. 1997. Reproduction of Diaeretiella rapae on Russian wheat aphid hosts at different temperatures. Entomol. Exp. Appl., 82: 159-166.
Bernal, J. S, M. Waggoner & D. Gonzalez. 1997. Reproduction of Aphelinus albipodus (Hyme- noptera: Aphelinidae) on Russian wheat aphid (Hemiptera: Aphididae) hosts. Eur. J. Entomol., 94: 83-96.
DeBach, P. & D. Rosen. 1991. Biological control by natural enemies (2nd ed.). Cambridge University Press, Cambridge.
Flanders, S. E. 1940. Environmental resistance to the establishment of parasitic Hymenoptera. Ann. Entomol. Soc. Amer., 33: 245—253.
Force, D. C. & P. S. Messenger. 1964. Duration of development, generation time, and longevity of three hymenopterous parasites of Therioaphis maculata reared at various constant temperatures. Ann. Entomol. Soc. Amer., 57: 405—413.
Gilstrap, E E. 1997. Importation biological control in ephemeral crop habitats. Biol. Control, 10: 23-29.
Gilstrap, EF E., D. Gonzalez & L. McKinnon. 1994. Biological control of Russian wheat aphid: a summary of 1998-1993. pp. 53-61. In Peairs, E B., M. K. Kroening, & C. L. Simmon (comps.). Proceedings of the sixth Russian wheat aphid workshop. Fort Collins, January 23-25, 1994. Great Plains Agricultural Council, Fort Collins.
Gonzalez, D. & E E. Gilstrap. 1992. Foreign exploration: assessing and prioritizing natural enemies and consequences of preintroduction studies. pp. 53-70. Jn Kauffman, W. C. & J. E. Nechols (eds.). Selection criteria and ecological consequences of importing natural enemies. Entomol. Soc. Amer., Lanham.
Greathead, D. J. & A. H. Greathead. 1992. Biological control of insect pests by insect parasitoids and predators: the BIOCAT database. Biocontrol News Info., 13: 61N—68N.
36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Hafez, M. 1961. Seasonal fluctuations of population density of the cabbage aphid, Brevicoryne bras- sicae (L.) in The Netherlands, and the role of its parasite, Aphidius (Diaeretiella) rapae (Curtis). T. Pl.-ziekten, 67: 445-548.
Huffaker, C. B., EF J. Simmonds & J. E. Laing. 1974. The theoretical and empirical basis of biological control. pp. 41-78. Jn Huffaker, C. B. & P. S. Messenger (eds.). Theory and practice of bio- logical control. Academic Press, New York.
Huffaker, C. B., R. EK Luck & P. S. Messenger. 1977. The ecological basis of biological control. pp. 560-586. In White, D. (ed.). Proceedings, XV International Congress of Entomology. Wash- ington, D. C., 19-27 August. Entomol. Soc. Amer., College Park.
Hughes, R. D. & G. EK Maywald. 1990. Forecasting the favourableness of the Australian environment for the Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae), and its potential impact on Australian wheat yields. Bull. Entomol. Res., 80: 165-175.
Lajeunesse, S. E. & G. D. Johnson. 1992. Developmental time and host selection by the aphid para- sitoid Aphelinus sp. nr. varipes (Foerster) (Hymenoptera: Aphelinidae). Can. Ent., 124: 565- 575.
Lee, J. H. & N. C. Elliot. 1998a. Comparison of developmental responses to temperature in Aphelinus asychis (Walker) from two different geographic regions. Southwest. Entomol., 23: 77-82.
Lee, J. H. & N. C. Elliot. 1998b. Temperature effects on development in Aphelinus albipodus (Hy- menoptera: Aphelinidae) from two geographic regions. Great Lakes Entomol., 31: 173-179.
Luck, R. F 1990. Evaluation of natural enemies for biological control: a behavioral approach. Trends Ecol. Evol., 5: 196-199.
Meats, A. 1989. Bioclimatic potential. pp. 241-252. In Robinson A. S. & G. Hooper (eds.). Fruit flies: their biology, natural enemies and control. Volume 3B. Elsevier, Amsterdam.
Messenger, P. S. 1959. Bioclimatic studies with insects. Ann. Rev. Entomol., 4: 183-206.
Messenger, P. S. 1971. Climatic limitations to biological control. Proc. Tall Timbers Conf. Ecol. Animal Control Habitat Mgmt., 3: 97-114.
Messenger, P. S. & N. E. Flitters. 1954. Bioclimatic studies of three species of fruit flies in Hawaii. J. Econ. Entomol., 47: 756-765.
Murdoch, W. W. & C. J. Briggs. 1996. Theory for biological control: recent developments. Ecology, 77: 2001-2013.
Murdoch, W. W., J. Chesson & P. L. Chesson. 1985. Biological control in theory and practice. Am. Nat., 125: 344-366.
Pearce, E. A. & G. Smith. 1990. The Times books world weather guide. Times Books-Random House, New York.
Stiling, P. 1993. Why do natural enemies fail in classical biological control programs? Amer. Entomol., 39: 31-37.
Trimble, R. M., L. R. H. Blommers & H. H. M. Helsen. 1990. Diapause termination and thermal requirements for postdiapause development in Aphelinus mali at constant and fluctuating tem- peratures. Entomol. Exp. Appl., 56: 61-69.
Walker, J. T. S., S. C. Hoyt, D. P. Carroll & G. V. Tangren. 1988. Influence of constant and alternating temperatures on wooly apple aphid (Homoptera: Eriosomatidae) and its parasitoid Aphelinus mali (Haldeman) (Hymenoptera: Aphelinidae). Melanderia, 46: 36-42.
Walter, H. & H. Lieth. 1967. Klimadiagram Weltatlas. VEB Gustav Fischer Verlag, Stuttgart.
Yu, D. S. 1992. Effects of photoperiod and temperature on diapause of two Aphelinus spp. (Hyme- noptera: Aphelinidae) parasitizing the Russian wheat aphid. Can. Ent., 124: 853-860.
Zar, J. H. 1996. Biostatistical analysis. Prentice Hall, Englewood Cliffs.
Received 8 Feb 2000; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 37-38, (2001)
A NEW SPECIES OF CORTICARINA FROM ARIZONA (LATRIDITDAE: CORTICARIIND)
FRED G. ANDREWS
California Department of Food and Agriculture, 3294 Meadowview Road, Sacramento, California 95832-1448
Abstract.—Corticarina arizonensis is described as new. It is compared to other North American species.
Key Words.—Insecta, Coleoptera, Latridiidae, Corticarina, Arizona.
In 1994 two specimens of Corticarina were swept from an oak tree at the base of a talus slope at approximately 9200 feet in the Barfoot campground in the Chiricahua mountains of Southeastern Arizona. Dissection of the lone male dem- onstrated a new species based on the unknown aedeagal shape. Recent collecting (1998) in the same spot yielded an additional twenty-one specimens. The same aedeagal shape was found when males from this collection were dissected. All specimens were collected within 100 feet of the original collection, but this time all were swept from conifers. Additional specimens were collected on Mt. Lem- mon, Arizona at an elevation of 7500 feet from the flower heads of dead cow parsnip (Heracleum lanatum Michaux) in the company of numerous Cortinicara gibbosa Gyllenhal. Numerous collections of Corticarina cavicollis Mannerheim and Corticarina fuscula Gyllenhal have been made in the Chiricahua Mountains and Southeastern Arizona at lower elevations without evidence of this new spe- cies. It is likely that this species is restricted to the higher mountain elevations. Corticarina arizonensis can be differentiated from all other species of Corticarina by the shape of the male aedeagus (Figs. 1 and 2). Corticarina arizonensis is the only North American species in which both the dorsal and ventral lobes of the aedeagus are sharply pointed. The aedeagus of the other North American Corti- carina are illustrated by Andrews (1985, 1992).
CORTICARINA ARIZONENSIS Andrews, NEW SPECIES
Description.—Length 1.33—-1.59 mm. Width 0.63—0.74 mm. Pronotal length 0.30—0.37 mm. Pronotal width 0.37—-0.44 mm. n = 43. Color dark brown to dark reddish brown; legs, antennal segments 2—6 lighter brown; surface shiny. Anten- nae 0.45—0.51 mm long, segment 8 longer than wide, segment 10 wider than long. Pronotum 1.11 to 1.47 times as wide as long; lateral margin slightly sigmoid in anterior one-half, widest in anterior one-half; postmedian depression weakly de- veloped; lateral depression narrow, weakly developed, limited to posterior one- half; dorsally evenly rounded from side to side; lateral margins smoothly arcuate, minutely serrate. Pronotal surface distinctly punctured, punctures generally sep- arated by approximately a puncture width. Elytra narrowly inflated, 1.13 to 1.75 times as long as wide. Elytral striae well defined; setae arranged in straight lines, each raised medially and generally terminating in next puncture; humeral callus distinct. Fully winged. Eyes fully developed, 77 facets in single eye examined. Male: protibia toothed on inner surface, tooth 45/100 from apex. Male genitalia as in Figs. 1 and 2.
38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Figures 1-2. Corticarina arizonensis aedeagus. Figure 1. Ventral view. Figure 2. Lateral view.
Types.—HOLOTYPE (Male): ARIZONA. Pima Co: Santa Catalina Mts., Mt. Lemon, Summerhaven, [X-17-1998, 32°26.26' N, 110°45.52' W, F Andrews & T. Eichlin. Deposited in the collection of the Califomia Academy of Sciences. PAR- ATYPES (23). Same data as Holotype (11); ARIZONA. Cochise Co. Barfoot Park, VIII-2-1994, EF Andrews & T. Eichlin (2); 10.6 mi. NW Southwest Research Station, Barfoot Cpg., [X-14-1998, 31°55.11' N, 109°16.47' W, E Andrews & T. Eichlin (10).
LITERATURE CITED
Andrews, E G. 1985. Corticarina scissa (LeConte), a valid species (Coleoptera: Lathridiidae). Coleop. Bull. 39: 147-149.
Andrews, E G. 1992. Two new species of Corticarina from coastal California (Coleoptera: Lathridi- idae: Corticariini) with notes on J. L. LeConte types. the Coleop. Bull., 46: 274-280.
Received 3 Feb 2000; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 39-44, (2001)
MELANOTRICHUS BOYDI, A NEW SPECIES OF PLANT BUG (HETEROPTERA: MIRIDAE: ORTHOTYLIND RESTRICTED TO THE NICKEL HYPERACCUMULATOR STREPTANTHUS POLYGALOIDES (BRASSICACEAE)
MICHAEL D. SCHWARTZ! AND MICHAEL A. WALL?
'% Biological Resources Program, ECORC, Agriculture and Agri-Food Canada, Ottawa, Ontario K1A OC6 Canada Department of Botany and Microbiology, Auburn University, 101 Rouse Life Sciences Building, Auburn, Alabama 36849-5407*
Abstract——Melanotrichus boydi, NEW SPECIES is described from the western foothills of the Sierra Nevada Mountains in California. The nickel hyperaccumulating species, Streptanthus po- lygaloides (Gray), Brassicaceae, an endemic to serpentine soils, is its only known host plant.
Key Words.—Insecta, Miridae, plant bugs, Melanotrichus, new species, nickel hyperaccumula- tors, serpentine soil.
As a consequence of its size, exceedingly diverse topography, climate, and geologic history, California supports a remarkably large and interesting flora (Munz & Keck 1973). Despite the pioneering work of E. P. Van Duzee and recent students of North American Muiridae, the plant bug fauna of California is still poorly known. Current studies by the junior author in serpentine soil exposures of California have revealed a new species of plant bug restricted to a nickel hyperaccumulator species of crucifer. In this paper we describe Melanotrichus boydi Schwartz & Wall, NEW SPECIES, provide illustrations of the male geni- talia, photomicrographs of the head, pretarsus, ostiolar peritreme, scalelike setae, a distribution map and discuss the first known example of bioaccumulation for a native herbivore on a metal hyperaccumulator. All measurements are based on ten specimens with the mean and range presented.
MELANOTRICHUS BOYDI Schwartz & Wall, New Species (Figs. 1-14)
Types.—Holotype, male: U.S.A. CALIFORNIA. ELDORADO Co.: NE of Col- oma, [900 m], 22 Jun 1998, M. A. Wall, ex Streptanthus poly[galoides|., M. Wall 98-105; deposited: California Academy of Sciences, San Francisco (CAS). Par- atypes: 1d, 12 same data as holotype except no host and M. Wall 1; MARIPOSA Co.: 16, 12 NW of Coulterville, [700 m], 23 Jun 1998, M. Wall (2); 36, 59 Lake McClure at hwy 49 [bridge], [900 m], 23 Jun 1998, M. Wall (9); 13, 52 NW of Mariposa, 29 Jun 1998, M. Wall (7); PLACER Co.: 16, 3¢ S of Sugar Pine Res[ervoir], [1270 m] 28 Jun 1998, M. A. Wall; TUVOLUMNE Co.: 436,42 S of Mocassin on hwy 49, [700 m], 11 Jun 1999, M. A. Wall; 4¢, 92 SW of Chinese Camp, Red Hills Rec Area, [470 m], 28 May 1999, M. A. Wall; 12 Chinese Camp, [470 m], 10 Jun 1997, M. A. Wall; 13¢ S of Chinese Camp in Red Hills Rec Area, [470 m], 9-13 Jun 1996, M. A. Wall. Paratypes deposited
* Currently Department of Ecology and Evolutionary Biology, University of Connecticut, Torrey Life Science Building, 75 North Eagleville Road, U-43, Storrs, Connecticut 06269-3043.
40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Figures 1-4. Scanning electron micrographs of Melanotrichus boydi. Figure 1. Head, lateral view. Figure 2. Ostiolar peritreme, lateral view. Figure 3. Scalelike setae on corium. Figure 4. Pretarsus, apical view.
at the American Museum of Natural History, New York, CAS, the Canadian National Collection of Insects, Ottawa and the National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C.
Description of Males.—Orthotylinae: Orthotylini. Length from clypeus to apex of membrane 5.74 (5.20—6.15) mm, maximum width across hemelytra 1.88 (1.60—2.10) mm. Head: width 0.93 (0.88— 0.99) mm, vertex width 0.39 (0.36—0.41) mm. Labium: length 1.14 (1.08—-1.23) mm, reaching apex of mesosternum, sometimes just extending to slightly beyond middle of mesosternum, but labrum usually reaching apex of mesosternum. Antenna: segment 1, length, 0.55 (0.50—0.60) mm; 2, 1.86 (1.73-2.15) mm; 3, 1.60 (1.45—1.83) mm; 4, 0.47 (0.45—0.49) mm. Pronotum: length 0.70 (0.63—0.79) mm, basal width 1.34 (1.23—-1.45) mm.
Generally green with greenish-yellow to orange-yellow on embolium, cuneus, mesoscutum, some- times scutellum, pronotum on calli and anterior of calli, head, antennal segment 1, extreme base of antennal segment 2, legs, and venter; black on antennal segments 2—4, apex of labial segment 4, tarsus, pretarsus; membrane of hemelytra infuscate with greenish-yellow veins; dorsum with inter-
2001 SCHWARTZ & WALL: NEW PLANT BUG SPECIES 41
Figures 5-13. Male genitalia of Melanotrichus boydi. Figures 5—7. Left paramere. Figure 5. Apex of shaft, apical view. Figure 6. Sensory lobe, apical view. Figure 7. Sensory lobe, lateral view. Figures 8 and 9. Right paramere. Figure 8. Dorsal view. Figure 9. Lateral view. Figure 10. Phallotheca, dorsal view. Figure 11. Detail of ductus seminis & secondary gonopore, lateral view. Figure 12. Detail of spiculum, lateral view. Figure 13. Ventral processes of genital segment, apical view.
mixed vestiture of moderately distributed, shining white, reclining simple setae and head, anterior of pronotum and hemelytra with scattered shining, silvery, sericeous scalelike, setae (Fig. 3).
Head broadly subtriangular; clypeus moderately produced and slightly rounded in dorsal aspect, anteriorly flattened with slightly rounded ventral margin in lateral aspect (Fig. 1); posterior margin of head wider than, and slightly removed from, anterior margin of pronotum; basal carina broad, gently rounded; eyes large, height greater than head height in lateral aspect; posterior margin gently curved anteriorly in dorsal aspect; vertex width equal to 1.5 width of eye. Antennal segment 1 with mod- erately distributed, reclining, black simple setae and two subapical black bristles; diameter about 1.5 as wide as segment 2; segment 3 & 4 slightly thinner than segment 2. Pronotum trapeziform, calli slightly swollen. Mesoscutum moderately broadly exposed. Hemelytra subparallel-sided, widest at apex of embolium; length of cuneus about 3X width. Venter unmarked; ostiolar peritreme (Fig. 2). Legs long; tibia with dark brown to black bristles, unmarked at base; claw (Fig. 4) gently curved with minute pulvillus and apically converging parempodium.
Genitalia: Left paramere (Figs. 5-7) broad, C-shaped, sensory lobe with truncate, broadly serrate apex; shaft broadly curved, gradually attenuate. Right paramere (Figs. 8 and 9) L-shaped, broad basally, strongly attenuate distally. Phallotheca (Fig. 10) strongly sclerotized, convoluted, aperture on right aspect. Vesica with narrow elongate ductus seminis (Fig. 11) and one basally thickened, distally attenuate spiculum (Fig. 12); apex of spiculum subequal to apex of secondary gonopore (Figs. 11 and 12). Ventral processes of genital segment (Fig. 13); right process larger than left process, strongly sclerotized with flattened dorsal flange, which protrudes posteriorly beyond margin of genital aperture.
Description of Females.—Similar to males except, eye smaller, head and vertex wider, and heme- lytral margin more rounded, length from clypeus to apex of membrane 5.86 (5.20—6.50) mm, maxi- mum width across hemelytra 2.01 (1.80—2.29) mm. Head: width 1.01 (0.98-1.15) mm, vertex width 0.49 (0.45-0.53) mm. Labium: length 1.26 (1.20—1.38) mm, reaching apex of mesosternum, sometimes just extending to slightly beyond middle of mesosternum, but labrum usually reaching apex of me-
42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Nevada
El Dorado
Madera
Figure 14. Distribution of Melanotrichus boydi. Areas filled in black represent distribution of serpentine soil within the foothills of the Sierra Nevada. M. boydi was either observed or collected at all numbered sites. Site numbers correspond with information presented in Table 1 (modified from Wall 1999).
sosternum. Antenna: segment 1, length, 0.58 (0.53—0.64) mm; 2, 1.85 (1.66—2.06) mm; 3, 1.49 (1.33- 1.58) mm; 4, 0.46 (0.45—0.48) mm. Pronotum: length 0.81 (0.71-0.91) mm, basal width 1.58 (1.35-— 1.75) mm.
Diagnosis.—Melanotrichus boydi NEW SPECIES is most similar to M. mistus (Knight) and M. stanleyaea (Knight) in size (almost all other species of Melan- otrichus are less than 5 mm in length) but is distinguished by the white simple setae on the dorsum, the black antennal segments 2—4, and the structure of male genitalia, especially the narrow apex of the right paramere and pointed vesical spiculum. Melanotrichus mistus has conspicuous black setae on the dorsum, yel- low to orange antenna, and male genitalia with the apices of the right paramere and vesical spiculum blunt. Melanotrichus stanleyaea has similar pale vestiture to M. boydi but the antenna is pale, and the apex of the right paramere is longer and more acutely attenuated.
Taxonomy.—Schuh (1995) and Kerzhner and Josifov (1999) treat Melanotri- chus Reuter as a subgenus of Orthotylus Fieber. However, given the absence of a phylogenetic analysis of the genus we choose to follow North American authors and describe our new species in the genus Melanotrichus. In Henry (1991: 455) M. boydi will key to M. concolor (Kirschbaum). In Knight (1968) this species will key to M. stanleyaea Knight, under Dichaetocoris Knight (cf., Knight 1968:
2001 SCHWARTZ & WALL: NEW PLANT BUG SPECIES 43
Table 1. Concentration of Ni found in sample of M. boydi collected at six sites throughout Cali- fornia (see Fig. 14). Concentrations represent values obtained from samples consisting of 15—20 in- dividuals of M. boydi from each site analyzed via atomic absorption spectrophotometry.
Site Ni content in ppm on location dry weight basis Site 1 735 Site 3 ail Site 4 789 Site 6 777 Site 7 718 Site 10 751
114) or as M. wileyae Knight (cf. Knight 1968: 117). The black antennal segments 2—4, distribution, and host plant are sufficient to distinguish M. boydi from these species.
Hostplants.—Streptanthus polygaloides (Gray), Brassicaceae, a winter annual, endemic to serpentine soils from the foothills of the western slopes of the Sierra Nevada Mountains in California is the host of both immature and adult stages. Wall and Boyd (in press) provided evidence to indicate that M. boydi is probably monophagic. At ten localities where this species was collected on S. polygaloides they sampled other locally abundant plants—a conifer, three woody dicots, four herbaceous dicots, including other species of Brassicaceae and another species of Streptanthus, and a monocot—for the presence of M. boydi and did not collect any.
Distribution.—Figure 14. In addition to the localities listed under Types, M. boydi was also collected from the following sites in late-May to mid-July: BUTTE Co.: N of Magalia, 800 m; CALAVERAS Co.: N of San Andreas, 300 m; FRESNO Co.: NW of Pine Flat Lake, 400 m; PLACER Co.: S of Washington, 1330 m. In the western foothills of the Sierra Nevada M. boydi is associated throughout the range of S. polygaloides from Butte County in the north to Fresno County in the south encompassing a wide range of elevation (330 m to 1330 m) within the foothills woodland and the yellow pine forest plant communities.
Etymology—Named to honor Dr. R. S. Boyd, Department of Botany and Mi- crobiology, Auburn University, who provided insight and encouragement to the junior author during his Master’s degree, and for his leadership in the study of hyperaccumulation ecology.
Discussion.—Melantrichus boydi is unique in several ways. Not only is it the only species of insect reported to specialize on a Ni hyperaccumulator, but it appears to accumulate Ni at levels one to two orders of magnitude higher than other insects found feeding on S. polygaloides (Wall 1999). This high level of Ni accumulation is concordant across the known range of M. boydi (Table 1). Bot- anists have long used a qualitative colorimetric test for identifying plants in the field that contain high levels of Ni. In this test, plant material is crushed onto filter paper permeated with the colorless chemical, dimethylglioxime, which reacts with the Ni in the plant and changes to a various shades of red (Reeves 1992). This same test consistently gives a positive result when M. boydi is crushed onto the filter paper. While perhaps only having novelty status, this colorimetric test adds another interesting element to the taxonomist’s arsenal for identifying this
44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
species. At the very least, the presence of high levels of Ni in M. boydi is con- sidered an autapomorphy for the species.
Including M. boydi n. sp., five North American mirids apparently specialize on brassicaceous hosts, all are members of the genus Melanotrichus. Melanotrichus albocostatus Knight is known from Cardaria costatus, Descurainia sophia (L.) Webb, and Sisymbrium irio L. Webb (Henry 1991), M. leonardi Kerzhner and Schuh is known from D. sophia (Kelton 1980), M. stanleyaea is known from Stanleya pinnata (Pursh) Britton (Knight 1968), and M. whiteheadi Henry is known from D. pinnata pinnata (Walt.) Britt. (Henry 1991). Interestingly, S. pin- nata is a hyperaccumulator of Selenium (Brooks 1998). Determining whether or not host preference for Brassicaceae has evolved in multiple lineages, or if these five species represent a monophyletic group within Melanotrichus requires a de- tailed phylogenetic analysis, which is beyond the scope of this present study.
ACKNOWLEDGMENT
The Pacific and Agri-Food Research Centre, Agassiz, British Columbia pro- vided support for MDS to study type specimens of E. P. Van Duzee, housed in the CAS. Support for this study was also provided by the Alabama Academy of Sciences to Michael Wall and by NSERC and FRBC grants to G. G. E. Scudder, Department of Zoology, The University of British Columbia, Vancouver, Canada. Two anonymous reviewers made critical comments on an earlier version of the manuscript.
LITERATURE CITED
Brooks, R. R. 1998. Geobotany and hyperaccumulators. Chapter 3. pp. 55—94. In Brooks, R. R. (ed.). Plants that Hyperaccumulate Heavy Metals. CAB International, Wallingford.
Henry, T. J. 1991. Melanotrichus whiteheadi, a new crucifer-feeding plant bug from the southeastern United States, with new records for the genus and Key to the species of eastern North America (Heteroptera: Miridae: Orthotylinae). Proc. Entomol. Soc. Wash., 93: 449-456.
Kelton, L. A. 1980. Two new species of Melanotrichus Reuter from western Canada and a description of the male of M. atriplicis (Heteroptera: Miridae). Can. Entomol. 112: 337-339.
Kerzhner, I. M. & M. Josifov. 1999. Cimicomorpha IT, Miridae. Volume 3. Jn Aukema, B & C. Rieger (eds.). Catalogue of the Heteroptera of the Palaearctic. The Netherlands Entomological Society, Wageningen.
Knight, H. H. 1968. Taxonomic review: Mirdae of the Nevada Test Site and the western United States. Brigham Young Univ. Sci. Bull., Biol. Ser., 9: 1-282.
Munz, P. A. & D. D. Keck. 1973. A California flora and supplement. University of California Press, Berkeley.
Schuh, R. T. 1995. Plant bugs of the world (Insecta: Heteroptera: Miridae). New York Entomol. Soc., New York.
Reeves, R. D. 1992. The hyperaccumulation of nickel by serpentine plants. pp. 253-277. In A. J. M. Baker, J. Proctor, and R. D. Reeves (eds.). The Ecology of Ultramafic (serpentine) Soils. Inter- cept, Andover.
Wall, M. A. 1999. Nickel accumulation in serpentine arthropods with emphasis on a species of Me- lanotrichus (Heteroptera: Miridae). M.S. Thesis, Auburn University, Alabama.
Received 29 Mar 2000; Accepted 5 May 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 45-46, (2001)
Scientific Note
ALTITUDINAL DISTRIBUTION AND PHENOLOGY OF THREE SPECIES OF CARRION BEETLES (COLEOPTERA: SILPHIDAE) FROM NEVADO DE COLIMA, JALISCO, MEXICO
The taxonomy of carrion beetles (Coleoptera: Silphidae) from México was revised by Peck, S. B. & R. S. Anderson (1985. Quaest. Ent., 21(3): 247-317) who recorded eleven species belonging to four genera. Other contributions, es- pecially for Jalisco State include: Volcan de Tequila (Navarrete-Heredia, J. L. 1995. Dugesiana, 2(2): 11—26); Sierra de Manantlan (Rivera-Cervantes, L. E. & E. Garcia-Real. 1998. Dugesiana, 5(1): 11-22), and La Primavera and Barranca del Rio Santiago (Navarrete-Heredia, J. L. & H. E. Fierros-Lépez. 1998. Duge- siana, 5(1): 49-50). This study was done in Jalisco by staff members from the Center for Zoological Research, University of Guadalajara to determine carrion beetle distribution. In this note, we describe our trapping results from the National Park Nevado de Colima, Jalisco, México.
Field work was done on the NW slope of the National Park Nevado de Colima, in the locality El Floripondio, Cerro Las Viboras, San Gabriel County, between 2200—3000 meters above sea level (m). We used carrion traps (model NTP-80) designed by Moré6én, M. A. & R. Terrén [1984. Acta Zool. Mex. n.s., (3): 1-47]. Six sites were selected: Cupressus forest (2300 m), oak-pine forest (2400 m), cloud forest (2600 m), Abies forest (2840 m), disturbed Abies forest (2920 m), and Abies-grass association (2950 m). One carrion trap was used for each site during one month, starting in April and ending in October 1998. Rotting squid was used as bait.
Three species of carrion beetles were collected: Oxelytrum discicolle (Brullé),
24 2,840 om ‘ 82Q Te 2,600 IHL F ae £0 we 2 & yi pO Bek
2.900 04
Nicrophorus | ; - i "mexicans
trophioris olds
a sits liscicolle
ie | Abies a distinbed
Cirpressus | oakcpine | i Wie ke, “# tly forest —s forest «== Ae
forest forest
SS
association —
Figure 1. Altitudinal distribution of three Silphidae species from El Floripondio, Jalisco, collected with carrion traps.
46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
250 -—
200 : 150 ‘oO iD) oq, NM = 100 a —
50 BR ROLES 0 "ANSI NN crac
@ Nicrophorus mexicanus Nicrophorus olidus
Figure 2. Abundance of Nicrophorus mexicanus and N. olidus from El Floripondio, Jalisco, col- lected with carrion traps between April and October, 1998.
Nicrophorus mexicanus (Matthews) and Nicrophorus olidus (Matthews). These species were represented by 743 specimens. Nicrophorus mexicanus was the com- monest species (628 specimens), and was collected monthly and distributed be- tween 2300—2950 m. Nicrophorus olidus was the second most common species, represented by 107 specimens, found between 2300—2600 m, but collected only during May—August (Fig. 1). Finally, only eight specimens of O. discicolle were collected in the Cupressus forest in June. Other Mexican localities demonstrating similar silphid abundance and distribution include: Volcan de Tequila, Jalisco (Navarrete-Heredia, 1995); Sierra de Manantlan (Rivera-Cervantes & Garcia- Real, 1998) and Cofre de Perote, Veracruz (Arellano, L. 1998. Dugesiana, 5(2): 1-16). In all of them, Nicrophorus species are well represented, mostly at higher elevations, whereas O. discicolle is less abundant and restricted to lower places.
Acknowledgment.—Dr. Rodolfo Novelo for review of this note, and Dr. Mar- garet K. Thayer helpful comments.
Hugo E. Fierros-Lé6pez and José Luis Navarrete-Heredia, Entomologia, Centro de Estudios en Zoologia, Universidad de Guadalajara, Apdo. Postal 234, 45100 Zapopan, Jalisco, México.
Received Nov 30, 1999; Accepted May 31, 2000.
PAN-PACIFIC ENTOMOLOGIST 77(1): 47-50, (2001)
FIRST RECORDS OF LEPTOPODIDAE IN WASHINGTON STATE (HEMIPTERA: HETEROPTERA) WITH NOTES ON HABITAT
R. S. ZAcK, C. N. LOONEY, M. E. HITcHcox, & J. P. STRANGE
Department of Entomology, Washington State University, Pullman, Washington 99164-6382
Abstract.—Patapius spinosus (Rossi) was found in several locations in Benton and Whitman Counties, Washington State. Adults were found from late January through late October with mating pairs and immatures being found only in the fall of the year. The bug was associated with cobblestone and basalt that was being used as roadgrade stabilization or in piles of such rock that had been left after construction activities.
Key Words.—Insecta, Leptopodidae, Hanford Site, Patapius spinosus (Rossi), Washington State.
Extant leptopodid bugs are primarily Old World in distribution with fossil ev- idence known from Mexico and Ecuador (Froeschner 1988). The first recording of a leptopodid from the United States was in California (Arbuckle, Colusa Coun- ty) where a single individual was found during the examination of tree protectors used to trap peach twig borer larvae on almond trees (Usinger 1941). Subsequent to this first finding, the species in question, Patapius spinosus (Rossi) has been found in Nevada, and Idaho (Brothers 1979). Patapius spinosus is currently known from California, Nevada, and Idaho in the United States, the Canary Is- lands, Europe, North Africa, and has been introduced into Chile (Froeschner 1988). Both Brothers (1979) and Froeschner (1988) provide excellent illustrations of the bug. Our finding of this species at several widely separated locations in Washington State is a significant extension of its previously known range and may indicate that the species is relatively widespread in the western United States.
Patapius spinosus appears to be rather generalized in its habitat selection. Both Usinger (1941) and Brothers (1971) found the species in areas removed from water sources; Brothers (1971) found specimens on the undersides of cobblestone in a mine-tailings dump. The sites from which we obtained specimens are com- parable to a mine tailings area except that our sites are located near water (which the tailings site may have been). Our first finding occurred at the Hanford Nuclear Site (Benton County) in southcentral Washington State. The Hanford Site is an area of native shrub-steppe vegetation with semi-arid climatic conditions that in- clude hot and dry summers and cold winters. Annual precipitation is less than 12 cm. Temperatures range from an average of 3° C in Jan to 33° C in July; tem- peratures of 30° C or above occur an average of 56 days per year (ERDA 1975).
Specimens were found on the undersides of cobblestones that had been depos- ited along the margins of a built-up gravel road that ran parallel to an alkaline pond (West Lake-—46°36.06’ N, 119°32.78’ W) located at 150 m in elevation. West Lake is the only naturally occurring pond on the Hanford Site. The pond is surrounded by an alkaline crust with no emergent macrovegetation along the shoreline. However, large areas of bullrush (Scirpus sp.) as well as various grasses and other vegetation occur in the area surrounding the pond. The size of the pond is a direct function of ground water elevation and fluctuates throughout long and
48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
short term periods depending on climate and seasonal weather conditions. On average, the pond encompasses 4 hectares. The first specimens were discovered while examining the undersides of the cobblestone on 15 Jan 1999. Subsequent to this finding, a more concerted effort was undertaken on 29 Jan during which several hundred rocks were turned and 30 adult specimens located. Again, on 12 Feb, several specimens were found; we continued to find specimens, sporadically, throughout the summer and into early September when the study, at this site, was discontinued. Only material from the first two collecting dates were taken and processed. Additionally, a single specimen was collected in a pitfall trap (14—26 Aug 1998) located in vegetation surrounding West Lake. Although no effort was made at these times to quantify our collections, we noticed that specimens were never found on the section of rock that was in contact with the soil (moist) and that there appeared to be a certain clumping phenomenon to their discovery. Although few rocks harbored specimens, multiple specimens were sometimes dis- covered under individual rocks. Specimens are deposited in the M. T. James En- tomological Collection, Washington State University.
In an examination of the M. T: James Entomological Collection, Washington State University, we located two specimens that had been collected along the Snake River at the McCauley Ranch that was located (area since flooded) between Almota and Penwawa Canyons (Whitman County) on 20 Mar 1971. A descriptive label on these specimens stated that they had been collected under rocks, as had our Hanford Site specimens. We therefore expanded our survey activities to sev- eral, apparently likely sites adjacent to the Snake River. These sites were all located in Whitman County, along the Snake River: Wawawai River road, ap- proximately 19 km SW of Pullman (46°36.91’ N, 117°22.51’ W); corner of Wa- wawai Grade road and Wawawai River road, approximately 17 km SW of Pullman (46°36.05’ N, 117°22.65' W); Almota, (Fig. 1) approximately 24 km W of Pull- man (46°42.22’ N, 117°27.98' W) and; Boyer Park (Fig. 2) (at Lower Granite Dam) approximately 22 km WSW of Pullman (46°40.64' N, 117°26.74’ W).
On 28 Feb 1999, sampling was conducted at the above-delineated Snake River sites. Records were kept of the number of rocks turned, the number of times leptopodids were found, and in what size “‘groups.’’ A total of 1610 rocks were turned with 94 leptopodids found. Specimens were found singly and in groups of up to 10 on the underside of a single rock. While no samples were taken during the summer, on 21 Oct 1999 the Snake River sites were again surveyed. At this time, no count of bugs was maintained and only a small number were collected and processed. However, the number of leptopodid adults was high and, indeed, appeared higher than during our winter sampling periods. Additionally, this was the only time that we found mating pairs and immatures. Again, leptopodids were found in small piles of cobblestone and basalt rock that had been used as road- grade support or had been left in small piles (often of several hundred rocks) after completion of road or walking path construction. These latter piles were often scattered and had been overgrown with grasses (Fig. 2). It is interesting to note that not every pile of rock contained the bugs but, when they were found, nu- merous rocks within the pile harbored them.
Rocks with leptopodids underneath always contained some open space below and were almost never wet. In the few instances where rocks with leptopodids below were wet, all of the surrounding rocks in the pile were also wet. In most
Figure 2. Pile of basalt rock at Boyer Park (Whitman County).
50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
areas, when wet and dry rocks were adjacent, it appeared that the insects favored the dry rocks. Rocks with leptopodids on them varied extremely in color and arrangement. There was no apparent pattern in size or color of rock and the number of specimens occurring on the underside of the rock. Some piles of rocks were embedded in grasses; others were on bare dirt. In no cases were leptopodids found on the steep piles of basalt rubble adjacent to roadwork or railroads. When the insects were found near railways and road cuts, they were always on the undersides of rocks arranged in a flatter and less congested manner.
It is interesting to speculate on how P. spinosus was introduced into Washing- ton. At both Hanford and the Snake River sites, the bugs were found under cobblestone, river rock that is used for stabilization of roadways. In all instances, the rock had been brought to the site from other areas that could not be discerned by us. However, at Hanford, the rock may have come from the adjacent Columbia River (approximately 3 km from West Lake) and, at the Whitman County sites, rock may have come from the adjacent Snake River. It may be that the bugs are associated with cobble habitat and are distributed as this rock is moved and used in construction efforts. Unfortunately, besides our minimal observations, we have no evidence of such movement.
ACKNOWLEDGMENT
This project was founded by The Nature Conservancy with awards from the U.S. Department of Energy, The Nature Conservancy of Washington State, and the Bullitt Foundation. We thank Dara Nunn and Jennifer Strange for aid in collecting field data.
LITERATURE CITED
Brothers, D. R. 1979. First record of Patapius spinosus in Idaho and Nevada (Hemiptera: Leptopod- idae). Great Basin Nat., 39: 195-196.
Energy Research and Development Administration (ERDA). 1975. Final Environmental Statement, Waste Management Operations. ERDA-1538. Volume 2. Energy Research Development Ad- ministration, Richland, Washington.
Froeschner, R. C. 1988. Family Leptopodidae Amyot and Serville, 1843. p. 166. Jn Henry, T: J. and R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden.
Usinger, R. L. 1941. A remarkable immigrant leptopodid in California. Bull. Brooklyn Entomol. Soc., 36: 164-65.
Received 24 Nov 1999; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 51-53, (2001)
Scientific Note
ERNOBIUS MOLLIS (L.) (COLEOPTERA: ANOBITDAE): AN EXOTIC BEETLE COLONIZES MONTEREY PINE, PINUS RADIATA D. DON, IN NORTHERN CALIFORNIA
In 1990, the first collection of the bark anobiid, Ernobius mollis (L.), west of Texas was made in Oakland, Alameda County, California from a living Norway spruce, Picea abies (L.) Karsten, planted as an ornamental tree (Seybold, S. J. & J. L. Tupy, 1993. Pan-Pacif. Entomol., 69: 36—40.). Ernobius mollis is native to northern Europe, but has been introduced into North America (Craighead, F C. 1950. Insect Enemies of Eastern Forests, USDA For. Serv. Misc. Pub. 657.) and the southern hemisphere (Casimir, J. M. 1958. Div. Wood Tech., For. Comm. New South Wales Tech. Notes, 2: 24—27 [Australia, New Zealand, and South Africa]). Seybold and Tupy (1993) speculated that E. mollis, known to colonize Monterey pine, Pinus radiata D. Don, in the southern hemisphere, might soon colonize the abundant urban plantings of P. radiata in the San Francisco Bay Area.
On 20 Sep 1993, a homeowner in Albany, Alameda County, California (Talbot Street) reported that she had observed insect damage in a variety of wooden articles in her home. On examination of her residence, none of the wooden articles of concern to the homeowner appeared to harbor active infestations of wood- destroying insects. However, examination of several pieces of barked P. radiata firewood stacked within the structure indicated that the small branch sections were infested with a wood-boring insect. These P. radiata branches had emergence holes through the bark surface. Peeling the already loose bark from the xylem revealed live larvae and pelleted frass in the bark-xylem interface as well as larval insect galleries etched in the xylem surface. The homeowner reported that the P. radiata branches had been cut approximately six months earlier (1.e., Mar 1993) from a standing tree in Fremont, Alameda County, California on Bud Court near the Highway 880 Mowry Avenue Exit. The branches were placed in a laboratory cage at the University of California at Berkeley at ambient indoor temperature (16° C to 29° C) and larval and adult E. mollis were periodically collected from the logs from Sep 1993 to Jun 1996. By Jun 1996 the bark-xylem interface had been completely obliterated and the remaining paper thin bark formed an easily damaged shell over the xylem.
This collection documents the presence of E. mollis in urban plantings of P. radiata in the San Francisco Bay Area. Although this collection record was from cut branches, future northern California reports of infestations from moribund P. radiata and perhaps other ornamental conifers should be expected. Through its association with moribund tree tissue, E. mollis may also play a role in the dis- semination of the pitch canker fungus, Fusarium circinatum Nirenberg & O’Donnell (Nirenberg, H. I & K. O’Donnell. 1998. Mycologia, 90: 434—458.), a fairly recently introduced and fatal disease of P. radiata and other pines in coastal northern California (McCain, A. H., C. S. Koehler & S. A. Tjosvold. 1987. Calif. Agric., 41: 22—23.). In Europe, E. mollis is documented to infest cones of Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, and giant sequoia,
32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
Sequoiadendron giganteum (Lindley) J. Buchholz (Roques, A. 1983. Les insectes ravageurs des c6nes et graines de coniféres en France, Institut National De La Recherche Agronomique Publication.). A native species of Ernobius [punctulatus (LeConte)] is known to infest cones of P. radiata (White, R. E. 1982. A Catalog of the Coleoptera of America North of Mexico Family Anobiidae, USDA Agric. Handbook No. 529-70, Furniss, R. L. & V. M. Carolin 1992. Western Forest Insects, USDA For. Serv. Misc. Publ. No. 1339.) and to acquire the pitch canker fungus through its association in the P. radiata cone with other cone-infesting insects (Hoover, K., D. L. Wood J. W. Fox & W. E. Bros. 1995. Can. Entomol., 127: 79—91.). Therefore, association of E. mollis with moribund P. radiata cone tissue and the bark-xylem interface of branch and stem material appear to be a likely means for this introduced insect to disseminate pitch canker disease in northern Califormia. As both the insect and the fungal pathogen have wider host ranges among conifers, E. mollis could play a role in transmission of the fungus to new hosts and to new regions such as the montane forests of the Sierra Nevada or the coastal forests of the Pacific Northwest.
Pinus radiata is frequently brought into coastal California homes as firewood, and this example illustrates how easily larval and adult E. mollis might enter homes. However, experiences from Europe, the southern hemisphere, and eastern North America suggest that because of its requirement for bark-covered sapwood, E. mollis will not be a pest in structures in northern Califormia unless it encounters bark-covered timbers, or boards with unfinished edges or bark-occluded knots. This example also illustrates that although an active infestation of E. mollis may be present in firewood in a structure, if damage to other, unbarked wooden items in the structure clearly occurred prior to manufacture, any relationship between E. mollis and the damage can likely be discounted.
Finally, the original colony of E. mollis collected 22 Feb 1990 in Oakland was maintained until Aug 1996 in a laboratory cage at ambient indoor temperature on the original P. abies branch and stem material. On 12 Jul 1996, seven live male and twelve live female adults were recovered from the cage. One pair of these adults was observed in copulo and live larvae were also present. On 6 Aug 1996, one live male, four live females, and two live larvae were also recovered from the cage before the logs were frozen and disposed of. Thus, multiple, consecutive generations of E. mollis successfully re-infested the same dry substrate for more than 6.5 years.
Record.—USA. CALIFORNIA. ALAMEDA CO.: 1.5 km SE Albany Hill, Tal- bot Street, Albany, 20 Sep 1993, S. J. Seybold, Pinus radiata.
Acknowled gment—I thank R. Kohl for bringing this occurrence of E. mollis to my attention; A. J. Blomquist (University of Nevada, Reno) for helping to process the specimens; D. L. Wood (University of California at Berkeley) and B. J. Cabrera (University of Minnesota) for critical reviews of earlier drafts of this report; and N. G. Rappaport (USDA Forest Service, Albany, California) for trans- lation of French literature. This work was supported by a cooperative research agreement (PSW-92-0014CA) between DLW and the Chemical Ecology of Forest Insects Project, Pacific Southwest Research Station, U.S. Department of Agricul- ture, Forest Service, Albany, California. Specimens of E. mollis recovered from P. radiata were deposited in the California Academy of Sciences Entomology Collection, Golden Gate Park, San Francisco, Califomia 94118.
2001 SCIENTIFIC NOTE 53
Steven J. Seybold, Departments of Entomology and Forest Resources, 219 Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, Minnesota 55108-6125.
Received April 9, 2000; Accepted July 6 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 54—56, (2001)
Scientific Note
RANGE EXTENSION OF PSENEO PUNCTATUS FOX AND NOTES ON PREDATION OF AN INTRODUCED SHARPSHOOTER, HOMALODISCA COAGULATA (SAY)
The tribe Psenini Bohart & Menke (Hymenoptera: Sphecidae: Pemphredoninae) was described in 1976 (Bohart, R. M. & S. Menke 1976. Sphecid wasps of the world, a generic revision, University of California Press, Berkeley, California, USA). It is a poorly understood group, and basic biologies are unknown. Psenini are small, slender, delicately constructed wasps that most commonly occur in association with damp shady situations. Some genera and species within the tribe are located in southern Califormmia (R. M. Bohart, personal communication). How- ever, only one species of Pseneo has been recorded in California, Pseneo longiv- entris (Cameron) (van Lith, J. P. 1975. Neotropical species of Psen and Pseneo [Hymenoptera: Sphecidae: Psenini]. Tijdeschrift voor Entomologie. 118: 1—41).
We became aware of a large aggregate of digging wasps on the grounds of the University of California Riverside in August 1998. Many of the nests seemed to be completed, closed, nests, but the majority had entrances open with provisioning occurring. Many nests were so close together that the mounds at the entrances were overlapping. The wasps appeared to have little difficulty finding their own nests, and no aggressiveness was observed between neighboring females. We counted 181 nesting sites within 30 plant propagation trays (45 x 45 x 10 cm) filled with U. C. mix #3 (Matkin, O. A. & P. A. Chandler. 1957. The UC soil type mixes. U. C. Berkeley, California Agriculture Experiment Station, Manual 23) to a depth of 6 cm. All trays were located within a 10 m? area in an open lathhouse on raised benches. The soil-filled trays were being temporarily stored prior to being used for seedling establishment in unrelated experiments. Nesting wasps were observed wherever the soil was exposed in any of the trays. We observed wasps active in 53 (29%) of the observed 181 nesting sites.
Specimens were collected and identified as P. punctatus Fox, a species found from North Dakota to southern Mexico. Features of the male, including the spe- cies-specific details of the antenna and genital capsule, clearly match those of typical P. punctatus. However, the Riverside wasps have extensive reddish col- oration on the legs in both sexes; in van Lith’s (1975) key they run to P. carolina, a southeastern U.S. species described exclusively from females (also originally described as a subspecies of P. punctatus). Specimens of putative male P. carolina from Texas raise questions as to the validity of the taxon, as they are nearly identical to the California specimens of punctatus (A. Finnamore, personal com- munication), and the resultant distribution of “‘carolina’’ would apparently bisect the known distribution of punctatus. Our suspicion is that P. punctatus is a species with a large distribution and substantial geographic variation, which has not yet been sampled adequately to resolve the limits of the various forms, including P. carolina, which was probably inappropriately elevated to species status by van Lith. Pseneo punctatus represents the second Pseneo species collected in Cali- fornia (Bohart, personal communication; Krombein, K. V. 1979. Superfamily
2001 SCIENTIFIC NOTE 55
Sphecoidea. pp. 1573-1740. Jn Krombein, K. V., P. D. Hurd, Jr., D. R. Smith & B. D. Burks [eds.]. Catalogue of Hymenoptera of America North of Mexico. 2: 1119-2209. Smithsonian Institution Press, Washington, D.C., USA). Voucher specimens are located at the University of California Riverside, Entomology Re- search Museum.
During our observations of P. punctatus over a period of several days we noted the following behaviors. Nest excavation was observed only in the afternoon. Nests were excavated with the mandibles and front legs. Soil was loosened by the mandibles, formed into a small uneven clump, and pushed backward beneath the body with the front legs. Usually the clumps of soil are thrown clear of the body by the initial thrust of the front legs. If not, the soil clumps are thrown clear of the body with thé assistance of the middle and hind legs. The female typically digs the burrow vertically straight down. As the wasp digs deeper, small clumps of soil clog the nest entrance and hide the digging female. Occasionally, as soil accumulates near the nest entrance, the wasp would back up and clear the entrance hole. During mound building, most wasps would occasionally tamp the soil down with their abdomens, presumably to prevent the nest entrance from becoming obstructed with excavated soil. Nest entrances were 6.50 + 2.35 mm (mean + SD, n = 16, range 5-12 mm) in diameter, and were sometimes hidden under fallen leaves or other debris.
Over a period of hours to days, females repeatedly provisioned their nests with large Homopteran sharpshooters (Cicadellidae). When the nest was fully provi- sioned, the female would repeatedly emerge and re-enter the nest entrance drag- ging clumps of soil with her forelegs back into the hole each time she entered. When the upper portion of the burrow was filled with soil, the female would walk over the top of the mound repeatedly scraping the surface with her forelegs and tapping it with her abdomen. At this point, it was no longer possible to discern that a burrow was present.
What was of particular interest to us was that P. punctatus was using both the native smoketree sharpshooter, Homalodisca lacerta (Fowler) (Homoptera: Ci- cadellidae), and the introduced glassywinged sharpshooter, Homalodisca coagu- lata (Say), as prey. Homalodisca coagulata became established in California dur- ing 1989-1990 (Sorenson, J. T. & R. J. Gill. 1990. Pan-Pacific Entomol., 72: 160-161). Ithas now been established in 11 counties and on numerous host plants including citrus, grapes, oleander and a wide variety of ornamental landscape and native plants. Currently, H. coagulata is the principle vector of Pierce’s disease of grapevines in the southern portion of California (Costa, H. S., M. J. Blua, J. A. Bethke, & R. A. Redak. 2000. [In press]. HortScience University of California, Office of the President. 2000. Report of the Pierce’s Disease Research and Emer- gency Task Force). The vast majority of prey we observed were H. coagulata adults. Pseneo punctatus’ natural prey most likely is H. lacerta, and we believe it has expanded its usable hosts to include the recently introduced sharpshooter.
When several of the nests were excavated, we usually observed three cells per nest, but it was difficult at times to accurately determine how many cells each nest contained. Many cells contained sharpshooters that were old, decayed and untouched. We could not determine if they were abandoned or if they were part of the normal provisioning of the nest and simply unused. Although the number of sharpshooters per cell varied, they commonly contained about 4 adults. In some
56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
cells, we observed eggs deposited between the front pair of coxae on the venter of a single sharpshooter. They were oblong, white, and slender, and on average they were 2.29 + 0.35 mm long (mean + SD, n = 5, range 3.0—4.6).
Wasp cocoons were also observed. On average cocoons were ellipsoidal, 14.50 + 1.34 mm in length, and 5.79 + 0.70 mm in width (mean + SD, n = 14, range 13-17, and 5—7 respectively). Cocoons were covered with small pieces of plant material and sharpshooter fragments (mostly head capsules and wings).
Occasionally, we observed wasps emerging from the soil as adults. Emergence holes were a mean of 3.64 + 0.50 mm in diameter (mean + SD, n = 11, range 3—4) and unlike active nest entrances, they were not marked by any debris or mounds of dirt.
Specimens Examined CALIFORNIA. RIVERSIDE Co.: Riverside, University of California cam- pus, lath house, 21 Aug 1998, Kathleen A. Campbell, swept, 2 females, deposited: University of California, Riverside, Entomology Research Museum; same loc., 25 Aug 1998, reared from larva, 1 female, reared from pupae, (6 males, 2 females), deposited: University of California, Riverside, En- tomology Research Museum; same loc., 27 Aug 1998, reared from larva, 1 female, deposited: Uni- versity of California, Riverside, Entomology Research Museum; Riverside, University of California campus, undeveloped area 2 meters west of the lath house, 11 Sept 1998, James A. Bethke, swept from eucalyptus, (1 male, 3 females), deposited: University of California, Riverside, Entomology Research Museum; Riverside, University of California campus, solid bin adjacent to Entomology Annex I, swept, 1 female, deposited: University of California, Riverside, Entomology Research Mu- seum.
Acknowled gment.—Authors thank R. Bohart for review of this manuscript, L. Petro for helpful observations, and R. Bohart and A. Finnamore for helpful com- ments.
James A. Bethke, Kathleen A. Campbell, Matthew J. Blua, Richard A. Redak, and Douglas A. Yanega, Department of Entomology, University of California, Riverside, California 92521.
Received 11 Mar 2000; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(1): 57-60, (2001)
Scientific Note
MONODONTOMERUS ARGENTINUS BRETHES (HYMENOPTERA: TORYMIDAE): A PARASITOID OF EUGLOSSA NIGROPILOSA MOURE (HYMENOPTERA:
APIDAE: EUGLOSSINAE)
Euglossine bees are the only group in the subfamily Apidoideae that do not have eusocial behavior. Euglossines display a broad spectrum of social interac- tions, from solitary to presocial, and are thus an important group for studying the evolution of eusociability in bees (Gardéfalo, C. A. 1985. Entomol. Gener., 11: 77-83). An important factor involved in the origin of social behavior is the effect of parasites and parasitoids on survival and reproduction (Roubik, D. W. 1989. Ecology and natural history of tropical bees [1st ed.]. Cambridge University Press. New York).
Although it is known that microhymenoptera are parasitoids of Euglossa (Zuc- chi, R., S. FE Sakagami & J. M. EK de Camargo. 1969. J. Fac. Sci. Hokkaido Univ., Series VI, Zool., 17: 271-380), little is known about their relationship with their hosts (Dressler, R. L. 1982. Ann. Rev. Ecol. Syst., 13: 373-394). Here I present data on the parasitic behavior of the microhymenopteran Monodontomerus ar- gentinus Brethes (Torymidae) and discuss its effect on the social structure of E. nigropilosa. 1 compared the percentage parasitism of M. argentinus on both an attended and an unattended nest of E. nigropilosa. This study was carried out between August 1994 and April 1995.
Euglossa nigropilosa is a communal bee of NW South America, the colonies of which have active bees all year. It is distributed between 700 and 1400 m in the Andes of Colombia and Ecuador. This species builds nests with an envelope, a resinous cover that protects the cells. Cells are of 10 X 6 mm, with soft resinous walls. I found several nests of E. nigropilosa in wood cavities of timber buildings at the Reserva Natural La Planada (RNLP), Colombia (77°24’ W, 1°5’ N). The nests contained colonies of up to 22 females (Otero, J. T. 1996. Bol. Mus. Ent. Univ. Valle. 4: 1-19).
Nests of E. nigropilosa were parasitized by M. argentinus, an ectoparasitic wasp that is known to attack the solitary bee Eufriesea nigrescens Friese, another euglossine, in the eastern part of the Andes in Colombia (Sakagami, S. FE & Strum. 1965. Insecta Matsumurana, 28: 83—97). This is the first record of a different host for M. argentinus. Samples of both species were deposited in the Museo de En- tomologia of Departamento de Biologia de la Universidad del Valle (Cali, Co- lombia).
With the objective to study bee behavior inside the nest, on 20 Sept. 1994 I placed a nest with four adult bees and 18 cells of E. nigropilosa in a wooden box 30 X 15 X 10 cm, with a glass plate covering the upper surface and a wood cover which remained in place during non-observation hours. The bees had free access through a one cm diameter hole. However, the adult bees abandoned the box nest. This undefended nest was compared to a nearby undisturbed nest, in which adult bees were still present, for levels of cell parasitism by M. argentinus.
58 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
This second nest provided a control for the degree to which adult bees can protect their nest from parasitism. On 11 Nov. 1994 I opened all the cells of the aban- doned box nest to check the level of parasitism of M. argentinus. For comparison, I opened 20 randomly chosen cells from the undisturbed nest to determine whether or not they were parasitized.
Comparison between the nests with and without females revealed that parasit- ism dropped significantly (x? = 12.73, P = 0.0004) with the presence of adults in the nest. The percentage of parasitism on the 18 cells in the box nest with female bees absent was 72%. However, the percentage for the control nest, which had between eight and 14 female bees, was only 15%, based on the sampled cells. In the box nest cells there was an average of 13.69 (+ 9.04 SD, n = 13) wasp offspring per cell. Individual wasps were found at all developmental stages: young larvae, mature larvae, prepupae (pupae without pigmentation); and pupae (black pupae).
I observed the behavior of 38 M. argentinus wasps inside the defended E. nigropilosa nest. Monodontomerus argentinus arrived at the nest entrance after a zigzag flight. The wasp flew around the entrance for up to five min, before landing at the entrance hole. Parasitoid wasps entered the nest slowly, using the same entrance hole that the bees used. Upon landing, the wasps drummed on the surface of the nest with their antennae, moving them fast and harmoniously. The wasps alternated each antenna in this process. The wasps advanced slowly and continued drumming until they arrived at the entrance hole. Once inside the nest, the wasps drummed with their antennae on the surface of a cell for up to five min. without moving any other part of the body. When a wasp found an appropriate cell, it continued drumming for several minutes before ovipositing. Oviposition lasted for up to two min. For this process the wasp raised her abdomen, exerted her Ovipositor and put it in contact with the resin wall of the cell. Occasionally the parasitoids withdrew their abdomen and continued drumming on the next cell.
Euglossa nigropilosa appeared to recognize M. argentinus as a natural enemy. These interactions were very strong. For example, a parasitoid, recently killed in a cyanide bottle, was left exposed in the nest. This body was torn apart by a resident bee. On first detecting the dead wasp, the bee became excited and circled around it, touching it with her antennae. It then bit the wasp in the abdomen and pushed it five cm away from the cell. The bee again found the dead wasp on the ground and bit it repeatedly for more than a minute. Following this attack the parasitoid lost two legs and had its wing and crushed abdomen nearly destroyed.
This strong reaction may, however, have been a response to the odor of the cyanide, and not the presence of the parasitoid body. During observations of attempts to parasitize a cell, I never observed the bees to perceive the wasp’s presence. The wasps proved very adept at escaping from the bees notice in the nest. When a parasitoid was approached by a bee, the wasp jumped to the ground, so evading detection, and remained there for several minutes before attempting to oviposit again.
Sakagami and Strum (1965) found an aggregation of nests of the solitary bee Euplusia longipennis in Colombia in 1956 (Eufriesea nigrescens sensu Kimsey, L. S. 1982. Systematics of bees of the genus Eufriesea. University of California press). These nests had 81 highly elaborate resinous cells of approximately 19 X 9 mm, two of which were parasitized by M. argentinus (2.5% parasitism). Each
Table 1. Length of bees and cells, and number of broods per cell of the parasitic wasp Monodontomerus argentinus in two different euglossine bees in Colombia.
Euglossa nigropilosa Eufriesea nigrescens Bee length (mm) 12 16 Cell length (mm) 10 19 Cell diameter (mm) 6 8 Number of broods by cell 13.7 (SD = 9.4; n = 13) 33.5 (SD = 2.1; n = 2)
AHLON OWILNAIDS 1007
6S
60 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)
parasitized cell had an average of 33.5 (SD + 2.1, n = 2) parasites. Eufriesea nigrescens and Euglossa nigropilosa were attacked by the same parasitoid, M. argentinus, but the two host species have different defense strategies. The solitary species, E. nigrescens, provides cells with thick protective walls. In contrast E. nigropilosa build cells with a thin, vulnerable wall. However, adult bees of E. nigropilosa may protect the nest from M. argentinus. In addition, the presence of a resinous envelope enclosing the nest may restrict the wasp’s entrance. These defenses of E. nigropilosa, however, appear to provide less effective protection from the parasitic wasps than the thick cell walls of E. nigrescens. Parasitism incidence was higher in E. nigropilosa, between 72% in box nest and 15% in the control nest, than in E. nigrescens (2.5%). There appears to be less parasitism when nests are protected by many active females. In this case, the parasitoids probably spent more time escaping from resident bees and were not able to ovi- posit in cells.
From these observations I suggest that the presence of active bees in the nest may affect the rate of M. argentinus parasitism of E. nigropilosa. Despite having vulnerable cell walls, E. nigropilosa has active bees throughout the year and the large colony size makes the presence of a female in the nest more likely, decreas- ing the risk of M. argentinus attack. In contrast, E. nigrescens is a seasonal bee with oviposition limited to only two months of the year. Females of Eufriesea nigrescens die after oviposition and they can not protect the cells. Thus an in- trinsic protective mechanism is needed, in this case a thick resinous cell wall that is difficult for the parasitoid to penetrate.
Acknowled gment.—I thank Dr. Philip Silverstone-Sopkin and Dra. Patricia Cha- c6n from Universidad del Valle and Dr. R. Ospina and G. Nates from Universidad Nacional de Colombia for their support; Dr. E. E. Grissell from USDA for iden- tification of wasps; Dr. J. Ackerman, Dr. W. O. McMillan, Dra. N. Flanagan and Dr. P. Bayman for comments in the manuscript; FES foundation for logistic sup- port in the Reserva Natural La Planada and the MacArthur Foundation for eco- nomic support.
J. Tupac Otero, Department of Biology, University of Puerto Rico, San Juan, Puerto Rico 00931-3360.
Received I Mar 2000; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST Information for Contributors
See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English, but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be retumed. Please maintain a copy of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.
Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in, nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF ANY FIGURES TO THE 8.5 X 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text pages (pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The title must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.
Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet 1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use: (Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith 1987: table 3).
Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194—198; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED (INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author of each species after its first mention.
Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are applicable.
Literature Cited. — Format examples are:
Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.
Blackman, R. L, P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB Academic Publishing, The Hague, The Netherlands.
Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.
Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.
Hlustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors are strongly encouraged to provide illustrations no larger than 8.5 X 11 in for easy handling. Number figures in the order presented. Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of the manuscript, and (5) journal. FIGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.
Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.
Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—161.). A short acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of the note with the affiliation underscored.
Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page charge fee notice with acknowledgment of initial receipt of manuscripts.
THE PAN-PACIFIC ENTOMOLOGIST
Volume 77 January 2001 Number 1
Contents
PUBLICATIONS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY ____-------------------
BRAILOVSKY, H. & E. BARRERA—The genus Notobitopsis Blote with the description of two new species (Hemiptera: Heteroptea: Coreidae: Coreinae: Mictini)
SCHEFTER, P. W. & K. A. JOHANSON—Three new species of Helicopsyche from Vietnam (Trichoptera: Helicopsychidae)
MILICZKY, E. R. & C. O. CALKINS—Prey of the spider, Dictyna coloradensis, on apple, pear, and weeds in central Washington (Araneae: Dictynidae)
BERNAL, J. S., D. GONZALEZ & E. DAVID-DIMARINO—Overwintering potential in California of two Russian wheat aphid parasitoids (Hymenoptera: Aphelinidae et Aphid- iidae) imported from Central Asia
ANDREWS, F. G.—A new species of Corticarina from Arizona (Latridiidae: Corticariini)
SCHWARTZ, M. D. & M. A. WALL—Melanotrichus boydi, a new species of plant bug (Heteroptera: Miridae: Orthotylini) restricted to the nickel hyperaccumulator Streptanthus polygaloides (Brassicaceae)
ZACK, R. S., C. N. LOONEY, M. E. HITCHCOX & J. P. STRANGE—First records of Leptopodidae in Washington State (Hemiptera: Heteroptera) with notes on habitat
SCIENTIFIC NOTES FIERROS-LOPEZ, H. E. & J. L. NAVARRETE-HEREDIA—Altitudinal distribution and phe-
nology of three species of carrion beetles (Coleoptera: Silphidae) from Nevado de Coli- ma, Jalisco, México
SEYBOLD, S. J.—Ernobius mollis (L.) (Coleoptera: Anobiidae): an exotic beetle colonizes Monterey pine, Pinus radiata D. Don, in Northern California
BETHKE, J. A., K. A. CAMPBELL, M. J. BLUA, R. A. REDAK, AND D. A. YANEGA—Range extension of Pseneo punctutus Fox and notes on predation of an introduced sharpshoot- er, Homalodisca coagulata (Say)
OTERO, J. T—Monodontomerus argentinus Brethes (Hymenoptera: Torymidae): a parasitoid of Euglossa nigropilosa Moure (Hymenoptera: Apidae: Euglossinae)
19
28 37
ao
47
45
51
54
a7
The
PAN-PACIFIC
ENTOMOLOGIST
Volume 77 April 2001 Number 2
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603)
The Pan-Pacific Entomologist
EDITORIAL BOARD
J. Y. Honda, Editor R. M. Bohart
Z. Nisani, Assistant to the Editor J.T. Doyen
R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta F. Parinas, Treasurer Warren E. Savary
Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary; and financial communications should be addressed to: Julieta F. Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Application for membership in the Society and changes of address should be addressed to: William Hamersky, Membership Committee chair, Pacific Coast Entomological Society, Dept. of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. Robert V. Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture, 1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con- tributors, and volume 73(4): 248—255, October 1997, for more detailed informa- tion. Information on format for taxonomic manuscripts can be found in volume 69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.
The annual dues, paid in advance, are $25.00 for regular members of the So- ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for institutional subscriptions or sponsoring members. Members of the Society receive The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes are available; see 72(4): 247 for current prices. Make checks payable to the Pacific Coast Entomological Society.
Pacific Coast Entomological Society
OFFICERS FOR 2001
Stanley E. Vaughn, President Vincent F. Lee, Managing Secretary Katherine N. Schick, President-elect Joshua J. Fairbanks, Recording Secretary Robert L. Zaparko, Treasurer
THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and addition- al mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
This issue mailed 2 April 2001
The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.
The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PAN-PACIFIC ENTOMOLOGIST 71(2): 61-67, (2001)
INVERSE DENSITY-DEPENDENT PARASITISM OF OPSIUS STACTOGALUS FIEBER (HOMOPTERA: CICADELLIDAE) BY GONATOPUS SP. (HYMENOPTERA: DRYINIDAE)
W. D. WIESENBORN 1410 Stacey Lane, Boulder City, Nevada 89005
Abstract—The relationship between Opsius stactogalus Fieber leafhopper density (abundance per plant-sample mass) and parasitism by the dryinid wasp Gonatopus sp. was examined. Branch- es cut from Tamarix ramosissima Ledebour shrubs, sustained by treated wastewater, were weighed and sampled for parasitized and unparasitized leafhoppers. Leafhopper abundance in- creased linearly as branch mass increased exponentially (mass®-®), in agreement with isometric scaling laws relating leaf abundance to branch mass. The proportion of leafhopper nymphs parasitized (18%) was greater than the proportion of adults parasitized (2.9%). Branches were more likely to contain one or more parasitized nymphs, signifying exploitation of the branch by Gonatopus, as nymph density increased. On branches containing at least one parasitized nymph, increasing nymph density was related to an increase in the number of parasitized nymphs but to a decrease in parasitism rate. Parasitism of O. stactogalus nymphs on branches exploited by Gonatopus was inverse density dependent. Gonatopus appears not to regulate populations of O. stactogalus or limit the leafhopper’s damage to T. ramosissima.
Key Words.—Insecta, Cicadellidae, Opsius stactogalus, Dryinidae, Gonatopus, Tamarix, para- sitism.
The tamarix leafhopper, Opsius stactogalus Fieber, is a small, cryptic insect primarily found on tamarisk, Tamarix spp. (Tamaricaceae) (Harding 1930, Liesner 1971). Tamarisk is a halophytic shrub or tree facultatively-dependent on shallow groundwater (Brock 1994, Di Tomaso 1998). Species of Tamarix are native to the Old World, occurring from the Mediterranean across southern Russia to east- ern Asia, and were imported to the USA in the early 1800s for use as ornamentals (Robinson 1965). Tamarisk since has spread beyond cultivation with the most- recent survey estimating the plant’s extent to exceed 350,000 ha in the western USA (Robinson 1965). Tamarisk’s invasiveness, water uptake, and low diversity of dependent wildlife has made it the target of classical biological control (For- nasari 1997). The taxonomy of New World Tamarix is unclear (Brock 1994, Di Tomaso 1998); eight species occur in North America (Baum 1967) with five species, four deciduous and one evergreen, currently recognized in California (Wilken 1993).
Opsius stactogalus is the most abundant insect on tamarisk (Harding 1930, Hopkins & Carruth 1954, Liesner 1971). The insect’s green color and size (length 0.81 mm for first-instar nymphs, 4.5 mm for adult females, Harding 1930) make it difficult to distinguish from tamarisk’s sessile, scalelike leaves (1.5 to 3.5 mm- long in Tamarix ramosissima Ledebour, Wilken 1993). The life cycle of O. stac- togalus is comprised of five nymphal instars, adults, and eggs inserted within stems (Harding 1930). Generations per year number three in Kansas (Harding 1930) and four in New Mexico (Liesner 1971). Like many other Cicadellidae, O. stactogalus is a fluid-feeder, and the aggregate feeding imposed by populations of the leafhopper can reduce tamarisk’s growth (Liesner 1971). Although the
62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
leafhopper was described as a new species in 1907 from collections in Texas, a later comparison of Meditteranean and North American specimens and review of the systematic literature revealed the 1907 species description as synonymous with an original description in Europe during 1847 (Harding 1930). Opsius stactogalus, like its host tamarisk, is native to the Old World.
Opsius stactogalus nymphs and adults are parasitized by the dryinid wasp Gon- atopus spp. (Harding 1930, Liesner 1971). Dryinidae occur worldwide and par- asitize only the homopteran suborder Auchenorrhyncha (Olmi 1984). Dryinid lar- vae are individually enclosed in a sac of exuviae that protrudes between their host’s thoracic or abdominal sclerites. Adult females of many Dryinidae species also are predaceous on their hosts (Olmi 1984). The only published records of Gonato pus parasitizing O. stactogalus in North America are of Gonatopus caroli Olmi and Gonatopus curriei Krombein (Guglielmino & Olmi 1997). One adult female Gonatopus reared from a parasitized O. stactogalus | collected at the present study’s locality was identified as G. caroli (M. Olmi, personal commu- nication).
The effect of parasitism by Gonatopus on the abundance of O. stactogalus is uncertain. Harding (1930) described parasitism rate by Gonatopus as low and unable to significantly affect rapid population increases of O. stactogalus. In con- trast, Liesner (1971) described parasitism rates (30—44% of leafhoppers parasit- ized) high enough to hinder his collecting leafhoppers for experimental trials. Increasing parasitism rate with increasing O. stactogalus density (density depen- dence) may regulate leafhopper populations (Turchin 1995) and decrease their detrimental effect on tamarisk. Parasitism rate decreasing with increasing leaf- hopper density (inverse density dependence) or unrelated to leafhopper density (density independence) would not regulate populations of O. stactogalus and limit the insect’s damage to tamarisk. A synopsis (Stiling 1987) of the relationships between insect-host density and parasitism rate (171 published accounts) found density independence most common (52%) followed by density dependence (25%) and inverse density dependence (23%). This study examines the relation- ship between density of O. stactogalus and parasitism rate by Gonatopus.
MATERIALS AND METHODS
The study site was located at the effluent discharge of the City of Boulder City, Clark County, Nevada, wastewater treatment plant. The treatment plant discharges approximately 2000 m? of secondary-treated municipal effluent per day equally into two parallel unlined southward-flowing channels approximately 0.1 km apart. Effluent discharged to each channel maintains a 2-m wide surface flow extending approximately 3 km downstream. Sustained by the effluent, and bordering both channels, are dense, linear stands of the deciduous tamarisk T. ramosissima al- ternating with clumps of cat-tail, Typhus latifolia L. (Typhaceae). The site lies at an elevation of 610 m within the Mojave Desert; creosote bush, Larrea tridentata (Sessé & Mocino ex Candolle) Coville (Zygophyllaceae), and bursage, Ambrosia dumosa (Gray) Payne (Asteraceae), are the dominant plant species surrounding the effluent-supported vegetation.
Leafhoppers on tamarisk were sampled at 12 points along the west edge of the east channel. Sampling points were located every 80 m along the channel begin- ning 80 m downstream of the effluent discharge. Three samples were taken at
2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 63
each point during the morning on 17-18 Jul, 16—17 Aug, 16—17 Sep, and 20-21 Oct 1999. Each leafhopper sample was taken by bagging a 1-m long tamarisk branch within a plastic trash bag (159 liter, 84 cm X 1.2 m, 76 pm thick) held open with a 52-cm diameter metal hoop. The bag was quickly swept over the branch and slipped off the hoop to constrict the bag around the proximal-end of the branch. The branch then was cut at the top of the bag, pushed into the bag, and the bag closed. Enclosed leafhoppers were killed with an aerosol insecticide (Raid® Concentrated Deep Reach Fogger, S.C. Johnson & Son, Racine, Wiscon- sin) containing 14 g of 1.7% cypermethrin. Approximately 10 min after treatment, the bag was held upright and the branch shaken for 30 sec. Dead leafhoppers shaken from the branch were collected into a 3-dram (15-mm diameter < 65-mm long) patent-lip glass vial. Plant mass supporting the collected leafhoppers was measured (+ 2 g) by removing the branch from the bag and weighing it with a 300-g capacity spring scale. Vial contents were examined with a microscope, and collected leafhoppers were sorted from plant debris and stored in 70% ethanol. Leafhoppers were examined at 30X and the number of parasitized and unpara- sitized nymphs and adults counted. Parasitized and unparasitized O. stactogalus in ethanol were deposited as vouchers at the University of California, Riverside, Entomology Museum.
Opsius stactogalus density was estimated by first determining the relationship between leafhopper abundance and plant mass. Leafhopper abundance (trans- formed In[Y + 0.5]) was regressed (SPSS version 6.1, Chicago, Illinois) against plant mass and life stage, nymph or adult coded as an indicator variable. Two regressions were performed with plant mass in grams and with plant mass trans- formed In(grams). Plant mass transformed In(grams) was selected, due to its great- er partial 7?, as the better linear predictor of leafhopper abundance and used to calculate leafhopper density. The interaction life stage < plant mass was added to the regression to test if the trends in abundance across transformed plant mass differed between nymphs and adults. For graphing, abundances of nymphs and adults in each sample were summed, transformed In(Y + 0.5), and regressed against transformed plant mass.
Parasitism of O. stactogalus was examined by first comparing (x? test) the proportions of parasitized nymphs and adults. Parasitism of nymphs was further examined at two spatial scales, the tamarisk branch and the individual leafhopper. To evaluate exploitation of tamarisk branches by ovipositing Gonatopus, the in- fluence of nymph density (dn[N + 0.5]/In[g]; NW = number of nymphs, g = grams plant mass) on the probability of a leafhopper sample containing at least one parasitized nymph was determined with logistic regression (Neter et al. 1996). A predicted change and 95% confidence interval in the odds of a Gonatopus ex- ploiting versus not exploiting a branch as nymph density increased was calculated from the regression coefficient and its standard error.
To evaluate parasitism of individual leafhoppers on branches exploited by Gon- atopus, the number of parasitized nymphs (G) in leafhopper samples with at least one parasitized nymph was transformed In(G + 0.5) and regressed against nymph density. One outlier leafhopper sample, collected on 17 Aug 1999, containing 35 unparasitized nymphs and 41 parasitized nymphs was excluded from the regres- sion due to its extreme influence (standardized residual = 3.4) on the fitted func- tion. Parasitism rate (R) on branches exploited by Gonatopus was quantified as
64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
«ep, «e@)
=~
Leafhopper Abundance, Y+0.5
ThY = 0.80 - 3.7X
Parasitism Rate, (G+ 0.5)/(N - G+ 0.5)
0.1 02 03 04 05 06 0.7 0.8 Plant Mass (g) Nymph Density, In(N + 0.5)/In(g)
Figures 1-2. Fig. 1 (left). Opsius stactogalus abundance (nymphs + adults) plotted on a logarith- mic scale against Tamarix-branch mass. Curve was fit by linear regression with plant mass transformed In(g). Fig. 2 (right). Parasitism rate of O. stactogalus nymphs versus nymph density (V = nymph abundance) on exploited branches containing = 1 parasitized nymph. Parasitism rate is the ratio of parasitized nymphs (G) to unparasitized nymphs plotted on a logarithmic scale. The size of each data point represents its final weight determined from iteratively reweighted least squares regressions.
the ratio of the number of parasitized nymphs to the number of unparasitized nymphs (N — G), transformed as an empirical logit (Agresti 1990), R = In[(G + 0.5)/(N — G + 0.5)] in each leafhopper sample. Iteratively reweighted least squares regressions (see logistic regression with repeat observations, Neter et al. 1996) of parasitism rate against nymph density were performed to produce a maximum likelihood estimate of the regression coefficient. The regression was unweighted in the first iteration, and the resulting predicted parasitism rates (R) were used to calculate predicted probabilities of parasitism, 7 = [exp(R)/[1 + exp(R)]. A weighted regression then was performed with leafhopper-sample weights calculated with N(m)(1 — tm). Predicted probabilities of parasitism from the weighted regression were used to recalculate weights, and the process was repeated (3 iterations) until the regression coefficient stabilized. Weights deter- mined in the final iteration were adjusted downward (multiplied by 34/51) to restore the unweighted error degrees of freedom. The leafhopper sample identified above as an outlier again was excluded (standardized residual = 3.2) from the analysis. A predicted change in the odds ratio of parasitized nymphs (+ 0.5) to unparasitized nymphs (+ 0.5) as nymph density increased was calculated from the regression coefficient.
RESULTS
Abundance of O. stactogalus was more linearly related to plant mass trans- formed In(grams) (partial 7? = 0.092; F = 29.8; df = 1,285; P < 0.001) (Fig. 1) than to plant mass in grams (partial 7 = 0.065; F = 20.5; df = 1,285; P < 0.001) and was greater (F = 9.1; df = 1,285; P = 0.003) in nymphs (634 leaf- hoppers) than in adults (309 leafhoppers). The partial regression coefficient (0.64 + 0.12 [SE]) for transformed plant mass therefore equaled the exponent of the power function, leafhopper abundance = f (plant mass®?°*). Trends in abundance
2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 65
across transformed plant mass did not differ (F = 0.47; df = 1,284; P = 0.5) between nymphs and adults.
The proportion of O. stactogalus nymphs parasitized (18.0%) by Gonatopus was greater (x? = 41.6; df = 1; P < 0.001) than the proportion of adults para- sitized (2.9%). Tamarisk branches supporting nymphs (99 of 144 total samples) were more likely (x? = 41.4; df = 1; P < 0.001) to be exploited by Gonatopus as nymph density increased. The odds of an ovipositing Gonatopus exploiting versus not exploiting a tamarisk branch increased 2.3-fold (95% confidence in- terval = 1.8 to 3.0) for a 2-fold increase, from 2 to 4 nymphs on a 100 g branch, in nymph density. The increase in the odds of Gonatopus exploiting a branch therefore was approximately proportional to the increase in nymph density.
On branches supporting at least one parasitized nymph (36 samples excluding outlier), increasing nymph density was related to an increase in the number of parasitized nymphs (F = 15.6; df = 1,34; P = 0.004) but to a decrease in the parasitism rate of nymphs (F = 30.9; df = 1,34; P < 0.001) (Fig. 2). The odds of a nymph being parasitized versus unparasitized decreased 38% with a 2-fold increase, from 2 to 4 nymphs on a 100 g branch, in nymph density. Parasitism
of O. stactogalus nymphs on branches exploited by Gonatopus therefore was inverse density dependent.
DISCUSSION
A diminishing rate of increase in O. stactogalus abundance, as the mass of the tamarisk branch supporting the population increased, is predicted by the scaling laws of isometric growth (Schmidt-Nielsen 1984). Tamarisk branches cut for leaf- hopper samples were composed of a central, non-photosynthetic stem supporting smaller, branching stems covered with sessile leaves. The central stem contained vascular tissue transporting xylem and phloem to and from the leaves, the portion of the branch supporting O. stactogalus. The capacity for this transport, limiting the abundance of leaves (and leafhoppers) supported by the stem, would have been proportional to the area of the stem’s cross-section. The mass of the stem, however, would have been proportional to its volume. As the stem grew, its cross- sectional area (A) would have increased more slowly than its mass (MV) according to the equation A = M*? or A = M°°’. This function approximates the observed relationship between O. stactogalus abundance and tamarisk-branch mass.
Greater parasitism of nymphs compared to adults, as observed in O. stacto- galus, also has been reported in other leafhopper species. Dryinids parasitizing eight species of leafhoppers inhabiting forage grass generally were found to attack nymphs more than adults (Buntin 1989). In contrast, Liesner (1971) observed higher rates of parasitism by Gonatopus in O. stactogalus adults (14—100%) than in nymphs (5—54%). Greater parasitism of O. stactogalus nymphs compared to adults is not due to different durations, and therefore exposure times, of the two life stages; both first through fifth instar nymphs and adults live an average of 1 month (Harding 1930). Gonatopus parasitizing O. stactogalus preferred nymphs over adults for hosts.
Increasing O. stactogalus nymph density was associated with increasing prob- ability of a branch being exploited by Gonatopus and with increasing abundance of parasitized nymphs. Gonatopus therefore appears to be attracted to, or arrested by (Vinson 1984), populations of O. stactogalus. The increase in abundance of
66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
parasitized nymphs, however, did not keep pace with the increase in nymph abun- dance, resulting in a parasitism rate that decreased as nymph density increased. Reproductive rates of parasitoids can lag behind those of their hosts due to lower fecundities or long host-handling times (Stiling 1987).
Inverse density-dependent parasitism of O. stactogalus by Gonatopus is con- trary to most insect host-parasite systems that have been studied (Stiling 1987). Decreasing parasitism rate with increasing host density may be common in Dryi- nidae, however, as it also has been observed in several dryinid species parasitizing grassland leafhoppers (Waloff 1975). The only other insect known to parasitize O. stactogalus in North America is the egg parasite Polynema saga (Girault) (Hymenoptera: Mymaridae) (Lawson 1929). Harding (1930) found approximately 25% of O. stactogalus eggs in Kansas parasitized by P. saga, and Liesner (1971) dissected P. (Barypolynema) saga from the leafhopper’s eggs in New Mexico. Opsius stactogalus therefore supports two parasite species separately attacking nymphs and adults or attacking eggs. In comparison, 34 of 50 cicadellid species collected in Kansas grasslands supported two or more species of parasites infect- ing nymphs and adults (Baldridge & Blocker 1980), and eggs of the leafhopper Circulifer tenellus (Baker) in Iran were found to harbor seven species of parasites (Walker et al. 1997). Although caged O. stactogalus were eaten by Chrysopidae larvae (Harding 1930) and spiders (Liesner 1971), and leafhoppers may be eaten by adult female Gonatopus, the contribution of predation to O. stactogalus mor- tality in nature is unknown. Inverse density-dependent parasitism of O. stacto- galus by Gonatopus may not only be due to intrinsic differences between the two species, such as fecundity rate, but also to interactions across several trophic levels (Hare 1992). The leafhopper’s abundant, predictable food source and low diversity of natural enemies may combine to increase its reproductive rate beyond that of its parasite.
Populations of O. stactogalus are not regulated (Turchin 1995) by the inverse density-dependent parasitism imposed by populations of Gonatopus. Uncon- strained population increase by O. stactogalus agrees with descriptions (Harding 1930, Hopkins & Carruth 1954, Liesner 1971) of the leafhopper occurring on tamarisk in high densities. Liesner (1971) also suggested populations of O. stac- togalus can be high enough to reduce tamarisk growth. Reduced growth expect- edly would result from the photosynthate drain imposed on the plant by high densities of the fluid-feeding leafhopper. Lack of population regulation by Gon- atopus permits unhindered herbivory by O. stactogalus, considered beneficial when viewed in the context of biological control against tamarisk. Populations of O. stactogalus and herbivory by the leafhopper may be limited only by temper- ature. Insect development and photosynthate extraction would slow during the fall as temperatures decline and would cease when 7. ramosissima and other decid- uous Tamarix drop their leaves and force O. stactogalus to overwinter in the egg stage within stems.
ACKNOWLEDGMENT
I am grateful to M. Olmi of the Universita degli Studi della Tuscia, Viterbo, Italy, for providing dryinid collection records and for identifying a specimen and to J. D. Hare of the Department of Entomology at UC Riverside for critiquing this manuscript. Appreciation is extended to the City of Boulder City, Nevada,
2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 67
for allowing the study on their lands and providing wastewater treatment plant discharge quantities.
LITERATURE CITED
Agresti, A. 1990. Categorical data analysis. John Wiley & Sons, New York.
Baldridge, R. S. & H. D. Blocker. 1980. Parasites of leafhoppers (Homoptera: Cicadellidae) from Kansas grasslands. J. Kansas Entomol. Soc., 53: 441-447.
Baum, B. R. 1967. Introduced and naturalized tamarisks in the United States and Canada (Tamari- caceae). Baileya, 15: 19-25.
Brock, J. H. 1994. Tamarix spp. (salt cedar), an invasive exotic woody plant in arid and semi-arid riparian habitats of western USA. Chapter 4. pp. 27-44. In deWaal, L. C., L. E. Child, P. M. Wade, & J. H. Brock (eds.). 1994. Ecology and management of invasive riverside plants. John Wiley & Sons, New York.
Buntin, G. D. 1989. Dryinid (Hymenoptera: Dryinidae) parasites of leafhoppers and planthoppers (Homoptera) in forage-type bermudagrass. J. Kansas Entomol. Soc., 62: 602-606.
Di Tomaso, J. M. 1998. Impact, biology, and ecology of saltcedar (Tamarix spp.) in the southwestern United States. Weed Technology, 12: 326-336.
Fornasari, L. 1997. Host specificity of Coniatus tamarisci (Coleoptera: Curculionidae) from France: potential biological control agent of Tamarix spp. in the United States. Environ. Entomol., 26: 349-356.
Guglielmino, A. & M. Olmi. 1997. A host-parasite catalog of world Dryinidae (Hymenoptera: Chry- sidoidea). Cont. Entomol. Int., 2: 165-298.
Harding, L. 1930. The biology of Opsius stactogalus Fieber (Homoptera, Cicadellidae). J. Kansas Entomol. Soc., 3: 7-22.
Hare, J. D. 1992. Effects of plant variation on herbivore-natural enemy interactions. Chapter 12. pp. 278-298. In Fritz, R. S. & E. L. Simms (eds.). 1992. Plant resistance to herbivores and path- ogens: ecology, evolution, and genetics. Univ. Chicago Press, Chicago, Illinois.
Hopkins, L. & L. A. Carruth. 1954. Insects associated with salt cedar in southern Arizona. J. Econ. Entomol., 47: 1126-1129.
Lawson, P. B. 1929. A leafhopper parasite—Polynema saga (Girault) (Hymenoptera, Mymaridae). Ann. Entomol. Soc. Am., 22: 130.
Liesner, D. R. 1971. Phytophagous insects of Tamarix spp. in New Mexico. M.S. Thesis, New Mexico State University, Las Cruces.
Neter, J.. M. H. Kutner, C. J. Nachtsheim, & W. Wasserman. 1996. Applied linear statistical models (4th ed.). McGraw-Hill, Boston, Massachusetts.
Olmi, M. 1984. A revision of the Dryinidae (Hymenoptera). Mem. Amer. Entomol. Inst., no. 37.
Robinson, T. W. 1965. Introduction, spread, and areal extent of saltcedar (Tamarix) in the western States. U.S. Dept. Int., Geol. Surv. Professional Paper, 491-A.
Schmidt-Nielsen, K. 1984. Scaling: why is animal size so important? Cambridge University Press, Cambridge.
Stiling, P. D. 1987. The frequency of density dependence in insect host-parasitoid systems. Ecology, 68: 844-856.
Turchin, P. 1995. Population regulation: old arguments and a new synthesis. Chapter 2. pp. 19-40. In Cappuccino, N. & P. W. Price (eds.). 1995. Population dynamics: new approaches and synthesis. Academic Press, New York.
Vinson, S. B. 1984. Parasitoid-host relationship. Chapter 8. pp. 205-233. Jn Bell, W. J. & R. T. Cardé (eds.). 1984. Chemical ecology of insects. Chapman & Hall, London.
Walker, G. P, N. Zareh, I. M. Bayoun, & S. V. Triapitsyn. 1997. Introduction of western Asian egg parasitoids into California for biological control of beet leafhopper, Circulifer tenellus. Pan- Pacific Entomol., 73: 236-242.
Waloff, N. 1975. The parasitoids of the nymphal and adult stages of leafhoppers (Auchenorryhncha: Homoptera) of acidic grassland. Trans. R. Entomol. Soc. London, 126: 637-686.
Wilken, D. H. 1993. Tamaricaceae: tamarisk family. p. 1080. In Hickman, J. C. (ed.). 1993. The Jepson manual: higher plants of California. Univ. Calif. Press, Berkeley.
Received 10 Mar 2000; Accepted 26 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(2): 68-70, (2001)
TAENIOGONALOS RAYMENTI CARMEAN & KIMSEY (HYMENOPTERA: TRIGONALIDAE) REARED AS A HYPERPARASITE OF STURMIA CONVERGENS (WEIDEMANN) (DIPTERA: TACHINIDAE), A PRIMARY PARASITE OF DANAUS PLEXIPPUS (L.) LEPIDOPTERA: NYMPHALIDAE)
ANTHONY R. CLARKE! AND MyYRON P. ZALUCKI
Department of Zoology and Entomology, The University of Queensland, Queensland, 4079, Australia
Abstract.—Taeniogonalos raymenti is confirmed as a hyperparasitoid of the tachinid Sturmia convergens which parasitises larval Danaus plexippus. Trigonalids are indirect parasitoids and in this case we have direct evidence that wasp eggs must have been laid on the caterpillar’s host plant, Asclepias fruticosa, before the secondary host, but not necessarily before the primary tachinid host, was present. Levels of hyperparasitism during our sampling period were very low at less than two percent.
Key Words.—Insecta, Sturmia, Tachinidae, Asclepias.
Trigonalid wasps are unique among the parasitic Hymenoptera as they oviposit onto foliage and the eggs are subsequently injested, along with the foliage, by larval sawflies or herbivorous caterpillars. The host relationships of larval trigon- alids are poorly known, but most appear to be either primary parasitoids of larval sawflies, or hyperparasitoids of hymenopteran and tachinid parasitoids (Weinstein & Austin 1991). The family has a cosmopolitan distribution, but is quite small with only 100 known species. The biology, host relationships and systematics of trigonalids are reviewed by Weinstein & Austin (1991) and Carmean & Kimsey (1998). One of the major messages in both those reviews is that trigonalids remain a very poorly known group, with little or no biological information for most species. This note supplies a new host record for a trigonalid species from South- east Queensland.
MATERIALS AND METHODS
Collection and Rearing.—As part of wider study, 222 wild, 5th instar Danaus plexippus (L.) caterpillars were collected from four sites in South-east Queensland from February to November 1998. Collections were made at Mt Crosby (27°32! S, 152°49’ E), The University of Queensland Veterinary Research Farm Pinjarra Hills (= Moggill Farm) (27°35' S, 152°53' E), Dunn’s Creek via Beenleigh (27°43' S, 153°09' E) and Pine River via Boondal (27°17' S, 152°55’ E).
Caterpillars were collected from either Ascle pias fruticosa L. or A. curassavica L., returned to the laboratory and reared individually on cut foliage of the host species they were collected on until they pupated. A further 33 caterpillars were obtained by placing D. plexippus eggs onto A. fruticosa plants, which did not carry existing D. plexippus eggs or larvae, at Moggill Farm and then immediately
‘Current Address: Australian School of Environmental Studies, Griffith University Nathan Campus, Qld, 4111.
2001 CLARKE & ZALUCKI: TAENJIOGONALOS REARED 69
Table 1. Fate of fifth instar Danaus plexippus (L.) larvae collected from sites in South-east Queens-
land. (P.1. = Paradrino laevicula Mesnil, S.c. = Sturmia convergens (Weidemann), W.s. = Winthemia sumatrana (Townsend).
No. of adult No. of
No. of larvae No. of larvae tachinids emerging trigonalids Collection site Date collected parasitised (P.1./S.c./W.s.) reared
Wild collections
Mt Crosby Feb 98 21 18 31 (25/1/5) 0 Dunn’s Ck Apr 98 31 18 25 (21/4/0) 0 Pine River May 98 2. 12 19 (16/3/0) 0 Pine River June 98 6 3 0 0 Dunn’s Ck June 98 22 19 36 (32/4/0) 0 Dunn’s Ck June 98 Pal 15 21 (21/0/0) 0 Dunn’s Ck June 98 40 20 24 (22/1/1) 0 Dunn’s Ck Nov 98 23 22 30 (17/13/0) 0 Moggill Farm Nov 98 21 19 40 (10/26/4) 1 Mt Crosby Nov 98 10 5 11 (6/5/0) 0 “Bag’’ reared
Moggill Farm Nov 98 33 33 48 (0/48/0) 6
bagging the plants with insect proof gauze. Caterpillars were harvested as 5th instars and then reared as above.
Identification and Voucher Material.—Trigonalids were identified by J. C. Car- dale, Australian National Insect Collection, CSIRO Entomology, Canberra. Voucher material has been deposited with the University of Queensland Insect Collection. Tachinids were identified by B. Cantrell, Queensland Department of Primary Industries, Brisbane. Two species of tachinid were routinely reared from monarch butterfly caterpillars during our research. We were able to determine which tachinid was the trigonalid host from the remnant pupal cases, which are noticeably different between the two tachinid species.
RESULTS
Danaus plexippus caterpillars were found to be heavily parasitised by tachinids, most commonly Sturmia convergens (Weidemann) and Paradrino laevicula Mes- nil. Both have been previously recorded at primary parasites of D. plexippus in South-east Queensland (Zalucki 1981). As we will be reporting on primary par- asitism elsewhere, we will not here discuss it further.
Seven trigonalid wasps emerged from seven tachinid pupal cases which had emerged from seven individual caterpillars. These were identified as individuals of Taeniogonalos raymenti Carmean & Kimsey (previously T. tricolor Rayment, see Carmean & Kimsey 1998). All but one of these specimens were reared from a single collection of caterpillars made from plants which had been bagged im- mediately after placing butterfly eggs upon them (Table 1). In all cases the tachinid host, based on pupal case identification, was S. convergens. Because plants had been bagged immediately following artificial butterfly egg placement, it confirms that both S. convergens and T. tricolor placed their eggs on the host plant (A. fmtticosa) prior to the presence of the primary host insect. The identity of the tachinid from which the seventh wasp was reared was not recorded, but the host plant was again A. fruticosa.
70 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Overall hyperparasitism rate was only 1.8% (n = 397 tachinids reared), but for the case where 6 wasps were reared from a single collection, the hyperparasitism rate was 13% (n = 46 tachinids reared).
DISCUSSION
Taeniogonalos is the most widely distributed genus of the Trigonalidae and Species in the genus have been reared previously as primary parasitoids of saw- flies, or a hyperparasitoids of tachinid and ichneumonid parasitoids of lepidoptera (Carmean & Kimsey 1998). This is the first rearing and oviposition substrate record for JT. raymenti, but it is not the first time the genus Sturmia or the sub- family Danainae have been associated as primary and secondary hosts of this genus. Hirai & Ishi (1995) recorded T. fasciata (Strand) (previously Poecilogon- alos fasciata) as hyperparasitising Sturmia bella (Meigen), a parasite of Parantia sita (Moore) (the tiger chestnut butterfly), in Japan.
ACKNOWLEDGMENT
This work was undertaken while Anthony Clarke was in receipt of a University of Queensland New Staff Grant and an Australian Research Council Large Grant. Dr. Jan Green and Ms. Michelle Tung assisted in collecting and rearing of the specimens. We thank Ms. Jo Cardale and Dr. Bryan Cantrell for the identifications.
LITERATURE CITED
Carmean, D. & L. Kimsey 1998. Phylogenetic revision of the parasitoid wasp family Trigonalidae (Hymenoptera). System. Entomol., 23: 35-76.
Hirai, N. & M. Ishi 1995. Host relationships of the tachinid fly, Sturmia bella (Diptera: Tachinidae), and the trigonalid wasp, Poecilogonalos fasciata (Hymenoptera: Trigonalyidae), parasitizing the Chestnut Tiger butterfly, Parantica sita (Lepidoptera: Danaidae). Appl. Entomol. Zool., 30: 241-244.
Weinstein, P. & A. D. Austin 1991. The host relationships of trigonalyid wasps (Hymenoptera: Tri- gonalyidae), with a review of their biology and catalogue to world species. J. Nat. Hist., 25: 399-433.
Zalucki, M. P. 1981. Temporal and spatial variation of mortality in Danaus plexippus L. Aust. Entomol. Mag., 8: 3-8.
Received 20 Mar 2000; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(2): 71-78, (2001)
NEW SPECIES OF CERACLEA (TRICHOPTERA: LEPTOCERIDAE: ATHRIPSODINI) AND A FIRST RECORD OF ADICELLA (TRICHOPTERA: LEPTOCERIDAE: TRIAENODINI) FROM FAR EASTERN RUSSIA
TATYANA I. AREFINA! AND JOHN C. MORSE?
‘Institute of Biology and Soil Sciences, Vladivostok, Russia Department of Entomology, Clemson University, Clemson, South Carolina 29634-0365, U.S.A.
Abstract.—Three new species of the genus Ceraclea, belonging to the subgenus Athripsodina: C. (A.) affinis, C. (A.) bilobulata and C. (A.) breviramosa from the southern Far East of Russia are described and illustrated. The genus Adicella, represented by a new species, is newly recorded for the Russian fauna.
Key Words.—Trichoptera, Leptoceridae, Ceraclea, Adicella, new species, new record, Far East- ern Russia.
Until now, 9 genera and 59 species of the family Leptoceridae are known from the Russian Far East (Arefina 1997; Vshivkova et al. 1997; J. C. Morse, L. Yang, and I. M. Levanidova, unpublished data). During a survey of the caddisflies (Tri- choptera) of the Ussuri River Basin, three Ceraclea species were collected that are new to science. Furthermore, a representative of the genus Adicella, which has not been previously known in Russia, was found.
The types of the new species are preserved in alcohol and held in the collections of the Institute of Biology and Soil Sciences, Russian Academy of Sciences,
Vladivostok. In the present work, the terminology generally follows that in the revision by Yang & Morse (1988).
CERACLEA (ATHRIPSODINA) DILUTA Group Ceraclea (Athripsodina) affinis, NEW SPECIES (Figs. 1-7)
Types.—Holotype male: KHABAROVSK TERRITORY (RUSSIA), Birushka
River, Ussuri River Basin, 23 Jul 1996, T. Arefina. Paratype: 1 female, same data as holotype.
Description—Length of forewing: male—7.9 mm, female—8.1 mm. Head and body with white and brown hairs mixed. Vertex of head, thorax and coxae brown, abdomen and legs lighter. Wings yellowish brown with setae darker.
Male Genitalia (Figs. 1-4).—Tergum IX (IX) with pair of small papillae near poster meson. Preanal appendages (pre. app) subtriangular, fused basally, each with acute apex. Tergum X (X) composed of triangular median lobe and pair of lateral lobes, each with blunt apex in dorsal and lateral views, lateral lobes extending slightly beyond median lobe. Main body of each inferior appendage (inf. app) Straight in lateral view with subapicodorsal lobe (sap. do) bent caudad; basoventral lobe (bv) of each inferior appendage nearly half as long as main body, obtusely angled about 110° from main body, stout at basal half then abruptly tapering to digitate apex; harpago (har) tiny; mesal ridge of inferior appendage (mes.rdg) produced in small, nearly trapezoidal process located slightly lower than harpago. Phallus strongly curved ventrad and constricted ventrally two-thirds distance from base; both paramere
spines (par) seta-like with right spine positioned near apex, left spine almost straight and situated in anterior half of phallus.
72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
2
Figures 1-4. Male genitalia of Ceraclea (A.) affinis NEW SPECIES in lateral (1) and dorsal (2) views; left inferior appendage (3) in caudal view; phallus (4) in lateral view. Abbreviations: bv = basoventral lobe of an inferior appendage; har = harpago; infapp = inferior appendage; mes.rdg = mesal ridge of an inferior appendage; par = paramere spines; pre.app = preanal appendage; sap.do = subapicodorsal lobe of an inferior appendage; IX, X = abdominal segments IX and X.
Female Genitalia (Figs. 5—7).—Tergum IX rounded apically, with pair of small subdorsal processes. Preanal appendages (pre.app) short, as long as broad, covered with short setae. Lamellae (lam) parallel- sided in dorsal and ventral views, semicircular in lateral view, setose ventrally. Each gonopod plate (go.pl; e.gon.IX of Nielsen 1980) with triangular caudal projection, plates approximate for most of their length, with triangular excisions between their caudal projections; gonopod plates slightly con- cave in middle with prominent mesal and lateral edges. Spermathecal sclerite (sp.sc) long, V-shaped, with lateral arms curved mesad near caudal end. Pair of broad sclerotized plates (sc.pl) suspended above spermathecal sclerite and extending cephalad nearly to two-thirds length of segment VUI.
Immature Stages.—Unknown.
Diagnosis.—Ceraclea affinis belongs to the C. (A.) diluta Group, which pres- ently includes C. diluta (Hagen), C. perplexa (McLachlan) and C. trilobulata Morse, Yang, and Levanidova. Male genitalia of the new species most closely resemble those of C. trilobulata in the trilobate tergum X, in the shape of its inferior appendages, and in the presence of a ventral constriction of the phallus. It differs from C. trilobulata in the much longer base of the phallus, with the position of the ventral constriction nearly two-thirds of the distance from the based of the phallus; in the apparently longer middle lobe of tergum X; and in the thicker base of each inferior appendage. Because the females of C. perplexa and C. trilobulata are unknown, it is not possible to separate them from the female
2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 73
pre.ap 7
(ox
Figures 5—7. Female genitalia of Ceraclea (A.) affinis NEW SPECIES (holotype) in lateral (5), dorsal (6) and ventral (7) views. Abbreviations: go.p! = gonopod plate of gonopod IX; lam = lamella; pre.app = preanal appendage; sc.pl = sclerotized plate of spermathecal sclerite; sp.sc = spermathecal sclerite.
of the new species; however, the apicolateral lamellae of C. affinis are shorter than those of C. diluta.
Distribution.—Known only from the type locality in Khabarovsk Territory (Russia).
Etymology.—Latin, “‘similar,’’ with reference to the resemblance of the new species with C. trilobulata.
CERACLEA (ATHRIPSODINA) ANNULICORNIS Group Ceraclea (Athripsodina) bilobulata, NEW SPECIES (Figs. 8-15) Types.—Holotype male) KHABAROVSK TERRITORY (RUSSIA). Ussuri River near Zabaikalskoe Village, 23 Jul 1996, T. Arefina. Paratypes: 7 males, same data as holotype.
Description—Length of forewing: male—5.5—6.2 mm, female—5.3-5.9 mm. Head with mostly
74 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Figures 8-11. Male genitalia of Ceraclea (A.) bilobulata NEW SPECIES (paratype) in lateral (8) and dorsal (9) views; left inferior appendage (10) in caudal view; phallus (11) in lateral view.
white hairs. Vertex of head and thorax yellow-brown, abdomen lighter, nearly white ventrally. Wings straw yellow with yellow-brown setae.
Male Genitalia (Figs. 8—15)—Tergum IX protruding at apical center. Preanal appendages very short and broad in dorsal view, separated basally, slightly shorter then tergum IX to slightly longer. Tergum X extending far beyond preanal appendages, broad basally, divided into two lobes nearly to base, each lobe with rounded apex; dorsal edge of tergum straight in lateral view, ventral edge of tergum upturned from middle, apex rounded. Main body of each inferior appendage slender, both harpago and subap- icodorsal lobe short; harpago nearly as long as semimembranous subapicodorsal lobe to slightly longer; basoventral lobe of each inferior appendage vestigial, very short, with single spine directed somewhat mesad; mesal ridge without project and bearing only one normal seta; main body with variable cau- dolateral lobe bearing two large setae. Phallus curved ventrad, with sclerotization extended full length ventrally; anterior portion shorter and broader than posterior portion; two paramere spines present, left paramere spine about same size as right, but retracted somewhat more cephalad.
Remarks.—Different individuals of the species vary in the shape of the preanal and inferior appendages and in size and location of the paramere spines of the phallus. In spite of the variability of these characters, the available specimens generally agree with the foregoing description.
Female and Immature Stages.—Unknown.
Diagnosis.—The male of this species resembles those of C. sibirica (Ulmer) and C. hastata (Botosaneanu) in the presence of a spine at the basoventral position of the inferior appendages; in the mesal ridge of each inferior appendage without
2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 75
14 15
Figures 12-15. Male genitalia of Ceraclea (A.) bilobulata NEW SPECIES (paratype) in lateral (12) and dorsal (13) views; left inferior appendage (14) in caudal view; phallus (15) in lateral views.
any projection; in tergum X lacking lateral processes; and in the sclerotization extending the full length of the phallus ventrally. However, tergum X is divided into two lobes in the new species whereas C. sibirica and C. hastata have no excision on tergum X. Ceraclea bilobulata differs from both C. sibirica and C. hastata and other known males of the C. (A.) annulicornis Group in the short and basally separated preanal appendages, in the very short basoventral lobe of each inferior appendage, in the mesal ridge of the inferior appendages having only one seta, and in possessing two phallic paramere spines nearly equal in length.
Distribution.—Known only from the type locality in Khabarovsk Territory (Russia).
Etymology.—Latin, “‘two-lobed,”’ with reference to the shape of tergum X in the male genitalia.
CERACLEA (ATHRIPSODINA) RIPARIA Group Ceraclea (Athripsodina) breviramosa, NEW SPECIES (Figs. 16—23)
Types.—Holotype male: KHABAROVSK TERRITORY (RUSSIA). Ussuri River near Zabaikalskoe Village, 23 Jul 1996, T. Arefina. Paratypes: 24 females, same data as holotype; Khabarovsk Territory (Russia), Kiya River, Ussuri River Basin, Ekaterinoslavka Village vicinity, 26 Jul 1996, T. Arefina, 4 females; Pri-
76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Figures 16-20. Male genitalia of Ceraclea (A.) breviramosa NEW SPECIES (holotype) in lateral (16) and dorsal (17) views; left inferior appendage (18) in caudal view; harpago and subapicodorsal lobe of left inferior appendage (19) in caudal view; phallus (20) in lateral view.
morye Region (Russia), Kabarga River, Ussuri River Basin, 28 Jul 1998, T. Tiu- nova, 2 females.
Description—Length of forewing: male—5.8 mm, female—5.5—6.0 mm. Head with white hairs on middle of front and vertex, side hairs brownish. Head and thorax yellow-brown, abdomen whitish, abdominal dorsum pale brown. Wings straw yellow with yellow-brown setae.
Male Genitalia (Figs. 16—20).—Tergum IX protruding at apical center. Preanal appendages nearly as long as broad, separated basally, about as long as tergum IX, obliquely truncate apically. Tergum X about 2.5X as long as preanal appendages, broad at base, tapering to blunt apex, with V-shaped cleft apically one-eighth of its length; tergum X with pair of smaller papillae near middle of tergum. Basoventral lobe of each inferior appendage shorter than main body of appendage, directed somewhat mesad, bearing single spine subbasally; triangular projection above basoventral lobe, nearly as long as spine in lateral view. Main body of appendage slender; harpago slightly longer than subapicodorsal lobe, each harpago setose, with two small spines at apex; each subapicodorsal lobe with several long setae along dorsal surface and membranous apex bearing two setae. Mesal ridge of each inferior appendage with two normal setae. Phallus strongly curved ventrad about half distance from base, with sclerotization extended full length ventrally; anterior portion of phallobase slightly broader and longer than posterior portion in lateral view. Paramere spines aligned, with apex of left spine slightly inserted in elliptical opening of base of right spine, as typical for Group; left spine with short and straight
2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 77
i eosSree§ eye
eee SS -as
Figures 21-23. Female genitalia of Ceraclea (A.) breviramosa NEW SPECIES (paratype from Ussuri River near Zabaikalskoe Village) in lateral (21), dorsal (22) and ventral (23) views. Abbrevi- ation: d.rdg = diagonal ridge of a gonopod plate.
dorsomedial projection; apical half of left spine sinuous, with upturned tip; right spine straight, curved ventrad at three-quarters distance from base, with tip turned slightly outwards, to right.
Female Genitalia (Figs. 21—23).—Preanal appendages broad but very short, as typical for Group. Lamellae setose ventrally, each with broadly rounded apex in lateral view. Gonopod plates widely separated anteriorly, not concave, each with well-sclerotized diagonal ridge (d. rdg); each plate with two caudal projections: mesal projection narrow and elongate, lateral projection rather large, oval in lateral and ventral views. Anterior part of the spermathecal sclerite narrow, broadens in middle, convex laterally in posterior part.
Immature Stages.—Unknown.
Diagnosis.—The male genitalia of this species resemble those of most species of the C. (A.) riparia Group in possessing two phallic paramere spines which are aligned, with the apex of the left spine inserted in an opening of the base of the right spine, but in this new species the left spine has a dorsomedial projection and an upturned tip. This is the only species with the C. (A.) riparia Group whose male: (1) lacks a pair slender lateral processes of tergum X; (2) has the inferior appendages each with such a short basoventral lobe; and (3) has the mesal ridge
78 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
of each inferior appendage bearing only two setae. The female of this species resembles those of C. riparia (Albarda), C. yangi (Mosely) and C. nankingensis (Hwang) in the shape of the caudal projections of the gonopod plates and of the spermathecal sclerite, but it differs from all known species of the C. (A.) riparia Group in possessing a diagonal ridge of each gonopod plate.
Distribution.—Known only from the type localities in Khabarovsk Territory and the Primorye Region (Russia).
Etymology.—Latin, “‘short-branched,”’ with reference to the short basoventral lobe of each inferior appendage in the male genitalia.
NEw RECORD Adicella sp. n. (Yang and Morse, unpublished data)
Material Examined. —KHABAROVSK TERRITORY (RUSSIA). Kiya River, Ussuri River Basin, Yekaterinoslavka Village vicinity, 26 Jul 1996, T. Arefina, 2 males.
Distribution.—Khabarovsk Territory (Russia), China.
ACKNOWLEDGMENT
The work was supported by the Russian Fund of Fundamental Investigations, grant 96-04-50388.
LITERATURE CITED
Arefina, T. I. 1997. Three new to the Russia species of the caddisflies (Trichoptera: Leptoceridae). Far Eastern Entomol., 47: 18.
Nielsen, A. 1980. A comparative study of the genital segments and genital chamber in the female Trichoptera. Det Kongelige Dansk Videnskabernes Selskab Biologiske Skrifter, 23: 1-200.
Vshivkova, T. S., J. C. Morse & L. Yang. 1997. Fam. Leptoceridae. pp. 154—202. In Lehr, P. A. (ed.). Key to the insects of Russian Far East V. Trichoptera and Lepidoptera Pt. 1, Vladivostok.
Yang, L. & J. C. Morse. 1988. Ceraclea of the People’s Republic of China (Trichoptera: Leptoceridae). Contr. Amer. Entomol. Inst., 23: 1-69.
Received 3 Aug 1999; Accepted 6 Jul 2000.
PAN-PACIFIC ENTOMOLOGIST 71(2): 79-89, (2001)
A FURTHER CONTRIBUTION TO THE AUSTRALIAN SYSTEMATICS OF THE TRIBE COLPURINI (HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE)
HARRY BRAILOVSKY
Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Aut6noma de México, Apdo Postal 70153, México D.EF, 04510, México
Abstract.—One new genus (Weirhygia) and four new species (Weirhygia faceta, Grosshygia formosa, Grosshygia lepida, and Grosshygia pisina) from Australia are described in the tribe Colpurini (Coreidae). A revised key to all known genera and species is included. Dorsal habitus illustrations and drawings of male genital capsule and female genital plates are provided.
Key Words.—Insecta, Hemiptera, Heteroptera, Coreidae, Colpurini, new genus, new species, Australia.
The Australian species of the tribe Colpurini have recently been reviewed by Brailovsky (1993), Brailovsky & Barrera (1996) and Steinbauer & Clarke (1996). During a visit to Queensland Museum, Brisbane, Australia, I found, a series of specimens of Colpurini that I first thought to be a previous known species of Grosshygia Brailovsky and Grosshygioides Brailovsky. A preliminary check of the head, pronotum, development of the hemelytra, general shape of the male genital capsule and abdominal sternite VII of the female revealed a new genus, and species and three new species of Grosshygia.
This additional information is made possible largely by the extensive collec- tions of G. B. Monteith and his colleagues.
One of the most striking developments in australian colpurini systematics has been the discovery of an extensive fauna living on the ground of tropical rainforest at high and low elevations of northern and northeastern Queensland. They live associated with piles of freshly dead leaves from recently fallen trees.
REVISED KEY TO AUSTRALIAN COLPURINI
1. Each side of head immediately in front of eye with long pointed spine Ochs Ae ete een ne aoa Sint, nha A ss Ae eS NE Pachycolpura manca Breddin 1’. Sides of head in front of eyes unarmed 2. Tylus projecting as single, large, acute spine 2'< Tylus globose, truncated or ‘Difid’ 2.5 5 2c. sate pe wnecace ee eres, 25) wae eed 5 3. Femora unarmed; callar region of pronotum conspicuously convex; pron- otal disc behind midline with transverse wrinkle; male genital capsule with small median projection ........... Acanthotyla fasciata (Walker) 3’. Femora strongly armed with long, sharp ventral spines; callar region of pronotum weakly convex to flat; pronotal disc without transverse wrin- kle; posteroventral edge of male genital capsule without median pro- ACCUIOM Ve YR, hee gen ad gt Eun eg Oene fore LD Snes 8 ee 4 4. Mandibular plate armed with short projection; posteroventral edge of male genital capsule convex, obtusely rounded ... Agathyrna praecellens Stal
80
4’.
125
Pass
U3:
13%:
14.
14’.
$5:
share
16.
16’.
THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Mandibular plate unarmed; posteroventral edge of male genital capsule
elongate and bifurcate ..... Woodwardhysgia bifida Brailovsky (in part) Antenniférous*tubercles armed” 5) 0520. h ake: oar needeees Sa peo ae ed 6 Antenniferous tubercles unarmed ................0 0c eee eee ees 14 Bucculae armed with obvious spine near middle third ............... 7
2 Bucculaeniitormiy-roundeg: (022s Oe ee vente ee ee 8
Rostrum reaching middle third of abdominal sternite V; body robust, longer than 10.00 mm; frontal angles of pronotum conspicuously prominent; scutellum as wide or wider than long .................
en rr ee ehh, aera ea a ore ra ie Acantholybas kirkaldyi Bergroth
. Rostrum reaching posterior margin of abdominal sternite IV; body shorter
than 9.00 mm; frontal angles of pronotum scarcely exposed; scutellum
clearly longer than wide ............. Acantholybas brunneus Breddin Micropterous, hemelytral membrane reduced to small flap; ocelli incon-
SPLCM OUST aly ye ee cote ee Cra eee te ea ewer a pee ele oe BUN eee ne an 2
. Macropterous to submacropterous, hemelytral membrane well developed;
ocelli clearly developed ....... Pachycolpuroides monteithi Brailovsky Head dorsally flat; abdominal sternite VII of female without plica or fissura ... Weirhygia faceta Brailovsky NEW GENUS NEW SPECIES
. Head dorsally convex; abdominal sternite VII of female with plica and
TUSSUGE Ae nase = Eke p iri ONS | ACE Rhea de Se aligt leche Foye Wc eects cade nese ne Maa 10 Apex of scutellum globose ............... Grosshygia nigra Brailovsky
= Apex Of seutelim SUDSCULED 22.0.8 ned lisiicdes sere) cwceeede 2 arena ee ates 11
Antenniferous tubercles each with external lobe arcuate, recurved, con- verging anteriorly, almost touching the basal joint of antennal segment De ese etch ge ea ee te ete Grosshygia lepida Brailovsky NEW SPECIES
. Antenniferous tubercles each with external lobe obliquely projecting, di-
VETSTI Seal LOLI@TU Ys ala ah Te Be ek leer the 5 btn he iden Seether aes too HARE 12 Head dorsally with vertex uniformly convex, without conical lobes; he-
melytral-menm Grane a WSent, oes A... ches ts eveyeet tei sa fer eee Ae Sori gc pect chap yt Nees c eeu RR Grosshygia pisina Brailovsky NEW SPECIES Head dorsally with a transverse depression separated into two conical
elevations; hemelytral membrane reduced to small flaps ........... 13 Antenniferous tubercles each armed with long lobe; antennal segment II
longer than 1.96 mm ................ Grosshygia lobatula Brailovsky Antenniferous tubercles each armed with short lobe; antennal segment II
shorter than 1.73 mm .............. Grosshygia monticeps Brailovsky Head dorsally convex; apex of scutellum globose ................... BUSINES ys tl A oe ee tre Grosshygia formosa Brailovsky NEW SPECIES Head dorsally flat; apex of scutellum subacute ..................... 15 Abdominal sternite VII of female without plica or fissura; frontal angles
of pronotum rounded, blunt, not produced ....................... 16 Abdominal sternite VII of female with plica and fissura; frontal angles
of pronotum produced forward as conical teeth .................. 20 External edge of gonocoxa I in lateral view with upper half conspicu-
ously exposed and lower half projected in a medium-sized convex lobe
2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 81
17. Posteroventral edge of male genital capsule with small V-shaped con- cavity, laterally enclosed by two shorter arms; gonocoxa I in caudal WITS Wi EC LOSE tee pbsanete nit Anke cop cc epee as Sciophyroides sortita (Horvath)
17’. Posteroventral edge of male genital capsule with U-shaped concavity, enclosed by two lateral medium-sized robust arms; gonocoxa I in cau- dal-view-opened. <9 in ee.. 2 eee, Scio phyroides sulcicrus (Breddin)
18. Body length longer than 10.30 mm; posteroventral edge of male genital capsule with large U-shaped concavity, laterally enclosed by two
strong divergent arms ............ Sciophyrella australica (Brailovsky) 18’. Body length shorter than 10.10 mm; posteroventral edge of male genital CASH le SHOLsAS TAB OVER. sacra ee AOR ay RR I cal tk Seger 19
19. Posteroventral edge of male genital capsule produced into medium-sized and robust lateral projections, enclosing deep U-shaped concavity; gonocoxa I in lateral view narrow .... Sciophyrella diminuta (Horvath)
19’. Posteroventral edge of male genital capsule with small V-concavity en- closed by two short globose arms; gonocoxa I in lateral view enlarged oP at bp hl dere, oe A Sciophyrella minuscula Brailovsky & Barrera
20. Mandibular plate armed with large prominent tubercle; pronotal disc with deep longitudinal furrow along midline; hemelytra truncate, membrane PLO s| ee eRe, Bienes Sota Grosshygioides mandibularis Brailovsky
20'. Mandibular plate unarmed; pronotal disc flat, without midline furrow; macropterous; hemelytral membrane well developed .............. Zz
21. Femora armed with two rows of ventral spines: tylus apically bifid
Rete ee Seon yAr cae ratte Oe Woodwardhygia bifida Brailovsky (in part) 21’. Femora unarmed; tylus apically globose ..................0000000e-
et ie Oa Lo rere ae Hygia (Australocolpura) sandaracine Brailovsky
WEIRHYGIA BRAILOVSKY, NEW GENUS
Type species.—Weirhygia faceta Brailovsky, NEW SPECIES.
Description—Head longer than wide (across eyes), pentagonal, nondeclivent, and dorsally flat; tylus unarmed, apically globose, extending anteriorly to and laterally higher than juga; juga unarmed, thick- ened, apically rounded, shorter than tylus; antenniferous tubercle armed with raised lobe, diverging anteriorly and rounded to quadrate apically; sides of head in front of eyes unarmed; genae and man- dibular plates unarmed; antennal segment I robust, thickest, slightly curved outward; segments II and Ill cylindrical, and slender; segment IV fusiform; antennal segment II longest, I shortest and III longer than IV; ocelli not raised; preocellar pit obliquely deep; eyes small, globular; postocular tubercle protuberant, globose; buccula rectangular, short, elevated, not projecting beyond antenniferous tubercle, without teeth; rostrum long, reaching posterior margin of abdominal sternite VI or anterior margin of VII; rostral segment IV longest, I shortest, and II longer than II. Thorax. Pronotum: Wider than long, trapeziform, non declivent, and bilobate; anterior lobe longer than posterior lobe, each with antero- lateral borders barely convex; collar wide; frontal angles projecting forward as conical teeth; humeral angles rounded, projected into rounded lobes, elevated, directed outward, higher than posterior pronotal disc; posterolateral and posterior borders almost straight; callar region weakly convex, separated me- dially by deep longitudinal furrow, which extends to posterior margin. Anterior lobe of metathoracic peritreme elevated, reniform, posterior lobe sharp, small. Legs: Unarmed; tibiae cylindrical, with lon- gitudinal sulcus indistinct. Scutellum: Triangular, flat, wider than long, with apex subacute. Hemelytra: Coleopteroid; clavus and corium fused into coriaceous pad, wings meeting each other along midline; hemelytral membrane reduced to small flap, reaching middle third of abdominal tergite III, thus leaving the abdominal terga exposed. Abdomen: Connexival segments practically at same level as abdominal segments; posterior angle of connexivum extending into short and robust spine; abdominal sterna with
82 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
medial furrow extending to posterior border of sternite VI. Integument: Body surface dull; ventral face of head, posterior lobe of pronotum, scutellum, clavus, corium, propleura, mesopleura and me- tapleura, acetabula and abdomen with scattered punctures, each puncture with short decumbent silvery bristle-like hair, intermixed with long erect bristles on antennal segments, legs, and abdominal sterna; dorsal surface of head, anterior lobe of pronotum, and connexival segments impunctate.
Male Genitalia—Genital capsule. Posteroventral edge simple, transversely barely arcuate, with shallow notch at midline (Fig. 3).
Female Genitalia.—Abdominal sternite VII without plica and fissura; gonocoxae I enlarged antero- posteriorly, in lateral view with external face obliquely straight, in caudal view open; paratergite VIII short, almost square, with spiracle visible; paratergite LX larged than paratergite VIII (Fig. 4).
Diagnosis.—The reduction of wings, the prominent postocular tubercle, the pronotal disk with deep midline furrow, and tylus apically globose might suggest a relationship with Grosshygioides Brailovsky.
In Weirhygia, the antenniferous tubercle armed, the mandibular plate unarmed, posterior angle of connexivum extending into short and robust spine, abdominal sternite VII of female without plica and fissura, and frontal angles produced for- ward into conical teeth. In Grosshygioides, the antenniferous tubercle is unarmed, the mandibular plate armed, posterior angle of connexivum unarmed, abdominal sternite VII of female with plica and fissura, and frontal angles projecting forward as small rounded teeth.
Distribution.—Only known from Australia.
Etymology.—Named for Tom Weir, distinguished Australian entomologist.
WEIRHYGIA FACETA BRAILOVSKY, NEW SPECIES (Figs. 3, 4, 7)
Types.—Holotype male: Australia. NE Queensland, Isley Hills, 1050 m, 17°03’ S 145°42' E, 30 Nov 1993, Cook, Monteith and Janetzki. Deposited in Queensland Museum, Brisbane, Australia. Paratypes: 1 male, 1 female; data: same as holotype. Deposited in the ‘‘Colecci6n Entomoldgica del Instituto de Biologia, UNAM, México, and Queensland Museum, Brisbane, Australia. 1 male; data: Australia: NE Queensland, Upper Isley Ck., 750 m, 17°03’ S 145°41’ E, 29 Nov 1993, Monteith, and Janetzki. Deposited in Queensland Museum, Brisbane, Australia.
Description—Male (holotype). Dorsal coloration: chestnut orange with inner face of antenniferous tubercle, dorsal aspect of postocular tubercle, frontal angles, short longitudinal stripe at middle third of posterior lobe of pronotal disk, and posterior angles of connexival segments II to VII yellow to orange yellow; antennal segments I to III chestnut orange, IV yellow with basal join chestnut; he- melytral membrane yellow with inner angle black. Ventral coloration: Chestnut orange with rostral segments I to IV, buccula, acetabula (punctures chestnut orange), anterior and posterior lobe of meta- thoracic peritreme, evaporative area, and genital capsule yellow to orange yellow; rim of abdominal spiracular peritreme black; punctures of abdominal sternite V to VII reddish orange; coxae yellow with dark brown spots; trochanters yellow and apically dark brown; femora yellow, and dorsally covered with dark brown to light chestnut brown spots; tibiae yellow with four to five red rings; tarsi yellow with diffuse reddish to dark brown marks.
Female—Dorsal coloration: head chestnut orange, sprinkled with small reddish tubercles and with following areas yellow: dorsal aspect of postocular tubercle and longitudinal stripe adjacent to eyes; antennal segment I chestnut orange, segment II dark yellow with apical third reddish orange, segment III reddish orange with basal joint yellow, and IV yellow with basal joint reddish orange; pronotum, scutellum, corium, clavus and dorsal abdominal segments with punctures reddish brown to dark brown; connexival segments reddish brown with posterior angle yellow; hemelytral membrane black to dark brown with middle third yellow. Ventral coloration: yellow with reddish brown to dark brown punc- tures and irregular spots scattered throughout the body; rostral segments I-II chestnut brown, HI-IV yellow (apex of IV chestnut orange); anterior and posterior lobe of metathoracic peritreme creamy
2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 83
Figures 1-2. Male genital capsule of Grosshygia spp. Figure 1. G. formosa Brailovsky, NEW SPECIES. Figure 2. G. pisina Brailovsky, NEW SPECIES.
Figures 3-4. Weirhygiafaceta Brailovsky, NEW GENUS, NEW SPECIES. Figure 3. Male genital capsule. Figure 4. Female genitalia.
yellow and evaporative area yellow; coxae yellow with diffuse dark brown spots; trochanters yellow with apex dark brown; femora and tibiae yellow, with four to five reddish orange to black irregular rings; tarsi yellow with diffuse reddish to dark brown marks; abdominal spiracular peritreme black. Measurements.—Male (female). Head length: 1.88 mm (1.98 mm); width across eyes: 1.52 mm (1.64 mm); interocular space: 1.00 mm (1.02 mm); interocellar space: 0.50 mm (0.46 mm); preocular distance: 1.30 mm (1.36 mm); antennal segments lengths: I, 1.12 mm (1.12 mm); IJ, 1.72 mm (1.76 mm); III, 1.40 mm (1.40 mm); IV, 1.20 mm (1.16 mm). Pronotal length: 1.72 mm (1.80 mm); width across frontal angles: 1.68 mm (1.84 mm); width across humeral angles: 3.32 mm (3.56 mm). Scutellar
length: 1.00 mm (1.04 mm); width: 1.28 mm (1.48 mm). Maximum width of abdomen: 5.04 mm (5.84 mm). Total body length: 10.20 mm (11.63 mm).
Etymology.—From the Latin, facetus, meaning elegant, fine.
GROSSHYGIA FORMOSA BRAILOVSKY, NEW SPECIES (Figs. 1, 5)
Types.—Holotype male: Australia. Queensland, Kroombit Tops, 65 km SW Gladstone (Sieved litter. Q.M. Berlesate no. 383, in rainforest), 1100 m, 22—26
Feb 1982, G. Monteith and G. Thompson. Deposited in Queensland Museum, Brisbane, Australia.
84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Figure 5. Dorsal view of Grosshygia formosa Brailovsky, NEW SPECIES.
Description—Male (holotype). Coloration: dark reddish brown; antennal segment I dark chestnut orange II, and III pale chestnut orange, IV yellow with basal joint pale chestnut orange; hemelytral membrane yellow; inner margin of acetabula, coxae, and trochanters dirty chestnut orange; femora chestnut orange with basal joint, and few subapical spots dirty yellow; tibiae chestnut orange with three to four irregular yellow rings; tarsi chestnut orange with dirty yellow reflections; upper margin
2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 85
of connexival segments black. Structural characters—Head: longer than wide across eyes, pentagonal, conspicuously convex dorsally; antenniferous tubercles unarmed; rostrum reaching anterior margin of abdominal sternite VII; vertex uniformly convex, without conical tubercles. Pronotum: slightly tra- peziform, bilobed, nondeclivent; anterolateral margins convexly rounded, moderately elevated, and slightly reflexed; callar region transversely nodulose; anterior and posterior lobe along the middle line with wide longitudinal depression. Legs: unarmed. Scutellum: apex globose. Hemelytra: micropterous, reaching posterior margin of abdominal segment I; hemelytral membrane reduced to small pads, sep- arated from each other, leaving the abdomen exposed mesally. Genital capsule: posteroventral edge simple, with median triangular expansion (Fig. 1).
Female—Unknown.
Measurements.—Head length: 1.86 mm; width across eyes: 1.34 mm; interocular space: 1.06 mm; preocular distance: 1.32 mm; antennal segments lengths: I, 1.12 mm; II, 1.44 mm; III, 1.38 mm; IV, 1.28 mm. Pronotal length: 1.28 mm; maximum width of anterior lobe: 2.28 mm; maximum width of
posterior lobe: 2.92 mm. Scutellar length: 0.72 mm; width: 1.18 mm. Maximum width of abdomen: 4.60 mm. Total body length: 10.30 mm.
Discussion.—Grosshygia nigra Brailovsky and G. formosa, are the only species of the genus with the scutellar apex globose. In the other known species, it is subacute. In G. formosa the head is longer than wide, the vertex is uniformly convex without conical lobes, the antenniferous tubercles are unarmed, and the posteroventral edge of male genital capsule has a median triangular expansion (Fig. 1). In G. nigra the head is wider than long, the vertex has two conical elevations, the antenniferous tubercles are armed with extremely long lobes, and the posteroventral edge of male genital capsule is transversely straight and entire.
Etymology.—From the Latin, formosus, meaning beautifully formed.
GROSSHYGIA LEPIDA BRAILOVSKY, NEW SPECIES (Fig. 6)
Types.—Holotype female: Australia. N Queensland, Mt. Barflie Frere, Swiftlet Cave, 900 m, 8 Dec 1990, Monteith, Thompson, Cook and Sheridan. Deposited in Queensland Museum, Brisbane, Australia.
Description.—Female. (holotype). Coloration: dark chestnut orange with following areas dark yel- low: dorsal aspect of postocular tubercle, rostral segments I to IV, posterior edge of connexival seg- ments and abdominal pleural sterna III to VI, diffuse reflections on abdominal sterna III to VII, and genital plates; head and scutellum reddish brown; antennal segments I to III chestnut orange with basal joint of III yellow; antennal segment IV yellow with basal joint chestnut orange; anterior and posterior lobe of metathoracic peritreme bright chestnut orange; coxae reddish orange; trochanters and tibiae chestnut orange fore and middle femora chestnut orange with diffuse yellow rings; hind femur yellow with apical joint and two or three incomplete chestnut orange rings; tarsi yellow. Structural characters. Head: longer than wide across eyes, pentagonal, conspicuously convex dorsally; antenni- ferous tubercles armed, lobes raised, arcute, recurved, converging anteriorly, apically subacute, almost touching basal joint of antenna segment J; ocelli absent; rostrum reaching middle third of abdominal sternite VI; vertex globose with thin transverse depression, dividing it into two elevations, the anterior one broader, conical and higher than posterior one which is slightly rounded. Pronotum: scarcely quadrate, bilobed, nondeclivent; anterolateral margins convexly rounded, moderately elevated, and slightly reflexed; callar region transversely nodulose; anterior and posterior lobe along the middle line with wide longitudinal depression. Legs: unarmed. Scutellum: apex subacute. Hemelytra: micropterous, reaching middle third of abdominal segment III; wings reduced to small pads, separated from each other, leaving abdomen exposed mesally.
Male—Unknown.
Measurements.—Head length: 1.80 mm; width across eyes: 1.72 mm; interocular space: 1.20 mm; preocular distance: 1.30 mm; antennal segments lengths: I, 1.00 mm; II, 1.60 mm; III, 1.04 mm; IV, 1.04 mm. Pronotal length: 1.60 mm; maximum width of anterior lobe: 2.20 mm; maximum width of
86 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
Figure 6. Dorsal view of Grosshygia lepida Brailovsky, NEW SPECIES.
posterior lobe: 2.68 mm. Scutellar length: 0.92 mm; width: 1.16 mm. Maximum width of abdomen: 4.84 mm. Total body length: 9.55 mm.
Discussion.—Grosshygia lepida and its closely related species G. lobatula Brai- lovsky and G. monticeps Brailovsky, has the head longer than wide, the scutellar
2001
Figure 7.
BRAILOVSKY: NEW AUSTRALIAN COLPURINI
Dorsal view of Weirhygia faceta Brailovsky, NEW GENUS, NEW SPECIES.
87
88 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)
apex subacute, and the hemelytra reaching the medial third of abdominal segment III. In G. lepida the external lobes of the antenniferous tubercles converget an- teriorly, conspicuously arcuate, recurved, and almost touching the basal joint of antennal segment I (Fig. 6), and the vertex has two conical elevations separated along middle third by a deep longitudinal depression. In the other two species the external lobes of antenniferous tubercles are obliquely projecting, diverging anteriorly, and the two anterior conical elevations of vertex are almost fused. Etymology.—From the Latin, lepidus, meaning pleasant, elegant.
GROSSHYGIA PISINA BRAILOVSKY, NEW SPECIES (Fig. 2.)
Types.—Holotype male: Australia. NE Queensland, 20 km WNW of Cape Trib- ulation (Site 2), 50 m, 23 Sep 1982, Monteith, Yeates and Thompson. Deposited in Queensland Museum, Brisbane, Australia.
Description—Male (holotype). Dorsal coloration: head reddish brown with dorsal aspect of pos- tocular tubercle yellow; antennal segments I to III chestnut orange with basal joint of III yellow; antennal segment IV yellow with basal joint chestnut orange; anterior lobe of pronotum reddish brown, posterior lobe chestnut orange, anterolateral margins yellow; scutellum dark chestnut orange; heme- lytral membrane chestnut orange; connexival segments bright reddish with posterior margins of seg- ments III to VI yellow; dorsal abdominal segments reddish brown. Ventral coloration: Head reddish brown, rostral segments I to IV and small spot close to eyes yellow; thorax reddish brown with acetabula dark reddish, pro, meso, and metapleura dirty yellow with punctures dark brown; coxae bright reddish; trochanters bright chestnut yellow; fore and middle femora chestnut orange with basal third, subapical third, and few scattered spots yellow; hind femur yellow with apical third and four irregular rings chestnut brown; tibiae and tarsi yellow; abdominal sterna dark reddish brown, scattered with yellow diffuse areas; pleural abdominal sterna bright reddish with posterior third of sterna III to VI yellow; genital capsule reddish brown with posteroventral margin pale orange yellow. Structural characters. Head: longer than wide across eyes, pentagonal, conspicuously convex dorsally; antenni- ferous tubercles each armed with short and robust lobe, obliquely projecting, diverging anteriorly and subacute apically; rostrum reaching anterior margin of abdominal sternite VII; vertex uniformly convex without conical elevations. Pronotum: slightly trapezoidal, bilobed, and nondeclivent; anterolateral margins convexly rounded, moderately elevated, and slightly reflexed; callar region transversely nod- ulose; anterior and posterior lobe along medially with wide longitudinal depression. Legs: unarmed. Scutellum: apex subacute. Hemelytra: micropterous, reaching posterior third of abdominal segment III; hemelytral membrane absent. Genital capsule: posteroventral edge simple, transversely concave, entire (Fig. 2).
Female-——Unknown.
Measurements——Head length: 1.82 mm; width across eyes: 1.48 mm; interocular space: 1.00 mm; preocular distance: 1.24 mm; antennal segments lengths: I, 1.14 mm; II, 1.84 mm; III, 1.14 mm; IV, 1.12 mm. Pronotal length: 1.48 mm; maximum width of anterior lobe: 1.88 mm; maximum width of posterior lobe: 2.28 mm. Scutellar length: 0.80 mm; width: 0.96 mm. Maximum width of abdomen: 3.68 mm. Total body length: 9.10 mm.
Discussion.—Like G. monticeps Brailovsky, the head is longer than wide, the scutellar apex is subacute, the antenniferous tubercles each is armed with a short lobe, and the total length of antennal segment II is shorter than 1.86 mm. In G. pisina, the vertex is uniformly convex without conical elevations, the membrane of the hemelytra is absent, and the