te-tatie 6 “« Pei rm hi One cope 2 : y Snsteiee Pag hye vibe etait Cheat bet BANOS CST f : , 2 Rute ade . é see fe Neath all Mitte hota ; Bs : i te ‘é WO "8 ee : ehake - : ee . eet neta piqued * F 4 Mpa i : ; F 3 Weise aever tees os Stitnces tp onecre shh : ‘ * ‘ 7 - > : E aA EEA NEA gh ating ( : Hs8 4 ; S :
A ; z ¢ eta Vege igre dnts res : . ‘ 7 , 7 $7 B 2 ¢ 2 * ’ Arise even
Pet AE, UR gs a af
cei : ‘ Pevdpahanat eae doe ie INN EM Sane . . ‘ ™ ~
Peas Pe haan bhoen wis
a
RA nnn RL AD AR ME et eu ear
1 wire j
VOL. 102 JANUARY 2000 NO. 1 @ ee (ISSN. 0013-8797)
+N ELaX PROCEEDINGS |
ea of the
j
N iad Ri
PUBLISHED QUARTERLY
CONTENTS
ASSIS-PUJOL, CRISTIANE VIEIRA DE and MICHEL LECOQ—Comparative study of sper- mathecae in eleven Rhammatocerus Saussure 1861 grasshopper species (Orthoptera:
Achadidae .Gomphnocerinacs Scyllimint) it yee, een eee na at eee ries cite ae ecm Keone ee aac ees 120 BROWN, JOHN W.—Revision of Lobogenesis Razowski and Odonthalitus Razowski (Lepidoptera:
Tortricidae: Tortricinae), with comments on their monophyly ....................0eeeeee eee ee 2 BUTLER, LINDA and JOHN STRAZANAC—Macrolepidopteran larvae sampled by tree bands in
temperate mesicund sernctorests injeastern, WiMlteGestates| siieeeiec one. sane sees sets neice 188 CICERO, MARY R. and EDWARD M. BARROWS—Milesiine flower flies (Diptera: Syrphidae)
in a central Appalachian broadleaf forest: Abundances, flight periods, and diflubenzuron.... 50
CRYAN, JASON R. and LEWIS L. DEITZ—Review of the New World treehopper tribe Stegaspidini (Hemiptera: Membracidae: Stegaspidinae): Ill: Flexocentrus Goding, SAVLOCENIFUSHS tal sand, O71 DEULSCTUSsD CLUZ ancy a ene tele terns ocd t= ee aicielacs pe iste re Aaah tata 82
DARSIE, RICHARD E.,, JR.—Description of the pupae of five species in the subgenus Armigeres, genus Armigeres Theobald, with a key to species of the known pupae of the sub-
PenUshUMiptekan GUC Ae amet ste satie en rh Mel vartele = ceuctiie Mie ara cistaevesieis NSE aol iam ae sl yt eueprite 108 GATES, MICHAEL W.—A new species of Cirrospilus Westwood (Hymenoptera: Eulophidae) fLOMMtAe SOULNWeSteEMNUMIted States anu MCxI GO ca), ac elt tee ae cra tele roo 'e ove aleve emis ievoe siassvayeinvere= 58
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota signifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey and
ASGray, (Asteraceae) in SOUthenne @alitOnntaet S10 yeie es «lor tente sie ah asia es viele else ona cle slgaiauaresasaler 69 HASTRITER, MICHAEL W.—Jordanopsylla becki (Siphonaptera: Ctenophthalmidae), a new SHECIES OM CanhOud MCU EVAMAMLESE SILC Mth ye my rac cto ercie ay hacteieei-lopiate slotelsiseris erm roe elem nate elerateret 215)
HOEBEKE, E. RICHARD, ROBERT A. BYERS, MIGUEL A. ALONSO-ZARAZAGA, and JAMES F. STIMMEL—Jschnopterapion (Chlorapion) virens (Herbst) (Coleoptera: Curculionoidea: Brentidae: Apioninae), a Palearctic clover pest new to North America:
Recognition featiressdisintoution:)and DIOMOMMICS Heke aic to clo sy claveciewc J+) eisioe ts eleharstais mie eerste 151 HUNG, AKEY C. F. and HACHIRO SHIMANUKI—Nucleotide sequence and restriction site analy- ses in three isolates of Kashmir bee virus from Apis mellifera L. (Hymenoptera: Apidae) ...... 178
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
OFFICERS FOR 2000 Davip G. FurtH, President MIcHAEL G. PoGuE, 7reasurer JOHN W. Brown, President-Elect THEODORE R. SCHULTZ, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho us B. WILLIAMS, Corresponding Secretary MiIcHAEL E. ScHAuFF, Past President
Jon A. Lewis, Custodian Davip R. SmitH, Editor
Publications Committee THOMAS J. HENRY Wayne N. MaAtTHIs
Gary L. MILLER, Book Review Editor
Honorary President Louise M. RuUssELL
Honorary Members KARL V. KROMBEIN RONALD W. HoDGEs DoNALD M. ANDERSON WILLIAM E. BICKLEY
All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168.
MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:30 P.M. Minutes of meetings are published regularly in the Proceedings.
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency).
PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington.
The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 699-700 OF THE JULY 1999 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October).
Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168.
Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168.
Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 10 January 2000
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 1-20
A REVISION OF THE ENDEMIC HAWAIIAN REDUVIID GENUS SAICELLA USINGER, WITH DESCRIPTIONS OF FOUR NEW SPECIES (HETEROPTERA: REDUVIIDAE: EMESINAE)
DAN A. POLHEMUS
Department of Entomology, MRC 105
, Smithsonian Institution, Washington, DC
20560-0105, U.S.A. (e-mail: polhemus.dan @ nmnh.si.edu)
Abstract.—The Hawaiian genus Saicella i
s revised, and four new species are described:
S. perkinsi from Oahu, S. lilinoe from Maui, S. kipahulu from Maui, and S. mulli from Hawaii. A key to species is provided, accompanied by distribution maps, habitus figures,
and illustrations of key characters.
Key Words: Reduviidae, Hawaiian Islands
Assassin bugs, in the family Reduviidae, are uncommon and elusive components of the native Hawaiian insect biota. Only two endemic genera, Saicella Usinger and Nes- idiolestes Kirkaldy, both in the subfamily Emesinae, are known from the islands, con- taining a total of five currently described species between them. Even with the de- scription of the additional new taxa now present in museum collections, it seems un- likely that the total native Hawaiian redu- viid fauna will exceed 15 species. This stands in contrast to the native Hawaiian Nabidae, which appear to fill many typical reduviid niches in the islands. The latter group contains 25 described species, with an equal or greater number awaiting de- scription. This is a striking reversal of fam- ily dominance in comparison to continental settings, where species of Reduviidae far outnumber those of Nabidae, and is a pe- culiarity of community organization in the Hawaiian archipelago and other isolated Pa- cific island groups. It is probably indicative of a difference in overwater dispersal abil- ities between these two groups of Heter- optera (Leston 1957), and of their relative capabilities to exploit underutilized ecolog- ical niches.
, Saicella, new species, biogeography
Here I revise the classification of Saicel- la, providing descriptions of four new spe- cies. The two previous species in this ge- nus, S. smithi Usinger (1958) and S. usin- geri Wygodzinsky (1966), were described from Maui and Kauai respectively. Two of the new taxa described herein are also from Maui, with the other two coming from the islands of Oahu and Hawaii. These new species now extend the known range of the genus throughout the archipelago, and it seems quite likely that further collecting will produce examples of this genus from Molokai and possibly Lanai.
Collections of Saicella have been sporad- ic at best over the last 100 years. The first two examples were taken by R. C. L. Per- kins on Maui in 1894 and 1896, during his collecting work for the Fauna Hawaiiensis. They seem to have escaped the notice of Kirkaldy, who authored the treatment of Heteroptera for this series, and were not formally described until 1958, when R. L. Usinger happened across one of the speci- mens in the British Museum, and subse- quently located the second in Honolulu, at the Bishop Museum. Usinger was eventu- ally able to collect a series of a second spe-
i)
cies on Kauai in 1961, which was described by Wygodzinsky (1966) in his monograph of world Emesinae. A few additional spec- imens were collected in the mid-1980’s by researchers from the Bishop Museum dur- ing biological surveys on Maui and Oahu, and a series of specimens was reared from eggs by hobbyist William Mull on Hawaii. Even so, the sum total of specimens col- lected between 1894 and 1994 amounted to less than 50 individuals, many of them im- matures.
This situation has changed markedly in the last several years, due to a better un- derstanding of the microhabitats preferred by Saicella species and improvements in collecting techniques. In particular, the use of small scale pyrethrin fogs targeted at mossy logs and tree trunks in areas of wet native forest has been especially effective. This technique produced 80 specimens from five different localities on Maui and Kauai in only nine months during 1998 and 1999, thereby more than doubling the adult captures from the previous hundred years, and revealing two new species in the pro- cess. Continued use of such local fogging may well prove Saicella to be typical, though cryptic, components of wet, high el- evation native forest communities on all the major Hawaiian volcanoes.
The comparatively high diversity of Sai- cella on Maui is an interesting biogeo- graphic anomaly, especially considering that all three Maui species have come from Haleakala, the mountain that forms the east- ern portion of the island, while none are yet known from the West Maui Mountains. The Haleakala species divide the mountain into three discrete areas of endemism, one in the area near upper Waikamoi Gulch, another on the central portion of the northern face in the upper Hanawi Gulch area, and a third in Kipahulu Valley (Fig. 12). These areas of Saicella endemism are roughly congru- ent with portions of the mountain retaining a surface of Kula Volcanic lavas with a mean age of 0.41 ma, which are surrounded by younger flows of the Hana Volcanics
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
that erupted in the late Pleistocene, 0.4 ma later (Clague and Dalrymple 1987, Langen- heim and Clague 1987). It is tempting to hypothesize that the patterns of speciation currently observed in Maui Saicella are the result of allopatry due to vicariance, with the eruption of the Hana Volcanics having isolated sections of the older Kula Volcanic surface, which then functioned as islands of rain forest habitat, but tests of this model must await a cladistic analysis. The Maui pattern is all the more interesting in light of the fact that no similar pattern of intra-is- land endemism is observed on the much older island of Kauai, where a single spe- cies, S. usingeri, occurs on both the Alakai Plateau and in the Makaleha Mountains, de- spite the isolation of these massifs from each other by sheer cliffs, deep valleys, and other topographic barriers easily as signifi- cant as those separating the three allopatric species on Maui.
METHODS
Synonymies given are nomenclatural only. All measurements are given in milli- meters. Characters typical of the genus that are noted in the generic description are not repeated in the individual species descrip- tions. To avoid confusion, tergite number- ing in the descriptions refers to visible ter- gites; the basal tergites of many Emesinae are reduced or hidden (see Wygodzinsky 1966), so that the location of actual tergite I is often difficult to ascertain.
Institutional abbreviation codes used in the Material Examined sections are ex- plained in the Acknowledgments section. In addition, information not found on original specimen labels but helpful in their inter- pretation is given in brackets in the Material Examined sections. Spellings for place names follow those found on the topo- graphic maps of the United States Geolog- ical Survey, and do not include diacritical marks. In the following discussions, the word ‘‘Hawaii”’ refers to the island of Ha- waii itself. If the entire Hawaiian island
VOLUME 102, NUMBER 1
chain is under discussion then the phrase ‘*Hawaiian Archipelago” is employed.
Genus Saicella Usinger 1958 (Figs. 1-13)
Type species.—Saicella smithi Usinger 1958: 440.
Description.—Small, micropterous redu- viids, length 4.5—6.0 mm (Figs. 1—2).
Body surface shining, portions of head, thorax, and basal abdominal segment often bearing short, appressed, wool-like pile. Ground color varying from pale green to yellowish brown, marked with red, dark brown, or black, legs often conspicuously annulate with dark brown or black.
Head short, divided into anteocular and postocular portions by deep transverse sul- cus, both portions strongly convex above, anteocular section with sides subparallel when viewed dorsally, postocular section semi-globular both dorsally and laterally; eyes small, set laterally, lying below dorsal surface of head; rostrum 3 segmented, bent at first intersegmental suture, cylindrical, length of segment I subequal to lengths of segments II and III combined, segment I bearing 2+2 spinelike setae to either side of midline, segment II with 1+1 similar se- tae (Fig. 3); antenniferous tubercles large, antennae inserted near anterior margin of head, lacking long hairs in both sexes; buc- cula with a pair of socketed spine-like setae, gena with 3—4 similar setae.
Pronotum covering anterior half of me- sonotum, divided into anterior and posterior sections by deep, forward curving trans- verse sulcus; anterior section subglobular, with a deep longitudinal median sulcus sep- arating two elevated lobes; posterior section transverse; anterior acetabulae opening for- ward and downward. Scutellum (exposed portion of mesoscutum) subequal in length to posterior section of pronotum, bearing a long, erect spine. Metanotum subequal in length to scutellum, also with a long, erect spine. Forewings consisting of short, slen- der pads, reaching at most to base of ab- domen, membrane absent.
Abdomen broadly attached to thorax, not constricted at base, connexival margins out- wardly convex, broadly arcuate, occasion- ally sculptured, dorsal tergites often bearing small tumescences posteromedially, ventri- tes broadly convex. Male with seventh vis- ible tergite triangular, elongate, often with a longitudinal medial keel, completely cov- ering genital segments when viewed dor- sally; ventrite VIII fully visible, large; py- gophore large, length equal to % total ab- domen, sclerotized dorsally, posterosuperi- or process narrowing suddenly on distal half, coming to a spine-like point; paramer- es small, club-like.
Foreleg stout, coxa with 2 long, stout, socketed spines basally, followed by 2-3 smaller spines distally; trochanter unarmed; femur with 5 long, stout, socketed spines on basal % of ventral surface (Fig. 3), in- termixed with long erect pale setae and set within a row of smaller spinules, length of the longest spines subequal to width of fe- mur, inner dorsal surface of femur bearing about 10 moderately long socketed spines; tibia slender, ventral surface with two lon- gitudinal combs of erect, pale, spine-like setae, small brushes of gold setae also pre- sent at tip on inner face, and on central sec- tion of outer face; tarsus two segmented, basal segment slightly shorter than second, both segments with numerous short setae on ventral surfaces, lacking specialized se- tae; claws curved, inner claw medially in- cised, outer claw with two small subbasal projections. Middle and hind legs slender and elongate, hind femur surpassing apex of abdomen, all segments bearing numerous very short, semi-erect, distally angling se- tae; middle and hind tarsi 3 segmented, these segments subequal in length.
Discussion.—Usinger (1958) considered the subfamilial placement of Saicella to be equivocal, due to the uniformly micropter- ous condition, and suggested that the genus might fall in either the Saicinae or Emesi- nae. Wygodzinksy (1966) provided detailed arguments for a placement in the Emesinae,
4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tribe Ploiariolini, an interpretation followed herein.
KEY TO THE SPECIES OF SAICELLA
1. Wing pads long; reaching to base of abdomen, surpassing posterior tip of scutellar spine when viewed from above (Fig. 2); Maui WE RHA ELD coeds ane de aT © smithi Usinger
— Wing pads shorter, not reaching to base of ab- domen, usually not surpassing posterior tip of scutellar spine (except in S. perkinsi, where this spine is vertical), definitely not exceeding base ofmetanotallispinei(Gicai)i yates 2
2. Antennal segment I distinctly annulate, with al- ternating dark and pale bands (Fig. 1) ..... 3
— Antennal segment I usually unicolorous brown, bearing at most a few faint indications of dark- ErpannuUlatlONSie. wiry js oe eee a one ae 5
3. First visible abdominal tergite with conical tu- mescence anteromedially; ground color pale green overlain with darker markings; Maui Gf tO ler aus Daca DI OPER Ley CECk oan 0, dyons fo S. lilinoe, n. sp.
— First visible abdominal tergite lacking a conical tumescence; ground color yellowish brown, overlain with darker brown or reddish mark- Ine sa Walalpandehlawalluewe emery) cars es ete 4
4. Wing pads extending to tip of scutellar spine: anterior lobes of pronotum dull, bearing patch- es of pale appressed hairs; abdomen with red markangsilaterallliyiKaualeyepeseis eats
F keagnied «yaad ht Mos shen S. usingeri Wygodzinsky
Wing pads tiny, hard to see, not extending past
base of scutellar spine; anterior lobes of pron-
otum glabrous, shining; abdomen without red markings laterally; Hawali...... S. mulli, n. sp.
5. Pronotum with numerous long, slender, erect, pilose hairs; hind tibia unicolorous medium brown, without well defined annulations; an- terior margin of first visible abdominal tergite strongly angled upward anteromedially; Oahu ed SE GES ict ch CaS res eel OUR Eee fh S. perkinsi, n. sp.
— Pronotum lacking long, erect, pilose hairs, bearing only patches of pale, appressed hairs; hind tibia with numerous black annulations; anterior margin of first visible abdominal ter- gite not strongly angled upward anteromedial-
liv; Mi alse. bpd Sokpalts ay clceresey = S. kipahulu, n. sp.
Saicella smithi Usinger (Figs. 2—3, 9, 12)
Saicella smithi Usinger 1958: 440.
Type.—Holotype, 2, from Haleakala, Maui, 5,000 ft., in The Natural History Mu- seum, London.
Diagnosis.—Recognized by the relative- ly long wing pads that reach beyond the
apex of the scutellar spine to the base of the abdomen (Fig. 2); absence of obvious an- nulations on antennal segments I and II; and absence of raised tubercles on the abdomi- nal tergites (Figs. 2-3, 9).
Redescription.—Micropterous female: General coloration yellowish brown, with darker brown markings on thorax and base of abdomen; legs multiannulate with dark brown; lateral portions and posterior margin of abdominal tergites sparingly marked with bright red.
Length 4.70 mm, maximum width (across abdomen) 1.50 mm.
Head \ength/width = 0.68/0.54, covered by a thick layer of short, pale, curling, re- cumbent setae; width of vertex 2.9 dorsal width of an eye (0.32/0.11); length of an- terior lobe of head 2.50 dorsal length of an eye (0.40/0.16); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 1.68 dorsal length of an eye (0.27/0.16); ocelli absent; length of antennal segments I-IV = 1.98/ 2.07/0.52/missing; rostrum length 0.85, reaching to bases of fore coxae; coloration of head yellowish brown, without contrast- ing dark markings; antennal segment I dark yellowish, distal % medium brown, basal half with very faint suggestions of 1 or 2 brown annulations, antennal segments II and III uniformly light brown.
Pronotum length (midline)/width = 0.57/ 0.63, covered by an obscure layer of short, pale, recumbent setae; coloration uniform golden yellowish brown; anterior section with lateral lobes shining, lacking patches of appressed setae; posterior section bearing a small, conical tumescence centrally.
Scutellum triangular, coloration uniform- ly golden yellowish brown, lateral angles bearing an obscure layer of short, pale, re- cumbent setae, central section produced into an erect, slender, backward angling spine posteromedially; length/width = 0.27/ Oals:
Hemelytra short, micropterous, reaching to base of abdomen, separated medially by an erect, slender, backward angling spine
VOLUME 102, NUMBER 1
fe
1
Fig. 1. Saicella lilinoe. Female, dorsal habitus (Young Sohn illustration).
6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 2-3. Saicella smithi, female (Arthur Smith illustrations). 2, Dorsal habitus. 3, Body and forelegs in lateral view.
arising from the underlying mesonotum, gested, covered by an obscure layer of this spine similar in size, shape and orien- short, pale, recumbent setae, posterior mar- tation to that arising from scutellum; wing gins rounded, membrane absent; coloration pads rugulose, with claval vein barely sug- yellowish brown.
VOLUME 102, NUMBER 1
Legs elongate, with fore coxa approxi- mately 3.5X as long as thick (0.63/0.18); fore trochanter bearing numerous slender erect gold setae; fore femur fusiform, over 5.7X longer than wide (1.26/0.22); colora- tion of legs yellowish brown with brown markings; fore femur yellowish brown ba- sally, with a broad brown annulation on dis- tal half; fore tibia with 3 brown annulations, knee pale, apex dark; middle and hind fem- ora each with 4 brown annulations, these annulations roughly equal in length to in- tervening pale spaces; middle and hind tib- iae with 5 and 7 brown annulations respec- tively, these annuli concentrated in the bas- al half of the segment; middle and hind tarsi uniform brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.26/0.99/0.06/0.20; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 2.14/2.87/0.06/0.06/ 0.06; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 2.86/4.16/0.05/0.05/0.05.
Abdomen yellowish brown, with lateral and posterior sections of first visible tergite darker brown, lateral sections of first through sixth visible tergites also bearing irregular bright red maculations; connexival margins broadly arcuate, abdominal tergites lacking raised tumescences posteromedial- ly, first through sixth visible segments dor- sally concave, with lateral margins curved upward and narrowly emarginate, posterior margin of sixth visible tergite bisinuate, seventh visible tergite broadly triangular, flat, with a very small indentation at ex- treme posterior apex (Fig. 9); all tergites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae.
Ventral surface yellowish brown, cov- ered by an obscure layer of short, pale, re- cumbent setae; abdominal paratergites red- dish.
Micropterous male: Unknown.
Distribution.—Maui (Haleakala, Waika- moi area) (Fig. 12).
Ecological associations.—Unknown.
Material examined.—HAWAIIAN IS- LANDS, Maui: 1 92, Haleakala, 5,000 ft
7
P30: mij. EEAEIV. 18943 RoC: I. Perkins (paratype, BPBM).
Discussion.—The description above was taken from a female paratype, housed in the Bishop Museum. It is one of the two orig- inal specimens of S. smithi on which Usin- ger (1958) based his description. The Ar- thur Smith figure accompanying Usinger’s (1958) description, reproduced in Fig. 2, is accurate in most respects, but depicts ob- vious dark annulations on antennal segment I that are not present on the Bishop Muse- um specimen examined by the author.
Both of the above specimens were taken by Perkins from “Haleakala, 5,000 ft.”’; based on Perkins’ field notes, the type lo- cality must lie somewhere upslope of Olin- da and west of Waikamoi Gulch. Much of the forest in this area has now been cleared for cattle ranching, but a tract is still pro- tected in The Nature Conservancy of Ha- wail’s Waikamoi Preserve. Repeated sur- veys in this remaining forest block, how- ever, have failed to produce further speci- mens of S. smithi. As a result, the male characters of this species remain unknown.
Saicella lilinoe Polhemus, new species (Figs. 1, 8, 12)
Diagnosis.—Recognized by the short wing pads that do not reach beyond the apex of the scutellar spine (Fig. 1); strongly annulate antennal segments I and II (Fig. 1); and presence of raised tubercles poster- omedially on the fourth through sixth visi- ble abdominal segments (Figs. 1, 8).
Description.—Micropterous male: Gen- eral coloration pale green, with darker brown markings on head, thorax and ab- domen; legs multiannulate with dark brown or black (Fig. 1).
Length 5.25 mm, maximum width (across abdomen) 1.80 mm.
Head \ength/width = 0.80/0.56, covered by a thick layer of pale, appressed setae; width of vertex 4.8 dorsal width of an eye (0.44/0.09); length of anterior lobe of head 2.4 dorsal length of an eye (0.45/0.19); eyes small, consisting of approximately 20
8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 4-9. 7, S. kipahulu. 8, S. lilinoe. 9, S. smithi.
ommatidia each; length of posterior lobe of head 1.63 dorsal length of an eye (0.31/ 0.19); ocelli absent; length of antennal seg- ments I-[V = 3.10/2.75/0.75/0.55; rostrum length 1.00, reaching to bases of fore coxae; coloration of head uniform medium brown,
Saicella species, dorsal view of female terminal abdomen. 4, S. mulli. 5, S. usingeri. 6, S. perkinsi.
without contrasting markings; antennal seg- ment I dark yellow, bearing 5 evenly spaced dark brown annulations, tip light; antennal segment II dark yellow, bearing 6 evenly spaced dark annulations, tip dark; antennal segments III and IV uniform medium
VOLUME 102, NUMBER 1
brown except narrowly pale at extreme ba- ses and tips.
Pronotum length (midline)/width = 0.75/ 0.50; anterior section yellowish green, be- coming embrowned toward pleurae, lateral lobes shining, lacking setae, intervening longitudinal sulcus bearing numerous ap- pressed pale setae; posterior section of pronotum milky greenish white, lacking se- tae, bearing a small, conical tumescence centrally.
Scutellum triangular, lateral angles black, central section pale green, produced into an erect, slender, backward angling spine pos- teromedially, this spine strongly curving when viewed laterally; a ring of pale, ap- pressed setae present around base of spine; length (including spine)/width = 0.32/0.27.
Hemelytra short, micropterous, reaching only to middle of scutellar spine when viewed from above, separated medially by an erect, slender, pale green, backward an- gling spine arising from the underlying me- sonotum, this spine similar in size, shape and orientation to that arising from scutel- lum; wing pads consisting of tiny, elongate flaps, venation and membrane absent; col- oration black centrally, margins brown.
Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (0.80/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 7.0 longer than wide (1.75/0.25); coloration of legs dark yellow with brown or black mark- ings; fore femur yellowish brown basally, with a broad brown annulation on distal half; fore tibia with 3 diffuse brown annu- lations on outer face, knee pale, apex nar- rowly pale; middle and hind femora each with 4 brown annulations, these annulations broad and diffuse basally, becoming smaller than width of intervening pale areas distal- ly; middle and hind tibiae each with 4—6 brown annulations, these annuli regularly spaced on middle tibia, more irregularly spaced on hind tibia, with a prominent black annulation on basal %4, followed by other smaller annulations in distal half of segment; middle and hind tarsi uniform me-
dium brown; lengths of leg segments as fol- lows: fore femur/tibia/tarsal 1/tarsal 2 = 1.75/1.37/0.09/0.25; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 3.50/4.70/0.09/0.09/ 0.09; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 4.80/6.60/0.10/0.10/0.10.
Abdomen pale green, yellowish centrally, this ground color overlain with irregular black markings on entirity of first visible tergite and posterolaterally on second through sixth visible tergites, the lateral dark markings on third through sixth visible tergites extending inward along posterior tergite margins to form transverse bands, anterior sections of third through sixth vis- ible tergites with blackish brown marks in the form of posteriorly directed triangles, this combination of markings giving the ab- domen a transversely striped appearance (Fig. 1); connexival margins forming incip- ient knobs at sutures, intervening sections weakly concave; first visible tergite with a small, conical tumescence anteromedially, third through fifth visible tergites with smaller, more rounded raised tumescences posteromedially, sixth visible tergite trian- gular, longitudinally keeled on _ posterior half; all tergites shining, covered by a sparse, obscure layer of very short, pale, re- cumbent setae.
Ventral surface of head and thorax black, abdominal venter greenish brown, covered by an obscure layer of short, pale, recum- bent setae; abdominal paratergites narrowly margined with pale green adjoining con- nexival margin.
Male genitalia with paramere stout, slightly curving, bearing a small lateral tab at tip; tip of pygophore narrowed and elon- gate, tapering evenly to truncate apex with a slightly expanded terminal cap.
Micropterous female: Length 5.70 mm, maximum width (across abdomen) 1.90 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posteromedial tumescences on abdominal tergites more highly developed, particularly on fifth vis- ible tergite, posterior margin of sixth visible
10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tergite broadly V-shaped, seventh visible tergite roughly triangular, rounded posteri- orly (Fig. 8).
Distribution.—Maui (Haleakala) (Fig. 12):
Ecological associations.—All specimens so far captured have been taken by applying a light pyrethrin fog to mossy tree trunks and root masses of ohia trees (Metrosideros polymorpha). These insects appear to prefer the sheltered, almost cave-like situations found in cavities beneath multi-stemmed Metrosideros and fallen logs, and multiple specimens are often taken at a single spot, indicating a tendency toward aggregation.
Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Maui, Haleakala, Poo Uli Cabin area, near head- waters of Kuhiwa Stream, Hanawi Natural Areay Reserve 5,200), ft VSs85) me]; 20°45.03'N, 156°07.40'W, 5-6 May 1998, CL 8324, D. A. Polhemus (BPBM). Para- types: HAWAIIAN ISLANDS, Maui, Ha- leakala: 7 3d, one 2, same data as holotype (USNM, BPBM); 14 6,9 @, same locality as preceeding, 5 May 1998, lot Ol, pyre- thrin fog on mossy ohia [Metrosideros po- lymorpha], J. K. Liebherr (CUIC); 11 36,5 ?, 5 immatures, State Fence Camp cabin, along headwaters of Heleleikeoha Stream, Hana Forest Reserve, 5,300 ft. [1,615 m.], 20°44.41'N, 156°06.12'’W, 11-12 May 1998, CL 8326, D. A. Polhemus, J. K. Lie- bherr and C. Ewing (USNM, BPBM); 5 6, 4 2, same locality as preceeding, 12 May 1998, lot 01, pyrethrin fog on mossy ohia [Metrosideros polymorpha], J. K. Liebherr (CUIC); 1 36, Haleakala National Park, Ki- pahulu Valley, West Rim below Kuiki, 2,090 m. [6,855 ft.], 14 May 1993, lot O1, sifting moss from ohia [Metrosideros po- lymorpha], J. K. Liebherr and A. C. Me- dieros (CUIC).
Etymology.—The name “‘lilinoe’’ refers to the Hawaiian goddess of mists, an ap- propriate name for this species of the cloud forests.
Discussion.—In addition to the charac- ters mentioned in the key and diagnosis,
Saicella lilinoe may be easily recognized by its green and black coloration in living in- dividuals, the striped appearance of the ab- domen (Fig. 1), and the glabrous anterior pronotal callosities.
This species appears to be distributed across the northern face of Haleakala, from the Koolau Gap eastward to Kipahulu Val- ley. This is an extremely wet cloud forest zone, with precipitation exceeding 9,000 mm (360 inches) a year. A single specimen is also at hand from the upper section of Kipahulu Valley, further around the eastern tip of the island near to the type locality of S. kipahulu; this is the only place where two species of Saicella are known to be poten- tially sympatric.
Saicella kipahulu Polhemus, new species (igs! 7; 12)
Diagnosis.—Recognized by the short wing pads that do not reach beyond the apex of the scutellar spine; unicolorous antennal segments I and II, and absence of raised tu- bercles posteromedially on the fourth through sixth visible abdominal segments.
Description.—Micropterous male: Gen- eral coloration medium brown to yellowish brown, with dark brown or reddish mark- ings on abdomen; legs multiannulate with dark brown.
Length 5.20 mm, maximum width (across abdomen) 1.25 mm.
Head \ength/width = 0.79/0.55, covered by a thick layer of pale, appressed setae; width of vertex 3.7 dorsal width of an eye (0.37/0.10); length of anterior lobe of head 4.2 dorsal length of an eye (0.50/0.12); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 2.42 dorsal length of an eye (0.29/ 0.12); ocelli absent; length of antennal seg- ments I-IV = 2.25/2.00/0.75/0.40; rostrum length 0.95; coloration of head uniform me- dium brown, without contrasting markings; antennal segment I medium brown, becom- ing slightly paler basally, lacking annula- tions; antennal segments II-IV uniform me-
VOLUME 102, NUMBER 1
= o D £ 1) =] a)
@)
S. perkinsi
OC)
eri on Kauai. 11, S. perkinsi on Oahu.
Distribution of Saicella species. 10, S. using
Figs. 10-11.
12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
dium to dark brown except narrowly pale at extreme bases and tips.
Pronotum length (midline)/width = 0.55/ 0.75; anterior section medium brown, lat- eral lobes bearing scattered, elongate patch- es of appressed gold setae, intervening lon- gitudinal sulcus also bearing appressed gold setae; posterior section of pronotum yellow- ish brown, with a raised longitudinal me- dian carina, set with scattered short, semi- erect Setae.
Scutellum triangular, medium brown, central section pale yellowish brown, pro- duced into an erect, slender, backward an- gling spine posteromedially, this spine strongly curving when viewed laterally; a ring of appressed gold setae present around base of spine; length (including spine)/ width = 0.35/0.30.
Hemelytra short, micropterous, reaching nearly to tip of scutellar spine when viewed from above, separated medially by an erect, slender, yellowish brown spine arising from the underlying mesonotum, this spine sim- ilar in size and shape to that arising from scutellum, but more vertical; wing pads consisting of narrow, elongate flaps, bear- ing a few short, erect gold setae, venation and membrane absent; coloration medium brown.
Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (0.75/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 6.0 longer than wide (1.50/0.25); coloration of legs dark yellow with brown or black markings; fore femur yellowish brown ba- sally, with a broad brown annulation on distal half; fore tibia with 3 diffuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown an- nulations, these annulations broad and dif- fuse basally, becoming smaller than width of intervening pale areas distally; middle and hind tibiae each with 6 brown annu- lations, these annuli regularly spaced on middle tibia, more irregularly spaced on hind tibia, with a prominent black annu- lation on basal %4, followed by other small-
er annulations in distal half of segment; middle and hind tarsi uniform medium brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.50/ 1.20/0.10/0.25; middle femur/tibia/tarsal 1/ tarsal 2/tarsal 3 = 2.60/3.70/0.08/0.08/ 0.08; hind femur/tibia/tarsal 1/tarsal 2/tar- sal 3 = 3.60/5.10/0.08/0.08/0.08.
Abdomen dark yellow, this ground color overlain with irregular black markings on basal % of first visible tergite and antero- medially on second through sixth visible tergites, the dark markings on these latter tergites extending outward along anterior tergite margins to form transverse bands, giving the abdomen a transversely striped appearance, additional irregular dark mark- ings present on connexival segments, su- tures between tergites and connexiva irreg- ularly blotched with red; connexival mar- gins evenly convex and curving, lacking knobs at sutures; first visible tergite with anterior margin raised to form a low, tri- angular tumescence anteromedially, second through fifth visible tergites lacking tumes- cences, sixth visible tergite elongate, trian- gular, tip swollen and rounded, pale; all ter- gites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae.
Ventral surface of head and thorax black, abdominal venter greenish brown, covered by an obscure layer of short, pale, recum- bent setae; abdominal paratergites narrowly margined with pale green adjoining con- nexival margin.
Male genitalia with paramere stout, bear- ing a small lateral tab at tip; tip of pygo- phore narrowed and elongate, tapering evenly to acuminate apex.
Micropterous female: Length 5.75 mm, maximum width (across abdomen) 1.80 mm. Similar to male in general structure and coloration; abdomen broader and more strongly expanded, posterior margin of sixth visible tergite nearly straight, seventh visible tergite triangular (Fig. 7).
Distribution.—Maui (Haleakala) (Fig. 12):
Ecological
associations.—The habitat
VOLUME 102, NUMBER 1
© = S. smithi
@) =S. lilinoe
[m| = S. kipahulu
Fig. 12. Distribution of Saicella species on Maui.
preferences of this species are very similar to those of S. Jilinoe, with individuals being taken from mossy tree trunks and root mas- ses in wet montane forests.
Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Maui, Haleakala, West Camp, upper Kipahulu Valley, Haleakala National Park, 6,400 ft. [1,950 m.], 20°43.36’N, 156°07.79'W, 18- 22 May 1998; C833 1D ArrPolhemus, B. H. Gagné and R. Takumi (BPBM). Par- atypes (all micropterous): HAWAIIAN IS- LANDS, Maui, Haleakala: 1 6, 8 2, same data as holotype (USNM, BPBM, CUIC); 1 3,1 &, rim of pit crater downslope from West Camp, upper Kipahulu Valley, Hale- akala National Park, 6,050 ft. [1,845 m.], 20°43.18’N, 156°07.73'’W, 19 May 1998, 13:00—17:00 hrs., CL 8333, D. A. Polhe- mus, B. H. Gagné and R. Takumi (USNM); 1 3,3 2, Kipahulu Valley, West Camp, Ha- leakala National Park, 1,850 m. [6,070 ft.],
28 February 1984, at night, EK G. Howarth (BPBM); | 2, same data as preceding but 12 July 1983, on tree trunk, E G. Howarth (BPBM); 3 2, same data as preceding but 17 July 1983, under wet ohia (Metrosideros polymorpha) bark, EF G. Howarth (BPBM).
Etymology.—The name “kipahulu”’ is a noun in apposition and refers to the Kipa- hulu Valley type locality.
Discussion.—In addition to the charac- ters given in the key and diagnosis, Saicella kipahulu may be recognized by its brown and black coloration in living individuals, and by the scattered patches of appressed golden hairs on the anterior pronotal cal- losities.
On the basis of current collections, this species appears to be confined to Kipahulu Valley, a deep, cliff-bound gap cutting the eastern face of Haleakala. All collections have come from the upper section of the
14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
valley, in the highest forest zone near the transition to the subalpine grasslands.
Saicella usingeri Wygodzinsky (Figs. 5, 10)
Saicella usingeri Wygodzinsky 1966: 408.
Types.—Holotype, d¢ and allotype & from Kawaikoi Ridge, Kokee, Kauai, in Bishop Museum, Honolulu.
Diagnosis.—Recognized by the annulate first antennal segment; relatively long wing pads that reach to the apex of the scutellar spine when viewed from above; yellowish brown coloration with red markings on the abdomen; and short length of the seventh visible female abdominal tergite, which is rounded posteriorly rather than broadly an- gulate.
Redescription.—Micropterous male: General coloration golden yellowish brown with slight greenish overtones, darker brown or red markings present on head, thorax and abdomen; legs multiannulate with dark brown.
Length 6.10 mm, maximum width (across abdomen) 1.45 mm.
Head \ength/width = 0.75/0.60, covered by a thick layer of wooly, appressed gold setae; width of vertex 3.1 dorsal width of an eye (0.37/0.12); length of anterior lobe of head 2.2 dorsal length of an eye (0.37/ 0.17); eyes small, consisting of approxi- mately 20 ommatidia each; length of pos- terior lobe of head 2.06 dorsal length of an eye (0.35/0.17); ocelli absent; length of antennal segments I-IV = 4.10/3.50/0.80/ 0.60; rostrum length 1.00, reaching to bases of fore coxae; anterior lobe of head dark yellowish brown, posterior lobe of head dark brown; antennal segment I dark yel- low, bearing 6 evenly spaced dark brown annulations of varying size, tip light; anten- nal segment II dark yellow, bearing 6 even- ly spaced dark annulations, tip dark except at extreme apex; antennal segments III and IV uniform medium brown except narrowly pale at extreme bases and tips.
Pronotum length (midline)/width = 0.78/
0.87; anterior section golden brown cen- trally, becoming embrowned toward pleu- rae, lateral lobes shining, bearing scattered patches of wooly, appressed gold setae, in- tervening longitudinal sulcus bearing nu- merous appressed gold setae; posterior sec- tion of pronotum yellowish brown, lacking evident setae, bearing a low longitudinal ca- rina medially rising to a small, conical tu- mescence posteromedially.
Scutellum triangular, golden yellowish brown, central section paler yellowish, pro- duced into an erect, slender, backward an- gling spine posteromedially, this spine an- gulate near middle when viewed laterally; a ring of pale, appressed gold setae present around base of spine; length (including spine)/width = 0.45/0.27.
Hemelytra short, micropterous, reaching to tip of scutellar spine when viewed from above, separated medially by an erect, slen- der, pale yellowish, backward angling spine arising from the underlying mesonotum, this spine similar in size, shape and orien- tation to that arising from scutellum; wing pads consisting of tiny, elongate pads, wid- ening somewhat posteriorly, venation and membrane absent; coloration medium brown, margins slightly lighter.
Legs elongate, with fore coxa approxi- mately 4.5 as long as thick (0.90/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 7.5 X longer than wide (1.87/0.25); coloration of legs dark yellow with brown or black mark- ings; fore femur yellowish on basal *%, with a broad brown annulation on distal % of outer face; fore tibia with 3 diffuse brown annulations on outer face, knee pale, apex dark; middle and hind femora each with 4 brown annulations, basal annulation broad and diffuse, all annulations shorter than in- tervening pale areas; middle and hind tibiae each with 10—11 brown to black annula- tions on outer faces, these annuli of des- creasing size distally but regularly spaced on middle tibia, more irregularly spaced and concentrated distally on hind tibia; middle and hind tarsi uniform medium
VOLUME 102, NUMBER 1
HAWAII
Fig. 13. Distribution of Siacella mulli on Hawaii.
brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.87/ 1.35/0.12/0.20; middle femur/tibia/tarsal 1/ tarsal 2/tarsal 3 = 3.75/5.40/0.07/0.07/0.07; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 5.00/8.20/0.10/0.10/0.10.
in
Abdomen golden yellowish brown with greenish overtones, this ground color over- lain with irregular dark brown to black markings anteromedially on all tergites, markings on first, second, third, and sixth visible tergites more extensive, these mark-
16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ings extending outward along anterior ter- gite margins to form transverse bands, giv- ing the abdomen a transversely striped ap- pearance; paratergites irregularly blotched with red on central sections, with dark markings on posterolateral angles; connex- ival margins broadly arcuate, lacking knobs at sutures; first visible tergite broadly raised anteromedially but not bearing a tumes- cence, all other tergites flat, lacking raised tumescences, sixth visible tergite elongate and triangular, longitudinally keeled on posterior half; all tergites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae.
Ventral surface yellowish brown mottled with dark brown, covered by an obscure layer of short, pale, recumbent setae; ab- dominal paratergites with spiracles creamy white.
Male genitalia with paramere club-like, bearing a small lateral tooth near tip; apex of pygophore strongly and suddenly nar- rowed, tip acuminate (see Wygodzinsky 1966: Figs. 124b, c).
Micropterous female: Length 5.50 mm, maximum width (across abdomen) 1.75 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posterior mar- gin of sixth visible tergite VI bearing a V- shaped indentation medially, seventh visi- ble tergite short, broadly rounded (Fig. 5).
Distribution.—Kauai (Alakai Plateau) (Fig. 10).
Ecological associations.—A series of this species was taken near upper Koaie Stream by applying a light pyrethrin fog to mossy, fallen ohia (Metrosideros polymorpha) trunks, in the same manner used to obtain the Maui specimens discussed previously. Wygodzinsky (1966) states that Usinger ob- tained his type series by beating tree ferns (Cibotium sp.). Label data from another specimen collected by Swezey indicates it came from Tetraplasandra, an arborescent native forest plant with broad leaves.
Material examined.—HAWAITIAN IS- LANDS, Kauai: 3 46, rainforest along
Koaie Stream at USGS gauging station, near crossing of Mohihi—Waialae Trail, 1;130° om: (3, 700] sili 22-06 47 ANE 159°35'18"W, 7 January 1999, D. A. Pol- hemus (USNM); 1 6, 1 2, Alakai Swamp Trail, Kawaikoi Ridge, Kokee, 18 August 1961, tree ferns, R. L. Usinger (holotype and allotype, BPBM); 2 3, Kualapa, 2,000 ft. [610 m.], 23 October 1979, S. L. Mont- gomery (BPBM); | 2, Kokee, 4-6 August 1961, Maa, Miyatake and Yoshimoto (BPBM); 1 3, 2 immatures, Kokee, 20 Au- gust 1925, on Tetraplasandra, O. H. Swez- ey (BPBM).
Discussion.—Saicella usingeri is endem- ic to Kauai; specimens are at hand from both the Alakai Plateau and the Makaleha Mountains, indicating that this species may occur on all the high mountain massifs of the island, including Namolokama and Mt. Kahili.
Saicella mulli Polhemus, new species (Figs. 4, 13)
Diagnosis.—This species may be recog- nized by the near-absence of wing pads, and by the unusual condition of the seventh vis- ible female abdominal tergite, which is di- vided into two lobes by a longitudinal me- dian sulcus (Fig. 4).
Description.—Micropterous male: Gen- eral coloration yellowish brown, with dark- er brown markings on head, thorax and base of abdomen; legs multiannulate with dark brown.
Length 5.50 mm, maximum width (across abdomen) 1.15 mm.
Head \ength/width = 0.70/0.55, covered by a thick layer of short, pale, recumbent setae; width of vertex 3.5 dorsal width of an eye (0.32/0.09); length of anterior lobe of head 2.36 dorsal length of an eye (0.37/0.11); eyes small, consisting of ap- proximately 20 ommatidia each; length of posterior lobe of head 2.63 dorsal length of an eye (0.29/0.11); ocelli absent; length of antennal segments I-IV = 3.10/2.75/ 0.85/0.60; rostrum length 0.98, reaching to bases of fore coxae; coloration of head yel-
VOLUME 102, NUMBER 1
lowish brown, darker along longitudinal midline on frons and vertex; antennal seg- ment I dark yellowish, distal ”; medium brown, basal 7% with 5 or 6 small brown annulations, antennal segments II and IV uniformly light brown.
Pronotum length (midline)/width = 0.72/ 0.69; anterior section medium brown cen- trally, lateral lobes shining, bearing scat- tered patches of wooly, appressed gold se- tae only anterobasally, intervening longitu- dinal sulcus bearing numerous appressed gold setae; posterior section of pronotum golden yellowish brown, lacking evident setae, broadly domed centrally but not pro- duced to a sharp tumescence.
Scutellum triangular, medium brown, central section pale yellowish, produced into an erect, slender, backward angling spine posteromedially, this spine angulate near middle when viewed laterally; a ring of pale, appressed gold setae present around base of spine; length (including spine)/ width = 0.26/0.27.
Hemelytra very short, micropterous, dif- ficult to discern, not reaching to base of scutellar spine, separated medially by an erect, slender, backward angling spine aris- ing from the underlying mesonotum, this spine similar in size, shape and orientation to that arising from scutellum but not as sharply angulate near middle when viewed laterally.
Legs elongate, with fore coxa approxi- mately 3.85 as long as thick (0.77/0.20); fore trochanter bearing numerous slender erect gold setae; fore femur fusiform, over 7.4X longer than wide (1.62/0.22); colora- tion of legs golden yellowish brown with dark brown markings; fore femur yellowish brown basally, with a broad, diffuse brown annulation on distal 4%; fore tibia with 3 dif- fuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown annulations, most basal of these an- nulations diffuse, all annulations roughly equal in length to intervening pale spaces; middle and hind tibiae with 8 and 10 brown annulations respectively, these annuli con-
17
centrated in the basal *% of these segments; middle and hind tarsi uniform brown; lengths of leg segments as follows: fore fe- mur/tibia/tarsal 1/tarsal 2 = 1.62/1.29/0.07/ 0.25; middle femur/tibia/tarsal 1/tarsal 2/ tarsal 3 = 3.10/4.50/0.10/0.10/10; hind fe- mur/tibia/tarsal 1/tarsal 2/tarsal 3 = 4.50/ 6.60/0.08/0.08/0.10.
Abdomen with lateral margins broadly ar- cuate, all tergites lacking raised tumescenc- es posteromedially, first through sixth visi- ble segments dorsally concave, paratergites curved upward and narrowly emarginate, posterior margin of fifth visible tergite broadly V-shaped, sixth visible tergite elon- gate, triangular, domed along longitudinal midline on posterior half; all tergites shin- ing, covered by a sparse, obscure layer of very short, pale, recumbent setae; colora- tion yellowish brown, irregularly mottled with darker brown.
Ventral surface medium brown, covered by a sparse layer of short, recumbent gold setae; abdominal paratergites light brown, spiracles whitish.
Male genitalia with paramere relatively slender, broadly curving; tip of pygophore narrowed and elongate, apex rounded.
Micropterous female: Length 5.30 mm, maximum width (across abdomen) 1.20 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posterior mar- gin of sixth visible tergite broadly V- shaped, rising to a knob-like tumescence posteromedially; seventh visible tergite short, bisected into two lobes along longi- tidinal midline by a deep sulcus, posterior margin broadly rounded (Fig. 4).
Distribution.—Hawaii (Volcano area) (Fig. 13).
Ecological associations.—Taken on mossy trunks of ohia trees (Metrosideros polymorpha).
Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Hawaii, reared from eggs laid by female taken at Puu Makaala, 3,700 ft. [1,130 m.], 10 Au- gust 1975, on mossy bark of ohia [Metros-
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ideros polymorpha], W. P. Mull (BPBM). Paratypes: HAWAIIAN ISLANDS, Ha- waii: 1 2, same data as holotype (BPBM).
Etymology.—Named in honor of Wil- liam Mull, who collected the only known specimens and reared numerous individuals through all life stages.
Discussion.—Saicella mulli is endemic to Hawaii; the only known series was reared from eggs laid by a single female captured at Puu Makaala, near Volcano. It seems likely that this species will also be found to inhabit windward Mauna Kea and the Kohala Mountains.
Saicella perkinsi Polhemus, new species (Figs. 6, 11)
Diagnosis.—Recognized by the long, pale, pilose hairs on the pronotum, and the absence of annulations on the middle and hind tibiae, and the enlarged, triangular form of the seventh visible male abdominal tergite (Fig. 6).
Description.—Micropterous female: General coloration golden yellowish brown, with darker brown markings on head, thorax and abdomen, red markings laterally on abdominal tergites; legs mul- tiannulate with dark brown.
Length 6.10 mm, maximum width (across abdomen) 1.50 mm.
Head \ength/width = 0.81/0.55, covered by a thick layer of appressed, wooly golden setae, interspersed with numerous long, pale, erect pilose setae; width of vertex 4.0X dorsal width of an eye (0.40/0.10); length of anterior lobe of head 3.1 dorsal length of an eye (0.44/0.14); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 2.6 dorsal length of an eye (0.37/0.14); ocelli absent; length of antennal segments I-IV = 4.10/3.75/0.95/0.70; rostrum length 0.97, reaching to bases of fore coxae; coloration of head uniform golden brown, without contrasting markings; antennal segment I uniform golden brown, without annula- tions; antennal segments II-IV medium brown, also lacking annulations.
-
Pronotum length (midline)/width = 0.81/ 0.90; anterior section golden brown, lateral lobes with thick linear patches of appressed, wooly, golden setae, intermixed with nu- merous long, pale, erect, pilose setae, inter- vening longitudinal sulcus also bearing nu- merous appressed gold setae; posterior sec- tion of pronotum golden brown, bearing nu- merous long, pale, erect, pilose setae, swollen centrally, not produced to a tumes- cence.
Scutellum triangular, lateral angles me- dium brown, central section golden brown, produced into an erect, slender, nearly ver- tical spine posteromedially, this spine straight when viewed laterally; numerous long, pale, erect pilose setae present around base of spine; length (including spine)/ width = 0.15/0.25.
Hemelytra short, micropterous, reaching to base of erect, slender, golden brown, spine arising nearly vertically from the un- derlying mesonotum, this spine similar in size, shape and orientation to that arising from scutellum; wing pads consisting of tiny flaps, widening slightly on posterior halves, venation and membrane absent; col- oration medium brown, set with a few long, pale, erect, pilose setae.
Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (1.0/0.25); fore femur fusiform, 7.4 longer than wide (2.00/0.27); coloration of legs golden brown with dark brown markings; fore fe- mur yellowish brown on basal %, with a broad brown annulation on distal %; fore tibia with 3 diffuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown annulations, these an- nulations roughly equal to length of inter- vening pale areas; middle and hind tibiae uniform medium brown, lacking annula- tions; middle and hind tarsi uniform medi- um brown; lengths of leg segments as fol- lows: fore femur/tibia/tarsal 1/tarsal 2 = 2.00/1.00/0.12/0.22; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 3.90/5.25/0.06/0.06/ 0.07; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 5.70/8.90/0.06/0.06/0.07.
VOLUME 102, NUMBER 1
Abdomen yellowish brown, this ground color overlain with irregular dark brown markings centrally on all tergites and pos- terolaterally on paratergites; connexival margins broadly convex, evenly curving; first visible abdominal tergite strongly flexed upward anteromedially, remaining tergites flat, lacking tumescences, poste- rior margin of sixth visible tergite with small V-shaped indentation medially, sev- enth visible tergite triangular (Fig. 6); all tergites shining, covered by a sparse, ob- scure layer of very short, pale, recumbent setae.
Ventral surface of head, thorax and ab- domen golden brown, covered by an ob- scure layer of short, pale, recumbent setae, intermixed on mesosternum with numerous long, pale, erect, pilose setae; abdominal paratergites irregularly marked with red bordering connexival margins.
Micropterous male: Unknown.
Distribution.—Oahu (Waianae Moun- tains) (Fig. 11).
Ecological associations.—Taken on mossy tree trunks at night.
Material examined.—Holotype, microp- terous 2, HAWAIIAN ISLANDS, Oahu, Waianae Mountains, Kaluaa Gulch, | mi. SE of Puu Hapapa, 700 m. [2,300 ft.], 10 October 1976, on mossy tree trunk, E G. Howarth (BPBM). Paratype: HAWAIIAN ISLANDS, Oahu: 1 2, Waianae Kai Forest Reserve, Kamaileunu Ridge, 720 m. [2,360 ft.], 19 October 1975, north side of ridge on tree trunk at night, W. C. Gagne (BPBM).
Etymology.—Named in honor of R. C. L. Perkins, whose pioneering collections of native Hawalian insects have given us an exceptional historical perspective regarding the islands’ insect fauna.
Discussion.—Saicella perkinsi is endem- ic to Oahu, and the only specimens so far known have come from the leeward slopes of the Waianae Range. This is one of the drier portions of the island, and it seems curious that this or a related species has not
19
been taken in the wet forests of the Koolau Mountains on eastern Oahu.
The long pilose hairs on the pronotum are present in one of the two specimens at hand, but mostly lacking in the other. It is presumed that in the latter case these hairs, being slender and delicate, were rubbed off during the specimen’s collection or subse- quent handling.
ACKNOWLEDGMENTS
This revision is based primarily on ma- terial collected by the author during field surveys supported by the Smithsonian Insti- tution’s Drake Fund (USNM), or held in the collections of the Bernice P. Bishop Muse- um, Honolulu, Hawaii (BPBM). It could not have been completed without the generous cooperation of the staff in the Department of Entomology at the latter institution, partic- ularly Dr. Scott E. Miller (now with ICIPE, Nairobi), David Preston, Dr. Francis G. Ho- warth, and Gordon Nishida. I also thank Dr. James K. Liebherr of Cornell University (CUIC) and Curtis Ewing, from the Univer- sity of Hawaii, for companionship and as- sistance in the field, and the staff of Pacific Helicopter Tours, Inc., particularly pilots Tom Hauptman and Eric Pacheco, who braved difficult weather over rugged terrain to convey us to sampling sites on the wind- ward slopes of Haleakala. Their skill and de- votion are greatly appreciated.
LITERATURE CITED
Clague, D. A. and G. B. Dalrymple. 1987. The Ha- waiian-Emporer volcanic chain. Part I. Geologic evolution, pp. 5-54. In Decker, R. W,, T. L. Wright, and P. H. Stauffer, eds., Volcanism in Ha- wail, Vol. 1. USGS Professional Paper 1350. U.S. Government Printing Office, Washington, DC. 839 pp.
Langenheim, V. A. M. and D. A. Clague. 1987. The Hawaiian-Emporer volcanic chain. Part II. Strati- graphic framework of volcanic rocks of the Ha- waiian Islands, pp. 55—84. Jn Decker, R. W., T. L. Wright, and P. H. Stauffer, eds. Volcanism in Ha- waii, Vol. 1. USGS Professional Paper 1350. U.S. Government Printing Office, Washington, DC. 839 pp.
Leston, D. 1957. Spread potential and the colonisation of islands. Systematic Zoology 6: 41—46.
20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Usinger, R. L. 1958. A new genus of Reduviidae from the Hawaiian Islands (Hemiptera). Proceedings of the Hawaiian Entomological Society 16: 437_ 440.
Wygodzinsky, PW. 1966. A monograph of the Eme- sinae (Reduviidae, Hemiptera), Bulletin of the American Museum of Natural History 133: ]— 614.
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 21-49
REVISION OF LOBOGENESIS RAZOWSKI AND ODONTHALITUS RAZOWSKI (LEPIDOPTERA: TORTRICIDAE: TORTRICINAE), WITH COMMENTS ON THEIR MONOPHYLY
JOHN W. BROWN
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Washington, DC 20560- 0168, U.S.A (e-mail: jbrown @sel.barc.usda.gov)
Abstract.—The Neotropical tortricid genera Lobogenesis Razowski and Odonthalitus Razowski are revised. Eight species are included in Lobogenesis: L. lobata Razowski (type species) from Costa Rica and Panama; L. penai, new species, from Cochabamba, Bolivia; L. magdalenana, new species, from Colombia and Venezuela; L. larana, new species, from Lara Province, Venezuela; L. contrasta, new species, from Cochabamba, Bolivia; L. antiqua, new species, from Cochabamba, Bolivia; L. peruviana, new species, from Peru; and L. varnicosa, new species, from Argentina. Nine species are included in Odonthalitus: O. lacticus Razowski (type species) from Durango, Mexico; O. bisetanus, new species, from Oaxaca, Mexico; O. improprius, new species, from Oaxaca, Mexico; O. conservanus, new species, from Jalisco, Mexico; O. orinoma (Walsingham), new combination, from Guerrero, Mexico; O. fuscomaculatus, new species, from Michoacan, Mexico; O. poas, new species, from Alajuela Province, Costa Rica; O. viridimontis, new species, from Puntarenas Province, Costa Rica; and O. regilla (Walsingham), new com- bination, from Guatemala. While the monophyly of Lobogenesis is well supported by characters of the male and female genitalia, Odonthalitus, as currently defined, may be paraphyletic with respect to Lobogenesis.
Key Words: Phylogeny, systematics, South America, Central America, Anopina, Euliini,
new species, genitalia
Razowski (1992) proposed Odonthalitus to accommodate the single species O. lac- ticus Razowski, described from two speci- mens (1 6, 1 @) collected in Durango, Mexico. In the same paper, he described the monotypic genus Lobogenesis for the new species L. lobata Razowski, represented by a single male from Costa Rica. Although the male genitalia are moderately divergent, these two monotypic genera share a variety of morphological features and are nearly identical in facies. During the course of continued studies on the systematics of the New World Euliini, I discovered numerous
undescribed and two described (misplaced) species that are morphologically and super- ficially similar to these two genera, stimu- lating an assessment of the phylogenetic re- lationship between the two genera and de- scriptions of the new species. A phyloge- netic hypothesis based primarily on structures of the male and female genitalia indicates that while Lobogenesis is mono- phyletic, its separation from Odonthalitus may leave the latter paraphyletic. Although similar in facies to Anopina Obraztsov, the male and female genitalia of Odonthalitus and Lobogenesis are highly
22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
divergent from all other Euliini, with a bi- furcate uncus in the male and greatly re- duced apophyses anteriores in the female. The subbasal attachment of the socius and the free (unjoined) distal arms of the gna- thos of Lobogenesis are reminiscent of some Sparganothini. However, these char- acter states are unlikely to be homologous with those found in Sparganothini, as the two groups have little else in common. Some females of Odonthalitus have bilobed papillae anales reminiscent of those found in some species of Apotoforma Busck (Tor- tricini) (see Razowski 1984, 1993); the greatly reduced apophyses anteriores also are similar in the two. However, because the groups share no other derived character states and male genitalia of the two are highly divergent, these similarities in fe- male genitalia are suspected to represent convergence rather than evidence of com- mon ancestry. If the bilobed papillae anales and greatly reduced apophyses eventually are demonstrated to be homologous _ be- tween Odonthalitus and Apotoforma, it would provide evidence for the placement of Odonthalitus and Lobogenesis in Tortri- cini rather than Euliini.
The purposes of this paper are to describe 7 new species of Lobogenesis and 6 new species of Odonthalitus, propose two new combinations in Odonthalitus, present a hy- pothesis of the phylogenetic relationships among the species, and modify the descrip- tions of the genera based on the new taxa.
MATERIALS AND METHODS
Specimens (n = 90) were obtained from the following institutions: The Natural His- tory Museum (BMNH), London, England; Instituto Nacional de Biodiversidad (IN- Bio), Santo Domingo, Heredia, Costa Rica; San Diego Natural History Museum (SDNHM), San Diego, California, U.S.A.; Essig Museum of Entomology, University of California, Berkeley (UCB), U.S.A; Na- tional Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C., U.S.A.
Dissection methodology follows that summarized in Brown and Powell (1991). Illustrations of genitalia were drawn with a camera-lucida attachment on a dissecting microscope. Forewing measurements were made with an ocular micrometer mounted in a dissecting microscope. Terminology for wing venation and genitalic structures fol- lows Horak (1984). Abbreviations and sym- bols are as follows: FW = forewing; HW = hindwing; DC = discal cell; n = number of specimens examined; ca. (circa) = ap- proximately; « = mean. Upper side refers to dorsal surface of wings, under side to ventral surface. Because most species are extremely similar in facies, not all are il- lustrated. In contrast, male and female gen- italia are diagnostic and are illustrated for each species. Dissections of the genitalia are required for accurate species determi- nations, and comparison with the illustra- tions is the best means for identifying spec- imens.
PHYLOGENY
A phylogenetic analysis was conducted on the 17 taxa recognized as species (plus a putative out-group—Anopina Obraztsov). The analysis was based on 23 morpholog- ical characters, 15 binary and 8 multi-state, that vary among the species of Lobogenesis and Odonthalitus, and exhibit shared, de- rived states at the species level. These in- clude 1 character of the head, 2 characters of the thorax, 15 characters of the male gen- italia, and 5 characters of the female geni- talia. Character state polarity was deter- mined using the out-group method. Because sister group relationships are poorly under- stood within Euliini (e.g., Brown and Pow- ell 1991), selection of an out-group was somewhat arbitrary. The genus Anopina Obraztsov was chosen because of its similar forewing pattern and long antennal cilia in the male. Horak’s (1984) assessment of tax- onomically significant structures in Tortri- cinae was used for determination of char- acter state polarity for characters that could not be determined convincingly using the
VOLUME 102, NUMBER 1
out-group (e.g., uncertain homology, both states present in out-group). The characters, along with their putative plesiomorphic and apomorphic conditions, are presented in Ta- ble 1. The distribution of the character states among the 17 taxa and the out-group is presented in Table 2; ‘*?” is used where the character state is unknown (e.g., char- acters for species represented by a single sex).
The data set was subjected to parsimony analysis using Hennig86 version 1.5 (Lip- scomb 1994), employing the “‘mhennig*”’ command. This algorithm constructs trees, each by a single pass through the data, by adding the taxa in a different sequence each time, and then applies branch-swapping to each of the trees. The analysis generated 6 equally parsimonious trees with a length of 48, a consistency index of 0.75, and a re- tention index of 0.85. Because over half of the taxa (n = 9) are represented by a single sex, there is a considerable amount of miss- ing data, which resulted in the equivocal placement of several taxa.
The cladogram illustrated in Fig. | rep- resents the “‘majority rule’’ consensus tree (see Lipscomb 1994) and is identical to one of the six trees. It is considered the best working hypothesis of the phylogeny of the species. The analysis demonstrates that the monophyly of Lobogenesis is well support- ed (i.e., in all 6 trees), while Odonthalitus, as current defined, may be paraphyletic (i.e., in all 6 trees). The hypothesis of the phylogeny is fairly concordant with the geographic distribution of the two genera: with the exception of L. lobata (from Costa Rica and Panama), Lobogenesis appears to be restricted to South America, while Odonthalitus is entirely Central American (including northern Mexico) in distribution.
Superficially adults of Odonthalitus are indistinguishable from those of Lobogene- sis. Synapomorphies for the two include the forewing pattern, loss of the male foreleg hairpencil, elongate cilia of the male anten- na, bifurcate uncus, and reduced apophyses anteriores. The monophlyly of Lobogenesis
23
is supported convincingly by the following: uncus bifurcations long, slender, and atten- uate apically; gnathos arms free distally or only weakly attached by membrane; socius large, subbasally attached, with dorsal lobe; and linear row of tiny spinelike teeth in api- cal region of valva. In contrast, Odonthal- itus Was portrayed as paraphyletic in each of the’6 ‘trees:
There are at least three possible ways to resolve the paraphyly. One solution would be to consider the entire clade as a single genus to which either Odonthalitus or Lo- bogensis could be applied (based on the choice of the first revisor). Another would be to include lacticus and regilla with the species considered Lobogenesis in Fig. 1, which would result in the synonymy of Lo- bogenesis with Odonthalitus, and to de- scribe a new genus for the remaining Cen- tral American species. The third would be to maintain Lobogenesis as shown in Fig. 1, restrict Odonthalitus to the species lac- ticus and regilla, and propose a new genus for the remaining Central American species. However, for the present the generic status of both Lobogensis and Odonthalitus are re- tained for a number of reasons. First, the extremely long, slender aedeagus of regilla and the notched papillae anales of lacticus are extremely similar to those of O. viridi- montis and O. poas, suggesting a closer phylogenetic relationship of these species than is portrayed in the cladogram. Second- ly, from a biogeographic perspective, one would expect /acticus and regilla (both from Mexico) to be more closely allied to Odonthalitus than to Lobogenesis. Finally, because the data set for the analysis con- tains so much missing data (i.e., several species are represented by a single sex), any classification based on the results of the parsimony analysis likely would be inher- ently unstable. Hence, a conservative ap- proach is employed that maintains the sta- bility of the generic nomenclature of the
group.
24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Characters for phylogenetic analysis of Odonthalitus and Lobogenesis; 0 = plesiomorphic state, 1— 3 = apomorphic states; figure references to apomorphic condition(s), where illustrated.
1. Head O—length of male antennal cilia equal to width of flagellomere 1—length of male antennal cilia 3 times width of flagellomere 2. Thorax O—male foreleg hairpencil present 1—male foreleg hairpencil absent 3. Forewing O—pattern variable 1—pattern with distinctive costal and basal patch 4. Uncus O—unmodified, without bifurcate distal portion
1—bifurcate in distal one-half (Figs. 2—8)
2—bifurcate from base (Figs. 14—18)
3—bifurcations widely separate at base (Figs. 19—20) 5. Uncus O—tips of bifurcation round or blunt
1—tips of bifurcation narrowed and pointed (Figs. 2—5)
2—1ips with narrow triangular enlargement (Figs. 6—8) 6. Gnathos Q—gnathos arms joined distally
1—-gnathos arms weakly joined distally (Figs. 14—16)
2—gnathos arms separate distally (Figs. 2—8, 19-20)
7. Gnathos OQ—arms without lateral process(es) 1—arms with lateral lobes (Figs. 17—18) 8. Socius O—simple, small, slightly digitate
1—large, with moderate dorsal lobe (Figs. 2—5) 2—huge, with large dorsal lobe (Figs. 6-8)
9. Transtilla O—simple bridge 1—lost (Figs. 17—20) 10. Valva Q—simple at base
1—with patch of curved spines at base of costa (Figs. 14-15) 2—with large hook at base of casta (Fig. 16) 3— costa with free hairy lobe near middle (Figs. 17—8) 11. Valva O—costa simple 1—costa with free flange ca. 0.67 distance from base (Figs. 3—5)
2— flange enlarged (Fig. 4) 12. Valva O—simple
1—dense patch of setae in middle of basal one-third (Figs. 4, 6-8) 13. Valva O—simple
1—curved, sclerotized line from patch of setae to apex (Figs. 6-8) 14. Valva Q—apex more or less rounded
1—apex attenuate, somewhat pointed (Figs. 14—18) 15. Valva O—unmodified
1—linear row of tiny spinelike teeth in apical region (Figs. 2—8) 16. Valva O—simple
1—with notch at venter (Figs. 19-20) 17. Aedeagus O—simple, unmodified
1—somewhat undulate (Figs. 3—5) 2— strongly bent, somewhat *‘J°’ shaped (Figs. 14—16)
18. Aedeagus O0—simple, unmodified
1—extremely long, slender (Figs. 17—19) 19. Sterigma O—simple
1—with scobinate lateral patches (Figs. 21—22) 20. Sterigma O—simple
1—rounded or v-shaped notch at ostium (Figs. 9, 12) 21. Sterigma O—simple
1—-with sclerotized arch surrounding ostium (Figs. 10-11, 13) 22. Papillae O—simple, parallel-sided
1—with a mesal notch (Figs. 23—25) 23. Apophyses ant. O—unmodified, moderately long
1—reduced in length (Figs. 21—22) 2— reduced to a short nub (Figs. 9-13, 25—26) 3— lost (Figs. 23-24)
VOLUME 102, NUMBER 1
Table 2. Distribution of character states among species; see Table | for explanation of character states (? =
missing data).
Outgroup 0000 0000 lacticus Tas} 0200 regilla dais 0200 poas Deda 0010 viridimontis RaW Eal 0010 orinoma Deer? 2222 fuscomaculatus Fo plee) BPP conservanus LI 0100 bisetanus ua 0100 improprius 112 0100 contrasta iltaleallslk 2202 peruviana ita ak 22:02 antiqua iBall 2202 penai abaya AL 1201 lobata ibaa 1201 larana alga Lace 1201 magdalenana algal aleACO)atk varnicosa PAM? De De SYSTEMATICS
Lobogenesis Razowski 1992
Lobogenesis Razowski 1992: 213; Powell et al. 1995: 144.
Type species.—Lobogenesis lobata Ra- zowski 1992, by monotypy.
Redescription.—Head: Antennal cilia ca. 3.0 times width of flagellomere in male, ca. 0.5 times width of flagellomere in fe- male. Labial palpus (segments II and III combined) ca. 1.5 times horizontal diameter of compound eye; segment II weakly up- turned, slightly expanded distally by scal- ing; segment III ca. 0.4 as long as II, smooth-scaled, well exposed. Maxillary palpus rudimentary. Frons with overhang- ing tuft of scales. Ocelli small or absent. Chaetosema present. Proboscis present. Thorax: Smooth-scaled. Legs unmodified, male without foreleg hairpencil. Forewing: Length 2.3—2.4 times width; length of DC about 0.55 times FW length; width of DC about 0.20 DC length; CuA, originates about 0.60 along length of DC; all veins separate beyond DC; chorda and M-stem absent. No upraised scale tufts; male with- out costal fold. Hindwing: Sc+R and Rs closely approximate; M, and CuA, closely approximate; M-stem absent; tuft of hair-
0000 0000 0000 000 1000 0101 0000 012 1000 0001 01?? iMeue 1300 0100 0100 Oalbs 1300 0100 0100 013 Dye ONe Dp Be 2200 002 222? 2 Bee 220 O01 0200 0100 2010 001 0100 0100 20/2? Ree 0100 0100 202-2 Due? 0001 1010 00?? Dee, 0001 1010 0000 02 0001 1010 00?? 222 0000 0010 00?? eae 0010 0010 1001 002 0021 0010 1000 NO? 0010 0010 1001 002 apie 2 2D? ??00 102
like scales along 1A+2A, originating near base of wing. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in female. Male genitalia: Uncus bifurcate from a common, slender stalk; each arm slender, attenuate distally or with triangular enlarge- ment at tip. Socius broad and long, with dorsal lobe, not fused to gnathos. Gnathos simple, non-dentate, arms narrow, separate distally. Subscaphium and hami absent. Transtilla a simple band or enlarged, ill-de- fined bridge. Valva moderately short, slight- ly attenuate distally; row of tiny spinelike teeth in middle of apical region. Pulvinus absent. Vinculum complete. Juxta a scler- otized plate. Aedeagus moderately long, usually weakly undulate; phallobase sim- ple; cornuti absent. Female genitalia: Pa- pillae anales narrow. Apophyses posteriores moderately long to short; apophyses anter- iores reduced to a membranous nub. Sterig- ma a simple sclerotized band. Ductus bur- sae moderately long, membranous. Corpus bursae frail, elongate, pear-shaped; spicules and signum lacking. Accessory bursa fre- quently present.
Diagnosis.—Superficially, adults of Lo- bogenesis are indistinguishable from those of Odonthalitus. The two can be separated by the following characters of the male gen-
26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Odonthalitus Lobogenesis = = aie : =) a = = wn 3 @ = STS, Ss = S > «(68 paees eS Sisige s a S Ss OTs = Siti PSS Sy So Saw eee Bes | = us Be tS te SO OmeS: Se Se Oe a ee eee) ee = 8 Ny ~ iS) Y Sy Sy Os = So 3S = SOS SS oss SS Soe pees Sr oS FG ae See, SS @ Ss S856 85 Ss See oS eo as 16-14 19 ics aoe 10' a ae 11! 5! 18 9 -17 12 23. 20 Tait ual Ball fc; ae eae niet 17 22 16 iS 9 8 le = 1 aa. 10 6! ao 6 q 3 2 1 Fig. 1. Consensus cladogram of phylogeny of Lobogenesis and Odonthalitus. Prime (') and double prime (") refer to successive steps in multi-state characters; minus (—) refers to character state reversal.
italia: 1) paired uncus arms (bifurcations) always from a common stalk in Loboge- nesis, either from a common stalk or sepa- rated basally in Odonthalitus; 2) distal tips of paired uncus arms (bifurcations) ex- tremely slender, pointed or triangular dis- tally in Lobogenesis, blunt or rounded in Odonthalitus; 3) arms of gnathos free dis- tally in Lobogenesis, joined distally in
Odonthalitus, either as a single process or weakly attached by membrane; and 4) so- cius attached subbasally, with a large dorsal lobe in Lobogenesis, socius attached basally in Odonthalitus.
Distribution and biology.—Lobogenesis is known from montane forest habitat from Costa Rica south to Argentina, from 1,100— 3,000 m elevation. Adults have been col-
VOLUME 102, NUMBER 1
lected at lights. Nothing is known of the early stages. Three species are known to be sympatric in Bolivia (at Cochabamba) and two in Venezuela (at Rancho Grande).
Remarks.—Species of Lobogenesis are rare in collections; the 8 species treated be- low are represented by 37 specimens, 14 of which represent one species. Three species are known only from males and one only from females. Modern systematic revisions of Neotropical tortricids (e.g., Brown and Powell 1991, 2000) typically result in an increase in the number of recognized spe- cies by a factor of 3—4; the present treat- ment results in descriptions of 7 new spe- cies to accompany the single previously de- scribed species.
Lobogenesis penai J. Brown, new species (Figss2; 32)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale cream; roughened above, pale cream. La- bial palpus pale cream mesally, mostly brown laterally. Antennal scaling bronze. Thorax: Tan with dark brown tegula. Fore- wing: Length 6.8 mm (n = 1). Upper side pale cream, with irregular, pale brown striae throughout; basal 0.2 dark brown; dark brown, triangular patch bordering costa ca. 0.45—0.60 distance from base to apex. Un- der side nearly uniform dark tan with faint indication of upper side markings. Hind- wing: Upper side white, with faint gray- brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 2 (drawn from USNM slide 88523; n = 1). Uncus slender at base, bifurcate in distal 0.7, with extremely long, slender spines subapically. Socius large, hairy, with conspicuous portion dorsad of attachment. Gnathos arms extremely slender, elongate, separate distally. Transtilla large, membra- nous, with narrow sclerotized arch in ven- tral portion. Valva broadest at base, rounded apically; basal 0.5 of costa sclerotized. Ae- deagus short, somewhat pistol-shaped, with
27
a short lateral thorn; phallobase with small rounded membranous cap.
Female. Unknown.
Type.—Holotype, ¢, Bolivia, Incachaca, Cochabamba, tropical cloud forest area, 2,100.1 27-W I/S-1X- 1956 (LL Pena, USNM).
Diagnosis.—The male genitalia of L. penadi are most similar to those of L. larana and L. lobata. They can be distinguished from those species by the much longer, more slender gnathos arms and the elon- gate, slender bifurcations of the uncus, with long spines subapically (Fig. 2).
Etymology.—The species name is a pa- tronym for Louis Pena, a noted Chilean en- tomologist and the collector of the holo-
type.
Lobogenesis lobata (Razowsk1) (Figs. 3; 9)
Lobogenesis lobata Razowski 1992: 215; Powell et al. 1995: 144.
Redescription.—Male. Head: Frons with sparse, smooth scaling below mid- eye, pale light brown; roughened above, pale tan. Labial palpus pale tan mesally, brown mixed with tan laterally. Antennal scaling brown. Thorax: Whitish, with brown tegula. Forewing: Length 5.5 mm (n = 2). Upper side whitish, with faint, sparse tan overscaling; basal 0.2 brown; brown semicircular patch bordering costa ca. 0.45—0.65 distance from base to apex; termen with irregular brown striae. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side whitish gray with slightly darker gray- brown mottling. Under side light gray- brown with darker mottling. Genitalia: As in Fig. 3 (drawn from USNM slide 89442, Panama; n = 2). Uncus narrow at base, bifurcate in distal 0.65, each arm slender, elongate, pointed distally. Socius with free dorsal lobe. Gnathos arms slender, weakly joined near middle, rounded distally, to- gether forming an H-shaped process. Tran- stilla a narrow sclerotized band with a pair
28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 2-5. Male genitalia of Lobogenesis; valvae spread, aedeagus removed. 2, L. penai. 3, L. lobata. 4, L.
larana. 5, L. magdalenana.
VOLUME 102, NUMBER 1
of dorsally projecting thorns at middle. Valva nearly rectangular, rounded apically; costa with free flange near middle; narrow sclerotized ridge across face of valva ca. 0.65 distance from base to apex; small lin- ear patch of short, spinelike teeth in apical region. Aedeagus weakly undulate; phal- lobase with large, rounded, membranous cap.
Female. FW length 5.0 mm (n = 2). Su- perficially as in male, except thorax dark brown and antennal cilia short. Genitalia: As in Fig. 9 (drawn from USNM< slide 88441, Panama; n = 2). Papillae anales un- modified. Apophyses posteriores moderate; apophyses anteriores extremely short, semi- membranous. Sterigma a sclerotized band, with U-shaped excavation at ostium. Duc- tus bursae relatively long, slender, frail. Corpus bursae rounded.
Type.—Holotype, 6, Costa Rica, Puntar- enas Province, 6 km S San Vito, 20/27-IV- 1967 (D. Veirs, UCB).
Additional specimens examined.— COSTA RICA: Puntarenas Province: Fea. Cafrosa, Est. Las Mellizas, P. N. Amistad, 1,300 m, 1 2, X-1990 (M. Ramirez & G. Mora, INBio). PANAMA: Cerro Cam- pana, nr. Chica, 1 6, 2/5-IV-1965 (S. S. & W. D. Duckworth, USNM); Cerro Cam- pana, 1 2, 11/14-VII-1967 (O. S. Flint, USNM).
Diagnosis.—Lobogenesis lobata is most similar to L. larana and L. magdalenana in several features of the male genitalia. It can be distinguished from those species by the paired thorns from the middle of the transtilla. The sexes were associated by their common occurrence at Cerro Cam- pana.
Lobogenesis larana J. Brown, new species (Figse4. 133)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, pale tan. Labial palpus whitish mesally, mostly brown lat- erally. Antennal scaling brown. Thorax:
29
Whitish with dark brown tegula. Forewing: Length 5.2—5.8 mm (* = 5.4; n = 4). Upper side pale whitish, with variable, faint brown striae and reticulations, particularly dense in distal 0.5; basal 0.2 brown; brown semicir- cular patch bordering costa ca. 0.45—0.65 distance from base to apex; termen with ir- regular brown line. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale whit- ish gray with slightly darker gray-brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 4 (drawn from USNM slide 89444; n = 3). Uncus narrow at base, bifurcate in distal 0.65, each arm slender, pointed distally. So- cius with dorsal lobe. Gnathos arms slender, separate. Transtilla a moderate, evenly sclerotized band. Valva nearly rectangular, rounded apically; an attenuate flange from costa just beyond middle, ending in free, rounded process; sacculus represented by undulate line of sclerotization; a dense patch of setae in basal 0.33 of valva; small linear patch of spinelike teeth in apical re- gion. Aedeagus relatively slender, slightly undulate; phallobase with small membra- nous cap.
Female. FW length 6.5—7.0 mm (* = 6.8; n = 6). Superficially as in male, except brownish overscaling of forewing conspic- uously more dense, larger average forewing length, and lacking elongate antennal cilia. Genitalia: As in Fig. 11 (drawn from USNM slide 87880; n = 3). Papillae anales slender, unmodified. Apophyses posteriores slender; apophyses anteriores reduced, slen- der, semi-membranous. Sterigma a simple band with membranous region surrounding ostium, bordered by inverted U-shaped sclerotized region. Ductus bursae moderate- ly long. Corpus bursae oblong.
Types.—Holotype, ¢, Venezuela, Lara, Yacambu National Park, 13 km SE Sana- re, cloud forest, 4,800’ [1,500 m], black- light, 4/7-III-1978 (J. B. Heppner, USNM).
Paratypes, 3 6, 10 2. VENEZUELA: 3 6, 9 @, same data as holotype. Aragua
30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Province: Rancho Grande, 1,100 m, 1 2, 1/ 7-VIII-1967 (R. W. Poole, USNM).
Diagnosis.—The male genitalia of Lo- bogenesis larana are most similar to those of L. magdalenana and L. lobata (Figs. 3— 5) in the overall shape of the valva, the lin- ear patch of spinelike teeth in the apical re- gion of the valva, and the slender bifurca- tions of the uncus. The three can be sepa- rated by the presence of a dense patch of setae in the basal 0.33 of the valva in L. larana and the mesal pair of spines from the transtilla in L. lobata, both features of which are lacking in L. magdalenana.
Remarks.—The genitalia of the female from Rancho Grande differ slightly in the shape of the sterigma from those from Ya- cambu.
Etymology.—The specific epithet refers to the Venezuelan province of Lara.
Lobogenesis magdalenana J. Brown, new species (Bigs 257412,030. 37)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, pale bronze- brown. Labial palpus whitish mesally, brown mixed with tan laterally. Antennal scaling brown. Thorax: Brown. Forewing: Length 5.0 mm (n = 2). Upper side pale whitish, with faint brown striae in distal 0.5; basal 0.2 brown; brown semicircular patch bordering costa ca. 0.45—0.65 dis- tance from base to apex. Under side uni- form dark tan with faint indication of up- perside markings. Hindwing: Upper side whitish with plae gray-brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 5 (drawn from BMNH slide 29065, Colombia; n = 2). Uncus narrow at base, bifurcate in dis- tal 0.65, each arm slender, elongate, point- ed distally. Socius with dorsal lobe. Gna- thos arms slender, apparently joined dis- tally by membrane. Transtilla a narrow sclerotized bridge, weakly arched mesally. Valva nearly rectangular, rounded apically; costa with roughened flange in basal 0.5;
narrow sclerotized ridge across face of val- va, terminating in free costal flange; linear patch of short, spinelike teeth in apical re- gion. Aedeagus weakly undulate; phallo- base with large, rounded, membranous cap.
Female. FW length 5.1—6.5 mm (* = 5.9; n = 3). Superficially as in male, except forewing length larger, forewing with brown overscaling more dense, and lacking elongate antennal cilia. Genitalia: As in Fig. 12 (drawn from BMNH slide 29064; Colombia; n = 3). Sterigma a sclerotized band, with membranous area surrounding ostium; ostium indicated by a sclerotized V- shaped notch. Ductus bursae moderately long. Corpus bursae oblong.
Types.—Holotype, 6, Colombia, Mag- dalena, Sierra Nevada de Santa Marta, San Pedro de la Sierra, 1,500 m, 14-VIII- 1973, Oxford Expedition to Colombia (BMNH).
Paratypes, 1 ¢6, 2 2. COLOMBIA: Mag- dalena, Sierra Nevada de Santa Marta, San Pedro de la Sierra, 1,500 m, 1 @, 6-VII- 1997 (BMNH); Sierra del Libano, 6,000’ [1,846 mj]; B® 2, V-1899 (HB. “Hin Smith, BMNH).
Additional specimens examined.—VEN- EZUELA: Aragua: 1 km south of Rancho Grande; 1d, 5-11-1976 (C2 Ms & Os! Flint, USNM); Rancho Grande, cloud for- est, 1,100 m,; 1 2, 30/31-I11-1978 (J. B: He- ppner, USNM); T. EK Amazonas, Cerro de Neblina, Camp VII, 1,850 m, 0°51'N, 65°58'""W, 1 &, 2/4-XTI-1984 (R. Brown, USNM).
Diagnosis.—Lobogenesis magdalenana is most similar to L. larana and L. lobata; the differences in male genitalia are dis- cussed above in the diagnosis of L. larana. The female genitalia can be distinguished by the shape of the mesal notch of the ste- rigma: V-shaped in L. magdalenana and more U-shaped in L. larana.
Remarks.—Male and female genitalia of the specimens from Venezuela, deviate slightly from those of specimens from Co- lombia: the linear row of fine spinelike
VOLUME 102, NUMBER 1
31
Figs. 6-8. Male genitalia of Lobogenesis: valvae spread, aedeagus removed. 6, L. peruviana. 7, L. antiqua. 8, L. contrasta.
32) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
teeth in the apical region of the valva con- sists of considerably fewer teeth in the male, and the antrum is slightly more rounded in the female. In addition, fore- wing length is conspicuously less in the fe- male from Rancho Grande. However, be- cause the genitalic differences are consid- erably less than that between other species of Lobogenesis, they are interpreted as geo- graphic, infraspecific variation within L. magdalenana. Additional specimens are necessary to determine whether or not this interpretation is correct.
Etymology.—The specific epithet refers to the province of Magdalena.
Lobogenesis contrasta J. Brown, new species (Figs. 8, 27)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, tan-brown; roughened above, dark cop- per-brown Labial palpus pale tan mesally, dark red-brown laterally. Antennal scaling brown. Thorax: Mostly dark brown. Forewing: Length 6.8—-8.0 mm (x = 7.2; n = 6). Upper side white, with faint, dif- fuse, pale yellow-tan overscaling; basal 0.2 dark brown with some red-brown scales; dark brown, semicircular patch bordering costa ca. 0.45—0.60 from base; two small, brown, triangular dots on costa between large costal patch and apex; ter- men with diffuse brownish striae, with a few pale yellow-green scales; a varibale, ill-defined line from dorsum near tornus. Under side nearly uniform tan-brown. Hindwing: Upper side dingy pale yellow with pale gray-brown mottling. Under side light gray-brown with darker mot- tling. Genitalia: As in Fig. 8 (drawn from USNM slides 68611 and 89440; n = 3). Uncus slender from broad, expanded cap on dorsum of tegumen, bifurcate in distal 0.5, with swollen, foot-shaped processes apically. Socius large, hairy, with con- spicuous portion dorsad of attachment. Gnathos comparatively slender, arms free distally, with shovel-shaped, attenuate tip.
Transtilla complete, more strongly scler- toized mesally. Valva broadest at base, slightly attenuate distally, with a pair of triangular lobes at costa, one at base, con- tiguous with transtilla, the second as a free rounded flange near mid-costa; a densely setose region in basal 0.33, con- tinuing toward apex as a long, curved, sclerotized line. Aedeagus relatively slen- der, slightly curved; phallobase with a large, rounded, membranous cap.
Female. Unknown.
Type.—Holotype, 6, Bolivia, Incachaca, Cochabamba, tropical cloud forest area, 2,100 m, 27-VIII/5-IX-1956 (L. Pena, USNM).
Paratypes, 5 ¢, same data as holotype.
Diagnosis.—Lobogenesis contrasta 1s most similar to L. peruviana and L. antiqua in size, forewing maculation, and male gen- italia. It be can be distinguished from L. peruviana by its simple aedeagus—that of L. peruviana has a distinct ventral thorn from near the middle. It can be separated from L. antiqua by the shape of the scler- otized ridge across the face of the valva (see Figs. 6-7), which is more strongly curved in the latter.
Etymology.—The specific epithet is an adjective, referring to the contrast between the pale ground color and the forewing pat- tern elements.
Lobogenesis peruviana J. Brown, new species (Figs. 6, 10, 29)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish and pale tan; roughened above, pale tan. Labial palpus pale tan mesally, brown mixed with yellow-tan laterally. Antennal scaling brown. Thorax: Mostly gray white, with red-brown prothoracic collar and brown tegula. Forewing: Length 7.5 mm (n = 1). Upper side silver white, with faint pale yellowish overscal- ing; basal 0.2 dark red-brown; dark red- brown semicircular patch bordering costa ca. 0.45—0.60 from base, second smaller
VOLUME 102, NUMBER 1 33
S—
S — O
SS
“
Figs. 9-11. Female genitalia of Lobogenesis. 9, L. lobata. 10, L. peruviana. 11, L. larana.
34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
patch at ca. 0.75 distance from base; ter- men with diffuse gray-brown striae, with a few pale green and yellow sclaes. Under side nearly uniform tan brown. Hindwing: Upper side whitish with pale grayish mot- tling. Under side light grayish with darker mottling. Genitalia: As in Fig. 6 (drawn from BMNH slide 29061; n = 1). Uncus slender from enlarged cap on dorsum of tegumen, bifurcate in distal 0.5; each tip bearing an enlarged triangular process, with two long, fine, subapical setae. Socius large, hairy, with conspicuous portion dor- sad of attachment. Gnathos arms compar- atively slender, separate distally, each with a shovel-shaped, attentuate tip. Transtilla complete, weakly U-shaped, with small upturned flap from venter near middle. Valva broadest at base, slightly attenuate apically, with a slender, digitate projection from costa at base; densely setose area in basal 0.33, extending toward apex as curved, sclerotized line. Aedeagus undu- late, moderately broad, with distinct thorn from venter near middle; phallobase with large, rounded, membranous cap.
Female. FW length 8.0 mm (n = 1). Su- perficially as in male, except lacking elon- gate antennal cilia. Genitalia: As in Fig. 10 (drawn from BMNH slide 29062; n = 1). Papillae anales unmodified. Apophy- ses posteriores moderate in length; apoph- yses anteriores reduced to short, slender, semi-membranous nub. Sterigma a broad, sclerotized band, with inverted U-shaped membranous region surrounding ostium; ostium represented by small U-shaped sclerite. Ductus bursae relatively short, slender, frail. Corpus bursae rounded, pear-shaped.
Types.—Holotype, ¢, Peru, Pillahuata, Cuzco, 2,600 m, 14/16-VIII-1982 (M. Mat- thews & M. Packer, BMNH).
Paratype, 1 @, same data as holotype (BMNBH).
Diagnosis.—Lobogenesis peruviana is the largest species in the genus and is most similar to O. contrasta. It can be distin- guished by the characters cited above in the
diagnosis for O. contrasta. Autapomorphies for L. peruviana include the triangular api- cal processes and setae of the distal portion of the uncus, the upturned mesal ventral portion of the transtilla, and the thorn from the venter of the aedeagus.
Etymology.—The specific epithet is an adjective referring to the country of its cap- ture.
Lobogenesis antiqua J. Brown, new species (Figs. 7, 28)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, tan- brown; roughened above, bronze-brown. Labial palpus pale tan mesally, mostly brown laterally. Antennal scaling brown. Thorax: Dark brown mixed with red brown. Forewing: Length 7.8 mm (n = 1). Upper side dingy white, with faint pale tan-yellow overscaling; basal 0.2 dark brown; dark brown semicircular patch bor- dering costa ca. 0.45—0.60 from base, two additional small costal spots between semi- circular patch and apex; termen with irreg- ular brownish reticulations. Under side uniform tan-brown with faint indication of upper side markings. Hindwing: Upper side whitish with pale grayish mottling. Under side light grayish with darker mot- tling. Genitalia: As in Fig. 7 (drawn from BMNH slide 29063; n = 1). Uncus slender from broad, enlarged cap on dorsum of tegumen; uncus missing distal one-half (assumed to be bifurcate). Socius large, hairy, with conspicuous portion dorsad of attachment; weakly sclerotized along outer edge. Gnathos arms comparatively slender, arms separate distally, each with a trian- gular tip. Transtilla complete, most strong- ly sclerotized mesally. Valva broadest at base, weakly attenuate apically, with two projections from costa, one at base contig- uous with transtilla, the second a free, rounded flange at ca. mid-costa; densely setose area in basal 0.33, extending toward apex as curved, sclerotized line. Aedeagus
VOLUME 102, NUMBER 1
Figs. 12-13.
nearly straight, relatively slender; phallo- base with broad, rounded, membranous cap.
Female. Unknown.
Type.—Holotype, 6, Bolivia, Yungas del Espiritu Santo, Cochabamba, 1888-89, Par- avicini Coll., BM 1937-383 (P. Germain, BMNH).
Diagnosis.—Lobogenesis antiqua is most similar to L. contrasta; superficially, it has a more yellowish tan forewing ground col- or. The male genitalia of L. antiqua can be
Female genitalia of Lobogenesis. 12, L. magdalenana. 13, L. varnicosa.
distinguished from those of L. contrasta by the conspicuously longer linear row of spinelike teeth in the apical region of the valva, the larger patch of setae in the basal 0.33 of the valva, and the more undulate sclerotized line extending from the setose patch to the apex of the valva (see Figs. 7— 8).
Etymology.—The specific epithet, from the word antique, is an adjective referring to the fact that the type specimen was col- lected more than 100 years ago.
36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Lobogenesis varnicosa J. Brown, new species (Figs= 1333)
Description.—Male. Unknown.
Female. Head: Frons with sparse, smooth scaling below mid-eye, whitish and pale tan; roughened above, pale tan. Labial palpus pale whitish mesally, light brown laterally. Antennal scaling brown. Thorax: Pale tan. Forewing: Length 6.5—7.2 mm (x = 6.8; n = 2). Upper side dingy white, with faint brownish overscaling and darker brown striae; basal 0.2 dark brown; dark brown triangular patch bordering costa ca. 0.45—0.60 from base, with a dark brown dash immediately below, near apex of DC. Under side nearly uniform tan brown. Hindwing: Upper side dingy whitish with pale gray-brown overscaling and mottling. Under side light grayish with darker mot- tling. Genitalia: As in Fig. 13 (drawn from USNM slide 89267; n = 2). Papillae anales unmodified. Apophyses posteriores moder- ate; apophyses anteriores reduced to short, slender, semi-membranous nub. Sterigma a broad, weakly sclerotized band, with in- verted U-shaped arch dorsad of ostium. Ductus bursae relatively long, undifferen- tiated from corpus. Corpus bursae long, moderately slender.
Types.—Holotype, 2, Argentina, Tucu- man, Ciudad Universitaria, 17-H-1959 (J. FE G. Clarke, USNM).
Paratype, 1 9, same data as holotype (USNM).
Diagnosis.—Superficially, Lobogenesis varnicosa is virtually indistinguishable from most other species in the genus. The genitalia are most like those of L. peruviana among described species. They are easily distinguished by the strongly sclerotized area around the ostium (see Figs. 10, 13).
Etymology.—The specific epithet is a manuscript name used by Josef Razowski for this species; its origin is unknown to me.
Odonthalitus Razowski 1992
Odonthalitus Razowski 1992: 208; Powell et al. 1995: 145.
Type species.—Odonthalitus lacticus Ra- zowski 1992, by monotypy.
Redescription.—Head: Antennal cilia ca. 3.0 times width of flagellomere in male, ca. 0.5 times width of flagellomere in female. Labial palpus (segments IT and III combined) ca. 1.5 times horizontal di- ameter of compound eye; segment II weakly upturned, slightly expanded dis- tally by scaling; segment III 0.3—0.4 as long as II, smooth-scaled, exposed. Max- illary palpus rudimentary. Frons with overhanging tuft of scales. Ocelli small or absent. Chaetosema present. Proboscis present, presumably functional. Thorax: Smooth-scaled. Legs unmodified, male without foreleg hairpencil. Forewing: Length 2.3—2.4 times width; length of DC about 0.55 times FW length; width of DC about 0.20 DC length; CuA, originates about 0.60 along length of DC; all veins separate beyond DC; chorda and M-stem absent. No upraised scale tufts; male with- out costal fold. Hindwing: Sc+R and Rs closely approximate; M, and CuA, closely approximate; tuft of hairlike scales along 1A+2A, originating near base of wing. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in female. Male gen- italia: Uncus bifurcate, each arm slender; arms either from a common stalk or wide- ly separated basally. Socius variable, usu- ally narrow, digitate, not fused to gnathos. Gnathos simple, non-dentate, arms nar- row, usually joined distally. Subscaphium and hami absent. Transtilla a simple, slen- der band, frequently reduced or membra- nous. Valva moderately short, somewhat triangular-lanceolate. Pulvinus absent. Vinculum complete. Juxta a sclerotized plate. Aedeagus long, slender, straight or curved; phallobase simple; cornuti absent. Female genitalia: Papillae anales narrow, variably notched mesally in a few species. Apophyses posteriores moderately short, anteriores reduced to a nub in most spe- cies. Sterigma a simple scobinate band. Ductus bursae moderately long, membra- nous. Corpus bursae frail, elongate, pear-
VOLUME 102, NUMBER 1
shaped or ovoid; spicules and signum lacking. Accessory bursa frequently pre- sent.
Diagnosis.—Adults of Odonthalitus are characterized by a white, cream, or pale tan forewing ground color with a simple pattern that includes a dark, trapezoidal, triangular, or semicircular patch near the middle of the costa and a dark basal patch. In a few spe- cies the pattern is variably obscured by darker overscaling, particularly in females. Males have elongate antennal cilia (ca. 3 times the width of the flagellomere) and lack the characteristic euliine male foreleg hairpencil (Brown 1990). Most species are extremely similar in facies, and examina- tion of the genitalia is required for accurate identification.
Superficially, adults of Odonthalitus are indistinguishable from those of Loboge- nesis. Characters that distinguish the two are discussed above in the redescription of Lobogenesis. In addition, a few species of Odonthalitus have a modification of the papillae anales in the female genitalia in which they are variably differentiated into a dorsal and ventral lobe separated by a weak indentation (i.e., viridimontis and paos) or distinct, deep notch (i.e., lacti- cus).
Distribution and biology.—QOdonthali- tus is known from montane oak-pine for- est habitat from Durango, Mexico, south to Costa Rica, at 1,000—2,600 m eleva- tion. Adults have been collected at lights. A single individual of O. paos was reared by J. Powell on synthetic diet from eggs deposited by a field-collected female, sug- gesting that the larvae are general-feeders. Two species are known to be sympatric in Costa Rica (at Aquiares, Cartago Prov- ince) and Mexico (at Paradero de Mi Ka, Oaxaca).
Remarks.—As with Lobogenesis, most species of Odonthalitus are rare in collec- tions; the 9 species treated below are rep- resented by 53 specimens. Five species are known only from the holotype—3 from males and 2 from females. Approximately
37
62% (n = 33) of the specimens examined represent a single species, O. viridimontis, that appears to be restricted to Costa Rica. The availability of specimens from Costa Rica can be attributed primarily to the ef- forts of parataxonomists associated with INBio, a program that is contributing sig- nificantly to our understanding of the Lep- idoptera fauna of the New World tropics; in addition, the work of Jerry Powell, Dan Janzen, and others have stimulated this ef- fort. The present treatment results in the de- scriptions of 6 new species and the reas- signment of 2 previously described species in Odonthalitus. The new combinations are proposed for 2 species previously consid- ered “‘Unplaced Euliini’ (Powell et al. £995).
Odonthalitus bisetanus J. Brown, new species (Figs. 14, 34)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale brown; roughened above, pale yellow. La- bial palpus whitish yellow mesally, brown laterally. Antennal scaling bronze-brown. Thorax: Pale tan. Forewing: Length 7.0 mm (n = 1). Upper side pale tan; basal 0.2 dark brown; dark brown, triangular patch near middle of costa, with vertex of triangle curving outward toward termen; three small irregular spots at costa between patch and apex; apex and termen with irregular brown striae. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale cream with pale gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 14 (drawn from JWB slide 650; n = 1). Uncus bifurcate from base. Gnathos and socius unmodified. Transtilla membra- nous mesally. Valva broadest at base, weak- ly attenuate distally; junction of costal base and transtilla with large hooklike process comprised of 4 strong, weakly curved spines; 2 large setae below hooklike pro- cess along inner edge of valva. Aedeagus moderately slender, nearly j-shaped, with
38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 14-16. Male genitalia of Odonthalitus; valvae spread, aedeagus removed. 14, O. bisetanus. 15, O. improprius. 16, O. conservanus.
greatly enlarged membranous cap on phal- Mpio. Yolox, Paradero de Mi Ka, 2,000 m, lobase. 5/14-XI-1980 (E. Welling, UCB). Female. Unknown. Diagnosis.—The group of long curved Type.—Holotype, ¢, Mexico, Oaxaca, spines at the base of the valva in the male
VOLUME 102, NUMBER 1
genitalia immediately distinguish Odon- thalitus bisetanus from all other species except O. improprius. These two species can be separated by the following: base of the uncus bifurcations rectangular in O. bisetanus and rounded in O. improprius; tip of the uncus bifurcations sightly flat- tened in O. bisetanus and pointed in O. improprius; patch of long spines from a lobe at the base of the transtilla in O. bis- etanus and from a narrow, attenuate flange in O. improprius; and a pair of large, strong setae just below the basal lobe of the transtilla in O. bisetanus lacking in O. improprius.
Etymology.—The specific epithet refers to the two setae at the base of the valva.
Odonthalitus improprius J. Brown, new species (Fig. 15)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, whitish. Labial palpus whitish mesally, brown laterally. Antennal scaling pale bronze-brown. Tho- rax: Pale tan. Forewing: Length 5.8 mm (n = 1). Upper side pale tan; basal 0.2 dark brown; dark brown, triangular patch near middle of costa, with vertex of triangle curving outward toward termen; two small triangular spots at costa between patch and apex; terminal region with diffuse brown striae. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale cream with pale gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 15 (drawn from JWB slide 699; n = 1). Uncus bifurcate from base. Gnathos and socius unmodified. Transtilla membra- nous mesally. Valva broadest at base, slightly attenuate distally; junction of costal base and transtilla with flat, claw-shaped flange with 4 strong spines from its venter. Aedeagus moderately slender, nearly j- shaped, with greatly enlarged membranous cap on phallobase.
Female. Unknown.
39
Type.—Holotype, 6, Mexico, Oaxaca, Mpio. Yolox, Paradero de Mi Ka, 2,000 m, 5/14-XI-1980 (E. Welling, UCB).
Diagnosis.—Odonthalitus improprius is extremely similar to O. bisetanus; the dif- ferences are detailed above in the diagnosis of the latter.
Etymology.—The specific epithet is Lat- in for inappropriate or improper.
Odonthalitus conservanus J. Brown, new species (Figs. 16, 21)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan-brown; roughened above, bronze- brown. Labial palpus pale whitish mesally, brown laterally. Antennal scaling pale bronze-brown. Thorax: Tan brown. Fore- wing: Length 5.0 mm (n = 1). Upper side whitish with irregular, fine, brown reticu- lations throughout; basal 0.2 dark brown; dark brown, triangular patch at middle of costa. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side light gray brown with slightly darker gray brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 16 (drawn from JWB slide 606; n = 1). Uncus bifur- cate from base, arising from narrow dorsal nub. Gnathos and socius unmodified. Tran- stilla a broad, weakly sclerotized plate. Val- va broadest at base, attenuate distally, with rounded excavation at costa ca. 0.33 dis- tance from base to apex; inner portion of excavated region bearing a large sclerotized hooklike process, with patch of fine setae from valva just ventrad of hook. Aedeagus moderately slender, nearly j-shaped, with greatly enlarged membranous cap on phal- lobase.
Female. FW length 6.5—6.8 mm (x = 6.6; n = 4). Essentially as described for male, except entire FW evenly overscaled with brown, nearly obscuring FW pattern. Gen- italia: As in Fig. 21 (drawn from JWB slide 607; n = 2). Papillae anales unmodi- fied, from bristly base. Papillae anales mod-
40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 17-18.
poas.
erately long, unmodified. Sterigma an irreg- ular, sclerotized band, with a bristly lobe at each postero-lateral margin; mesal portion of sterigma surrounding ostium somewhat diamond-shaped; ostium at anterior vertex of diamond. Ductus bursae membranous, undifferentiated from corpus bursae, slen- der, frail.
Types.—Holotype, 3d, Mexico, Jalisco,
Male genitalia of Odonthalitus; valvae spread, aedeagus removed. 17, O. viridimontis. 18, O.
Sierra de Manantlan, nr. Las Joyas, 1,800 m, 18-VII-1985 (J. Doyen, UCB).
Paratypes, 4 2 as follows: MEXICO: Jal- isco: same locality as holotype, 2 2, 16- VII-1985, 1 9%, 18-VII-1985 (J. Doyen, UCB, USNM); 11.6 mi [18.6 km] S of El Chante, 1 2, 3-IV-1987 (N. Bloomfield, SDNHM).
Diagnosis.—The male genitalia of Odon-
VOLUME 102, NUMBER 1
41
Figs. 19-20. Male genitalia of Odonthalitus; valvae spread. 19, O. regilla. 20, O. lacticus.
thalitus conservanus are most similar to those O. bisetanus and O. improprius. They can be distinguished from the latter two by the large hooklike process at the base of the valva, the absence of the patch of hooklike spines at the base of the transtilla, and the broad, shield-
shaped transtilla. The female genitalia are most similar to those of O. fuscomaculatus in the presence of a pair of scobinate lobes on the sterigma and the relatively unmodified (1.e., long) apophyses anteriores. Etymology.—The specific epithet refers
42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
to the “‘conserved”’ status of the type lo- cality of Las Joyas in the Sierra de Man- antlan, a global biosphere reserve in west- ern Mexico.
Odonthalitus orinoma (Walsingham 1914), new combination (Fig. 26)
Tortrix orinoma Walsingham 1914: 287. Anopina orinoma: Obraztsov 1962: 27; Powell et al. 1995: 142.
Redescription.—Male. Unknown.
Female. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, cream and brown. Labial palpus pale whitish yellow mesally, tan lat- erally. Antennal sclaing light brown. Tho- rax: Brown. Forewing: Length 7.5 mm (n = 1). Upper side pale cream with nearly uniform pale brown overscaling; basal 0.2 dark brown; triangular brown patch border- ing costa ca. 0.45—0.65 distance from base; costa between latter patch and apex with three small brown triangular spots; terminal region with irregular brown striae. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side pale gray-brown. Under side pale gray brown. Genitalia: As in Fig. 26 (drawn from BMNH slide 66598; n = 1). Papillae anales unmodified. Apophyses posteriores unmodified; apophyses anteriores greatly reduced. Sterigma a sclerotized band, with ill-defined scobinate plates on each side of ostium and a sclertoized, tubelike region mesally; ostium covered by rounded, semi- membranous plate. Ductus bursae long, slender, undifferentiated, gradually widen- ing to corpus bursae.
Type.—Holotype, 2, Mexico, Guerrero, Omilteme, 8,000’ [2,580 m], VI-1880 (H. H. Smith, BMNH).
Diagnosis.—QOdonthalitus orinoma is most similar to O. fuscomaculatus in facies and female genitalia. The two can be dis- tinguished by features of the female geni- talia—O. fuscomaculatus has conspicuous- ly longer apophyses, an undulate anterior
edge of the sterigma, and a shorter ductus bursae (see Figs. 22, 26).
Remarks.—The forewing pattern is illus- trated in Walsingham (1914) and the geni- talia in Obraztsov (1962). Obraztsov (1962) transferred “‘T.” orinoma to Anopina on the basis of the similarity in forewing pattern, indicating that “‘Until the male of orinoma is known, the systematic position of this species is somewhat doubtful.’”” The com- bination of forewing length and pattern, and the extremely short apophyses clearly in- dicate that it belongs in Odonthalitus.
Odonthalitus fuscomaculatus J. Brown, new species (Fig. 22)
Description.—Male. Unknown.
Female. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, pale tan brown. Labial palpus whitish mesally, tan laterally. Anten- nal scaling light brown. Thorax: Brown and pale tan. Forewing: Length 7.5 mm (n = 1). Upper side cream, with nearly uni- form brown overscaling and irregular brown striae; basal 0.2 dark brown; brown triangular patch bordering costa ca 0.45— 0.65 distance from base. Under side uni- form dark tan with faint indication of upper side markings. Hindwing: Upper side pale gray-brown. Under side pale gray brown. Genitalia: As in Fig. 22 (drawn from USNM slide 89449; n = 1). Papillae anales unmodified from bristly base. Apophyses posteriores and anteriores nearly equal in length. Sterigma with a pair of swollen, bristly lobes dorso-laterad of ostium; osti- um cup-shaped, sclerotized. Ductus bursae slender, undifferentiated from corpus bur- sae.
Type.—Holotype, 2, Mexico, Michoa- can, San Lorenzo, Rt. 15, km 206, 19-VII- 1966 (O. Flint & A. Ortiz, USNM).
Diagnosis.—Odonthalitus fuscomacula- tus is most similar to O. conservanus in structures of the female genitalia. Odon- thalitus fuscomaculatus can be distinguish by its larger, cup-shaped ostium, more slen-
VOLUME 102, NUMBER I
Figs. 21-23. Female genitalia of Odonthalitus. 21, O. conservanus. 22, O. fuscomaculatus. 23, O. poas.
43
44
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 24-26.
Female genitalia of Odonthalitus. 24, O. viridimontis. 25, O. lacticus. 26, O. orinoma.
VOLUME 102, NUMBER 1
der apophyses posteriores, and the more mesal position of the bristly lobes of the sterigma.
Etymology.—The specific epithet is an adjective referring to the dark overscaling of the forewing.
Odonthalitus poas J. Brown, new species (Bigs. 18523; 36)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, bronzy tan. Labial palpus pale cream mesally, bronzy brown laterally. Antennal scaling brown. Thorax: Dark brown, whitish tan at mid-dorsum. Forewing: Length 5.0—5.5 mm (x = 5.4; n = 4). Upper side white, with irregular brown transverse striae, particularly dense in distal 0.33; basal 0.2 dark brown; dark brown triangular or rhomboidal patch bor- dering costa ca. 0.45—0.60 from base, fre- quently with small, disjunct spot below lower apex of triangle; costa with a few small brown markings between patch and apex. Under side dark tan with faint indi- cation of upper side markings. Hindwing: Upper side dingy white, with pale gray- brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 18 (drawn from JWB slide 761; n = 2). Uncus moderately broad at base, bi- furcate in distal 0.75, with fine hairs. Gna- thos complex, with various rounded and an- gulate lateral processes and spines. Tran- stilla mostly membranous with setate basal portions. Valva broadest at base, attenuate apically, with excavation near mid-point of costa leaving a digitate, hairy, lobelike flange at termination of basal 0.5 of costa. Aedeagus slender, nearly evenly curved throughout; phallobase with ovoid membra- nous cap.
Female. FW length 6.0 mm (n = 2). Su- perficially as in male, except slightly larger average forewing length and lacking elon- gate antennal cilia. Genitalia: As in Fig. 23 (drawn from JWB slide 762, Costa Rica; n = 2). Papillae anales with shallow notch near middle. Apophyses anteriores absent;
45
posteriores short, slender. Sterigma a sim- ple, uniform, slightly spiculate band, with a broad subrectangular flap mesally covering ostium. Ductus bursae moderately broad, with patch of weak sclerotization, gently broadening into frail corpus bursae. Corpus bursae oblong.
Types.—Holotype, 6, Costa Rica, Car- tago Province, Rio Aquiares, nr. Santa Cruz, 9 km NW Turrialba, 1,500 m, 15-V- 1985 (J. Powell, UCB).
Paratypes, 3 6, 2 2. COSTA RICA: AI- ajuela Province: NE slope of Volcan Poas, 8 km N Vara Blanca, 1,400—1,450 m, 1 6, 6-VI-1988, 2 3, 19-VI-1988 (J. Brown & J. Powell, UCB), 1 2, 25-VII-1990, JAP 90G20 (S. Meredith & J. Powell, UCB), 1 2, 25-III-1992, JAP 92C50 (J. McCarty & J: Powell, UCB):
Diagnosis.—Odonthalitus poas is most similar to O. viridimontis. It can be distin- guished superficially from O. viridimontis by the brown, rather than yellow, transverse striae. The base of the uncus is much more narrow in O. poas and the costal lobe of the valva more pronounced (see Figs. 17—18).
Biology.—A confined female (JAP 90G20) produced two eggs. The larvae were reared on artificial diet; one success- fully pupated but did not eclose (J. Powell, personal communication). In general, spe- cies that are capable of completing devel- opment on artificial diet are general-feed- ers—the only clue to potential larval host plants of the group.
Etymology.—The specific epithet, a noun in opposition, refers to the collecting locality of all but the holotype on the slope of Mount Poas, Costa Rica.
Odonthalitus viridimontis J. Brown, new species (Bigs: 177.245 35)
Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, brown and pale yellow; roughened above, bronzy yellow. Labial palpus pale yellow mesally, reddish brown laterally. Antennal scaling brown. Thorax: Dark brown. Fore-
46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 27-37. Adults of Odonthalitus and Lobogenesis. 27, L. contrasta (3). 28, L. antiqua (d). 29, L. peruviana (2). 30, L. magdalenana (@, Bolivia). 31, L. varnicosa (2). 32, L. penai (3). 33, L. larana (2). 34, O. bisetanus (d). 35, O. viridimontis (3). 36, O. poas (3). 37, L. magdalenana (3, Venezuela).
VOLUME 102, NUMBER 1
wing: Length 5.0-5.5 mm (* = 5.2; n = 10). Upper side grayish white with faint yellow transverse striae; basal 0.5 dark brown; dark brown semicircular or trian- gular patch bordering costa ca. 0.45—0.60 from base, frequently with small hooklike marking from lower apex of triangle; ter- minal area with brown overscaling. Under side dark tan with faint indication of upper side markings. Hindwing: Upper side light gray-brown with slightly darker gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 17 (drawn from JWB slide 609, Costa Rica; n = 5). Uncus extremely broad in basal 0.5, arms widely divergent distally. Soci re- duced to tiny pads. Gnathos arms complex, with a slender, curved basal appendage. Valva broad at base, attenuate apically, with rounded excavation along costa in distal 0.5. Aedeagus extremely long, slender, bent at ca. 0.33 distance from base to tip.
Female. FW length 5.0—6.5 mm (x = 5.6; n = 8). Superficially as in male, except slightly larger average forewing length, fre- quently with more dense overscaling, and lacking elongate antennal cilia. Genitalia: As in Fig. 24 (drawn from JWB slide 793, Costa Rica; n = 5). Papillae anales flattened laterally, weakly notched near middle. Apophyses anteriores absent. Sterigma a simple band, dilated at middle to accom- modate ostium. Antrum sclerotized. Ductus bursae frail, slender, undifferentiated from corpus bursae.
Types.—Holotype, ¢, Costa Rica, Pun- tarenas Province, Monteverde, 1500 m, 11- VI-1988 (J. Brown & J. Powell, UCB).
Paratypes; 47 6, 15"2..\COSTA RICA: Cartago Province: Rio Aquiares, nr. Santa Cruz, 9 air km NW Turmialba, 1,500 m, 2 2, 16-V-1985 (J. Powell & P. Opler, UCB), 1 2, 8-VI-1988, 2 5d, 1 2, 10-VI-1988 J. Brown & J. Powell, UCB); Monumento Nacional Guayabo, A. C. Amistad, 1,100 m, 1 3, VI-1994 (G. Fonseca, INBio). Pun- tarenas Province: Monteverde, 2 6, 1 &, 18/19-V-1985 (J. Doyen, UCB), 2 2, 11- VI-1988, 1 3, 13-VI-1988 (J. Brown & J.
47
Powell, UCB), 1 6, 1 2, 22/24-VII-1990 (S. Meredith & J. Powell; UCB), 1 ¢6, 1 &, 29/31-III-1992, UV & MV lights (S. McCarty & J. Powell, UCB); Estacion La Casona, Res. Biol. Monteverde, 1,520 m, 1; fe, XI-1990 (N. Obando), 1 6,1 2, X-1991 (J. Saborio, INBio), 1 6, IX-1992, 1 3, V- 19933) ded 5° Te S-VIlIE1993," 1, = 1994 (all N. Obando, INBio, USNM); San Luis, Monteverde, A. C. Arenal, 1,000—1,350 m, 2 2, VI-1994 (Z. Fuentes, INBio); Buen Amigo, San Luis, Monteverde, 1,000— 15350m, 16) VI-1994) "1°92 XM-1996.(Z: Fuentes, INBio); 2 km E of Monteverde, 2 3d, 12-VI-1988, blacklight (J. Brown & J. Powell, UCB); Estac. Biol. Las Cruces, Rio Jaba, 6 km SE San Vito, secondary forest, 150mm ,e21ss.,. 220/211-1993 y G- “Powell; UCB).
Diagnosis.—The male genitalia of Odon- thalitus viridimontis are distinguished easi- ly from those of its congeners by the ex- tremely broad base of the uncus, the shape of the valvae, and the extremely long, slen- der aedeagus (Fig. 17). Although the shape of the aedeagus is similar to that of O. re- gilla (Fig. 18), the two are not similar in other features of the genitalia.
Etymology.—The specific epithet is a noun referring to the type locality of Mon- teverde, Costa Rica.
Odonthalitus regilla (Walsingham 1914), new combination (Fig. 19)
Tortrix regilla Walsingham 1914: 289. ‘‘Eulia”’ regilla: Powell et al. 1995: 146.
Redescription.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan-brown; roughened above, bronze- brown. Labial palpus pale whitish yellow mesally, bronze-brown laterally. Antennal scaling pale bronze-brown. Thorax: Tan- brown. Forewing: Length 7.2 mm (n = 1). Upper side pale red-brown; basal 0.2 dark brown, basal patch wider at costa than at dorsum; dark brown, irregularly triangular patch at middle of costa; a brown spot near
48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
apex of DC at vertex of costal triangular patch; apex and termen with brown band concolorous with costal patch. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side light gray brown. Under side light gray brown. Genitalia: As in Fig. 19 (photo- graph of BMNH slide 5794; n = 1). Uncus with a posterior nub, behind (anterad) which are a pair of extremely slender, elon- gate arms ( = bifurcations of uncus) from dorsum of tegumen. Gnathos swollen dis- tally. Socii short, broad. Transtilla membra- nous, ill-defined. Valva slender, nearly uni- form in width, with weak notch along ven- ter ca. two-thirds distance from base to apex; apex with strong, free, curved hook from weakly rounded base. Aedeagus long, straight, extremely slender.
Female. Unknown.
Type.—Holotype, ¢, Guatemala, Retal- huleu, Las Mercedes, 3,000’ [970 m], X/XI- 1880, (G. C. Champion, BMNH).
Diagnosis.—The male genitalia of Odon- thalitus regilla are moderately divergent from other species in the genus: the nearly parallel-sided valva and the clawlike pro- cess from the apex are unlike any other spe- cies. The long, slender aedeagus is reminis- cent of that of O. viridimontis, and the basal origin of the bifurcation of the uncus is sim- ilar to O. lacticus.
Remarks.—The adult is illustrated in Walsingham (1914). The forewing pattern, elongate antennal cilia, bifurcate uncus, and long, slender aedeagus indicate that it be- longs in Odonthalitus.
Odonthalitus lacticus Razowski 1992 (Figs. 20, 25)
Odonthalitus lacticus Razowski 1992: 208; Powell et al. 1995: 145.
Redescription.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, cream. Labial palpus whitish mesally, tan laterally. Antennal scaling light brown. Thorax: Brown. Fore- wing: Length 6.5 mm (n = 1). Upper side
cream; basal 0.2 dark brown; triangular brown patch bordering costa ca. 0.45—0.65 distance from base; costa between latter patch and apex with three small brown tri- angular spots; terminal region with irregular brown band. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale gray brown. Under side pale gray brown. Genitalia: As in Fig. 20 (drawn from UCB slide 3392; n = 1). Tegumen short, rounded. Uncus com- posed of two distinct, widely separated rods. Socius short. Gnathos long, slender, with accessory process near middle, arms separate distally. Valva relatively short, at- tenuate at apex, venter concave in basal 0.5; costa undifferentiated; sacculus weakly sclerotized. Transtilla a broad rectangular plate. Aedeagus large, curved, broadened apically. Female. FW length 6.0—7.0 mm (x = 6.3; n = 3). Essentially as described for male, except ground color in some specimens evenly dusted with pale brown, reducing definition of forewing markings. Genitalia: As in Fig. 25 (drawn from UCB slide 3312 and JWB slide 603; n = 2). Papillae anales with pronounced notch near middle, result- ing in bilobed appearance. Apophyses an- teriores represented by a short thorn; apophyses posteriores comparatively long. Sterigma a simple, broad, sclerotized band, with a pair of lateral incisions. Ductus bur- sae broad, moderately short, with fine lines of sclerotization. Corpus bursae short, rounded. Accessory bursa from narrow ductus arising near middle of ductus bursae. Types.—Holotype, 2, Mexico, Durango, 4 miles [6.4 km] west of El Palmito, 20- VII-1964, flight trap (J. Powell, UCB). Paratype, | d, same data as holotype. Additional specimens examined.—MEX- ICO: Sinaloa: 15 mi [24 km] west of El Palmito, black and white lights, 1 2, 18- VII-1964 (J. Chemsak & J. Powell, UCB); 8 mi [12.8 km] west of El Palmito, 6,000’, Io, 122X21975:9 at Hight) GycPowell; 3: Chemsak, T. Eichlin & T. Friedlander, UCB).
VOLUME 102, NUMBER 1
Diagnosis.—The male genitalia of Odon- thalitus lacticus are fairly divergent from other species in the genus. They can be dis- tinguished by the widely concave distal 0.5 of the venter of the valva, the widely sep- arate bases of the bifurcate uncus, and the relatively robust aedeagus (Fig. 20). The notch in the papillae anales of the female is similar to that of O. viridimontis and O. paos, but is much deeper.
ACKNOWLEDGMENTS
I thank the following for allowing me to examine material in their care: Jerry Powell (UCB), Eugenie Phillips (INBio), Kevin Tuck (BMNH), and David Faulkner (SDNHM). I thank the following for re- viewing the manuscript: Jerry Powell, Uni- versity of California, Berkeley, California; Richard Brown, Mississippi State Univer- sity, Mississippi State, Mississippi; William Miller, University of Minnesota, St. Paul, Minnesota; and E Christian Thompson and David Smith, Systematic Entomology Lab- oratory, USDA, National Museum of Nat- ural History, Washington, D.C. I thank J. Powell for providing me with the opportu- nity to conduct field work in Costa Rica and for comments on rearing attempts. The fig- ures were skillfully rendered by Susan Escher (Figs. 2—5, 9-18, 20, 24), Front Royal, Virginia, and Linda Lawrence (Figs. 6-8, 21-23), USDA, Systematic Entomol- ogy Laboratory, Washington, D.C.
49
LITERATURE CITED
Brown, J. W. 1990. Taxonomic distribution and phy- logenetic significance of the male foreleg hairpen- cil in the Tortricinae (Lepidoptera: Tortricidae). Entomological News 101: 109-116.
Brown, J. W. and J. A. Powell. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). University of California Pub- lications in Entomology 111. 87 pp.
2000. Systematics of Anopina Obraztsov (Lepidoptera: Tortricidae). University of Califor- nia Publications in Entomology. In press.
Horak, M. 1984. Assessment of taxonomically signif- icant structures in Tortricinae (Lep., Tortricidae). Mitteilugen der Schweizerischem Entomologisch- en Gesellschaft 57: 3—64.
Lipscomb, D. 1994. Cladistic Analysis Using Hen- nig86, version 1.5. George Washington Universi- ty, 122 pp.
Obraztsov, N. 1962. Anopina, a new genus in the Cne- phasiini from the New World (Lepidoptera: Tor- tricidae). American Museum Novitates 2082: 1— 39.
Powell, J. A., J. Razowski, and J. W. Brown. 1995. Tortricidae: Tortricinae, pp. 138—150. /n Heppner, J. B., ed., Atlas of Neotropical Lepidoptera, Checklist: Part 2, Hyblaeoidea-Pyraloidea-Tortri- coidea. Association for Tropical Lepidoptera. Sci- entific Publishers, Gainesville, FL.
Razowski, J. 1984. A list of the known Neotropical Tortricini moths (Lepidoptera, Tortricidae) with description of the first South American species. Revista Brasiliana Entomologica 28: 203-206.
. 1992 [1990]. On some peculiar Neotropical
tortricine genera (Lepidoptera: Tortricidae). SHI-
LAP Revista de Lepidopterologia 18: 209-215.
. 1993. Revision of Apotoforma Busck, 1934 (Lepidoptera: Tortricidae), with descriptions of four other Tortricini. Acta Zoologicae Cracovien- sia 36: 183-197.
Walsingham, Lord T. de Grey. 1914. Biologia Centrali- Americana. Insecta, Lepidoptera Heterocera, Vol. 6. British Museum (Natural History), London. 482 PP.
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 50-57
MILESIINE FLOWER FLIES (DIPTERA: SYRPHIDAE) IN A CENTRAL APPALACHIAN BROADLEAF FOREST: ABUNDANCES, FLIGHT PERIODS, AND DIFLUBENZURON
Mary R. CICERO AND EDWARD M. BARROWS
Laboratory of Entomology and Animal Behavior, Reiss Building Suite 406, Department of Biology, Georgetown University, Box 571229, Washington, DC 20057-1229, U.S.A (e-mail: barrowse @ gunet.georgetown.edu)
Abstract.—We used 20 Malaise traps to collect adult milesiine syrphids in four water- sheds in a central Appalachian broadleaf forest during early April through late September in 1991 through 1993. The traps captured 31 species in 12 genera. As a group, these flies flew throughout our annual sampling periods, their sample sizes varied with species and year, and their abundances peaked in late May or mid-June, depending on the year. For all species whose sample size was 210 individuals, male-female sex ratios varied from 0.1 through 3.0. We performed a split-plot analysis of flies captured from mid-May through early August, a period in each of the 3 years when traps captured at least one fly during a sampling period. This analysis did not find an effect of trap site or diflubenzuron on fly population sizes based on Malaise sampling, because these effects did not exist or
our sample size was too small to detect them.
Key Words:
The broad aims of our study are to learn more about the biology of syrphid flies in the tribe Milesiini and determine whether aerially-applied diflubenzuron affects pop- ulation sizes of these beneficial insects in a temporate broadleaf forest. Diflubenzuron (Dimilin®, Uniroyal) is an insect growth regulator used to control forest and other pest arthropods. We report which milesiine species are present in this forest and their flight periods, seasonal abundances, and sex ratios, based on sampling with Malaise traps. Further, we used a split-plot analysis to test the hypotheses that diflubenzuron and trap site affect the number of individ- uals captured.
Milesiine syrphids are beneficial insects in forests and other habitats, where they may be Batesian mimics of some stinging hymenopterans, honeydew consumers, food
Appalachian forest, diflubenzuron, Malaise traps, phenology, Syrphidae
for other organisms, nectarivores, polleni- vores, pollinators, and scavengers (Gilbert 1981, 1986; Maier 1982; Vockeroth and Thompson 1987; Waldbauer 1988; Owen 1991). We studied this syrphid taxon be- cause its members have diverse habits and are taxonomically well known and easily identified. Syrphids and bees are the major pollinators in many habitats.
Federal and state agencies have treated thousands of hectares of forests with diflu- benzuron to control gypsy moths, Lyman- tria dispar (L.), in the eastern United States (USDA Forest Service 1994). This pesti- cide also reduces the numbers of nontarget, beneficial forest insects including honey bees (Egger 1977), macrolepidopterans and nonlepidopteran mandibulate herbivores (Martinat et al. 1988); macrolepidopterans (Sample et al. 1993; Butler ect ale 1997);
VOLUME 102, NUMBER 1
stoneflies (Griffith et al. 1996), and yellow- jackets (Barrows et al. 1994). However, there are no published reports regarding the possible effects of aerially-applied difluben- zuron on forest syrphids.
MATERIALS AND METHODS
We studied syrphids in the 1,902-ha Fer- now Experimental Forest in Tucker County, West Virginia, over a 3-year period (1991— 1993). Detailed accounts of our materials and methods are published in Barrows et al. (1994) and Barrows (1995).
The Fernow Experimental Forest is from 533 to 1,112 m in elevation and has slopes up to 60%, a mean of 145 frost-free days, a mean annual precipitation of 147 cm, and mean temperature of 9°C (Anonymous 1987). Acer rubrum L., Betula lenta L., Fa- gus grandifolia Ehrhart, Liriodendron tuli- pifera L., Prunus serotina Ehrhart, and Quercus spp. are common trees in this for- est:
We used watersheds 1, 4, 7, and 13 as study areas. Watersheds 1, 7, and 13 are conterminous, and watershed 4 is about 200 m from watershed 7 (Adams et al. 1994, map). Watersheds | and 13, the difluben- zuron-treated plots (test plots) totaled 42 ha, and watersheds 4 and 7, the control plots totaled 63 ha. On 16 May 1992, the USDA Forest Service aerially applied diflubenzu- ron (Dimilin® 4L) with a helicopter at 35.1 g (AI) ha"! to watersheds 1 and 13.
During April through September, all 3 study years were drier than a 4l-year av- erage of 76 cm for the Fernow Experimen- tal Forest, 1991 and 1993 were warmer than average (15.9°C), and 1992 was cooler than average. During these months, mean tem- perature was 17.6, 14.9, and 17.1°C, and the mean precipitation was 56.9, 59.9 and 64.3 cm, in 1991, 1992, and 1993, respectively.
We captured flies from 29 April through 27 September 1991-1993 using five Townes-style Malaise traps (Townes 1972; manufactured by Golden Owl Publishers, Lexington Park, MD) placed 20—35 m apart on the ground in a transect in each of the
51
watersheds as illustrated in Barrows et al. (1994). Each transect ran across a valley and approximately perpendicularly to a sec- ond-order stream. From the vantage point of looking upstream, two traps were on the left side and two were on the right side of a central trap. Valley left sides had north- erly exposures, and right sides had south- erly exposures. The central trap of each transect was from 0 to 6 m from the edge of a stream. We placed all traps on the for- est floor, not on logging roads or in other artificial openings.
A trap was made of 1-mm-mesh, nylon gauze, a supporting frame, and a collecting head. A 0.95-liter jar, which was part of the head and contained 95% ethanol, collected the flies as well as thousands of species of other arthropods. Each trap was 1.2 m wide, 1.7 m long, 1.0 m high at its back, and 2.0 m high at its front (head end). Trap roofs were made of white gauze and baffles and sides were made of black gauze. We emp- tied our 20 traps every 10 d on the same Julian day in April through September in 3 yr, totaling 895 samples.
Falling tree limbs, mice, bears, and pos- sibly other animals occasionally damaged our traps; we repaired, or replaced, them as soon as we found damage. Five samples were incomplete because of trap damage and could not be used in our statistical an- alyses. We identified syrphids to genus us- ing keys in Vockeroth and Thompson (1987) and to species with the reference collection at the National Museum of Nat- ural History, Smithsonian Institution, and deposited a voucher collection of these flies in that Museum. For species with large enough sample sizes for analysis, we used x’ goodness of fit tests to determine wheth- er they had sex ratios that significantly dif- fered from male-female sex ratios of 1:1, 1: 2, and 2:1. To look for a possible difference in the number of syrphid individuals among years, we used analysis of variance. Finally, we used a split-plot analysis to look for possible diflubenzuron and trap-site effects on the number of captured flies (PROC
oy PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
_—
JE
O
= total = 661 1992 total = 349 total = 732
O
WY)
LLJ
<
Tp Figure 1
LL
=
=
LL
O
O
= 110 140 170 200 230 260 110 140 170 200 230 260 110 140 170 200 230 260 Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep
JULIAN DAY Fig. 1. Numbers of milesiines captured in each 10-day sample in the four watersheds, Fernow Experimental
Forest, 1991-1993.
MIXED, SAS Institute 1992). The treat- ments were pesticide (with the two levels application and nonapplication) and trap site (five levels or locations). We calculated the grand mean of the trap means of flies for eight sampling periods (mid-May through early August) per watershed per year. This was a period when most milesi- ines flew and there was at least 1 fly per sampling period in each of the 3 sampling years. In 1991, the preapplication year, mi- lesiine populations were higher in the ap- plication watersheds than in the nonappli- cation watersheds, possibly because the for- mer were better syrphid habitats. To adjust for this difference, we analyzed the differ- ences between the 1991 and 1992 and be- tween the 1991 and 1993 grand means, test- ing the null hypothesis that the difference between the application-watershed means of fly number equals the difference between the nonapplication-watershed means of fly number. To look for a possible trap-site ef- fect, we tested the null hypothesis that the mean numbers of flies from the five trap sites are equal.
RESULTS AND DISCUSSION
Species, sex ratios, flight periods, and abundances.—Traps collected a total of 1,742 milesiines belonging to 31 species in
12 genera (Fig. 1, Table 1—2). The Fernow Experimental Forest is within the previous- ly known ranges of all of these species; however, this is the first published species list of Milesiini from a central Appalachian broadleaf forest.
By the ends of 1991, 1992, and 1993, our traps obtained 84%, 94%, and 100% of the 31 milesiine species, respectively. Some species were rare in samples, and not all species were in samples every year. The six most common species comprised 76% of the 1,742 collected specimens.
We were unable to run our traps for more than 3 years, but Owen’s (1991) study sug- gests that our traps may have obtained the majority of the milesiine species in our study site during our three-season study pe- riod. She caught 43,749 individuals in 91 species and 41 genera, by using one Townes-style Malaise trap (Townes 1972) in her British garden for 15 successive years. The trap obtained the vast majority of its total number of syrphid species by the end of its fifth year of operation, rarely caught some species, caught markedly varying numbers of some species from year to year, and obtained a previously uncaught species during most years from the 5th through 15th years of operation. Bankows- ka (1980) found 25 species in seven mile-
VOLUME 102, NUMBER 1
Table 1. Abundances of milesiine syrphids and percentages of males collected in Malaise traps from late April through late September, in all four water- sheds in the Fernow Experimental Forest, West Vir- ginia, 1991-1993.
Number
Species Captured % Males* Blera analis (Macquart) 30 47° B. badia (Walker) 263 65° B. nigra Williston | 100 B. pictipes (Bigot) IS 53° B. umbratilis (Williston) l 100 Brachypalpus oarus Walker 102 47° Chalcosyrphus inarmatus (Hunter) 20 55? C. libo (Walker) 17 298 C. nemorum (F.) 6 7, C. plesius Curran 2 100 C. vecors Osten Sachen 3 33 Criorhina nigriventris Walton 9 22 C. verbosa Walker 36 Le Lejota aerea (Loew) 9 56 Prerallastes thoracicus Loew 123 Sk: Somula decora Macquart 25 24 Specomyia vittata Wiedemann 30 23 Spilomyia alcimus (Walker) 4 100 S. fusca Loew 9 22 Temnostoma alternans Loew 111 She T. balyras Walker 573 522 T. barberi Curran 21 10 T. trifasciatum Robertson | 27 T. venustum Williston 8 25 T. vespiforme (L.) 167 69: Teuchocnemis lituratus (Loew) 66 50° Xylota 78-1 Thompson 4 25 X. 78-3 Thompson 17 294 X. bicolor Loew DZD, 36! X. nebulosa Johnson | 100 X. quadrimaculata Loew 36 17
Total 1,742
“The proportion of males for all species combined is 47%.
> This sex ratio is not different than 1.0 or 50% males (df = 1, P < 0.05, x? goodness of fit test).
© This sex ratio is not different than 2.0 or 66% males (df = 1, P < 0.05, x* goodness of fit test).
4 This sex ratio is not different than 0.5 or 33% males (df = 1, P < 0.05, x? goodness of fit test).
sline genera in Polish broadleaf forests, compared to the 31 species in 12 genera reported in our study (Table 1).
In our study, male-female milesiine sex ratios varied from 0.1 through 3.0. Four species had male-female sex ratios that were not different from 1:2; seven species,
Bye) 1:1; and two species 2:1 (Table 1). In all species in all 3 years, 47% of the syrphids were males which is not different than a 1: 1 sex ratio (50% males) (df = 1, x? = 2.431, P = 0.119). Deviations from a 1:1 sex ratio in certain species may be due to factors including having an adult sex ratio that is actually different from 1:1 and one sex’s being more likely to be trapped than the other.
Milesiines flew throughout most of our monitoring period; however, the traps col- lected the vast majority (95%) of them from early May through late July (Fig. 1, Table 2). Traps acquired individual milesiine spe- cies for an mean of 59.4 + 5.6 SEM days (range 10—130 days, 31 species), obtaining all species except for Chalcosyrphus ne- morum (FE.) and Pterallastes thoracicus Loew, for 100, or fewer, days. This mean and ranges are approximations because our sampling intervals were 10 days.
Of the species with sample sizes of 10 or more individuals, three species peaked in abundance in early May; three in mid-May; one in late May; one in early June; eight in mid-June; and one in late June (Table 2). The main annual peaks were in late May 1991 and mid-June in 1992 and 1993 (Fig. 1). The troughs between the two large peaks in 1991 and 1993 coincide with rainy periods. Visnyovszky (1988) also found rain-related drops in Malaise-trap sample sizes in these flies in Hungarian apple or- chards.
Among flies in the traps, Temnostoma balyras Walker was the most common, fol- lowed by Blera badia (Walker), T. vespi-
forme L., Pterallastes thoracicus, and T. al-
ternans Loew (Table 1). The only published study of North American Milesiine flight periods is based on the faunas of Piatt and Mason Counties in central Illinois, both ap- proximately 40° north, and Emmet County in northern lower Michigan, approximately 45° north (Waldbauer 1988). This study in- volved high-fidelity mimics of stinging hy- menopterans which were sampled by hand netting. Our study site (approximately 39°
54
Mablen2:
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Abundances of milesiine syrphids and percentages of males collected in Malaise traps from late
April through late September, in all four watersheds, Fernow Experimental Forest, West Virginia, 1991-1993
Month and Sampling Period?
April May June July August September Species il La ee es eee ReGen ee oe SOL aoe Tinion sian Bren
Blera analis 0) 0) | 3 4 20 1 0) 1 Oo 0 O= © @ © @ B. badia 2 2 5 35 78 93 Sy) IS} l 2 0 O OY © @ O B. nigra 0) l 0) 0) 0) 0) 0) oO © O © © O © O B. pictipes O 0) 0) Th 1 4 1 oO © © © OO © O B. umbratilis 10) 0) 0) 0) 0) i 0) oO © © OO @ © O © O Brachypalpus oarus 12 59 17 11 0) 0) 1 oO OF @ O O- OD © O O Chalcosyrphus inarmatus 0) l 8 =) 4 1 oO © O O OO O O C C. libo l 4 4 4 0) 2 l l Oo OO © OF O&O © © C. nemorum O 0) l 1 0) ] | l Yo O © OL. O-O © C. plesius 0) 0) 0) | 0) 1 0) oO O © O © O @ O O C. vecors O 0) 0) 0) 0) 2 | oO Oo O O GO O O © © Criorhina nigriventris D ] 4 2 0) 0) 0) oO © © @ OO OY WO O C. verbosa 6 16 3) 0) 0) 0) 0) oO OO © GO © © O © Lejota aerea 0) 6 3} 0) O 0) 10) Oo Oo © & © O OF OD O Pterallastes thoracicus® 6) 0) l 9 NS) 41 sy alse ay a Del aril 0) Somula decora‘ 0 0) 0) 6 9 5 4 1 Oo OY O OG © © @ Specomyia vittata‘ 6 0) 7 16 2 1 0) tO @ © O O @ © @ Spilomyia alcimus* 0) 0) 0) 0) O 0) | oy Oo 2 OO OO O O @ S. fusca‘ 0) 0) 0) 0) 0) 0) 0) ye Or Be Oe O Temnostoma alternans® 0) 0) 6 29 13 60 16 3} 3 Oe th Oe OY. @ Temnostoma balyras* 2 17 28) 186" 136) 154" 7 437 192 lL © OO O © © © T. barberis 0 0) 1 5 6 6 2 YY 0 CO O @ © @ © O T. trifasciatum’ 0) 0) 0) 0) 5 4 | l YO OO OO O GO © @© O T. venustums 0) 0) l l 1 | 1 | x YO © © © O O @ T. vespiforme® 0) 0) 9 36 19 82 14 7 © © OO O OD @ ©@ Teuchocnemis lituratus 0) 3 25 21 6 6 3 OO 2 0) © O Oo O © @ Xylota bicolor 0) 0) 0) 0) 3 6 9 | BO O 1 @ ©. @ X. nebulosa 0) O 6) ] 0) 0 0 OF Os 0) OO -O) Or OF @ X. quadrimaculata 0) 0) 2 4 8 9 6 4 l fi Oa © © C X. n. sp. | 0) O 0) 0) 0) 1 2 0) l oO OY O O © NG id Coy 2 0) 4 3 4 2 3 0) 0) l OO 0 O O @ © Totals 31 NS 129 340) 31255507 SiG9 a7 OM SO A Gyr
“Sample peaks in species with =10 specimens are in bold face.
> | = April: Julian days 110-120; May: 2 =
230; 13° =
120-130; 3 = 130-140; 4 = 140-150; June: 5 = 150-160; 6 = 160-170; 7 = 170-180; July: 8 = 180-190; 9 = 190-200; 10 = 200-210; August: 11 = 230-240; September: 14 = 240-250; 15 = 250-260; 16 = 260-270.
210-220; 12 = 220-
© High-fidelity mimics of stinging hymenopterans (Waldbauer 1988).
north) shares three milesiine species with each of the Illinois sites and seven species with the Michigan site. Approximate mile- siine flight periods in West Virginia are similar to those in the Illinois sites. Their flight periods in West Virginia were earlier and longer than those in Michigan.
The mean numbers of milesiines cap- tured in nonapplication watersheds per trap per sampling period was 2.7 + 0.7, 1.7 +
0.7, and 3.4 + 0.7 SEM in 1991, 1992, and 1993, respectively, showing a difference in number of individuals among years (AN- OVA, df = 10, F = 5.94; P = 0.0199). Malaise traps are an excellent means for monitoring population sizes of insect taxa that are readily captured by these traps. However, they probably never collect target taxa in the exact proportions that they occur in their communities (Barrows 1986; Dar-
VOLUME 102, NUMBER 1
Nn Nn
150 e test plots
O control plots 120
Figure 2
110 140 170 200 230 260 110 140 170 200 230 260 110 140 170 200 230 260 Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep
JULIAN DAY
Fig. 2. Numbers of milesiines captured in each 10-day sampling period in application and nonapplication watersheds, Fernow Experimental Forest. The arrow indicates the time of pesticide spraying on application plots.
NO. OF MILESIINES CAUGHT
ling and Packer 1988; Archer 1990). Visual Trap-site and diflubenzuron.—Our split- counts of some syrphid species suggested plot analysis detected no diflubenzuron, or that a Malaise trap did not collect them in _ trap-site, effect on milesiines (P > 0.05, for the proportions that they occurred near the both factors) (Fig. 2, Table 3). However, trap (Owen 1991). this finding does not indicate that aerially-
Table 3. Split-plot analysis of possible diflubenzuron and trap-site effects on numbers of milesiine syrphids.
Treatments Year Mean + | SEM? Pe
No pesticide (1 and 13) 199] 42 = 0.90 not applicable No pesticide (4 and 7) 2.7 220.53
Pesticide (1 and 13) 1991 minus 1992 2.2 + 0.64a 0.2016 No pesticide (4 and 7) 0.9 + 0.64a
Pesticide (1 and 13) 1991 minus 1993 —0.3 + 0.98a 0.4642 No pesticide (4 and 7) 0.7 + 0.98a
Trap | 199] 2 6ie= (O:8i1a 02297 Trap 2 19+ 0.8la
Trap 3 2.0 = '0:8ia
Trap 4 3.1 == 10:8ila
Trap 5 3.4 + 0.8la
Trap | 1992 1.8 = 0.55a 0.3438 Trap 2 1.3, '0.55a
Trap 3 IpOy==O55a
Trap 4 IESe=="0i55a
Trap 5 2.9 = 0:55a
Trap | 1993 4.6. cb lda 0.8302 Trap 2 Sos) Labla
Trap 3 302 Telia
Trap 4 B32 ee ela
Trap 5 39° Talla
« This mean is the average number of syphids per 10-day sample per year per watershed. Within a year, means followed by the same letter are not different from each other (P = 0.05, least-squares-means test). These means are based on two control, or test, plots per year, or four traps per subplot per year.
® The null hypothesis is that the means within groups (pesticide treatment or trap-site treatment) are equal.
56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
applied diflubenzuron would never decrease milesiine populations in forests. We might not have detected a possible diflubenzuron effect because our sample size was small, milesiines might have entered our test wa- tersheds from untreated ones and masked a pesticide effect, or both. No apparent effect of trap site on milesiine sample size sug- gests that these flies were relatively evenly distributed in our sample area.
In conclusion, we report that 31 milesiine species occurred in a central Appalachian broadleaf forest and they were of markedly different abundances and sex ratios in Mal- aise-trap samples. Although, our study found no evidence that diflubenzuron af- fected milesiine population sizes, a more comprehensive study might find such an ef- fect. This pesticide remains on _ treated leaves for many months on plants and in leaf litter (Wimmer 1995), and larval mi- lesiines are scavengers which could be poi- soned by them. If forest managers continue to use diflubenzuron to control gypsy moths and other forest pests, more studies of this pesticide and syrphids are warranted.
ACKNOWLEDGMENTS
We are pleased to thank many people who helped with our study: Mary Beth Ad- ams, Pamela J. Edwards, and Fredericka Wood (USDA Forest Service, Parsons, WV); Zorayda C. Alviar, Holi D. Ieng, Jo- seph H. Neale, and James V. Tedesco (Georgetown University); Joseph E. Bar- rows, Julie N. Barrows, Melissa L. Bar- rows, and Robert E. Harrigan (Bethesda, MD); Donald A. M. Mackay (Pleasantville, NY), David R. Smith (USDA, Washington, DC); Amy H. Onken and Richard C. Rear- don (USDA Forest Service, Morgantown, WV); Mary J. Camp and William E. Potts (Statistical Consulting Services, USDA, Beltsville, MD); Janet D. Rupp (Golden Owl Publishers, Lexington Park, MD); and Deborah J. Smith (Falls Church, VA). Lin- da Butler (West Virginia University, Mor- gantown, WV), Chris T. Maier (Connecticut Agriculture Experiment Station, New Ha-
ven, CT), and an anonymous reviewer made insightful comments that improved this paper. EK Christian Thompson (System- atic Entomology Laboratory, USDA, Wash- ington, DC) kindly gave us invaluable help with syrphid identification and other mat- ters. This study was funded by Georgetown University and the Appalachian Integrated Pest Management Gypsy Moth Project of the Northeastern Area, State and Private Forestry, USDA Forest Service.
LITERATURE CITED
Adams, M. B., J. N. Kochenderfer, E Wood, T. R. An- gradi, and P. Edwards. 1994. Forty years of hy- drometeorological data from the Fernow Experi- mental Forest, West Virginia. USDA Forest Ser- vice, Northeastern Forest Experiment Station. General Technical Report NE-184. 24 pp.
Anonymous. 1987. Forest Research: Fernow Experi- mental Forest. U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station NE-INF-75-87. 12 pp.
Archer, M. E. 1990. The solitary aculeate wasps and bees (Hymenoptera: Aculeata) of an English sub- urban garden. Entomologist’s Gazettee 41: 129— 142.
Bankowska, R. 1980. Fly communities of the family Syrphidae in natural and anthropogenic habitats of Poland. Memorabilia Zoologica 33: 3—93.
Barrows, E. M. 1986. A hornet, paper wasps, and yel- lowjackets (Hymenoptera: Vespidae) in suburban habitats of the Washington, D.C., Area. Proceed- ings of the Entomological Society of Washington 88: 237-243.
. 1995. Chapter 8. Pollinating insects—Native species, pp. 66—76. Jn Reardon, R. C., ed., 1995, Effects of Diflubenzuron on Non-target Organ- isms in Broadleaf Forested Watersheds in the Northeast. USDA Forest Service FHM-NC-0595. 174 pp.
Barrows, E. M., S. S. Wolf, & D. M. Lynch. 1994. Diflubenzuron effect on yellowjacket (Hymenop- tera: Vespidae) worker numbers in a central Ap- palachian broadleaf forest. Journal of Economic Entomology 87: 1488-1493.
Butler, L., G. A. Chrislip, V. A. Kondo, and E. C. Townsend. 1997. Effect of diflubenzuron on non- target canopy arthropods in closed, deciduous wa- tersheds in a central Appalachian forest. Journal of Economic Entomology 90: 784—794.
Darling, D. C. and L. Packer. 1988. Effectiveness of Malaise traps in collecting Hymenoptera: The in- fluence of trap design, mesh size, and location. Canadian Entomologist 120: 787-796.
Egger, A. 1977. Die Nebenwirkungen von DIMILIN
VOLUME 102, NUMBER 1
auf Bienen—eine Untersuchung anlasslich einer Flugzeugapplikation im Forst. Centralblatt fiir das Gesamte Forstwesen 94: 65-72.
Gilbert, E S. 1981. Foraging ecology of hoverflies: Morphology of the mouthparts in relation to feed- ing on nectar and pollen in some common urban species. Ecological Entomology 6: 245-262.
. 1986. Hoverflies. Naturalists’ Handbooks No. 5. Cambridge University Press, Cambridge, UK, 66 pp.
Griffith, M. B., E. M. Barrows, and S. A. Perry. 1996. Effects of aerial application of diflubenzuron on
the emergence and flight of adult aquatic insects (Plecoptera, Trichoptera). Journal of Economic Entomology 89: 442—446.
Martinat, P. J., C. C. Coffman, K. Dodge, R. J. Cooper, and R. C. Whitmore. 1988. Effect of diflubenzu- ron on the canopy arthropod community in a cen- tral Appalachian forest. Journal of Economic En- tomology 81: 261—267.
Owen, J. 1991. The Ecology of a Garden: The First Fifteen Years. Cambridge University Press, Cam- bridge, England, 403 pp.
Sample, B. E., L. Butler, and R. C. Whitmore. 1993. Effects of an operational application of Dimilin®
ai
on non-target insects. Canadian Entomologist 125: 173-179.
SAS Institute. 1992. SAS/STAT software: Changes and enhancements, Release 6.07. Technical Report P-229. SAS Institute, Cary, NC, 620 pp.
Townes, H. 1972. A light-weight Malaise trap. Ento- mological News 83: 239-247.
United States Department of Agriculture Forest Ser- vice. 1994. Gypsy moth news 36.
Visnyovsky, E. 1988. Phenological study of the syr- phid flies caught by a Malaise trap, pp. 123-127. In Niemcezyk, E. and A. FE G. Dixon, eds., Ecology and Effectiveness of Aphidophaga. SPB Academ- ic Publishing, The Hague, The Netherlands.
Vockeroth, J. R. and F C. Thompson. 1987. Syrphidae, pp. 713-743. In McAlpine, J. F, ed., Manual of Nearctic Diptera, Vol. 2. Biosystematics Research Centre, Ottawa, Ontario.
Waldbauer, G. P. 1988. Asynchrony between Batesian mimics and their models. The American Naturalist 131: S$103-S121.
Wimmer, M. J. 1995. Chapter 16. Terrestrial Environ- ment, pages 23-30. Jn Reardon, R. C., ed., 1995, Effects of Diflubenzuron on Non-target Organ- isms in Broadleaf Forested Watersheds in the Northeast. USDA Forest Service FHM-NC-0595. 174 pp.
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 58-61
A NEW SPECIES OF CIRROSPILUS WESTWOOD (HYMENOPTERA: EULOPHIDAE) FROM THE SOUTHWESTERN UNITED STATES AND MEXICO
MICHAEL W. GATES
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
(e-mail: gates @citrus.ucr.edu)
Abstract.—Cirrospilus coachellae, n. sp. (Hymenoptera: Eulophidae), is described from the southwestern United States and Mexico. Comparisons are made with described species of Cirrospilus and differences between Cirrospilus and the closely related Za- grammosoma are discussed. Cirrospilus coachellae is the primary parasitoid providing control of the citrus peelminer, Marmara sp. (Lepidoptera: Gracillariidae), a cyclical pest
of citrus in the southwestern United States.
Key Words:
Eulophidae in the genus Cirrospilus Westwood are cosmopolitan in distribution and parasitize insects with cryptic feeding habits. Cirrospilus parasitizes leaf-miners, leaf-rollers and galling insects in the orders Lepidoptera, Coleoptera, Diptera and Hy- menoptera (Schauff et al. 1997, Boutek 1988). Species of Cirrospilus typically are larval ectoparasitoids, but also have been documented to utilize primary ichneumo- noid parasitoids attacking concealed insects and to directly attack several life stages of the primary host (Boucek and Askew 1968). Currently, there are over 300 species of Cirrospilus described worldwide, of which the majority occur in the Holarctic region.
Cirrospilus belongs to the subfamily Eu- lophinae, as characterized by the submar- ginal vein smoothly joining the parastigma, a postmarginal vein often longer than the stigmal vein, and a minimum of three setae on the dorsal surface of the submarginal vein (Schauff et al. 1997). Cirrospilus is characterized by having the funicle two- segmented, notauli complete, and the post-
Hymenoptera, Eulophidae, Cirrospilus, taxonomy, Chalcidoidea
marginal vein shorter than or equal in length to the stigmal vein (Schauff et al. 1997). Gordh (1978) recognized another closely related eulophine genus, Zagram- mosoma Ashmead, as distinct from Cirros- pilus on the basis of two characters: head with the vertex distinctly vaulted so as to extend above the compound eye in Zagram- mosoma, and the propodeum with a well developed median carina in Cirrospilus but only a weak carina in Zagrammosoma. LaSalle (1989) further separated these gen- era by noting in Zagrammosoma the notau- lus curves to meet the axilla anteriorly, thus not approaching the scutoscutellar suture, while in Cirrospilus the notaulus extends straight to the scutoscutellar suture. LaSalle discounted the propodeal carina character because of its inconsistency across taxa. Yefremova (1996) cited additional char- acters which serve to distinguish Zagram- mosoma from Cirrospilus. In Zagrammo- soma, the axilla is situated anterior to the posterior margin of the mesoscutum and typically is elongated. Cirrospilus has the axilla located posterior to the posterior mar-
VOLUME 102, NUMBER 1
gin of the mesoscutum, and the axilla is more triangular than the axilla of Zagram- mosoma. Based upon material examined by the author, Yefremova’s character appears sufficient to separate the two genera. How- ever, the question remains as to whether recognizing Zagrammosoma as distinct will eventually render Cirrospilus paraphyletic.
The species described here belongs to Cirrospilus, however it splits the diagnostic characters pertaining to the vaulted vertex and notaulus configuration. Cirrospilus coachellae has the vertex distinctly vaulted and the notaulus straight, while other mem- bers of Cirrospilus do not possess the vault- ed vertex (as seen in Zagrammosoma). Cir- rospilus coachellae has the axillar character used by Yefremova (1996) to differentiate Cirrospilus and Zagrammosoma.
Specimens used in this study were bor- rowed from or are deposited in the institu- tions referred to in the type material section. The acronyms are as follows:
BMNH: The Natural History Museum, London, U.K.
UCR. University of California, River- side, Entomology Research Mu- seum, Riverside, California, U.S.A.
USNM: National Museum of Natural
History, Smithsonian Institution, Washington, D.C., U.S.A.
Cirrospilus coachellae Gates, new species (Figs. 1—4.)
Type Material.—Holotype °@, card mounted, ““USA: CA: River. Co., Coachella Valley, NW Salton Sea, 5.VII.96, M. Guil- len, collector/Ex Marmara sp. peelmines on grapefruit,’ deposited in USNM.
Paratypes: same data as holotype, card mounted, (Total paratypes: 23 2, 12 6; de- posited in UCR, 8 2, 5 6; deposited in USNM, 8 2, 4 6; deposited in BMNH, 7 2, 3 3). Additional pointed material: *“Mex. Baja Cal. Sur, Las Barracas, 14-IV
59
1985/Coll. P. Debach, Pantrap’’, (4 2, de- posited in BMNH).
Diagnosis.—Cirrospilus coachellae can be distinguished from other members of the genus by the unique possession of a vaulted vertex, which is not seen in other species, and the notaulus not curving to intercept the axilla anteriorly.
Female.—Length 2.16—2.75 mm. Head yellow except for following, which are brown: three sets of stripes originating from occipital foramen, first set diverging to in- tercept lateral ocelli, second proceeding lat- erally to contact posterodorsal margin of eye, and third set proceeding ventrad and curving anterodorsally to meet ventral eye margin (Fig. 1); lower face with line ex- tending from between toruli to clypeal mar- gin; two stripes originating lateral to toruli and intercepting ventral half of malar sul- cus; frons with two tapering stripes located between anterodorsal eye margin and ex- tending half distance to lateral scrobal mar- gin; two spots between scrobal depression and medial eye margin extending to contact scrobe; scrobal depression; scape and ped- icel dorsally (Fig. 2). Mesosoma and me- tasoma yellow except for following which are brown: median stripe on pronotum, me- soscutum and scutellum; two pairs of stripes dorsolaterally and laterally on pron- otum, these fused anteriorly and diverging posteriorly; lateral lobe with stripe medial- ly; axilla with antero-medial spot; broad submedial stripes on scutellum; propodeum medially from one-third posterior margin to about four-fifths anterior margin, except transverse, teardrop-shaped yellow spots lateral of dorsellum; gastral tergite 1 (Gt1) with triangular spot surrounding petiolar in- sertion, distal point of spot approaching posterior edge of tergite; Gt2—4 with lateral tergal spots separate from medial spots on Gt2 and increasing fusion caudally to form a solid stripe on Gt4; Gt3—7 with broad, longitudinal stripe connecting transverse bands at intersegmental junctions (Fig. 3). Wings hyaline and venation yellowish ex-
60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Scutoscutellar suture
Figs. 1—4. tasoma, dorsal view. 4, Wing, dorsal view. Figs. 1, 4, scale bar = 1 mm. Fig. 2, scale bar = 50pm.
cept brown at junction of stigmal and post- marginal veins (Fig. 4).
Head: Quadrate, 1.1X higher than broad; distance between posterior ocelli 1.5-2.0X greater than distance between posterior ocellus and eye margin. Face fine- ly reticulate and moderately setiferous; oc- cipital carina absent; eyes reddish, eye height nearly 2 greater than malar space; antenna 8-segmented; scape and pedicel with fine longitudinal carinae medially;
Cirrospilus coachellae. 1, Head, anterior view. 2, Antenna, lateral view. 3, Mesosoma plus me-
pedicel longer than broad; one anellus; two funicular segments setose and lacking sur- face sculpture; Fl 1.0—1.1X as long as broad; F2 0.83—1.0X as long as broad; cla- va 1.7X as long as broad, tapering distally.
Mesosoma: Dorsum finely reticulate; mesoscutal midlobe about 1.2 as long as broad; scutellum quadrate to 0.9X as long as broad; propodeum reticulate with smooth median area; callus with few erect setae; coxae with fine longitudinal striations and
VOLUME 102, NUMBER 1
with a few setae; hind femur 3.0—5.0X as long as broad. Forewings reaching beyond tip of gaster, with submarginal vein 0.8— 0.9X length of marginal vein; stigmal vein 2.4—4.0X longer than postmarginal vein; all veins setose and stigmal vein more darkly infuscate than remaining veination; wing evenly setose distad of parastigma, except for small asetose area at apex of stigmal vein.
Metasoma: Gaster 1.1 longer than me- sosoma; petiole short; tips of ovipositor sheath visible dorsally.
Male.—Length 1.67—2.32 mm. Smaller than female, body striping often paler and less extensive than in female.
Variation.—Aside from differences in specimen size, the primary variation is in the coloration and striping patterns. Longi- tudinal striping on the mesosoma ranges from contacting the anterior and posterior margins of a tergite (e.g., pronotum, scu- tellum), to being represented only as a cen- tral dash or absent altogether (e.g., parap- sidal stripe). The gastral coloration varies primarily in thickness and extent of stripes.
Biology.—This species has been recov- ered in large numbers from the citrus peel- miner, a new species of Marmara (Lepi- doptera: Gracillariidae) (Guillen, Davis and Heraty, in prep.), in the Coachella Valley of Riverside County, CA. It is a gregarious ec- toparasitoid of the peelminer larva and up to 12 C. coachellae have been documented from a single individual (Guillen, personal communication). This same miner attacks the stems and leaves of Nerium oleander L. (Apocynaceae) and Gossypium sp. (Sola- naceae) in Arizona (M. Guillen, personal communication), which also support C. coachellae. Cirrospilus coachellae has also been recovered from another similar undes- cribed species of Marmara mining stems of tree tobacco, Nicotiana glauca Grah. (So- lanaceae).
61
Distribution.—This insect has been re- covered from Riverside County, California, Yuma County, Arizona, and Baja Califor- nia, Mexico.
Etymology.—This species is named for the Coachella Valley in the Colorado Desert of southern California where the type series was recovered.
ACKNOWLEDGMENTS
This work was partially supported by a UC-IPM grant to John Heraty (UCR). Ad- ditional support is provided by a grant from the Citrus Research Board (CRB) to John Heraty to survey native leafminer parasit- oids. Thanks also to John LaSalle (BMNH) for providing additional specimens of this species collected in Baja California Sur and to Arthur Fong of the California State Park System for permission to collect on state park system lands. Thanks to John Heraty who provided critical comments upon ear- lier drafts of this manuscript.
LITERATURE CITED
Bouéek, Z. 1988. Australasian Chalcidoidea (Hyme- noptera): A biosystematic revision of fourteen families with a reclassification of species. CAB International, Wallingford, England, 832 pp.
Bouéek, Z. and R. Askew. 1968. Palaearctic Eulophi- dae (excl. Tetrastichinae). (Hym. Chalcidoidea). Index of Entomophagous Insects. Le Francois, Paris, 260 pp.
Gordh, G. 1978. Taxonomic notes on Zagrammosoma, a key to the Nearctic species and descriptions of new species from California (Hymenoptera: Eu- lophidae). Proceeding of the Entomological So- ciety Washington 80: 344-359.
LaSalle, J. 1989. Notes on the genus Zagrammosoma (Hymenoptera: Eulophidae) with description of a new species. Proceeding of the Entomological So- ciety Washington 91: 230-236.
Schauff, M., J. N. LaSalle, and L. Coote. 1997. Chap- ter 10. Eulophidae, pp. 327—429. In Annotated keys to the Genera of Nearctic Chalcidoidea (Hy- menoptera), NRC Research Press, Ottawa, Ontar- 10, Canada.
Yefremova, Z. 1996. Notes on some Palaearctic and Afrotropical species of the genus Zagrammosoma (Hymenoptera: Eulophidae). Entomological Re- view 75(5): 163-171.
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 62-68
NEW RECORDS OF TWO XYLEBORUS (COLEOPTERA: SCOLYTIDAE) IN NORTH AMERICA
NATALIA J. VANDENBERG, ROBERT J. RABAGLIA, AND DONALD E. BRIGHT
(NJV) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, MRC 168, Wash- ington, DC 20560-0168, U.S.A. (e-mail: nvandenb @sel.barc.usda.gov); (RJR) Maryland Department of Agriculture, 50 Harry S. Truman Parkway, Annapolis, Maryland 21401, U.S.A. (e-mail: rabaglrj@mda.state.md.us); (DEB) Research Branch, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada K1A OC6 (e-mail: brightd@em.agr.ca)
Abstract.—New American locality records are given for two exotic ambrosia beetles (Coleoptera: Scolytidae) in the genus Xyleborus. Xyleborus pfeili (Ratzeburg), a widely distributed Old World species, is reported for the first time in North America, from three counties in Maryland. Xyleborus californicus Wood, of northern Palearctic origin, but previously established in the western United States, is reported for the first time from Maryland, Delaware, South Carolina and Arkansas. Diagnoses and descriptions are given for the two species, along with scanning electron micrographs of key characters. Modi- fications are made to a previous key to include these new additions to the eastern North
American ambrosia beetle fauna.
Key Words:
The introduction through commerce of exotic bark and ambrosia beetles (Coleop- tera: Scolytidae) poses a threat to our native forests and urban plantings. The adult bee- tles damage trees by tunneling and feeding in the cambium region just beneath the bark’s surface (true bark beetles), or by dril- ling into the sapwood and feeding on intro- duced fungal symbiotes (ambrosia beetles). Although scolytid beetles may promote the general health of a stand by culling overly mature or damaged trees (Atkinson et al. 1990), the effect of foreign species, released from controlling factors of their native en- vironment, can be unpredictable or even devastating (U.S. Congress, Office of Tech- nology Assessment 1993).
Within the past decade, scolytid intro- ductions and interceptions have increased concern over the effects of exotic species in
Xyleborus, Scolytidae, eastern North America, introductions
North America. Recently, several econom- ically important bark beetles have become established on this continent (e.g., Tomicus piniperda (L.) (Haack and Kucera 1993) and Hylastes opacus Erichson (Hoebeke 1994, Rabaglia and Cavey 1994)), or been collected at or near United States and Ca- nadian ports-of-entry (e.g, Ips typographus (L.), Cavey and Passoa 1993). Eight species of exotic ambrosia beetles have been estab- lished in eastern North America within this century (Wood 1977, 1982, Atkinson et al. 1990, Hoebeke 1991), and some of these have become significant pests. We report the occurrence of two additional exotic am- brosia beetles new to eastern North Amer- ica: Xyleborus pfeili (Ratzeburg) and X. cal- ifornicus Wood.
Xyleborus pfeili was first detected in North America in a mixed sample of Xy-
VOLUME 102, NUMBER 1
leborine ambrosia beetles submitted to one of us (NJV) at the USDA Systematic En- tomology Laboratory, Washington, D.C. The specimens were included along with a sample of frass collected from branch/ stump of paw paw, Asimina triloba (An- nonaceae), at the Wye Research and Edu- cation Center, Carmichael, Maryland, and were reported to have emerged from the host in June and July 1992. The lot con- sisted of 2 specimens of Xyleborinus saxe- seni (Ratzeburg), 3 specimens of Xyleborus affinis Eichhoff, and 5 specimens of another species which could not be identified using the keys to exotic and native North Amer- ican fauna in Wood (1982) or Atkinson et al. (1990). The 5 unknowns were recog- nized (DEB) as a new record for the Old World species, X. pfeili. Additional speci- mens were found (RJR) during bark beetle trapping surveys in Maryland from 1994— 1997, where yet another exotic species, Xy- leborus californicus, was detected for the first time. During manuscript preparation, the latter species was also found in Arkan- sas and South Carolina. Although it was first described from a population in Cali- fornia (Wood 1982), it is now known to be a northern Palearctic species (Siberia (S.L.Wood, pers. comm.), China, new in- terception in British Columbia (DEB)). De- scriptions, diagnoses, revised keys and scanning electron micrographs are provided for each of the newly reported exotic spe- cies to aid in identification. Material from the trapping survey has been deposited in the National Museum of Natural History, Washington, D.C.
Xyleborus pfeili (Ratzeburg) (Figs. 2, 4—6)
Distribution.—This species is native to Europe (Austria, Belgium, Bulgaria, former Czechoslovakia, France, Germany, Greece, Hungary, Italy, Poland, Romania, Spain, Switzerland, and former western USSR), Asia (China, Japan, Korea and Turkey) and Africa (Algeria and Morocco), and has been
63
introduced into New Zealand (Wood and Bright 1992).
Diagnosis.—Among_ eastern North American Xyleborus, specimens of X. pfeili with well-developed declivital tubercles are most likely to be confused with X. celsus Eichhoff and X. ferrugineus (FE). Xyleborus pfeili (Fig. 2) differs from X. celsus (Fig. 1) in its more gradual declivity with greater consistancy in the size of interstrial tuber- cles, and the straighter declivital striae 1. It differs from X. ferrugineus by the presence of tubercles on declivital interstriae 1. Spec- imens of X. pfeili with less developed de- clivital tubercles are most likely to be con- fused with X. volvulus (FE). They can be dis- tinguished by the larger size (X. volvulus females are generally less than 2.8 mm), the weakly sulcate elytra with distinctly raised interstriae 1 (Fig. 2), and the more abruptly formed declivity (Figs. 4—5).
Description.—Female: Length 3—3.6 mm, cylindrical, reddish brown, with legs paler yellowish.
Frons minutely reticulate, weakly shin- ing, coarsely shallowly punctate; longitu- dinal carina weak to obsolete.
Pronotum 1.2 times as long as wide, sides roughly parallel, summit near middle; anterior margin of pronotum arcuate, un- armed; anterior and posterolateral areas weakly reticulate and faintly shining, with moderate to weakly developed asperities and moderately long setae; median area posterior to summit devoid of asperities, more polished, with fewer setae and sparse scattered punctures.
Elytra about 1.8 times as long as wide; strial punctures on disc moderately large, each with very short recumbent seta, sepa- rated within row by approximate diameter of a puncture; interstrial discal punctures minute, more widely and less regularly spaced, each with stouter, longer, erect to semirecumbant seta; interstriae | with gran- ules sometimes developed in posterior half before declivity. Elytral declivity steep, oblique, linear in profile, occupying ap- proximately posterior 30% of elytra, weak-
64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-3.
ly sulcate, highly polished except for duller patches in depressed areas; rows of strial punctures less regular than on disc, deviat- ing around larger tubercles; 2 or 3 larger conical tubercles often present on interstriae 1 and 3; scattered smaller tubercles or gran- ules often occurring on some or all inter- striae; interstriae | slightly elevated.
Male: Not seen. Males of Xyleborus spp. are “‘exceedingly rare and flightless”’ (Wood 1982). Described and illustrated by Balachowsky (1949). Apparently similar to female, except for smaller size (2.1—2.6 mm long) and pronotal shape which is ogival in outline and concave in anterior %.
Variation.—Specimens vary in develop- ment and position of declivital armature, al- though specimens from the introduced pop- ulation (Figs. 2, 4—5) thus far show less variation than some of the Old World spec- imens examined. Specimens with reduced declivital tubercles show less variation in the relative size of these structures, and some specimens in a series from Brout-Ver- net, France, entirely lack declivital tuber- cles except at base.
Specimens examined (all 2, introduced population).—UNITED STATES: MARY- LAND: Cecil Co.: Elk Neck State Forest, 4 June 1997, ex Chalcoprax-baited funnel trap, R.J. Rabaglia. Kent Co.: Coleman, 4 June 1997, ex Chalcoprax-baited funnel trap, R.J. Rabaglia. Queen Anne’s Co.: Car- michael, Wye Research and Education Cen- ter, June-July 1992, R.N.Peterson, ex branch/stump of paw paw, Asimina triloba; Wye, 5 May 1995, ex ethanol-baited funnel trap, R.J. Rabaglia; Wye, 26 September
Posterolateral views of elytral declivity. 1, Xyleborus celsus. 2, X. pfeili. 3, X. californicus.
1996, ex ethanol-baited funnel trap, R.J. Rabaglia.
Xyleborus californicus Wood (Fig. 3, 7—9)
Distribution.—According to Wood (pers. comm.) this species belongs to a complex indigenous to Siberia and neighboring parts of northern Asia. Recently a specimen from China was intercepted in Vancouver, B.C. The species was originally described from a population established in California and Oregon, but its exotic status was never se- riously in dispute (Wood 1982). The sudden occurrence of X. californicus in the Mid- Atlantic region and southeastern United States further supports the exotic origin of this species. Each year its numbers and dis- tribution have increased.
Diagnosis.—Wood (1982) states that this species might be confused with pubescens Zimmermann. It can be distinguished by the more abundant pubescence, the smaller size and the lighter coloration. We found X. cal- ifornicus to be most similar to X. pellicu- losus Eichhoff except for the much smaller size and paler color. Both of the latter spe- cies have the same structure of the antennal club and similar body proportions, puncta- tion and vestiture. Xyleborus pelliculosus was not included in Wood’s key because its presence in the United States was reported subsequently (Atkinson et al. 1990).
Description.—The following description is from Wood (1982) with the addition of a phrase (in bold face) to describe the anten- nal club:
VOLUME 102, NUMBER 1
Figs. 4-9.
Female: Length 2.0—2.2 mm, 2.9 times as long as wide; yellowish brown.
Frons rather strongly convex; surface strongly reticulate, a few small granules from epistoma to upper level of eyes. Ves- titure of fine, sparse hair. Antennal club (Fig. 9) flattened, subcircular, posterior face solid or with a subapical suture and rows of setae visible on apical third.
Pronotum 1.2 times as long as wide; sides almost straight and parallel on basal two-thirds, rather broadly rounded in front; anterior margin unarmed; summit in front of middle; anterior slope steep, rather coarsely asperate; posterior areas strongly
4—6, Xyleborus pfeili. 4, Dorsal view. 5, Lateral view. 6, Detail of head showing left antenna. 7— 9, X. californicus. 7, Dorsal view. 8, Lateral view. 9, Detail of head showing left antenna.
reticulate, punctures small, shallow, rather close. Vestiture of fine, short rather abun- dant hair.
Elytra 1.7 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; disc occupying basal three-fourths; striae not impressed, punctures small, shallow, distinct, in rows, spaced by diameter of a puncture; inter- striae three to four times as wide as striae, almost smooth, shining, punctures fine, in indefinite rows in some specimens, distinct- ly confused on basal half in others. Decliv- ity steep, convex, general contours as in pu-
66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
bescens; strial punctures large, shallow, dis- tinct, their interior surfaces reticulate-gran- ulate; interstriae only slightly wider than striae, their punctures mostly replaced by minute granules on all interstriae, a few granules on 1, 3 and lateral areas; postero- lateral margin rounded, with an indefinite row of scattered granules. Vestiture of rath- er abundant, short, fine hair, distinctly lon- ger on margins of declivity.
Specimens examined (all 2 ).—UNITED STATES: DELAWARE: New Castle Co.: Wilmington, 21 August 1997, Delaware Dept of Agric. Coll. Sussex Co.: Redden State Forest HQ Tract, 18 May 1997, Mi- chael A. Valenti; Redden State Forest Ap- penzellar Tract, 23 May 1997, Michael A. Valenti. MARYLAND: Anne Arundel Co.: Odenton, 18 April 1994, R. J. Rabaglia; Somerset Co.: Wellington, 27 April 1994, R. J. Rabaglia. Calvert Co.: Lusby, 11 April 1995, ex Ipslure-baited funnel trap, R. J. Rabaglia. Caroline Co.: Idylwilde Wildlife Management Area, 12 May 1994, R. J. Ra- baglia. Cecil Co.: Elk Neck State Forest, 9 April 1997, ex Lineatin-baited funnel trap, R. J. Rabaglia. Charles Co.: Indian Head, 11 April 1995, ex Ipslure baited-funnel trap, R. J. Rabaglia. Harford Co.: Upper Cross- roads, 3 May 1994, R. J. Rabaglia. Kent Co.: Sandy Bottom, 9 April 1997, ex etha- nol-baited funnel trap, R. J. Rabaglia. Queen Anne’s Co.: Matapeake, 8 April 1997, ex Chalcoprax-baited funnel trap, R. J. Rabaglia; Wye Mills, 23 April 1997, ex ethanol-baited funnel trap, R. J. Rabaglia; Romancoke, 5 May 1997, ex Ipslure-baited funnel trap, R. J. Rabaglia. St.Mary’s Co.: Mechanicsville, 11 April 1995, ex Ipslure baited-funnel trap, R. J. Rabaglia. Talbot Co.: Longwoods, 12 May 1997, ex ethanol- baited funnel trap, R. J. Rabaglia. Worces- ter Co.: Pocomoke State Forest, 27 April 1994, R. J. Rabaglia. SOUTH CAROLINA: Aiken Co.: Savannah River Site, 7 July 1993, ex turpentine-baited bumper trap, R. D. Klaper Coll. Stephen Co.: Chattahoo- chee National Forest, 10 July 1996, ex lob- lolly-baited tent trap, C. A. H. Flechtmann
Coll. ARKANSAS: Pulaski Co.: Little Rock, 21 March 1997, ex oak stump, B. Baldwin Coll.
Discussion.—Atkinson et al. (1990) pub- lished a key to females of the eastern North American Xyleborus. The key is modified below in order to accomodate the newly re- corded species. Figure captions in italics re- fer to figures in the original publication. Note that X. validus Eichhoff in Atkinson et al (1990) has been removed from Xyle- borus; it is now placed in Euwallacea (see Wood & Bright 1992) and is not included in the modified key.
1. Anterior margin of pronotum usually armed by several coarse serrations (Fig. 8), however serrations absent or reduced in atratus (Fig. 2); body stout, less than 2.2 times as long as wide; mature color black 6423.5 fes.ci 3. gttracon os Shen ceeeietencne 2 — Anterior margin of pronotum unarmed by large serrations (Fig. 10); body slender, greater than 2.5 times as long as wide; mature color usually yellowish or reddish brown Posterolateral costa on declivity armed by 3-5 distinct tubercles. North-central Unit- ed States and Canada, south to Virginia. 2A8=3 SSMS ee eee ease: obesus LeConte — Posterolateral costa on declivity of uni- form height, may appear slightly undulat- ing, but without denticles (Fig. 9) ..... 3 Anterior margin of pronotum armed by numerous small subequal serrations (Fig. 8); declivital striae impressed, declivital interstriae less than 2 times as wide as striae (Figs: 45 9) ss nn oe) ao eee 4 — Anterior margin of pronotum armed by 2— 6 serrations, median pair conspicuously larger than others; declivital interstriae at least twice as wide as striae. North-central United States and Canada. 2.3—2.6 mm REL, She eMC: Eee ae es ee sayi (Hopkins) Anterior margin of pronotum armed by 6— 8 subequal serrations (Fig. 8); declivity flattened, interstrial setae subequal in length to width of interstriae (Fig. 9). North-central United States and Canada, Pacific Northwest of United States and Canaday228=3 25mm ween dispar (EF) — Anterior margin of pronotum with weakly developed serrations (Fig. 2); declivital interstriae 2 impressed, declivity slightly bisulcate; interstrial setae twice as long as width of interstriae (Fig. 4). Asian exotic,
2(1)
4(3)
VOLUME 102, NUMBER 1
(1)
6(5)
7(6)
8(7)
9(6)
10(9)
11(10)
Tennessee, Georgia, Maryland, West Vir- einias.3 Osmium: Ws ts ae atratus Eichhoff Declivity strongly concave with obtusely elevated margins on posterolateral areas (Fig. 14); sutural interstriae of declivity armed by 2—4 stout tubercles. Southern Texas to Central America. 3.8—4.2 mm
Ato tee 35 ioral Ce A oA eR horridus Eichhoff Declivity convex, flattened, or somewhat impressed near midline, but not concave
Denticles on some interstriae much larger than on others (Fig. 1) (Figs. /6-/7) ... = 7 Denticles on all interstriae (where present) more uniform in size (Fig. 2, 3) (Figs. 18— 22 ON Aaa apap soisy sci aus Seekers, Suc iar(acew Nee ANS 9 Declivity steep, flat, surface dull; stria | on declivity strongly curved away from midline, with 2 large, pointed, widely spaced tubercles almost on striae; smaller granules on all interstriae only at base or lateral areas of declivity, not on face, forming a circumdeclivital ring (Fig. 1) (Fig. 16). Eastern North America. 3.6—4.5 1100-0 Wigs oe RPC OREO cho tone celsus Eichhoff Declivity less steep, slightly impressed along midline, surface shining; interstriae 1 and 2 armed only at base by small tu- bercles; interstria 3 with 3 widely spaced denticles, the middle one conspicuously larserthantothers, (Hig. D7) csc nt = 8 Anterior portion of pronotum of female impressed, weakly sulcate. Southeastern United States 2:05275 mm .........
Anterior portion of pronotum of female convex, normal (Figs. /O—//), impressed and sulcate in males (Figs. //, /3). East- ern North America, Neotropics. 2.0—2.3
MIM ee acectay ar ret tee ae SE ne ferrugineus (F.) Surface of declivity opaque, dull (Figs. Ole PAD) PAID) Vecccamstt oe ace ae eR me ae 10
Surface of declivity shining (Figs. 2—3) (Figs. 18, 22) except for duller patches in depressed! areas ).quj eles = faa bel) 12 Declivity broadly sloping, occupying pos- terior 30—40% of elytra (Fig. 23), decliv- ity slightly tapered posteriorly; tubercles of interstriae | and 3 small but conspicu- ous (Fig. 20). Eastern North America, Neotropics. 2.0-2.7 mm ... affinis Eichhoff Declivity steep, occupying posterior 15% of elytra, apex blunt, not tapered; tuber- cles of interstriae 1 and 3 very small (Fig. aN ese tse 5, cy tee Stee cee Fane el Pater aries ee 11 Anterior portion of pronotum of female impressed, weakly sulcate. Eastern North
67
America} 2:3=2-4) mm? 0.3. fed. 28!
Fi bre event te ono planicollis Zimmermann — Anterior portion of pronotum of female convex, normal (impressed and sulcate in males). Eastern North America. 2.3—2.7
T2010 See tear gihetns orients xylographus (Say) 12(9) Interstrial setae on elytra and declivity placed in a single median row (Figs. 2, 4)
ape ac eae eh ak cide arabe chs fact kaa SP 13 — Interstrial setae on elytra and declivity in 2 or 3 randomly placed rows (Figs. 3, 7) Ne aes. Can Cea oe een care Ree ae 16 13(12) Elytral declivity convex, posterolateral ar- eas not subacutely elevated (Figs. 22, 24) Leche Seri bole Ae a Saran ete ae se ces 14
— Elytral declivity flattened, sloping, pos- terolateral areas subacutely margined (Eaig)RCHUSS FILS os whe Scapa Gs eee aes 15
14(13) Discal interstriae 2 times as wide as striae; some declivital tubercles with height and basal width greater than the diameter of strial punctures. Western North America to Honduras, New York to Georgia. 2.2—
Pe i237 1) 01 aga eo ae intrusus Blandford
— Discal interstriae less than 1.5 times as wide as striae; some declivital tubercles with height and basal width less than the diameter of strial punctures (Fig. 22). Eastern North America. 2.3—2.7 mm SASL, yd Oe eee eNO pubescens Zimmermann
15(13) Length 2.1—2.8 mm; eye with upper part above emargination wider than antennal club; southern Florida, Texas, widespread in Neotropical region .. volvulus (Fabricius)
— Length 3.0—3.6 mm; eye with upper part above emargination narrower than anten- nal club; introduced, Maryland ..... veh tee Biseo Sia ao aie oh pfeili (Ratzeburg)
16(12) Length 2.0-2.2 mm; color yellowish- brown; introduced, California, Oregon, Arkansas, Delaware, Maryland, and South G@arolina ou. a as ae californicus Wood
— Length 3.2 mm; color dark brown; intro- duced, Maryland, Pennsylvania .....
Ar Maio 5 bo IAB Sees pelliculosus Eichhotf
ACKNOWLEDGMENTS
We thank S.L. Wood, Brigham Young University, Provo, Utah for confirming the identification of X. californicus and provid- ing information about its possible origin; Lisa Roberts, Systematic Entomology Lab- oratory, for producing the scanning electron microscope images; C. L. Staines, Edge- water, Maryland, J. W. Brown and D. M. Anderson, Systematic Entomology Labo-
68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ratory, ARS, USDA, Washington, D.C. for their review of earlier versions of this man- uscript.
LITERATURE CITED
Atkinson, T. H., R. J. Rabaglia, and D. E. Bright. 1990. Newly detected exotic species of Xyleborus (Co- leoptera:Scolytidae) with a revised key to species in eastern North America. Canadian Entomologist 122: 92-104.
Balachowsky, A. S. 1949. Coléopteres: Scolytides. Faune de France 50. 320 pp.
Cavey, J. and S. Passoa. 1993. Possible new introduc- tion European spruce bark beetle. USDA Forest Service, Northeastern Area, Pest Alert. NA-TP- 18-93
Haack, R. and D. Kucera. 1993. Common pine shoot beetle. USDA Forest Service, Northeastern Area, Pest Alert. NA-TP-05-93.
Hoebeke, E. R. 1991. An asian ambrosia beetle, Am- brosiodmus lewisi, new to North America (Cole- optera: Scolytidae). Proceedings of the Entomo- logical Society of Washington. 93(2): 420—424.
. 1994. New records of immigrant bark beetles (Coleoptera: Scolytidae) in New York: attraction of conifer-feeding species to ethanol-baited trap logs. Entomological News 105(5): 267-276.
Rabaglia, R. J. and J. EF Cavey 1994. Note on the dis- tribution of the immigrant bark beetle, Hylastes opacus Erichson, in North America (Coleoptera: Scolytidae). Entomological News 105(5): 277— 279
U.S. Congress, Office of Technology Assessment. 1993. Harmful non-indigenous species in the United States, OTA-F-565 Washington, DC, U.S. Government Printing Office.
Wood, S. L. 1977. Introduced and exported American Scolytidae (Coleoptera). Great Basin Naturalist 37: 67-74.
. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs 6. 1356 pp.
Wood, S. L. and D. E. Bright. 1992. A catalog of Scol- ytidae and Platypodidae (Coleoptera), Part 2: Tax- onomic Index, Vol. A. Great Basin Naturalist
Memotrs 13: 1-833.
PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 69-81
LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF NEASPILOTA SIGNIFERA (COQUILLETT) (DIPTERA: TEPHRITIDAE) ON HEMIZONIA PUNGENS (HOOKER AND ARNOTT) TORREY AND A. GRAY (ASTERACEAE) IN SOUTHERN CALIFORNIA
RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: rgoeden @ucracl.ucr.edu)
Abstract.—Neaspilota signifera (Coquillett) is a bivoltine, monophagous fruit fly (Diptera: Tephritidae) apparently developing solely in the flower heads of Hemizonia pungens (Hooker and Arnott) Torrey and Gray in southern California. The egg, first-, second-, and third-instar larvae, and puparium are described and figured. The mouth- hooks of the first and second instars are bidentate, but tridentate in the third instar. The pair of flattened integumental petals are fused laterally with the well developed, stomal sense organs in the first instar; whereas, the integumental petals are papilliform and progressively more numerous in the second and third instars. The dorsal sense organ is well defined in all three instars. The six oral ridges in the second and third instars are dentate ventrally. The life cycle is of the aggregative type. Most eggs are laid singly between the chaff and ovules of preblossom flower heads and perpendicular to the receptacle. First instars feed on ovules, as do the second instars, which also feed on soft achenes in open flower heads, like all third instars. A third of the third instars examined also pitted the receptacles and apparently supplemented their diet with sap. Pupariation occurs inside the mature flower heads, but no protective cell is formed, as with congeners that overwinter as prepuparia. Instead, F, adults emerge from their cells in early summer (June) and either produce a partial second generation in late- blooming flower heads or pass the summer, fall, and winter in riparian habitats as long- lived adults. Surviving, overwintered adults aggregate the next year in early spring (March—April) on preblossom host plants to mate and subsequently oviposit. A Pter- omalus sp. (Hymenoptera: Pteromalidae) was reared from puparia as a solitary, larval- pupal endoparasitoid.
Key Words: Insecta, Neaspilota, Hemizonia, Asteraceae, nonfrugivorous Tephritidae, bi- ology, taxonomy of immature stages, flower-head feeding, monophagy, seed
predation, parasitoid
Revision of the genus Neaspilota (Dip- tera: Tephritidae) by Freidberg and Mathis (1986) facilitated determination of speci- mens reared from California Asteraceae (Goeden 1989) and stimulated several life- history studies, including those on N. viri-
descens Quisenberry (Goeden and Headrick 1992) and N. wilsoni Blanc and Foote (Goeden and Headrick 1999). This paper describes the immature stages and life his- tory of a third species from California, Neaspilota signifera (Coquillett).
70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
MATERIALS AND METHODS
The present study was based in large part on dissections of subsamples of flower heads of Hemizonia pungens (Hooker and Arnott) Torrey and Gray collected during March—May, 1995-1997 from the San Ja- cinto Wildlife Area at 390-m_ elevation, Lakeview, Riverside Co., administrated by the State of California, Department of Fish and Wildlife. One-liter samples of excised, immature and mature flower heads contain- ing eggs, larvae, and puparia were trans- ported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, pho- tography, description, and measurement. Five first-, 11 second-, and 15 third-instar larvae, and 11 puparia dissected from flow- er heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Ad- ditional puparia were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hy- drated to distilled water in a decreasing se- ries of acidulated EtOH. They were osmi- cated for 24 h, dehydrated through an in- creasing series of acidulated EtOH and two, 1-h immersions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a Philips XL-30 scanning electron mi- croscope in the Institute of Geophysics and Planetary Physics, University of California, Riverside.
Most adults reared from isolated puparia were individually caged in 850-ml, clear- plastic, screened-top cages with a cotton wick and basal water reservoir and provi- sioned with a strip of paper toweling im- pregnated with yeast hydrolyzate and su- crose. These cages were used for studies of longevity and sexual maturation in the in- sectary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Single pairs of virgin males and females obtained
from emergence cages also were held in each of six, clear-plastic, petri dishes pro- visioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observa- tions of their courtship and copulation be- havior.
Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden et al. (1998a, b), Goeden and Headrick (1992, 1999), Goe- den and Teerink (1997, 1998, 1999a, b, c), Teerink and Goeden (1998, 1999), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher speci- mens of N. signifera immature stages, adults, and parasitoids reside in my research collections.
RESULTS AND DISCUSSION Taxonomy
Adult.—Neaspilota signifera was de- scribed by Coquillett (1894). Freidberg and Mathis (1986) designated a male lectotype from Los Angeles, California, and pictured the unpatterned wing [as did Foote et al. (1993)] along with drawings of the lateral aspect of the head, male right foretarsus, epandrium and cerci, aculeus and its apex enlarged, and spermatheca.
Immature stages.—The egg, larvae, and puparium heretofore have not been de- scribed nor illustrated.
Egg: Twenty-five ova dissected from four, field-collected, mature females of WN. signifera plus nine eggs measured in situ in dissected, field-collected flower heads (Fig. 5A) were white, opaque, smooth, elongate- ellipsoidal, 0.72 + 0.01 (range, 0.54—0.85) mm long, 0.20 + 0.003 (range, 0.16—0.24) mm wide, smoothly rounded at tapered bas- al end; pedicel also tapered, 0.025 + 0.001 (range, 0.02—0.03) mm long.
All eggs found in flower heads were damaged during removal and unsuitable for
VOLUME 102, NUMBER 1
scanning electron microscopy. The egg of N. signifera is similar in shape to those of N. viridescens and N. wilsoni but longer and wider on average than that of N. viridescens (Goeden and Headrick 1992) and shorter and narrower than that of N. wilsoni (Goe- den and Headrick 1999).
First instar: White, elongate-cylindrical, bluntly rounded anteriorly and posteriorly (Fig. 1A), minute acanthae circumscribe in- tersegmental lines; gnathocephalon with pair of prominent, flattened, integumental petals dorsad to mouthhooks (Fig. 1B-1); mouthhook bidentate (Fig. 1B-2); median oral lobe laterally flattened, apically pointed (Fig. 1B-3); each integumental petal fused laterally with prominent, stomal sense or- gan (Fig. 1B-4). Posterior spiracular plate bears two ovoid rimae, ca. 0.01 mm in length (Fig. 1C-1), and four interspiracular processes, each with two to four branches, longest measuring ca. 0.01 mm (Fig. 1C-2).
The poor condition of my specimens of first instar N. signifera allowed few com- parisons with those of N. viridescens (Goe- den and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999); however, their general habitus is similar and their mouthhooks also are bidentate. Like N. wil- soni, but not N. viridescens, the integumen- tal petal is fused laterally with the stomal sense organ in first instar N. signifera, a condition first reported in Trupanea vicina (Wulp) by Goeden and Teerink (1999b).
Second instar: White, elongate-cylindri- cal, rounded anteriorly, truncated postero- dorsally (Fig 2A), minute acanthae circum- scribe intersegmental lines (Fig. 2B-1); gnathocephalon conical (Fig. 2B); dorsal sensory organ a well-defined, dome-shaped papilla with a basal pore sensillum on each side (Fig. 2C-1); anterior sensory lobe (Fig. 2C-2, D-1) bears the terminal sensory organ (Fig. 2C-3, D-2), lateral sensory organ (Fig. 2D-3), supralateral sensory organ (Fig. 2D- 4), and pit sensory organ (Fig. 2D-5); sto- mal sense organ prominent, ventrolaterad of anterior sensory lobe (Fig. 2D-6); mouth- hook bidentate (Fig. 2C-4); median oral
71
AccV SpotMagn WD /————4 1004m 10.0kV 3.0 266x 395
Acc.V Spot Magn 100 kV.3.0 6
Fig. 1. habitus, anterior to left; (B) gnathocephalon, antero- lateral view, |— integumental petal, 2— mouth hook,
First instar of Neaspilota signifera: (A)
3— median oral lobe, 4— stomal sense organ; (C) posterior spiracular plate, 1— rima, 2— interspiracular process.
lobe laterally flattened (Fig. 2C-5); single row of four, papilliform, integumental pet- als dorsal to each mouth hook (Fig. 2C-6, D-7); six oral ridges with long axes parallel in row lateral to mouth hooks, oral ridges
72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
AccV. Spot Magn WD /-——— |. 200 um Acc Spot Magn 10.0. kV. 3.0 _122x 37.2 : 10.0 kV 3:0 09x
Spot Magn
AccV Spot Magn WD Budo tha Atc:V 100kV 3.0 1484x 368 ‘ Wo 10.0 kV
pAcc.V Spot Magn WO kV 3.0 . 2478x
Fig. 2. Second instar of Neaspilota signifera: (A) habitus, anterior to left; (B) gnathocephalon, anterolateral view, |— minute acanthae, 2— oral ridge, 3— pit sensillum; (C) gnathocephalon, 1— dorsal sensory organ, 2— anterior sensory lobe, 3— terminal sensory organ, 4— mouthhook, 5— median oral lobe, 6— integumental petal, 7— oral ridge, 8— pit sensillum; (D) gnathocephalon, 1— anterior sensory lobe, 2— terminal sensory organ, 3— lateral sensory organ, 4— supralateral sensory organ, 5— pit sensory organ, 6— stomal sense organ, 7— integumental petal; (E) anterior thoracic spiracle, 1— minute acanthae; (F) posterior spiracular plate, 1— rima, 2— interspiracular process.
dentate along ventral margins (Fig. 2B-2, oblong papillae (Fig. 2E); minute acanthae C-7); pit sensilla circumscribe gnathoce- circumscribe anterior margin of prothorax phalon posteriorad of oral ridges (Fig. 2B- (Fig. 2B-1, B-1); lateral spiracular com- 3, C-8); anterior thoracic spiracle with three plexes not seen; posterior spiracular plate
VOLUME 102, NUMBER 1
bears three ovoid rimae, ca. 0.01 mm long (Fig. 2F-1), and four interspiracular pro- cesses, each with two to four branches, lon- gest measuring 0.009 mm (Fig. 2F-2).
The habitus of the second instar of WN. signifera is more like N. wilsoni (Goeden and Headrick 1999) than the barrel-shaped second instar of N. viridescens (Goeden and Headrick 1992). Unlike both N. viridescens (Goeden and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999), the dorsal sensory organ of N. signifera is well de- fined in the second instar. However, one dis- tinction between the first and second instar common to all three species of Neaspilota is the presence of oral ridges with dentate margins in the latter instar (Goeden and Headrick 1992, 1999). The stomal sense or- gans of the second instar of all three species are well developed and bear sensory struc- tures, variously described as conical in N. viridescens (Goeden and Headrick 1992), papillose in N. wilsoni (Goeden and Head- rick 1999), but verruciform in N. signifera (Fig. 2D-6). Also, the integumental petals of the second instars of all three species are papilliform and about four in number above each mouthhook in N. signifera (Fig. 2C-6, D-7); whereas, in the first instars these structures are broad, flattened, and paired (Goeden and Headrick 1992, 1999). The mouthhooks of the second instar of N. sig- nifera are bidentate, like those of N. wilsoni (Goeden and Headrick 1999): whereas, those of N. viridescens are tridentate (Goe- den and Headrick 1992). Similar, apparent interspecific differences in dentation were noted among mouthhooks of second instar Trupanea spp. (Goeden and Teerink 1999b and references therein). Finally, the inter- spiracular processes each bear two to four branches (Fig. 2F-2), not five to nine branches like those of N. viridescens (Goe- den and Headrick 1992), nor two to six branches like those of N. wilsoni (Goeden and Headrick 1999).
Third instar: White to pale yellow, with posterior spiracular plate dark brown to black, elongate-cylindrical, tapering anteri-
713
orly; posterior spiracular plate on caudal segment flattened and upturned dorsally ca. 60° (Fig. 3A), minute acanthae circum- scribe thoracic and abdominal segments an- teriorly, acanthae more numerous on pos- terior segments; gnathocephalon conical (Fig. 3B); dorsal sensory organ a well-de- fined, dome-shaped papilla (Fig. 3D-1), pit sensillum on each side at base of dorsal sensory organ (Fig. 3D-2); anterior sensory lobe (Fig. 3C-1) bears the terminal sensory organ (Fig. 3C-2), pit sensory organ (Fig. 3C-3), lateral sensory organ (Fig. 3C-4), and supralateral sensory organ (Fig. 3C-5); eight to 10 papilliform, integumental petals in double row above each mouth hook (Fig. 3C-6, D-3); six oral ridges laterad of mouthhook, oral ridges dentate along ven- tral margins (Fig. 3B-1, C-7); stomal sense organ prominent ventrolaterad of anterior sensory lobe (Fig. 3C-8, D-4); mouth hook tridentate (Fig. 3B-2, C-9, D-5); median oral lobe laterally flattened, apically pointed (Fig. 3C-10, D-6); prothorax circumscribed anteriorly with minute acanthae (Fig. 3B- 3); anterior thoracic spiracle on posterior margin of prothorax bears three to four ob- long papillae (Fig. 3E); spiracle of metatho- racic lateral spiracular complex absent or not seen, this complex otherwise consisting of vertical row of four verruciform sensilla and verruciform sensillum posterior to up- per verruciform sensillum (not shown, but otherwise like Fig. 3F); abdominal lateral spiracular complex consists of a spiracle (Fig. 3F-1), vertical row of four verruci- form sensilla (Fig. 3F-2), and a single ver- ruciform sensillum posterior to upper ver- ruciform sensillum in vertical series (Fig. 3F-3); caudal segment broadly circum- scribed by minute acanthae (Fig. 3G-1); ste- lex sensilla dorsolaterad and ventrolaterad of posterior spiracular plates (Fig. 3G-2), lateral stelex sensilla not seen; posterior spi- racular plate bears three ovoid rimae, ca. 0.029 mm in length (Fig. 3G-3), and four interspiracular processes (Fig. 3G-4), each with one to three, simple, pointed branches, longest branch measuring 0.009 mm; inter-
74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
| Acc: 10.
2 4
0
a bee 4 « we +
ee
Acc Spot Magn . WD. H-————-J 10.0kV 4.0 -2038x 36.3
~— OM e o O80 [G9 VIG 9, Pa So ww wy
Fig. 3. Third instar of Neaspilota signifera: (A) habitus, anterior to left; (B) gnathocephalon, anterolateral view, 1— oral ridge, 2— mouthhook, 3— minute acanthae; (C) 1— anterior sensory lobe, 2— terminal sensory organ, 3— pit sensory organ, 4— lateral sensory organ, 5— supralateral sensory organ, 6— integumental petal, 7— oral ridge, 8— stomal sense organ, 9— mouth hook, 10— median oral lobe; (D) gnathocephalon, anterior view, 1— dorsal sensory organ, 2— pit senillum, 3— integumental petal, 4— stomal sense organ, 5— mouth- hook, 6— median oral lobe; (E) anterior thoracic spiracle; (F) first abdominal spiracular complex, 1— spiracle,
VOLUME 102, NUMBER 1
mediate sensory complex with a medusoid sensillum (Fig. 3H-1) and a stelex sensillum (Fig. 3H-2) ringed by minute acanthae (Fig. 3H-3).
The habitus of the third instar of N. sig- nifera generally is like that reported for N. viridescens Goeden and Headrick (1992) and N. wilsoni (Goeden and Headrick 1999), except that the minute acanthae cir- cumscribe the body segments differently; i.e., in N. signifera, the anterior part of each segment is circumscribed; in N. wilsoni, all intersegmental areas and all abdominal seg- ments except the pleura are circumscribed; and in N. viridescens, the intersegmental ar- eas are free of acanthae. Unlike these two congeners, the dorsal sensory organ is well defined in the third instar of N. signifera (Fig. D-1) as well as in the second instar; however, I was unable to determine whether this also held for the first instar. The dorsal sensory organ is well defined in the first instar of both congeners. Thus, speculation by Goeden and Headrick (1992, 1999) that this intraspecific, differential degree of def- inition of the dorsal sensory organ may be a consistent generic character has now been invalidated by comparison with a third spe- cies of Neaspilota.
Additional bases for differention between instars were noted. The integumental petals in the third instar N. signifera are papilli- form and arranged in a double row above each mouthhook, but are less numerous and form a single row in the second instar. The integumental petals in the second and third instars of N. viridescens also are papilliform and situated similarly (Goeden and Head- rick 1992), but in N. wilsoni are fewer in the third instar (Goeden and Headrick 1999). The stomal sense organs of the third instars of all three species are well devel- oped and bear different sensory structures,
TS
described as several cone-shaped sensilla in N. viridescens (Goeden and Headrick 1992): as papilliform and pit-type in N. wil- soni (Goeden and Headrick 1999); and as verruciform (Fig. 3C—8), or what might be termed ‘“‘compound verruciform’’, as shown in Figure 3D—4. Like the second instar of N. signifera, the third instar has oral ridges with dentate margins, also reported in the second and third instars of N. viridescens (Goeden and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999). Counting only those oral ridges with dentate margins in a single row ventral to the stomal sensory organ, these apparently number six in the second and third instars of all three species examined to date; therefore, further com- parison with additional species may show this to be a consistent generic character. The third instars of Trupanea imperfecta, T. jo- nesi, T. nigricornis, T. pseudovicina, T. sig- nata, and T. wheeleri also bear serrated oral ridges (Goeden and Teerink 1997, 1998, 1999b, Goeden et al. 1998a, Knio et al. 1996, Teerink and Goeden 1998), but these oral ridges appear to be fewer in number, and not arranged in a more or less regular, vertical row laterad to the oral cavity, as in Neaspilota. Also, the mouth hooks of the third instars of N. signifera and N. virides- cens are tridentate (Goeden and Headrick 1992); whereas, those of N. wilsoni are bi- dentate (Goeden and Headrick 1999). Such interspecific differences in dentation are supported by our findings that the mouth hooks of third-instar Trupanea vicina are bidentate; whereas, those of 12 other con- geners examined from California are triden- tate (Goeden and Teerink 1999b and cita- tions therein).
Puparium: Mostly white to yellow, with posterior 2—3 segments grayish to black- ened posteriorly, broadly ellipsoidal and
<_ 2— verruciform sensillum, 3— verruciform sensillum; (G) caudal segment, 1— minute acanthae, 2— stelex sensillum, 3— rima, 4— interspiracular process; (H) intermediate sensory complex, 1— medusoid sensillum,
2— stelex sensillum, 3-minute acanthae.
76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Acc.V. ‘Spot Magn WD -—————{_ 500 um
10:0 kV 3.0 46x
Spot Magn - WD Tf 0. 306x535 6
Fig. 4. itus, anterior to left; (B) anterior end, 1— invagination scar, 2— anterior thoracic spiracle; (C) caudal seg- ment, |— rima, 2— interspiracular process.
Puparium of Neaspilota signifera: (A) hab-
smoothly rounded at both ends (Fig. 4A), minute acanthae circumscribe body seg- ments anteriorly; anterior end bears the in- vagination scar (Fig. 4B-1) and anterior thoracic spiracles (Fig. 4B-2); caudal seg- ment circumscribed by minute acanthae, two stelex sensilla, dorsad and ventrad of
posterior spiracular plates; posterior spirac- ular plate bears three broadly elliptical ri- mae (Fig. 4C-1), and four interspiracular processes, each with two to three branches (Fig. 4C-2); intermediate sensory complex consists of a medusoid sensillum and a ste- lex sensillum. Twenty-five puparia aver- aged 2.43 + 0.03 (range, 2.20—2.70) mm in length; 1.10 + 0.02 (range, 0.92—1.28) mm in width.
DISTRIBUTION AND Hosts
The distribution of N. signifera as mapped by Foote et al. (1993) restricted this tephritid to Arizona, California, Oregon, and Washington in the western United States North of Mexico. The distri- bution of its main, probably sole, host plant, H. pungens, apparently does not extend far into Mexico (Shreve and Wiggens 1964, Hickman 1993), nor, perhaps, does N. sig- nifera. Accordingly, Freidberg and Mathis (1986) listed only two collection records for N. signifera from northern Mexico, one from Baja California Norte, and one from Sonora.
Hemizonia pungens belongs to the sub- tribe Madiinae of the tribe Helenieae of the family Asteraceae (= Compositae) (Bremer 1994). I have not reared N. signifera from six other species of Hemizonia from Cali- fornia, involving a total of 11, 1-liter sam- ples of mature flower heads. Freidberg and Mathis (1986) also listed Lasthenia (as Baeria) fremontii (A. Gray) E. Greene as another host, apparently based on two sep- arate series of specimens labelled “‘ex’’ or “on” this plant and collected over 5- and 18-day periods, respectively, from Califor- nia. Lasthenia fremontii belongs to a dif- ferent subtribe Baeriinae of the Helenieae. I have not reared N. signifera, nor any other species of Neaspilota (only Trupanea spp.), from 18 samples of mature flower heads from three species of Lasthenia, but not in- cluding L. fremontii (a northern California species, Hickman 1993). Furthermore, 11 of the 12 described species of Neaspilota in California for which hosts are now known
VOLUME 102, NUMBER 1
(Goeden 1989 and unpublished data) have hosts belonging solely to the tribe Astereae, with only N. sp. prob. punctistigma Benja- min or near, besides N. signifera, having been reared by me from flower heads of a host in the tribe Helenieae (subtribe Pecti- dinae). In addition, the generalist species, N. viridescens, has now been reared from 21 host species in the tribe Astereae and a single host in the Tribe Senecioneae (Goe- den, 1989, Goeden and Headrick 1992, and unpublished data). Therefore, the above in- formation suggests that the host record for L. fremontii in Freidberg and Mathis (1986) is suspect, possibly based on sweep records for adults, which are problematic predictors of reproductive hosts (Headrick and Goe- den 1998), and that N. signifera is a true monophage on H. pungens.
BIOLOGY
Egg.—In 10, closed, preblossom, imma- ture flower heads of H. pungens, 12 eggs were inserted pedicel-last, usually between the chaff and ovules, and perpendicular to the receptacle (Fig. 5A). No flower head contained any floret damaged by oviposi- tion. The diameters of the receptacles of these 10 flower heads containing eggs av- eraged 3.64 + 0.13 (range, 2.42—4.56) mm, and these heads held an average of 1.2 + 0.1 (range, 1-2) eggs.
Larva.—Upon eclosion, the first instars usually tunneled into an ovule, or into a co- rolla before entering an ovule (Fig. 5B). One first instar was found feeding within each of 18, closed, preblossom flower heads, the receptacles of which averaged 3.63 + 0.18 (range, 2.0—5.13) mm in di- ameter. An average of only 1.3 + 0.3 (range, O—3) ovules was damaged in these 18 heads, or based on 93 + 4 (range, 52-— 150) as the average number of ovules/ achenes counted in 51 preblossom to post- blossom flower heads, about 1.4% (range, O-—3.2%) of the ovules were damaged. No receptacles within these 18 infested flower heads were abraded or pitted by feeding of first instars.
a
Second instars continued feeding on ovules in preblossom flower heads or in soft achenes in open flower heads (Fig. 5C). All fed with their bodies horizontal to and their mouthparts directed towards the receptacles within a series of adjacent ovules/soft achenes, but always well above the recep- tacles. Receptacles of 23 flower heads con- taining second instars were not fed upon and averaged 4.15 + 0.12 (range, 3.13- 5.15) mm in diameter. These flower heads contained an average of 1.3 + 0.1 (range, 1—3) second instars that had destroyed an average of 6.7 + 0.9 (range, 2—18) ovules/ soft achenes, as calculated for the preceding instar, about 7% (range, 2—19%) of the av- erage total of 93 ovules/soft achenes per flower head.
Third instars fed mainly on soft achenes in the centers of open flower heads, or in one case, a postblossom flower head. Thir- ty-four flower heads that averaged 4.09 + 0.14 (range, 2.85—5.70) mm in diameter contained an average of 1.3 + 0.14 (range, 1—4) third instars. An average of 25 + 2.4