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VOL. 90 JANUARY 1988 ‘NO. i 95, 90674 (ISSN 0013-8797)
(> PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY ot WASHINGTON °
b 05 1988 PUBLISHED apts “QUARTERLY
BURGER, J. F.—A new genus and two new species of Pangoniini (Diptera: Tabanidae) of zoogeographic interest from Sabah, Malaysia ................00.0 000 cece eee eee ee 12
CLEMENT, S. L. and T. MIMMOCCHI—Occurrence of selected flower head insects of CETIOUTEER SOISIIIAUS\AN Italy ANGUGLEECE | av be\s oh) lM b ahlasicters balers eizk, Al an lets eile wlele Kabres 47
DROOZ, A. T. and H. H. NEUNZIG—Notes on the biology of two Phycitines (Lepidoptera: Pyralidae) associated with Toumeyella pini (Homoptera: Coccidae) on Pine ............ 44
GOEDEN, R. D.—Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica (Dip- tera a Me DHnrTGde) Meee Sh Le ett ae eL Ee Era he RT PP ROME UN BERET MAD AS ML STS Ore i 57)
HALSTEAD, J. A.—Belaspidia longicauda, new species, the first Nearctic Belaspidia (Hyme- MOPLETEAGhalGididae) cat Wl RU asd) Suna polis he FEAL OP REPEAL der eat fet be Pe eek 87
HANSSON, C.—A revision of the genus Mestocharis and a review of the genus Grahamia (iymenopterd, BUlOophidazey sy Bitty tee Aaete e AWE eA RETRO URIBE NS eat erp: 28
HEYDON, S. L.—A review of the Nearctic species of Cryptoprymna Forster, with the description of a new genus, Polstonia (Hymenoptera: Pteromalidae) .........................----- 1
MAcDONALD, J. F.—New synonyms pertaining to Chelifera and generic key for North Amer-
ican Llemerodromiinae: (Diptera: |Empididae) 2.00. Waki joke ia eee AAD st 98 MATHIS, W. N. and T. ZATWARNICKI—Studies on the systematics of the shore-fly tribe
DAPini (Diptera Ephydridae) Mansy Week Ue k M ME Lite LORIE EM) Bel Le Bea TPM Niel i 106 NEAL, J. W., Jk.—Unusual oviposition behavior on evergreen azalea by the Andromeda lace
bug Stephanitis takeyai (Drake and Maa) (Heteroptera: Tingidae) .................... 52
OSWALD, J. D.—A review of the South Pacific genus Austromegalomus Esben-Petersen (Neu-
roptera: Hemerobiidae) with a description of a new species from Rapa ................. 55 PETERSON, B. V., M. VARGAS V., and J. RAMIREZ-PEREZ—Simulium (Hemicnetha)
hieroglyphicum (Diptera: Simuliidae), a new black fly species from Costa Rica .......... 76 RAMIREZ-PEREZ, J., B. V. PETERSON, and M. VARGAS V.— Mayacnephia salasi (Diptera:
Simuliidae), a new black fly species from Costa Rica ...........0...0. 000020 e cece eee 66
STARK, B. P., S. W. SZCZYTKO, and B. C. KONDRATIEFF—The Cultus decisus complex of Eastern North America (Plecoptera: Perlodidae) .................0...00..0 22200005.
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1988
F. EUGENE Woon, President WARREN STEINER, Program Chairman F. CHRISTIAN THOMPSON, President-Elect GEOFFREY B. WHITE, Membership Chairman RICHARD Rossins, Recording Secretary ANNE WEIBER, Custodian JOHN KINGSOLVER, Corresponding Secretary MAnyA B. STOETZEL, Delegate, Wash. Acad. Sci.
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PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 1-11
A REVIEW OF THE NEARCTIC SPECIES OF CR YPTOPRYMNA FORSTER, WITH THE DESCRIPTION OF A NEW GENUS, POLSTONIA (HYMENOPTERA: PTEROMALIDAE)
STEVEN L. HEYDON
Illinois Natural History Survey, Section of Faunistic Surveys and Insect Identification, 607 E. Peabody Drive, Champaign, Illinois 61820.
Abstract.—The genus Cryptoprymna Forster, herein reported from the Nearctic region for the first time, is represented by two species, the Holarctic species C. atra (Walker) and the Nearctic species C. dixiana n. sp. Cryptoprymna is redescribed. C. atra is reported herein to be a parasitoid of the pupal stages of syrphids on conifers. A new genus, Polstonia, is described with two included Nearctic species: P. quadriplana n. sp., the type species, and P. pelagocorypha n. sp. A modification of Graham’s (1969) key to the genera of the Sphegigasterini is presented to facilitate identification of the genus Po/stonia. Keys are
given to the Nearctic species of both genera.
This is the second, following Heydon and LaBerge (in press), in a series of papers re- vising the Nearctic miscogasterine Pter- omalidae. The specimens upon which this review 1s based were among the material submitted to me by various collections for that first paper, which was a revision of the genus Sphegigaster Spinola. Terminology and methodology follow those used by Hey- don and LaBerge (in press) except that de- scriptions of new species are based on the type-specimens, the “‘sensillae’’ of the fu- nicular and club segments are called mul- tiporous plate (abbreviated MPP) sensillae, and the specimens were examined under fluorescent light so may appear more green than described herein when viewed under incandescent light.
This paper contains the first Nearctic rec- ord for Cryptoprymna Forster. The Nearctic fauna of Cryptoprymna contains one Hol- arctic species, C. atra (Walker), and the new Nearctic species C. dixiana. Because the original description of the genus (Walker 1833) is now insufficient for distinguishing
this genus from several other similar genera erected since 1833, I am redescribing the genus based on my examination of three of the four described species. The original de- scription of C. atra is also very short so I am presenting a detailed diagnosis for dis- tinguishing this species from C. dixiana. This diagnosis 1s based on the Nearctic specimens of C. atra, which were compared by me with specimens from the Palearctic region from the British Museum of Natural History.
The new genus Polstonia is described and followed by descriptions of the two new Nearctic species included in the genus, P. quadriplana and P. pelagocorypha. The geo- graphic range of this genus extends into the Neotropical region since I have seen spec- imens from South America that belong to other species in this genus. None of these Neotropical species will be described here due to lack of sufficient material for a thor- ough study of the fauna of this region.
Both these genera key to the Sphegigas- terini in Graham (1969) and are similar in
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
having the anterior margin of the clypeus without any projecting denticles and retic- ulate petioles which are distinctly longer than wide. Relationships between these genera and the other genera of the Miscogasterinae will be discussed more fully in a later paper.
Cryptoprymna Forster
Prosodes Walker, 1833: 371, 374 (Preoc- cupied by Eschscholtz 1829). Type- species: Prosodes ater Walker 1833 (monotypy). Lectotype male in the BMNH. Brulle, 1846: 582-583. Gahan and Fagan, 1923: 121.
Cryptoprymna Forster, 1856: 52 (key), pp. 56, 59. Walker, 1872: 97, 98 (key, syn- onymy). Ashmead, 1904: 330, 332, 372 (key). Nikol’skaya, 1952: 252 (key). Schmiedeknecht, 1909: 375, 376, 380 (key, diagnosis). Gahan and Fagan, 1923: 41. Peck, Boucek, and Hoffer, 1964: 40 (key). Graham, 1969: 124, 140 (key, syn- onymy). Dzhanokmen, 1978: 77, 80 (key). Farooqiand Subba Rao, 1985: 260, 310G. Farooqi and Subba Rao, 1986: 285.
Cryptoprymnus: Thomson, 1878: 17, 22. Cresson, 1887: 75 (key). [Invalid emen- dation]
Walker (1833) erected the genus Prosodes in his Monographia Chalciditum. Forster (1856) pointed out that the name Prosodes had previously been proposed for a genus of tenebrionid beetles (Eschscholtz 1829), and he renamed the genus Crypfoprymna. There are presently three described species: C. atra (Walker 1833), a Holarctic species; C. africanus Boucek (1976), from southern Africa; and C. brama (Motschulsky 1863), from southern Asia. I add a fourth species, C. dixiana, from the southeastern United States. Ashmead (1896) described Crypto- prymna illinoensis from the Nearctic, but this species was transferred to Callitula Spi- nola (Delucchi 1955) and later synony- mized with Callitula cyrnus Walker (Burks 1975).
Description.— Color: Head, mesosoma, coxae, and petiole black, gaster dark brown. Wing hyaline.
Female.— Head transversely oval in an- terior view; 2x as wide as long; clypeus subareolate, anterior margin truncate: gena with broad concavity extending from mouth margin to lower orbit; eye glabrous; occiput concave or straight posteriorly, acarinate. Antenna inserted below middle of face, just above lower orbits; formula 1:1:2:6:3; scape slender, length 8 x width, reaching or nearly reaching median ocellus; club distinctly wider than F6, sutures oblique, with a patch of micropilosity on terminal or terminal two segments, terminal spine or projection ab- sent. Mandible 4-tooth, upper two smaller and approximated. Mesosoma compact, rounded dorsally in profile; pronotum with neck short, collar with sharp transverse ca- rina anteriorly and smooth except imme- diately behind carina; mesoscutum with no- tauli extending to its hind margin as impressed lines; scutellum as long as wide, frenal sulcus nearly obliterated; prepectus acarinate; mesopleuron with upper epime- ron smooth; propodeum as long as scutel- lum, strongly arched, median carina and plicae sharp, median panels alveolate with some rugae, spiracles round, nucha unde- veloped. Wing with basal cell and vein setu- lose or bare; speculum present; relative lengths of wing veins: marginal > post- marginal > stigmal; stigma unenlarged, only 2-3 as wide as stigmal vein. Legs with coxae relatively small; hind tibia with one apical spur. Petiole much longer than wide, cylindrical; areolate dorsally, strigose lat- erally; with basal flange laterally and ven- trally; lateral setal row sometimes present as a few weak setae anteriorly. Gaster ovate; T1 enlarged, nearly concealing succeeding terga, hind margin convex; hypopygium reaching to tip of gaster.
Male.—Similar to female, but antenna with funiculus parallel-sided, all segments elongate: club lacking area of micropilosity, palps unmodified.
VOLUME 90, NUMBER |
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Figs. 1-8. fore wing showing arrangement of dorsal setae along basal vein and admarginal setae. 4, Female head (dorsal view). C. dixiana n. sp. 5, Female propodeum and petiole. 6, Female head (anterior view). 7, Female head (dorsal view). 8, Female fore wing showing bare basal vein and arrangement of ventral admarginal setae.
Diagnosis. — Cryptoprymna can be distin- guished by the following unique combina- tion of characters: edentate clypeus; large area of micropilosity on club; carinate pronotum; propodeum arched and as long as scutellum, median carina and plicae de- veloped; elongate sculptured petiole; en- larged Tl; hypopygium extending to tip of
Cryptoprymna atra (Walker). 1, Female whole body. 2, Female propodeum and petiole. 3, Female
gaster, and loss of metallic coloration. Though these character states are all apo- morphic within the Miscogasterinae, none of these characters is unique to this genus. This combination of apomorphies 1s unique, however.
Biology.—Little is known of the biology of the species 1n this genus. Graham (1969)
4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mentions specimens of C. atra taken on Abies sp. and Pinus sylvestrus L. Among the Nearctic specimens of C. afra, is one from Stockholm, Maine, mounted with a syrphid pupa that has a lateral emergence hole and a label reading “beaten from fir.” A speci- men of C. dixiana n. sp. from Fort Pierce, Florida, was reared from a similar pupa, but there is no information on the plant source of this pupa. It seems likely that species of this genus are parasitoids of syrphids on co- nifers.
Key To NEARCTIC SPECIES OF CRYPTOPRYMNA FORSTER 1. Wing with basal cell and vein setate, ventrally with a patch of setae behind the marginal vein (Fig. 3). Occiput concave in dorsal view (Fig. 4). Petiole bare (Fig. 2) atra (Walker) — Wing with basal cell and vein bare, ventrally with only a single row of setae behind the mar- ginal vein (Fig. 8). Occiput straight in dorsal view (Fig. 7). Petiole with short lateral setal row (Fig. 5) dixiana Nn. sp.
Cryptoprymna atra (Walker) (Figs. 1-4)
Prosodes atra Walker, 1833: 375. Lectotype male (designated by Graham 1969) in the Westwood collection (BMNH) (not seen). Westwood, 1840: 68-69. Haliday, 1842: V, Plate C (figure). Walker, 1872: 94 (fig- ure); 1873: 371 (figure). Gahan and Fa- gan, 1923: 41, 121.
Cryptoprymnus cavigenaThomson, 1878: 22. Lectotype female (designated by Gra- ham 1969) in the collection of Univer- sitetets Zoologiska Institutionen, Lund (not seen).
Cryptoprymna atra (Walker): Schmiede- knecht, 1909: 380. Delucchi, 1955: 174 (synonymy). Boucek, 1961: 71 (distri- bution). Graham, 1969: 140-141 (biolo- gy, synonymy, distribution). Boucek, 1976: 14-15. Dzhanokmen, 1978: 80.
Diagnosis. —In addition to the characters given in the key, female C. atra differ from C. dixiana in the following ways: the trun- cate portion of the clypeus of C. atra has a
concave anterior margin and the anterior lateral corners are sharp while the anterior margin is straight and the corners rounded in C. dixiana; the antennal flagellum of C. atra is longer, 0.89 + (S.E.) 0.014 (n = 4) times as long as the head width compared with 0.81 times in C. dixiana; the antennal club of C. atra is more slender, 2.1 + 0.21 times as long as wide compared to 1.6 times in C. dixiana; and the wings of C. atra are longer, 2.2 + 0.13 times the mesosomal length versus 1.7 times in C. dixiana.
Biology.—The specimen from Stock- holm, Maine, was reared from the pupa of a syrphid which was “beaten from fir.”
Nearctic material examined (CNC, INHS, USNM): Canada. BRITISH COLUMBIA: Terrace, 8-VIII-1960, 1 ¢. NEW BRUNS- WICK: Acadia Experiment Station (Fred- ricton) 1-17-VH-1970, 1 4, 13-VIII-1970, 1 6. QUEBEC: Messines, 10-VII-1947, 1 4; Parke Reserve (near St. Eleuthere), 1 3-VIII- 1957, 1 6. United States. MAINE: Stock- holm, 6-VI-1955, 1 2°. MICHIGAN: Isle Royale, 3-7-VHI-1936, 1 °. OREGON: Saddleback Mt. (near Rose Lodge), 1 1-VIII- LOGIS le
The records of C. atra from Greenland cited by Bakkendorf (1955) are in error (Boucek 1961).
Cryptoprymna dixiana, NEw SPECIES (Figs. 5-8)
Description. — Holotype female: Color. Body black with mesosoma tinged blue, scutellum and propodeum tinged gold. An- tenna with scape brownish yellow; pedicel and flagellum brown. Legs reddish brown, femora and mid tibia with dark bands; tarsi light brown, pretarsus dark brown. Head and mesosoma with scattered, short (one half ocellar diameter) white setae.
Sculpture. —Clypeus, median area of face subareolate; face laterally and dorsally, frons, vertex finely alveolate; gena coriaceous. Mesosoma with pronotal collar with trans- verse row of punctures posterior to anterior
VOLUME 90, NUMBER 1
transverse Carina; mesoscutum alveolate, side lobes more finely so than median lobe; scutellum areolate. Gaster T1 polished; T7 coriaceous.
Structure.— Mesosomal length 0.85 mm. Relative lengths of head, mesosoma, gaster; 16:42.5:33. Head broadly oval in anterior view (Fig. 6), width 1.2 height (33:27), 2.1 length (33:16); clypeus with anterior margin nearly straight mesally, anterior cor- ners rounded; eye height 1.4 length (16: 11), 2.1 x malar length (16:7.5); POL 1.3 x OOL (8:6), 1.5 x LOL (8:4); occiput straight in dorsal view (Fig. 7). Antenna inserted a quarter of the way up eye; scape length 0.88 x eye height (14:16), reaching to median ocel- lus, slightly recurved; length of pedicel plus flagellum 0.82 head width (27:33); rela- tive lengths of segments (annelli omitted, club taken as a unit) scape = 14:3:2.5:3:3: 2.5:2.5:2:8; widths of F1, F6, club as 2:4:5; F1-2 elongate, F3-—4 quadrate, F5—6 trans- verse; MPP sensillae two thirds length of segment, arranged in single row; club length 1.6 x width (8:5), asymmetrically curved to outside, sutures oblique, area of micropi- losity extending to midway down C2, C3 pointed apically. Mesosoma with meso- scutal length 0.48 x width (12:25); scutellar length 0.93 width (13:14); propodeum with costula rugiform, nucha coriaceous, supracoxal flange drawn out over base of hind coxa. Wing length 2.4 x width (71:30); basal cell and basal vein bare (Fig. 8); costal cell with single row of setae; relative lengths of submarginal, marginal, postmarginal, and stigmal veins = 30:14:10:7. Petiole 0.96 x as long as propodeum (13:13.5) (Fig. 5); length 2.4 maximum width (13:5.5); with a central and diverging lateral carinae on basal one fourth; lateral setal rows present as a patch of a few setae anteriorly. Gaster 1.6 as long as wide (35:22); T5-6 with distal fringe of setae; hypopygium with short erect white setae.
Allotype male: Color. Similar to female but funiculus, club dark brown; femora dark brown, lighter distally. Structure. Antenna with flagellum parallel-sided, length of ped-
icel plus flagellum 1.3 x head width (39:31); scape length 0.80 x eye height (12:15). Pet- iole longer (petiole 1.2 propodeal length [14:12]), more slender (petiolar length 2.8 x width [14:5]). Gaster with terminal seg- ments glabrous.
Diagnosis.— Characters for separating C. dixiana from C. atra are given in the key to Nearctic species and in the discussion section for C. atra. C. dixiana can be dis- tinguished from C. africana by the same characters given in the key for distinguish- ing C. dixiana from C. atra, except that C. dixiana and C. africana both lack setae on the basal cell and basal vein. The straight occiput and lack of setae on the basal cell of C. dixiana distinguishes that species from C. brama.
Biology.—The allotype male from Ft. Pierce, Florida, is mounted with a syrphid pupa.
Etymology.—The name is a latinization of Dixie, referring to the southeastern United States distribution of this species.
Type material.— Holotype female is from Andrews, South Carolina, and was collected 8 May 1963 by R. D. Eikenbary (USNM). The allotype male is from Ft. Pierce, Flor- ida, and was collected 26 April 1955 by Holtzburg (USNM).
Polstonia, New GENUS
Type-species.—Polstonia quadriplana Heydon. The gender is feminine. It is my pleasure to name this genus in honor of Jane Polston with whom I have spent many hours collecting.
Description.—Color: Head, mesosoma, and coxae dark blue to green; metasoma dark reddish brown to black. Wing hyaline.
Female: Face slightly bulging viewed in profile; clypeus subareolate, anterior margin straight or slightly produced: genal concav- ity short, reaching only one fourth the dis- tance to lower orbit; eye bare, bulging in anterior view; ocellar triangle width 1.5 x length; occiput acarinate, moderately con- cave. Antenna inserted below middle of face,
6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
just above a line between lower orbits; for- mula 1:1:2:6:3; scape slender (length 7 x width), extending to mid ocellus or higher; funicular segments with MPP sensillae in a single row; club with ventral patch of mi- cropilosity and terminal spinelike protu- berance on C3. Mandible 4-toothed, upper two smaller and approximated or equally spaced. Mesosoma arched dorsally; prono- tum with collar lacking anterior transverse carina, smooth strip along hind margin oc- cupying a third to a half median length; mesoscutum with notauli present as shallow furrows, traceable to hind margin as strip of distinct texture; scutellum as long as wide, with 4-6 pairs of lateral setae, frenal sulcus obscure or absent; prepectus acarinate; me- sopleuron with upper epimeron smooth; propodeum with plicae and median carina complete and distinct, median panels al- veolate-rugose, nucha obscurely sculptured crescent. Wing with basal vein setate; specu- lum present; relative lengths of wing veins: marginal > postmarginal > stigmal; stigma small, width only 2-3 width of stigmal vein. Hind tibia with one apical spur. Pet- iole sculptured dorsally, length 2-3 x width; lateral setal row extending nearly entire length of petiole, setae projecting perpen- dicularly. Metasoma ovate, plicate ventral- ly near insertion of petiole; Tl and T2 sub- equal in length, distinctly longer than the succeeding terga, Tl with hind margin straight or sinuate.
Male: Similar to female but club lacking area of micropilosity, palpi unmodified.
Diagnosis.—The possession by Polstonia of an edentate clypeus, genal concavities, a thirteen-segmented antenna, the female an- tennal club with terminal spine and ventral patch of micropilosity, propodeum with distinct median carina and plicae, and an elongate and reticulate petiole makes this genus phenetically similar to Toxeuma Walker. It differs from Toxeuma by having an acarinate pronotal collar, notauli obscure posteriorly, frenal suture obscured, and lengths of Tl and T2 subequal. In these
characters, Polstonia resembles Sphegigas- ter. However, Sphegigaster species have a bidentate clypeus, never have the female antennal club with a terminal spine and only rarely with a ventral patch of micropilosity, their propodeum lacks the median carina and plicae, and the hind margin of gastral T1 is broadly concave. In contrast, Polston- ia species have an edentate clypeus, the an- tennal club in the female with a terminal spine and ventral patch of micropilosity, a distinct median carina and plicae on the propodeum, and gastral Tl has a nearly straight hind margin. The elongate petiole with complete lateral rows of setae that stick out perpendicularly is the one unique apo- morphic character defining this genus. Polstonia can be included in Graham’s (1969) key to the Sphegigasterini by mod- ifying the first half of couplet one so it goes to couplet la instead of 2, and then inserting the following couplet after the first: la. Clypeus simple (Fig. 9). Petiole with lateral
row of setae extending at least half its length (Figs, WiGand )2)) cs. 1s. . Polstonia Heydon
— Clypeus bidentate or tridentate. Petiole with
a short row of setae extending less than half its length
th
Key TO SPECIES OF POLSTONIA HEYDON
1. Petiole less than 2.3 x as long as wide, rounded dorsally, and areolate with reticulations only 2 as long as wide. Propodeum with area be- tween the basal foveae relatively smooth and divided into four equal sized regions (Fig. 11). Usually only the hind femur with basal dark
tened dorsally, and strigulate dorsally with re- ticulations three or more times as long as wide. Propodeum areolate between basal foveae, sublateral carinae weak or absent (Fig. 12). All femora with dark bands basally ............ Sees ceseee . pelagocorypha n. sp.
Polstonia quadriplana, New SPECIES (Figs. 9-11)
Description. — Holotype female: Color. Head, mesosoma, coxae dark green with coppery reflections; occiput, neck, pleural
VOLUME 90, NUMBER 1
Figs. 9-10. Polstonia quadriplana n. sp. 9, Female head (anterior view). 10, Female whole body.
regions, petiole darker; propodeum paler. Gaster black with greenish reflections. An- tenna with scape brownish yellow; pedicel brown; flagellum black. Mandible brownish yellow; teeth reddish brown. Legs brownish yellow; strong dark bands on mid tibia and hind femur with greenish reflections, fore femur with weak broad dark band: pretarsus darker. Wing veins pale brown. Head, dor- sum of thorax with distinct brown setae.
Sculpture.—Clypeus subareolate; face, frons, vertex, occiput alveolate; neck alveo- late; mesoscutum, scutellum, axilla coarsely alveolate; propodeum with median panels areolate-rugose; petiole finely areolate, cells 2x as long as wide; gastral terga 4-7 sub- imbricate.
Structure.— Mesosomal length 0.96 mm. Relative lengths of head, mesosoma, me- tasoma = 15:48:42. Head width 1.2 x height
Female propodeum and petiole.
8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(37:30) (Fig. 9), 2.5 x length (37:15); clypeus with anterior margin slightly produced; eye height 1.3 width (18:13.5), 2.0x malar length (18:9); POL 1.4x OOL (7:5), 2.3 x LOL (7:3). Antenna with length of pedicel plus flagellum 0.97 head width (36:37); relative lengths of segments (annelli omit- ted, club counted as a unit) scape = 17:6:5: 4:3.5:2.5:3:3:7.5; widths of Fl, F6, club as 3:4.5:5: funicular segments narrowed at bases; Fl-2 elongate, F3 quadrate, F4-6 transverse; MPP sensillae prominent; club length 1.5 < width (7.5:5), sutures oblique. Mesosomal length 1.7 x width (48:29); scu- tellar length 0.97 x width (15.5:16); dorsel- lum tilted nearly perpendicularly with re- spect to lateral metanota (Fig. 10), anterior edge regularly rounded, surface with little sculpture; propodeum with spiracles round, groove between basal foveae smooth and divided by the median and two short sub- median carinae into four subequal areas (Fig. 11). Wing length 2.2 x width (85:38); rela- tive lengths of submarginal, marginal, post- marginal, and stigmal veins = 35:18:16:11; basal vein marked by row of eight setae; basal cell with one seta distally on left wing. Petiole length 1.9 width (13:7); 1.1 x as long as propodeum (13:12); rounded dor- sally, median carina complete and sharp; length of setae in lateral setal rows a half to three fourths width of petiole. Metasoma broadly oval; length 1.4 x width (42.5:32.5); basal region with scattered setae laterally; T4-7 with submarginal row of setae.
Allotype male: Color similar to female ex- cept head and dorsum of mesosoma lacking coppery reflections; antennal flagellum brown. Sculpture similar to holotype. Me- sosomal length 0.98 mm. Antenna with pedicel plus flagellum 1.2 x head width (43: 35); relative lengths of segments 15:5:5:4.5: 4.5:4.5:4:4:9; width of Fl, F6, club as 3:3.5: 3.5. Petiole length 2.0 x width (14:7). Wings with basal vein having five setae on right wing and four on left.
Variation.—Female mesosomal length varies between 0.76 and 0.96 mm. Body
color varies from bluish black, to greenish black, to dark green with coppery reflections like the holotype. A female from Ohio has all the femora with dark bands basally. The groove between the basal foveae is some- times weakly sculptured posteriorly, but there is always a smooth strip of at least one spiracular outside diameter along anterior margin of propodeum. There are occasional specimens with two strong sublateral cari- nae on one side or the other. Male meso- somal length varies between 0.90 and 0.98 mm. The body color variation is similar to that of females.
Diagnosis. —In addition to the characters given in the key, the vertex of female P. quadriplana is usually nearly concolorous with the dorsum of the mesosoma; in P. pelagocorypha, the vertex is distinctly paler. The antenna of female P. quadriplana has each funicular segment narrowed basally, the prominent MPP sensillae give the seg- ments a coarse texture, and the club varies between 1.4 and 2.0 times as long as wide. The antenna of female P. pelagocorypha has cylindrical funicular segments with a rather smooth texture, and the club varies between 2.0 and 2.6 times as long as wide. In male P. quadriplana, the combined length of the pedicel and flagellum is between 4.1 and 4.8 times as long as the club length; it is between 3.6 and 3.9 times as long as the club in P. pelagocorypha. The dorsellum of P. quad- riplana 1s usually smooth and tilted nearly perpendicularly with respect to the metano- ta. In P. pelagocorypha, the dorsellum is usually alveolate and in nearly the same plane as the metanota. The lateral setae on the petiole are shorter in P. quadriplana, only a half to three fourths the width of the petiole (Fig. 11); they are nearly as long as the petiole width in P. pelagocorypha (Fig. 12). The petiole always has a distinct and complete median carina in P. guadriplana (Fig. 11); in P. pelagocorypha, the median carina may be lacking or incomplete, or there may be multiple fine longitudinal rugae (Fig. 12).
VOLUME 90, NUMBER 1
Biology.—The host(s) is unknown; how- ever, the numerous specimens from Nova Scotia were collected during a study in which apple trees were fumigated and the arthro- pods on them were collected on sheets be- neath the trees (W. R. M. Mason, pers. comm.).
Etymology.—The name comes from the Latin words quadrus, meaning fourfold, and planus, meaning flat or level, and refers to the four smoothish areas along the anterior margin of the propodeum, which are diag- nostic of this species.
Type material.— Holotype female is from Mt. Ste. Marie Low, Quebec, and was col- lected 20 September 1965 by J. R. Vock- eroth (CNC). The allotype male is from Cooper’s Rock State Forest (near Morgan- town), West Virginia, and was collected 22 June 1964 by O. Peck (CNC). Fifty-three paratypes are as follows (CNC, INHS, USNM): Canada. BRITISH COLUMBIA: Cultus Lake, 14-VII-1948, 1 @. NEW BRUNSWICK: Kouchibouguac National Park, 12-IX-1977, 1 2. NOVA SCOTIA: Aldershot, 4-VII-1952, 2 8, 1 9, 15-VH- 1952, 1 6, 7 9, 28-VII-1952, 11 9, 8-VII- 1952, 4 2, 18-VII-1950, 4 2, 9-IX-1950, 1 2. ONTARIO: Innisville, 18-VIII-1963, 1 2; Simcoe, 19-VI-1939, 1 92. QUEBEC: Lac Brulle, 15-VII-1946, 1 3, Mt. Ste. Marie Low, 20-IX-1965, 3 2; Old Chelsea, 3-VII- 1969, 1 4, S-VIII-1969, 9 2, United States. NEW YORK: Lake Placid, 15-VIII-1896, 1 2; Otter Lake (near Meridian), 25-VII- 1946, | ¢. OHIO: Barberton, 30-VI-1936, 1 9. VIRGINIA: Monterey, 22-VI-1964, | 2.
Polstonia pelagocorypha, New SPECIES (Fig. 12) Description. — Holotype female: Color. Head with face, frons dark green; vertex blue-green; occiput greenish black. Antenna with scape brownish yellow; pedicel, flagel- lum dark reddish brown. Mandible yellow- ish brown; teeth reddish. Mesosoma with
dorsum dark green; pleural area, coxae, pro- podeum, petiole, gaster bluish black. Legs brownish yellow except trochanters, basal two thirds of fore and mid femora, hind femur brown (hind femur with traces of me- tallic coloring); pretarsus black. Wing with veins pale brown. Head and mesosoma with pale brown setae.
Sculpture.—Pattern similar to P. guad- riplana except texture delicate, particularly on head, and petiole strigulate dorsally, cells three or more times as long as wide.
Structure.—Mesosomal length 0.92 mm. Head width 1.2 x height (34:27), 2.3 x length (34:14.5); clypeus with anterior margin slightly produced and reflexed: eye height 1.2 length (15:12), 1.7 x malar length (15: APOE <x OOL.(7.5:5), 2.1 x OOL GS: 3.5). Antenna with length of pedicel plus flagellum 1.0 x head width (34.5:34); rela- tive lengths of antennal segments (annelli omitted, club counted as a unit) scape = 15: 5:3.512.5:3:2.523:2.5:10:5; widths of Fl, F6, club as 3:3.5:4; funicular segments cylin- drical; MPP sensillae fine, club length 2.6 x width (10.5:4), sutures only slightly oblique. Mesosoma 1.7 as long as wide (46:26.5); scutellar length 0.86 width (12:14); dor- sellum in same plane as metanota, anterior margin scalloped, finely alveolate: propo- deum with groove between basal foveae subareolate, with short weak sublateral ca- rinae (Fig. 12). Wing length 2.2 x width (89: 41); relative lengths of submarginal, mar- ginal, postmarginal, stigmal veins as 32:18.5: 17:10; basal vein marked by row of nine setae on left wing; one seta in basal cell of left wing. Petiole length 3.2 x width (16:5), 1.3 as long as propodeum (16:12); flat- tened dorsally; median carina visible only in posterior third: length of lateral setae nearly equal to width of petiole (Fig. 12). Gaster fusiform, length 1.5 x width (35:23); succeeding terga withdrawn beneath T2 (specimens air-dried).
Allotype male: Color. Similar to holotype but paler, dorsum of mesosoma green with faint yellowish reflections, antennal pedicel
10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and flagellum brown, bands on femora dark but extending only one third length of mid and three fourths length of hind femora. Mesosomal length 0.84 mm. Antenna with pedicel plus flagellum 0.98 x as long as head width (32.5:33); relative lengths of segments as 13:4.5:4:3.5:3.5:3.5:3.5:3.5:9; parallel- sided, widths of Fl, F6, club as 3:3:3. Pet- iole length 2.6 width (16.6). Wing with basal cell bare except for a couple of setae adjacent to setal row on basal vein on right wing.
Variation. — The female mesosomal length varies between 0.77 and 0.96 mm. Body color varies from bluish black to dark green. The female from North Carolina has the vertex green; in the other females it is blue- green like the holotype. The groove between the basal foveae of the propodeum is some- times crossed by one or more weak carinae, but these are less than one spiracular outside diameter in length. The petiole varies be- tween 2.6 and 3.0 times as long as wide, and its dorsal surface is either acarinate, with weak or incomplete median carina, or with several long longitudinal rugae. The male mesosomal length varies between 0.67 and 0.94 mm. The color of the males varies from green with coppery reflections to bluish black. The male from Illinois has a brown- ish yellow flagellum and very weak dark bands on the femora. The petiole varies be- tween 2.7 and 3.0 times as long as wide.
Diagnosis.—For a detailed diagnosis see that of P. quadriplana above.
Biology.—The host(s) of this species is unknown.
Etymology.—The name is from the Greek words pe/agos, meaning sea, and koryphe, meaning top of the head, and refers to the sea-green vertex of the female.
Type material.—Holotype female (AMNH) is from 1.5 miles SW of Lolo Hot Springs, Montana, and was collected 22 July 1978 by N. L. Herman. Allotype male (CNC) is from Whiteface Mountain, New York, and was collected 19 July 1962, by J. C. Chillcott. Thirteen additional paratypes are
as follows (CMNH, CNC, FSCA, INHS, USNM): Canada. ALBERTA: Edmonton, 20-VI-1937, 1 6. NEW BRUNSWICK: Fundy National Park, 10-VII-1970, 1 9°; Kouchibouguac National Park, 10-XI-1977, 1 2. QUEBEC: Duchensay, 5-VII-1953, 1 @. SASKATCHEWAN: White Fox, 18-VII- 1944, 1 2. United States. ALASKA: Mata- nuska, 6-X-1945, | 4; Palmer, 1-VIII-1948, 1 6. ILLINOIS: MacLean Co., 30-V-1883, 1 2; Urbana, 10-VI-1928, 1 6. MICHIGAN: Manistee Co., 5-VII-1957, 1 6. NORTH CAROLINA: Lake Junaluska, 27-V-1954, 1 ¢. WEST VIRGINIA: Spruce Knob, 5-VIII-1960, 1 4, Weston, 13-18-IX-1938, Id:
ACKNOWLEDGMENTS
I thank the following persons for the loan of material: M. Favreau, American Mu- seum of Natural History (AMNH), New York, NY; Dr. G. Wallace, Carnegie Mu- seum of Natural History (CMNH), Pitts- burgh, PA; Dr. C. M. Yoshimoto, Canadian National Collection (CNC), Ottawa, ON; Dr. H. Weems, Florida State Collection of Ar- thropods (FSCA), Gainesville, FL; Dr. G. W. Byers, Snow Entomological Collection (SEC), Lawrence, KS; and Dr. E. E. Grissell, United States National Museum (USNM), Washington, D.C. I would like to express my appreciation to J. K. Bousman and Dr. G. L. Godfrey of the Illinois Natural His- tory Survey (INHS), Champaign, IL, and two anonymous readers for reviewing this paper. I would like to thank my advisor, Dr. W. E. LaBerge (INHS) for help and en- couragement, J. Sherrod for assistance with the illustrations, and the staff at the Center of Electron Microscopy at the University of Illinois at Urbana-Champaign.
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. 1904. Classification of the Chalcid flies or
VOLUME 90, NUMBER 1
the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected by Herbert H. Smith. Mem. Carneg. Mus. 1: i-1x, 225-551.
Bakkendorf, O. 1955. Notes on Icelandic and Green- landic chalcidoideous Hymenoptera. Entomol. Medd. 27: 135-162.
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1976. African Pteromalidae (Hymenoptera): New taxa, synonymies and combinations. J. Ento- mol. Soc. South. Afr. 39: 9-31.
Brulle, M. A. 1846. Les Chalcidites, pp. 547-605. In A. Lepeleltier de Saint-Fargeau and M. Lake, eds., Histoire Naturelle des Insects. Hymenopteres, Part IV. Paris.
Burks, B. D. 1975. The species of Chalcidoidea de- scribed from North America north of Mexico by Francis Walker (Hymenoptera). Bull. Brit. Mus. (Nat. Hist.) Entomol. 32: 139-170.
Cresson, E. F. 1887. Synopsis of the families and genera of the Hymenoptera of America north of Mexico, together with a catalogue of the described species and bibliography. Trans. Amer. Entomol. Soc., Suppl. Vol.
Delucchi, V. 1955. Notes sur les Pteromalides. Mem. Soc. R. Belge Entomol. 27: 171-175.
Dzhanokmen, K. A. 1978. [Identification of the in- sects of the European part of the USSR. Vol. 3. Hymenoptera. Second Part. Pteromalidae.] Opre- deliteli Faune USSR. No. 120: 57-228.
Eschscholtz, J. F. 1829. Zoologischer Atlas, enthal- tend Abbildungen und Beschreibungen neuer Thierarten, wahrend des Flottcaptains von Kot- zuebue zweiter Reise um die Welt, auf der Rus- sisch-Kaiserlichen Kriegschlupp Predpriaetie in den Jahren 1823-1826, Tritte Heft. Berlin.
Forster, A. 1856. Hymenopterologische Studien. 2. Chalcididae und Proctotrupu. Aachen.
Farooqi, S. I. and B. R. Subba Rao. 1985. Family Pteromalidae, pp. 254-263. Jn B. R. Subba Rao and M. Hayat, eds., The Chalcidoidea (Insecta: Hymenoptera) of India and the Adjacent Coun- tries. Part I. Reviews of families and keys to fam- ilies and genera. Oriental Insects, 19: 163-310.
11
—. 1986. Family Pteromaliae, pp. 279-306. In B. R. Subba Rao and M. Hayat, eds., The Chal- cidoidea (Insecta: Hymenoptera) of India and the Adjacent Countries. Part II. A catalogue of Chal- cidoidea of India and the adjacent countries. Ori- ental Insects, 20: 1-430.
Gahan, A. B. and M. Fagan. 1923. The type species of the Chalcidoidea or Chalcid-flies. United States National Museum Bulletin, 124: 1-173.
Graham, M. W. R. de V. 1969. The Pteromalidae of northwestern Europe (Hymenoptera, Chalcidoi- dea). Bull. Brit. Mus. (Nat. Hist.) Entomol. Suppl. 16: 1-908.
Haliday, A. H. vi, pls. A-P.
Heydon, S. L. and W. E. LaBerge. A review of the North American species of Sphegigaster Spinola north of Mexico with a review of the biology. J. Kans. Entomol. Soc. (In press.)
Motschulsky, V. de. 1863. Essai d’un catalogue des insectes de l’Ile Ceylan. Byull. Mosk. Obshch. Is- pyt. Prir. 36(3): 1-153.
Nikol’skaya, M. N. 1952. [The Chalcid fauna of the USSR (Chalcidoidea).] English translation— 1963, Israel Program for Scientific Translations. Jeru- salem.
Peck, O., Z. Boucek, and A. Hoffer. 1964. Keys to the Chalcidoidea of Czechoslovakia. Mem. Ento- mol. Soc. Can. 34: 1-120.
Schmiedeknecht, O. 1909. Hymenoptera family Chalcididae. Jn P. Wytsman, Genera Insectorum 97: 1-550.
Thomson, C. G. 1878. Hymenoptera Scandinaviae. 5. Pteromalus (Svederus) continuatio. Lund. Walker, F. 1833. Monographia Chalcidum. Art.
XLIII. Entomol. Mag. 1: 367-384.
1872. Notes on Chalcididae. Part VI. Hor-
moceridae, Sphegigasteridae, Pteromalidae, Elas-
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. 1873. Economy of Chalcididae. Entomolo- gist 6: 322-324.
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1842. [No Title] Entomologist 1: v—
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 12-19
A NEW GENUS AND TWO NEW SPECIES OF PANGONIINI (DIPTERA: TABANIDAE) OF ZOOGEOGRAPHIC INTEREST FROM SABAH, MALAYSIA
JOHN F. BURGER
Department of Entomology, Nesmith Hall, University of New Hampshire, Durham,
New Hampshire 03824.
Abstract. —A new genus of Pangoniini, Mesopangonius Burger, 1s described from Mount Kinabalu, Sabah, Malaysia. This is the first representative of this tribe known to occur in the Oriental Region. Although possibly derived from unspecialized Laurasian Pango- niini, its long, slender proboscis is characteristic of specialized genera of the tribe. Two new species, philipi and brackleyae are described in the genus, and a key is provided.
A recent small collection of Tabanidae from Mount Kinabalu, Sabah, Malaysia, by entomologists from the Smithsonian Insti- tution, Washington, D.C., yielded two re- markable new species in an undescribed ge- nus of the tribe Pangoniini. These are the first representatives of this tribe known from the Oriental Region, and are particularly in- teresting because they combine anatomical features of both generalized and specialized genera of Pangonuini.
Mesopangonius Burger, New GENUS
Type species.—Mesopangonius philipi Burger, Sabah, MALAYSIA, by original designation.
Medium-sized (12-16 mm long) rather slender to moderately stout-bodied Esen- beckia-like species with well-developed ocelli; eyes bare with no color pattern; fron- tal index 2.5-3.0. Antennal bases closely ap- proximated; flagellum subulate, bearing 8 annuli; basal annulation enlarged (Figs. 2C, 4C), about twice as long as high; apical an- nulation greatly elongated, about one-half
Scientific Contribution Number 1485 from the New Hampshire Agricultural Experiment Station.
length of remaining annulations combined. Proboscis slender, length 1.5-1.9 times height of head; labella long, slender and sclerotized (Figs. 2A, 4A); 2nd maxillary palpomere short and subcylindrical or somewhat flattened on outer surface, and bearing a shallow concavity. Legs long and slender, hind tibial spurs well-developed. Base of vein R, with a long spur, basal sec- tion of Cu bare or with 2-3 scattered setulae, Sc bare dorsally and ventrally. Female gen- italia (Fig. 2E-G): ninth tergite entire, rel- atively broad and heavily sclerotized lat- erally, narrowed and weakly sclerotized medianly; tenth tergite divided medially; cerci rounded apically, length and width subequal; eighth sternite shield-shaped, very weakly sclerotized; apical lobes of the an- terior gonapophyses deeply divided medi- ally, about as long as wide, distance between lobes about two-thirds width of individual lobes; arms of the genital fork with wing- like expansions apically; distal ends of sper- mathecal ducts membranous, unexpanded.
Mesopangonius philipi Burger, NEw SPECIES
Female (Fig. 1).—Length: body 12-14 mm; wing 13-15 mm. Front (Fig. 2B) yel-
VOLUME 90, NUMBER 1
Fig. 1.
Mesopangonius philipi sp. n., female. (8 x)
lowish brown laterally, slightly diverging below, with broad dark brown pollinose ridge in middle extending from subcallus to vertex, bordered by an irregular row of semi- erect black setulae; frontal index 2.8-3.2, divergence index 1.2—1.3. Ocelli large and prominent, borne on a conspicuous tuber- cle; vertex depressed below upper margin of head. Subcallus, gena and face brown pol- linose, gena sparsely clothed with brown hairs; face moderately produced, upper lat- eral surface with patch of brown hairs; beard rather sparse, with brown hairs anteriorly, pale yellowish ones posteriorly. Antenna (Fig. 2C) yellow brown, slightly darker api- cally; bases closely approximated, distance between them distinctly less than width of scape; scape and pedicel yellowish brown pollinose, black setose; flagellum subulate, with 8 annuli; basal annulation enlarged, about one-third broader than succeeding
annulation and bearing a dense tuft of black setulae at apex of upper margin, apical an- nulation three times length of penultimate annulation. Proboscis slender (Fig. 2A), length 1.7-1.9 times head height; labella long and slender, sclerotized. Maxillary palpus (Fig. 2D) with apical palpomere short and slender, subcylindrical, length less than one- fifth that of proboscis, bearing long, black semi-erect setae on outer surface, basal pal- pomere slightly broader than apical seg- ment, bearing long, black semi-erect setae. Eye bare, unpatterned (relaxed), rather coarsely faceted.
Mesonotum light brown, bearing semi- erect brown hairs, except pale yellowish white ones anteriorly near the head; noto- pleural lobe concolorous with mesonotum; scutellum slightly paler; pleuron paler yel- lowish brown, bearing pale yellow hairs, ex- cept dark brown ones posteriorly on mesan-
14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Antenna. (32 x) D, Maxillary palpus. (32 = and anterior gonapophyses. (60 x) G, Genital fork and caudal ends of spermathecal ducts. (90 x)
episternum. Legs slender, elongate, unicolorous pale yellowish brown, bearing mixed pale yellowish and dark brown hairs; apical spurs on hind tibia nearly as long as those on mid-tibia. Wing lightly brown tint- ed throughout; R, with long spur; cells r; and m, open to wing margin. Halter light brown.
Mesopangontius philipi sp. n., female. A, Profile of head. (16) B, Frontal view of head. (16x) C, ) E, Eighth, 9th tergites, cerci, dorsal view. (60 =) F, Eighth sternum
Abdomen pale greenish brown, with some yellowish tones intermixed but without def- inite pattern; tergite | slightly paler; tergites 5-7 slightly darker; all tergites bearing pre- dominantly dark hairs, with some pale yel- lowish ones intermixed anteriorly and lat- erally; ventral surface concolorous.
Holotype 2, MALAYSIA: Sabah; Kina-
VOLUME 90, NUMBER 1
Fig. 3.
balu National Park, Headquarters area, el. 1560 m, 9 Sept. 1983, G. F. Hevel & W. E. Steiner (National Museum of Natural His- tory, Washington, D.C. (NMNH)).
Paratypes, MALAYSIA: 2 8, 8, 13 Sept. 1983. Same data as holotype (NMNH; J. F. Burger Collection).
I take great pleasure in naming this species for the late Cornelius Becker Philip, inde- fatigable student of Tabanidae, who con- tributed much to our knowledge of Oriental Tabanidae.
Mesopangonius brackleyae sp. n., female. (8 x)
Mesopangonius brackleyae Burger, New SPECIES
Holotype female.—Length: body 15.6 mm; wing 16 mm (Fig. 3). Front (Fig. 4B) dark brown pollinose, slightly diverging be- low, middle with a poorly-defined raised ridge from subcallus to vertex, bearing an irregular median subshining black area and bordered by an irregular row of black set- ulae; frontal index 2.5, index of divergence 1.2. Ocelli large and prominent, borne on a
16
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4.
conspicuous tubercle at vertex. Subcallus concolorous with front. Gena and face gray pollinose, except face shining dark brown along lower margin; face moderately pro- duced, dorsolateral surfaces with patch of dark brown hairs; beard mostly pale yellow- ish, except dark brown hairs anteriorly. Scape and pedicel of antenna yellowish gray pollinose, antennal bases closely approxi- mated, distance between them distinctly less than width of scape; flagellum subulate (Fig. 4C), with 8 annuli, yellowish brown, apical annulation dusky brown; basal annulation conspicuously enlarged, twice as broad as second annulation, bearing 1-2 setulae on upper surface; apical annulation 2.5 times
Mesopangonius brackleyae sp. n., female. A, C, Antenna. (32x) D, Maxillary palpus. (32 =)
Profile of head. (16) B, Frontal view of head. (16x)
length of penultimate annulation. Proboscis slender (Fig. 4A), length 1.5 times head height, labella long and slender, sclerotized. Maxillary palpus (Fig. 4D) short, apical pal- pomere brown, distinctly flattened, length one-fourth that of proboscis, bearing long black setae on outer surface and narrow bare median concavity. Eye bare, unpatterned (relaxed), relatively coarsely faceted. Mesonotum and scutellum subshining dark brown, densely clothed with semi-erect yellowish hairs; postpronotal lobe reddish gray pollinose; notopleural lobe reddish; pleuron grayish pollinose, except mesanepi- sternum and anterior half of katepisternum with blackish tones, densely yellow pilose.
VOLUME 90, NUMBER |
Legs slender, elongate; coxa and femur dark brown, black pilose, except apex of femur paler; tibia pale brown, bearing yellow hairs; hind tibial spurs well-developed, subequal to mid-tibial spurs; tarsus basally concol- orous with tibiae, darker brown apically. Wing light brownish tinted on anterior half, subhyaline posteriorly; R, with long spur; cells r; and m, open to wing margin. Halter pale brown.
Tergite 1 of abdomen entirely pale yel- lowish brown, pale yellow pilose: tergites 2— 4 dark brown on basal three-fourths, con- trastingly paler brown on apical fourth, the dark and light areas bearing black and pale yellow hairs respectively; tergites 5-7 dark brown, black-haired, with conspicuous yel- low-haired posterior margins (Fig. 3); ven- tral surface of abdomen with similar pattern as dorsum, except sternite 2 predominantly light brown, and only basal halves of ster- nites 3-4 dark brown.
Holotype °, MALAYSIA: Sabah; Kina- balu National Park, Headquarters area, el. 1560 m, 9 Sept. 1983, G. F. Hevel & W. E. Steiner (National Museum of Natural His- tory, Washington, D.C.).
I take pleasure in naming this striking species for my good friend and colleague, Frances Brackley, a specialist of the Orchi- daceae, who first sparked my interest in dip- terous pollinators of alpine plants.
The following key will separate the species of Mesopangonius described above:
1. Abdominal tergites 2-5 dark brown, with paler brown apices and complete yellow-haired in- cisures; all femora dark brown, contrasting with light brown tibiae; apical palpomere of max- illary palpus somewhat flattened, outer surface with a bare, shallow concavity brackleyae, n. sp.
— Abdominal tergites light brown throughout, without strongly contrasting markings or hairs; all femora and tibiae concolorous light brown; apical palpomere of maxillary palpus narrow and subcylindrical, lacking a bare concavity on the outer surface .. philipi, n. sp.
DISCUSSION
Mesopangonius resembles Esenbeckia Rondani, a predominantly Neotropical ge- nus, but differs in having the basal annu-
17
lations of the flagellum not forming a par- tially-fused, enlarged plate, a more slender proboscis with very narrow, elongate label- la, and with cell r; of the wing widely open to the wing margin. The female terminalia also are similar to Esenbeckia, differing pri- marily in the broader ninth tergite (Fig. 2E), the more rounded cercus, and the more widely separated apical lobes of the anterior gonapophyses (Fig. 2F).
Mackerras (1955) subdivided the genera of Pangoniini into generalized (Group |) and specialized (Group 2) moieties. Those gen- era considered to be more generalized have the r; cell of the wing open, proboscis stout and subequal to head height, the labella dis- tinctly enlarged and unsclerotized, and the body usually slender or with the abdomen parallel-sided. Most of the genera included have a south or north temperate relict dis- tribution in montane or desert environ- ments. Fourteen of 18 genera in Mackerras’ Group | occur in coastal or desert North America (5), the mountains of Chile and Argentina (5), and in Australia (4). Two gen- era occur in Japan, and one each in Brazil and southern Africa.
Genera of Pangoniini considered to be specialized have cell r; closed, or strongly narrowed apically, proboscis slender, as long as to much longer than head height, labella narrow, sclerotized, and sometimes very long, and the body usually stoutly-built. These genera have a predominantly south- ern Palearctic, amphi-Mediterranean (Pan- gonius), or a new world tropical and sub- tropical (Esenbeckia) distribution. Mack- erras considered Austroplex Mackerras, from Australia, to be a link between the two groups because of its basally expanded an- tennal flagellum, but most of its features can be considered generalized.
Mesopangonius also has a preponderance of primitive features, eyes bare and coarsely faceted, cell r; of the wing open, relatively slender body (somewhat stouter in brack- leyae), basal flagellar annulations not con- solidated into an enlarged plate, relatively narrow, unspecialized maxillary palpus, the
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
long, slender legs, and the broad and un- divided ninth tergite. The principal spe- cialized features are the long, slender pro- boscis, the long, narrow, sclerotized labella, and the deeply divided and relatively widely- separated apical lobes of the anterior gon- apophyses. The preponderance of primitive features suggests that Mesopangonius 1s closer to the generalized group of genera, and that the elongate, narrow proboscis and labella may be an adaptation to a particular trophic niche. Mesopangonius also occurs in a montane “temperate” area on Mount Kinabalu at the northern end of the Crocker Range in Sabah, an environment similar to that where some genera of Pangoniini in Mackerras’ Group | occur.
Since Borneo is a continental island as- sociated with the Laurasian plate, Meso- pangonius may be derived from a general- ized Laurasian pangoniine stock. Genera of Pangoniini presently known from Eurasia, other than Mesopangonius, are Stonemyia Brennan (Japan, Southwestern Asia [Cau- casus], and possibly China), Nagatomyia Murdoch & Takahasi (Japan), and Pan- gonius Latreille (amphi-Mediterranean). Mesopangonius differs most conspicuously from Sfonemyia in having a more slender body, a longer, more slender antennal fla- gellum, with only the basal annulation en- larged and a much longer apical annulation, a longer, more slender proboscis with a long, narrow, sclerotized labella, shorter palpus, legs longer and more slender, ventral surface of scutellum without bristles, R, of the wing with a long spur, the larger, deeply-divided lobes of the anterior gonapophyses, and the caudal ends of the spermathecal ducts mem- branous and delicate (Fig. 2G). It shares few features with Nagatomyia other than the presence of ocelli, the slender body and the open cell r;. Mesopangonius differs from Pangonius in having a more slender body, ocelli larger and more prominent, basal an- nulation of flagellum broader, apical pal- pomere shorter relative to proboscis length, and with the outer concavity, when present,
very shallow, the legs more slender and elongate, cell r; of the wing open, and the lobes of the anterior gonapophyses larger and more deeply-divided medianly.
Mesopangonius bears little resemblance to generalized Pangoniini associated with the Australian plate, as one might expect from tectonic evidence. Ectenopsis Mac- quart has coarse eye facets and a narrow cylindrical apical palpomere, but otherwise has little in common with Mesopangonius. Some species of Fidena Walker, a genus with specialized features in the Scionini that has radiated extensively in the Neotropical Re- gion, have a proboscis configuration like that of Mesopangonius, but otherwise are quite distinct.
The only other representatives of the Pan- goniinae known from the Oriental Region are in the Philolichini (Philoliche Wiede- mann). Of these, only species in the sub- genus Buplex Austen of Philoliche share even a superficial resemblance to Mesopango- nius, and then only because they have ocell, and some species have a narrow, elongate proboscis. However, they are presently re- stricted to southern Africa. The Oriental species of Philoliche lack ocelli, have strong- ly produced faces, closed wing cells, lobes of the anterior gonapophyses more widely separated, and other features that clearly ex- clude them from close relationship with Mesopangonius.
It is remarkable that Mesopangonius re- mained uncollected and unknown for so long. Its chance discovery suggests that oth- er representatives of the Pangoniini possibly may be found in remaining “temperate is- lands” within tropical Asia, given sufficient patience and collecting, before such refuges disappear.
ACKNOWLEDGMENTS
I thank G. B. Fairchild, Bureau of Ento- mology, Florida Department of Agriculture, L. L. Pechuman, Cornell University, and H. J. Teskey, Biosystematics Research Centre, Ottawa, for their helpful comments
VOLUME 90, NUMBER | is)
about characters of the Pangoniini. I also LITERATURE CITED
thank Dz S. Chandler and Scott Sherman, Mackerras, I. M. 1955. The classification and distri- University of New Hampshire, for review- bution of Tabanidae (Diptera) II. History: Mor- ing the manuscript, and Tess Feltes, Ports- phology: Classification: subfamily Pangoniinae. mouth, New Hampshire, for preparing the Australian Journal of Zoology 3: 439-511.
illustrations.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 20-27
APTERONA HELIX (LEPIDOPTERA: PSYCHIDAE), A PALEARCTIC BAGWORM MOTH IN NORTH AMERICA: NEW DISTRIBUTION RECORDS, SEASONAL HISTORY, AND HOST PLANTS
A. G. WHEELER, JR. AND E. RICHARD HOEBEKE
(AGW) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania 17110; (ERH) Department of Entomology, Cornell University, Ithaca, New York 14853.
Abstract.—New distribution records for Apterona helix (Siebold), an Old World bag- worm moth detected in California in 1940 and in New York in 1962, are given for Michigan and Pennsylvania, and additional New York records are provided; populations in western and eastern North America are thought to represent separate introductions from Europe. Seasonal history, habits, and host plants of this parthenogenetic, polyphagous species are reported for populations observed in New York and Pennsylvania, and North American distribution and economic importance in Europe and North America are reviewed. Char- acters facilitating recognition of this psychid are provided, and the unusual helicoid or
snail-like larval case is illustrated.
Members of the psychid genus Apterona Milliére are endemic to the Palearctic re- gion, with 7 species occurring in the Med- iterranean area, west and central Europe, Asia Minor, Crimea, Caucasus, Iranian Pla- teau, and southcentral Asia (Kozhanchikov 1956). Apterona helix (Siebold), a parthe- nogenetic species, is widely distributed in central and southern Europe, ranging east to European USSR and Kirgiz SSR in cen- tral Asia and south to Iran (Strand 1912, Davis 1964).
The first confirmed North American re- cord of 4. helix was based on an infestation discovered at a private residence in Nevada City, California, in June 1940 (Kiefer 1940, Robinson 1953). In the western states, A. helix now occurs in the northern half of California (Eichlin 1985) and in portions of Idaho, Nevada, Utah (Davis 1964), Oregon (Every 1970), and Washington (Suomi 1986).
Apterona helix is also known in eastern North America, having been detected near Albany, New York (Loudonville), in June 1962 (Davis 1964); a second infestation was soon discovered in Albany. Eastern popu- lations are believed to be the result of an independent European introduction rather than to have originated from the established western U.S. populations (Davis 1964). Lenox, Massachusetts (Adamski 1984), is the only other eastern record.
The name 4. crenulella (Bruand) has been used in early (and some current) literature on this adventive species in North America. Some European workers considered helix to be a parthenogenetic form of crenulella, whereas others argued that they are distinct species. In revising the Western Hemi- sphere Psychidae, Davis (1964) retained he- lix asa “‘facultative, parthenogenetic form,” noting interfertility between crenulella and helix might be expected but that “many
VOLUME 90, NUMBER 1
questions remain unanswered.” Apterona helix is now accorded specific rank (Davis 1983, 1987).
Here we review its status as an econom- ically important species, give additional lo- calities for 4. helix in New York, and report Michigan and Pennsylvania as new state records. We summarize our observations on seasonal history, habits, and host plants in the East and give morphological characters allowing this immigrant species to be rec- ognized in the North American fauna.
ECONOMIC IMPORTANCE
The European literature indicates that 4. helix occasionally is injurious. It has been implicated in causing damage to apple, hor- ticultural crops, and olive (Rev. Appl. Ento- mol. (A) 3: 393, 1915; 4: 210, 1916; 56: 553, 1968).
In Utah, 4. helix has been observed skele- tonizing leaves of apple trees and causing extensive damage to many range plants; it sometimes injures various cultivated plants and becomes a nuisance when it congregates on the walls and windows of homes (Tib- betts and Knowlton 1952, Knowlton and Roberts 1968). There also are records of severe damage to cherry foliage in an or- chard (Knowlton 1961), leafmining injury to corn (Knowlton and Parrish 1965), and destruction of green color in three acres of barley and four acres of alfalfa (Knowlton 1966). In Idaho, 4. helix was extremely abundant in alfalfa and sweetclover, causing considerable skeletonizing of the foliage (Gittins 1958). Marshall (1970) recorded heavy damage to strawberry plants in Ne- vada. In California, where this insect has been called the garden bagworm, consid- erable damage to several commercial crops such as apple, cruciferous vegetables, and chrysanthemums and other plants grown for cut flowers has occurred when populations are high (Keifer 1947, Robinson 1953). Suomi (1986) reported that baby’s breath (Gypsophila, Caryophyllaceae) used in dried flower arrangements was so heavily infested
21
in one Washington county that plant ma- terial could not be shipped out of state.
There are no reports of damage by 4. helix to cultivated plants in eastern United States, although Adamski (1984) stated that cases were found attached to planted flowers and vegetables, ornamentals, and shade trees. In New York, the cases have attracted notice when they attach to houses (Davis 1964). Each year homeowners submit larval cases to Cornell University’s Insect and Plant Disease Diagnostic Laboratory; their con- cern is with large number of cases that ac- cumulate on houses and the paint that is sometimes removed when cases are pulled off (Klass 1983).
DISTRIBUTION IN EASTERN NorTH AMERICA
In addition to published records from the Albany, New York, area and Lenox, Mas- sachusetts, the following new records are available. Michigan, Pennsylvania, and some of the New York records are based on our collecting; other New York records (those without collector names and mostly lacking exact localities) were obtained from the Insect and Plant Disease Diagnostic Laboratory, Cornell University. Voucher specimens have been deposited in the insect collections of Cornell University and the Pennsylvania Department of Agriculture.
MICHIGAN: Kent Co., Wyoming, 5 May 1986, E. R. Hoebeke; Grand Rapids, 6 May 1986, ERH.
NEW YORK: Albany Co., nr. Colonie, Pine Bush, 30 June 1984, ERH and A. G. Wheeler, Jr. Broome Co., Binghamton, 23 July 1983, ERH and AGW. Chemung Co., Rt. 17 N. of Wellsburg, 27 June and | Au- gust 1982, AGW. Clinton Co., August 1983. Columbia Co., June 1985. Dutchess Co., Stanfordville, October 1984. Erie Co., Ton- awanda, 12 June 1983 and 22 June 1985, ERH. Essex Co., May, November 1977. Livingston Co., Dansville, June 1980. Greene Co., East Windham, June 1984. Monroe Co., Greece, 31 July 1982, ERH
22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and AGW. Onondaga Co., Solvay, 26 June and 4 September 1982, 14 May 1983, ERH and AGW. Ontario Co., Canandaigua, April 1979. Rensselaer Co., Troy, June 1973. Schenectady Co., August 1973. Tompkins Co., Ithaca, 20 April 1987, ERH. Ulster Co., May and September 1982. Wayne Co., Clyde, 25 June 1983, ERH and AGW.
PENNSYLVANIA: Erie Co., Erie, 11 July 1985, AGW. Lackawanna Co., Carbondale, 28 June 1985, AGW. Lebanon Co., I-81 at junc. I-78 NW of Jonestown, 20 August 1982 and March—August 1983, AGW. Mercer Co., Sharon, 29 July 1987, AGW. Susquehanna Co., Thompson, 28 June 1985, AGW.
Our collections of A. helix in Broome, Chemung, Erie, Monroe, Onondaga, Tomp- kins, and Wayne counties in New York; Erie, Lackawanna, Mercer, and Susquehanna counties in Pennsylvania; and Kent County in Michigan were made along or near rail- road right-of-ways. In fact, populations were nearly always found only in a small area adjacent to active or abandoned trunk lines. In Lebanon Co., Pennsylvania, A. helix ap- parently is restricted to less than a mile of road near the junction of interstate high- ways 78 and 81.
It seems reasonable to assume that spread of this flightless, parthenogenetic moth in North America is largely dependent on commerce, especially rail traffic. Mature larvae are known to leave low-growing hosts and to attach to a suitable, usually higher, substrate for pupation (Robinson 1953). A larva could easily attach its case to a rail car in storage and the pupa, female, eggs, or overwintering larvae could be transported many miles to initiate a new colony. In one California orchard a new infestation of J. helix was attributed to its introduction as cases attached to a “‘small private spray rig” that had been used in the infested area (Ar- mitage 1953).
SEASONAL HISTORY AND HABITS
To obtain information on phenology of this psychid in eastern North America, col-
lections were made in 1983 from a roadside planting and from guardrails at the junction of interstate highways 78 and 81 in Lebanon Co., Pa. Observations were made and cases collected on 30 March; 1, 13, 22, and 27 April; 5, 11, and 18 May; 2 and 21 June; 15 July; and 3 August. Populations at the site were not as large as the hundreds of old cases adhering to guardrails might suggest, and on several sample dates the bagworms were not easily found on vegetation; cases, which “resist weathering to an amazing de- gree” (Robinson 1953), probably persist for years in protected places.
Life stages present in the population were approximated by dissecting small numbers of cases (usually only 5-10) on each sample date and, for larvae, measuring widths of head capsules. Such measurements suggest- ed four larval instars: I, 0.28—0.30 mm wide (n = 40); II, 0.36-0.40 mm (n = 16); III, 0.48-0.54 mm (n = 10); and IV, 0.64—0.80 mm (n = 9).
As Robinson (1953) reported for A. helix in California, young larvae overwinter with- in the female’s pupal skin. Larvae appar- ently construct septa within this empty shell so that each is enclosed in its own cell (Davis 1964). In Pennsylvania during late March— early April, 14 occupied cases that were col- lected on guardrails and examined in the laboratory contained first instars. As many as 42 and 49 larvae emerged from a single case. Several cases were coated with clover mites, Bryobia praetiosa Koch, and their egg shells. When cases were brought into the laboratory, larvae soon emerged, began to construct their own cases from sand and soil of old cases, and to feed on foliage, the dam- age appearing as tiny, circular transparent areas (Fig. 1).
On 13 April, first instars, about | mm long and without cases, were active on guardrails, with a few larvae in small cases observed on nearby spotted knapweed, Centaurea maculosa Lam. (Asteraceae). These cases consisted merely of a few sand grains on the body. Some knapweed leaves
VOLUME 90, NUMBER |
Figs. 1-3. cases) feeding on black medic, Medicago lupulina, causing tiny, circular transparent areas. 2, Feeding injury by mature larvae to leaves of common mullein, Verbascum thapsus. 3, Feeding injury by mature larvae to black medic.
showed slight feeding symptoms similar to those observed in the laboratory. First in- stars were the only stage found in cases tak- en on 22 and 27 April (n = 7, 10) (Fig. 6). Of 5 larvae collected on 5 May, 3 were first and 2 were second instars; by 11 May only one first instar was present in a collection that contained 8 second instars; and only second instars were present on 18 May (n = 5). Cases observed during May were larger, consisting of a white silken sac impreg- nated with grains of sand and soil (Fig. 4a). Robinson (1953) described these cases as having the form of an inverted J or U. In May damage on knapweed foliage became more obvious.
Feeding damage to foliage by larval stages of Apterona helix. 1, Early-stage larvae (with small
Larvae feed mainly at night (Davis 1964). A larva feeds by using silk to fasten its case to a leaf and emerging through an opening near the bottom of the case (see Fig. 4b). It chews a hole in the adaxial or abaxial sur- face, inserts its head in the opening, and scrapes out tissue between the leaf surfaces, making a nearly oval mine. Davis (1964) noted that this injury closely resembles that made by lepidopteran larvae of the genus Coleophora (Coleophoridae). Two, three, or more windowlike areas were observed on some small leaves (Figs. 2, 3). According to Robinson (1953), fecal material is expelled through a lateral aperture in the upper or smallest whorl of the case.
24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
-)
Tas
a
Figs. 4-5. Larval cases of Apterona helix. 4a, Case of early-stage larva, consisting of small silken case impregnated with grains of sand and soil. 4b, Mature larval case, usually of 2! to 32 whorls; arrow indicates large basal opening through which the larva emerges to feed; scale line = 5 mm. 5, Aggregation of larval cases
on trunk of tree sapling.
Ten cases collected on 2 June contained third instars; all nine taken on 21 June yield- ed fourth instars (Fig. 6). Mature larvae oc- cupy the lower whorl (Robinson 1953, Da- vis 1964) of the helicoid or snail-like cases (Fig. 4b). On 15 July few active larvae were observed on knapweed, but closed cases were apparent and clumped on small trees (Fig.
5); several cases opened in the laboratory were found to contain pupae. Mature larvae also ascended guardrails for pupation, where current-season cases could not be distin- guished in the field from those of previous seasons. At other localities utility poles were used as pupation sites, with large numbers of old cases occurring in cracks and crevices.
VOLUME 90, NUMBER |
[Zan [Fee [WAR[ APR MAY | JUN | JUL | AUG] SEP | ocr | NOV] DEC]
GUEHWItTERING 1ST ST INSTARS NN J 1ST INSTARS SSS ~
.
2ND INSTARS
3RD INSTARS bie
4TH INSTARS
Fig. 6. Ist instars are overwintering inside the female case.)
Females were found in several cases col- lected on 3 August. Robinson (1953) re- ported that yellowish-white eggs (about 25/ case) were laid in the pupal skin that re- mains in the lower whorl and hatch in about 3 weeks; the diapausing larvae remain in- side the case through the winter.
Seasonal history of the Pennsylvania pop- ulation studied was nearly the same as that reported for 4. Ae/ix in northern California (Robinson 1953). Overwintered first-instar larvae became active in mid-April and fed until early to mid-July, with females ap- pearing to oviposit in August.
Host PLANTS
Spotted knapweed, the most frequently infested host at the study site in Lebanon Co., Pennsylvania, was the most common host plant observed along railroad lines in New York and Pennsylvania. White and yellow sweetclover, Melilotus alba Medik. and M. officinalis Lam. (Fabaceae), also were infested. The large population of A. helix at Solvay, New York, severely damaged these
1ST INSTARS
No" OVERWINTERING 1ST INSTARS NS
Inferred seasonal history of Apterona helix in Pennsylvania. (Note: stippling indicates period when
plants and several unidentified hosts. Species that showed occasional injury or moderate to heavy damage at only one or a few sites included green foxtail, Setaria viridis (L.) Beauv. (Poaceae); mugwort, Artemisia vul- garis L. (Asteraceae); black mustard, Bras- sica nigra (L.) Koch, and peppergrass, Lepidium virginicum L. (Brassicaceae); blueweed, Echium vulgare L. (Boraginace- ae); common mullein, Verbascum thapsus L., and yellow toadflax, Linaria vulgaris Mill. (Scrophulariaceae); common evening- primrose, Oenothera biennis L. (Onagra- ceae); cinquefoil, Potentilla sp. (Rosaceae); buckhorn plantain, Plantago lanceolata L., and broadleaf plantain, P. major L. (Plan- taginaceae); knotweed, Po/ygonum sp. (Po- lygonaceae); and crownvetch, Coronilla varia L., black medic, Medicago lupulina L., and alsike clover, Trifolium hybridum L. (Fabaceae).
Crownvetch was the dominant plant species at the Pennsylvania study site, but only slight feeding on leaflets of a few plants was observed. Along the railroad in New
26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
York common plants that generally were avoided were hedge bindweed, Calystegia sepium (L.) R. Br. (Convolvulaceae); lambs- quarters, Chenopodium album L. (Cheno- podiaceae); chicory, Cichorium intybus L., and horseweed, Conyza canadensis (L.) Cronq. (Asteraceae); and Queen Anne’s-lace, Daucus carota L. subsp. carota (Apiaceae). Apterona helix, although a polyphagous insect known from various wild and culti- vated plants (Robinson 1953, Davis 1964), fed mainly on low-growing herbs in New York and Pennsylvania. In California, how- ever, Robinson (1953) noted that “‘foliage of shrubs and trees were freely attacked later in the season.”’ We also observed some feed- ing on shrubs and trees at the study site and along railroad tracks, but these plants served primarily as sites for pupation (Fig. 5).
RECOGNITION FEATURES
The most characteristic morphological attribute of this small psychid moth is the larval case (Figs. 4a, 4b); it alone will enable recognition of this introduced bagworm moth in North America. The small spiraled case measures approximately 3—5 mm in diameter and 4-5 mm in depth. The mature larval case typically has 2’ to 3% whorls and is constructed of silk overlain with mi- nute earthen particles. There are three open- ings present in the case: a small apical open- ing, a large basal opening through which the larva is able to crawl and feed, and a large slitlike opening in the uppermost whorl through which the shrivelled female re- portedly emerges after oviposition (Davis 1964).
Males of A. helix are not known. The par- thenogenetic females are highly specialized, larviform, wingless or with reduced wings, with a small head with or without small antennal rudiments, with small pigment spots instead of eyes, and with leg rudiments without claws (Kozhanchikov 1956). The females never leave the spiral case.
ACKNOWLEDGMENTS
We thank Carolyn Klass (Cornell Uni- versity) for allowing us to use records of A. helix submitted to the Insect and Plant Dis- ease Diagnostic Laboratory, Gary Miller (Auburn University) for illustrating the lar- val cases in Figs. 4a and 4b, Alan Snook (Pennsylvania Dept. of Agriculture) for dis- secting cases and measuring head capsule widths of larvae, Maureen E. Carter (CU) for assistance in the field, James Stimmel (PDA) for helping make field collections and for reproducing Figs. 1-3 and 5, and Karl Valley (PDA) for reviewing an early draft of the manuscript.
LITERATURE CITED
Adamski, D. 1984. An unusual oviposition site for Amphipyra tragopogonis (L.) (Noctuidae). J. Lep- id. Soc. 38: 317-318.
Armitage, H. M. 1953. Current insect notes. Calif. Dep. Agric. Bull. 42: 39-41.
Davis, D. R. 1964. Bagworm moths of the Western Hemisphere (Lepidoptera: Psychidae). U.S. Natl. Mus. Bull. 244. 233 pp.
1983. Psychidae (Tineoidea), pp. 7-8. Jn R.
W. Hodges et al., eds., Check List of the Lepi-
doptera of America North of Mexico. E. W. Clas-
sey Ltd. & Wedge Entomol. Res. Found.
1987. Psychidae (Tineoidea), pp. 366-369. In F. W. Stehr, ed., Immature Insects. Kendall/ Hunt Publ. Co., Dubuque, Iowa.
Eichlin, T. 1985. Imported bagworm, Apterona cren- ulella. Calif. Plant Pest Dis. Rep. 4(4): 115. Every, R. W. 1970. A psychid moth (4pterona cren- ulella). U.S. Dep. Agric. Coop. Econ. Insect Rep.
20(40): 700.
Gittins, A. R. 1958. A psychid moth (Apterona cren- ulella). U.S. Dep. Agric. Coop. Econ. Insect Rep. 8(28): 607.
Keifer, H. H. 1940. Systematic entomology, pp. 241- 245. In D. B. Mackie, Chief. Division of Plant Industry, Bureau of Entomology and Plant Quar- antine. Calif. Dep. Agric. Bull. 29.
1947. Systematic entomology, pp. 168-173. In H. M. Armitage, Chief. Division of Plant In- dustry, Bureau of Entomology and Plant Quar- antine. 28th Ann. Rep. Per. Ending Dec. 31, 1947. Calif. Dep. Agric. Bull. 36.
Klass, C. 1983. Comments from diagnosticians. Entomology, pp. 3-4. /n Insect and Plant Disease
VOLUME 90, NUMBER |
Diagnostic Laboratory Annual Report, 1983. Coop. Ext. Cornell Univ., Ithaca, N.Y.
Knowlton, G. F. 1961. A psychid (Apterona crenu- lella). U.S. Dep. Agric. Coop. Econ. Insect Rep. 11(27): 598.
1966. A psychid (Apterona crenulella). U.S. Dep. Agric. Coop. Econ. Insect Rep. 16(22): 479, 482.
Knowlton, G. F. and D. S. Parrish. 1965. A psychid (Apterona crenulella). U.S. Dep. Agric. Coop. Econ. Insect Rep. 15(28): 744.
Knowlton, G. F. and R. S. Roberts. 1968. Range entomology, a subject of increasing importance. Utah State Univ. Entomol. Mimeo Ser. 133. 7 pp.
Kozhanchikov. 1956. Fauna of the U.S.S.R., Lepi- doptera, Vol. III, No. 2 (Psychidae). Zool. Inst. Akad. Nauk SSSR (n.s.), No. 62 (Translated from
27
Russian, 1969, Israel Program for Scientific Trans- lations, Jerusalem. 525 pp.).
Marshall. 1970. A psychid moth (Aplerona crenulel- la). U.S. Dep. Agric. Coop. Econ. Insect Rep. 20(25): 407.
Robinson, D. 1953. Garden bagworm, Apferona crenulella (-helix) in Nevada and Placer counties, California. Calif. Dep. Agric. Bull. 42: 25-33.
Strand, E. 1912. Psychidae, pp. 353-370. Jn A. Seitz, Gross-Schmetterlinge der Erde, Fauna palaearct. Bd. 2. A. Kernen, Stuttgart.
Suomi, D. 1986. Have you seen this bug?, pp. 4-5. In Co. Agent Entomol. Newsl. No. 2. Wash. State Univ., Pullman.
Tibbetts, T. and G. F. Knowlton. 1952. Apterona crenulella in Utah (Lep.: Psychidae). Entomol. News 63: 211.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 28-36
A REVISION OF THE GENUS MESTOCHARIS AND A REVIEW OF THE GENUS GRAHAMIA (HYMENOPTERA, EULOPHIDAE)
CHRISTER HANSSON
Department of Systematic Zoology, Lund University, Helgonavagen 3, S-223 62 Lund,
Sweden.
Abstract.—The Nearctic and Palearctic species of the genus Mestocharis Forster are revised. Three species are recognized, one exclusively Nearctic (M. tropicalis), one exclu- sively Palearctic (M. maculata) and one found in both regions (M/Z. bimacularis). Mesto- charis nearctica Yoshimoto is a new junior synonym of M. bimacularis. Hosts are known only for MM. bimacularis, a gregarious endoparasitoid in eggs of larger Dytiscidae (Co- leoptera). The two known species of Grahamia are reviewed and characters are presented to facilitate their separation. Both species are recorded as new to the Nearctic Region. The biology is known only for G. clinius, a parthenogenetic endoparasite of Haplodiplosis equestris (Diptera, Cecidomyiidae) in Europe.
Mestocharis has been treated in two rath- er recent papers, one dealing with the Eu- ropean (Boucek et al. 1963) and the other with the North American species (Yoshi- moto 1976). Boucek et al. recognized two species (M. bimacularis (Dalman) and ™. maculata (Forster)) and established two new combinations and three new synonyms. They dealt also with the intraspecific vari- ation of taxonomically important charac- ters and presented a key. Yoshimoto de- scribed two new species (V/. nearctica and M. tropicalis), compared them to the Eu- ropean species, and presented a key to the Nearctic species. The material I have at my disposal indicates that the intraspecific vari- ation described in those papers was under- estimated. The consequences of this under- estimation are that some characters used for separating the species are unsafe and that one of the species is invalid. The biology is known only for M. bimacularis (Jackson 1958, 1960, 1964), an endoparasite in eggs of larger Dytiscidae (Coleoptera). Two other Nearctic species were described as Mesto-
charis (M. wilderi Howard, M. williamsoni Girault), but both have been transferred to Pediobius Walker (Burks 1958: 68-69).
The genus Grahamia was erected to in- clude Entedon clinius Walker and Graham- ia tatrica Erdés (Erdés 1966). Later, Boucek and Askew (1968) catalogued the Palearctic Eulophidae and added new distribution rec- ords and corrected some previous host rec- ords for the genus. Hansson (1985) specu- lated that the two species might be the same because of high intraspecific variation in the length of flagellar segments, the only char- acter separating the two species, but made no definite decision regarding the validity of the two species. Because I have had access to a fairly large sample of these species from Europe and from North America, I con- clude that the two species are valid. A key is presented to facilitate their identification. The host is known only for G. clinius, a parthenogenetic endoparasite of Haplodi- plosis equestris (Diptera, Cecidomyiidae) 1n Europe.
Morphological terms used are explained
VOLUME 90, NUMBER 1
in Hansson (1985), the exception being POO, the distance between posterior edge of hind ocelli and occipital margin. Abbre- viations of museums and private collections used in the text were as follows: BMNH = British Museum (Natural History), London, England; CH = collection of the author; CNC = Canadian National Collections, Ottawa, Canada; DAFZ = Department of Agriculture and Forest Zoology, Hel- sinki, Finland; INHS = Illinois Natural His- tory Survey, Champaign, Illinois, USA; HNHM = Hungarian Natural History Mu- seum, Budapest, Hungary; LUZM = Lund University Zoological Museum, Lund, Swe- den; SMNH = Swedish Museum of Natural History, Stockholm, Sweden; USNM = Na- tional Museum of Natural History, Wash- ington, D.C., USA.
Genus Mestocharis Forster
Mestocharis Forster, 1878: 50. Type-species: Entedon bimacularis Dalman, 1820: 181 (= Mestocharis cyclospila Forster, 1878: 50), by original designation.
Diagnosis.—Species of Mestocharis are distinguished from other genera of Ente- dontinae by the following combination of characters: both sexes with two small and discoid anelli; antennal scrobes adjoining on horizontal line of frontal fork; mandibles tridentate; transverse pronotal carina ab- sent; postmarginal vein about as long as stigmal vein; anterior part of propodeum with two conspicuous indentations lateral- ly; anteromedian part of propodeum with a large and more or less triangular projection.
Remarks.—The monophyly of Mesto- charis is shown through the following syn- apomorphies: 1) anterior part of propo- deum with two conspicuous indentations laterally; 2) anteromedian part of propo- deum with a large triangular projection.
KEY TO THE HOLARCTIC SPECIES OF MESTOCHARIS
bho
2. Costal cell with a complete row of setae on underside; 2nd tergite smooth and shiny ne M. maculata (Forster) — Costal cell bare; 2nd tergite microreticulate (Fig. 1) hs ; 3 3. Seventh tergite 1.2-2.0= as long as width of base of same tergite (Fig. 8); scutellum smooth and shiny medially along its entire length M. tropicalis Yoshimoto — Seventh tergite 0.4-1.0= as long as width of base; scutellum with weaker reticulation me- dially, but not smooth (Figs. 1, 6) M. bimacularis (Dalman) 4. Costal cell with a row of setae on underside, antenna mainly testaceous = M. maculata (Forster) — Costal cell bare, apical part of scape and the antenna beyond usually infuscate M. bimacularis (Dalman)
Mestocharis bimacularis (Dalman) Figs. 1-7
Entedon bimacularis Dalman, 1820: 181.
Entedon arisha Walker, 1839: 121-122. Syn. Boucek et al., 1963: 5.
Mestocharis cyclospila Forster, 1878: 50. Syn. Bouéek et al., 1963: 5.
Mestocharis militaris Rimsky-Korsakov, 1933: 232, 244-245. Syn. Bouéek et al., 1963: 5.
Mestocharis nearctica Yoshimoto, 756-757. New synonymy.
Mestocharis bimacularis (Dalman), Bouéek et al., 1963: 5.
Diagnosis.—Costal cell bare; second ter- gite microreticulate; seventh tergite 0.4 to 1.0 as long as wide at base; scutellum re- ticulate all over; femora usually more or less infuscate.
Description.— Female: Entire antenna dark, except scape more or less pale at base. Face and clypeus golden-green, frons com- pletely golden-purple or golden-green below frontal fork. Vertex, mesoscutum and scu- tellum golden, coppery, golden-green or golden-red. Coxae dark and metallic, major part of femora dark and more or less me- tallic, tibiae varying from brownish to yel- low, base of tibiae occasionally dark, fore- tarsi dark, mid- and hindtarsi with basal
1976:
Figs. 1-8. Mestocharis spp. 1-7, M. bimacularis. 1, Habitus, female. 2, Front view of head, female. 3, Same, male. 4, Female antenna. 5, Male antenna. 6, Female gaster. 7, Male petiolus and gaster. 8, M. tropicalis, female
gaster. 30
VOLUME 90, NUMBER 1
three segments pale and with 4th segment dark. Forewing with a large fuscous spot just below stigmal vein, usually also with two much fainter spots below that spot (see re- marks for comments on these spots). Pro- podeum and gaster golden-green. Length of body: 1.6-2.6 mm. Ratios height of eye/ malar space/width of mouth opening = 3.1/ 1.0/2.0. Malar space 2.5-3.0x as wide as width of scape. Frons below fork with very weak reticulation and almost smooth, part between toruli and antennal scrobes raised In a conspicuous, blunt elevation. Frons above fork smooth and shiny in lower part, in upper part reticulate with low and narrow septa, and with small meshes. Horizontal line of frontal fork shaped like a V. Inner orbit of compound eye with | to 2 rows of setae. Vertex reticulate with very low and very narrow septa inside ocellar triangle, outside smooth and shiny. Ratios POL/ OOL/POO = 6.0/3.1/1.0. Occipital margin with a +strong carina behind ocellar tri- angle. Ratio width of head/width of thorax across shoulders = 1.1. Mesoscutum and scutellum reticulate with high and wide sep- ta, median part of scutellum with lower sep- ta. Forewing with speculum closed, base of submarginal vein with a short row of setae on underside. Propodeum with a strong and complete median carina, plicae complete or missing in anterior part of propodeum, pro- podeal surface rather strongly reticulate, also with irregular carinae. Propodeal callus with 6-9 and lateral part of propodeum inside spiracular sulcus with 3-10 setae. Petiolar foramen rounded. Petiolus transverse, me- dian part with considerably raised carinae, lateral corners protruding and sharp. Shape of gaster varying from ovate (Fig. 6) to lan- ceolate (Fig. 1). First tergite smooth and shiny, tergites 2-5 microreticulate in ante- rior 3/4, tergites 6-7 with stronger reticu- lation. Mean ratio length of thorax + pro- podeum/length of gaster = 0.82 + 0.104, n= 10.
Male: Scape pale with apical part dark, pedicel and two first flagellar segments oc-
31
casionally paler than remaining flagellum, which is dark. Face and frons golden-green or -blue, vertex golden-red. Thorax golden- green, -blue or -red. Coxae dark, remainder of legs yellowish, hindfemur usually dark at base. Forewing immaculate. Metallic col- oration much brighter than in female. Length of body: 1.3-1.8 mm. Frons above fork and vertex smooth and shiny. Ratios height of eye/malar space/width of mouth opening = 4.6/1.0/2.6. Malar space as wide as width of scape. Ratio width of head/width of tho- rax across shoulders = 1.3. Propodeal callus with 4-7 and lateral part of propodeum in- side spiracular sulcus with 2-7 setae. Petio- lus like in female, but varying in shape from transverse to as long as wide. Mean ratio length of thorax + propodeum/length of gaster = 1.06 + 0.087 n = 10.
Remarks.— The dark spots on the female forewings are apparently characters that de- velop with age. Newly emerged females have immaculate forewings, while the same fe- males 8-13 days later have clearly visible spots. In females that have lived 9-10 months the spots are exceptionally dark (Jackson 1964).
The shape of the female gaster is variable in this species, varying from ovate (Fig. 6) to lanceolate (Fig. 1). This character (ac- tually, the shape of last tergite) was used in Bouéek et al. (1963) to separate bimacularis from maculata: the last tergite was about halfas long as its basal width in bimacularis and as long or longer as its basal width in maculata. In Swedish specimens of bimacu- /aris last tergite varies from slightly less than half as long to as long as its basal width.
Yoshimoto (1976) separated nearctica from bimacularis through several characters presented in a table. However, these char- acters are either so variable intraspecifically that they have no taxonomic value or are misinterpreted. The propodeal carinae are variable characters. The median carina is usually strong, wide and complete, but in a few specimens it is narrower. The diagonal carina extending from the triangular pro-
32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
jection to the sides of the petiolar foramen varies from strong and complete to com- pletely missing. The hind margin of the first tergite is usually sinuate while the hind mar- gin of the second tergite varies from almost straight to sinuate. The characters of the male antenna of nearctica must have been misinterpreted by Yoshimoto. A male para- type of nearctica (Can., Ont., Ottawa 19.vu1.1939, O. Peck) that I saw showed the following characters: the scape is yellow with apical fourth infuscate and the first flagellar segment is about 1.8 x as long as wide. The measurements of the fifth flagellar segment are correct (1.7 x as long as wide), but this is also about the same size usually encoun- tered in bimacularis. Consequently I regard nearctica as a synonym of bimacularis.
Material examined.—Two paratypes of M. nearctica (1 2 and 1| 4, in CNC); Canada: British Columbia 2 2 (CNC, LUZM): New- foundland | ¢ (BMNH): Nova Scotia | 2 (CH); Ontario 1 2 (LUZM). Finland: 3 ¢ (DAFZ). Sweden: 40 ¢ 24 6 (CH, LUZM). USA: Michigan | ¢é (USNM). Lectotype 2 E. bimacularis (not seen) in SMNH.
Hosts.—This species is a solitary or gre- garious endoparasitoid in eggs of Dytiscidae (Coleoptera). The size of the egg restricts the number of wasps that can develop suc- cessfully, in larger eggs, e.g. those of Dytis- cus marginalis, up to 12 wasps have been reared, while in smaller eggs, e.g. those of Ilybius ater, only one wasp developed (Jack- son 1964). Imagines of /. bimacularis are most frequently encountered in pond- or marsh vegetation.
Distribution.— Widespread in Europe (Boucek and Askew 1968), in the Nearctic Region this species is recorded from both Canada (Alberta, Manitoba, Ontario and Quebec (Yoshimoto 1976); British Colum- bia, Newfoundland and Nova Scotia) and the United States (Michigan).
Mestocharis maculata (Forster)
Eulophus maculatus Forster, 1841: 41-42. Pleurotropis maculata (Forster), Erdos, 1956: 38-39.
Mestocharis maculata (Forster), Bouéek et al., 1963: 9.
Diagnosis.—Costal cell with a complete row of setae on underside: second tergite smooth and shiny; femora more or less pale testaceous. In other characters M. maculata is very similar to M. bimacularis, and the description of bimacularis is otherwise ap- plicable to maculata.
Material examined.—USSR: Moldavian SSR 1 4; Yugoslavia: Beograd | 2. Both specimens are from the Bouéek collection. Lectotype 2 E. maculatus (not seen) in the Forster collection in Vienna.
Distribution.—Europe: Czechoslovakia, Germany, Hungary and USSR (Bouéek and Askew 1968).
Mestocharis tropicalis Yoshimoto Fig. 8
Mestocharis tropicalis Yoshimoto, Ti Diagnosis.—Seventh tergite 1.2 to 2.0x as long as width of base of same tergite; scutellum smooth and shiny medially along its entire length; costal cell bare; femora pale. Males are unknown for this species. Remarks.—This species is similar to its two congeners but can be separated from them using the characters given in the key. The 7th tergite shows a great deal of intra- specific variation. I have seen only two spec- imens, both females, one from Florida (paratype) and one from Illinois. The spec- imen from Florida had ratio length/basal width of 7th tergite = 1.2, and the specimen from Illinois = 2.0. There is a great gap between these two measurements, but when more material turns up this gap may be filled. The shape of the gaster 1s more lanceolate than in M. bimacularis, which leaves no doubt that tropicalis is a valid species. Like M. bimacularis, tropicalis also has a fuscous spot just below the stigmal vein, but not the two fainter spots present in bimacularis. Material examined.— Paratype 2 (CNC); 1 2 from USA, Illinois, Champaign Co. (INHS). Holotype 2 (not seen) in CNC.
1976:
VOLUME 90, NUMBER 1
Distribution.—The United States (Flori- da, Illinois).
Genus Grahamia Erdos
Grahamia Erdés, 1966: 406. Type-species: Entedon clinius Walker, 1839: 90, by original designation.
Diagnosis. —Species of Grahamia are dis- tinguished from other genera of Entedon- tinae by the following combination of char- acters: all flagellar segments free; antenna with only one small and discoid anellus; mandibles tridentate; pronotal collar with- out transverse carina; costal cell narrow; postmarginal vein about 2 x as long as stig- mal vein.
Remarks.—The monophyly of Grahamia is shown through the single discoid anellus, a synapomorphy for the genus. Two things might argue against the value of this char- acter. First, it is clearly a reduction (the ple- siomorphic character state is three anelll, present in the closely related genus Chry- socharis Forster). Reductions are some- times regarded as poor apomorphic char- acter states. Secondly, this character state (one discoid anellus) occurs in other closely related genera (some Chrysonotomyia Ash- mead and Closterocerus Westwood). Never- theless, I prefer to keep Grahamiaa separate genus from Chrysocharis, the genus to which Grahamia shows closest morphological af- finity. Among the species-groups of Chry- socharis (sensu Hansson 1985), Grahamia comes closest to the mediana-group. Gra- hamia has, however, some characters that disagree with this assessment: its single dis- coid anellus of the antenna, all flagellar seg- ments free, and gallmidges as hosts. Chry- socharis has three anelli, two apical flagellar segments fused in species of the mediana- group, and leafminers as hosts.
KEY TO THE SPECIES OF GRAHAMIA (FEMALES)
1. First segment of flagellum about 2* and 4th segment about 1.5 = as long as wide (Fig. 12); malar space about 1.5= as wide as width of scape; metallic coloration of body compara- tively dull .. ..G. clinius (Walker)
— First segment of flagellum about 1.5 * and 4th segment 1 x as long as wide (Fig. 10); malar space narrower (as wide as width of scape); metallic coloration of body brighter
G. tatrica Erdés
Grahamia clinius (Walker) Figs. 11-12
Entedon clinius Walker, 1839: 90.
Chrysocharis clinius (Walker), Graham, 1959: 196.
“Chrysocharis”’ clinius (Walker), Graham, 1963: 204.
Grahamia clinius (Walker), Erdés, 1966: 407.
Diagnosis. — Flagellar segment | about 2 x and 4 about 1.5= as long as wide; malar space 1.5 as wide as width of scape; me- tallic coloration of body dull; larger species (1.6-2.1 mm).
Description.— Female: Entire antenna dark, scape occasionally paler in basal half. Face and clypeus golden-green. Frons and vertex metallic purple, below fork some- times golden-purple. Mesoscutum golden- green and scutellum metallic purple. Coxae dark brown and weakly metallic. Femora predominantly dark, fore- and midtibiae varying from pale to infuscate, hindtibia predominantly pale, tarsi + infuscate, 4th segment dark. Wings hyaline or weakly in- fuscate. Propodeum and gaster golden-green. Length of body: 1.6-2.1 mm. Ratios height of eye/malar space/width of mouth open- ing = 3.9/1.0/2.8. Malar space 1.5 x as wide as width of scape. Frons below fork reticulate with low and rather narrow septa, meshes small, above fork shiny and almost smooth. Horizontal line of frontal fork almost straight. Inner orbit of compound eye with one row of setae. Vertex reticulate with very low and very narrow septa. Ratios POL/ OOL/POO = 3.3/2.2/1.0. Occipital margin with a carina behind ocellar triangle. Ratio width of head/width of thorax across shoul- ders = 1.3. Mesoscutum reticulate with low and rather narrow septa, meshes small. Scu- tellum reticulate with low to very low and narrow to very narrow septa, 1.e. with finer
34
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
| 10 Nes) Ne: ii Sy Figs. 9-12. 9-10, Grahamia tatrica, female. 9. Habitus. 10, Antenna. 11-12, G. clinius, female. 11, Front
view of head. 12, Antenna.
VOLUME 90, NUMBER 1
reticulation than mesoscutum. Anterome- dian part of propodeum with a weak tri- angular fovea, propodeal surface +reticu- late, with or without a weak median carina. Propodeal callus with two setae. Petiolar fo- ramen rounded. Petiolus small and trans- verse. Gaster elongate, ratio length of tho- rax + propodeum/length of gaster = 0.67-
0.87. Remarks.—The species Tetrastichus
idothea Walker, 1844: 409, was regarded as a possible synonym of G. clinius by Graham (1961: 62).
Material examined.—Lectotype EF. cli- nius 2 (BMNH Type No. 5.2025); BRD: 1 2 ex Hapl. equestris (USNM). Canada: Brit- ish Columbia 3 2? (CNC, LUZM). Sweden: 1 2 (CH).
Host.—Grahamia clinius is known as an endoparasitoid in larvae of Haplodiplosis equestris (Diptera, Cecidomyiidae) (Baier, 1963/64, as Chrysocharis seiuncta). The sex ratio of reared G. clinius (2:6 = 100:1) (Baier 1963/64) suggests that the species propa- gates parthenogenetically.
Distribution. — Grahamia clinius is wide- spread in Europe (Boucek and Askew 1968), and now for the first time recorded from the Nearctic Region (Canada, British Colum- bia).
Grahamia tatrica Erdos Figs. 9-10
Grahamia tatrica Erdés, 1966: 407. Chrysocharis atripes Szelenyi, 1979: 178. Syn. Hansson, 1985: 97.
Diagnosis.—Flagellar segment | about 1.5 and 4 about | x as long as wide; malar space as wide as width of scape; metallic coloration of body brighter; smaller species (1.3-1.5 mm).
Remarks.—Apart from the distinguish- ing characters G. tatrica is very similar to G. clinius and the description of c/inius is applicable to tatrica.
Material examined. —Paratypes 2 2 G. ta- trica (HNHM Nos. 6061 & 6062); Canada: Nova Scotia | ? (LUZM),; Finland: 10 2 (CH,
35
DAFZ); Sweden: 2 2 (CH); USA: Michigan 1 @ (USNM), West Virginia | @ (LUZM). Holotype 2 (not seen) in HNHM.
Distribution.—Grahamia tatrica 1s re- corded from Europe (Finland, Hungary (Er- dds 1966), Sweden, Switzerland (Erdés 1966)), Canada (Nova Scotia) and the United States (Michigan, West Virginia). This species was previously not recorded from the Nearctic.
ACKNOWLEDGMENTS
I am grateful to Z. Boucek (BMNH), G.A.P. Gibson (CNC), S. L. Heydon (INHS), L. Huggert (same address as author), M. Ko- ponen (DAFZ), J.S. Noyes (BMNH), J. Papp (HNHM), M. E. Schauff (USNM), and C. M. Yoshimoto (CNC) for loan of type- and additional material. My work with this pa- per was partly financed through a grant from the O. Larsén Foundation.
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1966. Nonnulae Eulophidae novae Hunga- ricae (Hymenoptera, Chalcidoidea). Ann. Hist. Nat. Mus. Natl. Hung. Zool. 58: 395-420.
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1963. Additions and corrections to the Brit- ish list of Eulophidae (Hym., Chalcidoidea), with descriptions of some new species. Trans. Soc. Br. Entomol. 15: 167-275.
Hansson, C. 1985. Taxonomy and biology of the Palearctic species of Chrysocharis Forster, 1856 (Hymenoptera: Eulophidae). Entomol. Scand. Suppl. 26: 1-130.
Jackson, D. J. 1958. A further note on a Chrysocharis (Hym. Eulophidae) parasitizing the eggs of Dytis- cus marginalis L. and a comparison of its larva with that of Caraphractus cinctus Walker (Hym. Mymaridae). J. Soc. Br. Entomol. 6: 15-22.
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1964. Observations on the life-history of Mestocharis bimacularis (Dalman) (Hym. Euloph- idae), a parasitoid of the eggs of Dytiscidae. Opusc. Entomol. 29: 81-97.
Rimsky-Korsakov, M. N. 1933. Methoden zur Un- tersucherung von Wasserhymenopteren. Handb. Biol. ArbMeth. 9: 227-258.
Szelenyi, G. 1979. Four new species of Chrysocharis Forster, 1856 (Hymenoptera: Chalcidoidea: Eu- lophidae). Acta Zool. Acad. Sci. Hung. 25(1-2): 177-181.
Walker, F. 1839. Monographia Chalciditum. I. Lon- don. 333 pp.
1844. On the species of Chalcidites inhab- iting the Arctic Region. Ann. Mag. Nat. Hist. 14: 331-342, 407-410.
Yoshimoto, C. M. 1976. Synopsis of the genus Mes- tocharis Forster in America north of Mexico (Chalcidoidea: Eulophidae). Can. Entomol. 108: 755-758.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 37-43
GALL FORMATION BY THE CAPITULUM-INFESTING FRUIT FLY, TEPHRITIS STIGMATICA (DIPTERA: TEPHRITIDAE)
RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, California 92521.
Abstract.— Known heretofore only as a capitulum-infesting species, Tephritis stigmatica (Coquillett) also is reported here to develop in galls formed on branches and stems of Senecio douglasii (deCandolle) (Asteraceae) in southern California. Two generations of galls occur each year on S. douglasii, with immature larvae in juvenile, F, galls constituting at least part of the overwintering population. One or more larvae feed gregariously in interconnected, central longitudinal, short-branched feeding tunnels in the expanded pith. As many as six individuals, including five puparia, were found in one F, gall. Florets (including ovaries) were fed upon in ungalled capitula and F, and F, galls were formed on branches and stems of S. douglasii at some locations, but gall-formation is the much less common and less widespread mode of development. Galls are described and pictured.
Tephritis stigmatica (Coquillett) is the only tephritid among the 17 species in this genus currently described from North America north of Mexico for which the life history and behavior have been reported in any de- tail (Foote 1960a, Tauber and Toschi 1965, Foote and Blanc 1979). Tauber and Toschi (1965) studied this species reared from ca- pitula of Senecio integerrimus Nuttall (As- teraceae), acommon perennial shrub in cen- tral- and northeastern-montane California (Munz and Keck 1959, Munz 1968). In 1981, I first reared what appeared to be 7. stigmatica or an undescribed congener from galls on branches and stems of Senecio douglasii (deCandolle), another common perennial shrub in the same genus, but widely distributed below ca. 1500 m in Cal- ifornia (Munz and Keck 1959, Munz 1968). The capitula of S. douglasii also are widely infested by 7. stigmatica (Foote and Blanc 1963, Tauber and Toschi 1965, Wasbauer 1972).
Tephritis dilacerata Loew is fairly well known from studies by Berube (1978) and
Shorthouse (1980), but this European species is an obligate gall former in capitula and stems of its host, unlike the unique host- plant relationship reported herein. Several other species of Tephritis are known to form capitulum and stem galls; others to infest capitula without forming galls, but none to do both (Freidberg 1984). The purpose of this report is to describe gall formation by T. stigmatica as a unique alternative mode of development for this tephritid heretofore known only from capitula.
MATERIALS AND METHODS
Galls on branches and stems of S. doug- lasii from which adults recently had emerged and galls containing larvae or puparia were sampled at two locations in southern Cali- fornia south of Lamont Peak at Spanish Needle Creek, Sequoia Nat. Forest, Kern Co., on 24 VII 1984, 7 VIII 1984, and 3 III 1987; and 2 km south of Pearblossom, San Bernardino Co., on 22 IV 1985 and 12 I 1987.
Fully-grown larvae and puparia extracted
38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
from some of these galls were placed indi- vidually in glass shell vials with perforated plastic caps and held for adult emergence at room temperature in a covered bell jar at near-saturation R.H. The external mor- phology of adults reared from these and sev- eral other collections of galls were compared at magnifications of up to 50 to 7. stig- matica adults reared from capitula of S. douglasii and other Asteraceae sampled throughout southern and central California during 1980-86.
All host-plant identifications were made by or confirmed by A. C. Saunders, Curator of the Herbarium of the University of Cal- ifornia, Riverside. The plant nomenclature used is that of Munz and Keck (1959) and Munz (1968, 1974); the insect nomencla- ture, that of Foote (1959, 1960a).
RESULTS
Taxonomy.—The 26 4 and 32 2 reared from puparia dissected from branch and stem galls (18 4 and 29 2 were reared from the first Pearblossom sample) showed no consistent differences compared to adults of T. stigmatica reared from capitula. The lar- vae feed on the florets (including ovaries) but do not gall the capitula of S. douglasii (personal observation). Other than body size, body parts examined, measurements taken, and ratios calculated in this study (Table 1) were the same as those used by Quisenberry (1951) of Nearctic Tephritis, including T. stigmatica, and by Foote (1959) in his di- agnosis of the most recently described species of Tephritis.
Foote and Blanc (1963, p. 72) termed 7. stigmatica “the largest of the California Te- phritis species.” T. stigmatica adults reared from branch and stem galls on S. douglasii were larger on the average than adults reared from capitula of both S. douglasii and S. integerrimus (Table 1). The size of adults reared from capitula of these and other As- teraceae also varied considerably (Table 1). This probably reflected the different nutri- tional value of the capitula of varying size
and maturity (S. N. Thompson, in /itt. 1987), as well as the degree of larval development when the heads were sampled. A tiny male reared from a capitulum of Haplopappus venetus (Humboldt) Blake ssp. vernonioides (Nuttall) Hall, an uncommon and appar- ently nutritionally ill-suited host (as well as a new host genus and species record, Was- bauer 1972) at Cardiff-by-the-Sea, San Di- ego Co., on 15 x 1980, illustrates the ex- treme effect of host-plant unsuitability on adult size in 7. stigmatica (Table 1). Dif- ferences in mean head-part measurements as indices of body size differences (Foote 1960a) between males and females and among flies reared from galls versus capitula of the same or different host plants alone were insufficient to warrant description of the gall-forming flies as a separate species. F. L. Blanc, California Department of Food and Agriculture (Retired), Sacramento, con- firmed that specimens in Table | were 7. stigmatica (in litt. 1987).
Galls.— Tephritis stigmatica overwinters in southern California at least partly as im- mature larvae in F, galls formed mostly on low axillary branches on S. douglasii (Fig. la). Adults that emerge from heads in late summer and fall also overwinter. Young F, galls examined in February (mid-winter) 1n 1987 at Pearblossom and early March 1987 at Spanish Needle Creek were a third to half of full size. Most F, galls (Fig. 1b), like un- galled, elongating current season branches, showed darker (purple) coloration than galls and branch growth formed later in the year. The pigmentation could favor solar energy adsorption and facilitate larval and pupal development as well as adult emergence during cooler months of the year. Galls and ungalled branches that develop during the hot summer are mostly light green (Fig. 1a, 1d). As many as five axillary branches bore F, galls along one 10-cm basal section of stem (Fig. la). In contrast, F, galls usually were isolated on upper parts of aerial stems and usually contained only one or two te- phritids (Fig. 1d). Dissection of 72 juvenile
VOLUME 90, NUMBER |
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40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
overwintering F, galls yielded mostly 2nd instars and small to half-grown 3rd instars. A few fully-grown larvae and puparia also were found in F, galls collected in early March at Spanish Needle Creek. From one to five individuals in the same or different stages of growth, e.g. 2nd instars and pu- paria, were found together in overwintering F, galls within interconnected, longitudinal, short-branched feeding tunnels.
Fifty overwintered, fully-formed F, galls collected at Pearblossom in April, 1985, contained an average of 1.5 (range, 1-6) ful- ly-grown larvae and intact or empty puparia (Fig. 1c). All but one of the 50 galls were spindle-shaped (Fig. 1b, 1d). These galls either were sessile or born on pedicels of various lengths, up to 5 mm long. These F, galls tapered apically as a result of continued apical meristem growth and branch elon- gation distal to the gall; whereas, death of the apical meristem from larval feeding re- sulted in a club-shaped gall, or curved gall as a lateral branch assumed apical domi- nance (Goeden and Ricker 1981, Goeden 1987). Dissections suggested that multiple Ovipositions by one or more F, females in an axillary bud initiated the formation of a F, gall during the previous fall; whereas, a female emerging from these galls the fol- lowing spring probably oviposited in the apical meristem of an upright aerial stem to initiate F, galls. Capitula production was observed to be delayed on F, gall-bearing plants sampled at Spanish Needle Creek in early August, 1984.
Branch and stem galls are formed from expanded pith tissue, the branches and stems swelling to about three times normal di-
ameter (Fig. 1b, 1d). Vascular strands were deflected outward and followed the gall con- tour. Irregular masses of hypertrophied parenchyma cells lined the walls of the feed- ing chamber(s), resembling round, shiny globules which the larvae scored with mouthhooks while feeding. Frass lined the feeding tunnels of the late-stage 3rd instars (Fig. lc, 1g). Only part of the gall mass was consumed by a single larva (Fig. le, 1g); although, the interiors of smaller galls, es- pecially those containing two or more lar- vae, were largely consumed (Fig. 1b). Forty- two, spindle-shaped, overwintered, fully formed F, galls collected at Pearblossom in 1985 averaged 2.4 + 0.1 (range, 1.4—-3.2) cm in length and 6.5 + 0.2 (range, 4-10) mm in greatest diameter. These galls incor- porated as many as six nodes as indicated by the number of lateral branches arising thereon. The fully-grown larva extends its feeding tunnel distally and outward to, but not through, the epidermis leaving a round, thin, 2-mm dia., translucent window to the outside through which the emerging adult eventually exits (Fig. 1f, 1h, 11). Once the window is formed, the larva returns to its feeding tunnel and pupariates, often with the anterior part of the puparium projecting into the exit tunnel, and usually facing the window (Fig. 1g). The larval predecessor of one of 75 (1.3%) puparia examined in over- wintered Pearblossom galls tunneled basal- ly in constructing its exit tunnel. A maxi- mum of three windows was formed in one gall from Pearblossom that contained five puparia and one larva; otherwise, most galls contained a single window through which as many as three flies emerged. Exit holes
Fig. 1.
—
Galls of Tephritis stigmatica on Senecio douglasii: a, overwintering, juvenile, axillary-branch, F, galls
(0.8 x); b, fully-formed, dark-pigmented, spindle-shaped, F, gall with lateral window (2.1 =); c, same gall as in b opened to expose four puparia within (3.3); d, fully-formed, compound, F, gall showing lateral exit hole (0.8 x); e, same gall as in d opened to expose two empty puparia in interconnected, central-longitudinal feeding tunnel and two lateral exit tunnels (1.3 x); f, fully-grown larva just having completed an exit tunnel ending in circular window (4.5 «); g, puparium at juncture of short feeding tunnel and lateral exit tunnel (4.6 x ); h, closeup view of broken, epidermal window on gall surface (3.6); i, newly-emerged female reared from gall (7.6).
VOLUME 90, NUMBER 1 4]
42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
usually were constructed in the distal third or half of the overwintered Pearblossom galls at, below, or between the nodes (Fig. 1b, 1d, le, 1f, 1g, lh). Either one or both sexes of flies (Fig. 11) emerged from individual F, galls.
Among 39 F, galls collected at Spanish Needle Creek, only one (2.8%) was club- shaped, and the remainder, spindle-shaped. The 32 spindle-shaped F, galls from which flies had emerged in the field in the fall were 4.7 + 0.3 cm (range, 2.0-7.5) long and 8 + | (range, 5-10) mm wide at their widest. These spindle-shaped galls incorporated an average of 4 (range, 1-8) nodes. The 39 F, galls contained one or two, empty, or intact and parasitized, puparia in interconnected central-longitudinal feeding tunnels (Fig. le). Thirty-three of these galls had contained a single tephritid and six had contained two tephritids (Fig. le), which as noted above was considerably less than the number of individuals in F, galls from Pearblossom (Fig. lc). The feeding tunnels in these F, galls averaged 1.3 + 0.2 (range, 0.5-3.5) cm in length and were 2 mm wide. The exit tunnels averaged 3 (range, 2-5) mm in length and also were circular and 2 mm in cross- sectional width. Ten intact and mostly par- asitized empty puparia in these galls aver- aged 4.2 + 0.1 (range, 3.6—4.6) mm in length and 1.9 + 0.0 (range, 1.7—2.0) mm in width. An unidentified species of Eurytoma (Hy- menoptera: Eurytomidae) was reared from these puparia.
Galls were collected from S. douglasii at the following locations in addition to those noted above: 2 4 reared from galls collected south of Hesperia at Mojave River Forks, San Bernardino Nat. Forest, San Bernar- dino Co., on 21 IV 1981: 8 4 reared from galls collected at Orcutt, Santa Barbara Co., on 23 VI 1981; 1 dand | 2 reared from galls collected at Cajon Junction, San Bernardino Co., on 8 VII 1981. This represents only a small fraction of uncounted demes of S. douglasii examined, but found to lack galls during my many wide-ranging field trips
throughout southern California during 1981-1986.
DISCUSSION
Morphological study of 7. stigmatica from galls and capitula on S. douglasii to date has failed to support what otherwise might have been interpreted as sympatric speciation oc- curring on the same species of host plant. For example, Goeden (1987) reported that the somewhat rare native tephritid, 7ru- panea conjuncta (Adams), facultatively either galls the apical meristems or develops gregariously feeding on florets (including Ovaries) in capitula of its sole host plant, Trixis californica Kellogg (Asteraceae). This was the first published report of this facul- tative mode of development in nonfrugi- vorous Tephritidae (Zwolfer 1983, Freid- berg 1984, Price et al. 1986). However, with T. stigmatica, a member of the genus most closely allied to 7rupanea according to Foote (1959, 1960a, 1960b), development in ca- pitula or galls apparently are not mutually exclusive activities on S. douglasii at a par- ticular site and season, 1.e. both gall for- mation and feeding in capitula may occur simultaneously during summer into fall. However, gall formation apparently is the requisite mode of development for the F, generation on S. douglasii earlier in the year. And, I suspect that at least part of the over- wintered generation produced in branch (axillary bud) galls begun during the pre- ceding year, as well as overwintered adults produced in S. douglasii capitula, reproduce in summer at higher elevations in capitula of a succession of alternate hosts, e.g. S. integerrimus and S. triangularis Hooker (Wasbauer 1972), on which galls apparently are not formed (Tauber and Toschi 1965). This part of the life history of 7. stigmatica needs clarification.
Individual host plants were observed at Spanish Needle Creek on March 3, 1987, that bore current and previous year’s F, galls as well as F, galls on dead upright stems that terminated on branches with last year’s ca-
VOLUME 90, NUMBER |
pitula containing empty puparia. This pro- vided evidence of three successive genera- tions reproducing as gall formers augmented by a capitula-infesting generation on the same host individuals commensurate with the seasonal development described above. As previously noted, galls have not been found on S. douglasii at most locations where and when plants were examined in the field as opportunity allowed since 1981, indicating the absence of some unknown environmental requisite(s) for gall forma- tion by 7. stigmatica. This spotty incidence ofa gall-forming insect occupying only tiny, discrete, often distant, fractions of the total geographic range of its native host plant also was documented for the stem-galling moth, Carollela beevorana Comstock (Lepidop- tera: Cochylidae) by Goeden and Ricker (1981). In contrast, a congener of C. bee- vorana commonly forms galls on a related host plant throughout a large part of south- ern California (Goeden and Ricker 1986).
ACKNOWLEDGMENTS
My thanks to F. L. Blanc, R. F. Foote, A. L. Norrbom, and J. D. Pinto for their help- ful reviews of earlier drafts of this manu- script. The technical support of D. W. Rick- er, including the photography in Fig. 1, also is gratefully acknowledged.
LITERATURE CITED
Berube, D. E. 1978. Larval descriptions and biology of Tephritis dilacerata (Dip.: Tephritidae), a can- didate for the biocontrol of Sonchus arvensis in Canada. Entomophaga 23: 69-82.
Foote, R. H. 1959. A new North American species of Tephritis, with some observations on its generic position (Diptera, Tephritidae). Bull. Brooklyn Entomol. Soc. 54: 13-17.
1960a. The genus Jephritis Latreille in the
Nearctic Region north of Mexico: descriptions of
four new species and notes on others. J. Kans.
Entomol. Soc. 33: 71-85.
1960b. A revision of the genus 7rupanea in
America north of Mexico. U.S. Dep. Agric. Tech.
Bull. 1214. 29 pp.
43
Foote. R. H. and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect Surv. 7. 115 pp.
1979. New species of Tephritidae (Diptera) from the western United States, Mexico, and Gua- temala, with revisionary notes. Pan-Pac. Entomol. 55: 161-179.
Freidberg, A. 1984. Gall Tephritidae (Diptera), pp. 129-167. In T. N. Ananthakrishnan, ed., Biology of Gall Insects. Oxford & IBH Publ. Co., New Delhi, Bombay, Calcutta.
Goeden, R. D. 1987. Life history of Trupanea con- juncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan- Pac. Entomol. 63: 284-291.
Goeden, R. D. and D. W. Ricker. 1981. Life history of the gall-forming moth, Carollela beevorana Comstock, on the ragweed, Ambrosia dumosa (Gray) Payne, in southern California (Lepidoptera: Cochylidae). Pan-Pac. Entomol. 57: 402-410.
1986. Phytophagous insect fauna of the des- ert shrub Hymenoclea salsola in southern Cali- fornia. Ann. Entomol. Soc. Am. 79: 39-47.
Munz, P. A. 1968. Supplement to a California Flora. Univ. Calif. Press, Berkeley, Los Angeles, London. 224 pp.
1974. A Flora of Southern California. Univ. Calif. Press, Berkeley, Los Angeles, London. 1086 pp.
Munz, P. A. and D. D. Keck. 1959. A California Flora. Univ. Calif. Press, Berkeley, Los Angeles. 1681 pp.
Price, P. W., G. L. Waring, and G. W. Fernandes. 1986. Hypotheses on the adaptive nature of galls. Proc. Entomol. Soc. Wash. 88: 361-363.
Quisenberry, B. F. 1951. A study of the genus 7e- phritis Latreille in the Nearctic region north of Mexico (Diptera: Tephritidae). J. Kans. Entomol. Soc. 24: 56-72.
Shorthouse, J. D. 1980. Modification of the flower heads of Sonchus arvensis (family Compositae) by the gall former Tephritis dilacerata (order Diptera, family Tephritidae). Can. J. Bot. 58: 1534-1540.
Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior of Tephritis stigmatica (Co- quillett). Pan-Pac. Entomol. 41: 73-79.
Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Dip- tera: Tephritidae). Calif. Dep. Agric. Bur. Ento- mol. Occas. Pap. 19. 172 pp.
Zwolfer, H. 1983. Life systems and strategies of re- source exploitation in tephritids, pp. 16-30. /n R. Cavalloro, ed., Fruit Flies of Economic Impor- tance. Proc. CEC/IOBC Int. Sym., Athens, Greece, Nov. 1982, A. A. Balkema, Rotterdam.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 44-46
NOTES ON THE BIOLOGY OF TWO PHYCITINES (LEPIDOPTERA: PYRALIDAE) ASSOCIATED WITH TOUMEYELLA PINI (HOMOPTERA: COCCIDAE) ON PINE
A. T. Drooz AND H. H. NEUNZIG
(AD) 705 Jefferson Dr., Cary, North Carolina 27511; (HN) Department of Entomology, North Carolina State University, Raleigh, North Carolina 27695-7613.
Abstract. —The phycitines, Laetilia coccidivora (Comstock) and Ribua innoxia Heinrich were reared from slash pine, Pinus e/liottii elliottii Engelmann, infested by the striped pine scale, Toumeyella pini (King), in northern Florida. R. innoxia is new to the USA. It was found previously only in Cuba and Puerto Rico, where it reportedly fed upon fungus on
pineapple.
We report here the new distribution and record of association of R. innoxia, and its fall emergence and that of L. coccidivora in northern Florida. A comparison of the emer- gence patterns of the 2 species and significant differences in the appearance of their cocoons
are presented.
Scale-infested slash pine, Pinus elliottii el- liottii Engelmann, shoots were collected and caged on 30 September, 1985, at a pine seed orchard near White Springs, Florida, to in- vestigate the natural control factors of the striped pine scale, Toumeyella pini (King). Two species of phycitine moths emerged, Laetilia coccidivora (Comstock) and Ribua innoxia Heinrich. L. coccidivora is preda- ceous on numerous species of scale insects (Heinrich 1956). R. innoxia has not been reported from the USA, and information on its biology is fragmentary. It had been found only in Cuba and Puerto Rico and is con- sidered to be a scavenger or mycetophage (Heinrich 1956).
METHODS
A total of 100 scale-infested terminals within reach of the ground were cut from a number of pines. The foliage was trimmed to ca. 2 cm to fit the cages and the twigs were cut to 10 cm including the buds. The
twigs were caged individually in 5 cm diam. x 8.5 cm long clear plastic vials with snap lids. Water was not used with the twigs, as the scale crawlers were about to hatch. A 1 cm hole was cut out of each lid and a piece of fine mesh nylon was glued to cover the inside of the hole. Thus, aeration was pro- vided, yet scale crawlers and other insects that might emerge were retained. The in- sects were reared at ambient indoor tem- peratures. The phycitines were larvae at this time.
The cages were examined daily until no moths appeared over a 2-week period. Moths were removed each evening as they emerged and were frozen until they could be mounted and identified. The number and species of moths emerging by date were re- corded.
RESULTS AND DISCUSSION
Ninety-four L. coccidivora were reared from 46 twigs, and 18 R. innoxia came from
VOLUME 90, NUMBER |
Figs. 1, 2. 1, Cocoon of Laetilia coccidivora. 2, Cocoon of Ribua innoxia.
45
46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
24 twigs. No moths were found in 39 cages. Cocoons of the two phycitines differed greatly. The cocoon of L. coccidivora 1s sim- ilar to that made by many Lepidoptera that pupate above ground: an elongate, white, densely silked, but soft enclosure (Fig. 1). Larvae of R. innoxia formed a fragile, mesh or netlike cocoon composed of silk and frass (Fig. 2).
Emergence of L. coccidivora started on 18 Oct., with peak emergence between 27 Oct. and 9 Nov. and decreased through 17 Nov., with the last 2 moths appearing between then and | Dec. Moths of R. innoxia began to emerge on 4 Nov., more than 2 wk after L. coccidivora. Only | or 2 moths emerged occasionally until the last one appeared on 13)'Dec:
Holes in the integument of the scales, caused by feeding of L. coccidivora larvae, were common. The relationship between R. innoxia larvae and the scales was not es- tablished. Large amounts of “honeydew” were produced by the scales, and this sub- stance was infested with sooty mold, Cap- nodium sp.
Heinrich (1956) postulated that R. in- noxia larvae feed upon fungus; his type se- ries were associated with fungus on pine- apple, Ananas comosus (L.) Merrill, growing in Cuba. Our brief observations suggests a
mycelium-feeding habit for the larvae. The opinion that the larvae of L. coccidivora and R. innoxia have diverse feeding habits is supported by taxonomic studies of the adults (Heinrich 1956). Laetilia and Ribua are not closely related genera. Laetilia appears to be allied to some of the cactus-feeding phy- citines, whereas Ribua shows affinities to some of the stored products pests, partic- ularly the Indian meal moth, Plodia inter- punctella (Hubner). Further work is needed to establish with certainty the feeding habits and host of R. innoxia.
ACKNOWLEDGMENTS
We thank Douglass R. Miller, USDA, ARS, Systematic Entomology Lab., Belts- ville, MD, for identification of the scale, and Charles Lassiter, USDA, Forest Serv., Southeast. Forest Expt. Stn., Naval Stores and Timber Production Lab., Olustee, FL for help in collecting infested twigs. This 1s paper No. 11072 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, North Carolina 27695- 7601.
LITERATURE CITED
Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus. Bull. 207. 581 pp.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 47-51
OCCURRENCE OF SELECTED FLOWER HEAD INSECTS OF CENTAUREA SOLSTITIALIS IN ITALY AND GREECE
STEPHEN L. CLEMENT AND TIZIANA MIMMOCCHI
USDA, ARS, Biological Control of Weeds Laboratory —Europe, % American Embassy, Rome, Italy, APO New York 09794-0007; (SLC) USDA, ARS Plant Introduction Station, 59 Johnson Hall, Washington State University, Pullman, Washington 99164-6402; (TM) Via Baldo degli Ubaldi 59, 00167 Rome, Italy.
Abstract.—A 1984 survey was conducted in the south-Italian mainland and central Greece to locate sites where biocontrol specialists could collect insects of 6 promising biocontrol agents of yellow starthistle (YST), Centaurea solstitialis L., for use in host specificity tests. Four of these flower head insects were found: Urophora quadrifasciata and Terellia sp. (Diptera: Tephritidae) in Italy, and Eustenopus hirtus and Larinus curtus (Coleoptera: Curculionidae) in Greece. Urophora jaculata was the most abundant and ubiquitous species, but it is not a potential biocontrol agent because larvae will not develop in the heads of U.S. forms of YST. New and supplementary information on the distribution of several YST flower head insects and the extent to which they attack heads in Italy and
Greece are also presented and discussed in relation to published information.
Centaurea solstitialis L. (yellow starthis- tle [YST]) is a Eurasian winter annual or biennial plant that has spread to the United States where it is a weed on over 3 million hectares in some western states (Maddox et al. 1985, Maddox and Mayfield 1985). At- tempts to control YST biologically in the U.S. began in the 1960’s when a flower head gall fly, Urophora jaculata Rondani (Dip- tera: Tephritidae), erroneously called U. si- runaseva (Hering) in some earlier refer- ences, was introduced from Italy (White and Clement 1987). Repeated efforts to estab- lish this fly on U.S. forms of YST were un- successful. However, weed biocontrol workers discovered that a flower head wee- vil from Greece (Bangasternus orientalis (Cap.); Coleoptera: Curculionidae) will at- tack and complete its development on U.S. plants (Maddox and Sobhian 1987). This weevil, first released in western U.S. in 1985, 1s now established in California (Maddox et al. 1986).
We assumed that additional biological control agents would be needed to supple- ment the action of B. orientalis so in 1984 we surveyed YST in Italy and Greece to locate populations of promising agents, namely the tephritid flies Chaetorellia hexa- chaeta (Loew), U. sirunaseva, U. quadrifas- ciata (Meigen), and an undescribed Terellia species (= 7. cf virens (Loew) in Sobhian and Zwolfer [1985]), and the curculionid beetles Eustenopus hirtus (Waltl) (= E. cf abbreviatus Faust in Sobhian and Zwolfer [1985]) and Larinus curtus Hochhut. We targeted these six flower head species for study because our preliminary work and un- published reports at the USDA, ARS Bio- logical Control of Weeds Laboratory—Eu- rope (BCWLE), Rome, Italy, indicated they had restricted host ranges in southern Eu- rope. More than one species may be con- fused under the name quadrifasciata (1. M. White, pers. comm.) and some populations of C. hexachaeta may be separate species
48
Table 1.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Sites surveyed in Italy and Greece, collection dates, total number of flower heads of Centaurea
solstitialis collected and percent damaged by insects at each site, 1984.
Site
Sites Surveyed
Italy
5 km E S. Giovanni Rotondo, Promontorio del Gargano, Puglia Region (41°45'N, 15°55'E)
9 km S S. Giovanni Rotondo
12 km E S. Giovanni Rotondo
15 km NW S. Paolo di Civitate, Puglia (41°40'N, 15°20’E)
Castel del Monte, Puglia (41°40'N, 16°15’E)
10 km N Rome, Lazio Region (42°10'N, 12°15’E)
Greece
5.5 km W Agiokambos (39°45’N, 22°45’E)
Xiniada (39°10'N, 22°20’E)
ca. 6 km E Karpenissi (38°50'N, 22°50’E)
Hounti (38°45'N, 21°30'E)
ca. 2 km S Arta (39°10'N, 20°55’E)
10 km N Igoumenitsa (39°30'N, 20°15’E)
' Same plants were sampled each time.
(I. M. White, in press), but further taxo- nomic study is needed to clarify these pos- sible species groups.
The objective was to locate sources of in- sects of the aforementioned species for use in host specificity tests and to provide new and supplementary information on the ex- tent to which these and other species attack YST heads in southern Europe. Our ap- proach was to record the occurrence of each flower head species on single host popula- tions at several sites in the south-Italian mainland and central Greece. Information of this type is virtually nonexistent in the literature (Zw6lfer 1965, ZwGlfer et al. 1971, Sobhian and Zwoélfer 1985) on YST flower head insects in Europe.
METHODS
Samples were collected at twelve sites during the 1984 flowering season (Table 1).
i)
wn
No. No. % of
Plants Heads Heads Collection Dates Sampled Collected Damaged July 17 and Aug. 21 10! 454 37.89 July 17 and Aug. 21 20! 320) 818575 July 17 and Aug. 21 10! 273 24.54 July 17 52 155 14.02 July 17 and Aug. 21 16! 649 14.19 July 25, Aug. 14 10! 1627 18.19
and Sept. 7
Aug. 3 5 161 18.01 Aug. 3 5) 179 50.28 Aug. 4 5 105 41.90 Aug. 4 2 44 65.91 Aug. 5 10 86 ~—-:10.41 Aug. 5 6 258 19.77
Seven of the sites were sampled once during July and August, but the literature (Sobhian and Zwolfer 1985) and our unpublished data indicate that the six species we were most concerned with can be found on YST during these two months. Therefore, we are rea- sonably certain that our one-time collec- tions were sufficient to establish the pres- ence or absence of these insects at each site. It was convenient to sample five Italian sites more than once because other studies were being conducted at or near these sites. Sur- vey sites were roadside areas, embank- ments, and open fields (<3.5 ha) along roads. Each collection consisted of all heads in the flowering and seed formation stages (see Maddox 1981 for description of stages) from five or more randomly selected plants per site, except at one site where only two plants were available for sampling. At five Italian sites, the same plants were sampled two or
VOLUME 90, NUMBER 1
49
Table 2. Occurrence of flower head insects of Centaurea solstitialis at several sites in Italy and Greece, 1984. Relative Occurrence! Italian Sites Greek Sites Species 1 3 4 5 6 1 2 3 4 5 6 Diptera Tephritidae? Urophora jaculata Rondani te Ser a Ft TE SEED OE ee (OTE Oke x U. quadrifasciata (Mg.) * sia Terellia sp. ia aa =i Acanthiophyllus helianthi (Rossi) ae se ee Chaetorellia sp. nr. C. carthami Stack. is oe et xs Coleoptera Curculionidae?’ Bangasternus orientalis (Capiomont) 12) |) 22 Eustenopus hirtus (Waltl) eS * *
Larinus curtus Hochhut
Bruchidae Bruchidius tuberculatus (Hochhut)* Anobiidae Lasioderma sp. nr. haemorrhoidale (Illiger)> Lepidoptera Cosmopterigidae Pyroderces argyrogrammos (Zeller)°
*
'** Abundant (>30 specimens emerged). ** Low abundance (5-29 specimens emerged). * Very low abun- dance (1-4 specimens emerged). P = species present (see text for explanation).
> Identity of 4. helianthi was checked by comparison with specimens identified by Dr. R. H. Foote, former Research Entomologist, Systematic Entomology Laboratory, HBIII, USDA, Beltsville, Maryland. Other te- phritids were identified by Dr. I. M. White, C.A.B. International Institute of Entomology, London, England.
‘Identified by E. Colonnelli, Dipartimento di Biologia Animale e dell Uomo, Vaile dell Universita, Rome,
Italy.
‘Identified by Dr. M. L. Cox, C.A.B. International Institute of Entomology, London, England. Identified by Dr. R. Madge, C.A.B. International Institute of Entomology, London, England. ° Identified by Dr. R. W. Hodges, Research Entomologist, Systematic Entomology Laboratory, IIBIII, USDA.
three times (Table 1). Samples from each site were pooled to calculate the percentage of heads with insect-damaged seeds.
The relative occurrence of each species and the percentage of heads with damaged seeds and receptacle tissues was assessed for each site by rearing the insects and dissect- ing all of the heads in a laboratory at the BCWLE. A species was artibrarily rated as abundant, low in abundance, or very low in abundance according to the number of emerging adults (see Table 2). Reared in- sects were identified to species. Because less than 10% of the eggs of B. orientalis survive to the adult stage (Sobhian and Zwolfer
1985, Clement and Sobhian, unpub. data), we assumed that very few, if any, adults would be reared-out. Thus, we recorded the presence or absence of this weevil at each site by looking for its eggs, which are usually laid singly on leaflets near a flower bud (Bu 1-2 stages of Maddox 1981) and are covered by a characteristic black cap.
RESULTS AND DISCUSSION
The percentage of heads with insect-dam- aged seeds varied from 13.75-37.89% (av- erage of 20.43% of heads attacked) in Italy and 10.41-65.91% (average 34.38%) in Greece; less than 20% of the heads were
50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
damaged at 4 Italian and 3 Greek sites (Ta- ble 1). Species packing (no. of species) per site ranged from 2-6 (average 3.67) in Italy and 2-5 (average 3.0) in Greece (Table 2). In contrast, Sobhian and Zwéolfer (1985) re- ported average levels of resource utilization (% of heads attacked) and species packing of 36.9% and ‘tabove” 4 species for Italy, and 77.3% and 9.5 species for Greece, but these higher levels were based on samples from the south-Italian mainland and Sicily, and northern Greece. The average levels re- ported above for central Greece are com- parable to the ones Sobhian and Zwéolfer (1985) reported for Yugoslavia, Bulgaria and Romania (35.6% and 3.4 species). Thus, the collective evidence suggests that predispers- al seed predation by YST flower head in- sects is not markedly high in many areas of southern Europe, including Greece where Sobhian and Zwolfer (1985) and Zwéolfer (1985) reported that average levels of re- source utilization and species packing were significantly higher than they were in the western Mediterranean (Italy and France). High rates of parasitization of several species (Sobhian and Zwélfer 1985) might account for the fairly low levels of resource utiliza- tion in many areas.
In all, we found 11 species of seed pred- ators to be associated with YST heads (Ta- ble 2). Three of these, Acanthiophyllus he- lianthi (Rossi) (Diptera: Tephritidae), Lasioderma sp. nr. haemorrhoidale (Illiger) (Coleoptera: Anobiidae) and Pyroderces ar- gyrogrammos (Zeller) (Lepidoptera: Cos- mopterigidae) use plants in several genera as hosts, and 8, B. orientalis, E. hirtus, L. curtus (Coleoptera: Curculionidae), Terellia sp., U. quadrifasciata, U. jaculata Rondani, Chaetorellia sp. nr. carthami Stack. (Dip- tera: Tephritidae), and Bruchidius tubercu- latus (Hochhut) (Coleoptera: Bruchidae) ap- pear to be restricted to the genus Centaurea in the field (Sobhian and Zwéolfer 1985, Clement, unpub. data). Urophora jaculata was the most ubiquitous and abundant species, but this tephritid is not a candidate
biocontrol agent because it will not develop in the heads of U.S. forms of YST (White and Clement 1987). Chaetorellia sp. nr. car- thami was widespread in Italy; however, this species will form hybrids with C. carthami Stack., a pest of cultivated safflower, so its safety as a biocontrol agent has been ques- tioned by biocontrol workers (Sobhian and Zwolfer 1985). A third stenophagous species (i.e. restricted to Centaurea spp.), B. tuber- culatus, has been disqualified because adults were found in the heads of cultivated saf- flower, Carthamus tinctorius L., in northern Greece (Sobhian and Zwé6lfer 1985). Four of the 6 species that we set out to find were detected; U. quadrifasciata and Terellia sp. were represented in the guild of flower head insects in Italy while EF. hirtus and L. curtus were detected in Greece. None of these 4 species were abundant at any site (Table 2).
The failure of this survey to detect U. sirunaseva and C. hexachaeta was unex- pected because Sobhian and Zwo6lfer (1985) reported that both species occur throughout much of southern Europe. However, recent taxonomic studies (White and Clement 1987, White in press) have revealed a more restricted distribution for these tephritids. This new information on U. sirunaseva and C. hexachaeta, together with information from this survey and the literature on E. hirtus (Ter-Minasyan 1967, Fremuth 1982, Sobhian and Zwolfer 1985) and L. curtus (Zwolfer et al. 1971, Fremuth 1982, Sob- hian and Zwo6lfer 1985) suggest that none of these 4 species are rare but the 2 weevil species are better able to exploit YST over a much wider geographical area than are the 2 tephritid species. The seemingly broad eco- climatic tolerances of EF. hirtus and L. curtus would improve their chances for establish- ment in the western U.S. where YST occurs in markedly different climatic and vegeta- tional zones (Maddox 1981, Maddox et al. 1985, Maddox and Mayfield 1985, Roché et al. 1986).
In summary, this survey has enabled us to: pinpoint sites where biocontrol special-
VOLUME 90, NUMBER |
ists might be able to collect insects of 4 po- tential agents for use in host-specificity tests; clarify the Palearctic distribution of several YST flower head insects, including 6 species that are promising biocontrol agents; and contribute towards a better understanding of the extent to which YST flower heads are attacked by insects in southern Europe.
ACKNOWLEDGMENTS
We are grateful to the taxonomic spe- cialists (listed in Table 2) who identified specimens, I. White, D. Maddox and an anonymous reviewer for comments on the manuscript, and S. Craig for typing the manuscript.
LITERATURE CITED
Fremuth, J. 1982. Cleoninae aus der Turkei und den angrenzenden gebieten (Coleoptera: Curculioni- dae). Fragm. Entomol., Roma 16(2): 239-258.
Maddox, D. M. 1981. Introduction, phenology, and density of yellow starthistle in coastal, intercoast- al, and central valley situations in California. Agric. Res. Serv. U.S. Dept. Agric. ARR-W-20, July. 33 pp.
Maddox, D. M., A. Mayfield, and N. H. Poritz. 1985. Distribution of yellow starthistle (Centaurea sol- stitialis) and Russian knapweed (Centaurea re- pens). Weed Science 33: 315-327.
Maddox, D. M. and A. Mayfield. 1985. Yellow star- thistle infestations are on the increase. Calif. Agric. 39(1 1-12): 10-12.
Maddox, D. M. and R. Sobhian. 1987. Field exper- iment to determine host specificity and oviposi- tion behavior of Bangasternus orientalis and Ban- gasternus fausti (Coleoptera: Curculionidae), biological control candidates for yellow starthistle
51
and diffuse knapweed. Environ. Entomol. 16: 645- 648.
Maddox, D. M., R. Sobhian, D. B. Joley, A. Mayfield, and D. Supkoff. 1986. New biocontrol agent in- troduced for yellow starthistle. Calif. Agric. 40(1 I- 12): 4-5.
Roché, B. F., G. L. Piper, and C. J. Talbott. 1986. Knapweeds of Washington. Coop. Ext. Bull. 1393. Washington State Univ., Pullman, WA. 41 pp.
Sobhian, R. and H. Zwolfer. 1985. Phytophagous insect species associated with flower heads of yel- low starthistle (Centaurea solstitialis L.). Z. Ang. Ent. 99: 301-321.
Ter-Minasyan, M. E. 1967. Zhuki-dolgonosiki pod- semeistra Cleoninae fauny SSSR. Tsvetozhily 1 stebleedy (Triba Lixini). Nauka Publishers, Len- ingrad. 141 pp.
White, I. M. and S. L. Clement. 1987. Systematic notes on the Urophora Robineau-Desvoidy (Dip- tera: Tephritidae) species associated with Centau- rea solstitialis L. (Asteraceae, Cardneae) and other Palearctic weeds adventive in North America. Proc. Entomol. Soc. Wash. 89: 571-580.
White, I. M. In press. Taxonomic problems in the genus Chaetorellia Hendel. /n Cavalloro, R., ed., Proc. CEC-IOBC Int. Symp. on Fruit Flies of Eco- nomic Importance, 7-10 April 1987, Rome, Italy. Balkema, Rotterdam.
Zwolfer, H., K. E. Frick, and L. A. Andres. 1971. A study of the host plant relationships of European members of the genus Larinus (Col.: Curculioni- dae). Tech. Bull. Commonw. Inst. Biol. Control, No. 14, pp. 97-143.
Zwolfer, H. 1965. Preliminary list of phytophagous insects attacking wild Cynareae (Compositae) in Europe. Commonw. Inst. Biol. Contr. Tech. Bull., No. 6, pp. 81-154.
Zwolfer, H. 1985. Insects and thistle heads: Resource utilization and guild structure, pp. 407-416. Jn DelFosse, E. S., ed., Proc. VI Int. Symp. Biol. Contr. Weeds, 19-25 August 1984, Vancouver, Canada. Agric. Can.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 52-54
UNUSUAL OVIPOSITION BEHAVIOR ON EVERGREEN AZALEA BY THE ANDROMEDA LACE BUG STEPHANITIS TAKEYAI (DRAKE AND MAA) (HETEROPTERA: TINGIDAE)
JOHN W. NEAL, JR.
Florist and Nursery Crops Laboratory, Horticultural Science Institute, Agricultural Research Service, U.S. Department of Agriculture, Building 470, Beltsville, Maryland
20705.
Abstract.—The andromeda lace bug, Stephanitis takeyai (Drake and Maa), an adventive tingid from Asia, has a feeding host range in North America of twelve species in the Lauraceae, Salicaceae, Ericaceae and Styracaceae. Hybrid evergreen azalea (Rhododendron sp.) is reported as a new feeding and breeding host. Females oviposit in the midrib of the azalea leaf, a behavior different from that on its preferred host, Japanese andromeda (Pieris japonica). A comparison is made of oviposition site behavior between the androme- da lace bug, and S. pyrioides (Scott) the azalea lace bug.
Stenophagous lace bugs in North Amer- ica have been assigned common names based on the major reproductive hosts. Mi- nor feeding hosts usually involve congeneric species. Drake and Ruhoff (1965) report that ‘All species [of lace bugs] are rather highly specialized in their food habits, and gen- eration after generation live on the same kind of plant or closely related ones.”’ Host plants of the andromeda lace bug Stephani- tis takeyai (Drake and Maa) (= S. globuli- fera Matsumura), an adventive tingid from Asia (Schread 1953), have been reported by Drake and Ruhoff (1965), Schread (1953), Dunbar (1974), and Wheeler (1977) and world-wide include 12 species in Lauraceae, Ericaceae, Salicaceae and Styracaceae. Bai- ley’s (1951) review of New England tingids contains a report that a few S. takeyai were found in association with azalea lace bug, S. pyrioides (Scott), on a single deciduous azalea Rhododendron sp. in Connecticut. Bailey (1974) also reported S. takeyai on Rhododendron calendulaceum (Michx.)
Torr. Wheeler (1977) reported specimens in the U.S. National Museum of Natural His- tory from azalea at a nursery in Falls Church, Virginia, June 1969.
Dunbar (1974) reported that S. takeyai oviposits on the abaxial leaf surface, usually along the side of the midrib of the Japanese andromeda Pieris japonica(Thunb.) D. Don, and that overwintering eggs are found along the midrib but on occasion were distributed over the undersurface of the leaf.
I observed adults of S. takeyai in low numbers on evergreen and deciduous aza- leas throughout Prince George’s and How- ard counties, Maryland. These observations suggested that azalea is used by S. takeyai as a minor host, and prompted this study and report of insect behavior, host fitness and potential pest status.
MATERIALS AND METHODS
Containerized Japanese andromeda har- boring all stages of S. takeyai were pur- chased from a retail nursery in Burtonsville,
VOLUME 90, NUMBER |
te
Fig. 1.
shy ng “4
(A) Frass-covered eggs of S. takeyai placed lateral to the midvein of an andromeda leaf, (B) Eggs of
S. takeyai placed in the midvein of an azalea leaf, (C) Eggs S. pyrioides placed lateral to the midvein of an azalea
leaf. Arrow identifies one of several frass-covered eggs.
Maryland. Plants were transplanted to 11.6 liter (3 gal) containers, held in a heated greenhouse, and watered as needed. Stems of various lengths were removed periodi- cally to a bottle of water and placed in a gauze covered plexiglass cylindrical cage (32 cm high x 31 cm dia.) in a Sherer walk-in rearing chamber programmed at 26.1 + 1°C, with a photoperiod of 14:10 (L:D).
To test for host preference and acceptabil- ity, stems of the evergreen hybrid azalea ‘Martha Hitchcock’ (Rhododendron mu- cronatum x Shinnyo-no-tsuki) were placed in the cylinder contiguous to the infested andromeda cuttings. Leaves with adults and nymphs of S. takeyai found on the azalea during daily observations were transferred for isolation to a similar cage containing an uninfested second azalea cutting. Trans- ferred adults were allowed to feed and ovi- posit. Water was added to the bottle as needed. Both azalea and andromeda cut- tings were introduced weekly to the cylinder
with andromeda lace bugs, and the transfer of adults from andromeda to azalea was conducted over several weeks.
Leaves with eggs of S. takevai were com- pared against leaves with eggs of S. pyrioides removed from the same azalea cultiver used as host plants to maintain a greenhouse col- ony.
RESULTS
Tingids that leaf feed in an inverted po- sition defy gravity and deposit their fecal material on the abaxial surface at random. Preliminary tests confirmed that by remov- ing all frass by rinsing the leaf with warm running water, S. takeyai and S. pyrioides confined their egg laying either in or adja- cent to the midrib or at major lateral veins depending on host. Defecation by the fe- male on the operculum leaves a prominent mark at the egg site (Fig. 1, arrows). Stepha- nitis takeyai fed and oviposited on azalea cuttings. Originally some eggs on this host
54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
hatched, but most failed to produce second instars. Washing leaves prior to hatch to remove sticky plant exudate greatly reduced first instar mortality; nymphs developed to the adults when foliage was washed. The presence of plant exudates on the cuttings is a phenomenon due to the growing of aza- leas in the protective greenhouse environ- ment. Exudates are normally reduced by rain or rendered non-sticky by the accumulation of airborne particles.
Stephanitis takeyai oviposited only in and along the midrib in all azalea leaves ob- served (Fig. 1B), whereas S. pyrioides de- posited its eggs below and lateral to the mid- rib (Fig. 1C). Oviposition by S. takeyai on Japanese andromeda included periodic placement below and lateral to the midrib, but eggs were never inserted in the midrib (Fig. 1A). Andromeda leaves have conspic- uous midribs, but they are not raised as in other hosts such as azalea, possibly account- ing for this change in oviposition behavior. Wheeler (1977) found S. takeyai eggs in- serted in the midvein of spicebush, Lindera bezoin (L.) Blume, and sassafras, Sassafras albidum (Nutt.). This ovipositional site preference by the andromeda lace bug on azalea and other hosts is unusual, because when compared, the andromeda lace bug Oviposits on andromeda lateral to the un- raised midrib which is similar to oviposi- tion by the azalea lace bug below and lateral to the raised midrib on azalea. This behav- ior by S. takeyvai on azalea raises the ques- tion what would be the oviposition site on andromeda if the midrib were raised.
These findings determined that an ever- green azalea can be a suitable feeding-breed- ing host for S. takeyai. Further, it was found that females oviposit in and along the mid- vein of the azalea leaf which is different than when on andromeda. Fertilized eggs hatched and nymphs developed to adults normally. These results suggest that S. takeyai could develop to be a late season threat to azalea production. There is, however, no data to suggest that eggs of S. takeyai overwinter on azalea.
ACKNOWLEDGMENTS
Thanks are extended to Richard C. Froeschner (USNM) and Thomas J. Henry (USDA, SEL) for constructive comments and helpful suggestions on an early draft of the manuscript.
LITERATURE CITED
Bailey, N. S. 1951. The Tingoidea of New England and their biology. Entomol. Am. 31: 1-133. Bailey, N. S. 1974. Additional notes on Stephanitis takeyai in New England (Heteroptera: Tingidae).
Psyche 81: 534-538.
Drake, C. J. and F. A. Ruhoff. 1965. Lacebugs of the World: A catalogue (Hemiptera: Tingidae). Bull. U.S. Natl. Mus. 243: 1-634.
Dunbar, D. M. 1974. Bionomics of the andromeda lacebug Stephanitis takeyai, pp. 277-289. In Mem. Conn. Entomol. Soc. 25th Ann.
Schread, J. C. 1953. Control of the andromeda lace bug Stephanitis globulifera and the holly leaf miner Phytomyza ilicis. Conn. Agric. Exp. Stn. Bull. 568: 1-13.
Wheeler, A. G., Jr. 1977. Spicebush and sassafras as new North American hosts ofandromeda lace bug, Stephanitis takeyai (Hemiptera: Tingidae). Proc. Entomol. Soc. Wash. 79: 168-171.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 55-61
A REVIEW OF THE SOUTH PACIFIC GENUS AUSTROMEGALOMUS ESBEN-PETERSEN (NEUROPTERA: HEMEROBIIDAE) WITH A DESCRIPTION OF A NEW SPECIES FROM RAPA
JOHN D. OSwALD
Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York
14853-0999.
Abstract. —The hemerobiid genus 4Austromegalomus Esben-Petersen is reviewed. Aus- tromegalomus and its type species 4. brunneus are redescribed and A. insulanus is de- scribed as new. Figures, known distributions and a key to the two recognized species are provided. Several shared characters of the male ectoprocts, mediuncus and parameres suggest that the genera Austromegalomus, Conchopterella and Drepanacra are closely
related.
The genus Austromegalomus Esben-Pe- tersen, 1935, was proposed to accommo- date the single species 4. brunneus Esben- Petersen which was described in the same paper from three male specimens collected on the South Pacific island of Tahiti. Until now no additional specimens or species of Austromegalomus have been recorded in the literature. In this paper Austromegalomus insulanus is described as new, from 20 spec- imens collected on the island of Rapa lo- cated approximately 1200 km (750 mi.) SSE of Tahiti, and the genus Austromegalomus and the male of 4. brunneus are redescribed.
As with many early hemerobiid descrip- tions, the original descriptions of Austro- megalomus and A. brunneus are based al- most entirely upon venational characters. Austromegalomus insulanus is shown here to exhibit a wide range of intraspecific vari- ation in a variety of forewing venational traits and venation is judged inadequate to confidently separate the two species. The descriptions presented here emphasize characters of the male genitalia.
Intraindividual, as well as interindividu- al, variation in venational characters is
common in Austromegalomus. In tabulat- ing the variability of several forewing ve- national traits, both forewings of each in- dividual were scored for each trait. Estimates of mean forewing length were based on mea- surements of a single forewing of each spec- imen. Consequently, the sample sizes given in the species descriptions for venational traits are twice those given for estimates of mean forewing lengths.
Austromegalomus Esben-Petersen
Austromegalomus Esben-Petersen, 1935: 139. Type species: Austromegalomus brunneus Esben-Petersen, 1935: 140, by original designation.
Diagnosis. — Head: Temporal sutures well developed, marked internally by prominent costae; epicranial suture absent; labial palp three segmented, distal segment longest and with an apical subsegment, palpimacula present; maxillary palp five segmented, dis- tal segment longest and with an apical sub- segment.
Forewing: Length 5-9 mm, hind margin rounded, apex broadly pointed; costal area
56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. the first (proximal) oblique branch of the forewing radius (diagrammatic). 4-6. Female. 4, apex of abdomen (lateral view). 5, spermatheca and apex of bursa. 6, subgenitale (ventral view).
broad proximally, recurrent vein pectinate- ly branched; proximal half of subcostal space with 2 crossveins (the distal of these rarely absent); radius with 4-10 oblique branches; 2 well developed, posteriorly convergent, gradate series in outer half of wing.
Hindwing: Radius with 2 oblique branch- es; Cu2 frequently, though not always, traceable to near the posterior margin either as a distinct or indistinct vein or a row of setae; Outer gradate series well developed; inner gradate series with 1-3 crossveins or absent.
Male genitalia: Tergite nine a sclerotized arch, lateral lobes dilated ventrally; sternite nine in ventral view a remiform plate (Figs. 7, 13), shallowly arched in anterior view; ectoproct elongate oval, without narrowed projecting lobes; gonarcus with arms of
Austromegalomus insulanus. 1, venation of forewing and hindwing. 2-3, two possible states of
moderate size the greater part of which pro- ject free into body cavity, exposed surface limited to a narrow strip to which the medi- uncus and epimeres are fused; mediuncus a rigid plate dorsal to epimeres and para- meres, bilobed proximally and distally, lon- gitudinal midline shallowly depressed, proximal lobes strongly divergent and fused to gonarcus at a pair of widely separated points on opposite sides of gonarcus bridge, medial margins of proximal lobes and pos- terior margin of gonarcus bridge enclosing a triangular membranous fenestra; epimeres a pair of elongate strips of sclerotized mem- brane lying in the membranous sack sup- ported dorsally by the mediuncus, fused to gonarcus ventral to fusion of mediuncus with gonarcus; parameres with internal end of apophysis proxima enlarged in lateral view,
VOLUME 90, NUMBER 1
.2mm
[smn
9-12,15-18
7-8,13-14
Figs. 7-18. 7-12. Austromegalomus brunneus, male. 7, ninth sternite (ventral view). 8, apex of abdomen (lateral view). 9, gonarcus, mediuncus and epimere (lateral view). 10, gonarcus, mediuncus and epimere (dorsal view). 11, parameres (dorsal view). 12, parameres (lateral view). 13-18. 4. insulanus, male. 13, ninth sternite (ventral view). 14, apex of abdomen (lateral view). 15, gonarcus, mediuncus and epimere (lateral view). 16, gonarcus, mediuncus and epimere (dorsal view). 17, parameres (dorsal view). 18, parameres (lateral view).
external lobes narrow and linear in dorsal view but with their apices upturned in lat- eral view, medioventral surfaces of lobes sclerotized, laterodorsal aspects membra- nous.
Female genitalia: See below under A. In- sulanus.
Natural history and immature stages. — Unknown.
Distribution.—Known only from the French Polynesian islands of Tahiti and Rapa.
Etymology.—Name unexplained but al- most certainly from the Latin “australis,”
southern, and “.Wegalomus,” a hemerobud genus to which Esben-Petersen allied Aus- tromegalomus. Gender: masculine. Discussion.—Esben-Petersen diagnosed Austromegalomus by the branching ar- rangement of the first oblique branch of the radius (‘basal Rs” of Esben-Petersen). In Austromegalomus the vein track which an- teriorly parallels the median flexion line is nearly straight. The curvature of vein seg- ments confluent at forks along this track are usually somewhat asymmetric. The branch- es originating at these forks tend to form a linear series on the anterior side of the track
58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(Fig. 2). In an alternate state found in many other hemerobiid genera, the vein forks along this track are more symmetric giving the track a more or less undulate appearance (Fig. 3). Though these states are rather dis- tinctive when viewed as the opposite ends of a morphocline, intraspecific variation within A. insulanus encompasses both states. Too few specimens are available to ade- quately assess the degree of intraspecific variation of this trait in 4. brunneus. Fur- thermore, as pointed out by Esben-Petersen (1935) and Handschin (1955), similar asymmetric patterns of veins along this track are found in other hemerobiid genera (e.g. Drepanacra and Conchopterella). For these reasons this character cannot be used as a synapomorphy of Austromegalomus. Due to still unresolved questions concerning the homologies and polarities of diagnostic characters of the male genitalia, I have been unable to confidently identify any synapo- morphic characters for this taxon.
Key TO ADULT MALES OF AUSTROMEGALOMUS la. Emargination separating distal lobes of medi- uncus V-shaped (Fig. 16); gonarcus bridge ar- cuate in dorsal view (Fig. 16); epimeres short and narrow and not subtended by an acces- sory sclerite (Fig. 15) (Rapa) A. insulanus n. sp. 1b. Emargination separating distal lobes of medi- uncus quadrate (Fig. 10); gonarcus bridge quadrate (Fig. 10); epimeres long and broad and subtended distally by a small accessory sclerite (Fig. 9) (Tahiti) : .. A. brunneus Esben-Petersen
Austromegalomus insulanus, NEw SPECIES Figs. 1-6, 13-18
Diagnosis.— Diagnosed by characters in key couplet la. The longer forewing length and the alternating light and dark brown segments of the forewing longitudinal veins may also be diagnostic, though not enough specimens of A. brunneus are available to fully assess the potential overlap of these characters with those found in A. insulanus.
Description. — Forewing (Fig. 1): Length
6.09-8.48 mm (x = 7.06, N = 20); longi- tudinal veins mostly marked with alternat- ing light and dark brown segments, though several specimens (likely teneral) with ve- nation nearly evenly pale; membrane hya- line to brown, darker adjacent to dark vein segments. Venation (Fig. 1, N = 40): Num- ber of subcostal crossveins in proximal half of subcostal space = 2 (39 wings), 3 (1 wing); number of oblique radial branches proximal to stigmal subcostal crossvein = 6 (1 wing), 7 (6 wings), 8 (16 wings), 9 (14 wings), 10 (3 wings); number of inner gradate cross- veins anterior to cubitus = 10 (3 wings), 11 (7 wings), 12 (20 wings), 13 (8 wings), 14 (2 wings); number of outer gradate crossveins anterior to cubitus = 11 (2 wings), 12 (2 wings), 13 (8 wings), 14 (15 wings), 15 (13 wings); number of forkings of first oblique radial branch proximal to inner gradate se- ries = | (7 wings), 2 (29 wings), 3 (4 wings).
Male genitalia: Apex of abdomen as in Fig. 14. Gonarcus (Figs. 15, 16): gonarcus bridge arcuate in dorsal view. Mediuncus (Figs. 15, 16): distal pair of lobes contiguous medially at their bases. Epimeres (Figs. 15, 16): short and very narrow, weakly tanned and easily overlooked; not extending as far posteriorly as in 4. brunneus and not sub- tended distomedially by a pair of accessory sclerites. Parameres (Figs. 17, 18): internal end of apophysis proxima enlarged but not as prominently as in 4. brunneus; apices of external lobes tipped with a minute spine.
Female genitalia (Figs. 4, 5, 6): Gon- apophyses laterales remiform, styli arising dorsad of middle of sclerites; gonapophyses posteriores present as a pair of narrow rods; subgenitale present, attached to ventral body wall by a short membranous tube, apex emarginate; spermatheca composed of a darkly tanned bulb and a pair of ducts—a short duct joining the bulb to the bursa and a longer convoluted duct arising from the distal end of the bulb.
Etymology. —An adjective from the Latin “insula,” island, in reference to the island type locality.
VOLUME 90, NUMBER 1
Distribution.— Known only from the type series from the South Pacific island of Rapa (French Polynesia, Austral Islands).
Primary type material examined.— Male holotype (USNM). Verbatim label data: “Rapa/Anatakuri/Bay 28 XI 63,” “J. F. G. Clarke/Thelma M. Clarke,” ““USNM Loan/ USNM Loan,” ‘“Holotype/Austromegalo- mus/insulanus Oswald/J. D. Oswald 1987.” Condition: Excellent, no parts missing. Genitalia cleared and placed in a glycerin filled microvial pinned below the specimen.
Other material examined.— 19 paratypes. RAPA ISLAND: 2 6, Anatakuri Bay, 28.xi.1963 (Clarke) (USNM); 3 4, 2 2, Haur- ei, 15.x.-3.xii.1963 (Clarke) (USNM); | 4, 1 2, Mau Bay, 23.x.1963 (Clarke) (USNM); 2 2, Maugaoa, 244 m & 290 m, 18.1x.- 23.x1.1963 (Clarke) (USNM); | 4, 1 2, Man- gaoa [sic = Maugaoa] Pk., NE ridge, 305- 366 m, 6.vil.1934 (Zimmerman) (BPBM); 1 6, 2 2, Maurua, 61 m & 183 m, 25.1x.- 25.x.1963 (Clarke) (USNM); | 2, Mt. Oror- angi, SE valley, 183-244 m, 3.vu.1934 (Zimmerman) (BPBM), 2 2, Point Teak- aurae, 61 m, 7.x.1963 (Clarke) (USNM).
Note.— For a general account of the Clarke Expedition to Rapa, including collecting lo- calities and physiography, see Clarke (1971).
Austromegalomus brunneus Esben-Petersen Figs. 7-12
Austromegalomus brunneus Esben-Peter- sen, 1935: 140 (original description, fig- ures): Esben-Petersen 1937: 51 (listed); Handschin 1955: 9 (compared to Con- chopterella).
Diagnosis.— Diagnosed by characters in key couplet 1b. The shorter forewing length and the uniform brown coloration of the forewing longitudinal veins may also be di- agnostic, though not enough specimens of A. brunneus are available to adequately as- sess the potential range of intraspecific vari- ation in these characters.
Description. — Forewing: Length 5.37- 5.56 mm (x = 5.47, N = 2); longitudinal
59
veins uniformly brown, membrane also brown. Venation (N = 4): Number of sub- costal crossveins in proximal half of sub- costal space = 2 (4 wings); number of oblique radial branches proximal to stigmal sub- costal crossvein = 4 (3 wings), 5 (1 wing); number of inner gradate crossveins anterior to cubitus = 8 (1 wing), 9 (1 wing), 10 (2 wings); number of outer gradate crossveins anterior to cubitus = 11 (2 wings), 12 (2 wings); number of forkings of first oblique radial branch proximal to inner gradate se- ries = 3 (4 wings).
Male genitalia: Apex of abdomen as in Fig. 8. Gonarcus (Figs. 9, 10): gonarcus bridge quadrate in dorsal view; anterodorsal region of gonarcus arm broader in lateral view than in A. insu/anus. Mediuncus (Figs. 9, 10): distal pair of lobes separated medi- ally at their bases by a space about equal to width of each lobe. Epimeres (Fig. 9, 10): prominent, long and broad relative to 4. insulanus; apex of each epimere subtended medially by a small, weakly sclerotized and poorly delimited accessory sclerite. Para- meres (Figs. 11, 12): internal end of apoph- ysis proxima considerably enlarged in lat- eral view; apices of external lobes tipped with a minute spine.
Female: Unknown.
Etymology.—An adjective from the Latin “brunneus,” dusky or tawny, in reference to the brownish coloration of the body and forewing.
Distribution.— Known ony from the type series from the South Pacific island of Tahiti (French Polynesia, Society Islands).
Primary type material examined.— Male holotype (BPBM). Verbatim label data: “Society Is./1500'/Tahiti I.,” *“Fautaua Val./ IX-11-28,° “A. M. Adamson/Collector,” “Pacific Entomological Survey,” “TYPE 791,” “Austromegalo=/mus brunneus/é n. sp./det. Esben-Petersen.”’ Condition: Ex- cellent, only several small pieces of wings missing. Genitalia cleared and placed in a glycerin filled microvial pinned below the specimen.
60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Other material examined.—One male paratype (BPBM). Collection data same as holotype. A second paratype stated in the original description to have been retained by Esben-Petersen has not been traced.
PHYLOGENETIC POSITION OF AUSTROMEGALOMUS
The morphology of the male genital struc- tures of Austromegalomus suggests that it is closely related to the southern hemisphere genera Drepanacra and Conchopterella, which are known from the Australian region and the Juan Fernandez islands respective- ly. This evidence supports the conclusions of Esben-Petersen (1935) and Handschin (1955) based on venational characters. The following three shared traits appear to sup- port the hypothesis that these genera are closely related:
(1) The male ectoprocts are elongate oval without projecting narrow lobes. The ec- toprocts of many other hemerobiid genera are variously lobed.
(2) The mediuncus forms a rigid horizon- tal plate which is bilobed distally and at- tached to the gonarcus by a pair of widely divergent proximal arms. The full distri- bution of this state and its possible deriv- atives within the Hemerobiidae needs ad- ditional investigation.
(3) The parameres consist ofa prominent, anteriorly projecting apophysis proxima and a pair of small apical lobes. The medioven- tral surfaces of the apical lobes are sclero- tized, the dorsolateral surfaces membra- nous. The parameres of many other hemerobiid genera possess various other dorsal and/or lateral lobes and patterns of sclerotization.
Though one or more of the preceding characters may in the future prove synap- omorphic of a clade (Austromegalomus + Conchopterella + Drepanacra), at present, the polarities of these shared traits in rela- tion to their homologues found in other hemerobiid genera are not known with con- fidence. Consequently, firm conclusions
about the relative relationships among these three genera are presently impossible.
Currently available comparative analyses of important hemerobiid character com- plexes (e.g. wing venation and male geni- talia) are in most cases insufficiently de- tailed, with regard to hypotheses of homologies and/or polarities, to allow con- fident differentiation of synapomorphies and symplesiomorphies. Consequently, it has not been possible to fully assess the status of some Austromegalomus character states which might later prove to be useful indi- cators of phylogenetic relationships.
Several factors have contributed to the dilemma described above. First, no recent comprehensive revision of the Hemerobi- idae, with attention to character analysis, is available. Second, many terms widely em- ployed in the current nomenclature of neu- ropterous genital structures were originally proposed expressly as labels of convenience, without critical investigation of the homol- ogies of the labeled structures. Though some of these terms have apparently been applied to homologous structures (e.g. the gonar- cus), others have not (e.g. the mediuncus- arcessus). Uncritical application of existing genitalic terms has hindered the improve- ment of hypotheses of homology for some genitalic structures.
Most hemerobiid genera are currently di- agnosed, at least in part, on the basis of distinctive combinations of male genitalic characters. Given the importance of this character complex, additional comparative morphological studies are needed to clarify the homologies and polarities of genitalic characters. Until such analyses are under- taken, the phylogenetic position of Austro- megalomus, and many other hemerobiid genera, will likely remain unclear.
ACKNOWLEDGMENTS
I thank Oliver S. Flint, Jr., of the National Museum of Natural History (USMN), Washington, D.C., and Gordon M. Nishida of the Bernice P. Bishop Museum (BPBM),
VOLUME 90, NUMBER 1 61
Honolulu, Hawaii, for loaning material used | Esben-Petersen, P. 1935. Neuroptera from the So- in this study. I also thank James K. Liebherr ciety Islands. Bull. Bernice P. Bishop Mus. 142:
d 137-142. and Quentin D. Wheeler, Department of 1937. Check List of Neuroptera Planipennia
Entomology, Cornell University and an of Oceania. Occas. Pap. Bernice P. Bishop Mus. anonymous reviewer for providing com- 13: 49-60. ments on an earlier draft of this paper. Handschin, E. 1955. Los insectos de las Islas Juan Fernandez. 15. Neuroptera. Rev. Chil. Entomol. LITERATURE CITED 4: 3-20.
Clarke, J. F. G. 1971. The Lepidoptera of Rapa Is- land. Smithson. Contrib. Zool. 56: 1-282.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 62-65
LECTOTYPE DESIGNATION FOR EMPIS CHICHIMECA WHEELER AND MELANDER (DIPTERA: EMPIDIDAE)
WILLIAM J. TURNER
Department of Entomology, Washington State University, Pullman, Washington 99164.
Abstract.— A lectotype and six paralectotypes for Empis (= Lamprempis) chichimeca Wheeler and Melander are designated from the syntype series. Diagnostic leg features of the lectotype male are discussed and illustrated. Comments concerning the lectotype and
its presumed detached hind leg are provided.
The Neotropical genus Lamprempis Wheeler and Melander presently includes 22 species of metallic greenish blue to black flies with an evanescent anal wing vein and peculiarly ornamented legs. The often di- morphic sexes show presence or absence of pennate hair fringes and other modifica- tions of the legs. Several species are known from one sex only. Little information is available about the biology and habits for species of Lamprempis. Smith (1975) re- ports that one species, L. sazimae, occurs in great numbers in the highlands of Minas Gerais, Brazil, where it serves as an impor- tant pollinating agent for certain Umbelli- ferae and Eriocaulaceae growing in mead- ows at 1300 m above sea level.
The purpose of this paper is to report the interesting results of my study of the avail- able syntype series for Empis chichimeca Wheeler and Melander (1901: 368), the type species of Lamprempis, and to designate a lectotype for this species.
In 1981, while examining the A. L. Me- lander types of Empis Linnaeus at the Na- tional Museum of Natural History (USNM), I found several syntype specimens of E. chi- chimeca. The original series consisted of nine specimens (two males and seven females) collected by H. H. Smith in Amula, Guer- rero, Mexico. I could find only three female
specimens in the USNM type collection. Also present was the apparent right hind leg of a male, glued to a card rectangle and la- beled “type” in Melander’s hand. It had been attached on its anterior side and embedded in an unknown adhesive, but the characters of the exposed posterior surface are easily visible and match the description of the species provided by Wheeler and Melander (1901). At that time I supposed that the leg, which possesses characters sufficient for rec- ognizing the species, was the only portion remaining of one male, the remainder de- stroyed by pests or otherwise lost.
Later, in the collection of the American Museum of Natural History (AMNH), I dis- covered another part of the same syntype series (one male, three females). All speci- mens are in good condition. Interestingly, I found that the AMNH male is intact except for the missing right rear leg. After re-ex- amining the USNM point-mounted leg, I concluded that it likely represents the miss- ing leg from the AMNH specimen.
There is no indication when the leg of the male syntype was broken or removed from the otherwise intact specimen. One can only speculate why the leg was not kept with the associated male. The detached leg, however, possesses the diagnostic features of the species (see Smith 1975) and it serves as an
VOLUME 90, NUMBER |
a —
ra
Pil
Pes Pare SS ~~ \\ wae
<i
Fig. 1.
excellent reference even without the intact specimen itself. The leg may have permitted Melander to have the ideal “‘reference spec- imen” in his collection while Wheeler had the remainder of the specimen. Because I cannot find the other male in the syntype series, I assume that the detached leg may have served this function for Melander. Because this male and especially its de- tached leg bear the diagnostic features of the species, and nearly intact AMNH male is hereby designated the lectotype of Empis (= Lamprempis) chichimeca and its de- tached right rear leg (in the USNM) 1s sim- ilarly marked with my red lectotype label. I have illustrated the detached right leg (Fig. 1) along with the left hind leg (Fig. 2). The remaining six females of the known syntype series have been labeled paralectotypes. The male specimen (AMNH) selected as
63
2
Right hind leg (detached) of Empis (= Lamprempis) chichimeca, lectotype male, in posterior view.
lectotype is in excellent condition, except for the missing right rear leg, and bears the following label data: ““Amula, Guerrero, 6000 ft., Sept., H. H. Smith/W. M. Wheeler Collection/TYPE NO. | > AMNH [red la- bel]/AMNH, DIZ No. 918 [white label]/ LECTOTYPE, Empis (= Lamprempis) chi- chimeca Wheeler and Melander, des. W. J. Turner 1988 [red label, hand written].” The point-mounted leg (USNM) lacks a locality label but has the following label data: “E. chichimeca W & M TYPE [white label in Melander’s hand]/Cotype Lamprempis chichimeca W & M [red cotype label in Me- lander’s hand]/A L Melander Coll. 1961 [white label with green checked margin]/ LECTOTYPE (part), Empis (= Lamprem- pis) chichimeca Wheeler and Melander, des. W. J. Turner 1988 [red label, hand written].” Besides the lectotype male, I found that
64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2.
only two of the available females (AMNH) bear the same label data. The remaining four females (USNM, AMNH) have iden- tical labels but were collected in August. Only seven specimens (one male, six fe- males) from the original syntype series of nine specimens have been accounted for. The location of one additional male and female remain unknown. Because much of the insect material described in the Biologia Centrali Americana was subsequently de- posited in the British Museum (Natural History) (BM), I asked John Chainey, Cu- rator of Diptera (BM), to check for syntype specimens of this species. He was unable to locate any representatives of chichimeca under either Empis or Lamprempis in the BM collection. Further, there was no ref- erence made to the species in any lists of holdings by the museum. All of the known syntypes, now in either the USNM or AMNH, were originally in the collections of W. M. Wheeler or A. L. Melander re-
/
Left hind leg of Empis (= Lamprempis) chichimeca, lectotype male, in anterior view.
spectively, as indicated by the personal col- lection labels attached to the specimens. Only one USNM female lacks such a label probably because it was placed in that col- lection by the authors shortly after the species was described.
DISCUSSION
In 1901 Empis chichimeca was described by W. M. Wheeler and A. L. Melander, and placed into their new subgenus Lamprempis along with five other species from Mexico. Coquillett (1903) elevated Lamprempis to generic rank and designated EF. chichimeca as the type species. Although the diagnostic features for this species have never been illustrated, the species is easily keyed. Smith (1975) includes E. chichimeca in his ten- tative key to the described species of Lam- prempis and uses essentially the same word- ing as in the original description by Wheeler and Melander for describing the unique fea- tures of the hind leg: “Hind femora pos-
VOLUME 90, NUMBER |
teroventrally with two slender finger-like processes, with an emargination between them; hind tibia posteriorly with a stout scoop-shaped process truncated and flat- tened at the extremity; hind basitarsus in- crassate with an anterior projection tipped with two small black spines.”
The hind legs of this species are somewhat asymmetrical with minor differences in structures from left to right. Similar asym- metry can be found in the armature of the hind legs of males of Empis (Enoplempis) mira Bigot. In the case of E. chichimeca the right femur appears to have a small hooked process on the posterior surface near the base of the larger, digitate process. Proximal to the small hook is a low, irregular carina with toothlike projections running oblique- ly across the subbasal fourth of the hind surface. Unfortunately the leg 1s embedded in an adhesive glue matrix and the edge of the glue follows along the carina. On the left femur, in comparison, the small hook is lacking as is the oblique carina. The de- scription was likely made from the right (de- tached) appendage as it refers to the two, slender, fingerlike processes, probably the thicker digitate process and the small hooked one. I found that the similar digitate pro- cesses located medially on the posteroven- tral margin of both hind femora also differ from left to right in orientation, the left one being more linear, the right more oblique. One will also see from the illustrations that the general outline of each femur is different as well.
Both hind tibiae are moderately concave medially on both the anterior and posterior surfaces along nearly their entire length. The concavities appear natural and not simply artifacts of the legs having collapsed at death. Although the surrounding areas are heavily bristled, the depressed spaces remain essen- tually bare.
Smith (1975) indicates in couplet 14 of his key that E. chichimeca has only simple leg bristles. However, pennate bristles can be found in two irregular rows along the
65
entire dorsal surface of each hind tibia and flanked by fewer, less developed but still flattened bristles. An additional five or six pennate setae can be found at the extreme base of the tibiae ventrally while each femur bears a cluster on its inner and dorsal sur- faces apically. Although pennate leg bristles are not uncommon in females of some Em- pidinae (e.g., Rhamphomyia species), they are unusual in males and appear restricted to Lamprempis.
ACKNOWLEDGMENTS
I thank Lloyd Knutson, Systematic Ento- mology Laboratory, USDA, % National Museum of Natural History, Washington, D.C., and Pedro Wygodzinsky, Department of Entomology, American Museum of Nat- ural History, New York, New York, who made syntype specimens of Empis (Lam- prempis) chichimeca in their respective col- lections available for study. Dr. Knutson also provided work space and optical equip- ment in his office during my sabbatical stay in Washington, D.C. John Chainey at the British Museum (Natural History) checked for additional syntype material and his ef- forts are appreciated. Thanks also to Paul Arnaud, Jr., James Johnson and Norman Woodley who reviewed this manuscript. Scientific Paper No. 7622, Agricultural Re- search Center, College of Agriculture and Home Economics, Washington State Uni- versity, Pullman, Washington. The work was conducted under Project 9043.
LITERATURE CITED
Coquillett, D. W. 1903. The genera of the dipterous family Empididae, with notes and new species. Proc. Entomol. Soc. Wash. 5: 245-272.
Smith, K.G. V. 1975. A new species of Lamprempis Wheeler & Melander from Brazil, with a key to the described species of the genus (Diptera, Em- pididae). Pap. Avulso Zool. (Sao Paulo) 29: 21- 26.
Wheeler, W. M. and A. L. Melander. 1901. Supple- ment. Empididae, pp. 366-367. Jn Godman, F. D. and O. Salvin, eds., Biologia Centrali Ameri- cana. Insecta. Diptera. Vol. |. 378 pp., 6 pls. Lon- don.
PROC. ENTOMOL. SOC. WASH. 90(1), 1988, pp. 66-75
MAYACNEPHIA SALASIT (DIPTERA: SIMULIIDAE), A NEW BLACK FLY SPECIES FROM COSTA RICA
J. RAMiREZ-PEREZ, B. V. PETERSON, AND M. VARGAS V.
(JRP) Instituto de Biomedicina, P.O. Box 4043, Caracas, Venezuela; (BVP) Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % National Mu- seum of Natural History, NHB-168, Washington, D.C., U.S.A. 20560; (MVV) Centro de Investigacion y Diagnostico en Parasitologia, Universidad de Costa Rica, Ciudad Uni-
versitaria Rodrigo Facio, Costa Rica.
Abstract.—The female, male, pupa and larva of Mayacnephia salasi, new species, are described and illustrated. This genus is recorded from Costa Rica for the first time, and is now known from western North America, Mexico, Guatemala, Costa Rica and Panama. A key to the species of Mayacnephia known in the pupal stage is provided.
The genus Mayacnephia Wygodzinsky and Coscar6n (1973) was established to in- clude six Mesoamerican species that had been placed previously in the genus Cnephia Enderlein. Diaz Najera (1971) described another new species in the genus Cnephia that belongs in Mayacnephia, and J. L. Pe- tersen (1985) described a new species from Panama. B. V. Peterson (1981), using an expanded concept of the genus, assigned two species to it from western United States and an undescribed species from Canada. The species described below is the eleventh de- scribed species now assigned to the genus and the first to be reported from Costa Rica. We describe this new species to make its name available for biological studies cur- rently being conducted on black flies in Cos- ta Rica. We also provide a key to the known pupae of Mayacnephia, including the un- described species from Canada, and include distributions and references to published figures of these species.
Mayacnephia salasi Ramirez-Perez, Peterson, and Vargas, New SPECIES Figs. 1-18 Female (preserved in alcohol).— General body color dark brown. Length: body, 2.88 mm; wing, 3.48 mm.
Head: Lightly grayish pollinose. Frons (Fig. 5) narrow, nearly parallel sided, only slightly widening dorsally, about five times as long as width at narrowest point, about , width of head; slightly paler than occiput, densely covered with long, decumbent, pale yellow pile, and with a few black setae lat- erally. Clypeus concolorous or slightly light- er than frons; slightly longer than wide; densely covered with long, ventromedially directed, pale yellow pile interspersed with some dark setae laterally and ventrally. Oc- ciput densely covered with long, pale yellow pile and with a few dark setae mid-dorsally; postocular setae black, closely bending over eye margin. Antenna with nine flagello- meres; scape and pedicel pale yellowish, contrasting with rest of flagellum which is dark brown; pedicel and first flagellomere larger than other antennomeres and sub- equal in length and width; remaining flagel- lomeres subequal in length to each other but tapering in width distally; fine pubescence and longer setae dark. Mandible (Fig. 3) with 33-40 serrations. Blade of maxilla (Fig. 2) with 21-25 retrorse teeth. Palpus (Fig. 1) dark brown to black, proximal two palpo- meres pale brownish, distal two palpomeres slightly lighter brownish than palpomere three, all with black setae; palpomere five about 3 longer than palpomere three. Sen-
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sory vesicle as in Fig. 4, about '2 as long as its segment, proximally situated, its neck short, arising anterodorsally and extending vertically, with an enlarged ovoid mouth. Median proximal space of cibarium shal- low, broadly U-shaped, and without den- ticles; dorsolateral arms short, rather broad, sclerotized, inner surface of each arm with a rather extensive patch of minute setulae arising from dark granular bases.
Thorax: Antepronotal lobe concolorous with scutum, with dense, long, pale yellow pile interspersed with a few dark setae. Post- pronotum yellowish brown, distinctly paler than scutum, covered with long, semi-erect pale yellow pile. Scutum dark brown to blackish brown except lateral margins which are narrowly paler, and with a grayish pol- linose border extending around lateral and hind margins, posterior declivity broadly grayish pollinose; each anterolateral corner of scutum, adjacent to postpronotal lobes, a paler yellowish brown color which, in pos- terior view, extends posteriorly as a faint, narrow, submedian line, and with a similar faint, slender, median line or vitta that ex- tends to posterior declivity, these lines not visible in anterior view; scutum densely covered with short, recumbent, pale yellow pile that is longer along anterior and lateral margins and still longer posteromedially, also a few scattered dark setae present. Scu- tellum paler brownish than scutum, lightly grayish pollinose, densely covered with long, pale yellow and dark setae. Postnotum only faintly darker than scutellum, with a faint pollinosity. Anterior half of pleuron dark brown mottled with some paler areas, and distinctly paler brown on posterior half that is mottled with some darker areas; prester- nal lobe with moderately long pale yellow pile; anepisternal membrane distinctly paler than rest of pleuron; mesepimeral tuft dark. Wing (Fig. 6) membrane hyaline but with a light brownish tinge; veins yellowish brown. Base of C, stem vein, and other veins with dark pile; Sc setose ventrally; R, setose dor- sally; R,,; setose ventrally; cell bm present;
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C with spiniform setae as long as regular setae, apical half of R, with a few similar spiniform setae; fringe of anal lobe and ca- lypter pale yellow. Stem of halter and base of knob brownish yellow, rest of knob yel- low; stem with pale yellow pile. Legs rather uniformly dark brown; all coxae with both pale and dark setae but pale setae more nu- merous on fore and midcoxae, rest of setae on legs dark; hind basitarsus about seven times as long as broad. Calcipala short but distinct, broadly rounded; pedisulcus ab- sent. Claw only slightly curving from base, with a prominent, bluntly pointed, basal tooth that is wider than claw and over '2 as long.
Abdomen: Yellowish brown, basal scale (tergite one) dark brown, with a fringe of long, pale yellow pile; tergites broad, tergite two widest, others decreasing in width pos- teriorly; tergites yellowish centrally with darker brown margins, sparsely covered with short, pale yellow setae; pleural membrane paler and more yellowish brown, with both pale yellow and dark setae; sternites scarcely distinguishable; venter of abdomen pale brownish yellow, with mostly short, dark setae but with some scattered pale yellow setae. Terminalia as in Figs. 7-9. Anal lobe (Fig. 9) narrow dorsally, broadening ven- trally, widest at about midheight, broadly rounded ventrally, with a slight but distinct notch posteroventrally, not produced be- neath cercus, moderately setose. Cercus subrectangular, hind margin varying from strongly rounded to nearly straight. Hypo- gynial valves short, barely reaching to bases of cerci; valves subtruncate posteriorly, their medial margins lightly sclerotized; lightly setose. Stem of genital fork (sternite nine) (Fig. 7) long, heavily sclerotized, slightly more than '3 longer than arms; arms short, rather weakly sclerotized except for a short, heavily sclerotized rodlike extension on each side emanating from stem of genital fork; arm with a sclerotized subtriangular tooth- like process on anterior margin; arms rather broadly attached to tergite nine. Spermathe-
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ca (Fig. 8) kidney-shaped, moderately scler- otized, with a faint, loose reticulate pattern but without internal spicules; with only a small clear area at junction with spermathe- cal duct.
Male (preserved in alcohol).—General body color velvety dark brown to black. Length: body, 3.12 mm; wing, 3.18-3.45 mm.
Head: Frons and clypeus lightly grayish pollinose, with erect, black pile. Occiput with long, dark brown to black setae. Antenna entirely dark brown; first flagellomere an- gularly broadened distally, slightly longer than pedicel: fine pubescence pale yellow, longer setae black. Palpomere three darker than other palpomeres, all with black pile interspersed with a few more yellowish se- tae; palpomere five about 3 longer than pal- pomere three and about '2 longer than pal- pomere four. Sensory vesicle about ' as long as its segment; neck short, enlarging to form a round mouth.
Thorax. Antepronotum and postprono- tum concolorous, slightly paler brown than scutum; with some dark setae and some pale yellow pile having dark bases. Scutum with a light grayish pollinosity; densely covered with short, recumbent, pale yellow pile that is longer anteriorly, laterally and postero- medially. Scutellum brown, paler than scu- tum; densely covered with long, erect, dark setae and some decumbent, pale yellow se- tae. Postnotum concolorous with scutellum, lightly grayish pollinose. Pleuron brown an- teriorly, grayish pollinose, becoming paler brown medially and posteriorly; anepister- nal membrane brownish yellow; mesepi- meral tuft dark. Wing membrane hyaline but with a distinct brownish tinge, veins yellowish brown. Base of C, stem vein, and other veins with dark pile; Sc lightly setose
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ventrally; R, setose dorsally; R,,; setose ventrally; C and about distal 73 of R, with spiniform setae that are about as long as regular setae; cell bm present; fringe of anal lobe brownish yellow; fringe of calypter pale yellow. Knob of halter brown, stem yellow with pale yellow pile. Legs rather uniformly dark brown, with dark pile; hind basitarsus swollen, about 3.5 times as long as broad; calcipala short but distinct, rounded api- cally; pedisulcus absent.
Abdomen: Basal scale dark brown, with a fringe of long dark pile; tergites with dark- ened margins, paler medially, covered with short, brown pile; sternites concolorous with tergites, with long, dark setae. Terminalia as in Figs. 10 and 11. Gonocoxite (Fig. 10) subtriangular to conical, greatest length and width nearly equal, covered with pile on all but basal '4 to /s. Gonostylus short, about '’» longer than greatest width at base; taper- ing to a bluntly pointed, apical margin bear- ing two tiny terminal spines. Body of ventral plate of aedeagus (Fig. 11) subrectangular, broader than long, with a short, ventrally directed hirsute lip; in ventral view, apical margin slightly convex and shortly pro- duced nipplelike medially, lateral margins slightly concave just distal to junction with basal arms; basal arms bowed, nearly equal in length to body of ventral plate; median sclerite of aedeagus short, Y-shaped, stem variably longer than arms; aedeagal mem- brane rather densely covered with numer- ous groups of 8-10 minute setulae arranged in rows. Plate of endoparameral organ an elongate subtriangular shape, moderately sclerotized; arm moderately long, and twist- ing.
Pupa.—Length of specimens at hand 3.5 mm. Respiratory organ (gill) (Fig. 12) 1.62 mm long; consisting of four rather short but
—
Figs. 1-11.
Mayacnephia salasi. Figs. 1-9, female. 1, Maxillary palpus. 2, Blade of maxilla showing retrorse
teeth. 3, Tip of mandible showing serrations. 4, Enlarged view of sensory organ of third palpomere. 5, Front view of frons and ocular notches. 6, Portion of wing showing setation. 7, Genital fork (sternite 9). 8, Spermatheca. 9, Anal lobe and cercus. Figs. 10-11, male. 10, Gonocoxite and gonostylus (dorsal (inner) surface). 11, Ventral
plate of aedeagus, ventral view.
TRAMRE? PERL
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Figs. 12-18. Mayacnephia salasi. Figs. 12-14, pupa. 12, Respiratory organ (gill). 13, Frons. 14, Abdomen showing chaetotaxy on dorsal (d) and ventral (v) surfaces. Figs. 15-18, Larva. 15, Hypostoma. 16, Inner distal and subapical margins of mandible showing dentation. 17, Antenna. 18, Hypostomal cleft.
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cylindrical, inflated saclike filaments origi- nating from a common short, rather broad base covered with minute spicules; these saclike filaments are rather uniform in length and width, nearly transparent and have a minutely granular texture that is visible only at high magnifications. Head and thoracic integument glabrous; antenna of male ex- tending about '2 distance to hind margin of head; antenna of female extending about %4 or more of distance to hind margin of head; a single stout seta present near inner corner of each antenna, and two or three, some- what separated, shorter and more slender setae present along outer margin of frons at about midlength of antenna (Fig. 13). Dor- sum of thorax without any trace of integu- mental pattern; each side of thorax with about two anterodorsal and one postero- dorsal, and one anteroventral and one pos- teroventral long, simple trichomes, antero- dorsal trichomes stoutest. Chaetotaxy of each lateral half of abdominal tergites as follows (Fig. 14): tergite one with five or six fine setae; tergite two with four or five fine setae and four stouter hooks; tergites three and four each with 2-5 minute setae and four anteriorly directed spines along pos- terior margin; tergite five with about seven minute setae; tergite six with two or three minute setae and a row of minute, poste- riorly directed spinules along anterior mar- gin; tergite seven with three minute setae and an anterior row of minute spinules; ter- gite eight with two minute setae and an an- terior row of minute spinules; tergite nine with a few minute spinules anteriorly, and two long caudal spines situated on two slightly swollen convexities, these spines slightly curved, tips divergent, each with a long, stout seta near base posteriorly. Chae- totaxy of each lateral half of sternites as fol- lows: sternite three with three or four weakly sclerotized hooklets and one fine seta; ster- nites four and five each with four or five hooklets; sternite six with three well-devel- oped hooks; sternite seven with two well- developed hooklets, and one pale, medial
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oval area; sternite eight with one strong hooklet, and one fine seta; sternite nine with two fairly strong setae in striated mem- brane, plus a strong seta at base of caudal spine, otherwise bare; sternites three to eight each with a variably sized but distinct patch of minute spinules. Striated pleural mem- brane on each side of: segment five with two fine setae one of which is in a platelet-like sclerite; segment six with one hooklet and one fine seta in a platelet-like sclerite; seg- ment seven with one hooklet; segment eight with one fine seta; segment nine with eight stout hooks that may be simple, bifurcate, or grapnel-shaped; intersegmental membra- nous area between segment eight and nine with three short but distinct nipple-like bumps on each half. Cocoon a loosely wooven, saclike structure without any def- inite shape, and covered by detritus. Larva (mature, with fully developed res- piratory histoblasts).— Length 6.5-8.5 mm. General body color pale creamy brown; in- tersegmental lines narrow, slightly lighter than rest of abdomen. Head capsule pale yellowish brown; head spots pale brown but darker than surrounding fulvus area, an- teromedian and posteromedian spots slen- der, elongate, the two sets of spots well sep- arated, first and second anterolateral spots roundish, about equal in size and distinctly separated, posterolateral spots slightly dark- ened and somewhat obscure; eye spots small. Postocciput with broad gap dorsally, en- closing small cervical sclerites. Antenna (Fig. 17) pale brownish; about 4 as long as stalk of labral fan; proportions of segments (basal to apical) 1:7.7:2.6. Labral fan with 25-33 (av. 29) primary rays. Hypostoma as in Fig. 15, with 13 teeth arranged in three main groups of 4 + 3 + 4 plus a small tooth on each side of base of median tooth; median tooth long, subequal to longest lateral teeth of each side; each lateral group of teeth sim- ilar in structure to median group, consisting of one main tooth and a smaller tooth on each inner and outer margin, and a short, more ventral, lateral tooth; outer lateral
72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
margins of hypostoma with 3-6 weak ser- rations and two or three long and one or two short hypostomal setae, with longest seta reaching tip of longest hypostomal tooth. Hypostomal cleft (Fig. 18) poorly de- fined, a broad, shallow, U-shaped excava- tion extending about ¥ distance to base of hypostoma. Hypostomal bridge distinctly longer (25:18) than hypostoma. Mandible (Fig. 16) with one large apical tooth, three stout preapical teeth followed by a series of six or seven more seta-like teeth, and two outer teeth; inner subapical ridge with about 16-25 fine serrations. Maxillary palpus about 2.5 times as long as width at base. Lateral plate of proleg extending about ' or more length of apical segment; irregularly subquadrate to subtriangular, greatest width and height nearly equal; lightly sclerotized, with about 20 very slender rod-like exten- sions projecting distally toward bases of hooks; circlet of apical hooks arranged in about 17 rows. Segment eight of abdomen with two short, broadly rounded tubercles that extend about , to '4 depth of abdomen below their points of attachment. Anal pa- pillae with three simple lobes; minute rectal setulae present lateral to anterodorsal arms of anal sclerite. Anterodorsal arms of anal sclerite about '2 as long as posteroventral arms; anterior arms slender, posterior arms considerably broader; sclerotized platelike junction of arms bearing 8-10 fine setae, these setae short but conspicuously longer than rectal setulae. Posterior circlet of hooks consisting of about 8-10 hooks in about 62- 65 rows.
Types.— Holotype, 2 (mounted on five slides), temporary stream (#45a) located at one side of the road between km 96 and 97, near La Georgina on route from Cartago to San Isidro del General, Provincia San José, Costa Rica, November 14, 1983, C. R. Méndez and A. Solano V.
Paratypes.—1 9°, 1 6, same data as type except preserved in alcohol and terminalia of both specimens mounted on slides; 1 4, same data except November 1, 1983 (mounted on three slides); 1 6, same data
except November 6, 1983 (mounted on three slides); 1 2, same data except November 12, 1983 (mounted on four slides); 2 99, 1 é (all pinned), same data except October 28, 1983, A. Solano V. and H. Mayreno; | 4(mounted on six slides), same data except October 24, 1983, and | ¢(mounted on six slides), same data except November 7, 1983, A. Solano V. and H. Mayreno; 2 pupae, same data except October 28, 1983 (one pupa mount- ed on two slides, the other pupa mounted on four slides) ; 30 pupae (four mounted on slides), same data except October 28, 1983, A. Solano V. and H. Mayreno; 13 pupae, same data except July 14, 1986, A. Solano V.and H. Mayreno; | larva, same data ex- cept June 7, 1983 (mounted on five slides): 1 larva, same data except June 17, 1983 (mounted on six slides); 12 larvae (five mounted on six slides each), same data ex- cept July 14, 1986, A. Solano V. and H. Mayreno.
Holotype deposited in the collection of the U.S. National Museum of Natural His- tory, Washington, D.C. Paratypes are de- posited in the U.S. National Museum of Natural History, and the entomology col- lection of the Department of Parasitology, University of Costa Rica.
This species is dedicated to Eng. Luis An- gel Salas F., a distinguished acarologist, and Professor Emeritus at the School of Agron- omy, University of Costa Rica.
Biological notes.—All available speci- mens of M. sa/asi came from the same tem- porary stream (numbered 45a) at the type locality. This small, shallow, clean-water stream is situated at an elevation of 3150 meters in the bottom ofa deep, steep-sloped gulley in an area partially to heavily shaded by forest, and with a moderate amount of vegetation on the banks. The stream is about one meter wide, with a bed of rocky sections and muddy areas. The water flows at a slow to moderate rate, has a temperature that ranges between 10-14°C, and pH values are between 7.1 and 7.8. Larvae are present in the stream throughout the year.
Remarks.—Wygodzinsky and Coscaron
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(1973) recognized two groups of species in Mayacnephia based on the chaetotaxy of the abdomen and the form of the respiratory gills of the pupae. The new species described here belongs to their ““apomorphic” group having a reduced number of filaments and without at least an apical filamentous por- tion to the filaments, as well as having a larger number of sternal setae and hooks. Mayacnephia salasi has only four simple, inflated, saclike filaments in the respiratory organ, and has more numerous spines on the sternal sclerites (sternite five with about 5+5, sternite six with about 3+ 3, and ster- nite seven with about 2+ 2) characteristic of this group of species. The recently described M. fortunensis Petersen (1985), from Pan- ama, also belongs to this group. Mayacnephia salasi appears to be most similar to M. grenieri (Vargas and Diaz Na- jera) in the adult and pupal stages. The fe- male of M. sa/asi can be most easily differ- entiated from M. grenieri by the antenna, which has a yellowish scape and pedicel that contrast with the dark brown flagellum (in grenieri the antenna is entirely yellow ex- cept for the black first flagellomere), the hal- ter with its yellow knob and brownish yel- low stem (knob black, stem whitish), and the genital fork whose arms diverge from the stem in a broad V-shape and have rect- angular plates that are wider than long, each plate with a small, sclerotized, toothlike process on its anterior margin (arms of gen- ital fork diverge from the stem at nearly right-angles so the posterior space between the arms 1s broadly U-shaped, the plates are shorter, broader and subrectangular, and each has a prominent tooth-like process). The male of MM. sa/asi can be distinguished from M. grenieri by the all black antenna (in grenieri the antenna is yellow except for the black first flagellomere), the pale yellow, recumbent pile of the scutum (recumbent pile light brown), halter brown with a yellow stem (halter entirely black), and by the shape of the ventral plate of the aedeagus which, in ventral view, is more rectangular with a more broadly rounded distal margin, and
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longer basal arms (more triangular in ven- tral view, with narrower and more pointed apical margin and much shorter basal arms).
Larvae of M. salasi can be distinguished from M. grenieri by the following combi- nation of characters: length 6.5-8.5 mm (larvae of grenieri range from 8.0—9.0 mm), pale creamy brown color (distinctly yellow- ish but more opaque especially posteroven- trally), antenna about *4 length of stalk of labral fan (antenna as long as stalk of labral fan), labral fan with 25-33 primary rays (40- 45 primary rays), hypostoma with 13 apical teeth (9 apical teeth), hypostomal bridge distinctly longer (25:18) than hypostoma (hypostomal bridge and hypostoma nearly equal in length), and posterior circlet with 62-65 rows of hooklets (76 rows of hooklets in grenier!).
The species of Mayacnephia, as with some other groups of Simulidae, are most easily differentiated on the basis of the number and shape of the filaments of the pupal re- spiratory organ (gill). The following key to the pupae of the species of Mayacnephia will differentiate M/. sa/asi from the other described species of the genus. The pupa of M. osborni (Stains and Knowlton), a species described from California, does not appear in the key because it is unknown. We are unable to prepare a reliable key to the other stages of the species of Mayacnephia for lack of specimens. Dalmat (1955) and Diaz Najera (1962) provided keys that include the larvae of various species now included in Mayacnephia, and Dalmat provided a key to the males and females of the three species known from Guatemala.
KEY TO SPECIES OF MAYACNEPHIA PUPAE
1. Respiratory organ with two long, swollen tu- bular filaments arranged in the form of a V (figs. 17-18 in Dalmat 1949; fig. 1OB in Wy- godzinsky and Coscaron 1973). Highlands of Guatemala, Mexico (Chiapas) aguirrei (Dalmat)
— Respiratory organ with three or more fila- ments of varying form and arrangement 2
2. Respiratory organ with three long, swollen, tubular filaments (fig. 20 in Diaz Najera 1962). Mexico (Oaxaca) ....... mixensis (Diaz Najera)
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10.
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Respiratory organ with four or more filaments
OL WAT YIN) S POU oa ao rsycs cs, y's Sac eaters agshnyays cours 3 . Respiratory organ with four’ alarients ree A Respiratory organ with six or more filaments BH eigee cehatiat JL dss Bee 5 Filaments short, inflated and saclike with broadly rounded tips, all arising from a com- mon base (fig. 12 herein). Costa Rica ...... eae ic Se EI ch ae salasi n. sp.
Filaments longer, swollen but not greatly in- flated or saclike, tubular with pointed apices, arranged in two pairs (fig. 22 in Vargas and Diaz Najera 1948; fig. 9A in Wygodzinsky and Coscaron 1973). Mexico (Veracruz) .
. grenieri (Vargas and Diaz Najera) Respiratory organ with six long, swollen, tu- bular filaments that are rounded distally, an- terior two filaments single, posterior two fil- aments branching into two petiolate pairs (fig. 4 in Coleman 1953). California stewarti (Coleman) Respiratory organ with seven or more fila-
ments of varying form 6 Respiratory organ with seven or eight fila-
MGT: creas pecs cetitrs Haseeno ara e 7 Respiratory organ with iy l- 15 filaments 9 Respiratory organ with seven filaments 8
Respiratory organ with eight filaments (fig. 27
in Currie 1986). Western Canada Me
; .(unnamed species X) Respiratory organ n with four swollen, tubular filaments, three of which, in turn, give rise to
two slender filaments distally, and the fourth which has a single slender terminal filament
(fig. 8A in Wygodzinsky and Coscaron 1973). Highlands of Guatemala roblesi (Leon) Respiratory organ with four swollen, tubular, clavate filaments; anteromedial and postero- medial filaments unbranched although the former sometimes with a variably developed thumblike hump on its mesal surface; antero- lateral filament branching into two petiolate filaments, and posterolateral filament branch-
ing into three filaments (fig. 8 in Petersen 1985). Panama _fortunensis Petersen Respiratory organ with 11-12 filaments . 10 Respiratory organ with 14-15