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Life history and descriptions of adults and immature stages of aciurina semilucida (bates) (diptera: tephritidae) on chrysothamnus viscidiflorus (hooker) nuttall in southern California PDF

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Preview Life history and descriptions of adults and immature stages of aciurina semilucida (bates) (diptera: tephritidae) on chrysothamnus viscidiflorus (hooker) nuttall in southern California

PROC. ENTOMOL. SOC. WASH. 98(4), 1996, pp. 752-766 LIFE HISTORY AND DESCRIPTIONS OF ADULTS AND IMMATURE STAGES OF ACIURINA SEMILUCIDA (BATES) (DIPTERA: TEPHRITIDAE) ON CHRYSOTHAMNUS VISCIDIFLORUS (HOOKER) NUTTALL IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN AND JEFFREY A. TEERINK Department of Entomology, University of California, Riverside, CA 92521, U.S.A. Abstract.(cid:8212)Aciurina semilucida (Bates) is a rare, highly localized, tephritid fly forming pubescent, axillary bud galls on branches of Chrysothamnus viscidiflorus (Hooker) Nuttall in southern California. Documentation is provided that the wing patterns of the adults are very strongly sexually dimorphic in southern California, but much less dimorphic in some other parts of its range, e.g. in Idaho, where it previously has been reported only from C. nauseosus (Pallen) Britton. The egg, first-through third-instar larvae, and puparium are described, illustrated, and compared with other southern California Aciurina spp. The egg has a long pedicel, with a knoblike anterior apex. The second and third instars have abdominal terga IIJ(cid:8212)V expanded into a dorsal ambulatory lobe. This ambulatory lobe and the large open gall cavity within which the larva feeds are characteristics shared only with A. michaeli Goeden among southern California Aciurina spp. studied to date. Key Words: Insecta, Aciurina, Chrysothamnus, biology, galls, regional sexual dimor- phism, adult taxonomy, immature taxonomy, parasitoids Six Aciurina spp. (Diptera: Tephritidae) urina species ...(cid:8221) (Foote et al. 1993). In- forming galls on Chrysothamnus or Bac- stead, A. semilucida (Bates) is the rare, lo- charis spp. (Asteraceae) in southern Cali- calized, and distinct species in California fornia have been studied by us to date. This that we describe herein. paper follows those on A. thoracica Curran (Headrick and Goeden 1993), A. ferruginea MATERIALS AND METHODS (Doane), A. michaeli Goeden (Goeden and Our field studies were conducted on Teerink 1996a), and A. idahoensis Steyskal Chrysothamnus viscidiflorus (Hooker) Nut- (Goeden and Teerink 1996b), in a series tall (Asteraceae) at three locations in Deep projected to end with A. trixa Curran [as Spring Valley, the southernmost extension interpreted and reinstated by Dodson and of the Great Basin biome (Munz 1974, Hall George (1986)], which is the most common 1991), located northeast of the Westgard species of Aciurina in California (D. H. Pass in the White-Inyo Mountain Range, Headrick, RDG, and JAT, unpublished Inyo Co., California. Galls containing lar- data). Steyskal (1984) erroneously synony- vae and puparia of A. semilucida were sam- mized A. semilucida (Bates) with A. bige- pled from plants in Payson Canyon at 1770 loviae (Cockerell) and A. trixa (Dodson and m at the southwest end of the Valley, north George 1986) to make A. bigeloviae (cid:8220) of Deep Spring Lake at 1720 m towards the the most widely distributed and most vari- center of the Valley, and above Antelope able of all the Chrysothamnus-feeding Aci- Spring at 1695 m on the north side of the VOLUME 98, NUMBER 4 V3 Valley during March and April, 1993 and modified by Dodson and George (1986). 1994. Excised galls were transported in Morphological terminology and telegraphic cold-chests in an air-conditioned vehicle to format used to describe the immature stages the laboratory and stored under refrigera- follow Headrick and Goeden (1993) and tion for subsequent dissection, photogra- Goeden and Teerink (1996a, b) and our ear- phy, description, and measurement. Two lier works cited therein. Means + SE are first-instar, five second-instar, and 18 third- used throughout this paper. Voucher speci- instar larvae and two puparia dissected mens of reared adults of A. semilucida re- from galls were preserved in 70% EtOH for side in the research collection of RDG; pre- scanning electron microscopy (SEM). All served specimens of eggs, larvae, and pu- other fully grown larvae and puparia were paria are stored in the research collection of placed in separate, glass shell vials stop- JAT. pered with absorbant cotton and held in hu- midity chambers at room temperature for RESULTS AND DISCUSSION adult and parasitoid emergence. Specimens Taxonomy for SEM later were hydrated to distilled wa- ter in a decreasing series of acidulated Adult.(cid:8212)Aciurina semilucida was de- EtOH. They were osmicated for 24 h, de- scribed (as a Tephrella) by Bates (1935) hydrated through an increasing series of from five males and seven females reared acidulated EtOH, critically point dried, from galls on Chrysothamnus (as Bigelovia) mounted on stubs, sputter-coated with a sp. from Riparia, Washington. The host gold-palladium alloy, and studied with a plant of A. semilucida was further identified JEOL JSM C-35 SEM in the Department of as C. nauseosus (Pallen) Britton in Idaho Nematology, University of California, Riv- by Novak et al. (1967). Curran (1932) erside. erected the genus Aciurina and designated Most adults reared from isolated puparia the type species as A. trixa. Steyskal (1984) were individually caged in 850-ml, clear- synoptically revised the genus, and Foote et plastic, screened-top cages with a cotton al. (1993) adopted his revision, which syn- wick and basal water reservoir and provi- onymized both A. trixa and A. semilucida sioned with a strip of paper toweling im- with A. bigeloviae. Unfortunately, Steyskal pregnated with yeast hydrolyzate and su- (1984) and Foote et al. (1993) were un- crose. These cagings were used for longev- aware that several North American species ity studies and oviposition tests in the in- of Aciurina have sexually dimorphic wing sectary of the Department of Entomology, patterns, e.g. A. ferruginea, A. michaeli, and University of California, Riverside, at 25 + A. idahoensis (Goeden and Teerink 1996a, 1°C, and 14/10 (L/D) photoperiod. Virgin b, unpublished data). Furthermore, A. ida- male and female flies obtained from emer- hoensis shows regional variation in this di- gence vials as well as field-swept adults morphism, i.e. being strongly dimorphic in were paired in clear-plastic petri dishes pro- California, but less so in Idaho (Goeden and visioned with a flattened, water-moistened Teerink 1996b). pad of absorbant cotton spotted with honey Before redescribing the adults of what (Headrick and Goeden 1991) for direct ob- we believe are A. semilucida in California, servations, videorecording, and _ still-pho- we provide our reasons for identifying them tography of their courtship and copulation as such. First, A. semilucida was reported behavior to be reported elsewhere (Head- by Wangberg (1981) from galls on C. nau- rick and Goeden, unpublished data). seosus similar to those found by us on C. Plant names used in this paper follow viscidiflorus in California; however, in Ida- Munz (1974); tephritid nomenclature and ho, A. semilucida, like A. idahoensis (Goe- terminology follow Foote et al. (1993) as den and Teerink 1996b), is much less sex- 754 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ually dimorphic. Steyskal (1983) believed much different wing pattern (i.e. the Cali- that A. semilucida (cid:8220)(cid:8216)... was no more than fornia population has strongly sexually di- A. bigeloviae with a strongly reduced wing morphic wing patterns). The wing pattern pattern.(cid:8221)(cid:8217) Accordingly, he synonymized A. of these southern California females (Fig. semilucida with A. bigeloviae, along with 1F) is similar to that of A. Jutea (Coquillett) A. trixa, the resulting species representing as pictured and described in Steyskal what he termed *... the most widely dis- (1984) and Foote et al. (1993). Aciurina lu- tributed and most variable of all the Chrys- tea is a rare but widely distributed species othamnus-feeding species, that is, the typi- that has been reported only from C. visci- cal group of Aciurina.(cid:8221)(cid:8217) Dodson and George diflorus (Wangberg 1981, Steyskal 1984, (1986) concluded, however, that A. trixa is Foote et al. 1993). However, in both female a separate and distinct species forming California A. semilucida and A. lutea nei- smooth galls on C. nauseosus; whereas, A. ther hyaline mark in cell m crosses vein M bigeloviae forms (cid:8220)(cid:8216)cottony(cid:8221) galls on this (Fig. 1F), as does the apical mark in A. bi- same host plant. Neither A. bigeloviae nor geloviae and A. trixa (Headrick, RDG, and A. trixa apparently exhibited sexually di- JAT, unpublished data). Furthermore, A. L. morphic wing patterns, although consider- Norrbom (1996, in litt.) reports that three able wing pattern variation was reported af- males of A. lutea in the U.S. National Mu- ter extensive field and laboratory studies by seum collection (from northern California, Dodson and George (1986) in Arizona, Nevada, and Utah) have the same wing pat- Colorado, New Mexico, and Utah. How- tern as the female holotype (i.e. like Foote ever, only A. trixa apparently occurs on C. et al. 1993, Fig. 115), and thus are quite nauseosus in California, as no cottony galls different from California males of A. sem- of A. bigeloviae have ever been found on ilucida. Moreover, A. semilucida has a this widely distributed host plant extensive- densely gray microtrichose scutum with ly studied by us throughout its range in white to yellow, stout setulae (see below); southern California (D. H. Headrick, RDG, whereas, these three A. /utea males have a and JAT, unpublished data). Only A. trixa mostly bare, shiny scutum with slender, was reared from smooth galls on C. nau- black setulae (Norrbom, in litt., 1996). seosus at our study site, where A. semilu- Also, in A. lutea, the subapical hyaline spot cida was found only on C. viscidiflorus. in cell dm touches or approaches vein M, Figure 1A and 1B respectively show the but is well isolated from vein CuA, (Norr- right wings typical of voucher specimens of bom, in litt., 1996); whereas, cell dm touch- male and female A. semilucida reared from es CuA, in A. semilucida (see below). Fi- galls on C. nauseosus collected from near nally, Wangberg (1981) pictured and de- Lewiston, Nez Perce Co., Idaho, where scribed the gall of A. /utea as a simple, but Wangberg (1981) studied this species. Fig- abrupt, woody swelling with thickened ure 1C and 1D respectively depict a male walls on older stems as well as current and female with the more darkly patterned year(cid:8217)s stem growth. Thus, its reported gall condition and the variation found among is definitely different from that which we these same voucher specimens examined describe below. from Idaho. Compare the aforementioned Knowing now that sexually dimorphic wing patterns with those of a male (Fig. 1E) wing patterns occur in some species of Aci- and female (Fig. 1F) that we call A. semi- urina (Goeden and Teerink 1996a, b), and lucida reared from similar galls on C. vis- that this dimorphism may occur or nearly cidiflorus in southern California. The wing be absent within different parts of the range patterns of the males from California and of a single species (Goeden and Teerink Idaho compare favorably; whereas, the fe- 1996b), we conservatively chose to identify males from southern California have a the flies with which this paper is concerned VOLUME 98, NUMBER 4 WD F E Fig. 1. Right wings of Aciurina semilucida. (A) Light form of male from Idaho; (B) light form of female from Idaho; (C) dark form of male from Idaho; (D) dark form of female from Idaho; (E) male from California; (F) female from California. as A. semilucida. Accordingly, males (Fig. terns found among both males and females 1A, 1C) and females (Fig. 1B, 1D) of A. of A. semilucida from Idaho (Fig 1A(cid:8212)D), semilucida examined from Idaho and males but only among males (Fig. 1E) from Cal- from California (Fig. 1E) generally appear ifornia. Similarly, the most hyaline wing like the two most hyaline wing patterns pattern of the three patterns for A. bigelov- (>50% hyaline) among the four patterns il- iae illustrated by Foote et al. (1993, Fig. lustrated for A. bigeloviae by Steyskal 124) is generally illustrative of the wing (1984, Fig. 15-16). Put another way, these patterns of males and females of A. semi- two illustrations in Steyskal (1984) respec- lucida from Idaho and the males from Cal- tively represent the light and dark wing pat- ifornia. This suggested relabeling parallels 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that applicable to the wing pattern of male setulae or absent; one postsutural dorsocen- A. idahoensis from California, which close- tral slightly posteriorad of pre-sutural su- ly resembles that illustrated for A. notata in pra-alar, at half the distance to the supra- Steyskal (1984) and Foote et al. (1993) [but alar bristle; two notopleurals; one each of see discussion in Goeden and Teerink post-alar, intra-alar, and acrostical; scutel- (1996b) of inconsistencies between pub- lum with one basal, prominent, scutellar lished reports on A. idahoensis and A. no- bristle; katepisternal bristle well developed. tata|. The males of A. idahoensis from Cal- Coxae, femora, tibiae, and tarsi yellow; fore ifornia and males of A. semilucida both femur, unlike male, unthickened, but also have reduced wing patterns and mostly hy- bearing loose (cid:8220)(cid:8216)comb(cid:8221) of long, anteroven- aline wings, but are readily separated by the tral setae. Wings (Fig. IF) mostly dark much more reduced pattern in the wings of brown, with marginal hyaline incisions and the former males (Goeden and Teerink internal marks as follows: cell be wholly 1996b). yellow; cell c with basal and median dark Females from California(cid:8212)(cid:8212)Apparently, or hyaline spots, if the latter, also with females of A. semilucida in southern Cali- small, apical, marginal hyaline incision; cell fornia have a wing pattern (Fig. 1F) that r, with one or two hyaline spots along cos- heretofore has neither been recognized nor ta, if one, the spot extending halfway across described. Or they have been subsumed un- cell r,, if two, the proximal spot extending til now, like other species noted above, as to vein R,,3, the distal spot across cells r, one of the dark patterned wing forms of A. and r,,3, to vein R,,; (Fig. 1F) or across r,, bigeloviae or, conceivably, A. lutea, as dis- r>,, and r,,, to vein M; cell r,,; with a dis- tinguished and illustrated by Steyskal tinct bulla; cell dm with prominent preapi- (1984) and Foote et al. (1993). The follow- cal hyaline spot touching both veins M and ing description of the California form of A. CuA,, just CuA,, or neither vein; cell m semilucida female is based on three females with median, hyaline, wedge-shaped mark individually reared from galls collected at reaching or nearly reaching anteriorly- our Payson Canyon study site. Figure 1F is bowed vein M and_ sub-basal, hyaline, a photograph of the right wing excised from wedge-shaped spot half as long; vein dm-cu one of these three females. bowed apicad; cell br with prominent Head: White with golden yellow anten- preapical spot; cell cua, with one or two nae, frons, and ventral occiput. Three pairs marginal hyaline incisions, apical incision of frontal bristles, basally yellow, light absent or reaching CuA, and contiguous brown distally; two pairs of orbital bristles, with preapical spot in cell dm, and promi- the anterior pair light brown, posterior pair nent basal incision starting basad of and light yellow; inner vertical bristles and crossing vein A,+CuA, and extending an- ocellar bristles well developed and light teriorly to or nearly to CuA,; alula and ad- brown; other bristles wholly or basally jacent base of anal lobe hyaline. white or light yellow. Abdomen: Shiny, reddish-brown, except Thorax: Dark brown, mesonotum cov- for tergite 6, which may be dark brown, ered with fine grey pollinosity and many, oviscape shiny black; tergum sparsely cov- prominent, pale to golden yellow setulae ered with fine, light yellow setulae. posteriorad as far as and including all but Males and Idaho females.(cid:8212)The wings of shiny, posterior margin of dark brown or 12 males of A. semilucida examined from black scutellum. Mediotergite black, shiny California and 40 males and 20 females ex- medially, grey pollinose dorsally and later- amined from Idaho characteristically are ally. All of the following paired bristles are >50% hyaline (Fig. 1A(cid:8212)E), and are de- present and yellow-brown: One postprono- scribed here collectively: Cell be wholly tal; scapular bristles indistinguishable from hyaline; cell c wholly hyaline or yellow or VOLUME 98, NUMBER 4 ST with a small central brown spot; brown or longated dorsal petals dorsad of mouth lu- yellow-brown band covering veins bm(cid:8212)cu men (Fig. 2B-2); paired dorsal sensory or- and r(cid:8212)m; apices of cells r,,, and r,4,5 nar- gans dorsomediad of anterior sensory lobes, rowly brown; brown spot arising midway each consisting of dome-shaped papilla on vein CuA,; usually brown crossband (Fig. 2B-3, 2C-1); anterior sensory lobes running from apex of cell r, across vein bear terminal sensory organ (Fig. 2C-2), pit dm-cu, varying in width, sometimes inter- sensory organ (Fig. 2C-3), lateral sensory rupted in cell r,,,, or more commonly, bro- organ (Fig. 2C-4), and supralateral sensory ken or attenuated at or before vein M. organ (Fig. 2C-5); stomal sense organs ven- These spots connected or variously embel- trolaterad of anterior sensory lobes (Fig. lished in darker wing patterns (Fig. IC, 2D-1); lateral sensillum located laterad of 11D): anterior sensory lobe (Fig. 2B-4); ventro- Immature stages.(cid:8212)Egg: Fifteen ova of A. lateral sensillum ventrolaterad of stomal semilucida were white, opaque, smooth; sense organ (Fig. 2D-2); mouth hooks bi- with an elongate-ellipsoidal body, 0.52 + dentate, teeth conical (Fig. 2D-3); median 0.007 (range, 0.46(cid:8212)0.56) mm long, 0.21 = oral lobe laterally flattened, tapering api- 0.008 (range, 0.16(cid:8212)0.26) mm wide, smooth- cally (Fig. 2D-4); prothorax smooth, minute ly rounded at tapered posterior end; with an acanthae ventrally (Fig. 2B-5); anterior tho- elongate, anterior pedicel, 0.25 + 0.008 racic spiracles dorsolaterad on posterior (range, 0.20(cid:8212)0.30) mm long, 0.03 (range, margin of prothorax with three oval papil- 0.02(cid:8212)0.04) mm wide medially, and 0.05 lae (Fig. 2E); metathoracic lateral spiracular (range, 0.04(cid:8212)0.06) mm wide at knoblike, complex consists of an open spiracle (Fig. anterior apex. ~ 2F-1), and two verruciform sensilla (Fig. The eggs of A. semilucida are stalked 2F-2); abdominal lateral spiracular complex like those of A. ferruginea and A. michaeli consists of an open spiracle and a single (Goeden and Teerink 1996a) and A. ida- verruciform sensillum; caudal segment hoensis (Goeden and Teerink 1996b), all bears the posterior spiracular plates (Fig. four of which differ greatly in appearance 2G); posterior spiracular plates with three from the short-pedicelled, partly reticulated oval rimae, ca. 0.042 mm in length (Fig. egg of A. thoracica (Headrick and Goeden 2G-1), and four interspiracular processes, 1993). The respiratory function of the elon- longest measuring 0.01 mm (Fig. 2G-2); gate pedicel of these eggs was first sug- compound sensilla ventrad of posterior spi- gested for A. michaeli by Tauber and Taub- racular plates consist of a stelex sensillum er (1967) and was supported with morpho- (Fig. 2H-1) and a verruciform sensillum logical and biological data by Goeden and (Fig. 2H-2); stelex sensilla surround margin Teerink (1996a). The egg bodies and pedi- of caudal segment in 2-dorsal, 4-ventral ar- cels of A. idahoensis and A. semilucida are rangement. shorter on average than these parts of the Aciurina semilucida closely resembles A. eggs of both A. ferruginea and A. michaeli michaeli in general, third-instar habitus (Goeden and Teerink 1996a, b), with A. (Goeden and Teerink 1996a). Both species semilucida having the shortest pedicel of are enlarged dorsally and have a distinct the three species. hump, which differentiates them from other Third instar: Superficially smooth, pyri- Aciurina species studied thus far, i.e. A. form, tapering anteriorly, truncated poste- thoracica (Headrick and Goeden 1993), A. riorly, abdominal segments III-V enlarged ferruginea (Goeden and Teerink 1996a), A. dorsally, minute acanthae ventrally (Fig. idahoensis (Goeden and Teerink 1996b) 2A); gnathocephalon conical, rugose pads and A. trixa, (Goeden, unpublished data). mediad of anterior sensory lobes and later- Aciurina semilucida and A. michaeli are ad of mouth lumen (Fig. 2B-1); pair of en- very similar in the types, sizes, and place- 15KU 446 @ be kul, x veoe. KU ¥Abo Saal 1 Fig. 2. Third instar of A. semilucida. (A) Habitus, anterior to left; (B) gnathocephalon, anterior view, 1(cid:8212) rugose pads, 2(cid:8212)elongated dorsal petals, 3(cid:8212)dorsal sensory organ, 4(cid:8212)lateral sensillum, 5(cid:8212)minute acanthae; (C) anterior sensory lobe, 1(cid:8212)dorsal sensory organ, 2(cid:8212)terminal sensory organ, 3(cid:8212)pit sensory organ, 4(cid:8212)lateral sensory organ, 5(cid:8212)supralateral sensory organ; (D) gnathocephalon, ventrolateral view, 1(cid:8212)stomal sense organ, VOLUME 98, NUMBER 4 759 ment of sensory structures. They do how- terior spiracular plates (Fig. 3F-2) consist ever, differ in the number of sensilla in the of a stelex sensillum and a verruciform sen- lateral spiracular complex. Aciurina mi- sillum (Fig. 3H); stelex sensilla surround chaeli has only one verruciform sensillum margin of caudal segment in 2-dorsal, in the metathoracic lateral spiracular com- 4-ventral arrangement (Fig. 3F-3). plex and no sensilla in the abdominal lateral The second instar is enlarged dorsally in spiracular complex (Goeden and Teerink abdominal segments III(cid:8212)V as in the third 1996a). Aciurina idahoensis which co-oc- instar. The gnathocephalon is similar in curs with A. semilucida on C. viscidflorus, shape and sensory structures. The anterior has the same number of sensillum in the thoracic spiracles, lateral spiracular com- lateral spiracular complex as A. michaeli. plex and the posterior spiracles are consis- (Goeden and Teerink 1996b). tent between the two instars. The second Second instar: Superficially smooth, pyr- instar differs slightly from the third in the iform, minute acanthae ventrally (Fig. 3A); compound sensilla on the caudal segment. gnathocephalon conical, smooth, rugose One pair in the second instar consists only pads mediad and laterad of anterior sensory of a stelex sensillum. This was also ob- lobes (Fig. 3B-1); dorsal petals dorsad of served in A. idahoensis, where the second mouth lumen (Fig. 3B-2); paired dorsal sen- instar contained only one sensillum in one sory organs dorsomediad of anterior sen- of the compound sensilla (Goeden and sory lobes (Fig. 3B-3, 3C-1); anterior sen- Teerink 1996b); A. ferruginea also showed sory lobes bear the terminal sensory organ slight differences in the compound sensilla (Fig. 3C-2), pit sensory organ (Fig. 3C-3), in the second and third instars (Goeden and lateral sensory organ (Fig. 3C-4), and su- Teerink 1996a). pralateral sensory organ (Fig. 3C-5); stomal First instar: Superficially smooth, ovoi- sense organs ventrolaterad of anterior sen- dal, minute acanthae circumscribe interseg- sory lobes, near mouth lumen (Fig. 3B-4, mental lines; gnathocephalon conical, 3C-6); mouth hooks bidentate, teeth conical smooth (Fig. 4A); paired dorsal petals dor- (Fig. 3B-5); median oral lobe laterally flat- sad of mouth lumen (Fig. 4A-1); dorsal sen- tened, tapering apically (Fig. 3B-6); protho- sory organs dorsomediad of anterior sen- rax smooth (Fig. 3B); anterior thoracic spir- sory lobes (Fig. 4A-2, 4B-1); anterior sen- acles consist of three oval papillae (Fig. sory lobes bear the terminal sensory organ 3D); metathoracic lateral spiracular com- (Fig. 4B-2), pit sensory organ (not ob- plex consists of an open spiracle (Fig. served), lateral sensory organ (Fig. 4B-3), 3E-1), and two verruciform sensilla (Fig. supralateral sensory organ (Fig. 4B-4); sto- 3E-2); abdominal lateral spiracular complex mal sense organs indistinct, laterad of consists of an open spiracle and a single mouth lumen (Fig. 4A-3); mouth hooks bi- verruciform sensilla; caudal segment bears dentate, teeth conical (Fig. 4A-4); median the posterior spiracular plates (Fig. 3F-1); oral lobe laterally flattened (Fig. 4A-5); posterior spiracular plates with three oval prothorax smooth; anterior thoracic spira- rimae, ca. 0.01 mm in length (Fig. 3G-1), cles absent; lateral spiracular complex not and four blade-like interspiracular process- observed; caudal segment obscured in SEM es, longest measuring 0.008 mm (Fig. specimens. 3G-2); compound sensilla ventrad of pos- The first instar is not expanded dorsally << 2(cid:8212) ventrolateral sensillum, 3(cid:8212)mouth hooks, 4(cid:8212)median oral lobe; (E) anterior thoracic spiracle; (F) metathorax, lateral spiracular complex, 1(cid:8212)spiracle, 2(cid:8212)verruciform sensilla; (G) caudal segment, posterior spiracular plates, 1(cid:8212)rima, 2(cid:8212)interspiracular process; (H) compound sensillum, 1(cid:8212)stelex sensillum, 2(cid:8212)verruciform sensillum. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Second instar of A. semilucida. (A) Habitus, anterior to left; (B) gnathocephalon, anterior view, 1(cid:8212) rugose pads, 2(cid:8212)dorsal petals, 3(cid:8212)dorsal sensory organ, 4(cid:8212)stomal sense organ, 5(cid:8212)mouth hooks, 6(cid:8212)median oral lobe; (C) anterior sensory lobe, 1(cid:8212)dorsal sensory organ, 2(cid:8212)terminal sensory organ, 3(cid:8212)pit sensory organ, 4(cid:8212)1ateral sensory organ, 5(cid:8212)supralateral sensory organ, 6(cid:8212)stomal sense organ; (D) anterior thoracic spiracle; VOLUME 98, NUMBER 4 761 as in the third and second instars. The gen- eral habitus is more ovoid than pyriform. The gnathocephalon is smooth, lacking ru- gose pads. The anterior sensory lobes and stomal sense organs are small and indis- tinct. The gnathocephalon in A. semilucida first instars is similar to that of A. ferrugi- nea and A. idahoensis (Goeden and Teerink 1996a, b). Puparium: Smooth, elongate-reniform, reddish-brown (Fig. 5A); anterior end bears anterior thoracic spiracles (Fig. 5B-1) and invagination scar (Fig. 5B-2); caudal end bears posterior spiracular plates; posterior spiracular plates with three oval rimae, Ca. 0.021 mm in length (Fig. 5C-1), and four interspiracular processes with 1(cid:8212)3 branches, longest measuring 0.01 mm (Fig. 5C-2). Nineteen averaged 3.94 + 0.17 (range, 3.06(cid:8212)4.77) mm in length and 1.93 + 0.07 (range, 1.44(cid:8212)2.34) mm in width. DISTRIBUTION AND HOSTS Aciurina semilucida was known only from its type locality in Washington (Bates 1935) and from Nez Perce Co. in Idaho Fig. 4. First instar of A. semilucida. (A) Gnatho- (Wangberg 1981). Although its distribution cephalon, anterolateral view, 1(cid:8212)dorsal petals, 2(cid:8212)dor- probably parallels those of its known host sal sensory organ, 3(cid:8212)stomal sense organ, 4(cid:8212)mouth plants, C. viscidiflorus and C. nauseosus, hooks, 5(cid:8212)median oral lobe; (B) anterior sensory lobe, 1(cid:8212)dorsal sensory organ, 2(cid:8212)terminal sensory organ, both of which are common throughout the 3(cid:8212) lateral sensory organ, 4(cid:8212)supralateral sensory or- western United States (Munz 1974), it ap- gan. parently exists in localized, discontinuous demes, whose semi-isolation probably fa- cilitated the strong dimorphism at the ultimately show these southern and north- southwestern periphery of its distribution in ern flies to be distinct and each to have spe- southern California reported herein. Aciuri- cialized on only one of these two hosts, as na semilucida, like A. idahoensis, has only apparently has happened in California with been found in California at three locations A. ferruginea, A. michaeli (Goeden and in Deep Spring Valley, which lies close to Teerink 1996a), A. idahoensis (Goeden and the Nevada border. Alternatively, further Teerink 1996b), and A. semilucida on C. taxonomic study of the galls, hosts, and viscidiflorus versus A. trixa on C. nauseo- reared A. semilucida adults from other lo- sus in California (unpublished data). How- cations in the western United States may ever, oligophagy is known among Aciurina (cid:8212) (E) metathorax, lateral spiracular complex, 1(cid:8212)spiracle, 2(cid:8212)verruciform sensilla; (F) caudal segment, 1(cid:8212)pos- terior spiracular plate, 2(cid:8212)compound sensilla, 3(cid:8212)stelex sensillum; (G) posterior spiracular plate, 1(cid:8212)rima, 2(cid:8212) interspiracular proeess; (H) compound sensillum, stelex sensillum.

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