LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF NEASPILOTA WILSONI BLANC AND FOOTE (DIPTERA : TEPHRITIDAE) ON HAZARDIA SQUARROSA (HOOKER AND ARNOTT ) E. GREENE (ASTERACEAE) IN SOUTHERN CALIFORNIA

13 Pages·1999·6.4 MB·

by  R D Goeden

PROC. ENTOMOL. SOC. WASH.
101(4), 1999, pp. 897-909
LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF
NEASPILOTA WILSONI BLANC AND FOOTE (DIPTERA: TEPHRITIDAE) ON
HAZARDIA SQUARROSA (HOOKER AND ARNOTT) GREENE
E.
(ASTERACEAE) IN SOUTHERN CALIFORNIA
Richard D. Goeden and David H. Headrick
(RDG) Department of Entomology, University of California, Riverside, CA 92521,
U.S.A. (e-mail: [email protected]); (DHH) Department ofCrop Science, California
Polytechnic State University, San Luis Obispo, CA 93407, U.S.A. (e-mail: dheadric@
polymail.calpoly.edu)
—
Abstract. Neaspilota wilsoni Blanc and Foote is a univoltine, monophagous fruit fly
(Diptera: Tephritidae) developing in the flower heads ofHazardia squarrosa (Hooker and
Amott) E. Greene in southern California. The egg, first-, second-, and third-instar larvae,
and puparium are described and figured. The egg pedicel is conical and completely cir-
cumscribed by many, shallow aeopyles of various sizes. The mouth hooks of all three
instars are bidentate. The intersegmental areas of the first instar bear minute acanthae of
a heretofore unreported form, i.e., cylindrical and apically bearing one to five prongs.
Also newly reported, the caudal segment of the first instar bears stelex sensilla, each
basally ringed with one to three, upright, pointed acanthae. The integumental petal is
laterally fused with the stomal sense organ in the first instar, but separate in the second
instar. The larvae feed mainly on the ovules and soft achenes as first and second instars;
however, as third instars, they usually extend their feeding into the receptacle and addi-
tionally feed on sap that collects in the shallow scars. The nonfeeding prepuparium over-
winters in a protective cell that occupies much ofthe excavated flower head and is formed
of floret, pappus, and achene fragments impregnated with excess sap and liquid feces that
harden when dry. A few prepuparia pupariate and emerge as adults in the fall and over-
winter, but most pupariate in late winter to early spring, and emerge as adults that aggre-
gate in summer on preblossom host plants to mate and subsequently oviposit. A single
male specimen of Eurytoma veronia Bugbee (Hymenoptera: Eurytomidae) was reared
from a puparium of N. wilsoni as a solitary, larval-pupal endoparasitoid.
Key Words: Insecta, Neaspilota, Hazardia, Haplopappus, Asteraceae, nonfrugivorous
Tephritidae, biology, taxonomy of immature stages, flower-head feeding,
monophagy, seed predation, parasitoids
Revision ofthe genus Neaspilota (Diptera: scribes the immature stages and hfe history
Tephritidae) by Freidberg and Mathis (1986) of a second species from Cahfomia, Neas-
facilitated determination of specimens reared pilota wilsoni Blanc and Foote.
from California Asteraceae (Goeden 1989),
Materials and Methods
and stimulated several life-history studies, in-
eluding one on N. viridescens Quisenberry The present study was based in large part
(Goeden and Headrick 1992). This paper de- on dissections of subsamples of flower
—
898 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
heads of Hazardia squarrosa (Hooker and each of six, clear-plastic, petri dishes pro-
Amott) E. Greene [= Haplopappus squar- visioned with a flattened, water-moistened
rosus (Hickman 1993)] collected during pad of absorbant cotton spotted with honey
1989-1995 from three locations in San Di- (Headrick and Goeden 1994) for observa-
ego Co. in the manner described by Goeden tions of their courtship and copulation be-
W
(1985, 1989, 1992): of Dogpatch along havior.
State Hwy. 94 at 760-m elevation; Los Ter- Plant names used in this paper follow
renitos N of Interstate Hwy. 8 and junction Hickman (1993) and Bremer (1994); te-
with State Hwy. 79 at 1,000 m; Kitchen phritid names and adult terminology follow
Creek Road above Cibbetts Flat Camp- Foote et al. (1993). Terminology and tele-
ground at ca. 1,300 m, Cleveland Nat. For- graphic format used to describe the imma-
est. One-liter samples of excised, immature ture stages follow Goeden et al. (1998a, b),
and mature flower heads containing eggs, Goeden and Headrick (1992), Goeden and
larvae, and puparia were transported in Teerink (1997; 1998; 1999a, b, c), Teerink
cold-chests in an air-conditioned vehicle to and Goeden (1998, 1999), and our earlier
the laboratory and stored under refrigera- works cited therein. Means ± SE are used
tion for subsequent dissection, photogra- throughout this paper. Voucher specimens
phy, description, and measurement. Six of N. wilsoni adults and immature stages
eggs, four first-, three second-, and four and its parasitoid reside in the research col-
third-instar larvae, and three puparia dis- lections of RDG.
sected from flower heads were preserved in
70% EtOH for scanning electron micros- Results and Discussion
Taxonomy
copy (SEM). Additional puparia were
placed in separate, glass shell vials stopped Adult. Neaspilota wilsoni was de-
with absorbant cotton and held in humidity scribed by Blanc and Foote (1961) from a
chambers at room temperature for adult and male from Jacolitas Canyon, Fresno Co.,
parasitoid emergence. Specimens for SEM California. Foote and Blanc (1963) and
were hydrated to distilled water in a de- Foote et al. (1993) pictured the unpattemed
creasing series of acidulated EtOH. They wing, as did Freidberg and Mathis (1986)
were osmicated for 24 h, dehydrated along with drawings of the lateral aspect of
through an increasing series of acidulated the head, male right foretarsus, epandrium
EtOH, critically point dried, mounted on and cerci, aculeus and its apex enlarged,
stubs, sputter-coated, and studied with a and spermatheca.
—
JEOL JSM C-35 SEM in the Department of Immature stages. The egg, larvae, and
Nematology, University of California, Riv- puparium heretofore have not been de-
erside. scribed nor illustrated.
Most adults reared from isolated puparia Egg: Twenty-three ova dissected from
were individually caged in 850-ml, clear- five, 40+ days-old A'^. wilsoni plus 10 eggs
plastic, screened-top cages with a cotton dissected from field-collected flower heads
wick and basal water reservoir and provi- were white, opaque, smooth, elongate-ellip-
sioned with a strip of paper toweling im- soidal, 0.93 ± 0.009 (range, 0.82-1.00) mm
pregnated with yeast hydrolyzate and su- long, 0.24 ± 0.006 (range, 0.18-0.25) mm
crose. These cages were used for studies of wide, smoothly rounded at tapered basal
mm
longevity and sexual maturation in the in- end (Fig. lA); pedicel conical, 0.02
sectary of the Department of Entomology, long, completely circumscribed by different
University of California, Riverside, at 25 ± sized, mostly circular, shallow aeropyles,
rc, and 14/10 (L/D) photoperiod. Single through which the spongy inner layers of
pairs of virgin males and females obtained the chorion are readily visible (Fig. IB).
from emergence cages also were held in The egg ofN. wilsoni is similar in shape.
VOLUME 101, NUMBER 4 899
Fig. 1. Egg ofNeaspilota wilsoni: (A) habitus, anterior end to left; (B) pedicel, aeropyles.
but about 40% longer and 30% wider than (Fig. 2A-7); median oral lobe laterally flat-
that of N. viridescens (Goeden and Head- tened, apically pointed (Fig. 2A-8); pit sen-
rick 1992); moreover, the latter species has sillum laterad of dorsal sensory organ (Fig.
fewer aeropyles that are irregularly and less 2B-8); four verruciform sensilla circum-
densely spaced around its otherwise simi- scribe gnathocephalon medially (Fig. 2A-
larly-shaped and -sized pedicel. The eggs of 9); metathoracic lateral spiracular complex
12 species of Trupanea from California with a spiracle (Fig 2C-2) and four, verti-
studied to date, including T. femoralis cally aligned, verruciform sensilla poster-
(Thomson), T. nighcornis (Coquillett), and iorad (Fig. 2C-3); caudal segment with two
T. wheeleri Curran that occur in symphagy stelex sensilla, dorsolaterad and ventrolater-
with N. wilsoni in heads of H. squarrosa ad ofposterior spiracular plates, each stelex
(Goeden 1985, 1992, 1997), have pedicels sensillum basally ringed with one to three,
circumscribed by only one or two rows of upright, pointed acanthae (Fig. 2D-1); pos-
aeropyles (Goeden and Teerink 1999b and terior spiracular plate bears two ovoid ri-
mm
references therein). mae, ca. 0.006 in length (Fig. 2D-2),
First instar: White, elongate-cylindrical, and four interspiracular processes, each
bluntly rounded anteriorly and posteriorly with five to nine branches, longest measur-
mm
(not shown); minute acanthae, pointed to ing ca. 0.01 (Fig. 2D-3); intermediate
cylindrical and apically one- to five- sensory complex with one stelex sensillum
pronged, circumscribe intersegmental lines (Fig. 2D-4).
(Fig. 2A-1, C-l)s gnathocephalon smooth, The first instar is similar in general hab-
lacking rugose pads, but with pair ofprom- itus to that of N. viridescens (Goeden and
inent integumental petals dorsal to mouth- Headrick 1992). However, unlike N. viri-
hooks (Fig. 2A-2, B-1); dorsal sensory or- descens, the dorsal sensory organ of the
gan a dome-shaped papilla (Fig. 2A-8, B- first instar ofN. wilsoni is well defined (Fig.
2); anterior sensory lobe (Fig. 2A-4, B) 2A-3, B-2), as is the anterior sensory lobe
bears the terminal sensory organ (Fig. 2A- and integumental petal. The reported poor
5, B-3), pit sensory organ (Fig. 2B-4), lat- definition of N. viridescens may have re-
eral sensory organ (Fig. 2B-5), and supra- flected specimen condition, as the same dis-
lateral sensory organ (Fig. 2B-6); stomal section and preparation procedures as well
sense organ ventrolaterad of anterior sen- as scanning electron microscope were used
sory lobe (Fig. 2A-6, B-7), integumental with both species (but see descriptions of
petal fused laterally with stomal sense or- later instars below). Also, the pit sensory
gan (Fig. 2A-6, B-7); mouthhook bidentate organ, not visible with N. viridescens (Goe-
900 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. FirstinstarofA', wilsoni: (A)gnathocephalon, gnathocephalon,anterolateralview, I-minuteacanthae,
2- integumental petal, 3- dorsal sensory organ, 4- anterior sensory lobe, 5- terminal sensory organ, 6- stomal
sense organ, 7- mouthhook, 8- median oral lobe, 9- campaniform sensillum; (B) anterior sensory lobe, 1-
integumental petal, 2- dorsal sensory organ, 3- terminal sensory organ, 4- pit sensory organ, 5- lateral sensory
organ,6-supralateral sensoryorgan,7-stomalsenseorgan, 8-pitsensillum;(C)metathoracicspiracularcomplex,
1- minute acanthae, 2- spiracle, 3- campaniform sensillum; (D) caudal segment, 1- stelex sensilla, with basal
acantha, 2- rima, 3- intrespiracular process, 4- intermediate sensory complex, stelex sensillum.
den and Headrick 1992), is clearly seen in apparently are reduced to four in N. wilsoni
N. wilsoni (Fig. 2B-4). A fused integmental (Fig. 2D-1, D-4), two sensilla less than the
petal and stomal sense organ also was re- six reported for California Trupanea (Goe-
ported in first instar Trupanea vicina (Goe- den and Teerink 1999b and references
den and Teerink 1999b). therein). As another difference, the inter-
The cylindrical, apically pronged acan- spiracular processes of N. viridescens have
thae that circumscribe the intersegmental up to four branches (Goeden and Headrick
lines are the first of this type reported from 1992); whereas, N. wilsoni possesses up to
tephritid larvae, as are the lateral stelex sen- nine branches (Fig. 2D-3). Needless-to-say,
silla on the caudal segment that are basally comparative studies of first instars of ad-
ringed with upright, pointed acanthae. The ditional Neaspilota spp. are needed to better
10 stelex sensilla reported to surround the clarify these contradictory, apparent or real
posterior spiracular plate of the first instar distinctions, which hopefully will be forth-
of N. viridescens (Goeden and Headrick coming in our next several papers.
1992) are suspect in number. These sensilla Second instar: White, elongate-cylindri-
VOLUME 101. NUMBER 4 901
cal, tapering anterioriy, rounded posterioriy lindrical, like the first instars of both spe-
(not shown), minute acanthae circumscribe cies. Another distinction between the first
intersegmental lines (Fig. 3A-1, D-l)s and second instar of N. wilsoni is the pres-
gnathocephalon conical (Fig. 3A); dorsal ence of seven oral ridges with dentate mar-
sensory organ a dome-shaped papilla (Fig. gins in the latter instar, also pictured and
3A-2, B-1); anterior sensory lobe (Fig. 3A- equal to, but unquantified until now (un-
3, B) bears the terminal sensory organ (Fig. published data), for the second instar of N.
3A-4, B-2), lateral sensory organ (Fig. 3B- viridescens (Goeden and Headrick 1992).
3), supralateral sensory organ (Fig. 3B-4), The stomal sense organs of both species
and pit sensory organ (Fig. 3B-5); stomal bear three sensory structures, described as
sense organ prominent ventrolaterad of an- conical in N. viridescens (Goeden and
terior sensory lobe (Fig. 3B-6, C-1); mouth- Headrick 1992), but which appear papilli-
hook bidentate (Fig. 3A-5, C-2); median form in N. wilsoni (Fig. 3B-6, C-1). The
oral lobe laterally flattened (Fig. 3C-3); two mouth hooks of the second instar ofN. vir-
pit sensilla laterad of dorsal sensory organ idescens are tridentate (Goeden and Head-
(Fig. 2B-7); labial lobe with two pore sen- rick 1992); whereas, those ofA^. wilsoni are
silla (Fig. 3C-4); single row ofintegumental bidentate (Fig. 3A-5, C-2). Similar, appar-
petals dorsal to mouthhook (Fig. 3A-6, B- ent interspecific differences in dentation
8, C-5); most oral ridges lateral to mouth- were noted among mouth hooks of second
hooks dentate along posterior margins (Fig. instar Trupanea spp. (Goeden and Teerink
3A-7, C-6); pit sensilla circumscribe gnath- 1999b and references therein). Finally, the
ocephalon medially (Fig. 2A-8); anterior stelex sensilla on the caudal segment of the
thoracic spiracle with three to four oblong second instar of N. wilsoni are not basally
papillae; lateral spiracular complex of me- ringed by pointed acanthae (Fig. 3F-1), as
sothorax with a spiracle (Fig. 3D-2) and in the first instar (Fig. 2D-1); the interspi-
three verruciform sensilla in vertical row racular processes each bear two to six (Fig.
(Fig. 3D-3); caudal segment with at least 3E-2), not five to nine branches (Fig. 2D-
two stelex sensilla (not shown), dorsolater- 3), respectively; and the intermediate sen-
ad and ventrolaterad of posterior spiracular sory complex consists of a medusoid and a
plates; posterior spiracular plate bears three stelex sensillum (Fig. 3F-1, 2), not just a
mm
ovoid rimae, ca. 0.014 in length (Fig. single stelex sensillum (Fig. 2D-4). In con-
3E-1), and four interspiracular processes, trast, the interspiracular processes of the
each with two to six branches, longest mea- second instar of N. viridescens were noted
mm
suring 0.011 (Fig. 3E-2); intermediate to be two-branched at most (Goeden and
sensory complex with a stelex sensillum Headrick 1992). Again, comparative studies
(Fig. 3F-1) and a medusoid sensillum (Fig. of second instars of additional species of
3F-2). Neaspilota are needed in order to help dis-
Few differences were noted between the tinguish real from apparent, intra- and in-
first and second instars, besides the acqui- terspecific differences.
sition of prothoracic spiracles and another Third instar: Mostly white, with last two
rima on the posterior spiracular plate nor- or three segments darkened, elongate-cylin-
mal to all tephritid larvae (Headrick and drical, tapering anteriorly, posterior spirac-
Goeden 1998). The dorsal sensory organ is ular plate on caudal segment flattened and
less well defined in the second instar (Fig. upturned dorsally ca. 60°, minute acanthae
3A-2, B-1) than in the first instar (Fig. 2A- circumscribe all intersegmental areas and
3, B-2). However, the habitus ofthe second all abdominal segments except pleura (Fig.
instar of N. viridescens was described as 4A, F)s gnathocephalon conical (Fig. 4B);
barrel-shaped (Goeden and Headrick 1992); apparently six oral ridges laterad of mouth
whereas, that of N. wilsoni, is elongate cy- lumen, all with serrated margins (Fig. 4B-
902 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Second instarofNeaspilota wilsoni: (A) gnathocephalon and prothorax, anterolateral view, 1- minute
acanthae, 2- dorsal sensory organ, 3- anterior sensory lobe, 4- terminal sensory organ, 5- mouth hook, 6-
integumental petal, 7- oral ridge, 8- pit sensilla, (B) anterior sensory lobe, 1- dorsal sensory organ, 2- terminal
sense organ, 3- lateral sensory organ, 4- supralateral sensory organ, 5- pit sensory organ, 6- stomal sensory
organ, 7- pit sensillum, 8- integumental petal, (C) gnathocephalon, ventrolateral view, 1- stomal sense organ, 2-
mouthhook, 3- median oral lobe, 4- labial lobe, 5- integumental petal, 6- oral ridge; (D) metathoracic spiracular
complex, 1- minute acanthae, 2- spiracle, 3-campaniform sensilla: (E)caudal segment, 1-rima,2- interspiracular
process; (F) intermediate sensory complex, 1- stelex sensillum. 2- medusoid sensillum.
VOLUME 101, NUMBER 4 903
^~i
904 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1); dorsal sensory organ a dome-shaped pa- tures, described as several cone-shaped sen-
pilla (Fig. 4B-2, C-1), pit sensillum laterad silla in N. viridescens (Goeden and Head-
of dorsal sensory organ (Fig. 4C-2); ante- rick 1992), but which appear papilliform
rior sensory lobe (Fig. 4B-3, C) bears the and pit-type in N. wilsoni (Fig. 4C-7). Un-
terminal sensory organ (Fig. 4C-3), pit sen- like the second instar of N. wilsoni, the
sory organ (Fig. 4C-4), lateral sensory or- third instar apparently has six oral ridges,
gan (Fig. 4C-5), and supralateral sensory all with dentate margins, the same as par-
organ (Fig. 4C-6); stomal sense organ tially pictured, but unquantified, and also
prominent ventrolaterad of anterior sensory now reported for the third instar of N. vir-
lobe (Fig. 4C-7); mouth hook bidentate idescens (Goeden and Headrick 1992 and
(Fig. 4B-4); median oral lobe not clearly unpublished data). This may represent a
visible; prothorax circumscribed anteriorly loss from the second instarofone oral ridge
with minute acanthae (Fig. 4B-5); anterior in the third instar of both species; however,
thoracic spiracle bears three oblong papillae distinguishing and accurately counting
(Fig. 4D); metathoracic lateral spiracular these structures is rendered problematic by
complex with a spiracle (Fig. 4E-1) and degree and angle of exposure of the gnath-
four verruciform sensilla (Fig. 4E-2); ab- ocephalon as well as by a current lack of a
dominal lateral spiracular complex with a precise definition for and defined morpho-
spiracle (Fig. 4F-1), three verruciform sen- logical limits to these structures (Tesky
silla (Fig. 4F-2); caudal segment circum- 1981). The third instars of Trupanea im-
scribed by minute acanthae; stelex sensilla, perfecta, T. jonesi, T. nigricornis, T. pseu-
dorsolaterad, laterad and ventrolaterad of dovicina, T. signata, and T. wheeleri also
posterior spiracular plate; posterior spirac- bear serrated oral ridges (Goeden and Teer-
ular plate bears three ovoid rimae, ca. 0.013 ink 1997; 1998, 1999b; Goeden et al.
mm
in length (Fig. 4G-2), and four inter- 1998a; Knio et al. 1996a; Teerink and Goe-
spiracular processes (Fig. 4G-1), each with den 1998). Also, like the second instar, the
three to four branches each with one to mouthhooks of third instar N. wilsoni are
three, apical teeth, longest branch measur- bidentate (Fig. 4B-4); whereas, those of N.
ing 0.012 mm; intermediate sensory com- viridescens are tridentate (Goeden and
plex with a medusoid (Fig. 4H-1) and a ste- Headrick 1992). Such interspecific differ-
lex sensillum (Fig. 4H-2). ences in dentation are supported by our
The habitus of the third instar of N. wil- findings that the mouth hooks ofthird instar
soni generally is like that reported for N. Trupanea vicina are bidentate; whereas,
viridescens by Goeden and Headrick those of 12 other congeners examined from
(1992), except that minute acanthae circum- California are tridentate (Goeden and Teer-
scribe all intersegmental areas and all ab- ink 1999b and citations therein). The ab-
dominal segments except the pleura in the dominal lateral spiracular complex is like
former species; whereas, in the latter spe- that pictured and described for third instar
cies the intersegmental areas are free of Stenopa affitiis Quisenberry (Goeden and
acanthae. Like the second instar of N. wil- Headrick 1990).
soni, the dorsal sensory organ is less well Puparium: Mostly white to yellow, with
defined in the third instar (Fig. 4B-2, C-1) posterior two-three segments grayish to
than in the first instar (Fig. 2A-3, B-2). The blackened posteriorly, broadly ellipsoidal
dorsal sensory organ in the third instar N. and smoothly rounded at both ends, minute
viridescens also is not well defined, but this acanthae circumscribe intersegmental lines;
is not a generic character (unpublished anterior end bears the invagination scar
data), as proposed by Goeden and Headrick (Fig. 5A-1) and anterior thoracic spiracles
(1992). The stomal sense organs ofthe third (Fig. 5A-2); caudal segment circumscribed
instars of both species bear sensory struc- by minute acanthae (Fig. 5B-l),'two stelex
VOLUME 101, NUMBER 4 905
Fig. 5. Puparium ofNeaspilota uilsoni: (A) anterior end, 1- invagination scar. 2- anterior thoracic spiracle,
(B) caudal segment, 1- minute acanthae, 2- stelex sensillum, 3- rima, 4- interspiracular process.
sensilla, dorsad and ventrad of posterior soni, nor any other species of Neaspilota,
spiracular plates (Fig. 5B-2); posterior spi- has ever been reared from the genus Co-
racular plate bears three broadly elliptical reopsis in California (Goeden 1989), even
rimae (Fig. 5B-3), and four interspiracular during intensive and extensive field study
processes, each with 3 to 4 branches (Fig. of two principal associates of Coreopsis
5B-4); intermediate sensory complex with spp., Dioxyna picciola (Bigot) (Headrick et
a medusoid sensillum and a stelex sensil- al. 1996) and Trupaneajonesi Curran (Goe-
lum. Six puparia averaged 2.75 ± 0.16 den et al. 1998a).
(range, 1.99-3.11) mm in length; 1.24 ±
mm Biology
0.03 (range, 1.12-1.32) in width. —
Egg. In 17 closed, preblossom, apical
Distribution and Hosts
or penultimate, immature flower heads of
The distribution ofN. wilsoni as mapped H. squarrosa, 19 eggs were inserted be-
by Foote et al. (1993) restricted this tephri- tween or through the phyllaries, usually into
tid to California in the United States north or between the inner phyllaries, and per-
of Mexico. The distribution of this true pendicular or at a slight angle (ca. 15°) to
monophage probably also extends south- the receptacle (Fig. 6A). All eggs were ovi-
ward into northern Baja California like that posited pedicel-last. Only one flower head
of its only known host plant, H. squarrosa contained a floret damaged by oviposition.
(Freidberg and Mathis 1986, Goeden 1989, The diameters ofthe receptacles of 17 flow-
Foote et al. 1993), commonly known as er heads containing eggs averaged 1.8 ±
"saw-toothed goldenbush" (Hickman 0.2 (range, 0.85-3.75) mm, and these heads
1993), which belongs to the subtribe Soli- contained an average of 1.1 ± 0.1 (range,
dagininae in the tribe Astereae (Bremer 1-2) eggs—.
1994). We here reiterate our belief(Goeden Larva. Upon eclosion, first instars tun-
1989) reported by Foote et al. (1993) that neled through an inner phyllary into the
the unconfirmed record for Coreopsis cal- apical end of an ovule. An average of 1.3
liopsidea Gray in Freidberg and Mathis ±0.2 (range, 1-3) first instars was found
(1986), which they suspected was a host, feeding within each of 10, closed, preblos-
probably is not valid, owing to its distant som flower heads, the receptacles of which
mm
relationship to Hazardia; it belongs to the averaged 1.6 ± 0.2 (range, 1.1-2.0) in
tribe Heliantheae, subtribe Coreopsidinae diameter. These infested heads contained an
M
(Bremer 1994). Furthermore, neither wil- average of 17 ± 1 (range, 15-18) ovules.
906 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON