Quametopia , a new genus of Nearctic Anthomyzidae ( Diptera ) , with description of two new species , immature stages and life history

Quametopia gen. n. is established on the basis of recent phylogenetic analysis of taxa formerly assembled under the genus Mumetopia Melander, 1913 to include M. terminalis (Loew, 1863) and two new closely related Nearctic species of Anthomyzidae. The new genus is diagnosed and its phylogenetic relationships discussed. Quametopia terminalis (Loew, 1863) comb. n. is transferred from Mumetopia and redescribed based on revision of the type material (lectotypes of Anthophilina terminalis Loew, 1863 and its synonym Mumetopia nitens Melander, 1913 are designated) and other extensive material. Quametopia clintonia sp. n. and Q. amplistylus sp. n. (both from Canada, USA) are described and relationships of all three Quametopia species discussed. Immature stages of Q. terminalis and Q. clintonia sp. n. obtained by means of adult-to-adult rearing are described (1stand 2nd-instar larvae for the first time in the family Anthomyzidae) and illustrated. Keys to adults, eggs, larvae and puparia of Quametopia species are presented. Biology (habitat and host-plant associations, life history) of Q. terminalis and Q. clintonia sp. n. are studied in detail and their ecological separation demonstrated. Distribution of all Quametopia species is reviewed.


INTRODUCTION
The recent study by Rohá ek & Barber (2009) on the phylogenetic relationships of species associated under the genus Mumetopia Melander, 1913 revealed that neither of its two other known species, viz.M. nigrimana (Coquillett, 1900) and M. terminalis (Loew, 1863), are congeneric with the type species Mumetopia occipitalis Melander, 1913.The species M. nigrimana (Coquillett, 1900) and a number of its undescribed relatives proved to form a distinct unnamed genus-group taxon which belongs, as does the genus Mumetopia in a new restricted concept, to the Chamaebosca group of genera as defined by Rohá ek & Barber (2009: 212) forming there a sistergroup to Stiphrosoma Czerny, 1928.On the other hand, M. terminalis had been considered unrelated to the Chamaebosca clade and was excluded from the genus Mumetopia as a species whose generic affiliation and relationships required further detailed study (see Rohá ek & Barber, 2009).
Moreover, the examination of extensive material of Mumetopia terminalis from various parts of the Nearctic Region revealed that there were, in fact, three closely allied species involved under this name.To confirm their validity, successful rearing experiments were performed which obtained all preimaginal stages and first data for habitat and host-plant preferences of two of these species.These discoveries also raised the necessity to revise all available type specimens of Anthophilina terminalis Loew, 1863 (transferred to Mumetopia by Melander, 1913: 294) and of its synonym Mumetopia nitens Melander, 1913 so that these names can be correctly applied to the species recognized here.
An analysis of morphological characters (particularly those of the male and female terminalia) found that this species complex cannot be affiliated with any of the described genera of Anthomyzidae.Therefore, a new genus Quametopia is established below to accommodate these three species and its phylogenetic affinities are discussed on the basis of analysis of a set of characters used as criteria in diagnoses of anthomyzid genera as standardized by Rohá ek (2006Rohá ek ( , 2009)).All three members of Quametopia are (re)described in detail, including preimaginal stages (the first-and second-instar larvae are described for the first time in the family Anthomyzidae) in two of them, keyed, and their relationships, biology (including life history and host-plant preferences) and distribution discussed.

MATERIAL AND METHODS
The material examined is deposited in institutional and private collections as follows: AMNH -American Museum of Natural History, Division of Invertebrate Zoology, New York, NY, U.S.A.; BYUC -Brigham Young University, Monte L. Bean Life Science Museum, Provo, UT, U.S.A.; CASC -California Academy of Sciences, Department of Entomology, San Francisco, CA, U.S.A.; CNCI -Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa, ON, Canada; CSCA -California State Collection of Arthropods, Sacramento, CA, U.S.A.; DEBU -University of Guelph Insect Collection, School of Environmental Sciences, University of Guelph, Guelph, ON, Canada; EMUS -Utah State University, Logan, UT, U.S.A.; INHS -Illinois Natural History Survey, Champaign, IL, U.S.A.; LEMQ Abdomens of a large number of specimens were detached and genitalia dissected.After examination, all dissected parts were put into plastic tubes containing glycerine and pinned below the respective specimens; this is indicated in the text by the abbreviation "genit.prep.".
Quametopia clintonia sp.n. was first suspected to be a distinct species when swept in wooded areas where I. capensis was absent.Collection efforts subsequent to 21 July 2009 were directed at identifying the host plant and, despite the relatively sparse occurrence of this fly and considerable heterogeneity of the plant communities, Clintonia borealis (Aiton) Raf.(Liliaceae) (Fig. 105) emerged as the most probable host plant.
Field collections of litter and soil, and stems of Impatiens capensis.Samples of litter and soil were taken in the spring of 1999 and 2002 at the Bristol Place Park (Sault Ste.Marie, Ontario) site where adults of Q. terminalis occur in a stand of I. capensis.On 10 May 1999, five samples of ~13×18 cm were cut from the patch and maintained in plastic boxes at 20°C, 50-70% RH, and 16L : 8D photoperiod (standard conditions).On 25 May 2002, a horticultural bulb planter was used to cut 14 cores of ~5.8 cm diameter each.The cores were pushed into plastic cylinders, placed in small plastic bags, and maintained at 25°C, 30-50% RH, and 16L : 8D photoperiod (the relative humidity within the plastic bags was considerably higher).
A small sample of dried, fallen stems of I. capensis from the previous season was collected from the Birchwood Park (Sault Ste.Marie) site on 1 May 2010.The stems were transferred to plastic boxes (Fig. 5) and held in standard conditions until 31 July 2010.A slightly larger sample of stems was collected on 17 May 2010 and handled in the same fashion.A subsample of 62 stems (only those including roots, Figs 6, 7) was dissected in search of immature Q. terminalis.The maggots found were transferred to thawed (from frozen supply) sections of I. capensis stems until pupariation (larvae and puparia were handled as described below).
Rearing of Q. terminalis.On 24 June 2002, 12% and 24& from the Bristol Place Park site were divided between two small flight cages, each with a cylindrical soil core with I. capensis plants, provided with a bee pollen/sucrose mixture (1 : 1 by volume, ground in a coffee grinder, sprinkled on the dampened screen), regularly misted with water, and held in standard conditions.By early July, eggs were observed on damaged leaves of the I. capensis plants.The soil cores with plants were then replaced by a plastic Petri plate (9 cm diameter) with wet white sand and I. capensis leaves on the surface.By 22 July, the Petri plates with wet sand and rotting leaves had many larvae but since high mortality was observed, excess liquid was decanted off, and many larvae preserved in ethanol.Subsequently, for oviposition and for general rearing, short sections (~3 cm) of I. capensis stems were split longitudinally and laid on the sand, outside surface facing up, in place of leaves.Larvae readily colonized these stems, rasping the inner flesh and also wedging themselves between the outer epidermis and the inner tissue.
A cup cage (plastic drinking cup with screen bottom, a stoppered hole in the side, inverted on a Petri plate lid filled with moist sand) replaced the flight cages when only 4% and 6& remained (12 August 2002).Oviposition was later restricted to a small 5 cm-diameter Petri plate (Falcon ® 1006, Becton Dickinson Labware, Lincoln Park, NJ, USA) on the floor of the cage that was in turn lined with filter paper, the small plate supplied with moist sand and short sections of split stems of I. capensis (Figs 1,2).7. Rearing methods of Quametopia species: 1 -laboratory equipment for adult maintenance and oviposition (A -stem of Impatiens capensis, B -cut and split stem, C -pieces of split stem on sand in small (internal) Petri plate, D -bottom (external) Petri plate lined with filter paper, E -cup with screen bottom and lateral hole, F -rubber stopper, G -mixture (1:1) of ground sucrose and bee pollen, H -assembled cup cage); 2 -cup cage with adults of Q. terminalis used for oviposition; 3 -Petri plates with pieces of both Impatiens and Clintonia borealis for oviposition-choice trial (egg arrowed); 4 -a piece of dead Clintonia leaf with 3 larvae and 1 puparium inside (arrows); 5 -rearing boxes with dead stems of Impatiens; 6 -dead stems of Impatiens with roots (arrow), used for dissections; 7 -detail of hollow roots containing overwintering dipterous larvae (arrow).Photo: K.N.Barber.
The small oviposition plate was replaced on a 3-4 day cycle and the eggs cleaned with a 2% solution of formaldehyde, rinsed in water, and transferred to damp filter paper in another small Petri plate.The plates of eggs were held under standard conditions, checked daily, and the filter paper moistened as required.Neonate larvae were transferred to split stems of I. capensis in small Petri plates (no sand), the stems checked daily, hydrated and replaced as required as the larvae developed.
Puparia were cleaned in dilute bleach, rinsed in water then individually held in small plastic pill vials at 20°C.Larvae not pupariating by November were held over a simulated winter by placing them in small Petri plates with sections of I. capensis stems and maintained at 10°C, 12L : 12D, 3 November to 9 December, and then constant dark at 2°C until 17 March 2003, then returned to standard conditions.The plates were checked, cleaned, and the stems replaced occasionally (from frozen supply) during the overwintering period.Other rearings of Q. terminalis used 5% and 5& and 4% and 4& collected at Birchwood Park, 29 July and 27 August 2010, respectively, principally to provide additional eggs and larvae.
Rearing of Q. clintonia sp.n.On 23 August 2009, live adults of Q. clintonia sp.n. were collected from near Searchmont, Ontario (Whitman Dam Road -2%, 2&, Ranger Lake Road -1&).Cup cages and small Petri plates with wet sand and oviposition substrates were used and held at standard conditions.
Based on an oviposition trial (see below), short sections (~3 cm) of C. borealis petioles were chosen as the initial larval rearing substrate.Neonate larvae were transferred to petioles where they readily entered the conductive channels of the severed ends.Surviving late-second instars were transferred to pieces of C. borealis leaf beginning 24 September.Pieces of leaf (~3×3 cm) were variously frozen then thawed, lightly heated, or allowed to "season" in water for a day or more before presentation.The limp leaf material adhered to the small Petri plate and larvae readily penetrated between the leaf surfaces along the severed edges (Fig. 4).The leaf pieces were hydrated as required.Additional rearings of Q. clintonia sp.n. used 2% and 2& collected at Batchawana Provincial Park, 20 September 2009, and 7% and 5& collected at Lake Superior Provincial Park, 29 August 2010, principally to provide additional eggs and first-instar larvae for study.
Oviposition choices -Q.terminalis and Q. clintonia sp.n.On 23 August 2009, 2% and 2& of Q. terminalis were collected from Birchwood Park (Sault Ste.Marie) for comparison with those of Q. clintonia (above).Small Petri plates with moist sand and sections of plant substrates were used in the two cup cages of 2% and 2& each for Q. clintonia and Q. terminalis and the lone female of Q. clintonia in a third cup cage.Choices of oviposition substrate were: a section of I. capensis stem (~3 cm longitudinally split, outer surface up); C. borealis petiole (~3 cm of V-shaped basal section of leaf, inverted with ventral surface up); and a piece of C. borealis leaf blade (~1×3 cm, dorsal surface up) (Fig. 3).Two oviposition periods of 3 days each were pooled for a total of 6 days beginning from 24 August.
Etymology.The new genus is named Quametopia (= an abbreviated conjunction of Qua[m + Mu]metopia, gender feminine) because its type species was originally included in Mumetopia Melander, 1913, a similarly looking [Latin quam = like, as] genus of Anthomyzidae.

Diagnosis
(1) Head slightly higher than long.
The comparison of the characters listed in the above diagnosis of Quametopia with those known in all other described genera of Anthomyzidae, contrary to our expectations, resulted in finding that although Quametopia surely does not belong to the Chamaebosca clade, it is obviously more closely allied to it than Cercagnota Rohá ek & Freidberg, 1993 (recognized as sister group to the Chamaebosca clade by Rohá ek & Barber, 2009: 212-213), particularly because of the similarly formed female postabdomen.The following synapomorphies demonstrate this relationship: Female T7+S7 dorsomedially membranous or divided; female S8 short and posterodorsomedially (although not deeply) incised (however, in contrast to the members of the Chamaebosca clade, it is posteromedially protruding as in the unrelated genus Amygdalops Lamb, 1914 -cf.Rohá ek, 2004Rohá ek, , 2008)); posterior internal sclerites of female genital chamber well developed; spermathecae with surface spines (although different from those in representatives of the Chamaebosca group of genera).Also the formation of the female tergosternum T7+S7, where the original S7 is partly delimited, is more similar to that in some members (Stiphrosoma and Mumetopia nigrimana-group, see Rohá ek & Barber, 2009) of the Chamaebosca group of genera than to that of Cercagnota where S7 is completely integrated into the tergosternal ring without traces of its original boundary.Considering the characters discussed above, Quametopia can be considered a sister group to the Chamaebosca group of genera instead of Cercagnota (but for differences in the male genitalia see below).
Only a few of the male genital characters may indicate affinity of Quametopia to the Chamaebosca clade, e.g. the sclerotized and dark-pigmented dorsal section of the aedeagal part of folding apparatus (occurring in several genera of the Chamaebosca clade but also in Cercagnota) or the prolonged dorsolateral corners of the medandrium (being, however, much longer in some genera of the Chamaebosca clade).However, generally, the synapomorphic features of the male genitalia of the Chamaebosca group of genera are lacking in Quametopia where only the most plesiomorphic condition can be found, viz. in postgonite (slender and simple, with only single seta), filum of distiphallus (bare and formed by 2 ribbon-shaped sclerites), saccus of distiphallus (at most with sparse microspinules and flat tubercles), male cercus (simple, not projecting), caudal process of transandrium (absent), gonostylus (disparate and micropubescent).
The genus Quametopia includes three species, all occurring in the Nearctic Region: Q. terminalis, Q. clintonia sp.n. and Q. amplistylus sp.n.Although their morphological differences are relatively small, some of them (mainly those in genitalic structures of both sexes) proved to be stable and enabled confident identification of these closely related species.Moreover, the recognized species (at least as regards Q. terminalis and Q. clintonia sp.n.) seem to differ significantly in their biology (habitat preference and host-plant association).Quametopia terminalis is most different from the remaining two species and apparently forms a sister-taxon to the Q. clintonia + Q. amplistylus pair which shares several synapomorphies, e.g.shortened posterior lobe of gonostylus, reduced anterior lobe of pregonite, elongate dorsal pair and reduced lateral pair of internal sclerites in female genital chamber.The species of Quametopia are keyed and treated in fur-ther detail according to their presumed relationships below.
posterior spiracle lacking peritreme but having 2 spiracular slits.Interestingly, the unpaired median mouthhook has hitherto not been recorded in other families of Opomyzoidea but, following Ferrar (1987), the 1st-instar larvae were only described in three of its families, viz. in Agromyzidae, Aulacigastridae and Opomyzidae (the latter is considered to be the sister-group of Anthomyzidae, cf.Rohá ek, 1998) all of which have normally paired mouthhooks in this larval instar; the only exception is an extremely modified larva of Phytobia cambii (Hendel, 1931) (Agromyzidae), a cambium-miner, where all instars have an unpaired median mouthhook.The 2nd-instar larva, on the contrary, is already amphipneustic and its cephalopharyngeal skeleton (including paired mouthhooks) is similar to that of 3rd-instar larva but its anterior spiracle is simply clavate (with single slit) and the poste-rior spiracle has 2 spiracular slits as in that of the 1stinstar larva but with a well-developed peritreme.
Eggs of Quametopia species clearly have a surface structure different from those hitherto known in Anthomyzidae.Both the longitudinal elevated ribs (ridges, ledges) and the finely reticulate chorion surface have already been described in Anthomyza macra (cf.Rohá ek, 1986) but the number and position of these ribs is different (8 in Anthomyza macra, 4 in Quametopia spp.) as also is the extent of reticulation between them.The fine reticulate chorion surface structure on the ventral side of the egg also occurs in Anagnota coccinea (cf.Rohá ek & Freidberg, 1993) while longitudinal ribs (laterodorsal and dorsal) are also developed in the afrotropical Barbarista guttata (see Rohá ek, 1993).However, eggs of Quametopia spp.differ from all above-Figs 17-21.Quametopia terminalis (Loew), male (Canada: Ontario): 17 -apex of filum, subventrally (widest extension); 18aedeagal complex, laterally; 19 -hypandrial complex, ventrally (postgonite omitted); 20 -ditto, laterally; 21 -transandrium, caudally.Scales: Fig. 17  mentioned taxa in having transverse reticulation between ribs also on the dorsal side and even laterally.
The 3rd-instar larva is known in several genera of Anthomyzidae but only structures of the cephalopharyngeal skeleton, anterior and posterior spiracles were described in sufficient detail to allow some comparison.The cephalopharyngeal skeleton in Quametopia spp.has mouthhooks most similar to those of Stiphrosoma sabulosum (Haliday, 1837) (see Nye, 1958), Paranthomyza nitida Meigen, 1838) (see Rohá ek, 2009) and Anagnota bicolor (Meigen, 1838) (see Rohá ek & Freidberg, 1993) in having its middle part distinctly projecting ventrally; it differs, however, from these species by the posterior part being also extended ventrally so embracing the dental sclerite both anteriorly and posteriorly.Only the mouthhook of A. bicolor is posteriorly somewhat dilated but in this species this end is also deeply incised posteriorly (cf.Rohá ek & Freidberg, 1993: Fig. 23).The intermediate sclerite (im) of Quametopia differs from those known in other genera in having the ventral connecting bridge divided into two parts, a short more sclerotized and darker posterior and a long and paler anterior.Only P. nitida (see Rohá ek, 2009) has this structure somewhat similar.The latter species and A. bicolor also have the most similar pharyngeal sclerite differing only in the larger dorsal apodeme on the ventral cornua.The anterior spiracular process of Quametopia spp.resembles most those of Anthomyza and Anagnota in having a palmate shape with 5-6 papillae.Note: S. sabulosum may also be similar in this structure having 3-5 (usually 4, but Nye, 1958 illustrated only 3) finger-like papillae.The posterior spiracle is most similar to that of P. nitida (cf.Rohá ek, 2009) but of the same general structure as in other Anthomyzidae.It is to be stressed that the dorsolateral bulges of the last abdominal segment (in front of posterior spiracles) could be an additional diagnostic character of Quametopia because it can also be recognised in the puparium (see below) and was hitherto not found in puparia (posterior part of 3rd-instar larva has only been known in Typhamyza, see Rohá ek & Freidberg, 1993) of other genera (Anagnota, Anthomyza, Paranthomyza, Typhamyza).

Redescription
Male.Total body length 1.68-2.79mm; dark to blackish brown, relatively shiny; only extremities and fore head largely yellow (as in Fig. 10).Head slightly higher than long, subquadrate in profile but anteriorly somewhat rounded, bicolourous, yellow and dark brown.Frons dark brown posteriorly, yellow anteriorly; its yellow anterior third to almost half reaching at least to anterior (shorter) ors but usually distinctly behind it on orbit and yet more so medially and is darker (ochreous) posteriorly but bright to light yellow anteriorly (around bases of antennae).Frontal triangle large, reaching to level of anterior ors, brown or with paler brown anterior corner, sparsely greyish brown microtomentose anteriorly up to anterior ocellus and more densely so on ocellar triangle, but with bare shiny spots in posterior corners on both sides of ocellar triangle; narrow areas (stripes) between frontal triangle and orbits more densely grey microtomentose, duller and, consequently, seemingly lighter than adjacent areas.Frontal lunule reduced but visible, light yellow.Orbit posteriorly and around insertion of ors narrowly bare and shining, otherwise microtomentose, yellow anteriorly, brown posteriorly.Occiput slightly concave, dark to blackish brown, distinctly (but not densely) dark grey microtomentose except for bare and hence shining medial spot above foramen.Face yellow (sometimes darker yellow medially) and whitish microtomentose, dull (Fig. 8).Parafacialia and gena yellow and densely silvery white microtomentose; parafacialia internally and gena ventrally orange ochreous to pale-brown margined.Postgena and adjacent ventrolateral part of occiput pale brown.Mouthparts yellow to whitish yellow including palpus but clypeus brown or dark brown and some sclerites (anterior in particular) of proboscis also darkened (Figs 8,11).Cephalic chaetotaxy: pvt small and weak, strongly convergent; all macrosetae on frons very long but relatively thin; vti and oc distinctly longest of cephalic setae, at least as long as frons; vte and posterior ors subequal, long but shorter than vti or oc; 2 ors, posterior long, anterior ors variable in length but always markedly shorter (one-third to slightly more than half length) than posterior; 1 microsetula in front of anterior ors; 1-2 (rarely 3) pairs of minute medial microsetulae in front of frontal triangle; vi about as long as posterior ors but finer; subvibrissa well developed, up to three-quarters of vi length but markedly weaker; usually 6-9 short peristomal setae; about 3 small (one longer) setae on postgena with adjacent ventrolateral part of occiput finely setulose; postocular setulae (about 10) short, in single, relatively dense row.Eye very shortly and sparsely pilose, suboval; its longest (oblique) diameter about 1.4 times as long as shortest; smallest genal height 0.09 times as long as shortest eye diameter.Palpus whitish yellow only very basally brownish darkened, with 3-4 ventral setulae in addition to usual preapical seta.Antenna geniculate, yellow to whitish yellow (Fig. 11); 1st flagellomere lightest, with whitish cilia on anteroventral side about as long as those of arista.Arista about 1.9 times as long as antenna, brown with paler (ochreous or pale brown) two basal segments and distinctly long-ciliate.
Thorax slightly narrower than head.Mesonotum with scutum blackish brown, pleural sclerites especially ventrally, and sometimes humerus (= postpronotum) and notopleural area, brown.Mesonotum sparsely brownish grey microtomentose and shining; pleural part of thorax with somewhat denser microtomentum and rather subshiny.Subscutellum more densely microtomentose and duller than scutellum or metanotum.Thoracic chaetotaxy: 1 distinct but fine hu (as long as or longer than posterior npl); 2 npl, anterior long (almost as long as anterior dc but finer), posterior short; 1 prs well developed although thin, subequal or slightly shorter than hu; 1 sa, as long as prs; 1 pa ranging from slightly shorter than anterior npl to slightly shorter than posterior npl; 2 strong postsutural dc, anterior dc long (slightly longer and thicker than anterior npl), posterior dc very long, longest of thoracic setae (slightly longer than apical sc); a long row of 5-7 dc microsetae in front of anterior dc; ac microsetae sparse, in 4 rows on suture but lateral rows reduced, represented by only 1-2 (rarely 3) microsetae, so only medial rows longer and reaching beyond anterior dc; 2 sc, basal short and weak, apical sc long but slightly shorter and weaker than posterior dc; ppl reduced to pale hair-like microseta; 2 long stpl (posterior longer and thicker) plus several (3-7) upcurved setulae close (between, in front and below) to them in addition to several downcurved setae in ventralmost corner of sternopleuron.Scutellum moderately long, rounded triangular, with distinctly convex disc.Legs pale yellow to yellowish white (coxae in particular), only terminal segment of all tarsi dark brown.f1 with ctenidial spine as long as to distinctly longer than maximum width of t1 and inserted distinctly apicad of longest seta of posteroventral row; long setae in both posterodorsal and posteroventral rows relatively fine; fore    vein dm-cu slightly to distinctly shorter than apical portion of CuA1, the latter almost reaching wing margin; basal medial (bm) and posterior cubital (cup) cells closed; A1 short, not reaching wing margin; anal lobe and alula well developed.Wing measurements: length 1.77-2.78mm, maximum width 0.63-0.96mm; Cs3 : Cs4 = 1.16-1.72;r-m\dm-cu : dm-cu = 2.18-3.20.Haltere with large whitish knob and yellowish stem.
Female (Fig. 10).Similar to male unless mentioned otherwise.Total body length 1.89-3.06mm.Head colouring markedly different from that of the male (see Figs 9, 12).Frons largely dark brown, with yellow area restricted to anterior fourth, posteriorly maximally reaching to anterior ors, usually ending just above lunule.Face ochreous brown to brown, dull; bordering stripe of parafacialia and gena dark brown and sharply contrasting with yellow, silvery microtomentose remainder of gena and parafacialia.Mouthparts with dark brown clypeus and brownish palpus and proboscis, only labellae pale ochreous to whitish yellow.Also antenna distinctly darkened, with brownish inner side of pedicel (usually) and 1st flagellomere, the latter also brownish darkened on outer side below insertion of arista, otherwise orangeyellow.Lateral rows of ac microsetae usually longer, composed of up to 5 microsetae.Ctenidial spine on f1 strong, generally longer than in male, much longer than maximum width of t1; f3 without posteroventral row of erect setae.Wing measurements: length 2.06-3.20 mm, maximum width 0.65-1.15mm; Cs3 : Cs4 = 1.20-1.45;r-m\dm-cu : dm-cu = 2.17-3.00.
Abdomen somewhat broader and preabdominal terga more transverse and shorter than in male; T2-T5 similarly coloured in male.T2 distinctly shorter than T3 but less so than that of male.T3-T4 subequal in length, T5 usually slightly longer.T1-T2 microsculptured as in male but T3-T6 with fine transverse microsculpture reduced, almost glabrous.T3-T4 less densely though similarly microtomentose; T5 without microtomentum, bare.S1 reduced to form transverse bare strip-like sclerite having 3 darker pigmented parts (laterally and medially).S2-S5 light brown, each usually with paler anterior part, S2-S4 of subequal width, S2 sometimes and S5 regularly somewhat wider than S3-S4.S2 slightly wider than long, S3-S4 slightly longer than broad or as long as broad, S5 widest and slightly transverse but narrower (and paler) than S6.Setosity of preabdominal terga and sterna similar to that of male.
Postabdomen (Figs 23-25).T6 dark brown except for narrow anterior and posterior marginal stripe, comparatively long and strongly tapered posteriorly, relatively densely but finely setose (Fig. 23).S6 trapezoidal to suboblong with rounded corners, wider posteriorly, slightly lighter brown and with setae finer than those on T6.T7 and S7 fused to form tergosternum T7+S7 but with narrow (ventrolateral) incisions indicating the boundary between orginal T7 and S7 (see Figs 24, 25).T7+S7 dorsomedially seemingly divided (with unpigmented membranous part wider anteriorly and narrowed posteriorly -see Fig. 23), laterally longest, darkest and with longest setae, ventrally (original S7) finely setose only in posterior paler-pigmented half (Fig. 24).7th spiracle embedded in tergosternum T7+S7 near anterior margin ventrolaterally (Fig. 25).T8 (Fig. 23) brown, slightly transverse, with strongly rounded anterior corners and posterior margin narrowly pale-pigmented, with fine sparse setae in posterior corners.S8 (Fig. 24) short, with poorly defined anterior margin, posteromedially more or less bulging and posterodorsomedially incised (visible in caudal view -Fig.22), finely setose and entirely micropubescent (see also Figs 24,28).Internal structures of genital chamber (Figs 27, 28) distinct, formed by complex of 3 pairs (dorsal, lateral and ventrocaudal) of relatively short and partly fused sclerites and by 1 anterior, slender, relatively long and broad, dorsally bent annular sclerite.Ventral receptacle forming (Fig. 30) membranous, hyaline (and poorly visible), short, anteriorly strongly bent tube.Accessory gland inconspicuous (Fig. 30), vesiculate with some grains in surface, set on subterminally slightly dilated and ringed duct.Spermathecae (1+1) elongately irregularly pyriform to spindle-shaped, thus with both ends tapered; distal end of spermatheca more or less strongly curved, rugged and provided with several blunt processes, particularly on apex; proximal end subcylindrical, with a crown of blunt erect spine-like tubercles (Figs 26,30); duct without sclerotized cervix on its insertion in spermatheca.T10 (Fig. 23) short and transverse, bent and dark-pigmented, with 2 very long posteromedial setae and reduced micropubescence concentrated around bases of the latter.S10 much larger than T10, rounded pentagonal (Figs 24,29), with anterior part (about as long as posterior part) bare, posteromedially somewhat bulging; posterior half of S10 densely micropubescent and with relatively long but fine setae at posterior margin.Cerci short, narrow in dorsal view but broad in lateral view, with rich but rather short setae (the longest apical with lateral setae subequal in length) and very short micropubescence .

Preimaginal stages
Egg rather slender, elongate (Figs 35-37, 41-43), 0.71-0.83mm long, 0.16-0.19mm wide (maximum width), with both ends tapered, with dorsal side (dorsal side determined by position of freshly laid egg on substrate, anterior end by position of neonate larva before hatching) more convex than ventral side in lateral view.Colour whitish but chorion hyaline, with relatively simple sculpture, formed by 4 (2 lateral and 2 laterodorsal) pronounced longitudinal ribs of finely granulose structure and by very fine and tenuous transverse reticulation between them, forming 3 rows of meshes ventrally (Fig. 37), 2 dorsally (Figs 35, 42) and 1 laterally (Figs 36,43).Ventral egg area (Fig. 37) with 3 rows of strongly transverse meshes (1:3.5 to 1:5.5) is margined by finely granulose but hardly elevated line; between this line and lateral rib there is an additional narrow row of subquadrate to square-shaped meshes (1:1 to 1:1.5).Laterodorsal and/or lateral ribs can relatively often be malformed (interrupted, connected or split), similar to that in Q. clintonia on Fig. 44.Micropylar area lies on ventral side of anterior (more acute) end of egg with poorly differentiated micropyle situated in a subterminal depression.Note: The previous descriptions of eggs in Anthomyzidae (for references see above in the Discussion of pre-imaginal stages) incorrectly recognized dorsal side of egg as that in which micropyle is situated, i.e. the less convex side of the egg.
Cephalic segment simplified compared to those of 2nd and 3rd instars and only indistinctly anteroventromedially furrowed so cephalic lobes less distinct; facial mask with a few (3 complete) denticulate oral ridges (Fig. 50)."Antenna" (= sensory organ) on dorsal side of cephalic lobe two-segmented as in later instars but basal segment reduced to a short ring.Oral atrium developed but the unpaired mouthhook (see below) is not projecting through it.Posterior part of cephalic segment ventrally ornamented by only some 5-6 transverse rows of minute spinules.Prothoracic segment simple, lacking anterior spiracles.Cephalopharyngeal skeleton situated inside the cephalic and all thoracic segments.
Last abdominal segment (Figs 46, 51) with dorsolateral bulges in front of posterior spiracles and protruding anal area similar to those of later instars.Posterior spiracles (Figs 51, 52) formed as a button-like projection on protruding stump-like process, lacking distinct peritreme, perforated by only 2 oval spiracular slits in subradial position and bearing 4 finely palmately branched (less than in later instars) interspiracular processes.
2nd-instar larva.Body whitish hyaline to dirty white, of the same shape as in 3rd-instar larva but smaller and more slender (Fig. 53).Measurements: length 1.57-2.78mm, maximum width 0.22-0.38mm.Two thoracic and all abdominal segments with creeping welts as in 3rd-instar larva but arranged in only 3-6 rows.Both anterior and posterior spiracles developed.
Cephalic segment similarly formed as in 3rd-instar larva, including "antenna" (Fig. 56), cephalic lobes and oral atrium but facial mask with fewer oral ridges and transverse rows of minute spinules (only 5-7) on posteroventral part of cephalic segment.Prothoracic segment laterally with retractable anterior spiracles being simply clavate, with single spiracular slit on tip (Fig. 58).
Cephalopharyngeal skeleton (Figs 54-55) generally similar to that of 3rd-instar larva but differing (except for size) as follows: paired mouthhooks with distal hook less pointed but basally bearing distinct, slender and acute tooth (see Fig. 55) and less projecting ventrally in the middle and posteriorly.Dental sclerites more slender.Intermediate sclerite (Fig. 54) as in 3rd instar but its ventral connecting bridge with posterior part indistinctly separate and pale-pigmented as is anterior part.Epistomal plate hyaline and its structure poorly visible.Pharyngeal sclerite (Fig. 54) with more sinuate parastomal bars, otherwise closely resembling that of 3rd instar but with slightly shorter dorsal cornua and ventral trough with fewer longitudinal ridges.
Last abdominal segment (Fig. 53) with similarly formed dorsolateral bulges in front of posterior spiracles and bulging anal area as in 3rd instar.Posterior spiracles (Fig. 57) with only 2 oval spiracular slits (as in 1st-instar larva) but with distinctly developed spiracular plate surrounded by peritreme set on protruding stump-like process (as in 3rd-instar larva); also 4 multiply branched interspiracular processes resembling more those of the latter.
Cephalic segment (Figs 64, 65) rather simple, weakly sclerotized, divided by anteroventromedial furrow into left and right cephalic lobes.Facial mask (Fig. 65) with numerous fine oral ridges composed of small cuticular thornlets.Each cephalic lobe dorsally with very minute two-segmented sensory organ, the so-called "antenna".Ventrally, between cephalic lobes, is the oral atrium (Fig. 65) with projecting apices of mouthhooks.Posterior part of cephalic segment ventrally ornamented with about 10 transverse rows of minute spinules.Prothoracic segment (Figs 64, 65) laterally with partly retractable anterior spiracles being terminally palmate and terminating regularly in 6, relatively short and thick, finger-like projections (for detailed structure see Fig. 62).Inside the cephalic and all thoracic segments there is the cephalopharyngeal skeleton (Fig. 64, cps).
Cephalopharyngeal skeleton (Figs 60-61) generally structured as in other saprophagous acalyptrate larvae.The paired mouthhooks (mh) of moderate size, laterally flattened, with simple, slightly bent distal hook, in the middle and posteriorly distinctly projecting ventrally to lateroventrally (Fig. 61).Dental sclerites (ds) below mouthhooks well developed, canine-shaped, projecting ventrally and lying between processes of mouthhooks.In addition, there is a pair of minute, closely attached sclerites (= ?accessory oral sclerites) between proximal parts of mouthhooks (see Fig. 61).Intermediate (= hypostomal) sclerite (Figs 60,61,im) situated between mouthhooks and pharyngeal sclerite and separate from both of them, H-shaped in ventral view (Fig. 61), with connecting part between lateral rods bipartite, with anterior part longer and pale, posterior part short-transverse and darkpigmented.Epistomal plate rounded subtriangular, palepigmented to hyaline, situated anteriorly to anterodorsally between anterior arms of intermediate sclerite, of rather complex structure with small rounded ?perforations (see Fig. 61).Pharyngeal sclerite (Fig. 60) forming the largest part of cephalopharyngeal skeleton.Its paired anterior projections (parastomal bars) very slender and lying closely above intermediate sclerite.Its paired dorsal (dcr) and ventral (vcr) cornua well developed, dark-pigmented only anteriorly (Fig. 60).Dorsal cornua anterodorsally connected by simple (non-perforated), rather short dorsal bridge (db); more posteriorly free and becoming attenuated and hyaline.Ventral cornua somewhat longer than dorsal cornua, connected by weakly sclerotized ventral, downward-curving trough which is provided with fine longitudinal ridges (Figs 60, 61) serving as filter for concentration of larval food.No window in either dorsal or ventral cornua but each of the latter provided with indistinct, small and unpigmented dorsal apodeme (da).
Last abdominal segment of larva (Figs 66, 67) characterised by dorsolateral bulges in front of posterior spiracles and by lateroventrally protruding anal area embedding slot-like anus in the middle and being surrounded by fine rows of spinules both anteriorly and posteriorly (Fig. 67).Paired posterior spiracles (Fig. 66, psp) formed by sclerotized plate on shortly protruding stump-like process (Fig. 63).Spiracular plate surrounded by more sclerotized rim (peritreme) and perforated by 3 oval or ovoid spiracular slits (ss) in irregularly radial pattern and provided with 4 finely palmately (multiply) branched interspiracular processes (isp) arising among spiracular slits.
Puparium (Figs 68-70) elongately barrel-shaped, only slightly wider than high, with more tapered anterior end, less tapered and rather rounded posteriorly.Measurements: length 2.42-3.26mm, maximum width 0.58-0.89mm, maximum height 0.55-0.83mm.Integument of (empty) puparium ochreous yellow (in the middle, lightest ventrally) to brown (anteriorly and posteriorly), intensively finely transversely wrinkled and ribbed in most of abdominal segments while roughly, densely and rather irregularly ribbed anteriorly (cephalic and thoracic segments) and posteriorly (caudal abdominal segment).Larval features mostly visible on puparium but modified by sclerotization process of puparium, usually somewhat reduced.Segmentation of body generally visible.Abdominal segments (except for last one) with shallow but distinct dorsolateral and ventrolateral crescent-shaped impressions .
Anterior end of puparium strongly tapered and distinctly dorsoventrally compressed and equal to thoracic (plus integrated cephalic) segments of larva.First visible segment (= prothoracic + cephalic) dorsolaterally with sclerotized (larval) anterior spiracles each forming a short, palmately branched projection with 6 short papillae; segment proper characteristically sculptured, anteromedially hardly or very shallowly emarginate, dorsally (Fig. 99) with a group of coarse twisted ridges medially and wrinkled margins (with rest relatively smooth), ventrally (Fig. 100) with distinctive blackish, palmately branched ornamentation having its (darkest) base relatively narrow and sharply demarcated.Also 2nd and 3rd (meso-and metathoracic) segments dorsally with distinctly coarser sculpture than on following (i.e.abdominal) segments, the former with dense twisted and partly net-like connected ridges, the latter with ridges more transverse and anteriorly with several rows of very fine spinules.Cephalopharyngeal skeleton of 3rd-instar larva situated inside of anterior part of puparium, affixed to its ventral wall.
Posterior end of puparium less narrowed, convex and densely ribbed 101), ventrally with characteristic sculpture surrounding larval anus (Fig. 101) including rows of very fine spinules, posteriorly behind anus relatively smooth but strongly sloping, with pair of elongate tubercle-like swellings, and dorsolaterally with still visible bulges (though reduced against those of larva).Posterior spiracles on short stump-like processes (Fig. 101) being set on relatively broadened subconical protrusions of posterior abdominal segment.Larval adornment (transverse bands = creeping welts of fine spinules) of cuticle well preserved both on dorsal and ventral sides of abdominal segments.
Discussion.Quametopia terminalis is the most distinct of the Quametopia species.Apart from structures in the male and female genitalia, it can be confidently recognized in the female sex by the darkened 1st antennal flagellomere (see Figs 9,12).Differences in other external features are small and often overlapping (for details see descriptions of Q. clintonia sp.n. and Q. amplistylus sp.n.) and, therefore, it is suggested to identify the males of Quametopia species using characters of gonostylus and internal genitalia (aedeagal complex) as listed in the above key.Also the pre-imaginal stages of Q. terminalis differ from those of Q. clintonia sp.n. in lacking the pronounced ledge defining the ventral perimeter of the egg, in the formation of the cephalopharyngeal skeleton of the 1st-instar larva, anterior spiracles and cephalopharyngeal skeleton of both the 2nd-and 3rd-instar larvae, and in the surface structures of the anterior and posterior segments of the puparium (see key).Moreover, Q. terminalis also differs from Q. clintonia sp.n. in its biology, being strictly associated with I. capensis (see below) in undergrowths of moist forests and their edges or openings.
Mumetopia nitens Melander, 1913 was synonymized with Q. [Mumetopia at that time] terminalis by Sabrosky (1965).Melander (1913) did not recognize that some parts of the head of Q. terminalis are sexually dichroic and described its female with darkened antennae, face and mouthparts as a different species, M. nitens.It is, however, peculiar that he was mistaken in this case because, in fact (see other material examined) there are both males and females of Q. terminalis in his material collected in the same locality and on the same date as the (female) type specimens.
Biology.Adults of Q. terminalis are commonly found in stands of I. capensis in Ontario (Canada) where they can be seen sitting on leaves, and hovering amongst stems or above the plants when disturbed.Most collections by the authors were made in shaded or semi-shaded habitats (Fig. 73), usually on moist soil (at least in the spring), where I. capensis occurred or occasionally where it may have been overlooked as a minor component.Where I. capensis stands are strong and dominant, this fly can be very abundant and its choice of host plant quite evident.However, no observation of oviposition was ever made in the field so finer details of life history remain unknown (but see note below of egg on damaged stem).
Adults of Q. terminalis have been caught as early as 16 May (Plummers Island, MD) and as late as 18 September (Sault Ste.Marie, ON; 2 October, Middlesex Co., MA but Malaise trap start date not recorded) which suggests two generations.The majority of specimens have been taken with sweep nets with two records from Malaise traps.
Field collections of litter and soil, and stems of I. capensis.Of the five rectangular samples from 1999 (~1170 cm 2 of substrate), three yielded a total of five Q. terminalis adults.Two males emerged 17 days and three females emerged 18, 19, and 23 days after collection at 20°C.Of the 14 cylindrical cores from 2002 (~370 cm 2 of substrate), four yielded a total of five adults of Q. terminalis.The one male and four females emerged between 13 and 16 days later (all found the same day after a 3-day interruption of observation, all but one dead) at 25°C.These preliminary observations indicated that Q. terminalis was resident in these sites and overwinters in the litter and/or soil below I. capensis.
Thirty-six of the 38 immature specimens of Q. terminalis and T. trisulcata dissected from the stems were found in the lower 3 cm of the plant and mostly in the hollow lateral roots (Figs 7,72).From Table 1, it can be seen that 26 flies emerged from the dissected stems and thus represent immatures overlooked during the initial dissection.A second search of a portion of this material (one box with 19 of the 26 adults) uncovered 11 empty puparia and four additional desiccated larvae.These 11 puparia were comprised of three Q.terminalis (one in a lateral root, and two loose) and eight T. trisulcata (one in the main root, one in a lateral root, five in a stem internode, and one loose).
The dissected stems of I. capensis also yielded small numbers of Cecidomyiidae, Sciaridae, and a community that appears to be based on the stem-boring Pristerognatha agilana (Clemens, 1860) (Tortricidae) (MacKay, 1959;Gilligan et al., 2008).A total of 82 adult P. agilana emerged and more larvae and pupae were encountered in the dissections.Parasites associated with the caterpillars or pupae included Braconidae, Ichneumonidae, and Chalcidoidea.A single platygastrid also emerged but its host is not known.
A single, serendipitous observation of a field-collected egg was made when preparing for rearings.A section of I. capensis stem collected 29 July 2010 was removed from the freezer in early September and a flattened (empty or damaged) egg was found attached to the outer wall of what appeared to be insect feeding damage (Fig. 41).This shallow injury did not penetrate to the hollow interior of the stem but fluid from the freeze-damaged tissues readily leaked into the depression indicating a possible entry point for a hatching larva -no such larva was found in the section of stem.
The association of Q. terminalis with its putative host plant is based on several observations: repeated collections of adults from pure and mixed stands; laboratory rearing from adult to adult upon presentation of the stems of I. capensis; and rearing of adults from litter/soil and stem samples.Dissection of maggots from overwintered stems, and particularly roots, coupled with a single observation of an egg on a damaged stem, provides a finer degree of detail and certainty.
Collectively, these data indicate that at least some Q. terminalis larvae overwinter within the roots (and pos-sibly stems) of I. capensis, primarily or exclusively as third instars.The relationship with T. trisulcata is not known but their combined life history may include the use of younger plants earlier in the season.Self-thinning is known to occur in dense stands of the annual I. capensis (see Schmitt et al., 1987) which would provide a ready source of decomposing plant tissue for ovipositing flies.Alternatively or additionally, mechanical damage caused by vertebrates, wind, or the larval tortricid moths, in particular, could provide access to the plant tissues or hollow stem interior -the frass of the latter also could be an important resource for the maggots.
Rearing of Q. terminalis.Egg production in the flight cages was not enumerated but the last 4% and 6& transferred to the cup cage yielded ~200 eggs from 18 August until 13 September 2002 whereupon the 5 surviving adults were killed, 112 days after their collection from the field.This suggests a capacity for a protracted flight period on the level of the individual flies.
The first puparium was observed on 4 August 2002, 41 days after initial setup of adults and about one month after the first observation of eggs.Puparia that were produced from 5 August to 31 October in 2002 yielded 18 adults (7%, 11&), 12 days (%, n = 4) or approximately 11 days (&, range = 9-12, n = 10) after pupariation when held at 20°C.These adults were generally undersized measuring ~84% (males) of the body length of wildcaught adults from Bristol Place Park.Of the puparia resulting from overwintered larvae, 29 adults (19%, 10&) were produced from 12-25 April and these required about 14 days (range = 13-15 for both sexes) at 20°C to eclose.These adults were also undersized measuring ~90% of wild-caught adults (males n = 7).
The smaller rearing from 2009 provided some additional information.Eggs required 6 days to hatch at 20°C.Of the 35 eggs laid between 24 and 27 August, 10 puparia were produced between 14 November and 24 January (other larvae died, were lost, or were preserved).There could have been some delays in development as a result of the late season despite the laboratory holding conditions.Since all adults (7%, 3&) emerged 15-16 days later, there was no obligatory puparial diapause which supports the stem-dissection results that suggest this species overwinters as mature larvae.The 10 adults produced in 2009 (at least 89-161 days after egg hatch) were not undersized but slightly larger than the wildcaught adults used to compare with the 2002-2003 rearings and this length of time suggests no more than two generations would be possible.Immediate and exclusive use of split stems of I. capensis in a moisture-controlled (no wet sand) Petri plate is thus recommended for this species (Fig. 71).
Oviposition choices.Table 2 summarizes the results of the oviposition choice trial.Females of Q. terminalis showed a strong preference for I. capensis by laying 43 eggs on the stems compared to a single egg on the petiole of C. borealis.
Distribution.The distribution of this species (as Mumetopia terminalis) was summarized by Sabrosky (1965): New Hampshire, Michigan to Quebec, south to Virginia.Actually, there are very few original records of this species: Loew (1863) described it from "Carolina" [this was for unknown reasons changed to "New Hampshire" by Melander (1913) and this correction, as an error made by Loew, was also accepted in Sabrosky (1965)], Melander (1913) described it as M. nitens from Massachusetts, Johnson (1925) recorded it from Maine, New Hampshire and Massachusetts (but at least his record from Maine: Mt.Desert is based on misidentification of Q. clintonia sp.n., see below), Procter (1946) probably only repeated the misidentified record from Mt. Desert, Winn & Beaulieu (1932) reported it from Quebec and Marshall et al. (2001) from Ontario.Other parts of the distribution range presented by Sabrosky (1965) were apparently based on unpublished records known to him.Although most of the above records obviously refer to Q. terminalis, some may be partly based on misidentifications of the other two species described below.The general distribution of the species is detailed here on the basis of material examined: Canada (AB, ON, QC) and USA (CT, DC, IL, IN, MA, MD, ME, MI, NC, NH, NJ, NY, PA, VA, WV).This is the most commonly collected species of Quametopia and it is glaringly absent from the maritime provinces of Canada.The single specimen from Alberta is quite baffling as is the absence of records through Manitoba and Saskatchewan.The host plant, I. capensis, is very broadly distributed in North America with notable absence only in Montana, Wyoming, the southwest USA, California and the far north of Canada and Alaska (United States Department of Agriculture, 2010) (present in Labrador, S.J. Meades, pers.comm.).This would seem to be a case of insufficient collection effort directed at I. capensis as Q. terminalis is very easily collected if it is present (in contrast to Q. clintonia sp.n. that can be much more difficult to sweep out of C. borealis).

Description
Male.Total body length 1.94-2.58mm; body shape and colouration closely resembling that of Q. terminalis, thus blackish brown, relatively shiny, with legs and fore head largely yellow.Head similarly formed and coloured as in Q. terminalis.Frons with anterior yellow area (being somewhat darker ochreous posteriorly) usually reaching up to level of posterior ors but only more medially, in dull narrow areas between frontal triangle and orbits.Frontal triangle large, slightly longer than wide, not reaching to level of anterior ors (thus slightly shorter than in Q. terminalis), brown, sparsely greyish brown microtomentose only in anterior corner and on ocellar triangle, otherwise glabrous and shiny; narrow areas between frontal triangle and orbits dull and grey microtomentose as in Q. terminalis.Orbit narrowly bare, shining and brown posteriorly (up to anterior ors), yellow and microtomentose in front of the latter.Occiput as in Q. terminalis but less densely microtomentose and more shiny (besides bare medial spot above foramen) also laterally.Face pale to dark yellow and dull as in Q. terminalis.Parafacialia and gena light yellow and silvery white microtomentose; parafacialia (internally) with narrower and paler, gena (ventrally) with wider and darker brown marginal stripe.Mouthparts whitish yellow including palpus and proboscis but clypeus (small and often hidden) brownish.Cephalic chaetotaxy as in Q. terminalis but pvt sometimes crossed, anterior ors usually about half length of posterior and postocular setulae dorsally (behind vte) longer.Eye with longest (oblique) diameter about 1.5 times as long as shortest; smallest genal height 0.10 times as long as shortest eye diameter.Palpus entirely whitish yellow, with same chaetotaxy as that of Q. terminalis.Antenna as in Q. terminalis but 1st flagellomere with whitish cilia on anteroventral side longer than those of arista.
Thorax closely resembling that of Q. terminalis in shape, colour, microtomentose pattern and even chaetotaxy.Legs as in Q. terminalis, yellow, only apical (terminal) segment of all tarsi dark brown, more rarely also preapical segment of fore tarsus brown.f1 with ctenidial spine longer than maximum width of t1; long setae in both posterodorsal and posteroventral rows fine but longest posteroventral seta (near ctenidial spine) often markedly thicker than others.t2 with ventroapical seta yet shorter (usually only as long as maximum width of t2) and slightly curved.f3 in distal two-thirds with denser posteroventral row of 12-15 erect and more distinctly thickened setae.Wing (Fig. 32) very similar to that of Q. terminalis including colour of membrane and veins.R4+5 long, usually slightly bent; M subparallel to R4+5, very  slightly bent or indistinctly sinuate, apically slightly divergent from R4+5.Cross-vein r-m situated slightly to distinctly in front of discal (dm) cell; dm cell elongate and formed as in Q. terminalis; cross-vein dm-cu usually distinctly shorter than apical portion of CuA1; all other veins and cells as in Q. terminalis.Wing measurements: length 2.22-2.70mm, maximum width 0.71-0.89mm; Cs3 : Cs4 = 1.29-1.55;r-m\dm-cu : dm-cu = 2.38-3.29.Haltere of the same colour and form as that of Q. terminalis.
Abdomen.Preabdominal terga T2-T5 as in Q. terminalis, completely dark brown, T1 paler brown than remaining terga, at least anterolaterally.T1-T5 with very fine transverse microsculpture and similarly greyish microtomentose as those of Q. terminalis but this microtomentum usually denser and covering larger medial area on T3 and T4 in both reaching or almost reaching to posterior margin (hardly so in T4 of Q. terminalis); also T5 with wider microtomentose stripe along anterior margin; consequently, bare areas of terga smaller and hence abdomen generally less shiny than in Q. terminalis.Preabdominal sterna becoming very slightly wider (S5 widest) and darker posteriorly; S1 as in Q. terminalis; S2 slightly wider than long; S3 and S4 subquadrate, usually as long as broad or S4 slightly wider than long; S5 largest, distinctly wider than long (thus transverse), with anterior corners more or less rounded.T6 similarly reduced and with bipartite pigmentation as in Q. terminalis.S6-S8 resembling those of Q. terminalis in shape, pigmentation and microtomentose pattern.S6 with 0-1 (usually 0) and S7 without setae; S8 large, with numerous setae.
Genitalia.Epandrium (Figs 74,75) similar to that of Q. terminalis but with setae more sparse and anal fissure narrower and more rounded dorsally.Cerci as in Q. terminalis, shortly setose and with apices attached medially.Medandrium (Fig. 74) with dorsolateral corners less projecting and ventrolateral arms less divergent; the latter with 3 setulae, 1 usually distant from 2 others.Gonostylus (Figs 78,(129)(130)(131)(132)(133) bilobed, with longer anterior clavate lobe similarly setose to that of Q. terminalis but with micropubescence reaching in distal part of its outer side; posterior lobe markedly shorter than that of Q. terminalis, always shorter than half length of anterior lobe, and regularly with some microsetulae at dorsal margin; ventral incision between both lobes shallow (Fig. 78).Internal genitalia very similar to those of Q. terminalis but differing as follows.Hypandrium (Fig. 77) slightly more robust and with internal membranous hypandrial lobes more projecting dorsally.Transandrium with somewhat thicker dorsal ledge; basal membrane similarly armed with fine spinules (Fig. 76) but lateral finely granulose area narrower (Fig. 77).Pregonite (Fig. 77) yet more reduced than in Q. terminalis, with anterior lobe almost indistinct in lateral view; 3 (usually) to 4 setae present on internal side of pregonite.Postgonite (Fig. 77) usually straighter, regularly with 1 fine proximal seta as in Q. terminalis, rarely with a similar seta in distal third of anterior margin.Aedeagal complex (Fig. 79).Phallapodeme similar to that of Q. terminalis but its apical part with lateroventral corners reduced.Aedeagal part of folding apparatus similarly sclerotized and armed but dark tubercles often covering larger area ventrally; connecting sclerite with thicker spinules distally.Phallophore with somewhat larger (expanded) anterior part.Distiphallus with flat posteroventral lobe setulose mainly in distal third (Fig. 79).Middle part of distiphallus as in Q. terminalis.Saccus of distiphallus somewhat larger and with more spinules, particularly on right side (Fig. 79).Filum similarly formed to that in Q. terminalis but its membranous apex with only very fine denticles at tip (Fig. 80).Ejacapodeme somewhat larger than that of Q. terminalis, with digitiform projection longer than body of ejacapodeme (Fig. 79).
Female (Fig. 104).Similar to male unless mentioned otherwise.Total body length 1.98-2.88mm.Head colouring different from that of the male but less distinctly than in Q. terminalis.Frons with anterior yellow area similar to that of male.Face ochreous to pale brown, greyish microtomentose and dull; bordering stripe pale brown on parafacialia, darker brown on gena.Mouthparts with blackish brown clypeus; palpus with brownish outer side (darkest basally) and ochreous yellow inner side and apex; proboscis ochreous to pale brown, only labellae dirty whitish yellow.Antenna not darkened, entirely yellow or orange-yellow (Fig. 104) as in male.Lateral rows of ac microsetae usually longer, also prs seta longer than in male.Ctenidial spine on f1 distinctly longer than maximum width of t1; t2 with ventroapical seta longer than maximum width of tibia but usually slightly bent (as in male); f3 without posteroventral row of erect setae.Wing measurements: length 2.26-2.96mm, maximum width 0.75-1.03mm; Cs3 : Cs4 = 1.19-1.65;r-m\dm-cu : dm-cu = 2.61-3.66.
Abdomen somewhat broader and preabdominal terga more transverse and shorter than in male (as in Q. terminalis); T1-T6 with fine transverse microsculpture reduced, less distinct than in male.Microtomentose pattern of T1-T4 similar to that of male, T5 and T6 almost without microtomentum.S1 reduced to transverse bare strip-like sclerite, similar to that of Q. terminalis but with less distinct tripartite pigmentation pattern.S3-S4 of subequal width, S2 and S5 usually slightly wider than S3-S4.S2 somewhat wider than long, S3-S4 as long as broad (or S4 slightly wider than long), S5 somewhat wider than S4 and slightly transverse, usually narrower than S6.

Preimaginal stages
Egg similar to that of Q. terminalis, 0.75-0.84mm long, 0.20-0.22mm wide (maximum width), thus slightly thicker on the average .Both ends tapered but usually less acutely than in Q. terminalis.Colour whitish, chorion hyaline with very similar sculpture as in latter species, thus formed by 4 (2 lateral and 2 laterodorsal) pronounced longitudinal ribs and by very fine and tenuous transverse reticulation between them, forming 3 rows of meshes ventrally (Fig. 40), 2 dorsally (Figs 38,45) and 1 laterally (Fig. 39).However, in contrast to Q. terminalis, ventral reticulated area with 3 rows of transverse meshes is margined by a distinctly elevated ledge (visible in Fig. 40) almost forming a low additional rib, being separated from lateral rib by a narrow row of subquadrate (almost square-shaped) meshes.In contrast, the row of meshes between lateral and laterodorsal rib is distinctly wider and meshes are transverse (1:2 to 1:3).Sometimes eggs with abnormal formation of ribs (Fig. 44) occur among normal eggs.The anomalies (split or interrupted ribs) can affect both lateral and laterodorsal ribs and can be relatively frequent.Micropylar area poorly visible, forming ventral subterminal depression on anterior (more acute) end of egg.
Larva.1st-instar larva.Body shape, colour and structures closely resembling those of Q. terminalis but differing as follows.Measurements: length 0.87-1.59mm, maximum width 0.14-0.22mm.Cephalic segment (Fig. 89) with only 3 complete oral ridges as in Q. terminalis but ventral transverse rows of minute spinules at posterior margin of cephalic segment more reduced (3-4, some incomplete).Posterior end (last abdominal segment) hardly different from that of Q. terminalis including shape, structure and armature of posterior spiracles.Cephalopharyngeal skeleton (Figs 87,88) also similar to that of the latter species but dorsal cornua more slender and shorter than ventral cornua, unpaired mouthhook shorter and less acute, parastomal bar thicker (particularly anteriorly) and intermediate sclerite with ventral plate anteriorly acute-angled (subtriangular).
2nd-instar larva.Similar to that of Q. terminalis, only differing in some detail of the anterior spiracle and cephalopharyngeal skeleton.Measurements: length 1.60-2.94mm, maximum width 0.22-0.42mm.Anterior spiracle (Fig. 91) simply clavate as in Q. terminalis but somewhat narrower, with ellipsoid slit.Cephalopharyngeal skeleton (Fig. 90).Mouthhooks differing from those of Q. terminalis in having more robust distal hook, reduced but sharply pointed denticle on its base (Fig. 92), and longer and more projecting middle and posterior processes.
Dental sclerite longer and more slender (Fig. 92).Intermediate sclerite (Fig. 90) similarly formed to that of 3rd instar, hence differing from that of Q. terminalis in having anterior part of ventral connecting bridge darkpigmented.Dorsal cornua distinctly shorter than ventral cornua and its anterior part (below dorsal bridge) with somewhat darker striped pigmentation.Ventral cornua with larger (although unpigmented) dorsal apodeme, thus more similar to that of 3rd-instar larva.
3rd-instar larva.Very similar to that of Q. terminalis including shape, colouring and armature but differing as follows.Measurements: length 3.17-4.92mm, maximum width 0.49-0.76mm, when extended.Anterior spiracles terminally palmate as in Q. terminalis but usually with only 5 (less frequently with 6) finger-like projections (see Fig. 95).Posterior spiracles hardly different from those of Q. terminalis.Cephalopharyngeal skeleton (Figs 93,94) generally similar to that of Q. terminalis but mouthhooks with middle projection longer and projecting more laterally (cf.Fig. 94), dental sclerite longer and more robust, intermediate sclerite (Figs 93,94) thicker and with connecting ventral bridge with anterior part dark-pigmented and posterior short-transverse protruded part anteriorly more acute and darkened only medially.Epistomal plate weakly sclerotized and pale-pigmented, of about rhomboid shape and of different (though poorly defined) structure (see Fig. 94).Pharyngeal sclerite (Fig. 93) differing from that of Q. terminalis in pigmentation (having darker stripes in anterior part), dorsal cornua distinctly shorter than ventral cornua and the latter provided with larger unpigmented dorsal apodeme.
Puparium (Figs 102, 103, with mature adult inside) of similar shape to that of Q. terminalis (slightly wider on the average) differing only in some detail in structures of anterior and posterior ends (not to mention larval cephalopharyngeal skeleton preserved inside).Measurements: length 2.86-3.26mm, maximum width 0.77-0.91mm, maximum height 0.66-0.83mm.Anterior end of puparium with somewhat deeper incisions between segments.First segment (= prothoracic + cephalic) dorsolaterally with anterior palmately branched spiracular processes, each with 5-6 short papillae; segment proper anteromedially usually with distinct (often deep) emargination, dorsally (Fig. 96) with somewhat finer sculpture in the middle, ventrally (Fig. 97) with dark palmately branched ornamentation having its blackish base broader.2nd and 3rd (meso-and metathoracic) segments dorsally (in large central area) with coarse ribbed structure usually less dense than in Q. terminalis, ventrally, on the contrary, more dense than in the latter species.Posterior end of puparium (Fig. 98) ventrally with fewer ribs on precaudal segment and area posterior to anus differently ribbed, with several rib-shaped tubercles.Posterior spiracles on short stump-like processes (Figs 98,(102)(103) without broadened subconical bases.Larval adornment (creeping welts) on ventral side of caudal abdominal segment with somewhat larger spinules.larvae, 6 puparia + 7 empty puparia: all lab-reared on Clintonia borealis, using adults from CANADA: Ontario: nr.Searchmont, Whitman Dam & Ranger Lake Roads; Batchawana Provincial Park; Lake Superior Provincial Park, Crescent Lake (Ba, DEBU).
Discussion.Quametopia clintonia sp.n. is most closely allied to Q. amplistylus sp.n. as demonstrated by the similarly abbreviated posterior lobe of gonostylus, reduced anterior lobe of pregonite, finely denticulate apex of filum of distiphallus and very similar internal sclerotization of the female genital chamber (reduced lateral sclerites, prolonged dorsal sclerite).Moreover, females of both species have yellow antennae, concolourous with those of males, in contrast to sexually dichroic antennae in Q. terminalis.While males of Q. clintonia sp.n. can be easily distinguished from those of Q. amplistylus sp.n. by the shape of the anterior lobe of gonostylus and some additional details in the aedeagal complex (see the key above), the females of both species are so similar that sometimes it is difficult to separate them confidently (particularly when internal sclerites in the female genital chamber are in unusual, i.e. other than rest, position).
Based on the material examined, it was found that sometimes both Q. clintonia sp.n. and Q. amplistylus sp.n. can occur syntopically.Quametopia clintonia sp.n. proved to be strictly associated with C. borealis in Ontario (for detail see below) but the host plant(s) of Q. amplistylus sp.n. remain(s) unknown.As mentioned above, Q. clintonia sp.n. differs from Q. terminalis in some structures of all pre-imaginal stages (see the key and in the above description) as well as in the host-plant and habitat association.This can supposedly be also true for Q. amplistylus sp.n. as it is more closely related to Q. clintonia sp.n. than Q. terminalis.
Biology.Compared to Q. terminalis, it was more difficult to typify and predict suitable habitat for Q. clintonia sp.n. as adults were never found in numbers sufficient to localize microsites in 2009.Once collections in Greenwater Provincial Park (2009) suggested the presence of this second species in Ontario, dominant understory plant species were recorded in subsequent specimen label data.Subsequently, C. borealis (Fig. 105) surfaced as the only species that was consistently present at capture sites of this fly.Some sites included virtual monoculture stands of the clonal C. borealis but, paradoxically, these proved not to be the most productive sources of adults when compared to the heterogeneous sites at Greenwater Provincial Park and Dubreuilville.While most adults were swept from mixed vegetation under canopy, single records implicating rotting mushrooms and a bleeding maple stump were located in forest habitat harbouring C. borealis.Records mentioning sedges and ferns have likely overlooked the C. borealis component.However, the large collection from Lake Superior Provincial Park (Crescent Lake, 2010) represents the strongest field evidence for this host association where the dominance of C. borealis was clear.Individual rotting leaves of this plant were frequently observed at this site (Fig. 105) and are presumed to be the oviposition and developmental sites.
Adults of Q. clintonia sp.n. (Fig. 104) have been collected as early as 13 June (Lake Superior Provincial Park, ON) and as late as 20 September (Batchawana Provincial Park, ON).Compared to Q. terminalis, this species is under-collected but might also manage two generations per year.Adults have been taken on sticky traps in mixed forest, by flight interception, pan, and pitfall traps, but mostly by sweeping in various forested areas with a range of ground cover including C. borealis (where records were made).
Rearing of Q. clintonia sp.n.Adults in the cup cages were presented with choices of plant parts (see next section below and Table 2) or, later, with only petioles of C. borealis.Most eggs hatched 6 days after oviposition but some as late as 13 days.Most larvae died during early stages within the petioles but surviving larvae transferred to leaf material (Fig. 4) survived and developed quite well.The first two puparia were produced on 30 October and 1 November but then died.Of the 13 puparia formed, 7 produced adults (3%, 4&) from 8 November to 1 December, a range of at least 65-88 days after egg hatch.Pupariation times ranged 15-19 (17.4 ± 1.72 s.d., n = 7) days at 20°C.The resultant adults were not measured but were not noticeably undersized.The last larva was quite lethargic and was killed 21 December, more than 100 days after hatching.It is recommended that leaf material be used as a larval substrate for rearing this species.
Oviposition choices (Table 2).The two females of Q. clintonia sp.n. in one cage laid 39 eggs (excludes nine eggs laid on the Petri plate or the sand) exclusively on C. borealis surfaces, 25 on the petiole and 14 on the leaf.The single female of Q. clintonia sp.n. was less discriminating with 11 eggs laid on the petiole and 9 on the leaf of C. borealis but also 2 eggs laid on the split stem of I. capensis.This is strong behavioural evidence of an ecological separation of Q. clintonia sp.n. and Q. terminalis which is congruent with field observations of habitat selection.
Distribution.Of the former records of "Mumetopia" terminalis, only the one by Johnson (1925) from Maine (Mt.Desert) proved to be based on misidentified Q. clintonia sp.n. (see type material).Based on material examined, this species is known from Canada (NL, NS, ON, QC) and USA (IN, ME, NH, NY, VT).This is a slightly more northern distribution than that of Q. amplistylus sp.n. with which it sometimes co-occurs (NS, NH, see below).The known distribution of the presumed host plant, C. borealis, extends broadly from southern Labrador west to Minnesota and south to Pennsylvania, then further south only at higher elevations to Georgia (Utech, 2002).With a directed sampling from the host plant, it is most likely that Q. clintonia sp.n. will be found in a larger area within the range of C. borealis especially since the current records in the USA seem to be restricted to the northern tier (as in Q. terminalis).It is tempting to speculate that Q. clintonia sp.n. and Q. amplistylus sp.n. share the same host plant as their combined distributions are suggestive of that of C. borealis and, as previously mentioned, they have been found in the same traps in Nova Scotia.Quametopia terminalis can occur within 50 m of Q. clintonia sp.n. in Ontario (Sault Ste.Marie, Pancake Bay Provincial Park) but is separated by host plant.

Description
Male.Total body length 2.22-2.75mm (2.69-3.02mm when dried from specimens preserved many years in alcohol); body shape and colouration similar to those of both relatives, thus blackish brown (brown to reddish brown in some type specimens originally preserved in alcohol), relatively shiny, with legs and fore head largely yellow.Head formed, coloured and bristled as in Q. clintonia sp.n. unless mentioned otherwise.Frons with anterior yellow area often medially reaching up to level of posterior ors and sometimes also covering anterior (microtomentose) corner of frontal triangle.Frontal triangle longer than wide, usually not reaching to level of anterior ors, coloured and microtomentose as that of Q. clintonia sp.n.Orbit with narrow shiny areas as in allies but its brown posterior part reaching slightly in front of posterior ors, thus yellow anterior part larger.Face often darker (dirty yellow to ochreous) than in relatives (not visible in faded specimens originally preserved in alcohol).Parafacialia and gena as in Q. clintonia sp.n. but lighter margined (dark yellow on parafacialia; pale brown on gena).Mouthparts yellow including palpus (lighter) and proboscis (darker) but clypeus brown.Cephalic chaetotaxy as in relatives but pvt often longer and only slightly convergent to almost parallel; anterior ors variable in length, ranging from one-third to three-fifths of length of posterior ors; postocular setulae dorsally longer as in Q. clintonia sp.n.Eye with longest (oblique) diameter 1.3-1.4times as long as shortest; smallest genal height 0.10 times as long as shortest eye diameter.Palpus pale to bright yellow.Antenna with whitish cilia on anteroventral side of 1st flagellomere not longer than those of arista.
Thorax resembling that of both relatives in shape, colour, microtomentose pattern and chaetotaxy but ac microsetae more numerous, in 4 rows reaching posteriorly to level of anterior dc.Legs as in Q. clintonia sp.n., yellow, with apical (terminal) segment of all tarsi dark brown and usually also preapical segment of fore tarsus brown.f1 with ctenidial spine longer than maximum width of t1; t2 with ventroapical seta slightly longer than maximum width of t2 and slightly curved.f3 in distal two-thirds with posteroventral row of 12-14 erect (as in Q. clintonia sp.n.) setae but they are longer and finer (resembling more those of Q. terminalis).Wing (Fig. 33;Fig. 34 with an aberrant venation having surplus cross-vein in dm cell) closely resembling those of both congeners including colour of membrane and veins.R4+5 long, slightly bent as in Q. clintonia sp.n.; M subparallel to R4+5, very slightly bent, apically parallel or very slightly convergent to R4+5.Cross-vein r-m situated around middle of dm cell (ranging from slightly in front of it to slightly behind it); dm cell formed as in congeners; cross-vein dm-cu shorter than apical portion of CuA1; all other veins and cells as in Q. terminalis.Wing measurements: length 2.38-3.00mm, maximum width 0.81-1.11mm; Cs3 : Cs4 = 1.30-1.85;r-m\dm-cu : dm-cu = 2.24-3.33.Haltere as in Q. terminalis.
Abdomen.Preabdominal terga T2-T5 similar to those of both allies, dark brown, only T1 with anterolateral corners paler.T1-T5 with fine transverse microsculpture and similarly greyish microtomentose to those of Q. terminalis thus with microtomentum covering smaller area on T4 only reaching to one-third or half its length medially; T5 with narrow microtomentose stripe at anterior margin; thus, bare areas of terga larger and abdomen as shiny as that of Q. terminalis.Preabdominal sterna S2-S5 becoming slightly wider (S5 widest and usually darkest) posteriorly; S1 as in Q. terminalis; S2 slightly wider than long; S3 and S4 subquadrate as in Q. clintonia sp.n.; S5 largest, widest, distinctly transverse.Postabdomen (Fig. 106): T6 similarly reduced (but sometimes longer), bare and with bipartite pigmentation as in Q. terminalis.S6-S8 resembling those of both congeners in shape, pigmentation and microtomentose pattern.Both S6 and S7 without setae; S8 large, with numerous setae.
Genitalia.Epandrium (Figs 107,108) broader than those of Q. terminalis and Q. clintonia sp.n., but with setae more dense than in latter species and anal fissure usually narrower and more triangular than in both these relatives.Cerci resembling those of Q. terminalis, shortly setose and with apices attached medially (less distinct in Fig. 107 because of everted anus in this specimen).Medandrium (Fig. 107) more similar to that of Q. clintonia sp.n. in having shorter dorsolateral corners but with ventrolateral arms as divergent as in Q. terminalis; the latter with 2-3 setulae.Gonostylus (Figs 109,(134)(135)(136)(137) bilobed‚ differing from those of relatives in having anterior clavate lobe longer and posteriorly strongly dilated; its posterior lobe of only half width of the anterior lobe and as short as that of Q. clintonia sp.n., usually without microsetulae at dorsal margin.Internal genitalia resembling those of congeners but differing in some detail as follows.Hypandrium (Fig. 111) more similar to that of Q. clintonia sp.n., with internal membranous hypandrial lobes more projecting dorsally.Transandrium medially more narrowed, with thin dorsal ledge; basal membrane similarly armed with fine spinules (Fig. 113) but lateral finely granulose area larger than in both relatives (Fig. 111).Pregonite (Fig. 111) with anterior lobe reduced and with 3 to 4 setae as in Q. clintonia sp.n.Postgonite (Figs 110,111) also very similar to that of Q. clintonia sp.n., but slightly longer and, hence, relatively more slender, with 1 fine proximal seta.Aedeagal complex (Fig. 110).Phallapodeme generally more robust than those of both congeners, and its apical part with small acute lateroventral corners.Aedeagal part of folding apparatus with dorsal narrow blackish part usually more robust and dark tubercle-like armature more dense ventrally; connecting sclerite well developed, pigmented and distally provided with both fine and thicker spinules.Phallophore more similar to that of Q. terminalis.Distiphallus with flat posteroventral lobe somewhat more triangular (Fig. 110) in lateral view, distinctly setulose.Middle part of distiphallus as in relatives.Saccus of distiphallus yet larger than in Q. clintonia sp.n. and internally provided with a few (4-6) small pale-pigmented tubercle-like spines in addition to fine spinules on surface (Fig. 110).Filum composed of usual pair of dark ribbon-shaped sclerites but dorsal (anterior) of them dilated and basally split; both these sclerites seem to cross below membranous apex which is distinctly dilated (lanceolate to somewhat truncate, see Fig. 112) with a few flat denticles at tip.Ejacapodeme well developed, with body as long as digitiform projection being hardly dilated terminally (Fig. 110).
Female.Similar to male unless mentioned otherwise.Total body length 2.38-2.65 mm (2.81-3.26mm when dried from alcohol-preserved specimens).Head colouring only slightly different from that of the male.Face ochreous to pale ochreous brown (much faded in alcohol-preserved specimens), grey microtomentose and dull; bordering stripe on parafacialia and gena darker, pale brown to brown.Mouthparts with dark brown clypeus; palpus ochreous to pale brown (but faded to dirty whitish yellow in alcohol-preserved specimens), darkest on outer side basally, lighter on inner side and apex; proboscis ochreous to pale brown, labellae dirty yellow.Chaetotaxy as in male but pvt slightly to distinctly convergent.Antenna not darkened, entirely yellow or orange-yellow as in male.Lateral rows of ac microsetae often longer, reaching far behind level of anterior dc.Ctenidial spine on f1 distinctly longer than maximum width of t1; t2 with ventroapical seta longer than in male; f3 without posteroventral row of erect setae.Wing measurements: length 2.58-3.02mm, maximum width 0.85-1.03mm; Cs3 : Cs4 = 1.21-1.62;r-m\dm-cu : dm-cu = 2.21-3.05.
Abdomen somewhat broader and preabdominal terga more transverse and shorter than in male; fine transverse 322  Postabdomen (Figs 114,118) most similar to that of Q. clintonia sp.n.The (very slight and partly overlapping) differences are as follows.S6 wider (more transverse) and usually more densely setose.Tergosternum T7+S7 slightly more transverse, with more setae on ventral part (original S7).T8 usually slightly wider anteriorly, with broadly rounded posterior corners (Fig. 114).S8 (on Fig. 118 with posterior side oriented ventrally due to exposed genitalia) with posteromedial bulge and (more dorsal) incision, as in relatives.Internal structures of genital chamber (Figs 117,120) very similar to those of Q. clintonia sp.n., but dorsal pair of posterior sclerites shorter and anterior annular sclerite usually longer.Ventral receptacle (Fig. 120) also most similar to that of Q. clintonia sp.n. with duct basally wider (often poorly visible because of membranous, hyaline nature of this structure).Accessory gland (Fig. 116) possibly larger than in relatives, elongately vesiculate with a set of stalked globuli inside, on ringed but hardly dilated duct.Spermathecae (1+1) most resembling those of Q. clintonia sp.n. but usually more robust (with larger maximum diameter) and distal tapered end of spermatheca usually shorter, less curved or straight and provided with more projections (Fig. 119).T10 (Fig. 114) often longer (less transverse) than that of Q. clintonia sp.n.S10 differing from that of Q. clintonia sp.n. in having anterior bare part smaller (Fig. 115) and strongly abruptly separated (cf.Fig. 120) from posterior micropubescent part.Cerci short, similar to those of relatives, usually wider in dorsal view (Fig. 114) but this much depends on the condition of the genitalia of the respective specimens (cerci are in different position during copulation, oviposition and at rest).

Figs
Figs 1-7.Rearing methods of Quametopia species: 1 -laboratory equipment for adult maintenance and oviposition (A -stem of Impatiens capensis, B -cut and split stem, C -pieces of split stem on sand in small (internal) Petri plate, D -bottom (external) Petri plate lined with filter paper, E -cup with screen bottom and lateral hole, F -rubber stopper, G -mixture (1:1) of ground sucrose and bee pollen, H -assembled cup cage); 2 -cup cage with adults of Q. terminalis used for oviposition; 3 -Petri plates with pieces of both Impatiens and Clintonia borealis for oviposition-choice trial (egg arrowed); 4 -a piece of dead Clintonia leaf with 3 larvae and 1 puparium inside (arrows); 5 -rearing boxes with dead stems of Impatiens; 6 -dead stems of Impatiens with roots (arrow), used for dissections; 7 -detail of hollow roots containing overwintering dipterous larvae (arrow).Photo: K.N.Barber.
rounded tube.Accessory gland inconspicuous as in Q.

TABLE 1 .
Quametopia terminalis and Tricimba trisulcata and associated parasites from intact overwintered stems and a dissected subsample of stems of Impatiens capensis, May-June 2010.

TABLE 2 .
Number of eggs laid by Quametopia terminalis and Q. clintonia sp.n. in a choice trial comparing leaf and petiole of Clintonia borealis and stem of Impatiens capensis, 24-30 August 2009.