A New Species of Megaselia Rondani (Diptera: Phoridae) Reared from Larvae of Moths (Lepidoptera: Limacodidae)

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A New Species of Megaselia Rondani (Diptera: Phoridae)Reared from Larvae of Moths (Lepidoptera: Limacodidae)Author(s): Teresa M. Stoepler and R. Henry L. DisneySource: Proceedings of the Entomological Society of Washington, 115(1):85-95.2013.Published By: Entomological Society of WashingtonDOI: http://dx.doi.org/10.4289/0013-8797.115.1.85URL: http://www.bioone.org/doi/full/10.4289/0013-8797.115.1.85

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PROC. ENTOMOL. SOC. WASH.

115(1), 2013, pp. 85–95

A NEW SPECIES OF MEGASELIA RONDANI (DIPTERA:PHORIDAE) REARED FROM LARVAE OF MOTHS

(LEPIDOPTERA: LIMACODIDAE)

TERESA M. STOEPLER AND R. HENRY L. DISNEY

(TMS) Department of Biological Sciences, George Washington University, 2023 GSt. NW, Ste. 340, Washington, DC 20052, U.S.A.; Present address: AHS Jr. AgriculturalResearch and Extension Center, Virginia Tech, 595 Laurel Grove Rd., Winchester,Virginia 22602, USA; (e-mail: [email protected]); (RHLD) Department of Zoology,University of Cambridge, Downing Street, Cambridge; CB2 3EJ, U. K. (e-mail: [email protected])

Abstract.—Megaselia larvivora Disney, n. sp. is described from a series rearedfrom the caterpillars of Acharia stimulea Clemens and Lithacodes fasciola Herrich-Schaffer feeding on Quercus alba Linnaeus. The fly closely resembles M. fisheri(Malloch), whose recognition is clarified. Observations from the rearing ofMegaselialarvivora Disney, n. sp. are reported.

Key Words: Acharia stimulea, description, facultative parasitoid, Lithacodes fasciola,Nearctic, new record, slug caterpillar

DOI: 10.4289/0013-8797.115.1.85

Although the larvae of many speciesare saprophagous, the biologically diversedipteran family Phoridae also includesfungivores, kleptoparasites, parasitoids,predators, and even herbivorous species(Disney 1994, Feener and Brown 1997).Of all the dipteran parasitoids, phoridshave the most taxonomically-broad hostrange including lepidopterans, beetles,vespid wasps, bees, ants, termites, spidersand myriapods (Feener and Brown 1997,Coupland and Barker 2004). Among thebest-known parasitoid phorids are thosethat specialize on adult ants, includingleaf-cutter and fire ant workers; larvaedevelop in the host’s head, often leadingto decapitation (Feener and Brown 1997).There are also several reports of phoridsreared from caterpillars, pupae and adultsof Lepidoptera (Disney 1994). In many of

these cases, the phorid species rearedwere known saprophages (e.g.,Megaseliagiraudii [Egger],M. rufipes [Meigen],M.scalaris [Loew] and Puliciphora pulexDahl), and the hosts were already deadwhen attacked (Eyles 1965, Disney 1994).In most other cases, the host was likelymoribund or diseased when attacked(Balduf 1928, Disney 1994), however afew appear to be true cases of para-sitoidism. For example, M. chlumetiaeDisney has been observed attackinghealthy caterpillars of the mango shootborer,Chlumetia transversa (Disney et al.1992).Megaselia chlumetiae lays eggs onthe host’s integument, the larvae hatchwithin hours to days, penetrate the host’sintegument and develop internally (Disneyet al. 1992). In the case of M. chlumetiae,pupation occurs within the host remains;

in other species, pupation may occur out-side the host (Disney et al. 1992, Disney1994). However, the natural history ofmost phorid genera is poorly understood.

Limacodidae (Lepidoptera) is a familyof forest moths whose larvae are knownas “slug caterpillars” for their mode oflocomotion which involves a high degreeof ventral contact with the leaf surface(Epstein 1995). Some of these unusuallarvae are brightly colored and possessstinging spines, while others are mor-phologically and behaviorally cryptic.Among the most notable features of lima-codid biology, species of limacodid larvaeare exceptionally polyphagous in their useof host plants, although they prefer to feedon smooth leaves (Lill et al. 2006), andlarval development is extremely pro-tracted, typically requiring two or moremonths from egg hatch to cocoon for-mation (Stoepler 2012). The parasitoidcomplex associated with limacodid larvaehas been recently circumscribed as part ofa long-term study on the ecology of theLimacodidae by John Lill and his lab(Gates et al. 2012).

In 2011, one of us (TMS) reared a spe-cies of Megaselia Rondani (Metopininae:Diptera: Phoridae) from the larvae of twospecies of these moths, Acharia stimuleaand Lithacodes fasciola. These were sentto Dr. Brian Brown (Natural HistoryMuseum of Los Angeles County) whopassed them on to RHLD. The fly provedto be a new species closely resemblingMegaselia fisheri (Malloch 1912).Megaselialarvivora Disney, n. sp. is described be-low and details of its larval habits arereported.

MATERIALS AND METHODS

As part of a field study on the ecologyof host caterpillar-parasitoid interactionsin the Limacodidae (“slug caterpillars”),Lithacodes fasciola Herrich-Schaffer and

Acharia stimulea Clemens larvae werereared from laboratory colonies andexposed to parasitism in the field. Labo-ratory colonies originated from mothscollected from the forest at lights at night;these moths were mated in the laboratory,and hatching caterpillars were immedi-ately transferred to fresh, excised whiteoak leaves (Quercus alba L.). Larvaewere reared at 25 °C in closed 540 mlplastic deli containers with moistenedfilter paper discs and fresh leaves collectedfrom the forest, replaced as necessary, forseveral weeks. Larvae were then deployedonto white oak saplings at Little BennettRegional Park in Clarksburg, MD, a 15km2 secondary forest composed of pri-marily beech, oak and hickory. The bodylength of each larva was measured withcalipers to the nearest 0.1 mm immedi-ately prior to deployment. These sentinellarvae were exposed to natural enemieson saplings for 5 days in two separatecohorts (L. fasciola: 225 caterpillars ex-posed Aug. 17 – 22, 2011; A. stimulea:180 caterpillars exposed Aug. 25 – 30,2011). At the end of each exposure pe-riod, each sapling was exhaustivelysearched and all remaining larvae wererecovered (n = 126 L. fasciola and 94 A.stimulea). Recovered larvae were mea-sured once again and examined for signsof parasitism. Healthy-appearing larvaewere placed in mesh sleeve cages (100 x100 mesh/inch) on each sapling, wherethey were measured and monitored forsigns of parasitism weekly. Larvae thatappeared parasitized (discolored, notfeeding or moving) were collected at eachcensus, placed in individual, tightly-sealedplastic rearing containers, and fed freshleaves until either the larva pupated, died,or a parasitoid emerged. Emerged flieswere preserved in 95% alcohol. RHLDsubsequently mounted these on slides inBerlese Fluid (Disney 2001). Voucherspecimens are deposited in the University

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON86

of Cambridge, Museum of Zoology(UCMZ) and in the Smithsonian NationalMuseum of Natural History (NMNH).

RESULTS AND DISCUSSION

Rearing observations.—Five exposedlimacodid larvae (n = 2 Acharia stim-ulea, n = 3 Lithacodes fasciola) sub-sequently yielded Megaselia larvivoralarvae, pupae and/or adults (2.1 and 2.4%of recovered A. stimulea and L. fasciolalarvae, respectively; Table 1). Each ofthese larvae was recovered from a sepa-rate white oak sapling, however all fivesaplings were within the same area of thestudy site (within ca. 50 m of each other).After the 5 day period of field exposure,all of the larvae were alive and 4 of the 5appeared healthy for ca. 1 – 3 weeks (andin one case 6 weeks, Table 1) beforeshowing signs of parasitism (discolor-ation, not feeding, not moving). The re-maining L. fasciola larva appearedparasitized immediately after the 5 dayfield exposure period (Table 1). Four ofthe five host larvae completely liquefiedquickly after the first signs of parasitism.From the liquid remains, M. larvivoralarvae appeared, pupated away from hostremains 11 – 14 days later and adultsemerged singly 3 – 7 days following pu-pation (approximately 30 days after hostswere collected from the field). Eachlimacodid host yielded severalM. larvivoralarvae and 0 – 11 adults survived to emerge(Table 1). The remaining L. fasciola larvadid not liquefy as did the other hosts, butinstead appeared to become “mummified”by a braconid parasitoid (possibly Triraphisdiscoideus Cresson; Kula et al. 2009).Two M. larvivora larvae hatched fromthis mummified host, but failed to pupate(Table 1). We are confident thatMegaselialarvivora parasitism is highly unlikely tohave occurred in the laboratory given thateach host larva was reared individually in

a closed, plastic deli container with atightly sealed, plastic lid (540 ml, Fabri-Kal, Kalamazoo, MI) from the periodimmediately following collection fromthe field until M. larvivora larvae wereobserved in the rearing containers.

Given that four of the five limacodidlarvae that subsequently yielded M. lar-vivora flies appeared healthy after theperiod of field exposure, it appears thatin at least some cases M. larvivora actsas a facultative parasitoid, likely attackingliving, but diseased or otherwise immu-nocompromised caterpillars (Eggletonand Belshaw 1992). We are unable todetermine whether M. larvivora femalesoviposit onto healthy caterpillar hosts.However, based on what is known aboutphorid parasitoid biology, we suggest thatthe most likely course of events is thatduring field exposure, these caterpillarswere first infected with a pathogen (e.g.,a virus or bacteria) or were previouslyparasitized by another parasitoid, but didnot display visible symptoms of infectionto the human observer. We posit thatMegaselia larvivora eggs were then laidon the victims, the larvae entered them,and the post-feeding third instar larvaethen left the host to pupariate. Consistentwith this hypothesis, the observed “liq-uification” of four of the larvae is similarto the symptoms of commonly-observedvirus infection (Stoepler, pers. obs.) andthe mummified L. fasciola larva was al-most certainly previously parasitized bya braconid wasp in the field.

Megaselia fisheri (Malloch 1912)

As the new species closely resemblesM. fisheri, the recognition of this speciesis clarified before proceeding to a de-scription of the new species.

Aphiochaeta fisheriMalloch 1912: 463 (female).Aphiochaeta inaequalis Malloch 1912: 464

(male and female). Not Brunetti 1912: 511.

VOLUME 115, NUMBER 1 87

Table

1.Megaselialarvivora

emergence

andlimacodid

caterpillarhost

records.Specim

enTS11-858containstheholotype.

Allem

erged

adultsaredepositedin

the

University

ofCam

bridge,Museum

ofZoology,England,exceptfor1maleand1femaleparatype(both

from

TS11-858)whicharedepositedin

theSmithsonianNational

Museum

ofNaturalHistory,Washington,D.C.

ID

Hostspecies

(Lim

acodidae)

Dates

ofhost

exposure

infield

Daysuntilappeared

parasitized

after

hostrecovered

Hostbody

length

when

recovered(m

m)

Hostfate

Megaselialarvivora

hatch

andem

ergence

(n)dates

Totalnumber

and

sexofem

erged

M.larvivora

adults

TS11-611

Lithacodes

fasciola

8/17–8/22/11

08.9

Liquefied

Larvae(2)9/5/11

1—♂

Pupae

(2)9/16/11

Adults(1)9/26/11

TS11-706

Lithacodes

fasciola

8/17–8/22/11

95.1

Mummified

Larvae(2)Dateunk.

N/A

Larvaefailed

topupate

TS11-724

Lithacodes

fasciola

8/25–8/30/11

97.0

Liquefied

Larvae(5)9/2/11

2—

1♂,1♀

Pupae

(5)9/16/11

Adults(2)9/19/11

TS11-852

Achariastimulea

8/25–8/30/11

42

19.7

Liquefied

Larvae(2)Dateunk.

N/A

Pupae

(2)Dateunk.

Adultsfailed

toem

erge

TS11-858

Achariastimulea

8/25–8/30/11

21

8.0

Liquefied

Larvae(?)Dateunk.

11—

3♂

8♀

Pupae

(?)Dateunk.

Adults(11)9/28/11


Aphiochaeta tertia Brues 1915: 134. Nom.nov.

Megaselia fisheri (Malloch). Brues 1950: 71.

Malloch (1912) described this speciesfrom two females under the nameAphiochaeta fisheri and from both sexesunder the name A. inaequalis. Aphiochaetainaequalis was a name preoccupied andso it was subsequently renamed A. tertia.Borgmeier (1966) then synonymised M.(A.) tertia with M. fisheri. Brian Brownkindly arranged for the loan of the femaletype of M. fisheri and the male holotypeand a female paratype ofM. tertia. Thesehave been remounted on slides.

Malloch had distinguished his two‘species’ by a difference in the size ofthe proboscis of the females and thecolor of their postpedicels. However, thepostpedicels of the type ofM. fisheri andthe female paratype of M. tertia are es-sentially the same yellowish brown. Themaximum width of the labrum of thetype of M. fisheri is about 0.17 mm andabout 0.18 mm in paratype female ofM. tertia, despite this species supposedlyhaving the smaller proboscis. However,the lobes at the rear of abdominal ster-num 8 of the female (see below), and otherfeatures not considered by Borgmeier,confirm his synonymy of M. tertia withM. fisheri.

When Borgmeier (1966) re-describedthis species, he devoted an entire page tothe female but only a fifth of a page tothe description of the male. The onlyfigure provided by either Malloch orBorgmeier is a figure of the female’swing. Because the anterior scutellar bristlescompared with those of the posterior pairare relatively longer in the female than isthe case with the males, the females arekeyed in Borgmeier’s Group VI. As M.fisheri clearly belongs to the M. giraudii(Egger) – M. densior Schmitz speciescomplex, its female was included in the

key to the European species of this com-plex (Buck and Disney 2001), despitebeing Nearctic, and an amplified descrip-tion was provided that included figuresof the abdominal sterinite 7 and the un-usually closely approximated lobes at therear of sternum 8.With regard to Europeanspecies of this complex, further specieshave since been recognized (Brenner2006, Disney and Pagola-Carte 2009,Disney et al. 2010, Disney 2011). TheNearctic members of this complex re-main poorly known.

With their shorter anterior scutellars, themales belong to Borgmeier’s Group VII.However, due to an oversight, Borgmeierfailed to include the male in his key to thespecies of Group VII. In this key forGroup VII, species couplet 2 dividesspecies into thosewith the costal cilia thatare ‘short’ as opposed to those in whichthey are ‘long’. Borgmeier (1964), in PartII of his monograph, defines ‘short’ asbeing < 0.1 mm and ‘long’ as being 0.1mm or more. However, in this species (asin many others) the cilia at the end ofcostal section I are long, but those onsection 3 are short. By treating the costalcilia as ‘long’, the males run to couplet 11on page 4, where the distinction is basedon females alone. However, the modifiedovipositor segments of the female ofM. compressa Borgmeier immediatelyexclude M. fisheri. Furthermore theunknown male ofM. compressa, like itsfemale, will have the antial bristles ofthe frons closer to the anterolaterals thanto the upper supra-antennals (as opposedto being about midway between thesetwo bristles) and will have only 2 (asopposed to 3) bristles on the axillary ridgeof the wing. The last two features willalso distinguish M. fisheri from the sec-ond option at couplet 11, M. straminipes(Malloch).

Additions to the description of themale are given below.


Male.—Postpedicel with numerousSPS vesicles whose greatest widths are0.007 – 0.012 mm. Palp about 0.19 – 0.20mm long, with 7 bristles (the longestbeing about 0.7 times as long as palp)and with about 10 hairs. Labrum strawyellow to light brown and 0.17 – 0.18mm maximum width. Labella each witha dozen to 20 short spinules on lowerface. Notopleuron of thorax with onlytwo bristles and no cleft. Hypopygium asFig. 1, with the epandrium and hypan-drium being largely brown, apart fromthe pale posterior lobes of the latter. Analtube is yellow. Abdominal tergites with

T1 yellow, T2 – T4 largely brown, T5yellow but brown toward sides and T6largely yellow. Some stronger hairs post-erolaterally on T2 and longer, bristle-like,hairs at rear of T6. Venter with hairs onsegments 3 – 6, with longer, bristle-like,hairs in middle of rear margins of 5 and 6.Front tarsus with a posterodorsal hairpalisade on all five segments and 4 and5 subequal in length. Dorsal hair palisadeof mid tibia extends about 0.8 times itslength. The longest hairs below the basalhalf of the hind femur are longer thanthose of the anteroventral row of the distalhalf. Wing 1.9 – 2.0 mm long. Costal

Fig. 1. Megaselia fisheri male, left face of hypopygium. Scale bar = 0.1 mm.


index 0.50. Costal ratios 3.4: 1.9: 1. Costalcilia (of section 3) 0.12 – 0.13 mm long.

Vein Sc does not reach vein 1. A small hair

at base of vein 3. With 4 axillary bristles,

the outermost being 0.12 mm long.

Megaselia larvivora Disney,new species

Recognition.—The female is imme-diately distinguished from that of M.

fisheri by the largely yellow abdominal

tergites and by the lobes at the rear of

abdominal sternum 8 being well sepa-

rated (Fig. 5). In the keys of Borgmeier

(1966), the males run to 11, lead 2 on

page 4, as does the omitted M. fisheri

(see above). Like the latter species, it has

a longer anal tube than M. straminipes

(Malloch). The males of M. larvivora

and M. fisheri closely resemble each

other. They differ in that M. larvivora

is smaller than M. fisheri, whose wingsexceed 1.5 mm in length whereas thoseof M. larvivora are less than this. Other-wise the posterolateral region of the leftside of the epandrium differs and thedifferentiated longer bristle of this side isrelatively stronger in the new species(cf. Figures 1 and 3).

Description.—Male.—Frons brown,clearly broader than long, with 60 – 80hairs and dense but extremely fine mi-crotrichia. Supra-antennal bristles (SAs)very unequal, with the upper pair almosttwice as long as the lower pair and muchmore robust. The antials a little lower onfrons than anterolaterals, and a littlecloser to the upper SAs than either isfrom an AL, which is very slightly higheron the frons than the upper SAs. Pre-ocellars a little closer together than eitheris from a mediolateral bristle, and the fourbristles at about the same level. Cheek

Figs. 2–3. Megaselia larvivora male. 2, SPS vesicles of postpedicel. 3, left face of hypopygium.

Scale bars = 0.1 mm.


with 3 – 5 bristles and jowl with two thatare longer and more robust. The sub-globose postpedicels yellowish brownwith a paler arista, and with at least 40subcutaneous pit sensilla (SPS) vesiclesas Fig. 2. Palps pale yellow, and viewedfrom above about a third as broad aslabrum, with 7 – 8 bristles (the longestbeing about 0.38 times as long as palp)and 6 – 8 hairs. Labrum yellow and about3 times as wide as a palp. Labella coloredas palps and with fewer than 20 shortspinules below each. Thorax yellow toyellowish brown on top and paler on sides(apart from brown on pteropleuron). Twonotopleural bristles and no cleft in frontof these. Mesopleuron bare. Scutellumwith an anterior pair of small hairs anda posterior pair of bristles. Abdominal

tergites with T1 yellow, T2 – T4 largelybrown, T5 yellow with brown patchestoward sides and T6 yellow. The hairsa little longer toward sides of T2 andlongest at hind margin of T6. Venter paleyellow, and with hairs on segments 3 – 6of which those on segment 6 are longest.Hypopygium largely brown apart fromthe subequal pale lobes at the rear of thehypandrium, with a yellow anal tube, andas Fig. 3. Apart from brown patch on midcoxa and tip of hind femur, legs yellow.Fore tarsus with posterodorsal hair pali-sade on segments 1 – 5 and 5 a littlelonger than 4. Dorsal hair palisade of midtibia extends about 0.7 times its length.Hairs below basal half of hind femurlonger than those of anteroventral rowof outer half. Hind tibia with 11 – 12

Figs. 4–9. Megaselia larvivora female. 4, Dufour’s crop mechanism. 5, lobes at rear of abdominal

sternum 8. 6, furca. 7, tergite 7. 8, sternite 7. 9, tergites 5 and 6. . Scale bars = 0.1 mm.


differentiated posterodorsal hairs, allbut the most basal 3 – 5 being robust,and spinules of apical combs simple.Wings 1.28 – 1.43 mm long. Costal index0.46 – 0.48. Costal ratios 3.2 – 4.8: 1.7 –2.4: 1. Costal cilia (of section 3) 0.08 –0.11 mm long. Hair at base of vein 3 small.With 3 axillary bristles, the outer beingabout as long as costal cilia. Sc not reachingR1. Veins light brown, but thin veins 4 – 6a little darker. Membrane lightly tingedgray (just evident to naked eye when

viewed against a white background).Haltere with yellow knob.

Female (Fig. 10).—Head similar tomale except that labrum is at least 5times as broad as a palp, and 1.3 – 1.5times wider than diameter of postpedicel.Thorax as male except the anterior scu-tellars are 0.68 – 0.84 times as long as theposterior pair. Abdominal tergites 1 – 6essentially yellow (Fig. 10). Some hairstoward the sides of T2 and at the rear ofT6 longer than the rest. T5 – T6 as Fig. 9.

Fig. 10. Megaselia larvivora female, right face. Scale bar = 1 mm.


T7 brown and as Fig. 7. Venter yellowishgray and with hairs below segments 3 – 6.Sternite 7 brown and as Fig. 8. Postero-lateral lobes at rear of sternum 8 as Fig. 5.Cerci pale yellow, 0.09 mm long and 0.02mm wide (i.e., about 4.5 times as long asbroad) and with 6 relatively short hairsand a very long pre-apical hair that isabout 0.11 mm long. Furca brown, butnot dark, and as Fig. 6. Dufour’s cropmechanism as Fig. 4. Legs similar tomale but the longest hair below base ofhind femur only as long as those of an-teroventral row of outer half. Wing asmale except length 1.7 – 1.9 mm. Costalindex 0.46 – 0.49. Costal ratios 4.5 – 6.2:2.0 – 3.8: 1. Costal cilia 0.09 – 0.12 mmlong. With 3 – 4 axillary bristles, with theoutermost longer than costal cilia. Oth-erwise it and haltere as male.

Type material.—Holotype male:U.S.A.,Washington D. C., George WashingtonUniversity, reared from larva of Achariastimulea, exposed on 25.viii.2011 for5 days; adult flies emerged 28.ix.2011,T. M. Stoepler, deposited in the Univer-sity of Cambridge, Museum of Zoology,England. Paratypes: 2♂, 8♀ as holotypeexcept 1 male and 1 female deposited inthe Smithsonian National Museum ofNatural History, Washington, D.C.; 2 ♂,1 ♀, as holotype except reared from lar-vae of Lithacodes fasciola exposed on 17.viii.2011, the flies emerging 19 & 26.ix.2011.

Etymology.—This species is namedfor its feeding biology (larva-feeding).

ACKNOWLEDGMENTS

TMS thanks Dr. John Lill (AssociateProfessor, George Washington University)for his support of her studies, DanielaAndriamalala and the Hormiga lab(GWU) for assistance with and use ofphotomontage software, and WendyHanley (Park Manager, Little Bennett

Regional Park) for facilitating research atthe study site. We thank Martin Hauser

and two anonymous reviewers for com-

ments on the manuscript. RHLD’s studies

of Phoridae are currently supported by

grants from the Balfour-Browne Trust

Fund (University of Cambridge) and

the Systematics Research Fund of the

Linnean Society and the Systematics

Association (UK).

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