I). L. W?gner et al,
Introduction With its fiery orange forewings, iridescent blue-black hindwings, and 10-centimeter (cm) wingspan, the regal fritillary was one of Ne\; England's most magnificent grassland insects. Although always locali' distributed, it formerly occurred in all six New England states (Scud Status Update and Life History Studies 1889, Denton 1900, Klots 1951, Opler 1983, 1992). Preferred hahita included marshes and swamp edges, wet meadows, fields, pastures. on the Regal Fritillary (Lepidoptera: and native grasslands (Denton 1900, Weed 1926, Clark 1932, Clark a" Nymphalidae) Clark 1951, Kiots 1951, Schweitzer 1992, Glassberg 1993; see also 1). An examination of the numbers of specimens in collections and accounts in the literature indicates that the species reached its greatc. - abundance in New England in Massachusetts on the sandplain grass- David L. Wagner, Matthew S. Wallace, lands of Martha's Vineyard, Nantucket Island, and other offshore George H Boettner, and Joseph S. Elkinton islands (Scudder 1889, Jones and Kimball 1943). On the mainland an: on Block Island, RI, the species was commonly associated with wet meadows and human-maintained grasslands created by mowing, grazing, and other agricultural practices. As in other Speyeria. the larvae are violet specialists. New England's regal fritillary colonies were associated with ovate-leaved violet (V/olafimbriatula), lance- leaved violet (V lanceolata), common blue violet (V papllionacen and birdfoot violet (V pedata) (Schweitzer 1987, Cassie et al. in pre Abstract D. Schweitzer pers. comm.). Eastern populations of the regal fritillary (Speyeria idalia) have been declining since Adult males begin emerging in mid- to late June, followed one to at least the late 1940s. New England populations disappeared from north to south, with two weeks later by the first females (Scudder 1889, Clark and Clark the last viable colonies occurring on island sandplain grasslands and heathlands off the 1951, Barton 1993, 1994). Adults are long-lived, with some individu - coasts of Bhode Island and Massachusetts. Extensive field surveys in 1992 and subse- surviving up to 90 clays in the wild (Barton 1993). Although female'- quent efforts have failed to locate any colonies in New England. Females are perhaps the most fecund of all butterflies. In this study, eight captive females laid more than 1.300 are believed to pair shortly after emerging from the chrysalis, few e eggs per female (range 227-2,494), and egg hatch ranged from 19 to 78 percent. The are laid until the latter half of August or early September. Eggs hatch addition of less than 5 percent raw albumin to the honey-water diet of three adult after two to three weeks, and the first instars immediately enter females in 1993 coincided with an increase in daily egg output and hatch. One cohort of diapause, presumably without eating (Edwards 1879, Maynard I 88(" laboratory-bred larvae was almost entirely lost to a nuclear polyhedrosis virus. Young Scudder 1889, Weed 1926, Mattoon et a!. 1971). Feeding begins in violet (Viola) leaves were suitable for the establishment of first instars, but mature foliage was not; the later resulted in 100 percent mortality of first instars. spring, and larvae complete their development by late May orJune We discuss reasons for the regal fritillarys decline and make management recommen- pupal stage lasts an additional two-and-a-half to four weeks (Edward'- dations. Because females frequently oviposit away from the host plant, dense violet 1879, Maynard 1886, DLW unpubi. data). colonies should be especially advantageous for the establishment of the minute first- It is not known exactly when the regal fritillary began its decline instar larvae. Given the long life of adults and their propensity for nectar, we think that New England, but a letter from amateur lepidopterist John Bakeles'- - the availability of late-summer nectar will be essential in efforts to maintain or reestab- lish this striking insect. the correspondence of Alexander B. Klots (Homer Babbidge Lihrar Special Collections, Univ. of Connecticut) indicated that viable coli nies in Connecticut were already disappearing by the late 1940s. TL last known Connecticut colony, located on the Roxbury-Bridgewat. town line, disappeared in 1971 (R. Muller pers. comm.). The speck- demise evidently proceeded from north to south, with the last main - land sightings occurring in the 1970s and early 1980s. By the mid- 1 980s only six populations remained, all on offshore islands: Block - 261 262 I). L Wa ,ur et (f 1, RdJ I ;';ti1!i; Butterfly In Neu' l'ng!wid Conanicut ishinds in Rhode Island and \Lirt ms Vi lt\ iii, santuekct, Naushon Island, and No \lans I and in Ia'-,sar liuseits ( I ether 1), S hweit,er unpuhl. data. Ibis pattern l distppearing Ironi north to south was repeated in New York. where the. at populat ions were recorded in the I 980s on long Island. INc last New lngland regal fritillary was seen by Iupidoptcrist Iarr\ ( alI in I O) I m the hon h end of Block Island. loctised searches ot the six ilorementioned islands in July and August 1902 yielded no sightings, nor have any hcen reported since. In December 199 I a meeting was org;tnuied by Scott \lelvin, an endatigered-specics biologist with the \lassaeluisctts l)ivision of Fisheries and \Vildli he, ho review the status of the tegal Iritillary in New Fogland and New Yi)rk. 'Ihie working group ineludt'd stall from the U.S. Fish and WiIdhik Service, Massachusetts I )ivision oh Fisheries and WikUife, Natural 1 heritage Programs in Massachusetts. Rhode Island, and Connecticut, The Nature ( onservancv, and Massachusetts Audu bon Society; unhersitv entomologists: and other informed parties. The group recommended a two-step plan ol action: first, to establish a captive breeding protocol (if any remaining hutterl lies were genetic stock from indigenous populatu ins e mId he used to reestablish the species) and second, to studs management issues. nlv later did participants learn t hat ii ic rcgat liii dlar\ v as already extirpated from New England arid that breeding stock would have to he acquired from outside the region. 13el4 w we discuss results from our efforts in I 902- I °93 and 1993- 199i to establish rearing protocols for this britten lv and i)osSil)lC reasons for the species decline. We discuss brief N two management issues: violet density and nectar av,tilabiflt
Methods Females were obtained from colonies in Penns lvania (1992; n = 3), Missouri (1993; n 2', and Iowa (19Q. ii = 3). lach female was individually housed in a 12-liter (1) cardboard ice cream carton covered with netting and placed on a bench near a south- or east-facing win- dow, Dried violet leaves and strips of ( rumpled toweling were pro- vided as ovipositional uhstrates, and oiihaiiiers were misted with water two or three thntcs a week to si inulate rainfall Feniales were
- removed and fed once a day with a sugar or hione' solution using the feeding stations of Maiioon et al. (1971), Once a teniale had rcpeatedl retracted her proboscis or 15 minutes had l)155e(l, she was gently rinsed off with water and returned to her (ontainec, In 1992 a small portion of raw chicken-egg albumin (le's than 5 percent by volume)
264 26-a, Regal Fr/f/Flair /ii!tcifiy in New Fiih,'iid 1). L. ll'ai,oicrctaj, was added to the diet at the onset of egg-laying; in 1993 this was done moistened filter paper had been added. l:ft.sl le.i us were provided 20 and 22 days after oviposition had commenced. daily, although a portion of the older leaves was mlw ;n's left as well. Newly deposited eggs were collected every day, placed in plastic vials, and misted once a week to levatc humidities. Eclosion was monitored daily; newly hatched first inst ars were t ranskrred to steril- Results ized, moistened wooden blocks, placed In a l)L15t1( bag, and held in a 'lime three regal fritil!arv females e illected 2() August 1 ')2 in Penn- refrigerator at 2 to 5° C (Mattoon et a!. I °7 I). Blocks were remoistcned sylvania laid an average of I .-'t 0 eggs (range )(- 1 .8 mtie die two every six to eight weeks, Larvae in moldy bli K (5 were transferred to tcmales captured in Missouri in late August I 003 laid an ii ei'agt' of blocks that had been aut iclaved 1 11 S eggs (range 3m I 2, I) m, and tIme three females apt ured in In the hope of establishing a !aboral n'y colony that would produce Iowa in late August and earls- September 00 laid au a' erage of' multiple generations each year, we prevented first inst arc from enter- 1,l5'.3 eggs (range 22.2 tO). 'lime females laid eggs oii the sides of ing diapause during the winter by exposing them to long day lengths the cardboard containers, nylon screening, 1 riinipk'd pafltt. and dried and warm temperatures (\lattoon ('t at. }Y" I I l)iapausing first instars violet k-ayes lhe 'vere t-spcciall' I)! 0 plat C o\ a iii protected were placed on ne violet caves in glass petri plates with moistened t'i'eviCes and seams. filter paper and held below an incandescent light to raise temperatures Captive females laid eggs oer a four- to ix week period, with peak to 25 to 30° C fbr 1't lit,urs each day Fhe fitter paper was kept moist to production occurring in the first bnn' weeks ol ovipositional activity ensure that the humklits remaiiietl high. and a new violet leaf was (Fig 2A) \ f'ter the first week in )etober, egg p duction wa modest. added daily. Larvae exposed to this regimen usualh- began feeding The last eggs were laid on I i October (I 002) amid 28 October I 993), within a few hours to two days a small percentage of stragglers took All females died within If) da s t>f their last oviposit i( iiial date, up to 20 days to begin ficding). Larvae h.-d on all four of the violets we W- noted marked fluetuat ions in dail egg product ion in both years offered: ovate-leaved violet, common blue violet, birdfoot violet (all of of the siud' In he second year of the study, albumin was added to the which grow wild in New England), and domestic cultivars of the three- diet of One female tin day 20 18 september 1 003 md to the diet of colored violet (V tricolor). two oilier females on dt' 22 2(1 epteniher U)O1,) 'l'lOs addition was Adult offspring Irom midwestern stock obtained in 1993 were followed by an increase in t'eettnditv (Fig. 211). l-gg liat Ii from our successfully paired in glass enclosures in January 1994. A second eight captive females ranged fri nit 10 tii $ percent ( sunimiuig across generation 0 larvae was obtained in February, and this cohort was all vmpositioiial dates for a gi en female) - Percentage egg hatch was immediately "forced through to the adult stage. imilai' for out- three collections: iowa OS pci'ccnt (H 2. \licsouri 62 Two species of violets associated with regal fritillary populations in pei' cut (n = 2), and Peilns\ lyania pe' cut (u 3)- t tcaltliv eggs New England, hirdfo it and vate-leaved vi ilets, were used in a h st w etc white in' cream colt n'ed w hen laid and darkened to a frosted gra' and leaf preference studs Wikl collected l)ltntS of these two specieS as the catci-pllat-s mat tired - Eggs that failed tii I iucl i were of ten velli nv and other violets were grown in the greenhouse at the Univcrsit\ of and collapsed The last eggs to be laid w'err often inviable. As in egg Connecticut. Larval establishment and survivorship were followed for production, Wi' noted considerable fluctuat U )ns in egg hatch - lii 1003 three size-age ctassc of leaves. For ovate-leaved violet, sue-age elases an increase in hatch coincided w ith the ;mddit h in of albumi-min to the were assigned as follows you ii g leaves were I to 3 cm long, pale green, female's honey-water diet, alt hough this ohsor ation wa largely (luc to and had the bases of the leaves curled over the midrib; intermediate the lw'm'fortnance of a single ft-male (Fig. ). leaves were 2 to 4 cm long, had begun to darken, and were mostly In 1083, larval development arid suivivorship were h llowcd for a uncurled; and mature leaves were 2 to 0 cm long and were fully single cohort of 80 larvae t fiat were st arted on r)tiinion blue violet darkened and expanded. For hirdihot violet, leaf ages were assigned as shortly after the larvae emerged from eggs in October The first five lOlli)WS young leaves were less than I millimeters (mm) long and still instars were of' similar duration, lasting iron an average if 3 0 days to had the lobes drawn together; intermediate hea\ es were 1 S to 20 mm 6.6 days. 'ilie sixth and final larval instu- was mitch longer, lasting an long, had begun to darken, and had edges that \s'ere partially uncurled average of I I days. and not touching the midrib; and mat ure leaves were fully darkened After noting that early inslars seemed to pretcr lc,mc s that wi-re (lot with the lobes separated. Rearing was done in glass petri plates to which yet 1u11 expanded, we set up an expert flint n examine larval cstab- lishimerti amid snrvivorslup nut leaves representing three si/c-age classes
265 200 I,). L ii (/,1.iWi' (1 Ii!. J(tlXUl .rriitiiarj' t,ITr('rf 1)' In IVelm" Fiigla'nd
00
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"Off (OfJOrtOfl day
.H,iire 3. Pcr /it1((c /11/11/i f)) d'o!c for /f 'c ;l/y(fj cs/ci'ii (opt turd rei'a/ fit/i//ar,e.s ( I 'P1 0 ,. nc 'i, 'cc ,,jai,In '1' o/i/ü 'c l/P'r 1 'tI 'cn f/ma/c t;cc in (I/U0/li/0 moc B)' i/ar '). oiii' /on,oli' /1(I(1.St(IJ)/O'(l l(l}'l11. //1' ítí' Ji, tU'o Iid ,v/oppcl. u/fni,n,;j /1(15 tj(/(/('(/ f) f//i' u/ic! Ii the' fbi '' ciii', 'ii iiiu iint/c,' oil iluiis 2(1 ,pu/ 22. Id' fi-oni ovate-leaved and hirdlcon violets Rot Ii v otng and itcrmt'diale a-.', fco%('(( ',Itt?)J, h ).ir a) ('stablislitIi('nt, tint iii )ar I(' survivi'ul niot'e than 1 3 d;tv, oil flat ore Jea ('S ol cit her o k't (Tahk 1). J,ar\ ac often reniained and continued to tis'cl f/li (fill'- tO thr('e-dav-( (1(1 leaves even after they had wilted. Discussion Jln' 2, dl eggs laid Nv one ()f(l(il' captive regal frifillarit s horn )Ut'i \Vere more hail twice the 1'( 1 nis titwiher 1< no\Vli tm i he latch Nv ati buttet') ft Species (I 20(1 h an ldtthi 'S chiec le Spot IJipIfl'drj'a IOi' edifIh-l: Lahinc flOS. It s p05//iNk' that ('(Ii gt'ualc'F lccutiditics ligure 2. Regal Ir/tillarj' /t'lpoS/tiofl. (A) .lIeau (la/I) eg,i pl'(XlUCt/(i// migh three l'ennsyli'ania-caplured feiiiaks (I 9)2) (i/bull? (('05 athl('d 10 th('i)' ha c heeit rt-alrscd had OUr captive lt'iiiales been given access to living d'' jhnn (lie first le' o/ co'iJ?(si!iOII Iwo feia/cs ii'en' la'f,n (/imug/i t/ie Howrs, soil extidates, and 'ft hi' nat ui'al substrates. ( .ornpared with f/is! (Uo i/u/cc; fhii'e I/males from Se/il. Il/It/I Oct.; tu'o fm/n to /2 Oct.; other hut tert lies, (lie rerah I 't lar' lc'ars to he a sveepstakt s and' 011)' (11W 1lI('1i(l/tJi (ii) )Ic(l?1 110/It ('L,0 /iiuiiliictioii for tJjiio' mn,ufi.ctt'ilI' Strategist' (hat. has plat.ed a l'('t11itfl1l 011 di'.' iittiiih&'r iii eggs U can (bets on 1/! and 2(1 captured fenzale.c ( f.H), albimmnai U'(mS (Id(/('(l II, Iheti produce. however, lint all eggs hatch. II )r I lie ('igh) wild-'. aught f/lrec/ei?iu/lesfI'r1lfl 2 Sefi!. One female u'ac /ai'i,li t/lrollç'/l flu' t/iI too dales: ,Sepr. auth 0 (Ic!.; tmmo from ( to / I Oct., dm1 olil)' one the rea/lec 267 L.. , , ( I Rrgo! / ritillary Butterfly in New Ei tg1 (/1(1 TahF' 2. Jan 'aT (le('elupln en! u/tue rt', i/Jill/I/ar)' .1 (('I/V, 1 / 8(11,11-i '(11' I(' 0 Tab/i' 1. /taI>1/s/lIHrn/ / firi-hitar iit,'af fri/i/lw) 1(111w' 11 (u'at('-lpaved fore I 'iii! of diupa;i.ie in em/I' () thei 19')3 I tee teal) - I;? 1ii(';ii;(eS( ('ut V (('1 (/11(1 l)i(I/Vf/ i i'Ir S/ui * /F 11)1 1S')ll?S)'/i 'a iiiu (1 '))2 /t'i;ialm. Te;, ((1111/) 1I'US lLc'(I (I) i'(//S(-' ,i'au'int,' t('il ?/)('i'(TtlIi'(',S (I ) aJipi'u.vi ;iiiII('l;' 2 -2" i iii 1(1';;' t ;'h '/ u ' i Ii .j liii (Ii I-I'; I; ' 1/( f(oe: ' , ,ni, p ;#b 'i-i ii ed jute, Si age (u mulat I ye a;a'f liiiill(1('. N Mean SD Iort Survival Number Surviving Day 1 Day 5 Day 9 Day 13 (d ) (d) (", 1st instar 86 "1) O' .IR' tc,Iv('d vk,it ______4.8 3.6 50 youniieiv5s 10 5 4 4 2d Instar 43 5.9 1.8 0 50 intermediate leaves 10 10 9 9 3d instar 43 4.1 1.3 5 48 mature lres 10 4 0 0 4th instar 41 3.6 1.2 0 48 birdloot violet Sthlnstar 41 6.6 1.7 0 48 young i"ves 10 5 5 5 6thinstar 41 17.1 27 37 32 intermediate leaves 10 6 5 3 Pupa 28 - 31 21 mttura eaves 10 4 1 0 Adult 18 - - "- 21 'Percent succumbing in animal dung. crc infertile or failed to hatch for sonic miter reason. Although adult and salts showed little i merest in ctntpltitg. ( )l the SO-phis admIts bred l( ins nutrition of our captive females or improper incubation cotidut In 1)ecember (199 ) and !anuai'v ( I 9t ), on! t v pairings were may explain our low hatch, similar levels ot egg taihire have been Ot)t.llrle(l over the four \Ve('kS the adults were housed together. ( )ttr ;bacrved in the Mormon fri illar\ (Spej'eiia inurinonia, ( L. Boggs inability to achieve mating in the winter remains the principal hurdle Pt't'5 comm.), to establishing a captive breeding program for this lititlr'rt lv U is likely that most mortality occurs in the first insiar, vli ich must stn'viv a six- to seven-month diapause having ted only on a portion of Reasons fir the Regal Fritillary's Decline it egg chorion (Edw.trrk 18'9. SciRkier I 880, Weed I 926, Mattoon et The regal fritillarv's decline has al. 19 I). Our lab-reared larvae consumed onl enough o the ehorion been noted range ide. There is only one viable eastern colony, in central Pentisylva nia; t lie range of the to tree themselves It'om the egg. Feniak's arc itidiserinunate egg layers, midwestern populations .115(1 to be ci mt raci ing ovipositing on grass soil, anti tinder pebbles, presumal)lv iii the tp'irs ii; ing he species western and northern boundaries Sc'liildk tiecki I ')$6 Nagel vicinity of violets hut onk i arelv plaeinU eggs ott viok'is (Setidder et aJ. 19'-) I , ( )pler 1 p92, Scliwcit,er 1 092, 1 99 ', (!assh'rn ui 1889. Clark 1932, Kots I 'cott 9%(', Ilartoti I O0, 13. Barton pets. Swengel I 993) l'xplanat ions tot' (lie decline include habitat trtgnicnta con-tm.); thus, the task 1)1 locating new '. tok') leaves in the spring must tion and conversion, lire, hurricane impact, pesticide tli'iit - collecting. he difficult for the tiny first instars. Flooding, mold, desiccation, competition with Other .SJx't'erfa. and t lie introduet h m and'or spread predators, and starvation undi nibiedl take high tolls, and the high of a pitrasit( iid r pathogen ( Nagel et a!. 1991 , Scliweitit-r I 992. I 99 fecundity of the regal tritillary helps compensate ibr the losses suffered Svengel I 993). Perhaps .ill ot these tat'tors have ont ribiitc d ii> the in the first instar. Lveit when l'irst-instar larvae were supplied with (k'flhis(' ol one or more populations, vet only the first. and possibh' the fresld\ picked loliage, losses were high SO' iii out' lab colony (Table 2). last, are apt to apply i'angcwide 'I lie regal fritillarv v as ,t likeN bench It was exceedingly difficult In our siudv to get captive-bred adults to ciary ot the extensive agriculture practiced across the eastern I pair in the winter. Adults housed in a flight cage (2 x 2 x 2 meters Em]) 'nitcd States and s 'uthern ( 'anada in t lie earl) twentieth t, entut'v. Past tires, and provided with high temperatures and humidities, natural light 270 269 I). L. Wagner et al, Regal / 'iThIfiiy Butterfly in New England hippolyta) has recovered in m near cxii ii I i ni (1 Ia nit i.it I 1 )H f'>r xample, are favor.tble area for bitt growth and have often 1989, Hammond and McC4rk Ic I 0') 1. It is i nlp ut .itit 1' ca nt i larval pro'. itied habitat for eg.iI fritillaries, c'p&ciafl when bordering establishment and survivorship are di ft'i entithi'. at icc tl by cltti'cni marshy lands or ponds, It is likcl3 that this grassland hutirf1y peaked violet species. in abundance in the \ortheast during ( ohonial times wheu much of Males and females of all fritillaries arc avid ne tar lccdcrs 1 lowe the region had been cleared for en ps and pasture (Clark 1914, Bell 1975, Dornfeld 1980, Ferris and Brown 1981). In the '.lu'm..tt li'itillary I 98) Conversely, as New England has reforested, suitable habitat for (Boggs and Ross 1993) and fritillary (.pe ei'ia :ureu the butterfly has diminished and become bncreacingly fragmented. Hammond and McCorkle 19')l), adult diet is known In fi t both \.igei et al. (1991) recorded an average daily movement of 0.07 longevity and fecundity. Regal tritihlaries feed on nectar throughout kilometers (km) for 23 recaptured regal fritillaries on the Rowe Sanctu- their long flight season, Iron hunt through September ( barton I 0') ary in Nebraska. Barton (1993, 1994) found that malts and females 1994). Summer mowings that remove nectar sour. es surli as thistle occasionally flew long distances: in 1993 the average distance moved (Cirsium), milkweed and butterfly weed Aslpios). i rottwecd by 22 marked adults identified as dispersers was 3 t kin. One male (Vernonia), and others (Scudder 18R9, Weed 1920. Nagul ci ;II. I)91, flew 15.8 km in a 12 hour period. In expansive taligrass prairies, Schweitzer 1992, Barton 1993, 1991) would force aduht. to m'vc away where the regal fritillary reaches its greatest abundance, such move- from emergence sites. Jflner et al. (1992) noted that a cnlon ot regal ments would not carry it away from suitable habitat as often as would fritillaries in Ohio disappeared after milkweed was eliminated at the he expected in the East where grasslands are usually small and isolated site. In Pennsylvania, an abundance of thistle sterns to he especially by forest. Perhaps the regal fritillary persisted longer on New England's important for the weliare of the regal fritillary (U. Barton pens. offshore islands because dispersing adults were repeatedly turned back comm.). by open ocean. Given the descriptive nature of otii cLiia, it is dill icuir to (Ofl(. hide In the spring of I we lost more than 80 percent of a second- with certainty that the addition of egg albumin to thi daily nectar" generation cohort of middle- to late-instar larvae to a nuclear polyhe- solution provided to our regal fritiliary It-males in ceased fec'und:tv and drosis virus (NPV). Prior to this we had seen very low mortalities in hatch (Figs. 2B and 3). Even if our results were coincidental nectar second to fifth instars (Table 2). The role that this or other pathogens quality and quantity must be essential to the rcproducti'. t' SUc('cs" of play in low egg hatch, the deformation of wings, or other aspects of such a long-lived butterfly. Management plans hoc the regal lrctilhary the species' decline remains unknown, but certainly patbogens need should include provisions to ensure the ;ivailahilit v .1 nectar at ,r near to be considered since a natural enemy such as a parasitoid or patho- those sites with highest violet densities. Acces'-. to nectar, cspe iahly in gen could account for the observed pattern of decline. Because patho- late summer when flowers can be very scarce in New liigiatid grass- gens tend to act in a density-dependent fashion, a highly specific agent lands, may prove & ritkal in etlorts to reestablish or manage tlii would be expected to occur at very low incidence or to disappear species. entirely as host populations dwindled (Anderson and May 1980). If regal fritillaries shared pathogens with other fritillaries or more dis- tantly related Lepidoptera, however, the disease agent could remain Acknowledgments abundant and cause continued high mortality, even as regal fritillary populations declined to extinction (see Holt and Lawton 1994). G. Leslie and R. Wilson of The Butterfly Place, Westford, MA, generously provided time, effort, space, and resuurc us and eagerly Reestablishment and Management Considerations: shared their knowledge of butterfly rearing. Many helped v. ilit the Violets and Nectar daily maintenance of our lab colonies: j. Adams, C. Boettner, 1,. Crowingshield, V. Giles, J. Henry, C. Iloule, j. P 'well, [). Wakoluk, and Compared with other butterflies, lcmale regal fritillaries appear to G. Witkus. The suggestion to add albumin to Ihic adult diet wa made be r-selection strategists with regard to their fecundity and oviposi- by P. Hammond. B. Barton and D. Schweitzer shared man' ohser a - tional behavior. Because females only occasionally oviposit directly tions and unpublished data with us. We are indebted to thou who onto violets, dense violet growth should promote survivorship of the helped us obtain gravid females: B. Barton and T. Wilkinson (Pennsyl- vulnerable first instars. Tree clearing, mowing, and thatch removal vania), R. Johnson (Missouri), and T. Orwig (Iowa). The figures were have been employed successfully to favor the proliferation of violets prepared by J. Henry. M. Mello located and dug up many of the violets on the coastal prairies where the Oregon silverspot Speyeria zerene 272 271 ItL',I.NFI I) iiI(H 1)101(1/i) i,1 \1f' 1/i,h1/lt! I). L. Wii,,('r et (71, used in the study. Two anonymous reviewers made helpful suggestions Holt, R. I) , andJ. H. I lwtori. 1994 Flie Ct ologn al uilsi qitenres of shari 1 n,ir.il on an earlier draft of the manuscript. P. flunwiddic, R. liiser, P. enemies. Ann Re'.. Itol Syst. 2. 1i)5.5)(j (uiJstcin, M. MdIo, C. Raith, 1. Sinimons, and V 'niith coiidticted Howe, W. Ii. 197. Tlit hittterflies of Not Ii .511(1 ii,, Donhk ili'. & ( ii , Ciii En (ii N \ the P)92 tick! citrvcvs of the six former island colonies. Special thanks Iftner, P. C.,j. A, 51mev .tndj. V. Calhoun. I ))2 Pullet lit 'itid sIilupe I ii, to S. Melvw and C. Raithel who sparked us :ill into action with their State Univ., i olittithits ,tkc-up tall in 1991. This work was supported by a contract grant lotte', F \I , and C P Kimhall, 1(14') Ihu I epitlopt('ra ot 'hititueket and \l,irl.'', (no. )2SMMI 1 to DLW and cooperative agreement (no. 11-1ô.(h)09.1575 \'iiiev,ird. N,oit itt ket \larta \Ittt lull \ssis, \.anuitt Ret 'ii to JSF') from the Massachusetts Division of Rshcrics and '"ik1Rfe, Kiots, A. It I I A I k Iii guide ti , the hi itt cub lies of Nt tub h A ire rica east ot tie great plains llotiglitotu Mitt liii, Post, in Labine P A Id(uk lie poptubaliot biologi if lie hunet II'. / ofu/u4u' i'. eu//ui VIII. Lirerature Cited Oviposition and its relation to patterns ci us iposutioul ii oH liuituerl Its I solution 22 .1 ndcrsi, R. M., and R. M. M.u 1Q80, Infectious isa' nd population ccles of forest itisee IS, 'n'ieflce 210' i)-'H--o() I Matloon, S (I , TI I) Davis, and 0. Spencer 10" 1. lieu ing te linqites It species of Spei'erki I Nt Iliphalidute). J. Lepid. Sin 2 247-2% Barton, TI )'), I'iekl studs 1 In' regal frItill:tri, S/n' vrii idali Report u I iepartment of tile Defense ,iiul Flie N.ittire ( tisrvain' I Iarr l'njrg P\. Maynard, C. j. 1 88h. Die butterflies of New rngland. lit .tdlie Whitteri, T1stot't, Nagel, H 0., T N igi I te tiga he and N I H ti ke rt I OP I Regal PIt) ft t' liii) It ti estiflaittiOn Barton, B. 199f. I he stabs il tIn, regal frttillarv. Report to Department of the Defense and The Nn ore .ons( us .t nii I Ian shii rg PA and natural liistu,r out Rowe Sant titan' Nebraska Prairie 'it 2 I iS-152, Bell, M. 1985. The face 1 1 onner ti iii people, g ologi', .ntd the land. State (Ieol. Nat, Opler, P. A. 108') ( ottntv atlas I eastern 1 'nited Stirs l'otti'rt lies 0 iI)I)82), 115. Fish lIst. "urv. Conn. Bull Witch, Set' WasI tin guilt I) I Boggs C. L., and Oplur, I' A. 15102. A field guide to eastern huiterb lies. hluuitghton Miff hit, ban C I. Ross. IOb Hie tkect of adult boil liinlttion on lift' history traits II ',/i'i' rjj t')r)Ilo,i,a I e nh pitra N' inphahid.u I. leok 'ai I: -4 41 Sch iklkne kt, ( I'. I 5186 Pastern regal tnt Ili r ( ç. cia/i,, i, \u'ws I ('pu) Si Sift I ii (ssn, It ( \\ Leahv, id I) Is \X'altoii. eds In R is. The huttertlii.s of Massachusetts I 'JKh. and other New England states \lass.n liusetts Audubon Soe,, Uncoin. Schv'eitzer, P. F. 15187. lEnient stewat'dslup abstract for S/iu'i'eu'ui u/iluu, Report to The Clark, A H. 19 2. Flie butterflies ot the District of columbia and vicinity. U.S. Nat!. Mus, N.ut.ure ('onservatn. v lIsten, \1 S Bull. 15" Schv'eitzer. P. F. 1)12 Sfc'tii'/a lila/ia the regal ft'ltill.o Result 1 ,t global status stlrves Report to 1 5 1 isP Wildlife Set's , n& or), NH Clark, A. H., and L. I I lirE 1db 1. The hutterl lies of Virginia. Smithson. Misc, Collect. vol. 116, no.". Schsvett,er, I 1 I' I itt)') Regal lOt i I aries tithe east A iii Ittit teui'l, Feb., p 9 Clark G. 1. 1914. A histot V ob I rinnueticui 0. P. Putnam's Sons, New York. Scott,j. A 10)/b 'l'lu. butterflies of North .Sniero .u Stanbord I dv Press, Stanford, CA. Dettioti, S. F. 1900 Moth'. nd kuttt rIles ii' the United States east of tile Rocky Moun. Scudder 5, It 1880 thu hitttrrfles I bitt astern I nited Situ's .ini ittauja with special lain" tlradllc Whidilen, Ii isti n i'efei'en e to 'ew I'itgland, S. II h tithE r, I ,nttht'de MA florithid, F. J. 19130. The hittiert Tt&s il Oregon. Timber Press, Forest Grove. ( >R. Swetig l.A. 1993. Regal fritillaries. mattE' royalty. Stit liuttert I. I tb., pp. 4-9 Edwards W Ti. l89. Dcs.'t iption 'it the preparatory stages of Argywils iiliilla Drury. Weed, C M, 1926. Butterflies worth knowing Duuibled.i'. 1.ig and (.o., Garden (.ity, 'an I.iit. 11: 21 21) NY. U. I)., and F. M. lli'own, 1981 ltinterflies of the Rocky Mountain slates Oklahoma 'wv. Press, Norman. (,la,s.shetg, J Butterflies titrougli binoculars. Oxford Univ. Press, Oxford, U.K. .DavidL. 1ligni'i': De,f)a1'tfl/r'/l! o,f Tco1o,t,'p (1/1(1 I'i'otii!fo;un'j' Bii1çi'. Hanintond P C, IdlI" he ilogleal l"tv& stigation on Viola a4unca. U.S. Forest Serv., t!'J-BO.X i3, I ti/i'e;city of Co;,nc'clicut, 5tort'.s', (1' (162(d) O i ,insl,rvi \jI!. Fotesi lot'. tills, lit. Hammond, P. C 198K. l.Lllogtl.al investigation on Viola adunca uS. Forest Serv., Sinslaw NatI, Forest, ( otyalli',, OR i11'atthew ,S' U 'allat'e: De/)(l;'t;funt qfLctnlogv rind .1, 'ofl(lu)nhir I' Flanintiind, P. C. 1989. 1991) 1 'tfi(i management plans for tile Oregon silverspot butterfly. Biology, UJiox 'ij, lini,'ers,ti' of Connect u ut, Storrs, CT 0(,2O)j043, Forest Serv,, Siuslaw NatI 1 orest, ( orvillis, OR Hammond, P C , and P. V ,\li'I. irl, It' 1)1 1991 tntroduction of the Oregon silverspot butterfly Speyer/e :e,i'ite hiipol1'ta) on Fairview Mountain. U.S. Forest Serv., George H. Boettner,' Department o/'Entomolo jemnalil flail, "iu',lai Nat!, Forest, I .ori'.tIljs. 1 l University of M'as.cacliusetts, Amherst, MA 01003 273 274