The taxonomy, larva and ecology of Agrotis buchholzi (Noctuidae) with a new sibling species from North Carolina

2004·4.8 MB·

by  Dale F Schweitzer

^VTHSOTW^
SEP 03
2004
Journal
of
T
HE Lepidopterists' OCIETY
Volume 58 2004 Number2
JounmloftheLepidopterists'Society
58(2),2004,65-74
THE TAXONOMY, LARVAAND ECOLOGYOFAGROTIS BUCHHOLZI (NOCTUIDAE) WITH A
NEW SIBLING SPECIES FROM NORTH CAROLINA
Dale F. Schweitzer
NatureServe&TheNatureConservency, 1761 MainStreet,PortMorris, NewJersey08349,USA
AND
Timothy L. McCabe
NewYorkStateMuseum,Albany,NewYork12230,USA
ABSTRACT.AgrotisbuchholziisoneoffourLepidopteraspeciesbelievedtobeendemictotheNewJerseyPineBarrens.Itoccursprimarily
inrecentlyburnedorexceptionallyxericorsterileareaswhereitssolelarvalfoodplant,Pyxidantherabarbulata(Diapensiaceae),occursinopen-
ingsintheshrublayer.Adultscanbequitecommonlocally.Therearetwobroodsapproximatelytwomonthsapartwiththefirststartingabout
lateMay. Hibernationis asprepupallarvaeinthesand.ThelarvaissimilartothatototherspeciesofAgrotis.Adultsareveryactiveandfeed
butdieirnaturalfoodsourcesarenotknown.Asiblingspecies,AgrotisCarolina,newspecies,iscloselyassociatedwithPbarbulatainsouth-
easternNorthCarolina. Itsrangeresemblesthatofanotherendemic,Hemipachnobiasubporphyrea. ThecombinedrangesofthesetwoAgro-
tisareverysimilartothoseofanunnamedCyclophora (Geometridae) andSpartiniphagacarterae Schweitzer(Noctuidae)andtheirhabitats
commonlyoverlapinbothstates. Fireisacrucial factorin formingandmaintaininghabitatforallofthesespecies.A. buchholzi maybecome
imperiledbecauseofadeclineofwildfires.A.Carolina,butnotA.buchholzi,isthrivinginhabitatsmaintainedbyprescribedburns.
Additionalkeywords: Sandplains,bivoltine,larvalhibernation,Agrotisobliqua,Agrotisstigmosa,monophagy.
Since its description very little has been published intensive collectingfrom 1969 to 1977. In 1977James
about the habitat and nothing about the life historyof Madenjian collected 46 specimens on die West Pine
Agrotis buchholzi (Barnes & Benjamin) (Noctuidae) Plains east ofChatsworth and a few about nine miles
andits foodplanthasnotbeen reported. Forbes (1954) to the west in Lebanon State Forest, both in Burling-
knewthis species onlyfrom the two types from Lake- ton County, during fieldwork forhis masters diesis at
hurst, NewJersey and indicated (correctly) that their Rutgers University, where a few specimens currendv
dates suggested two annual generations. Franclemont reside. Most ofhis specimens were discarded. During
(1956) reported the collection ofthe first substantial late May and early August 1980 DFS found die
series, andhis collectioncontained (as of1974) 34col- species to be comparably abundant at the West Pine
lected from 31 July to 15 August, 1955 and 19 col- Plainssite. Otherrecordsindie 1980s (all DFS and/or
lected 28 May to 10 June, 1956. Several collections W. Cromartie) were from a burn scar near Atsion,
J.
have a few specimens taken by Otto Buchholz, another nearPapoose Branch andan unbumedsitein
Fredrick Lemmer and Joseph Muller, all vicinity of the East Pine Plains, all in Burlington Countv, and at
Lakehurst, but only Franclemont had encountered it the Lakehurst Naval Air Station in Ocean CountvJ. In
in numbers. The late W. Cadbury III had a speci- 1994 DFS collected several on die approach zone of
J.
men from Browns Mills (now in Schweitzer coll.) AtlanticCityInternationalAirport, EggHarborTown-
which extended the range southwest into adjacent ship in northern Atlantic Counts; NewJersev, increas-
Burlington County. ing the known range to parts ofdiree counties. That
More recently one ofus (TLM) collected a speci- small population may originally have been a bit dis-
men nearAtsion, BurlingtonCounty, on25 May1976, junct and is quite isolated now. Another was taken
butDFS failedtoencounteritatnearbyBatsto during about 3 km to die soudi byThomas Hupfin 2000. ap-
66 Journalof the Lepidofterists' Society
parently a stray from the 1994 site as no intervening Foodplant and larval behavior. In 1980 DFS ob-
foodplant is known. In 1994, 1995 and 1996 DFS lo- tained eggs from two females, as had Franclemont
cated enormous populations in the Impact Area at previously. No suitable foodplant was found although
Fort Dix Military Reservation, in Ocean County, with some larvae fed on a species of Hud.sonia (Cistaceae)
47 being taken in two blackJight traps the night of12 but were promptly poisoned by it. In 198S we found
August 1994 alone. Lesser numbers were taken on all ofover 100 species of plants, includingasomewhat
several other nights in June and August. The known wiltedpieceofPyxidanthera andseveralgrasses, tobe
distribution thus encompasses parts ofthree contigu- unacceptable. By the end ofthe 198S field season it
ouscounties. Itcouldturn upinextremeeasternCam- was obvious to DFS that Agrotis buchholzi could be
den Countyafewkilometerswest ofAtsion. obtained consistently by placing light traps in large
Although the Pine Barrens region has always been concentrations of Pijxidanthera barhulata. Thus it
mapped by botanists to include a much larger area seemed obvious that either this plant or an abundant
(e.g., Stone 1911, Harshberger 1917, McCormiek associate, such as Leiophijllum buxifolium, would
1970), habitatforA. buchholzi andmostotherspecial- prove tobe the larval foodplant.
ized Pine Barrens Lepidopteraisverylimited southof On4August 1989, fiveworn females werecollected
the Mullica River, becoming more so because ofthe alivefromthree 15-wattblacklight setups onthe Lake-
elimination of large scale fires and development in hurst Naval Station by DFS and Thomas Breden. All
eastern Atlantic County. Contiguous large tracts of five were fed the next day andplaced in sealed plastic
true pine barrens habitats are confined to the region boxeswithfoliageoftheabovethreeplants.Allfemales
from about Lakehurst to Batsto. Pyxidanthera barbu- laidsomeeggs, twoofthem asubstantialnumber. Most
lata Michx., the foodplant reportedbelow, is scarce in eggswerelaid6to9August. Themajorityofeggswere
Cumberland County (e.g., Moore 1989) and infre- placed on the foliage orsteins ofthe Pyxidanthera but
quent, usually in small numbers, in most ofAtlantic some were placed on paper or on the Leiophyllum.
County, having disappeared from former stations Eggs hatched on about the seventh day after oviposi-
around Pomona since the 1970s probablybecause of tion andthe first fewlarvaewere found to feed readily
lackoffire (W. Cromartie pers. com. to DFS). on fresh Pijxidanthera, but not at all on Leiophijllum,
J.
Hudsonia orXerophyllum (Liliaceae). Herbaceous di-
Life HistoryofAgrotis Buchholzi
cotswerenottriedthistimebecausemostA. buchholzi
Phenology. Agrotis buchholzi has two broods with habitatsarecomposedalmostentirelyofwoodyspecies
adults present for about two to diree weeks each year and lichens, sometimeswith grasses orsedgespresent.
between 15 Mayand 15 June; and for a similarlyshort Cuttings ofPijxidanthera remained fresh for over
periodbetweenabout 17Julyand 15August. Adultsare twoweeks inplasticcontainers; larvaewereleftundis-
nocturnal and flyalmost exclusivelybefore midnight in turbedexcepttoaddnewfoliageandto makeobserva-
bothbroods. Theyhave notbeen observedatdusk. tions. One group of eggs was transferred to potted
Adultbehavior. Captiveadultsareextremelyactive Pijxidanthera before hatching and larvae allowed to
andalertevenwhendisturbedindiedaytime. Theyare feed on this foraboutamonth until the foodplantwas
alsoveryactiveatcollectingsheets. Theyapparentlydo severely defoliated by the third instars. Larvae were
not live morethan aweek. Adults are difficultto get in thereafter all placed in sealed plastic containers and
perfectcondition as theyquicklybecomebatteredand fedfreshcuttings as needed. Cuttingsweresometimes
worn, apparently from moving rapidly through fallen storedinarefrigeratorforuptotwoweeksbeforeuse.
pineneedles,cinders, twigs, deadleavesandashes.The The potted plant was covered with thin muslin, but
moths arebest collected atlights. thiswasprobablyanunnecessaryprecautionasthelar-
Adults do feed, and occasionallythe summerbrood vae showed notendencytoclimb.
adults come to sugar bait. Spring brood adults avidly Once feeding commenced, larvae were very slug-
nectar from Leiophijllum buxifolium Berg. (Ell.) blos- gish and usually remained hidden in accumulated
somsincaptivity, andwouldusuallyhaveeasyaccessto frass. The frass appeared to be quite sticky and be-
this and avariety ofotherflowering low shrubs in the came mixedwith sandgrains formingpilesadheringto
field. However, we have not found any on flowers at the foliage or stems ofthe foodplant. Bythe third in-
night. Summer brood adults would usually not have star,larvaeonfoliageactuallytouchingthesandtended
access to flowers but mightbe ableto locate rottingor tohideintheuppermostsandgrainswhilenotfeeding
damaged berries of Ericaceae. Adults readily accept andsome others often movedto the base ofdieplants
honey or maple syrup solutions and overripe fruit in orthe soil ratherthan remain amongtheirfrass. Later
confinement. instars became more subterranean and the last two in-
Volume 58, Number 2 67
stars emerged from the soil onlyto feed. Feedingwas
almost always at night except during cold weather in
Octoberwhen diumal feeding;was observed.
Although cuttings ofLeiophyllum and several Eri-
caceae were frequently added with the foodplant
when feeding the older larvae, these plants and the
grass Calamovilfa brevipilis were never eaten. On 13
September 1989 DFS collected a single third instar
larva on Pyoddanthera near Atsion. It is virtually cer-
tain that the larvae feed exclusivelyon this plant.
Reared A. buchholzi larvae reached full size out-
doors by earlyto mid October, but continued to feed
forafewmoreweeks. Firstbroodlarvaeobviouslyma-
turefastersincethesecondbroodadultsappearwithin
twomondisoffirstbroodadults. Allof65larvae (from
twofemales) countedon 15 Septemberreachedmatu-
rity. However, 10 died in late October or November,
probably from a pathogen. By 10 November, all sur-
vivors appeared to have completed feeding and most ©
were apparentlyin an earlypre-pupal condition. Most
hadformed cells in the moistsand, but sometimes ap-
peared briefly at the surface. The containers holding
thehibernatinglarvaewereplacedinathickstyrofoam
box, andburiedinpeatmossandleftoutsideonashel- FIGS. 1-4. Agrotis buchholzi larval head features. 1, Head,
teredporch. Decemberof1989provedtobethecold- frontal aspect (most head setae worn or broken off). 2, Ocellar
est Decemberon record in the region andwhen sea- oarrarlanfgaceem.enAltl,llaartevraaelrvieeawr.ed3,exHyopvoisphoanrvPnygxeiadlanctohmeprlaexf.ro4,mMNaanvdailblAei,r
sonable conditions returned at the end ofthe month StationnearLakehurst, NewJersey.
the peat was frozen and all but six of the 50 larvae
were dead. These six were removed to a refrigerator
until 8 March 1990. Three appeared at the surface Larval description ofAgrotis buchholzi. First instars were
later that month and formed new pupal cells at the notexaminedindetail.Theyhadnoobriouspatternandmovedin
the looping fashion typical for first instar Noctuidae. Second and
surface. All three pupated in April and eclosed in mid thirdinstarswere slightlyreddishwith littlepatternotherthanill-
May. Theydid not feed after November. defineddorsalandlateralpalelines.Virtuallyallpatternswerelost
Accordingto McCabe (1981)dierelatedA. stigmosa bythepenultimateinstarandtheolderlarvaewerearadieruniform
Morrison [asAgrotisvolubilis Harvey] alsodiapausesas blackishbrownmuchlikethecaterpillarofA.stigmosaillustratedbv
amature larvaby late summer and overwinters in that McCabe(19S1).Thereappearedtobesixinstars,butcloseobserva-
tionofyounglarvaewasdifficultandthenumberofinstarswasnot
state. He too had difficulty overwintering the larvae. establishedwithcertainty.
Crumb (1956) reported a prolonged spring and sum- Thediapausingprepupallarvaewerepreservedafterdievdied.
merlarvaldiapauseinA. venerabilisWalker, acommon This madeforlessthanpristinepreservedlarvaewithconsiderable
speciesin grassyplaceswith adults in earlyautumn. erosionandbreakageofsetae.Consequently,notallsetalbaseshad
The apparent monophagy of A. buchholzi is un- theirsetaeintactand,therefore,arenotrepresentedintheillustra-
tions.
usual, butnotunique, forthegroup ofgenera (includ- Last(sixth?)instarlarva. General:Theaverageheadwidth(as
ingEuxoa) combinedasAgrotisbyForbes (1954). The viewed from the face) 2.1 mm; average totalbodvlengdi (headto
undescribed Mesembragrotis illustrated by Handfield tail,naturalrestingposture)25.1 mm;abdominalprolegspresenton
r(i1p9e9n9i)nigs msoeendosphoafgothues ignraNssewCaJlearmsoevyiolfnaflborweevirpsilainsd. o3rrddintahlrmoeusgohse6rtihes;ansdpir1a0ctlheAsSeg0m.e3n3tms;msheitgaheosnimapvleer;agcero(cnh=et1s0)a.Cuonli--
oration (living material): Headwith abrownpavementpatternto-
Richiaacclivis (Morrison),whichis similaringenitalia, wards front of face only. Bodv brown-black with a conspicuous
adult and larval appearance, phenology and behavior, lateral series ofblackspots, twoon each segment, comprisingthe
is alsoaspecialiston flowers andseeds ofagrass, Pan- spiracleandtheLIpinaculum.Headcapsule(Fig. 1):Epicranialsu-
icumvirgatumL. atleastinNewJersey(DFS). Ourlar- ture0.27mmlong;heightofIrons0.97mm;secondadfrontal(Af2)
vae ofTricliosilia manifesto (Morrison) accepted only anteriorand 1stadfrontal (Afl)posteriortoapexoffrons; coronal
puncturesAa,pa,pb,and3ultraposterior(2risibleinfrontalaspect!
oaksbutapparentlybecomemorepolyphagousafterhi- presentasillustrated. Hypopharyngealcomplex(Fig.3):spinneret
bernation (Crumb, 1956),whenoakis unavailable. shorterthanlabia]palpus,apexlackingsetae;stipularseta(S)atan-
68 Journalofthe Lepidopterists' Society
Relationships. Agrotis buchholzi fits in a group
that includes A. obliqua B. Smith)(Fig. 10). These
(J.
moths are allverysimilar in appearance and male and
female genitalia, although the latter typically has the
orbiculardarkthroughout, unlikethe lighter ringseen
inA. buchholzi. The food plant ofA. obliqua has not
been established, but it occurs in the absence ofDia-
pensiaceae. The polyphagousA. volubilis, A. stigmosa
and A. venerabilis are also morphologically close to
these three. Agrotis buchholzi also has a sibling
species, previouslytreated in collections as asouthern
disjunct, whichwe considerdistinct.
Agrotis Carolina Schweitzer & McCabe,
new species
(Figs. 7-9, 14, 16, 18, 19)
Adults:Maleandfemalesimilar(Figs.7,8).Vertexofheadared-
dishbrown; frons andtipsofpalpi lightbrown;antennaesubpecti-
nate; tegulae with a black band; patagia with silvery gray scales,
lighterthandisk;abdomenuntufted;wingsvioletgraywithamline
wavy,bowedoutconsiderablybelowfold,withsubtlepinkishcastto
basalareain fold; orbicularround, reniform kidneyshaped, region
betweenorbicularandreniformandcostadarker; medianlineusu-
allypresentthough sometimes lacking;pm linedentate,evenlyex-
curved;terminallineabrokenseriesofblackspots;fringeconcolor-
ouswithground;wingexpanse (springbrood) =32-38mm(n=55,
Figs. 5-6. Agrotisbuchholzi larvalbodysegments. 5, Protho- average =34mm),secondbroodwingexpanse=26-30mm(n = 9,
rax, semidiagrammatic (most body setae worn or broken off). 6, average=29mm).
tFihresrtaafbrdoommitnhealNasveaglmeAnitr.SAtlalticoantenrepialrlaLraskerheuarrsetd,exNeovwisJoernsePyy.xidem- blunMtallyepogienntietdal(Fiiag.(F1i6g)s..H1a4r.p1e6,pr1e8s)e:ntV,allveensgtnheaarbloyutpahraallflewli-dstidhedo,f
valve, bluntly rounded and directed slightly towards outside. Cu-
terodorsalregionofprementum;distalregionofhypopharynxbears cullarspineslooselyspaced,with roomenough foradditionalspine
numerousveryfinespines, followedbyasingle rowof14-15large between manybases (Fig. 14). Evertedvesicawith chitinizedbasal
spines that extend to the posteriorapex ofprementum. Mandible band(Fig. 18).Remainderofvesicaanelongatetube,slightlycurved,
(Fig. 4)withinnerridges indistinct, lackingbasaltooth. Prothorax andswollentowardsend.
(Fig. 5): cervical shield well sclerotized; subdorsal setae (SD1 & Female genitalia (Fig. 19): Ovipositorlobes short, unmodified.
SD2)ontheshield;prespiracularsetae(LI&L2)andsubventralse- Anteriorapophysesextendhalfwaytobranchofappendixbursa.Duc-
taelackingpinaculi. Firstabdominalsegments (Fig. 6): subventral tus bursanearlytwice length ofanteriorapophyses. Corpusbursaa
setae (SV1 & SV3) approximate; LI posteriortospiracle; SD2 an- long tube with swollen apex. Appendix bursa much longer, narrow,
terodorsaltospiracle;spiracle0.31 mmhigh.Ab2-6with3subven- coiledonceuponitself,withgraduallyswollenapex. Ductusseminalis
tral setae; SD2 setal base anteriorto spiracle. Ab7&8 with only 1 issuesfromapexofappendixbursa.Thelengthyappendixbursacoils
setainsubventralgroup.Tarsalclawsarewithoutabasaltooth.The inwardlyoveritself.Thisisevidentwhendiebursaisinflatedinalco-
hind coxae are approximately the femur length apart. Crochets a holandwassketchedatthisstageindiedissection.Theappendixbursa
uniordinalmesoseries.Actualcrochetcountswerenotpracticalbe- hadtobeaccurately"arranged"indiethickermountingmedium.
cause ofthe state ofpreservation. Material examined. Ten speci- Holotype: Male, North Carolina, Brunswick Co., vie. Supply.
mens, Lakehurst Naval Air Station, Ocean County NewJersey, 6 Green Swamp Preserve, drysavanna, 12 April 1991, S. Hall & D.
December1989,fromovaoffemalecollectedanddeterminedbyD. Schweitzer. —
F. Schweitzer. Paratypes (68 allNorthCarolina,approximatelya3:1maleto
Agrotisbuchholzi larvae differ fromA. stigmosa (see McCabe female ratio): (6) Brunswick Co., vie. Supply, Green Swamp Pre-
1981) as follows: headcoronalpuncture (pb)outoflinewith setae serve, dry Savannah, 11 June 1991, 12 April 1991, S. Hall & D.
P2andPI;the LI pinaeulus, onthe firstabdominalsegment, less Schweitzer; McCabeslides2567, 2663,2664, 2879; (3) Brunswick
thanhalfitswidthfromthepinaeulusL2(thisdistanceistwiceLi's County, 2 mi. SWBoilingSprings Lakes,28June 1994,J. B. Sulli-
width inA. stigmosa) andthe SV3pinaeuluswell formedandnear van,R. Broadwell,&B.Smith;(1)BrunswickCounty,MOTSU,In-
toSV1. tersection offirebreaks 32 & 34, 11 April 1994, J. B. Sullivan, R.
MultiplerearingsofA. venerabilishaveshownvariabilityinthe Broadwell. & B. Smith; (23) Carteret County, Millis Savannah,
sizeoftheLIpinaculum.IfthisholdsforA.buchholziitcouldaffect 34.45.19N-76.59.23W,29 March 1998, T McCabe, McCabeslide
the apparent distance between it and the L2 pinaeulus. If run 3721;(18)OnslowCounty,CampLejeune,4April1995,S. Hall;(3)
throughCrumb'skeys.A, buchholzi appearsclosesttoA. gladiaria, PenderCounty, 3.2 mi. N. Maple Hill, AngolaCreek Preserve, 13
butCrumb(1956)didnothavetherelatedA. obliqua asitslarvais April1991,S.Hall&D.Schweitzer,McCabeslide2568;(8)Pender
unknown. County,HollyShelter,OldRd./BJ Rd„5April 1995,S. Hall.
Volume 58, Number 2 69
13
Figs.7—14. Adults. 7,AgrotisCarolina, firstbroodmale, Holotype,NorthCarolina(BrunswickCo.,vie. Supply, GreenSwampPreserve,
drySavannah, 12April1991,S. Hall&D. Schweitzer).8,AgrotisCarolina, female.NorthCarolina(PenderCount}',3.2mi.N.MapleHill.An-
golaCreekPreserve, 13April 1991, S. Hall& D. Schweitzer).9,AgrotisCarolina, secondbrood. NorthCarolina(BrunswickCo.,vie. Supply
Green Swamp Preserve, dry Savannah, 11 June 1991, S. Hall & D. Schweitzer). 10, Agrotis obliqua, female, Labrador (Lower Brook,
5S5c.h1w5ei.t5z2e-r6)0..521.22,9A,gr1o5tJiuslybu1c9h9h3o,lzJ.i,Bofpepm)a.le1.1,NAegwrotJleirsbeuych(hOoclezai,nmCaoluen,tyN,ewLaJkeerhsueryst(ONc.eAa.nS.C,ouenxtoyv,isF,orrteaDriexdilmatpeacAtprairle,a,eel1.22AuJguunset11999904.. DD..
Schweitzer). FIGS. 13-14. Cucullarspines. 13,Agrotisbuchholzi(samedataasFig. 15). 14,AgrotisCarolina(samedataasFig.16).
Deposition oftype material. Holotype male is deposited in buchholzi (34mmvs. 27mm forfirstbrood,29mmvs. 27mm for
the US National Museum of Natural History, Washington, D.C. thesecondbrood). Sizeanddistributionwillalloweasydetermina-
Paratypes are in that collection, the NewYork State Museum, the tion, althoughalmostalladultsarepalerandmuch morestrikingly
Schweitzercollection,the Sullivancollectionandthe McCabecol- marked thanA. buchholzi orA. obliqua.Agrotisobliqua (Fig. 10)
lection. occurs in boreal habitats and typically has the orbicular dark
Diagnosis. Agrotis Carolina resembles a large version ofA. throughout.ThemorphologicaldifferencesbetweenA.Carolinaand
70 Journalofthe Lepidoftemsts' Society
FlGS. 17-18. Aedoeagus. 17, Everted aedoeagus of Agrotis
buchholzi (samedataas Fig. 15). 18, EvertedaedoeagusofAgrotis
Carolina(samedataasFig. 16).
fifththevolumeofthebursa.Veryoldandwornspecimenspossess
particularly fragile appendixbursae and made forpoorand nearly
worthlessslidemounts.
Figs. 15-16. Male genitalia. 15, Valves ofAgrotis buchholzi,
McCabeslide2569, NewJersey(OceanCounty, LakehurstN.A.S., Distribution. We believe thatAgrotis Carolina will
4August 19S9, D. Schweitzer. 16,ValvesofAgrotisCarolina, Mc- provetobeconfinedtoeastern North Carolinaorper-
Cabeslide2663,NorthCarolina(BrunswickCounty,GreenSwamp
Preserve, 11June 1991,S. Hall). haps barelyinto South Carolinawith its apparent food-
plant (Radford et al. 1964). We have seen it from
Brunswick, Pender, Onslow and Carteret Counties in
A.buchholziincludegenitalicfeaturesaswellashabitus.Malegen- theOuterCoastalPlain; Stephen Hall(pers. com. 2001)
italia:ThebasesofthecucullarspinesinA. buchholzi (Fig. 13)are also has taken it in Hoke and Haniett Counties in the
mostlycontiguous,thoseofA. Carolina(Fig. 14)aretypicallywidely fall-line Sand Hills region ofthe innercoastal plain.
osfpatcheedstpoatcehse.exAtebnutlibtowuosulpdrobteubpeosrsainbclee,tonefaitrantohethbearsaslpienendinomfatnhye Populations ofA. Carolina in the outercoastal Plain
vesica, is opposite (180°) the basal comutus inA. Carolina and at areassociatedwithPi/xiclantlwrabarbnlata var.barbu-
more ofaright angle (270°) tothebasalcomutusinA. buccholzi. lata,whichisthefoodplantofA. buchholzi in NewJer-
Agrotisbuchholzi alsohas an indentation (a"valley") between the sey. Specimens collected in the sand hills were associ-
baseofdievesicaandtheoriginoftheelongatedistalportionofthe atedwiththe endemicvarietybrevifolia,whichdiffers
v(essoicmae.fTehaitsuriensdecnatnatniootnbiesvniotewseedenifitnhCeavreosliicnaa.isIanc'cpiedrefnetcatl'leyvpeiresricoensd in both its much more xeric sand ridge habitat and in
orseveredshortofthefinalnarrowingresultinginapoorlyinflated morphology. NeitherAgrotisbuchholzi norA. Carolina
vesica)themostdistalportionofdievesicadisplaysaswellingbefore occurs in thetinypinebarren remnantnearZuni,Vir-
diefinaltapering(seeFig. 17).Theapexofthisswellingdoesnotre- ginia that harbors the only significant population of
curveevenasfarasthebulbousvesicalbaseinA. Carolina,butre- Pyxiclanthera between New Jersey and North Car-
bcuucrhvheoslztio.aUnndfotrhtruonuagtheltyhethilsenlagstthfoefattuhreehaaesdoteoabgeusviperwoepderduirniAn.g olina. This conclusion is based on repeated collecting
dissectionasthetrueorientationisdistortedbymountinginthevis- efforts byboth authors and theVirginia Natural Her-
cousmountingmedium.Femalegenitalia:A.buchholziispropor- itage Program.
tionatelysmallerthanA. Carolina(photosareatthesamemagnifica- BiologyofAgrotis Carolina.Whilediisspecieshas
tion); appendixbursaswollen abruptlyat tip inA. buchholzi (Fig. notbeenreareditisvirtuallycertaindiatitfeedsondie
2u0m)ethdaitstoifncAt.ioCnarasolwienlal.gTrahdeuaslwloylleennlatrigpeodf(tFhieg.ap1p9)e.ndTihxerbuerissaaovfoAl.- same plant asA. buchholzi, namelyPyxidanthera bar-
Carolina is approximately one third the volume ofthe bursa. The bnlata including variety brevifolia. As with A. buch-
swollen tip ofthe appendix bursaofA. buchholzi is less thanone- holzi, A. Carolina is taken virtually at will by placing
Volume 58, Number 2 71
pine lowlands. See McCormick (1970) or Forman
(1979) forageneral description ofthese communities.
Large numbers can alsobe found aroundthe edgesof
and to some extent in reedgrass (Calamovilfa brevip-
ilis) savannas. Habitats forA. Carolina include mesic
wiregrass savannas, pine flatwoods and other fre-
f V quently burned open pinelands. All normal habitats
forboth species can be characterized as fire disclimax
coastal plain pinelands although the Atlantic CityAir-
port habitat is maintainedbyannualwintercutting.
Fire frequencies vary from essentiallv aprescribed
burn everywinter at some North Carolina sites (e.g.,
formerly at Green Swamp) to only two or three wild
fires per century in a few NewJersey sites. As agen-
eral rule one or more fires per decade will usuallvbe
20 required to maintain most high qualitvhabitats in op-
timum condition foreither species. AndrewWindisch
(pers. com. to DFS 2001) suggests aregime ofaverage
Figs. 19-20. Female genitalia. 19,Agrotis Carolina, McCabe to moderately severe wildfires at intervals of 10-30
slide2568, NorthCarolina, PenderCounty,3.2mi. W. Maple Hill, years probablyallows both Pyxidanthera andA. buch-
AngolaCreekPreserve, 13April 1991. 20,Agrotisbuchholzi, Mc-
Cabe slide 2570, NewJersey, Burlington Count)', Atsion, 16 May holzi topersistindefinitelyinthecore NewJerseyPine
1985,D.Schweitzer. Barrens landscape (see alsoWindisch 1999). Pyxidan-
thera appeared to be still increasing on the Atsion
lighttrapsatpatchesofthatplant. UnlikeA. buchholzi, burn scar in late 1989, oversixyears afterthe intense
weare notaware ofstraystaken awayfrom thatplant. July 1983 fire, andhadnotdeclinednoticeablybySep-
Given the longer growing season in southeastern tember 1991. By Julv 2001 it was still common in the
NorthCarolinaonewouldexpectthreebroodsofA. Car- more open areas but greatly reduced overall byheavy
olina with the first two about a month earlier dian the pine regeneration (DFSpers. obs.). Observations from
correspondingbroodsofA. buchholziin NewJersey. In- Fort Dix suggest A. buchholzi numbers peak about
deedthere arebroodspeakingin April andJune andan one to five years after fires. Prescribed burning, as
apparentlymoreprotractedthirdbroodwithdates from presently practiced in New Jersey, does not create
1 August to 14 September (Stephen Hall pers. com. suitable habitat for A. buchholzi since tire sites in-
2001). The first brood starts in late March, abit earlier volved alreadyhave a dense canopywhich these light
than might be expected considering the phenology of winterfires do not reduce.
thetworegions.Theflightseasonofthespringbroodof Therearenodirectdataon the immediate effects of
A. Carolina overlaps substantiallywiththe floweringpe- firesoneitherA. Carolina orA. buchholzibutbothhave
riod for Pyxidanthera whileA. buchholzi flies after die been collectedinnumbers inspringfollowingdormant
foodplant has flowered. We thus suspect that adult A. season fires indicatingobvious in situ survival aswould
Carolina obtainnectarfrom thelarvalhostplant. beexpectedforundergroundlarvae. While fewinsects
from a centimeter below the soil surface to the tree
EcologyofAgrotis Buchholzi,
tops would be likelyto survive in duffto crown infer-
A. Carolina and Foodplant nos in New Jersey wildfires, direct mortalitv ofover-
Pyxidanthera barbulata is a prostrate mat forming winteringlarvaeofeitherAgrotisshouldbevervlowin
plantofthelargelyarctic-alpinefamilyDiapensiaceae, most prescribed burns and in less intense wildfires.
with showy flowers appearing in March or April. It However, captiveA. buchholzi larvaemovedtodiesoil
grows ingaps inthe shrub layerinopen pinelands and surface to pupate bylate March (A. Carolina probably
flourishes for several years after fires on sterile white does soin February),which couldleavediemvulnera-
sands, such as the Atsion soil series in New Jersey. ble tointensewildfires duringthe springfire season.
Such soils are generallyverydryatthe surface,butof- In addition to direct mortalitv- larvae would starve
ten,probablynearlyalwaysin North Carolina, theyare and/or females be forcedto emigrate iffires killeddie
moistjustbeneath. Pyxidanthera foliage. Top kill of diis plant was ob-
Typical habitats forA. buchholzi are usually dwarf served in March followingwinterbums in atleastone
pine plains (Forman 1979) and recently burned pitch North Carolina site (DFS pers. obs.), but not at some
72 Journalofthe Lepidopterists' Society
others. Top kill is common in New Jersey fires. Most Lepidopteraspecies. Atotal ofeight species are found
patches resprout after light fires, and light fires may only in the pinelands ofsou—thern New Jersey and/or
well stimulategermination. Summerfires,whichoccur southeastern North Carolina thecombined rangesof
inveryfrequentlyburned parts ofFort Dix, mayburn Agrotis buchholzi andA. Carolina.
slowlyalongdie ground surface orthrough the shrubs Apartiallistofotheruncommon to rare speciesthat
and consume only the outer parts of the apparently sometimes co-occur with A. buchholzi includes the
succulent Pyxidanthera mats, leaving the centers in- now imperiled Atrytone arogos arogos (Boisduval &
tact. Larvaeindiesepatcheswouldhavegoodsurvival. LeConte), Hesperia attains slossonae (Skinner), Hi/-
Such fires are not nowtypical elsewhere in the range pomecis buchholzaria (Lemmer), Heterocampa varia
ofA. buchholzi, in part because of accumulation of (Walker), Catocalaherodiasgerhardi Barnesand Ben-
more fuel between infrequent fires. They may be jamin, Catocalajair Strecker,andthe previouslymen-
more typicalinA. Carolina habitats and mayalso have tioned unnamedMesembagrotis species.
been more typical in pre-settlement NewJersey. Un-
bumed refugiashouldalwaysbe leftin anyprescribed Conservation Status
burns and are clearlyneededwith some ofthe hotter
NewJerseywildfires. Agrotis buchholzi was under review on the Candi-
Associated species. Both of these Agrotis occur date (C2) List ofthe United States Fish and Wildlife
with other rare and extremely range-restricted Lepi- Service under the United States Endangered Species
doptera.Agrotisbuchholzi usuallyoccurswidisomeor Actaspublishedinthe Federal Registeron 15 Novem-
all ofthe other the Pine Barrens endemics and near ber 1994. Sincedie rangewas given as NewJerseyand
endemics. Crambus daeckellus Haimbach (Pyralidae) North Carolina, this listing; included Agrotis Carolina.
is easilyfoundin its habitats on Ft. Dix and Lakehurst The C2 listwas abandoned forpolitical reasons in the
Naval Air Station within five meters ofpatches ofXe- mid-1990*s andnodecisionwas madeonthesespecies.
rophyllum asplwcleliodes (L.) Nutt. (Liliaceae) (DFS At present neitherAgrotis buchholzi orA. Carolina
pers. obs.) but has notbeentaken elsewhere since be- are in immediate danger ofextinction but given pre-
fore 1960. Two still unnamed Crambidia (Arctiidae, sent trends offire suppression in NewJerseyandout-
Lithosiinae) species described and keyed by Fran- righthabitatloss andfragmentationin NorthCarolina,
clemont in Forbes (1960) bring the list of apparent and dependence on fortuitous management practices
Pine Barrens endemic moths to four species. Both in both states, neither species can be considered se-
Crambidia occur at some A. buchholzi sites. One of cure. Eidier could become threatened in the foresee-
the Crambidia is common and widespread in the re- able future. Very little, ifany, non-militaryland is ap-
gion, while the other may prefer habitats similar to propriately managed for A. buchholzi and, in the
those usedbyA. buchholzi. absence of appropriate management, the fire trends
Spartiniphaga carterae Schweitzer (Noctuidae), discussed belowthreaten its long-term survival. Both
which is a borer in the grass Calamovilfa brevipilis species have extraordinarily small ranges for moths,
(Torr.) Scribn., and an unnamed geometric! consis- each being known from only from a single state and
tentlymisidentified (e.g., Forbes 194S) as Cyclophora threeandsixcountiesrespectively.Agrotisbuchholziis
or Cosymbia culicaria (Guenee)1, which apparently still locally common in the northern halfofthe New
feeds mostly on Leiophyllum, have ranges similar to Jersey Pine Barrens in Burlington and Ocean Coun-
the combined range ofAgrotis buchholzi and A. Car- ties. It does not appear to occupy isolated, largely
olinaandareoftenmicrosympatricwith them.Agrotis roadside, patches of the foodplant in Monmouth
buchholzi was collected in the same light trap sample Count)' or around Batsto in southern Burlington
as a paratype of Abagrotis cryptica LaFontaine Countv.
J
(knownfromonlytwo specimensfrom the NewJersey A very substantial portion ofthe total range ofA.
Pine Barrens and one from Michigan). The now se- buchholzi is on lands owned bythe State ofNewJer-
verelyimperiledvenus flytrap cutworm,Hemipachno- seyanddie United States Military Sendees. Nearlyall
bia subporphyrea (Walker), is essentially endemic to ofdie rangeofA. buchholzi falls underthe jurisdiction
the eastern part ofdie range ofA. Carolina. With four ofthe NewJerseyPinelands Commission, which con-
and two respectively New Jersey and North Carolina veys somelevelofhabitatprotectionbutnoassurances
appearto lead eastern US states in terms ofendemic ofmanagement. The Military, The US Forest Service,
die state ofNorth Carolina, and The Nature Conser-
1BasedonexaminationofthetypesintheUnitedStatesNational vancyown substantial habitats forA. Carolina. Mostof
MuseumbyDFS. theoriginalhabitatforA. Carolina has beendeveloped
Volume 58, Number 2 73
or converted to agriculture or pine plantations while verypossibleinthenextfewdecadeswithoutappropri-
most ofthe range ofA. buchholzi still supports more ate management practices or an increase in moderate
orless natural, but increasingly fire-suppressed, vege- intensity wildfires. Since wildfires occasionally affect
tation. thousands ofhectares (maximum single fire recorded
Bvfarthe largestpopulations (basedon numbers in about38,000hectaresinApril-May1963),A. buchholzi
trap samples and extent of optimum habitat) of A. couldbecomequiteimperiledbyanymajorrangecon-
buchholzi are on Fort Dix and are maintained byfires traction or fragmentation which would leave it at risk
from militaryexercises widi someprescribedburning. ofhavingasingle hot fireburn all ofits habitats simul-
Recentfireintervalshave rangedfrom lessthanayear taneously.
tothreeyears in much ofthis habitat, with afewparts A Carolina is morewidespread (atleastsixcounties in
unbumed for five years or longer. Any major curtail- twoseparateareas)butitsmajorhabitatsarewidifnama-
mentorcessationofmilitaryactivitieson Ft. Dixwould trix ofa much more intensively developed and agricul-
be aserious potential threattoA. buchholzi, although tural (including pine farming) landscapes and die food-
moderate relaxation offire frequency (perhaps to two plant is much more localized than in New Jersey. P.
orthreeperdecade)wouldprobablybenefitsomerare barbulatavarietybarbulata iscurrentlyconsideredasun-
associates, especiallyAtnjtonearogosarogos. common (S3) by North Carolina Natural Heritage Pro-
Most ofthe range ofAgrotis buchholzi is relatively gram and die endemic Sand Hills variety brevifolia is
undeveloped, with vast tracts of forest, woodland, consideredEndangered(SI). Atpresentallmajorhabi-
scrub and substantial berry farming. Wildfires are re- tats with the foodplant that have been checked still
sponsible forthe existence ofmost remaininghabitats havethe moth. However, itwas not found at two small
for A. buchholzi and several other specialized Pine previouslyfire-suppressed remnant Pyxiclanthera sites
Barrens Lepidoptera. Givnish (1981), Forman and in Weymouth Woods State Natural Area in Moore
Boemer (1981), Buchholz and Zampella (1987) and County, within 20 kilometers ofthe Ft. Braggpopula-
Windisch (1999) all documentdrat acreageburnedby tion (Stephen Hall pers. com. May2001). Large tracts
wildfires has declined in recent decades, and Harsh- probablysupporting metapopulations on thousands of
berger (1917) describes a fire swept, cut over, and hectares of habitat exist, including in the Croatan
much less forested regionthanis presentnow. NationalForest, somemilitarylandssuchas Camp Le-
The location intheWest Pine Plains where Maden- jeune, Fort Bragg, and MOTSU, Green Swamp Pre-
jianand SchweitzercollectedA. buchholzi in 1977and serve, and Holly Shelter State Gamelands, but most or
1980 has since closedin noticeably. Parts ofthe Plains allareprobablyfunctionallyisolated from odiers.
have now not burned for 50-70 years instead ofthe Agrotis Carolina and its habitats are being perpetu-
optimum 5—15yearinterval, resultinginclosedwoody ated by current prescribed burning practices, usuallv
strataofshrubbypines, oaks andheaths, andamarked dormant season burns, on actively managed state and
decline in Pyxiclanthera and subshrubs such as private conservation lands as well as bycombinedacci-
Corema conradii Torr. and Arctostaphylos uva-ursi dentalandprescribedfireson militarylands.Treecover
(L.), forbs, and native grasses (Windisch 1999). When in these habitats tends to be naturallysparse. Perhaps
fires finally do occur after prolonged (>30-40 years) selective thinning combined with prescribed bums
exclusiontheytendtobeextremelysevereandmayre- would restoreA. buchholzi habitats in NewJersey.
duce or eliminate Pyxiclanthera plants and seedbanks We recommend that managing agencies for lands
(Windisch 1999). The increasedtime between and in- harboringAgrotis buchholzi orA. Carolina cooperate
creased severity of fires are leading to a loss of this withfutureinvestigators,includingamateurcollectors,
plant and its seeds in some areas ofdie Pine Barrens, to the extent practical. Both species generallvoceupv
eveninthecore,aswellasinperipheralareas (Windisch habitats ofhundreds to thousands ofhectares and are
1999). Nevertheless, intervening landscapes between among the more numerous Noctuidae in them andso
majorA. buchholzi habitats stilltypicallydohavesome are obviously not at risk from ordinarv collecting
Pyxiclanthera at least along sand roads and in distur- methods. It is specificallv recommended that where
bances, andstill experienceoccasionalwildfiresonthe collectingpermits are required on certain stateorfed-
scale ofafewhundredto afewthousandhectares. eral lands, they be made readilv available with mini-
While metapopulation dynamics are almost cer- mum restrictions other than reporting requirements.
tainlydeteriorating, femaleshavebeen collectedabout Reports from collectors would pro\ide useful updates
one and three kilometers out ofhabitat and so do dis- on the status ofA. buchholzi and A. Carolina and re-
perse. Metapopulations probablystill function except peated failures to find diese easily sampled modis
in eastern Atlantic County. Metapopulation collapse is wouldbeagoodindicatorofdeclines. Theonlyknown
74 Journalofthe Lepidopterists' Society
population of either species that might be small Fohbes,W.T. M. 1954. LepidopteraofNewY'orkandneighboring
enough to be vulnerable to occasional blacklight trap- states. Paret3, Noctuidae. Mem. Cornell Univ. Agric. Exp. St.
329:1^33.
ping is the isolated one nearAtlantic CityAirport and FRANCLEMONT, G. 1956.Taxonomicandbiologicalstudiesofthe
J.
current inventory efforts there (by DFS) aimed at NoctuoidandGeometroidmothsofthe NewJerseyPine Bar-
other taxa avoid its limited habitat during the flight rens, pp. 206-207. In 1956 yearbook ofthe American Philo-
sophicalSociety. ReportofCommitteeon Research.
season. Forman, R. T. T. (ed.). 1979. Pine Barrens: ecosystem and land-
scape.AcademicPress, NewYork.601 pp.
Acknowledgments Forman, R.T.T. & R. E. Boerner. 1981. Firefrequencyandthe
Pine Barrens of New Jersey. Bull. Torrev Botanical Club
MuchofSchweitzer's fieldworkwasconductedforThe Nature 108:34-50.
oCfonDseefrevnasnecytoutnhdeerNecowntJrearcstesyfNraotmurtahleHUenriitteadgeStPartoegsraDme.paSrutpmpeonrtt GlVNIrSeHns,oTf.NJ.e1w98J1e.rsSeey.roEtvinoyl,utgieoongr3a5p:h1y0,1-a1n2d3.fireonthe Pine Bar-
forthe 1994samplingat AtlanticCityAirportandforpreparation HANDFIELD, L. 1999. Le Guide des Papillons du Quebec. Version
ofthis manuscriptcame fromthe United States FishandWildlife Scientifique. Broquet.982pp.
Service, Pleasantville, NewJerseyoffice. Schweitzer's 1980 field- Harshberger, W. 1916.ThevegetationoftheNewJerseyPine-
workwassupportedbyagrantfromtheXercesSociety.Fieldassis- Barrens, anJ.ecological investigation. Christopher Souer Co.,
tanceofThomas Breden, Stephen Hall, WilliamJ. Cromartie,An- Philadelphia.329pp.
nette Scherer, and Thomas Hupt was greatly appreciated. North McCabe, T L. 1981. The larva ofAgrotisvolubilis Harvey (Lepi-
Carolinacollectinglocalitieswerepickedonthebasis ofinforma- doptera: Noctuidae). NewYorkEntomol.Soc.89(2):59-64.
otinonthferobamsitsheofNcoorntvherCsaartoiloinnsawiNtahtuPrralogHrearmitsatagfef.PWreogrthaamnkfiJleossaenphd McCovrenmtiocryk.,RJe.s1e9a7r0c.hThReepJ.Poirnte2B,aNrreewns:JearsperyelSitmaitnearMyuesceoulomg,icTarleinn--
Boppofthe NewYorkState Museum forthespecimen ofAgrotis ton. 100pp.
obliqua. Thanks to Patricia Kernan, staffartist for the New York MOORE, G. 1989. Checklist ofthe vascular flora ofCumberland
State Museum, for her illustrations ofthe larva. McCabe's 2003 County, NewJersey. Bartonia.55:25-39.
fieldwork, conducted injackpine barrens in NewYork, was sup- Radford,A. E., H. E.Ahles, &R. C. Bell, 1964. Manualofthe
portedbyagrant from the Biodiversity Research InstituteofNew vascularfloraofthe Carolinas. Univ. ofNorth Carolina Press,
YorkState.SpecialthanksareextendedtoRogerMeyersandRoger ChapelHill, 1183pp.
WSimnidtihsocfhFtp.roDviixdefodrmthuecirhaisnssiisgthatncientdourtihneghfaibeitladtwodrykntahmeirec.sAinndNreeww SMITHN,eJw.JBe.rs1e9y10S.taTtheeMiunsseecutsmofforNe19w09J.erPspe.y.14A-n8n8u8a.lreportofthe
Jerseyandprovided manyusefulcomments duringpreparationof STONE,W. 1911.TheplantsofsouthernNewJerseywithespecialref-
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JerseyStateMuseumfor1910.PartII.Pp.21-828, 129plates.
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