Journal of the Lepidopterists' Society 57(2),2003, 107-112

THE LARVA AND PUPA OF PERMAGNARIA PACK. (GEOMETRIDAE)

DAVID L. WAGNER DepaJiment of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269, US A. E-mail: [email protected]

ERIC W HOSSLER Depariment of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269, USA, and Department of Anatomy and Cell Biology, J.I-l. Quillen College of Medicine, East State UniverSity, Johnson City, Tennessee 37614, USA

AND

FRED E. HOSSLER Department of Anatomy and Cell Biology, J.H. Quillen College of Medicine, East Tennessee State University, Johnson City, Tennessee 37614, USA

ABSTRACT, LaJvae of were reared to maturity on red (Quercus ntbra). The larva and pupa of this rare eastern geometrid are described and illustrated. Diagnoses and photographic images of late instar huvae are provided for three members of the genus: Lytmsis permagnrl'ria, L. sinuosa, and L. l.tnitaria. Additional key words: , , , twig mimicry.

Lytrosis perrnagnaria (Pack.) has been regarded as in a solution of sugar and water, began laying pale one of the rarest of eastern macrolepidopterans. At the green eggs aft er two days in captivity. First ins tars time Forbes (1948) completed his work on the wander actively, often covering large distances, before 'Geometridae of and Neighboring States,' settling to feed. Captive 1st instar larvae accepted red the species was known only from the holotype (a fe­ (Quercus rubra), scrub (Q . ilic~folia ), and white (Q. male from Missouri). Up until a few years ago there alba ) as well as hickory (Carya spp.). The follow­ were only two specimens in the United States National ing larval description is based on two pickled larvae Museum. Rindge (1971) characterized it as being "an (onc pre-overwintering caterpillar preserved 29 No­ extremely rare species." Ferguson, an authority on the vember 2000 and one mature, post-overwintering North American Geometridae, had never seen this caterpillar preserved 17 May 2001) and 58 larval pho­ species alive before we arranged for him to visit tographs (of three pre-overwintering caterpillars and Goshen, , in 1999. But like so many rare or­ two post-overwintering caterpillars). The pupa was ganisms, in the right localities at the right time, L. per­ preserved on 27 May 2001. Adult, larval, and pupal rnagnaria can be common. At Goshen we occasionally vouchers and slides (transparencies) are deposited at observed more than a dozen individuals at light on the University of Connecticut. nights in early June. This species is distributed from Cranial and body setae of Lytrosis perrnagnaria are Georgia to eastern north to Missouri, Indiana, very short and inconspicuous. Because we had but two northeastern Tennessee, and central Virginia. Herc we larvae, and a Single last instar, our setal mappings must describe and illustrate the last instar larva and pupa for be regarded as tentative. This is particularly true of the the first time, distinguish the larva from congeners, cranial setae and minute body setae that were some­ note several morphological similarities in the imma­ times difficult to locate, ture stages of Lytrosis and Euchlaena, and provide Description. Last Instar Larva. Length: 40 mm (probably at­ brief observations on the 's life history. taining lengths of .50 mm; n = 1). Head (Figs . .5-11, 14-16, 23) somewhat quadrate, with dark spot at top and pale band down each side of triangle; third stemma enlarged; all setae short, espeCially P, METHODS AND RESULTS L, and A setae over dorsum of head (MD setae were not observed). Body (Figs. 1- 4, 12, 13). (Note: in our preserved specimens, the Lytrosis perrnagnaria was seen in the vicinity of the posterior balf of each segment is enlarged, espeCially that of AI. ) shale pit, southeast of Lake Merriweather, on the Ground reddish brown in pre-overwintering larvae and smoky gray­ brown in mature post-overwintering larvae, fading to tan in alcohol; property of the Boy Scouts of America Camp, south­ trunk with numerous brown spots and shmi undulating, often dou­ east of Goshen, Rockbridge Co., Virginia. A female, bled stripes and broken lines of varied width; integument rough with collected at light on 9 June 2000, held in a brown pa­ many shallow pits. Middorsal and subdorsal stripes poorly differen­ tiated; supraspiracular stripe perhaps most evident of all body per bag with a wet cotton ball that had been immersed stripes, espeCially on A3- A6; midventral and subventral stripes pres- 108 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

~ , '- DI , "- , \, *' D2 , \ ./ , , , ~ MDI " '-. I

r .I I SOl ! , \ SD2' 0 'a' '0 I , , 0 LI '0 '0 ~ lU I " l ~ J ,I " / , L3 , " / I' ~ ~ I " " SV setae \ \ ~ , /' ..... ~ .. , ~t} . " .' ". ! f i 0, T ' V '-. f­ 2 TI TIl At A2 A6 A7

3

1'I(:s. 1-4. Last ins tar larva of Lytrosis permagnaria. 1, Habitus. 2, Chaetotaxy; setae associated with thoracic legs not shown; SD2 minute and indicated on ly by its pinaculum. 3, Dorsal view, A9- AlO. 4, Lateral view, A7-AlO .

ent on abdominal segments. Most conspicuous markings include A8 raised. Anterior face of thoracic legs brown. Crochets: 50-55 on black oblique lines on A7 and All below each spirade and small anterior proleg (Fig. 13), 61 - 66 on anal proleg, mostly of two obliqll" line above spiracle on A8 (which is a continuation of the lengths; intercalated fleshy lobe of Forbes (1948) absent. Hypoproct obliq"" line that starts on A7). Horizontal black lim' across anterior and paraproct large, latter nearly one-half the length of anal plate proleg. Dorsum of A9 and AlO marked with incomplete middorsal and extending well beyond body; hypoproct sllhequal to paraproct, line. Spiracular peritreme thinned dorsad and ventrad; Tl and pOinted (Figs. 3, 4, 21). Chaetotaxy (Figs. 2- 4); setae brown, short, A6-AS spiracles enlarged, those on A6 and A7 lowered, and that of often less than one-half the height of spiracle on TI. Two SD and VOLUME 57, NUMBER 2 109

FIGS. 5- 13. SEM images of Lytrosis permagnaria. 5, Head, lateral (scale = 500 ~m). 6, Head, dorsofrontal (scale = 500 ~m). 7, Head, ven­ tral, prolegs removed (scale = 0500 ~m ) . 8, \1axillolahial complex (scale = 0500 ~m). 9, Maxilla (scale = 100 ~lrn). 10, Maxillary palpus (scale = 100 ~m). 11, Antenna (scale = 100 ~m). 12, Mesothoracic claw (scale = 200 ~m ) . 13, Crochets on A6 proleg (scale = ,500 ~m ) .

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FIGs. 14-16. Lytrosis perrnagnaria head. 14, Dorsofrontal. 15, Lateral. 16, Mandibles. llO JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

'. .,~,;~?< :2~'~' ~~ _: . ;.

20

Flt;s. 17-20. Pupa of Lytmsis pemwgnaria. 17, Ventral. 18, Lateral. 19, AS-AlO, ventral. 20, AS-lO, dorsal.

two L setae closely situated on T1. 01 from small wart on AI-AS; DISCUSSION 02 on Al and A.5 from small , often yellowed warts (in living individ­ uals ). S02 minute. LI behind spiracle on Al-AS, but displaced All Lytrosis caterpillars are twig mimics (Figs. downward on An; L2 below and cephalad on AI-AS; L3 grouped 21-26). This is most apparent in Lytrosis sinuosa with SV setae on AI- A2. An with five SV setae. SV setae on A7-A9 and all setae on AlO lengthcncd and paler. L3 and SV setae arising whose texture, coloration, and patterning closely re­ frol\l raised swellings on A7 and AS; A9 with Land SV setae arising sembles that of a Quercus (especially a white oak) twig from large fl eshy warts, SD2 and V extremely reduced. Anal platc (Figs. 24-25; Wagner et al. 2002). It is believed that with fo ur pairs of setae (Figs. 3, 4). Pupa, Length 20 mm, width .5 ..5 mm (n = 1; Figs. 17-20). middle ins tar Lytrosis larvae spend the winter exposed Fusiform, very dark and shiny, deeply rugous over anterior Y:J of on bark-two of three Lytrosis permagnaria that we wings and dorsum of thorax and head. Labrum hemispherical, sleeved on Quercus rubra in October 2000, survived length 0.64 of width. Labial palpus short, tonguelike, nearly as long as wide. Probosds extending just beyond prothoracic leg. Protho­ the winter in eastern Tennessee (Johnston City). racie femur not visible. Mesothoracic leg reaching just beyond an­ McGuffin (1981) stated that Lytrosis unitaria over­ tenna. Metathoracic leg exceeding mesothoracic leg and wing, winters as a 5th instar. In the same work, McGuffin re­ reaching antelior margin of A.5. Mesothoradc spiracle raised, elon­ gate, undercut posteriad. Length of TIll and A1 slIbequal. TI-TIII ported that L. unitaria has up to 9 instars- the largest with 2 setae; Al with 1 seta; A2-A3 with 2; A4 with 3; A.5-A6 with 4, number for any geometrid of which we are aware. SV on prononnced swelling; A7 -AS with :3 setae; A9 with 2 setae and There are four species of Lytrusis in eastern United dark pit cephalad of L seta. AlO cremaster consisting of 4 thickened, recurved pairs of setae and one enlarged pair of caudal hooks (Figs. States (Rindge 1971, Ferguson 1983), only one of 19- 20). which, Lytrosis unitaria (H.-S.) is widespread and com- VOLUME 57, NUMBER 2 III

FIGS. 21-26. Larvae of Lytrosis. 21, Lytrosis pennagnaria, overwintering larva. 22, Lytrosis permagnaria, mature larva. 23, Lytrosis per­ magnana, mature larva, head. 24, Lytrosis sint/osa , overwintering larva. 25, Lytrosis sinuosa, mature larva. 26, Lytrosis "nitaria, mature lalva.

man. The three others are scarce or only locally com­ larged subdorsal swellings on Al and enormous subdor­ mon. Two of the four, Lytrosis heitzmanomm Rindge sal swellings on A5; in addition there are subventral and L. sinuosa Rindge, were not desclibed until 1971- swellings on A2. In Lytrosis permagnaria the D2 setae which is remarkable in that Lytrosis are among the on Al and A5 arise from small, often orange-yellow largest eastern geomcters, with wingspans exceeding 5 walts; the body lacks conspicuous ridges or swellings. cm. Larvae are known for three of these. In Lytrosis The immature stages of Lytrosis share several simi­ unitaria the 02 setae arise from a transverse ridge on larities with members of the genus Euchlaena. Com­ Al that is less than one-fourth the segment length and mon features include the D2 setae arising from warts A5 has conical projections that bear the 02 setae. In or ridges on Al and A5; the black prespiraeular dashes, Lytrosis sinuosa the D2 setae arise ii'om grossly en- best developed anterior to the spiracle on A4- A6; the 112 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

presence of 5 SV setae on A6; the black horizontal line the three sites. In captivity L. permagnaria larvae ac­ on the anterior proleg; the humped dorsum of A8; cepted Quercus alba, Q. ilicifolia, Q. rubra, and a and, according to McGuffin (1981), a similar pupal cal­ Carya species. Survivorship was higher on Querucs, losity (=mesothoracic spiracle). Larvae of the two gen­ perhaps because picked oak foliage holds up longer. era may be distinguished as follows: the crochets num­ Lytrosis unitaria, the best known member of the ber >50 in Lytrosis; there are oblique (black) lines on genus, has been recorded from Acer, , A 7 (below spiracle) and A8 (above and below spiracle) , Pinus strobus, Prunus, Quercus, Rosa, and that are more apparent in Lytrosis than any of the Eu­ (McGuffin 1981, Wagner et al. 2002, DLW chlaena that we have examined (six eastern species); unpublished data). Wild hosts are unknown for Lytru­ the D setae are approximately one-half of the spiracu­ sis sinuosa, but captive individuals have been reared lar height (in Euchlaena the D setae are subequal to from both Acer negundo and Quercus (Wagner et al. the spiracular height); and lastly, both the paraprocts 2002). Host data for the related, and more well studied and anal proleg are proportionately larger in Lytrosis. genus, Euchlaena, indicate that its members are The close pairing of the two SD setae and the proxi­ widely polyphagous on woody plants (McGuffin 1981, mate grouping of the L setae on TI also may be diag­ HandHeld 1999, Wagner et al. 2002). It seems unlikely nostic for Lytrosis. This condition does not occur in that the moth's scarcity will be explained by an unusual (Minot) (McGuffin 1981) or host association-we leave it to others to discover why (Drury) (DLW specimens). such a widespread, unspecialized feeder, remains one Rindge (1971) noted that adults of Lytrosis permag­ of the East's rarest . naria possessed the most primitive features of any of the four members in the genus: i.e., the male has a ACKNOWLEDGMENTS metatibial hairpencil and the vesica has separate The late Douglas Ferguson provided many helpf1l1 comments spines that are exerted on the right, anterior to the on a previous draft, encouragement, and companionship at Goshen. John Richard Heitzman shared several useful observa­ apex of the aedeagus. The unremarkable larvalmor­ tions on Lytrvsis spedes in Missouri. Shawn Kennedy prepared phology of L. permagnaria supports Rindge'S posi­ the larval haGitus, pupal draWings, and assisted with the prepara­ tion-its body is unwarted and more closely resembles tion of the figures; Virge Kask the photographiC plate. Dale that of a Euchlaena than either L. unitaria or L. sinu­ Schweit7.er kindly sent us th e livestock of Lytrosis SinH!!S(l. The fig­ ured Lytrosis unitaria caterpillar was collected by Keith Hartan osa (Figs. 21-26). and photographed by Valerie Giles. Support for this paper came Given Lytrosis permagnaria's overall scarcity in the from the U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team, Cooperative Agreements 01- East, we are puzzled by its abundance at Goshen, Vir­ CA-11244225-215. ginia. Nothing impresses us as exceptional about the locality and indeed we probably would have passed on LITERATURE CITED blacklighting at the site, had we not known that L. per­ FERGUSON , D. C. 1983. Geometridae, pp. 88-107. In Hodges, R. magnaria had been collected along the road in previ­ w: et at. (ed. ), Check list of the of America north ous years. The Goshen colony strikes us as undistin­ of Mexico. E. w: Classey Ltd. and The Wedge Entomolgical guished botanically; woody plants growing in the Research Foundation. Cambridge Univ. Press, Cambridge, United Kingdom. vicinity of our sheets and traps include Acer rubrum, FORBES, w: T. M. 1948. The Lepidoptera of New York and neigh­ Amelanchier sp., Cary a sp., Comus sp., Nyssa sylvat­ boring states. II. Geometridae, Sphingidae, Notodontidae, Ly­ ica, Platanus occidentalis, Quercus rubra, Quercus mantriidae. Memoir 274. Cornell University Agricultural Ex­ periment Station, Ithaca. New York. 263 pp. alba, Sassafras albidum, and Tsuga canadensis. Both H ANDFIELD, L. 1999. Le guide des papillons du Quebec. Broquet Lytrosis unitaria and L. sinousa fly with L. permag­ Inc., Bouchelville, Quebec, Canada, 982 pp. naria at Goshen during early June. J. R. Heitzman MCGUF FIN , W. C. 1981. Guide to th e Geometridae or Canada (Lepidoptera). III. Suhfamily . 3. Memoirs of the (pers. com.) informs us that all four Lytrosis species Entomological Society of Canaela No. 117. 153 pp. may fly sympatrically in the Ozarks. RI NDGE, F. 1971. A revision of the rnoth genus Lytrosis (Lepidoptera, Lytrosis permagnaria has been reported to be lo­ Geornetridae). Arn eliean Museum Novitates 2474. 21 pp. cally common in northeastern Georgia by James WAGNER , D. L., D. C. FERGUSON, T. L. MCCABE & R. C. REARDO N. 2002. Geometroid caterpillars of northeastern and Appalachian Adams and in Cheaha State Park, Alabama by Tim forests. US FS Technology Transfer Bulletin, FHTET-2001-10. McCabe. Both of these localities and Goshen are low 239 pp. elevation or foothill Appalachian forests-to the best Received for publication 3 August 2002; revised and accepted 19 De­ of our understanding, no unusual plant is common to cember 2002.