Steenstrupia ZOOLOGICAL MUSEUM UN IVERSITY OF COPENHAGE N

Volume 6 (16) : 253 - 289 November 25 , 1980

Description of a new genus and two new species of from South America, with discussion of phylogeny and biological observations (: Neopseustoidea) by D. R. Davis

Department of Entomology National Museum of Natural History Smithsonian Lns titution, Washington D. C. 20560, USA and E. S. Nielsen

Zoological Museum, University of Copenhagen Universitetsparken 15, DK-2100 Copenhagen 0, Denmark

A b s tr act. One new genus, Synempora, and two new species, Synempora andesae and Apopiania penai, of the Ncopseustidae are described from temperate South America. The systematic position of the family Ne:opse ustidae is reviewed: The family represents the sister-group of the l'·'eolepidoptera, and a new suborder, Neopseustina, is proposed to ac­ comodate it. A cladistic analysis of the four genera of the family is given; it demon­ strates monophyly of a group comprising the two South American genera, Synempora gen. n. and Apopiania Davis, 1975. Also included are biological observations dealing with diurnal and nocturnal flight activity. resting posture, and bamboo or its debris as the possible larval feeding substratum.

I . Introduction

In a revision of the archaic family Neopseustidae, Davis (l975b) recog­ nized three genera and seven species, of which all species except one were restricted to the southeast Asian areas: Taiwan, India: Assam, Burma and China: Szechuan. The unique South American species, Apoplania chilensis 254

Davis, 1975, was known only from the northernmost temperate areas in Chile, the provinces Nuble, Linares, and Maule, and has not been re corded since. Recent collecting activities in the Patagonian region of South America by the above authors as well as by Dr. O. S. Flint of the Smithsonian Institution and Mr. L. E. Pena of the Institu to de Molina, has resulted in the discovery of one new ge nus and two new species of Neopseustidae. Sufficient series of these taxa are now available to permit several new observations to be offered re­ garding the systematics and biology of this interesting family. Davis' fieldwork was supported by a grant from the Smithsonian Insti­ tu ti on's Fluid Research Fund. His efforts were concentrated in the Nothofagus -Araucaria forests of southern Chile during the months of January and Feb­ ruary 1979_ He was greatly assisted in the field by his wife Mignon Davis, his assistant Bimta Akerbergs and Mr. Luis E. Peria G. Nielsen's fi eldwork was undertaken together with members of the staff of the Department of En tomology, Zoological Museum , Copenhagen, partici­ pating in The Danish Scientific Expedition to Patagonia and Tierra del Fuego 1978-79, an expedition predominantly operating in the Argentine andine temperate forest areas (Nielsen 1980). We wish to express our deep appreciation to Mr. Luis E. Pena G. and his associates at lnstitu to de Molina for courtesies too numerous to be mentioned. 1 We are indebted to Dr. N. P. Kristensen, ZMUC, for valuable discussions; like­ wise we wish to thank Ms . B. Akerbergs, USNM , for preparing most line illu­ strations, and Mr. O. Martin, Mr. B. W. Rasmussen, and Mr. R. Nielsen, ZMUC, and Mr. J. Fuglsang Nielsen for help in preparing the remaining illustrations.

The following ab breviations are used for deposition of material: BMNH British Museum (Natural History), London, England. CSI RO Commonwealth Scientific and Industri al Research Organization, Division of Entomology , Canberra City, Australia. IPCN Instituto Patagonico de Ciencias Naturales, S. M. de los Andes, Argen ti na. LEP Collection of Luis E. Pena G., Instituto de Molina, Santiago, Chile. MACN Museo Argentino de Ciencias Naturales, Buenos Aires , Argentina. MARC Department of Scientific and Industrial Research , Mt. Albert Research Center, Auckland, New Zealand. USNM United States National Museum of Natural History, Smithsonian Institution, Washington, D. C. ZMUC Zoological Museum, University of Copenhagen , Copenhagen, Denmark. r igs 1, 2. Synempora andesae sp. n. 1: Male, holotype. 2: Female, paratype. Scales: 1,2: 3 mm. 2 255

rate areas in Chile, :orded since. South America by Isonian Institution in the discovelY of ient series of these to be offered re-

Smithsonian Insti­ in the Nothofagus . January and Feb­ ~ Mignon Davis, his

Jers of the staff of )penhagen, partici­ d Tierra del Fuego Argentine andine

E. Pena G. and his 1 is to be mentioned. lie discussions; like­ ning most line illu- R. Nielsen , ZM UC , ing illustrations. rial:

~ngland. earch Organization , ralia. ;. M. de los Andes,

~ Molina, Santiago,

os Aires, Argentina. ~search , Mt. Albert istory , Smithsonian

agen, Copenhagen, lara type. Scales: 1,2: 2 256

3 !J Fig. 3. Apoplal1 ;a pClwi sp. n. Male, holotype. Scale: 3 mm. (j 2. Descripti ns of the new taxa

Synempora ge n. n.

T ype- species: Synempora (1ndesae sp . n.

Adu I t (Figs 1, 2). Relatively small , moderately heavy- bodied , with fully scaled, moderately broad wings . Antennae usually exceeding length of forewing sli gh tly. H e a d (Figs 4- 11). An tennae long, approximately 0.9-1.2 the length of f re­ wing. Scape long, intercala.ry scleri te present. Flagetl um segmen ts with p ilifonn 4 scales arranged in longi tu dinal rows of 2-4 contiguous raised sockets (Figs 7- 9) ; Figs 4-6. S 5: Mandible. longitudinal ri dges of scales with scutes, fl utes distinct (F ig. 1 1); segments with numerous short sensill a trichodea (Figs 9 , 1 0), m ~ U1 y long sensil1a tricho dea (Fig. 8) and few sensilla chae tica along distal margin (Fig. 10): some sensi/­ 1979): 0.3 1a of the two first mentioned types wi th tip divided. Eyes somewhat reduced, frons and • in teroclilar index (Davis 197 5a) : 0.81; supraocular index (Kristensen & Nielse n demes wee 257

ed moths, with fully g length of forewing

~ the length of fore­ ;ments wi th piliform d soc kets (F igs 7-9); Figs 4-6. Synempora andesae sp. n., head and head structures. 4: Head, anterior view. 5: Mandible. 6: Maxilla. Scales: 4: 0.5 mm; 5,6: 0.5 mm. Fig. II); segments long sensilla trich o­ ig. 10); some sensi!­ 1979): 0.30; eye index (Powell 1973): 1.0. Chaetosemata well developed on ; somewhat reduced , frons and vertex. Mandibles relatively large; both adductor and abductor apo­ Cristensen & ielsen demes well developed (Fig. 5). Galeae with scattered scales on the proximal 258 third, otherwise naked, short, subequal to entire labial palpus and about 0.5 the length of maxillalY palpus. Maxillary palpi five segmented with fourth segment the longest and slightly exceeding length of fifth. Labial palpi three segmen ted, nearly reaching apex of fourth segment of maxillary palpus; second and third segments approximately equal in length. Prementum elongate, sep­ ara ted and divergent for most its length with free arms nearly as long as basal segment of labial palpus. T h orax. Proti bial epiphysis present but extremely reduced in size (Fig. 12), its length approximately X3 its greatest width; metathoracic tibia with basal spines arising from distal th ird. Forewings (Fig. 14) slightly more slender th an in Apoplallia, forewing index : DAD; venation approximately similar to that of

Apoplania, bu t euA l diverges gradually from CuA2 from the poin t of fur­ cation in both fore- and hindwing, not abmptly as in Apoplania. Pregenital abd ome n . Relatively short and broad. Seventh sternite of m ale with a sligl tly raised median oval pad bearing a few scattered short, ac ute spines. Female with fenes trae of fourth sternite vestigial (Fig. 30); seventh stem ite heavily sclerotized, strongly curved to produce a large, median, caudal lobe bearing numerous m inute spines. A large ventral pocket present between seven th and eight stern ites. Male gen italia (Figs 23-25)1. Uncus well developed, exceeding length of valvae, divided almost its entire length into two inwardly curved lobes. Tegu­ mental lobes triangular, relatively short. Socii prominent, densely setose. Gn athos a small median plate bearing several stout caudal spines. Valvae rela­ tively short; simple and undivided. Vinculum a relatively narrow ring ventral­ ly with a pai r of well-developed, slender, widely separated anterior apophyses. Anellus a darkly sclerotized , symmetrically lobed, spinose selerite dorsad an d laterad to slender, rodlike aedeagus. Juxta likewise a deeply bilobed sclerite with a closely associated pair of acute lobes situated more laterally and ventral­ ly. Female genitalia (Figs 30, 32, 34). Posterior apophyses stout, relatively

I The terminology here applied in the description of the male and female genitalia is in accordance with Davis (1975b). The true nature of these generally highly specialized structures of the Neopseustidae is still insufficiently understood.

Figs 7-1l. Synempora andesae sp. n., antennal structures. 7: Flagellum segment with scales. 8: Flagellum segment with scales partly removed. 9: Detail of 8: Scale sockets and short sensialla trichodea, some with tip divided (arrow). 10 : Detail of 7; ch: sensillum chaeticum, It: long sensillum trichodeum, st: short sensillum trichodeum; note: divided tip of It (arrow). 11: Detail of lamellar flagellum scale. Scales: 7,8: 0.08 mm; 9-11: 0.004 mm. 259 palpus and abou t 0.5 :gmen ted wi ili fou rtil n. Labial palpi three J(illary palpus; second nentUI11 elongate, sep­ 11early as long as basal lced in size (Fig. 12), Dracic tibia with bas al Itly more slender than ately similar to iliat 0 :om the point of fu r­ Ipialzia.

~venth sternite of male scattered short , acu te ~al (Fig. 30); seventh 1 large , median , cau dal ocket present between l. exceeding lengili of 'Y curved lobes. Tegu­ inent, densely setose. lal spines. Valvae rela­ y narrow ring ventral­ ed anterior apophyses. )se selerite dorsad an d leeply bilobed selerite e laterally and ven tral ­ hyses stout, relatively and female genitalia is in nerally highly specialized

Flagellum segment with Iii of 8: Scale sockets and ~ etail of 7; ch: sensillum ichodeurn; note: divided ~s : 7, 8: 0.08 rnm; 9-11 : 260 short, approximately twice the length of ninth tergite; caudal end of ovipositor abruptly narrowing to a relatively broad trilobed apex. Ninth tergite divided medially and forming a pair of lateral lobes. Ductus spermathecae relatively straight to vesicle; utriculus beyond vesicle elongate, sinuate, and terminating in a slightly enlarged receptacular gland; lagena present. Ductus bursae very short, with membranous walls. Corpus bursae completely membranous, small, not extending cephalad further than seventh segment. Etymology. The generic name is derived from the Greeksynemporos which means fellow traveller, in reference to its sister group Apoplania (wanderer). It is to be treated as feminine .

Synempora andesae sp. n.

Adult (Figs 1,2). Wing expanse: d, 15-16 mm; 9 , 18-20mm. Head (Fig. 4). Vestiture willte to pale grey, sometimes heavily suffused with brown in female; dark brown to nearly black in male. Antennae with 93-107 segments; scape pale grey to brown in female, dark brown in male; flagellum light brown. Maxillary palpi and labial palpi greyish to brown in female, dark brown in male. Thorax. Dorsum dark brown in female with a median caudal tuft of elongate white scales; tegulae dark brown with lighter brown apices; male with dorsum and tegulae uniformly darker brown, nearly black. Venter thinly scaled, greyish in female , more brown in male. Prothoracic legs pale brown in female with darker brown banding evident at apex of femur, three dark bands on tibia, and a single band around each tarsal segment; banding darker in male. Meso and metathoracic legs paler in color, with similar banding. Forewings brownish to fuscous in female mottled with lighter shades of brown and white; tornal and anal areas often with white scaling predominate; forewings of male mostly dark fuscous, marked with irregular black and metallic spots and with a sparse 12 scattering ofwhitish scales. Hindwings uniformly fuscous in female with a short Figs 12, 13. Protit uniformly brownish fringe. penai sp. n. Scales : Wing scales (Figs 16, 17). Scale cover on dorsal wing surface dense. Micro­ trichia present. Forewing scales gradually expanding in width from base out­ Pregenital abd wards, apical margin with a medial notch. Metallic scales not different in shape formly straminec from nonmetallic ones. Longitudinal ridges with scu tes, flu tes distinct. In ter­ Male genitalia ridge pia tes relatively large, with a marginal thickening, the marginal flute, and their length, the : from one to three irregular placed flutes in center of plate, thus giving a pretzel­ Gnathos elongate like appearance (Fig. 17). Areas enclosed by marginal flutes mostly nonper­ approximately 8· forated. Scales of hindwing similar to those of forewing but apices rounded Anellus deeply d instead of notched. lateral lobe witt 261 lal end of ovipositor iinth tergite divided mathecae relatively ~te , and terminating Ductus bursae very membranous, small, csynemporos which lania (wanderer). It

Imm. eavily suffused with Itennae with 93-107 ~ in male; flagellum own in female, dark

Idal tuft of elongate ; male with dorsum mter thinly scaled, ,ale brown in female dark bands on tibia, <[ker in male. Meso ~orewings brownish and white; tornal ings of male mostly ~ and with a sparse 12 13 female with a short Figs 12, 13. Protibia with epiphysis (arrow) . 12: Synempora andesae sp. n. 13: Apopiania penai sp. n. Scales: 12, 13: 0.2 mm. rface dense. Micro­ dth from base out· Pregenital abdomen. Sparsely clothed with slender, pilifof!TI scales: uni­ ~ differen t in shape formly stramineous in female, brown in male . Ites distinct. Inter­ Male genitalia (Figs 23-25). Lobes of uncus widely separated for most of marginal flute , and their length, then converging caudally; tapering gradually to subacu te apices. IUS giving a pretzel­ Gnathos elongate, consisting of a narrow plate tenninating in a connate mass of es mostly nonper­ approximately 8-10 stou t setae . Anterior apophyses of vinculum very slender. lut apices rounded Anellus deeply divided into a pair of bilobed, densely spinose lobes; the most lateral lobe with its apex curving anteriorly. Juxta complex, with a deeply 262 divided dorsal scleri te bearing a pair ofspinose lateral anns, and a more ven trad, more membranous sheet bearing a lateral pair of hornlike lobes. Aedeagus reduced to a slender sclerotized rod dorsal to the ejaculatory duct. Male reproduc tive system (Fig. 22). Simplex ejaculatory duct with a posterior slender portion abou t twice as long as aedeagus, abruptly widening into a broad portion about twice as long as slender part; duplex ejaculatory ducts broad and short; accessory glands broad and short; vesicula seminalis similar broad, slightly longer than accessory glands. Broad ejaculatory ducts, accessory glands and vesicula fonning a twisted H-shaped structure. Vasa deferentia long and slender, somewhat irregular in width and fonning an an­ terior curve. Testes small, separate and oval structures.

Female genitalia (Figs 30, 32, 34). Apex of ovipositor with a relatively blunt median lobe and a bilateral pair of shorter, more acute lobes. Ninth ter­ gite deeply divided into a pair of mostly lateral pads on either side of ovi­ positor. Spennatheca extremely long, approximately X2 the length of bursa copulatrix. Corpus bursae reduced, only slightly larger than ductus bursae and approximately equalling posterior apophyses in length. Material examined. Holotype 0 : Argentina: Chubut Province, Esquel, Lago Menendez, El Sagrario Puerto, 550 m, 21. ii. 1979 (Mision Cientifica Danesa, Sta. 49) (ZMUC). Paratypes: Argentina: Same locality as holotype: 30,20. ii. 1979 (ZMUC); 150, 199 , 21. ii. 1979 (ZMUC); 10, 19, 21. ii. 1979 (BMNH); 10, 19, 21. ii. 1979 (CSIRO); 19,21. ii. 1979 (LEP); 10, 21. ii. 1979 (IPCN); 19 ,21. ii. 1979 (MACN). Chile: Osorno Province, Parque Nacional de Puyehue, Aquas Calientes to 2 km S., 600 m: 19,6. ii. 1978 (c. and O. Flint) (USNM), 10, 19, 10.-22. ii. 1979 (D. and M. Davis, B. Akerbergs) (USNM). Additional material preserved in alcohol and not to be included in the type series: Argentina: same locality as holotype: 10, 19 , 20. ii. 1979; 160, 119, 21 . i i. 1979 (ZMUC). Distribution (Fig. 36). Only known from the above mentioned two locali­ ties which both are situated in the Valdivian Cordillera forest area (tenn of Pena 1966). Habi ta 1. The entire Argentinian series was collected in the very moist forests of El Sagrario Puerto between the western end of Brazo Norte of Lago Menen­ dez and Lago Cisne in Parque National Los Alerces, west of Esquel. These forests are very dense and rich in plant species; however, collecting in the more diverse parts of the forest was unsuccessful for S. andesae. Instead, a habitat characterized by a dominance of Nothofagus dombeyi (Mirb.) Blume with a very dense understorey of Chusquea culeou E. Desv. (Fig. 43) produced the Figs 1­ entire series. Scale: ~ 263 ns, and a more ventrad, SC2 RI uike lobes. Aedeagus SCI tory duct. R5

~--~.... aculatory duct with a .,;:,,-_...­ gus, abruptly widening rt; duplex ejaculatory M3 ort; vesicula seminaJis road ejaculatory ducts, CuAI laped structure. Vasa Ith and forming an an­

,sitor with a relatively acute lobes. Ninth ter­ on either side of ovi­ R5 )(2 the length of bu rsa than ductus bursae and 14

Province, Esquel, Lago sion Cientifica Danesa, SC2 R, :,20. ii. 1979 (ZMUC); SCI R2'3 79 (BMNH); 10, 19, R. 21. ii. 1979 (IPCN); !, Parque NacionaJ de R5 . _. ___J. 1978 (C. and O. Flint) - ... -.... - M, I.kerbergs) (USNM). --- M2 be included in the type " . , M3 lii.1979; 160, 119, CuA, 1A+2A CuP CuA2 mentioned two locali­ Sc 'a forest area (term of R,

the very moist forests Norte of Lago Menen­ R. lest of Esquel. These collecting in the more ae. Instead, a habitat (Mirb.) Blume with a 15 ~ ---+I Fig. 43) produced the Figs 14, 15. Wing venation . 14: Synempora andesae sp. n. 15: Apop/ania penai sp. n. Scale: 2 mm. 264

Figs 16, 17 . Synempora andesae sp. n., wing and scale structures. 16: Forewing upperside vestiturej scales removed on lower part; note: microtrichia and medial notch in the apical margin of scales. 17: Detail of lamellar scale; note: pretzellike structures on inter-ridge plates. Scales: 1-6: 0.1 mOl; 17: 0.002 mOl.

The few specimens from Chile were mostly collected along the Rio Chan­ 1eufu, near dense stands of Chusquea.

Apoplania penai sp. n.

Adult (Fig. 3). Wing expanse: d , 17-19.5 mm; 9 , 16 mm. Head. Vestiture dull white to very pale stramineous. Eyes large, interocular index approximately l.2, eye index l.l. Antennae uniformly stramineous, ap­ proximately l.1-1.2 the length of forewing, 93-114 segmented; vestiture similar to that described for A . chilensis, arising from raised sockets arranged in

Figs 18-21. Apopiania penai sp . n., wing and scale structures. 18: Forewing upperside vestiture; scales removed on lower part; note: microtrichia, acute lamellar scale, and pili­ form scale (arrow). 19: Detail of lamellar scale. 20: Detail of lamellar scale at higher magnification. 21. Lamellar scale with oblique crests. Scales: 18: 0.04 mOl; 19-21: 0.002 mm. 265

Ires. 16: Forewing upperside ld medial notch in the apical tikc structures on inter-ridge

·ted along the Rio Chan-

Imm. Eyes large, interocular liformly stramineous, ap­ 4 segmented; vestiture 'aised sockets arranged in

es. 18: Forewing upperside lcute lamellar s ales and pili­ il of lamellar scale at higher ales: 18 : 0.04 mm; 19-21: 266

scattered longitudinal oriented rows of 2 to 6 sockets; socket rows concen­ trated toward distal end of each cupulifonn flagellomere; irregularly inter­ spersed between socket rows are solitary sensory sensilla with an enlarged socket. Maxillary palpi stramineous with dark brown scales concentrated at distal third of third, most of fourth, and at base of fifth segments. Labial palpi stramineous with dark brown scales scattered over most of distal two segments.

Thorax. Dorsum pale brown, tegulae stramineous, venter whitish to stra­ mineous. Prothoracic and mesothoracic legs mostly dark brown dorsally with scattered stramineous scales; tarsi appearing ringed with stramineous; most of ventral surfaces stramineous. Metathoracic legs paler with tarsal banding less evident. Protibial epiphysis prominent in size, approximately 0.4 the length of tibia (Fig. 13). Wing venation illustrated on Fig. IS; forewing index: 0.46. Forewing translucent, thinly scaled, mostly a dull white color, mottled with ( irregular bu t symmetrically arranged patches and slender transverse streaks of dark brown, giving the forewing a somewhat barred appearance; most promi­ \ nent spots of forewing consisting of a subapical, crescent shaped brown bar from costa and fading toward tennen; a less prominent brown bar from distal third of costa; and an obscure, oval patch of black scales at apex of discal cell. Hindwings less heavily marked, nearly transparent, with a slight darkening at

apex of cell and numerous transverse bars of dark brown presenting an overall Fig. 22. SYI barred pattern. cale; ag : a CI ejacul atory d Wing scales (Figs 18-21). Forewing scales medially with lateral margins parallel, distally with margins tapering to an acute point, thereby obliterating a reduced, di distinct apical margin. Black scales fonning dark spot on cross-vein R-M in large, aeu te forewing broader and shorter than nonnal scales and similarly acute. Longitu­ witllOut 101 dinal ridges disappear just at scale margin. Flutes distinct, inter-ridge plates an terior ap relatively small (Figs 19, 20). Dorsal side of wings beyond relatively short branou s she acute scales also provided with scattered, relatively long scales (Fig. 18). Ultra­ lateral to a structure of long and short scales similar. Some short scales with oblique acute lobes crests overlaying flutes (Fig. 21). Hindwing scales similar in structure to fore­ scleri te nar wing scales. a median rl Pregeni tal abdomen. Sparsely clothed with slender, pilifonn scales; strami­ consisting I neous above and more whitish ventrally. Female with a pair of large oval distal third fenestrae present near caudal margin of fourth sternite; seventh sternite heavily aedeagus bl sclerotized, with over 30 minute, peg-like setae concentrated over apex of processes d median lobe. A large ventral pocket present between seventh and eighth ster­ directed eel nites (Fig. 31). F emale g' Male genitalia (Figs 26-29). Uncus broad and rounded, with a short pair rounded ffii of blunt caudal processes separated by a shallow median cleft. Socii reduced sally and la to a pair oflateral setal patches. Tegumen without caudolaterallobes. Gnathos Ninth stem 267

ts; socket rows concen­ Imere ; irregularly in ter­ VD ensi11a wi th an enlarged TS 'n scales concen tra ted at h segments. Labial palpi VS ;t of distal two segments.

, venter whitish to stra­ lark brown dorsally with lth stramineous; most of with tarsal banding less imately 0.4 the length of ; forewing index: 0.46. VS hite color, mottled with der transverse streaks of TS ppearance; most promi­ VD :scent shaped brown bar nt brown bar from distal ties at apex of discal cell. 22 'ith a slight darkening at Iwn presenting an overall Fig. 22. Sy nempora andesae sp. n. Reproductive system of male: schematic drawing to scale; ag: accessory gland~ , ed: ejaculatory duct , se: simplex ejaculatory duct, sd: duplex Illy with lateral margins ejaculatory duct, ts: testes, vd : vas deferens, vs: vesicula seminalis. Scale: 0.5 mm. 1t, thereby obliterating a reduced, divided into a ventral and a dorsal part, bearing a series of relatively ot on cross-vein R-M in . large, acute spines arranged in two median rows. V alvae rel atively long, simple, imilady acu teo Longitu­ without lobes or processes; apex evenly rounded. Vinculum with a pair of 'stinct, in ter- ridge plates anterior apophyses, approximately 0.5 the length of valvae. Anellus a mem­ beyond relatively short branous sheet dorsally wi-th a prominent pair of heavily sc\erotized lobe arising g scales (Fig. 18). Ultra­ lateral to aedeagus; each lobe with three or two (the medial reduced) small Lort scales wlth oblique acute lobes arising from caudal margin of distal h al f Juxta a relatively broad Lilar in structure to fore­ sclerite narrowing noticeably caudad; caudal margin irregular, deeply serrated; a median row of 4- 5 minu te spines arising ventraHy from caudal half. Aedeagus , pilifonn scales; strarni­ consisting of a prominent, heavily sclerotized rod sharply curved dorsally at [th a pair of large oval distal third and from that point projecting caudad beyond gonopore; apex of seventhsternite heavily aedeagus broad and flattened , heavily armed with a pair of short spinose apical entrated over apex of processes directed laterad and a pair of much larger setose lanceolate processes seven th and eigh th s ter­ directed cephalad. Female genitalia (Figs 31, 33, 35). Apex of ovipositor with a bluntly ,nded, with a short pair rounded median lobe. Ninth tergite hoodlike, largely covering ovipositor dor­ an cleft. Socii reduced sally and laterally; caudal margin deeply divided exposing apex of ovipositor. -olateral lobes. Gn athos Ninth sternite divided medially into two symmetrical ventrally concave scler­ 268

AE- ---I-'::.!+

24

H4;i"-- AE

Figs 23-25. Synempora andesae sp. n., male genitalia. 23: Lateral view. 24: Ventral view. 25: Aedeagus and juxta in lateral view. ae: aedeagus, an : anellus,juh:juxta, hornlike lobe, jus:juxta, spinose arm. Scale: 0.5 mm.

ites of highly irregular form. Ductus bursae relatively long, extending anteriorly to sixth segment, without internal folds or thickened walls. Corpus bursae elongate, approximately X3 the length of posterior apophyses, and slender.

Material examined. Holotype 0: Chile: Osorno Province, Parque Nacional Figs 26-29 : Ap de Puyehue, Aguas Calientes to 2 km south, 600 meters, 10-22.ii.1979 (D. and 28: Aedeagul, l ~ M. Davis, B. Akerbergs), USNM 76464 (USNM). anellus, ju: juxt, 269

AE --\'~

AN , Ii,( , --~ '\'Y~ " , ~ "\~~".), I / .~:t. '/...... ~ .....~ 26

./ , , JUH v JUS ~ AE 27

T lv, 24: Vcntf

:ending an teriorly , Corpus bursae , and slender. 1

Parque Nacional Figs 26-29: Apoplania penai sp. n" male genitalia. 26: Lateral view. 27: Ventral view. Lii,1979 (0. and 28: Aedeagus, lateral view. 29: Aedeagus and anellus, dorsal view. ae: aedeagus, an: anellus, ju :juxta. Scale: 0 .5 mm, 270

Para types: Argentina: Rio Negro Province, Lago Nahuel Huapi, Puerto Blest, 770 m: 20, 17.xii.1978; 10, 26.xii.1978 (Mision Cientifica Danesa, Sta. 8) (ZMUC). Chile: Malleco Province, Cordillera Nahuelbuta, Cabreria, 1100 me­ ters: 10, 21.i.l977 (L. Pena) (LEP); 10, 9-15.i.1977 (USNM); 10, 19, 15 -20.i.1977 (USNM); Cordillera Nahuelbuta, Pichinahuel, l300 meters: 10, -, 9-15.i.1977 (USNM). Osorno Province, same data as holotype: 10 (BMNH), 1o (CSIRO), 1o (MARC), 90 (USNM); 1 km S. Aguas Calientes, Rio Chan­ leufu: 10, 8-9.ii.I978, (c. & O. Flint) (USNM); Valdivia Province, Sto. Domingo, Rincon de Piedra, ca. 23 km SE Valdivia: 10, 3.ii.l975 (E. Krahmer) (USNM)_ Described from a total of 22 males and 1 female. Additional material preserved in alcohol and not included in the type series: Argentina: Rio Negro Province, Lago Nahuel Huapi, Puerto Blest 770 m: 6 0 , 7.xii.l978 to 3.i.1979 (Mision Cientifica Danesa, Sta. 8) (ZMUC). 1 Comparative notes. The adults of Apoplania chilensis Davis and A. penai 30 new species are very similar superficially. In maculation they may be distin­ III guished by the heavier brown scaling at the apex of the forewing and along the termen of the hindwing in A. chilensis, as well as by the rather diffused black spot at the apex of the cell in A. penai; the latter is absent in A. chilensis. In contrast, the male genitalia of the two species differ considerably. Another means of easily distinguishing the two species, particularly helpful in the r- females, is by the wel1 developed epiphysis of A. penai (Fig. l3), as compared to the vestigial condition in A. chilensis (Davis 1975b, fig. 48). (/ - ' ­ The wing scale morphology of A. penai differs somewhat from that of A. chilensis, which has the lamellar wing scales obtuse, with an apical margin distinct from the lateral margins of the scale. This is also the case in Ne­ ~ opseustis calliglauca Meyrick, 1909, in which the distal margin is much wider than in A. chilensis (Davis 1975b). As described above, the scales in A. penai taper distally into an acute pOint. The oblique crests which are observed over­ 31 laying the flu tes on some of the scales (Fig. 21) seem to be remnan ts of a her­ ringbone crest pattern. This pattern has previously been recorded only from F igs 30, 3 L the non- Glossatan su borders Zeugloptera and Aglossa ta (Davis 1978, Kristensen segmental nI & Nielsen 1979). The hindwing scales are similar in structure to the forewing 31: Apop/u/l scales. Dis tr ibu ti on (Fig. 36). This species appears restricted to Chusquea-Notho­ all from tl' fagus habitats in certain montane areas of Patagonia between 370 and 41 0 Puerto Ble south latitude. [n the coastal mountains of Chile, it has been found to range and shrubs; from almost sea level near Valdivia to 1300 meters in the Cordillera Nahuel­ strongly cI buta. [n the Andes of Chile and Argentina it has only been found in the lake culeou E, region at two sites less than 100 airline km apart. diverse flo was that .4 Habitat. The series of A. penai collected in Argentina consisted of 9 males, collected i 271

[ Huapi, Puerto Blest, IV V VI VII :ifica Danesa, S tao 8) , Cabreria, 1100 me­ \ JSNM); 10, 19, 15 \~ \ ~ :1, 1300 meters: 10, : C l )Iotype: 10 (BMNH), 'l II J ~ Calientes, Rio Chan­ I I Idivia Province, Sto. r il( "\ '.­ ii.1975 (E. Krahmer) ) 'T ' I .~ led in the type series: f /;f to Blest 770 m: 6 0, V/, 1/ ~ ZMUC). --- r s Davis and A. penai 30 l they may be dis tin­ III V VI VII 'rewing and along the IV rather diffused bl ack :n t in A. chilensis. In \ \ ,nsiderably. Another I( \ I ilarly helpful in the ._ ,J . )L ~ ig . 13), as compared IS. .T' 48). ,hat from that of A. ( ith an apical margin ·also the case in Ne­ largin is much wider ,he scales in A. penai :h are observed over­ 31 )e remnants of a her­ recorded only from lvis 1978, Kristensen Figs 30, 31. Postabdomen and female genitalia in lateral view; Roman numerals denote segmental numbers; note: fenestrae on fourth sternite. 30: Synempora andesae sp. n. lure to the forewing 31:Apop/ania penai sp. n. Scales: 30, 31: 0.5 mm.

o Clmsquea-Notho­ all from the very moist woods at Puerto Blest at the western end of Brazo :ween 37 0 and 41 0 Puerto Blest of Lake Nalhuel Huapi. The forest here is rich in species of trees leen found to range and shrubs; however, the area where the A. penai specimens were collected is ! CordiYIera Nahuel­ strongly dominated by Nothofagus dombeyi (Mirb.) Blume and Chusquea :n found in the lake culeou E. Desv. Collections from more open areas with a richer and more diverse flora did not produce any Neopseustidae at all. The general impression was that A. penai was rare at this locality, because only a few individuals were lnsisted of 9 males, collected in spite of great efforts with from one to five traps operating nearly 272 every night during the flying period. All specimens were collected in UV light This super traps. above the In Chile, a total of 20 specimens of A. penai have been collected at three Eriocranii different montane sites representing both sides of the central valley. Luis Pena, terms rep to whom this species is dedicated, first collected the species at two camp sites Heteronet: situated in the higher reaches of the Cordillera Nahuelbu ta, a costal range rising This s to 1400 meters and located west of the city of Angol. The wet forests here are Neolepidc dominated by extensive stands of Araucaria araucana (Molina) C. Koch and proboscis Nothofagus obliqua (Mirb.) Blume, the branches of which are typically fes­ (i. e., lac tooned with epiphytic lichens. The bamboo genus Chusquea is present but Myoglossc largely restricted to certain forest clearings and streams borders. Further south Within in the low hills east ofValdivia, a single male was collected by Ernesto Krahmer following at an elevation of less than 200 meters. Although this area has been greatly arm in pr o disturbed by agriculture and lumbering, Nothofagus and Chusquea still remain putedly v quite common. The largest number (14) of specimens collected thus far are suggested from Aquas Calientes in the Parque Nacional de Puyehue, a densely forested be a possi area less than 100 airline kilometers from the lake Nahuel Huapi collecting site cuit to elf in Argentina. The forests near Aquas Calientes is rich in species (Fig. 44): (Kristense Nothofagus dombeyi (Mirb.) Blume and Eucryphia cordifolia Cav. are abun­ well-devel dant, as are Drimys winteri Forst, Myrceugenella apiculata (DC.) Kausel and retained : Myrceugenia planipes Berg. Chusquea culeou E. Desv. and C montana Phil. the pre-IT are also common but are largely encountered bordering forest clearings and ductor 111 river banks. The forests here are thus somewhat similar to that of the Puerto mandibul Blest area bu t there Eucryphia and C montana are missing. As many as 4 sepa­ of the ne rate UV light traps and 4 malaise traps were operated in a total of 8 and 7 dif­ dae so fa ferent sites, respectively, within an area of less than 4 square kilometers. The il1 the ad t: site which 'produced the greatest number of A. penai per night consisted of a The r.. dense forest with a heavy undergrowth of ferns and blyophytes bu t little or deserve CI no Chusquea . Thirteen of the specimens of A. penai were collected in light excavate ; traps and one was captured in a malaise trap. As is generally true for most The (\" nocturnal , collecting for neopseustids yielded best results during those probably nights subject to a light rainfall. As was observed for the collections in Argen­ arms are tina, the field work in Chile suggested that A. penai was rare at all localities it ctetidae, was found to occur. remnants excavatec 3. The systematic position of the Neopseustidae tidae , se­ (Kristense The Neopseustidae have previously been regarded as a member of the sub­ some 01>' order Dacnonypha by several au thors, bu t no significant synapomorphies have i. e. , man so far been discussed for the Neopseustidae and the other constituent families also pres. of this suborder. Davis (l97Sb) stressed the isolated position of the Ne­ potamidal opseustidae and proposed a new superfamily, Neopseustoidea, for the family. and HydJ 273

collected in UV light This superfamily was recently (Davis 1978: 36) regarded as representing a grade above the level of the Dacnonypha (as Eriocranioidea, comprised of the families ~en collected at three , and Lophocoronidae), thus in cladistic tral valley. Luis Pena, terms representing the possible sister-group of (= + cies at two camp sites (= Frenata)). a, a costal range rising This suggested sister-group relationship of the Neopseustidae and the le wet forests here are Neolepidoptera is now substantiated by two possible synapomorphies: (1) Molina) C. Koch and proboscis with intrinsic musculature and (2) presence of "normal type" scales ich are typically fes­ (i. e., lacunate, perforated) on the wings, and the ensuing clade is termed Isquea is presen t bu t MyogJossata (Kristensen & Nielsen 1980b). :Jrders. Further sou th Within the MyogJossata, the Neopseustidae represent the grade in which the j by Ernesto Krahmer following two lepidopterous ground- plan character states are retained: (1) Free area has been greatly arm in prothoracic furca slender from base and arising dorsally from an undis­ Chllsquea still remain pu tedly ventral posi tion (Davis 1975b, fig. 2). Recently Kristensen (1 978b) collected thus far are suggested the lateral displacement of the free arm of the prothoracic furca to Ie, a densely forested be a possible neolepidopteran autapomorphy, a character which may be diffi­ I Huapi collecting site cult to evaluate in species with a broad base of the free prothoracic furcal arm in species (Fig. 44): (Kristensen & Nielsen 1979). (2) Mandibular adductor and abductor muscles iifolia Cav. are abun­ well-developed. The mandibular muscles in those Neolepidoptera which have lata (DC.) Kausel and retained rudimentary mandibles are much reduced, while Neopseustidae, as and C. montana Phil. the pre-myogJossatan groups, have retained well-developed adductor and ab­ ~ forest clearings and ductor muscles. As discussed by Kristensen & Nielsen (l980b) the reduced to that of the Puerto mandibular musculature is undoubtedly related to the adetecticous condition g. As many as 4 sepa­ of the neolepidopterous pupae. Even if the immature stages of the Neopseusti­ a total of 8 and 7 dif­ dae so far are unknown, the presence of well -developed mandibular muscles luare kilometers. The in the adults indicate that the neopseustid pupae are decticous. : night consisted of a The Neopseustidae share with the Eriocraniidae some characters which rophytes but little or deserve comments. These are (1) presence of dorsal tentorial arms, (2) deeply ere collee ted in light excavate anteroventral margin of IX in male and (3) reduced ventral diaphragm. rrerally true for most The two first-mentioned characters seem to be genuine symplesiomorphies, : results during those probably pertaining to the lepidopterous ground-plan. The dorsal tentorial collections in Argen­ arms are present in several homoneurous groups: Eriocraniidae, Acanthoptero­ rare at all localities it ctetidae, Mnesarchaeidae and the hepialoid families. In many other groups remnants of the dorsal arms are present as short processes. Segments IX is excavated in males of the Agathiphagidae, Eriocraniidae, Acanthopterocte­ tidae, several (Philpott 1927), Anomoses hylecoetes Turner (Kristensen 1978b; this species also has a small medial anterior extension), J l1ember of the sub­ some , many and representatives of the , mapomorphies have i. e., many Meessiinae. Moreover, a deeply emarginate ventral margin of IX is constituent families also present in representatives of primitive Trichoptera, i. e., in several Philo­ lOsition of the Ne­ potamidae (Annulipalpia), some Rhyacophilidae and many Glossosomatidae dea, for the family. and Hydroptilidae (Integripalpia). The reduction of the ventral diaphragm in 274

\ \ \ I I I I I I I I I / \ I I \ I \ / \ \./ J \

f

32 1 Fig. 32. Synempora andesae sp. n ., female genitalia, ventral view. Scale: 0.5 mm. the Eriocraniidae and its absence in the Neopseustidae are discussed by Kristen­ sen & Nielsen (J980a), and regarded as parallel specializations. Apart from leduced ventral diaphragm, the Neopseustidae and Eriocraniidae do not to our knowledge share any derived traits that will contradict the assign­ ment of the Neopseustidae to the . There is thus a strong evidence of a sister-group relationship between the Neopseustidae + Neolepidoptera, in which case the Neopseustidae represent a clade with suborder status for which we hereby propose the name Neopseustina 2 .

2 Footnote added in press. Kiippers & Speidel (1980) have discussed the systematic position of the family Neopseustidae and full comments on their phylogenetic interpre­ tations are included in Kristensen & Nielsen (l980b). Kiippers & Spcidel also recognize the monophyly of the Neopseustidae and they formally proposed the suborder name Archifrenata to accomodate the family. We regard this name to be misleading because the Neopseustidae in all characters belongs to the non-frenate grade of Lepidoptera. 33 Therefore we would reject the suborder name Archifrenata, and uphold the name Neo­ pseustina proposed in our present paper. Fig. 33. A,c 275

\ \ \ \ \ I I I

.v. Scale: 0.5 mm.

Je discussed by Kristen· ntions. ;tidae and Eriocraniidae ill contradict the assign· s thus a strong evidence ': : te + Neoiepidoptera, in border status for which

t.····. discussed the systematic Ici r phylogenetic in terpre­ t &. Speidel also recognize Dosed the suborder name to be misleading because .., tc grade of Lepidoptera. 33 ld uphold the name Neo­ Fig. 33. Apop/ania penai sp. n ., female genitalia , ventral view. Scale: 0.5 mm. 276

4. The systematic position of Synempora

In the discussion of the phylogeny of the three neopseustid genera known at the time of his revision, Davis (l975b) recognized the genus Archepiolus3 Mu tuura to display the highest number of plesiomorphic character states, and Neopseustis Meyrick and Apoplania Davis to represent an overall more special­ ized pair of genera. In the cladogram on Fig. 37 we have further evaluated and substantiated the phylogeny of the presently known four genera.

The apomorphies I and 2 are shared by all four genera, and are consequently regarded as autapomorphies of the family Neopseustidae: 1. Segments of antennal flagellum strongly cupuliform. Amongnon­ ditrysian lepidoptera a similar condition is only known from some members of the possessing submoniliform flagellar segments (Kristensen & Nielsen 1979). 2. Male sternum VIl with a medial,conical ,s pinose process. A simi­ lar process has not been reported from any other palaeolepidopterous group. As we find the assignment of Archepiolus to the Ne opseustidae suffiCiently well founded on the basis of characters I and 2, we have not undertaken the task of reexamining Archepiolus schmidi Mu tuura, 1971, for characters 3-1l. We tllerefore cannot decide whether these are autapomorphies of the Ne­ opseustidae or synapomorphies ofNeopseustis + (Synempora + Apoplania). 3. Proboscis specialized. Each galea with a functional, closed food-chan­ nel , and the two galeae held together by "zip-scales" (Figs 38, 39) and curved spines on the medial galeal surfaces (Kristensen & Nielsen in prep.). The "zip-scales" are visible in the light microscope. 4. Monocondylar scape -pedicel articulation. Kristensen (1968) de­ scribed this feature from Neopseustis meyricki and stressed its uniqueness among primitive Lepidoptera. The sanle mode of articulation has now been observed in Apoplania chilensis (Davis 1975b, fig. 29), A. penai and Synem­ pora andesae. Figs 34, 3 ~ 5. Antennae as lo n g as or longer than forewings. A similar condition plallia pelle among non-ditrysian Lepidoptera is only found in the heteroneurous mono­ trysian , while it occurs in a limited number of apparently phylogene­ 6. Wing~ tically remote groups of Di trysia. Zeugloph and 0.3 5

3 Footnote added in press. KUppers & Speidel (1980) have synonymized Nemalocenlropus index pfl Hwang, 1965 and Archepiolus Mutuura, 197 1, without examination of representatives of Iidae are either of the two genera. Davis examined the extant type of the fortner genus in Beijing, The inde: China and will elsewhere formally confirm the above mentioned synonymy. the avail s 277

)pseustid genera known at d the genus Archepiolus 3 ,rphic character states, and :nt an overall more special­ have hHther evalu ated and fou r genera.

nera, and are consequently idae: upuliform. Among non­ ~own from some members geUar segments (Kristensen spinose process . A simi­ nlaeolepidopterous group. Neopseustidae sufficiently e have not undertaken the 1971, for characters 3 -1 1. . I tapom orphies of the Ne­ 'nell/pora + Apoplania). Ictional, cl osed food -chan­ ," (Figs 38, 39) and curved & Nielsen in prep.). The

Kristensen (1968) de­ Id stressed its uniqueness articulation has now been 29), A. pellai and Synem­ Figs 34, 35. Apex of ovipositor , ventral view. 34: Synempora andesae sp . n. 35: Apo­ ings. A similar condition plania penai sp. n. Scales: 34 , 35: 0.1 mm. the heteroneurous mono­ r of apparently phylogene­ 6. Wings relatively broad and short, forewing index about 0 .4. In Zeuglop tera, Aglossata and Dacnonypha the forewing index is between 0.29 and 0.35 , while in some groups of it reaches 0.4, but this high index probably cannot be an exoporian ground-plan character, as the Hepia­ 'nonyrnized Nemalocenlropus mination of representatives of lidae are currently regarded subordinate in the Exoporia (Kristensen 1978a). Df the former gellu s in Beijing, The index of Archepiolus varies between 0.35 and 0.38 pending on which of led synony my . the available illustrations has been used for the measurements. 278

7. Upper forewing surface only provided with one type of scales, interpreted by us as "normal type" scales. The shape of the scales varies considerably as described above, but the ultrastructure seems uniform. As reported by Kristensen (1978a), three of the hepialoid families similarly lack the "primitive type" scales.

8. Lamellar scales on upper surface of wings with circular or ellip­ tic perforated or nonperforated plates in the inter-ridge areas. A somewhat similar structure is only observed in one other lepidopterous group, Micropterigidae - Heterobathmiinae, where the inter-ridge plates are unper­ fora ted (Kristensen & Nielsen 1979).

9. Third axillary sclerite strongly melanized, seen as distinct black dot at base of forewing. We have not observed this character in other Lepidoptera.

10. Costal margin of hind wing withou t frenular bristles. These bristles are reported from the Agathiphagidae, Micropterigidae, Eriocraniidae, Lophocoridae (Common 1973), and Mnesarchaeidae (Philpott 1922), while o modified frenular bristles are absent in the hepialoid families (Philpott 1925, 40 Kristensen 1978a). Frenular bristles are also present in many heteroneurous non- di trysians.

11. Mesothoracic apophyses forming an internal bridge (Davis 1975b). This character is to our knowledge not known in any other lepido­ pterous group .

The absence of a ventral diaphragm has been reported from Synempora andesae (Kristensen & Nielsen 1980a), which also has the abdominal inter­ ganglionic connectives paired (Kristensen & Nielsen 1980c). Both these features are considered apomorphous among primitive Lepidoptera, but since representatives for the other neopseustid generd have not been examined for these characters, it is not possible to state whether they apply to the whole family or to subordinate groups only.

Characters 12-13 are interpreted as au tapomorphies of the genus Archepiolus: o 42 12. In tercalary sclerite between antennal scape and pedicel absent. The presence of this sclerite is a feature of the lepidopterous ground-plan (Kristensen 1978a) and is known from all higher groups of non- and several groups of Ditrysia.

Fig. 36. Distribution of Synempora andesae sp. n. and Apoplania penai sp. n. The range of Apopiania chilensis Davis is app. 200-250 km N of Victoria (see Davis 1975b, map 2). 279

.'lith one type of scales, CHILE The shape of the scales varies b.. ucture seems uniform. As b.. aloid families similarly lack S. andesae VICTORIA. * A. penai ~s with circular or ellip­ the inter-ridge areas. A ~ other lepidopterous group, Iter- ridge plates are unper­ I 50 KM I

d, seen as distinct black ALUMINE• rved this character in other frenular bristles. These cropterigidae, Eriocraniidae, idae (Philpott 1922), while oid families (Philpott 1925, sent in many heteroneurous

internal bridge (Davis known in any other lepido­

reported from Sy nempora S. C. de o has the abdominal inter­ BARILOCHE lisen 1980c). Both these itive Lepidoptera, bu t since .ave not been examined for er they apply to the whole

!; of the genus Archepio!us: o 42 ape and pedicel absent. lepidopterous ground-plan groups of 11 0n-Di trysia and

'opial1ia pCl1ai sr. n. The range ARGENTINA aria (see Davis 1975b, map 2). 280

13. Valva divide d. The valva is otherwise simple and undivided in Ne­ opseustidae (Davis 1975b).

}----~ 27-30

24-26

}------~22 - 23 Figs 38, 39. 18-21 galea at the I channel indie 0.01 mm ; 3"

Characters }------~ 14 -17 para + Ape 14. Chaet 12-13 In Archepi. similar to \" 15. ForeV' Archepia[UJ 3- 11 margin. T plesiomorp

~~~------? ------~ 16 . Fore ll yond ere pterous gr' 1-2 this state iE 17. Fore ..

Fig. 37. Cladogram demonstrating the monophyly of the Neopseustidae and the phylogeny scribed by of the neopseustid genera. Numbers refer to characters discussed in the text, section 4. ptera. As Filled circles: apomorphies, open circles: plesiomorphies. 1942). 281 and undivided in Ne­

27-30

24-26

23 Figs 38, 39. Apop/onia penoi sp . n., proboscis structures. 38 : Dorso-medial surface of 21 galea at the level of the food-channel; "zip-scales" at bottom and assumed slit to food­ channel indicated by the arrow. 39: "Zip-scales" at higher magnification. Scales: 38: 0.01 mm ; 39: 0.005 mm.

Characters 14-17 are regarded as synapomorphies of Neopseustis + (Synem­ pora + Apoplania). 14. Chaetosemata on head pre sent on prominently swollen areas. In Archepiolus these areas are only slightly elevated (Davis 1975b), a condition similar to what is observed in most other palaeolepidopterous groups. 15. Forewing and hind wing with R2 and R3 totally coalesc ent. In Archepiolus these two veins are " stalked", forking between cross-vein and margin. The condition of five radial forewing veins reaching the margin is the 3-11 ~ plesiomorphic state known from all palaeolepidopterous groups. 16. Forewing with M[ and M2 basally coalescent ("stalked") be­ yond cross-vein in ter-M. In Archepiolus and most other palaeoJepido­ pterous groups the three medial veins are free beyond the inter·M cross-vein; this state is therefore regarded as primitive. 17. Forewing with an anal pocket. An anal pocket similar to that de ­ Iistidae and the phylogeny scribed by Davis (l975b) is not known from any other homoneurous Lepido­ ed in the text, section 4. ptera. A somewhat similar structure is reported from some Hepialidae (Tindale 1942). 282

Characters 18· 21 are au tapomorphies ofNeopseustis: metafl lumen. 18. lA+2A in forewing distally strongly curved towards CuP. A the lu similar curvature of 1 A + 2A is not known from any other Neopseustidae or archip other homoneurous Lepidoptera. (Davis 19. Uncus a moderate lobe. In other Neopseustidae the uncus is large and to the hood shaped or produced into two prominent elongate lobes. tip is ; 20. Teguminal lobes long. In all other species of Neopseustidae (see Davis is int 1975b) the lobes are relatively much shorter. To ou. sent, tI 21. Intromittent organ highly specialized. The aedeagus is absent and functionally replaced by a pair of parameres (see discussion by Davis 1975b). 23. GJ blunt!). Characters 22 and 23 are synapomorphies of the genera Synempora + Apo­ plania: Charae: 22. Anteromedial process from metathoracic furcal stem directed 24. Se ventrally, reaching almost to or fused with intercoxallamella. As NeopsE described by Davis (1975b, fig. 6), the anteromedial process from the meta­ little. ­ thoracic stem is prominent, directed ventrally and distally fused to the in ter· coxal lamella. This fusion, described from Apoplania chilensis, has now also 25 . Fe been confirmed from A. penai by serial sectioning. Seen in lateral view the margil and the short Sl

26. In struCtl In Apo, simple 1

Characi 27. Zo on for any ot!·

28. Ae

29. Sp Neopse ondary

30. An 40 In spec Fig. 40. Synempora andesae sp. n., meta thoracic furca in lateral view; fs : furcal stem, long al mp: anteromedial process, pa : primary furcal arm. Scale: 0.2 mm . ground 283

metafurca of the two Apoplania species looks similar, having a relatively small rved towards CuP. A lumen between the stem and anteromedial process and a large zone between yother Neopseustidae or the lumen and dorsal margin of primary furcal arms. Whereas in Neopseustis archiphenax Meyrick, 1928, the an teromedial process is very poorly developed (Davis 1975b, fig. 5), in Synempora it is a long slender process almost parallel dae the uncus is large and to the furcal stem and reaching almost to the intercoxal sclerite to which the e lobes. tip is attached by a weak connective (Fig. 40). This condition in Synempora . Neopseustidae (see Davis is interpreted as a trend towards the fused condition observed in Apoplania . To our knowledge the fused condition is unique in Lepidoptera, as when pre­ sent, the anteromedial process is directed anteriorly or anteroventrally. 1e aedeagus is absent and cussion by Davis 197 5b). 23. Gnathos with short, stout spines . In Archepiolus the gnathos is a bluntly serrate lobe and in Neopseustis a slender uncinate process. ~nera Synempora + Apo­ Characters 24-26 are au tapomorphies of the genus Synempora: furcal stem directed 24. Sexual dimorphism very pronounced (see description above). In ntercoxallamella. As Neopseustis and Apoplania the wing pattern and size of the sexes differs very II process from the meta­ little. The female ofArchepiolus is unknown. istaJly fused to the inter­ ia clzilensis, has now also 25. Forewing lamellar scales of Synempora wi th a notch in apical Seen in lateral view the m argi n. The apical margin of Neopseustis calliglauca Meyrick is almost straight and the scales of Apoplania are either produced in to an acu te point or have a short straight margin. Scales ofArchepiolus have not been examined.

26. Inter-ridge plates on forewing lamellar scales with pretzellike stru c ture , i. e. , one or more flutes present inside the marginal flute. In Apoplania and Neopseustis calliglauca each of the plates is surrounded by a simple marginal flu te only.

Characters 29-30 are au tapomorphies of the genus Apoplania: 27. Zone between M and CuA proximal to inner cross-vein M-CuA on forewing without scale cover. A similar condition is not observed in any other Neopseustidae.

28. Aedeagus long, rodlike and heavily armed at tip.

29. Spermathecal lagena absent. The lagena is present in females of Neopseustis and Synempora, wherefore the absence here is regarded as a sec­ ondary specialization in Apoplania.

30. Anteriorly directed anterolateral lobes of vinculum very broad . In species of Archepiolus, Neopseustis and Synempora these lobs are relatively teral view; fs: furcal stem, long and slender, wherefore this condition is ascribed to the neopseustid 1m. ground-plan. 284

The following charac ter is one whose significance cannot yet be evalu ated: In both the forewings and hindwings of Neopseustis and Apoplania, CuA, di­ verges strongly from CuA2 at the CuA,/CuA2 bifurcation point, the resulting two veins being almost parallel shortly beyond this point. In Archepiolus and Synempora the two veins gradually diverge from the point of bifurcation. The strongly diverging state is known also from Lophocoronidae (Common 1973), whereas in Eriocraniidae and Acanthopteroctetidae both conditions are ob­ served (Davis 1978), and in some species the condition is different in fore­ and hind wings. The strongly diverging condition is apparently often linked with the presence of a distal cross-vein M3 - CuA,. As the distal cross-vein M - CuA 1+ 2 is proximal to the CuA bifurcation point in the forewing and totally absent in the hind wing of the Neopseustidae, it is not possible to state whether the strongly diverging condition in Neopseustis and Apoplania is a secondary specialization (and thus a convergence) or a retained plesiomorphic state, the specialized state being that observed in Archepiolus and Synempora.

5. Biological observations on Neopseustidae As noted by Davis (1975b) information available on Neopseustidae indicated the adults to be primarily crepuscular or nocturnal, based on the fact that they are readily attracted to light and had not been seen active during the day_ A primarily nocturnal habit is further suggested by the enlarged eyes of most 41 neopseustid species, a feature not typical of most families of non-ditrysian moths. The eye index of most neopseustid species is approximately 1. 2, which agrees with Powell's (1973) measurements on other nocturnal moths. Synempera andesae has been observed in Argentina to be rather active during the day. Approximately 10 specimens were observed active around noon or early afternoon on bright sunny days. They were flying from one to two meters above the ground relatively fast and undulating, with the antennae held dorsally over the head in a posterolateral position. Consequently, the specimens strongly resembled flying adelids and were difficult to observe owing to the sun-shadow mosaic pattern in the rain forest. When not flying they probably rested on the upper leaves of Chusquea culeou or on the lower branches of trees. The light-colored females probably closely resemble bird droppings when resting. In both Argentina and Chile, S. andesae was also somewhat active at night and was attracted to ultraviolet collecting lights. This "dual" activity is not too surprising, considering the somewhat intermediate size range of the eyes in this species (eye index, 1.0).

Figs 41, 42. Synempora andesae sp. n., male resting on leaf of Nothofagus dombeyi. 41: Dorsal view; note: subapical inflecton of wings (arrow). 42: Antero-lateral view. Scales: 41,42: app. 3 mm. 285 annot yet be evaluated: In

and Apoplania, CuA l di­ rcation poin t, the resulting point. In Archepiolus and ~ point of bifurcation. The oronidae (Common 1973), e both conditions are ob­ dition is different in fore­ is apparently often linked . As the distal cross- vein Joint in the forewing and :, it is not possible to sta te ;eustis and Apoplania is a Ir a retained plesiomorphic 'chepiolus and Synempora.

n Neopseustidae indicated oased on the fact that they active du ring the day. A he enlarged eyes of most families of non-ditrysian approximately l.2, which .octumal moths. ntina to be rather active Iserved active around noon re flying from one to two ng, with the antennae held msequently , the specimens : to observe owing to the . not flying they probably 'on the lower branches of resemble bird droppings

~omewhat active at night 'his "dual" activity is not :e size range of the eyes in

eaf of NOlilo/agu s dombeyi. i'). 42: Antero-lateral view. 286

fron t. 11: leg is helC mid and h As sta' neopseust inserted i suspected h abitat fc Ev iden host for a Becaus: been encc these 111 0 1 ally unsu· stan d of or its det Moreover o f the grc ofbambo The Bt ly it is p Among tI fOigs 43, 44. Habitats of Synempora allde sae sp . n. and Apoplama penai sp. n. 43: Be­ laIld, btl t tween W-end of Brazo Norte of Lago Menendez and Lago Cisne, Parque Nacionnl "Los m os t like: Alerces", Chubut, Argentina, where S. andesae was collected. Area dominated by of bambc Chusquea culeou (unders torey), Nothofagus dombeyi and Fitzroya cupressoidea (over­ dict the 11 storey). 44: Parque Nacional de Puyehue, Osorno, Chile, where both the two new neo­ pseustid species were collected. Area with Chusquea montana, C. culeou (und erstorey), nothing is Eucryphia cordijolia and Nothojagus dombeyi (overstorey). it might Neopseus especiall_ The typical resting position is illustrated in figures 41 , 42. As can be seen, tributionc the wings are steeply reflexed a.T1d the hind wings entirely hidden under the forewing. This is the typical resting posture observed in the most primitive Lepidoptera - the primitive stegopterous posture of Graham (1950). A char­ acteristic feature of the resting Synempora andesae is the subposterior in­ flexion of the wings (Fig. 41). In the resting specimen the body and the dorsum of the wings are held in a position almost parallel to the surface of the substratum, the thorax being slightly elevated. The palpi are appressed to the head: segmen t 2 and 3 of the maxillary palpi are appressed to the fron t of the head and 4 and 5 to the fron t and ventral part of the head, wi th the flexing taking place between segments 1-2 and 3-4. The labial palpi are curved under the head in a position lateral to the maxillary palpi, being appressed to the 287

front. The antennae are directed dorso-laterally and slightly curved. The fore­ leg is held at the level of the head and not stretched directly forward and the mid and hind legs are directed backwards. As stated by Davis (1975b), nothing is known about the biology of any neopseustid species, although it was suggested that the eggs were probably inserted in to relatively firm tissue of the plant host. The larvae were thus suspected of being endophagous. It was further observed that the preferred habi tat for Apoplania was in forests dominated by Nothofagus and Ch usquea . Evidence is now beginning to indicate that the latter may actually be the host for at leas t some Neopseustidae. Because (1 ) Sou th American Neopseustidae have, wi th one exception, only been encountered in habitats associated with Chusqllea and attemp ls to collect these motlu in areas of more diverse flora with li ttle or no bamboo were gener­ ally unsuccessful; and (2) the highest single day's catch (i. e., 53) was in a stand of al most pure Chusquea culeou, it appears probable that th is bamboo, or its debris, could be the food plant for the Sou th American Ne opseustidae. Moreover, the present known range of the eopseustidae lies within the range of the grass subfam ily Bambuseae. Thus, it i hypothesized that other genera of bamboos may serve as hosts for the old world genus Neopseustis. The Bambuseae possesses a broad pan tropical d is tribution, although present­ ly it is poorly represen ted in the floras of continental Africa and Australia. Among the grasses there is in general little evidence of direct dispersal across Ipop/allw pellai Sp. n. 43 : Be­ land, bu t bambusoids are ge nerally considered the group in which su ch dispersal ) Cisne, Parqu e acional "Los most likely could ha Ie occurred (Raven & Axelrod 1974). Present knowledge collected. Area dominated by of bamboo and neopseustid biogeography does therefore not seemingly contra­ I Filzroya (1lpressoidea (over­ dict the hypothesis of Neopseustidae being associated with bamboo. As almost where both the two new neo­ lalia, C. culeou (understorey) , . nothing is known abou t ecological tolerance and requiremen ts ofNeopseustidae, it might be such factors which govern the relatively limited distribution of Neopseustidae compared to that of the bamboo. Further collecting activities, especially in tropical Sou th America and Asia might easily clarify the dis­ 'es 41, 42. As can be seen, tribu tional pattern of Neopseustidae. entirely hidden under the rved in the most primitive , Graham (1950). A char­ 'Oe is the subposterior in­ ecimen the body and the arallel to the surface of the palpi are appressed to the 'pressed to the fron t of the the head , with the flexing bial palpi are curved under pi, being appressed to the 288

6. References Powell, J. It Gelccll Common, I. F. B., 1973: A new family of Dacnonypha (Lepidoptera) based on three new Raven, P. A species from Southern Australia, with notes on the Agathiphagidae. -J . AllSt. ent. menl>. Soc., 12: 11-23. Tindalc, N. Davis, D. R., 1975a: A review of the West Indian moths of the family Psychidae with de­ Hepiul scriptions of new taxa and immature stages. - Smithson. Contr. Zool. , 188: i-iii, 1-66. 1975b: Systematics and zoogeography of the family Neopseustidae with the proposal of a new superfamily (Lepidoptera: Neopseustoidea). - Smithson. Contr. Zool., 210: i-iii, 1-45. Manuscript 1978: A revision of the North American moths of the superfamily Eriocranioidea with the proposal of a new family, Acanthopteroctetidae (Lcpidoptcra). - Smith­ son. Contr. Zool., 251: i-iv, l-13l. Graham, M. W. R. de Y., 1950: Postural habits and colour-pattern evolution in Lepi­ doptera. - Trans. Soc. Br. Ent., 10: 217-232. Kristensen, N . P., 1968: The skeletal anatomy of the heads of adult Mnesarchaeidae and Neopsell stidac. - Ent. Meddr, 36: 137-151. 1978a: A new familia of from South America, with remarks on the phy­ logeny of the subordo Exoporia (Lepidoptera). - Ent. germ., 4: 272-294. 1978b: Observations on Anomoses hylecoetes (Anomosetidae), with a key to the hepialoid families (Insecta: Lepidoptera). - Steenstrupia, 5: 1-19. Kristensen, N. P., & Nielsen, E. S., 1979: A new subfamily of micropterigid moths from South America. A contribution to the morphology of the Micropterigidae, with a generic catalogue of the family (Lepidoptera: Zeugloptera). - Steenstrupia, 5:69-147. 1980a: The ventral diaphragm of primitive (non-ditrysian) Lepidoptera. A morpho­ logical and phylogenetic study. - Z. Zool. Syst. EvolForsch. 18: 123-146. 1980b (in press): Intrinsic proboscis musculature in non-ditrysian Lepidoptera­ : Structure and phylogenetic significance. In Cederholm, L. (ed.): Advances in systematics and phylogeny. - Ent. scand. suppl. 1980c (in press) : Abdominal nerve cord configuration in adult non-ditrysian Lepido­ ptera. - Int. J . Insect Morph. Embryo!. (in prep.): Double tube proboscis configuration in neopseustid moths (Lepidoptera: Neopseustidae) . Kiippers, P. Y. & Speidel, W. , 1980: Die systematische Stellung der Neopseustidae. - Atalanta, Miinnerstadt, 11: 55-65. Nielsen, E. S., 1980: Entomology. In Madsen, H. B., Nielsen, E. S. & 0dum, S. (eds): The Danish Scientific Expedition to Patagonia and Tierra del Fuego 1978-79. - Geogr. Tids., 80: 1-27. Pena G., L. E., 1966: Ensayo preliminar para divisar Chile en regiones entomofaunisticas, basadas especialmente en la familia Tenebrionidae (Col.). - Revta univ. Santiago 50-51: 209-220 [in Spanish, English translation in Postilla, 97: 1-17, 19661. Philpott, A., 1922: The wing-coupling apparatus in the Mnesarchaeidae. - N. Z. Jl Sci. Technol., 5: 275-276. 1925: On the wing-coupling apparatus of the Hepialidae. Trans. ent. Soc. London, 1925: 331-340. 1927: The genitalia of the Mnesarchaeidae. - Trans. Proc. N. Z. Inst., 57: 710-715, pI. 57. 2 9

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m) Lepidoptera. A morpho­ Jrsch.18: 123- 146. non-ditrysian Lcpidoptera­ ~derholm, L. (ed .): Advances pI. adult non-ditrysian Lepido­

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