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TENSE. U GET ONS peventy heise oie us eho ana eres uss ctu St clchioies wr saa ra cuaasstererig siewraw smal 1 CENUSH DO CCULALI Ui LB DILD Ret eyeraie sashes eosletenrelel aay sy ey ateeenchaselick giavenaine io ois ovaas + Key to the species based on coloration and markings ............... 15 Key to the species based on male genitalia ..............-.2:..++- DS Key to the species based on female genitalia ....................:- 30 Sechlonwl—— Species mln toll Oiamemie crseicictacus cielo arae lctonsyceevews tra @ eit 37 ISCKCCIMGLIFIEE PUSHEOUGL BPMN ooo 00cc 0000000 a000GcD000G0000 38 Bucculainpes solidagimella new Species) 4-54-44 4000 46 66 242 39

LB CCMAINES TRORUCTEG, IBSTENEIN 3 55500000000000000000004000006 41 BUGGKMinanagnellan Ghamperseeer aaa: 42

mee naninee (HOG NOH IBSEN 35000 0000c000000000000000000C 44 Isxniocibeniwas VORGMIG, WN SUXSIES oc00cc0c00b0G00000000000000 45 Bucculatrix sumulans new SpecieS ...........0.ecee cece eees 47 Bnceulioninice Tansee OsemwnwSeS scoccoaosb0ceanc00u0c000000 48 Bucculatrix parvinotata new species ..........-++-++e eee 49 ISCCHNOIMES OCMMBTCUE WAT SVESIES coascscccuscc00000000G0 49 Bueculatrin viguierae Mew Species’ .........-.+-++55+++--s- 50 Bucculatrix micropunctata new species .............-0+00-e 51 ISCO FIL? WMISIOUG TRG SNESIOS 5 og0c0cavgcgsodc0050000K0 52 Bucculatrix seneciensis new specieS .......+.-2-+ eee ee eee 54 Bucculatrix bieristata New SpeCieS ............--.--+++s-+- 55 IBCCHMON AEG CRINGE WI GAO 355500c000000d00000G0000000 56 SectionmUl= species) 17 to04) a aecneee ace cues deca asae geen eos 58 sinccullaine QMowclwGcs VAG > oo0000000000000000000000000 59 BOCES OCLUFISHGQIG IBSEN 550606000 00000000G0000008 60 Bucculatrix eurotiella Walsingham .............-+..+-+---- 61 ISnCeCMMonirige UCMEOTICOS® WMI 5 55000000000000000000000000 63 SCCUMOITES CHCCMCTUAD WEG? SNSSIGS cooocaeccesags00000000 64

NCCU MAL CCFUUOUNS IBBGIN 50000000ed0Gea0de0aac00000000 65 Bucculatrix separabilis new specieS .........-- eee eee ee eee 66 Bucculatrix brunnescens new species ..........-.-+2 eee eee 68 Bucculatrix evanescems new Speci€S ........-+--e eee ee eee 68 Bucculatrix benenotata new species .........0.0.ee ee eevee 69

nee Monn pWOCCOSG IRENE 5 5000000000008000000008n000706 70 INCE AES WOUTPCISHOS HON SUSSISS soccacsco0080008a805000 72 Bucculatrix franseriae new species ...........++-+-+++++--- 73 Bucculatrn stamtonelia Chambers 4.5-- 2520-0742 06-455+o- 74 Bucculatrix wmmaculatella Chambers .............++-+-++6- 76 Biunccnlonrusr comalia OWES oo0c0500000000000000000000000 77 Bucculatrix kimballi new species ........-...++--++eseeees 80


CONTENTS IS CUMMITTIGE COQUG BEBO ccococecnos cco 0s don DD SCboOOOONS 81 Bucculatrix ambrosiaefoliella Chambers ............---+..- 83 Bucculatrix palidula mew species ~....-..+.5..05545-. 495546 86 Bucculatwiralaentolasnewaspeclesm epee een pO ee o eee oor 87 IENOCIMOUAIE CUOMO? MEK FNSSIES socoocaccvoscco00s0000006 88 BucculatmaxangustatasmneynandsS olla yanmar er 89 NCCC? CUMING BW SVGISS oocccavcvccdcvcvcucssoc0g. 91 Bucculatniae plucnede MewaSpecles) Haeenne rete roe 92 IB OCCUNOMTUES CLYXOMOTOGNO IAB, coccovoavvccda0ec0v0b0cCDE 94 Bucculairix polymuvae new Species ..-..--.---.0+2---2500- 95 JEM CCUI TIES, SICCOSO WW SOSEIES. 5 oco0caccada00gccov000006 97 Bucculatrix submiens Walsingham .........-......-+--+:- 98 MCCUE SCTUOUCHOS IBN oc 000000060000 0000000 0000006 99 LENCCHM AE CHS IRAN sonccacsccocvgd Deven oaco0CenooE. 101 CCW WOCCOTOSE WHEW SUEGIES osccocasacn000Gca000006 103 ICC? CSOG BAG 5 o0000c0ccv0s00000g000000000000 104 SUCCES UAUDSOCTSOLG, IBFABIN 5 oc0000accacacccagvsa00006 105 ISMECMCirics IOCWQIAM Gs IRUBSS coc ccc odcnccon00 00000000000" 106 IBNCCWIOE SHMCOHOTIG BIEN 50000000000 b0ev00a000000000C 107 ISNCCMC Hiss UGDINO DET SOECICS oaccccccaccgocda0c0sag000s 109 IBOCCOMONTIES CHUCCUIG IBENEIN 50 000000000000000000000000006 111 Bucculatris, tmdenticola new Species) 4.4 on ed oe aoe 113 ISNCCHAU TIES SHOCHUDUNS WY SUSSIES 2c00cacccnsnnccon00000% 1S LNCCUWONTIES SCOPSE WRG SVECISS. oscccovccccccocscocuneoa00ce 116 Bueculatrax, angustisquamella Braun’ 4254-44504 6 +2 ooo soe 117 ISIC CTIGHITES COMMMDUCHIO HY SUSSIES 5 a0 o0000000000c0000000¢ 118 OCCA NE? SOTOTCUIG WHY SHEHES co cancucceccs000c0000000% 119 IBCCIM ORAS THOT DOMUCHANG IBTENOIN 5 ooc000nc0ca0n000e00000008 120 TNC CHHCNIUEE CLIFOSHOMIOHOL INNY SOSSIGS 5o0c00cccac0000ag00006 IZA Bucculainianenceliacinewaspe cles ane neee eee ere 122 IBnecubon EG Ven IME 55050500000 0000000000000000 00006 125 Sacsion UM =SHAGHES OD covcoccvcsoscccccvvvg0csncavc00 000006 126 Buceulanwarsponovelella Bi Scka ase een reno cee 126 Saciion IW—=snesies CO 1 90 coccascdoavvogeacncscacnoccncbns 128 iBucculatmempacrandellas Gnamlbers mann iene tee e 129 IBNCCHMCMTES (UOCTHONG IBUSOX o60500000c00ecacnD02a0 0000008 132 IBC COM OME? COMUOLUHL INN, SUSSCSoo000000000a000200000006 133 Bucculatrix platyphylla new species ............-+-+-+-+---- 134 Bucculatrax, ochisupfiusa mew, Species sane ee ene oe ore 135 IBC CUNO OSE tre OSCLONG CQUSINENS oo000000000000en0000000800 136 Bucculatrix qunquenotella Chambers ..................--- 138


COTWCONG WET SUSSIES o50000cccscvccvesca00uccs 140


Bucculatris, 2zopnopasia new Species 3.-..2.s0s.o6-ssne ses 142 BucoulanpclutigquosellarZellen sae nice een eee 144 BUGCKLOLK ta COnOnareHan@leMmens meee cin nie: 145 Bucculatri canadensisella| Chambers 2-5-2460 eens sees see 147 BUCCuLatKiaINUprOUISa Mew, SPECIES) aaa. 1. s nee ee as 149 Bueculatriapolyinia news SPEGleS sees ateccs eee cates seae.: 151 BuGceulatnem uteela Ghambersunn atari ciao 153

IB KCC UIAMES (FACOOMG. WEN SOYSSES 5560050000000000000000008 155 Bucculatrix. paropiila new, SpCGleS aasas- sas s5650 65 226 oO 157 ENCCNTALS YOGUCES IBENEIN 5 oan conoga0eK0 0g 0nGDGD0G DC GOS 158 Iicentniaee COPMSAAICIIG 0? SISSIES 6ooc0000ccK0cc0cob0b bee 160 IRENA CUOTUN IG NOT BUSSES 5060000000 0000n000000ee 162 IBMCCMGITULE COPE WHR SNASIES soccccccvs0ccugcb0g0000n0GS 163 Bncenteuimer COM MUG NNGVATICK oocaacnpecrocevcdcog0G0D DOWN 163 inecnlonncs OCHS NIGABOS goncaoncddnoccos0090900000006 165 leincculonnge Cunsianio. Wikswpi@lehe Go500c00000000c0g0000G0906 167 incenilenima CGC WET SHSSIES cooursaaasovgcv00c000K0n06 169 Sacifion Ws rscnes Oil 1) GE) cea ancics ovo actos cpa moe oie 171 Bie CMNGiT Ae CUCHCMNGD WY SUSSOS o60¢0000000005005b05000K6 171 neers CHS[MNCN WEY SOESES o5000000000000000000000C 173 Bnccilenrss COMOWMMANG) \BSEIBEN 51 oodc00000a0020000b0000 000" 174 SECHOIM Vel =—SPEClest OA mrdarne saiagaae ome dots aor eines a tcnaeee ae 175 Binccutionnrase jrommajQlocio QWi0SMS 4o550c000000c0000000000006 175 Section valli=— Species OS pesca tects sce essay ato es oi sane eer sets ae 178 Bnecniiatrize WECAMG ICSC occcocdcsccces0ccvdg00c000K0bCE 179 Saciion WIL =snacies GOW 89 sacosccosonsnnovagsnusaosccoods 180 BUCCI OUGCIGCHDIIG XENI 5 556500050000000000G000000 180 Buccs gossspudia Morrill .505c00c89000ee 900860000000 182 Bucculatris sphaeralceae new species ..................--- 184 Binccuiloimize Menino GimaHg IBWSE 5 o0c00600080000004000000000 185 List of the North American species of Bucculatrix ............. 188 PEE AMER TBE D re payers caacniens cee erste et ene thts ds Avsss best suelo outed narra) «cyte 189

EERELANATION Om JPIGUIRIOS <..ds 6 sala aco. aa cid ob eee Uae clei nio ela Sree Os 190









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Bucculatrix is one of the most easily recognized genera of Micro- lepidoptera. The elongate pointed face, tufted head, basal eye-cap of the antenna, and, in the male, the notched first segment of the flagellum assure recognition by casual examination.

In the early stages, species of this genus may be recognized by the short, very narrow, almost thread-like mines, and’ in the later stages by the feeding-pattern on the leaf made by the exposed larvae.

About 200 species are now known, occurring in all continents ex- cept New Zealand, with about three-fourths of the species in the north- ern hemisphere. About one-half are within the area of this mono- graph. In number of species, the genus is best represented in warmer regions, particularly in arid regions, where any collection is likely to contain representatives of it, some of them probably undescribed.

A total of 99 North American species are included in the genus. Of the 54 species listed in McDunnough’s Check List of the Lepidop- tera of Canada and the United States of America, Part II, Microlepi- doptera (1939), eight (capitialbella Chambers, albicapitella Chambers, rileyi Frey & Boll, crescentella Braun, chrysothamm Braun, tetrella

a INST iivi ion


Braun, althaeae Busck, pertenuis Braun) are reduced to synonymy. Bucculatrix submitens Walsingham, previously known only from Mex- ico, is here recorded from Arizona. Bucculatrix gossypiella Morrill, of Mexico, is included because of the probability of its introduction in the Southwest. Bucculatrix needhami Braun was recently (1956) de- scribed. Fifty species are described as new. Examples of all of the species here treated have been examined, with the exception of niveella Chambers and wmmaculatella Chambers of which no types are in exist- ence; these latter two species are assigned to their probable positions.

In connection with the evaluation of Chambers’ types (e.g. the series of luteella and packardella at the Museum of Comparative Zool- ogy) the following quotation is of interest. “‘ But a few years ago I began to make a collection to be preserved as types of all my species... . Unfortunately, during my absence in Colorado, the greater part of this collection was destroyed. One or more specimens of the greater num- ber of species were fortunately preserved, and most of the other species can be supplied. This collection is now in the Cambridge Museum. It contains types—pinned and spread—of something over 200 species.” (Chambers, Canad. Ent., IX, 39, 1877.) There is thus no assurance that his “‘types”’ actually represent the species described. Such un- certainty is apparent in the presumed types of B. packardella and B. luteella in the Museum of Comparative Zoology.

In the preparation of the slides of genitalia certain precautions must be observed to insure the retention of the tufts and patches of special- ized scales on the intersegmental membrane between the seventh and eighth segments and on the eighth abdominal segment in the female. Such tufts of scales often constitute the only reliable characters for species separation. They are loosely attached and tend to float off en masse in spite of the utmost care; it is therefore advisable to make measurements and sketches during the progress of the work. In the female, the minute size and the transparency of the abdominal wall per- mit examination of the bursa copulatrix and signum im situ, and it is not necessary and usually not desirable to separate the anal segments from the rest of the abdomen. However, if this is desired, the separa- tion should be made between the sixth and seventh segments, retaining segment 7 connected to the genital segments. In the species of Section IV, the position of segment 7 overlying the basal half of segment 8

a o*)


has been retained in slides and drawings to show the natural position of the specialized scale structures attached to the intersegmental mem- brane and lying ventral to the ostium. The degree of telescoping of segment 8 into segment 7 may vary on the slides, and thus the apparent positions of the fringing scales of segment 7 and the specialized scale patches on the intersegmental membrane may vary in relation to one another and to the specialized scale tufts on segment 8. In the males of Section IV, no attempt was made to spread the harpes; in spreading, the thread-like vinculum may be broken; the twisting of the harpes to a ventral position is shown to best advantage unspread. Because of the weak articulation of the harpes on the vinculum in other sections, they may be lost easily. Unless otherwise noted, all slides were made by the writer.

With the exception of the setal map of the larva of Bucculatrix canadensisella Chambers, all figures were drawn by the writer.

All genitalia drawings were made by transmitted light, using a binocular microscope (fitted with a micrometer disc) for the gross structure, with details added by examination under a compound micro- scope. The degree of magnification of the figures of genitalia was determined by the amount of detail. The figures of less specialized female genitalia are one-half the scale of males. Small inset figures of specialized characters to a higher magnification may accompany the larger figures; such details are often visible only under the compara- tively high magnification of a compound microscope. In so far as pos- sible, ventral structures are shown by full lines, dorsal by broken lines. In particular this refers to female genitalia where characters of value are present on the dorsal surface of segment 8. It will be noted that the bursa copulatrix is omitted in most of the figures; it is impossible to show details of the complete signum without a higher magnification and larger figures than is desirable.

A total of over 2500 specimens has been examined in the study of the genus. Included are some 750 specimens in my own collection, many of them reared, and including types, allotypes or paratypes of 19 new species; over 900 specimens in the United States National Mu- seum, including types, allotypes or paratypes of 30 new species; over 250 specimens in the Canadian National Collection, including types, allotypes or paratypes of 9 new species; some 150 in the collections

MEM. AMER. ENT. Soc., 18.


in the Academy of Natural Sciences of Philadelphia, including the Clemens’ types and the Darlington Collection; the collections of Cornell University, some 250 specimens, including a recent collection of a large amount of material from the Southern Appalachians, an area scarcely represented in other collections, and several species from Arizona; the Cornell collections have contributed types and type material of 7 new species; the collection of A. E. Brower of Augusta, Maine, with rep- resentatives of three new species; Charles P. Kimball of Barnstable, Massachusetts and Sarasota, Florida submitted material from New York and Massachusetts, and a less extensive collection from Florida, including type material of three new species; from J. R. Eyer, State College, New Mexico, from the Los Angeles County Museum, from C. L. Remington, Yale University, and from J. McDunnough, Halifax, Nova Scotia, I have received material for study. To all of these, grate- ful acknowledgement is made. To Dr. J. F. Gates Clarke of the United States National Museum, for slides of types, for notes from literature not available to me, and for the many courtesies extended throughout the progress of the work, I express my sincere thanks.

Under Literature Cited (p. 189) is a short list of books and papers referred to in the text; of these, Friend’s paper on Bucculatrix can- adensisella Chambers (1927) is a valuable detailed study of a typical species of the genus.

Abbreviations when used in the text in referring to the location of material are as follows: A.F.B.Coll. (A. F. Braun Collection) : A.N.S.P. (Academy of Natural Sciences of Philadelphia) ; B.M. (Brit- ish Museum); C.N.Coll. (Canadian National Collection); M.C.Z. (Museum of Comparative Zoology); U.S.N.M. (United States Na- tional Museum). Location of material from other collections is listed without abbreviation or with obvious abbreviations.

Genus Bucculatrix Zeller

Bucculatrix Zeller, 1839. Isis, XXXII, 214. Generotype, albedinella Zeller (= boyerella Duponchel).

Ceroclastis Zeller, 1848. Linn. Ent., III, 295, t. 2, fig. 47. Generotype, nigri- comella Zeller.

Face smooth, obliquely produced into a point extending well below the eyes (fig. 1), head tufted with long hair-scales, a forward-directed section of the tuft


attached to the vertex, the upward- and backward-directed section attached to the posterior part of the epicranium; tongue short, naked, maxillary palpi rudi- mentary, labial palpi minute, one-segmented, sometimes visible as a slender fila- ment (fig. 1) ; antennae shorter than the wing length, basal segment of antenna (scape) enlarged forming an eye-cap clothed with projecting scales, and anteri- orly fringed with long bristly setae; first segment of flagellum long, and in male typically with a deep notch (slight in some species) ; eye subglobose, large in some desert-inhabiting species.

Fore wings (Plates III and IV) lanceolate, more or less acuminate, costa and subcosta fused into a strongly sclerotized marginal vein, retinaculum a curved membranous hook in both sexes; radius obsolescent near base, R, from before middle of cell, Rz, Rg and R4 from near end of cell, Ry closely approxi- mate to or connate or stalked with the stalk of R; +My, or rarely absent; media two-branched, M, stalked with R;, rarely separate; cubitus unbranched, running through the middle or near to middle of wing, the cell thus chiefly in the upper half of the wing, cubitus sometimes obsolescent or absent beyond cell (figs. 20, 23); Ist anal, the fold; A, not forked, strong, reaching margin beyond middle of dorsum; Ag short, closely approximating the wing margin.

Hind wings lanceolate, from less than half the breadth of the fore wing to nearly as wide; frenulum of male a single strong seta, frenulum of female, two closely associated, or sometimes partially fused setae; Sc + R, to near middle of costa, R, to costa near apex, media two-branched, cell open, cubitus unbranched ; two anal veins often distinct.

Posterior tibiae with long hairs above and below, the upper decumbent, the latter pendant; middle pair of spurs articulating near base of segment, the inner twice or three times the length of the outer, apical pair shorter, inner twice the length of the outer.

Abdomen of the male in most of the species with an eversible scale sac, which is a mid-dorsal invagination of the body wall in the suture between the second and third segments (fig. 177, et al.), the distal circular area clothed with scales; the walls of the sac form the stalk when the sac is protruded and the scales expanded (fig. 82a) ; when expanded it resembles a flower-head of a Com- posite, the scales of different proportions and lengths in the several species. It may serve as an alluring organ.

Male genitalia. Although specialization within the sections and groups has resulted in apparent great diversity of structure, certain characters (or their modifications) are, with few exceptions, common to all the species of the genus: harpe weakly articulating with vinculum or closely associated with anellus and partially fused with it, of various shapes, commonly more or less inflated, with rounded apex (representing cucullus) usually defined by setal armature, or, harpe sometimes deeply bilobed (a specialization), the lobes distinguished by differences of setal armature, basal angles of costa produced as free arms, saccu- lus not defined; transtilla present in ambrosiaefoliella, then articulating with the

MEM. AMER. ENT. Soc.. 18.


basal costal arms of harpe, and perhaps in aimsliella represented by a narrow sinuate band; anellus a broad or slender cone, often with lateral supporting sclerotizations ; aedeagus more or less cylindric, often sinuate, or with apex pro- duced beyond the aperture, aperture often elongate and armed with opposing teeth, proximal end rarely dilated and semi-globose, entrance of penis dorsal at the proximal end, cornuti rarely present; vinculum a mere thread (Plates XXIII to XX XIX) or a broader band rarely anteriorly greatly produced, often with a narrow thin median perpendicular plate; a definitive gnathos rarely present in completeness, lateral arms seldom present, gnathos most often undifferentiated, or represented merely by a minutely spinulose strip ventral to the anal tube for which the term subscaphium (sensu Diakonoff, 1954) is here adopted; socii usu- ally two erect setose lobes, rarely reduced and nearly obsolete; uncus absent, ex- cept in a few species. Segment 8 modified in one section of the genus (Plates XLIV and XLV).

Female genitalia. The most distinctive characteristics of the female geni- talia are the signum, consisting of a series of spined ribs, usually forming a ring nearly or quite encircling the bursa copulatrix near its posterior end, and the single pair of apophyses, those of the ninth segment (a second pair, those of the eighth segment, initiated in some specialized species) ; position of ostium vari- ous, opening either near center of sclerotized basal half of segment 8, or (in Section VIII) at the posterior margin of the sclerotized basal half of segment 8, or at the anterior margin of segment 8, or in the intersegmental membrane be- tween segments 7 and 8; specializations may include the fringing of the poste- rior margins of the sclerotized section of segment 7 with specialized scales, the development of tufts or rows or patches of specialized (non-striated) scales on the intersegmental membrane and on segment 8, sclerotized outgrowths on seg- ment 8, and finally extreme modification of the inner margins of the ovipositor lobes for rasping or piercing—the rasping rods—and the transfer of the func- tion of the lobes to the terminal portion of the vagina, with its specialized vagi- nal setae (Braun, 1958) ; ductus bursae usually sclerotized for a greater or less length near ostium; inception of ductus seminalis adjacent to ostium, or more generally at the junction of the sclerotized section with the more anterior mem- branous section; in B. platyphylla only, a posterior lobe of the bursa copulatrix receives the ductus seminalis. Signum near posterior end of bursa and usually constricting it.

The preceding brief and of necessity inadequate description of the genitalia scarcely suggests the great diversity of structure within the genus. Genitalic structure is often correlated with food-plant groups and is the basis for division of the genus into sections.

Among the known food plants of Bucculatrix are representatives of some twenty-five plant families. In our area, members of the Com- positae are probably hosts to nearly two-thirds of the species; trees or


shrubs of the amentiferous families to a fifth or more of the species. The larvae of a few species feed upon members of each of several plant families.

The species may be grouped into two main divisions on the basis of larval habits. (1) The typical division, which includes the majority of the species (Sections II to VIII). In this division, the larvae are leaf-miners in the first two instars and part of the third, and in the fourth and fifth instars (with few exceptions) feed exposed, usually on the under surface of the leaf, skeletonizing it or sometimes completely consuming the leaf substance. leaving irregular holes. A few species in this division are miners throughout larval life, never feeding ex- ternally. (2) A division apparently confined to this continent, in which the larvae, so far as known, are gall-formers, feeding within the gall throughout larval life, or, if miners in the earlier instars, stem borers in the later instars (Section I).

In those species comprising the typical division of the genus, the egg may be deposited on either the upper or the lower surface of a leaf. Eggs are usually flattened ovoid, and cemented to the leaf surface by an adhesive material which encircles them in a narrow band (figs. 35, 36, 37). An apparent exception to this general shape is the egg of Bucculatrix thurberiella Busck, which is described as elongate, pro- jectile shaped with about 10 projecting ridges and stands perpendicular to the leaf” (Morrill, 1927). The surface of the egg is variously sculptured; the hexagonal pattern of sculpturing is most common, and is that found in species of the amentiferous plant feeders (fig. 37) ; the sculpturing may take the form of longitudinal ridges converging to- ward the micropylar end, or the egg may be irregularly ridged in a more or less concentric pattern, the ridges breaking into knobs toward the micropylar end (fig. 36). Upon hatching, the larva enters directly into the leaf tissue.

The mines are narrow, almost thread-like tracks with parenchyma entirely consumed, and vary in length in different species from two or three to five or six centimeters, or even more in thin-textured leaves. They may at first follow a vein, abruptly turning out onto the leaf blade (fig. 52a), or be contorted (fig. 54b), or long and irregularly winding (fig. 43). A darkening of the leaf adjacent to the earliest portion of the mine, observed in some species, suggests a resting period preceding

MEM. AMER. ENT. Soc., 18.


resumption of feeding. In those few species which are miners through- out larval life, the mines of the fourth and fifth instars broaden into characteristically shaped blotches (figs. 45, 56). In the linear mines, the frass is deposited as a central blackish line, with the grains of ex- crement tending to appear as separated particles; a small blotch at the beginning of the mine contiguous to the egg-shell is free of frass; here the frass is contained within the egg-shell.

The first and second instars and part of the third are passed in the mine. On leaving the mine, without further feeding, the larva spins the flat ‘‘ moulting cocoon,” usually a thin smooth sheet of silk within which it moults. After this moult, the larvae are external feeders. A second moulting cocoon similar to the first but slightly larger is spun at the end of the fourth instar. In general, the exposed larvae feed on the underside of the leaf, skeletonizing small patches of leaf, which may have a characteristic shape and appearance, recognizable from field ex- perience. In the fifth instar, in some species, the entire leaf tissue may be consumed, the small holes margined by veinlets. When the leaf is jarred, the larva may drop down on a silken thread. In the last in- stars, the larvae of B. divisa, feeding on the leaves of Balsamorrhiza, of B. arnicella on Arnica cordifolia, and of B. salutatoria on Artemisia tridentata enter the leaf through circular holes and mine out the leaf tissue, with only head and thorax inside the mine. Such mines resem- ble Coleophora mines, but can be recognized by the position of the cir- cular entrance holes at one edge of the mine (fig. 48b), instead of near the center.

In the two instars of the leaf-mining stage, the larva is flattened, apodal, the head lying in a horizontal plane, but as it is a tissue feeder, there is no great modification of mouth parts. The first and second moults take place within the mine; during the third instar the larva leaves the mine. In this instar and in the two following, the structure is that of the usual tissue-feeding larva (fig. 32). The body is cylin- drical, the head nearly vertical; three pairs of thoracic legs are present, the prothoracic smaller than the meso- and metathoracic legs and fur- nished with but a single claw. The prolegs are comparatively long and slender, present on abdominal segments 3, 4, 5, 6, and 10, the abdom- inal prolegs bearing two transverse rows of uniordinal crochets, the anal prolegs a single transverse row. ‘The anal prolegs in dorsal aspect


(fig. 33) are slender and diverging, presenting a good field recognition character. The last instars of those species which are miners through- out larval life (e.g. angustata) agree in general with the corresponding instars of the typical exposed-feeding larva, except that the head lies in a more nearly horizontal plane. The setal pattern of the fifth instar (fig. 34) is constant throughout this division of the genus with but minor variations; it will be noted from the setal map that, in this divi- sion of the genus, the setae are comparatively long.

The other main division of the genus (Section I) includes such species as Bucculatrix fusicola Braun, B. needhami Braun, B. viguierae new species, which are gall-formers and feed throughout larval life within the gall, and such species as B. solidaginiella new species and B. cuneigera Meyrick, which are leaf-miners in the early instars, and stem- borers in the later instars. In the gall-former, B. needhami, Dr. Need- ham’s studies (1948) show that a non-feeding instar is interpolated between the last feeding instar and the pupal stage. This condition is probably present in all of the gall-formers, although to date no care- ful studies have been made of other species. In all observed instances, the full-fed larva passes the winter in the gall, emerging from the gall in the spring by a minute circular aperture, and spinning a typical Buc- culatricid cocoon. Of the stem-borers, the mine of B. cuneigera only has been observed; in this instance the larva spins, in the end of the mine, a dense flat circular chamber, similar to the moulting cocoon of the species of the typical division of the genus, in which it passes the winter, emerging in the spring to crawl to the tip of a growing shoot and bore into it.

Bucculatrix needhami Braun may be taken as an example of larval structure in the gall-forming members of the genus. The larva in the last feeding instar, and still within the gall when full-fed, differs from the fifth instar larva of the leaf-mining and exposed-feeding species in several notable characters (fig. 31). The body is moniliform, with deep constrictions between the segments; the head is small, spinnerets non-functional; all three pairs of thoracic legs bear but a single claw; prolegs absent, their position merely indicated by a flattish area; setae are minute, almost microscopic. The larva is not capable of locomo- tion; “it did not creep, but lay on its side with the front end thrown back in a J-shaped hook, the head at the tip of the hook. It spun no

MEM. AMER. ENT. Soc., 18.


silk, not even enough to hold back the pellets of frass’’ (Needham, Journ. N. Y. Ent. Soc., LVI, 43-50, 1948). In this condition the larva passes the winter in the gall. In the spring a moult takes place within the gall, and the larva which emerges is of the normal lepidop- terous form with functional spinnerets and long setae; it agrees with the typical fifth instar exposed-feeding larva in structure, except that a single claw is present on all three pairs of thoracic legs. Here was a non-feeding instar, interpolated between larval and pupal stages; a clear case of hypermetamorphosis (Needham, /.c.). This non-feeding larva gnaws a hole through the wall of the gall, emerging and spinning a typical ridged Bucculatricid cocoon. For additional details refer to Dr. Needham’s paper in the Journal of the New York Entomological Society.

In the stem-borers of this division of the genus, larvae of which are leaf-miners in the early instars, the change to normal lepidopterous form must occur with the moult taking place at the end of the mine, since the power of active locomotion is necessary to reach the growing tip of the food plant.

The ridged cocoon is characteristic of the genus Bucculatriy, and is one of the most beautiful and intricate structures to be observed in the Lepidoptera. The manner of its spinning is briefly described below ; for a detailed description of the process, the paper by R. E, Snodgrass in the Report of the Smithsonian Institution, 1920 (1922) entitled The ribbed-cocoon maker of the apple,” pages 496 to 509, and Plates 2 and 3, should be consulted. The larva spins, on whatever substratum is to serve as the location of the cocoon, a thin oval mat of silk; in some species a palisade”’ of upright silken poles, each consisting of several united silk strands, previously encloses the area (fig. 57). The cocoon is Spun in two sections, the two sections joined on meeting; spinning begins at what is to be the posterior end. The ridges are formed by the projecting ends of a series of loops spun from side to side, the larva backing away as the work progresses. At about two-thirds or three- fourths of what is to be the final length of the cocoon, the larva reverses its position and commences to spin at the other end of the mat, grad- ually enclosing itself and bringing the two sections together. The ridges seldom meet exactly and the joint is usually discernible and often conspicuous. Within this ridged and somewhat open structure, a close-woven inner lining is spun. To complete the entire structure may


require an incredibly short time—a few hours, or more often, as much as a half day. The proportions of the cocoons of the several species may vary from rather broad and stout to elongate and slender; the number of ridges is constant (with some slight variation) for the spe- cies. Shape and number of ridges may thus be diagnostic characters of a species (Plates VII, VIII, IX). In color, cocoons vary from pure white to dark brown. The pale green color of the cocoon of Bucciula- trix flourensiae new species is unique in the genus.

The ridges of the cocoon are ill-defined in some species or are oc- casionally wanting. Cocoons spun by parasitized larvae are often not characteristic of the species and may be abnormal in some respect, for example, smooth instead of ridged.

Pupae (figs. 38, 39, 40) with the appendages free from the body wall; abdominal segments 3-7 movable in the male, 3—6 in the female ; the tenth abdominal segment with a projection on each side ending in a short strong spine, and a small dorsal tubercle at its anterior margin bearing a pair of minute spines; on dorsal surface of abdominal seg- ments 2—7, a row of minute strong spines along the anterior margins; the hind wings usually concealed by the fore wings, but sometimes their tips visible; foreshadowing the genital openings of the imago is the single genital opening of the male on the ninth segment (fig. 39) ; and the two genital openings of the female on the eighth and ninth segments respectively (fig. 40).

On emergence, the pupa is thrust through the anterior end of the cocoon, exposing half or more of its length. The skin splits trans- versely between vertex and prothorax and longitudinally along the prothorax and mesothorax (fig. 42a).

The majority of the species have but one generation in a year; two, or even three generations occur in some of the oak-feeding species, and in some Composite feeders. The winter is passed in the pupal state in species of temperate latitudes or more or less humid regions; it is probable that the period of emergence in arid regions is dependent on the season of precipitation. Some few species may hibernate in the imaginal state.

The percentage of parasitism is unusually high; a third or more of the larvae of any rearing may prove to be parasitized; sometimes as high as 90 percent of the larvae may be parasitized.

MEM. AMER. ENT. Soc., 18.


Wing expanse of the imagoes varies from 4 mm. to 14 or 15 mm. in our largest species, members of the gall-forming section of the genus (Section 1).

Except in Section I (the gall-formers and stem-borers), character- ized by very oblique or longitudinal streaks on a white ground ( Plate I, figs. 3, 4, 5, 6), the wing pattern of the imagoes conforms to a gen- eral type, which is however often obscured by expansion or contraction or obsolescence of marks. This pattern consists of an alternation of dark and white (or whitish) oblique bars or streaks from costa; the first of these dark areas from base to about one-third, followed by a pale bar, a second dark bar, a second pale bar separated from a whitish bar or streak near apex by the darker color or a dusted area; on the dorsal margin, a little distad of the first costal bar, a white or whitish area separated from a second whitish area or streak by a darker area which (in most species) includes a patch of black or black-tipped raised scales, sometimes conspicuous, which is one of the identifying characters of the genus. This series of marks may perhaps be most clearly recognized in some species of Section IV ( Plate II, figs. 15, 16,. 17, 18). Any one or more of these bands or streaks may be obscured or somewhat displaced by dark dusting, or be obsolescent ; for example, in a white species, the darker bands are reduced or obsolescent, and the wing would then be described as white, with darker or dusted streaks (figs. 7, 8, 13, 14) ; in other species, the condition is reversed, and the extent of dark marks increased (figs. 11, 17, 18); all gradations may exist between these various modifications of the general wing pattern (figs. 9, 10, 12). A pale median basal streak is sometimes present. There is thus an apparent great diversity of wing markings.

In general, resemblance of wings can not be assumed to indicate relationship; variation within a species may be greater than differences between species, and unrelated species may superficially resemble one another. For example, Bucculatrix sexnotata Braun and B. callistricha new species are both dark brown with silvery marks; the former is a Composite feeder and belongs to Section II, the latter, on Corylus, to Section IV, the two unrelated, as evidenced by examination of geni- talia. Genitalia must be examined for certain determination except in the most distinct species.


Bucculatrix is an isolated genus, without near relationship to any existing genus. It has generally been associated with the Lyonetiidae, but is sometimes regarded as constituting a separate family, Buccula- trigidae. The median position of the strong cubital vein (lower mar- gin of the cell) in the middle or above the middle of the wing, and the tendency for the veins to disappear by obsolescence rather than by coalescence are distinctive Lyonetiid characters, and thus ally Buc- culatrix to the Lyonetiidae whether or not the genus is regarded as a separate family.

Characters supporting family rank for the genus are the more prim- itive pupa with appendages free in contrast with the obtect pupae of the reduced genera of the Lyonetiidae, the antennal structure of the male, the eversible scale sac of the male abdomen, the characteristic signum and the single pair of apophyses of the female, the larval structure and habits and the unique cocoon.

The diversity of genitalic structure and the accompanying special- ization, with similar specializations in widely separated geographic areas, indicates an ancient genus with a long period of development and differentiation. This points to wide distribution in ancient times with specializations characterizing the several sections of the genus devel- oped before the period of isolation.

There is no evidence to substantiate any of the proposed theories of the phylogenetic origin of Bucculatrix. Various conjectures have been made. Meyrick (Proc. Linn. Soc. New South Wales, VII (Se- ries 2nd), 1892, p. 601) makes the following statements: Probably a development of the Nematobola group, but no immediate connection can be made out.... The peculiar larval habits may be compared with those of Nematobola” [now assigned to the Yponomeutidae]. The larval structure and habits as described for Nematobola candescens Meyrick (J.c., p. 593) are very similar to those of Bucculatrix. The antennae of the male in Comodica Meyrick (Lyonettiidae) with deep notch immediately above basal joint” (/.c., p. 561) suggests a possible relationship to Bucculatrix. Markings of several of the species in- cluded in Comodica are similar to those of some species of Bucculatrix. Later, Meyrick (1927) derives the Lyonetiidae, including Bucculatrix, from the Tineidae as “a specialised development of the Tineidae.”’

MEM. AMER. ENT. Soc., 18.


Philonome Chambers, stated by Forbes (1923) to be hardly dis- tinct from Bucculatriv”’ is shown by genitalia, which are Lyonetiid, to be unrelated to Bucculatrix.

Examination of genitalia has disclosed an unusual number of sib- ling species.1 These may be either allopatric or sympatric species. Such pairs of species may be characterized by essentially similar geni- talia which however differ in the more minute morphological charac- ters. Some such pairs, in addition to morphological differences, may be separated by differences of habitat requirements. Among the sibling species are Bucculatrix variabilis Braun and B. separabilis new species, which feed on the same food plant at the same time; these had been considered varieties of a single species; genitalia demonstrate their re- productive isolation. Bucculatrix arnicella Braun and B. tridenticola new species are almost indistinguishable in the imaginal state; by geni- talia they are distinct; one is a forest species, the other, a species of the sagebrush desert. Some pairs of species (as B. evanescens new species and 6. benenotata new species) are easily separated by wing marks, but the similar and often unique character of the genitalia indicate very close relationship.

In the present treatment of Bucculatrix, the species are grouped into eight sections :

Tat Species ml =hG ee cree res | ere pe ow

I: Spectres 764 even. he eee ne ere po, 59 DEL, Species 165) 38-2. eae teeny er eee p. 126 IVE Species” 66-90! 2 aaa ae eee arene p. 128 Wi “Species; QS 935 es ee eee > 1/7 Il Wile Species O40 at een ray are era area M/S Wilk “Species “95: 22 Sires = ican as eee DO; 7s Vale “Species 96=99 2s Ben. egret: anne p. 180

These sections are based primarily on genitalic structure, which is how- ever often correlated with food plant groups.

1 Sibling species are defined as ‘“ Pairs or groups of closely related species which are reproductively isolated but morphologically identical or nearly so” (Methods and Principles of Systematic Zoology, Mayr, Linsley and Usinger, 1953).



A section or subsection or group can not be assumed to be derived from another. Each must have originated independently from ances- tral stock, although evolutionary trends may be followed in some sec- tions or groups. Highly specialized characters of the genitalia have developed in each section, distinct from those of other sections, often resulting in great diversity of form within a section. Within each section there may be several ramifications, each culminating in species with specialized characters. Divergence of structure from the general type of the section or group is regarded as a specialization; the most complex (in genitalia structure) are considered to be the most special- ized. Thus the degree of specialization determines the sequence of species.

Key to the Species of Bucculatrix Based on Coloration and Markings *

1. Ground color of fore wings white, creamy white or pale, i.e., the greater area of wing light, the markings formed by spots or streaks of darker or dark-tipped scales; the basic pale ground color sometimes obscured by dusting of dark-tipped scales, the wings thus sometimes appearing dark

Ground color of fore wings not white; whitish ocherous, ocherous to dark brown or black, or sometimes irrorate. (Included here are species of which the general aspect of the ground color is ocherous, although the scales may shade from white at base through ocherous to brown or TUGSOES Bie HOS) Adoneeonescloctuceo ome coke uaa cum ame toe crc ocrar 60

2. Ground color clear white or creamy white, often lustrous, with little or no dusting except that the scales of the marks may be dark-tipped (if

dusting present minute and scarcely evident) ................... 3 Basic ground color white, but often obscured by dusting of dark-tipped SCAMES S:6.0'5.6 Grd eeollaee ee Gag ELGG ST Ose tore sare Ree RS Lo se acter me ot 48

3. Size usually large (8 to 15 mm. wing expanse) ; wings (of some species ) almost wholly white or creamy white, markings if any, at least in part longitudinal; mostly gall-formers or stem-borers in larval state .... 4

2 Because of the similarity of many species, sometimes unrelated, and variation within a species, this key, except in the case of species with distinct and well-defined markings, can serve only as a partial aid to determination. In variable species there are some individuals in which the general color may appear dark even though the basic ground color (i.e. bases of scales) is white. Although the aspect of two species may be different, the characters are often too intangible to be expressed in a key. Genitalic characters are the only c